Goodyera repens

Table of Contents



SPECIES: Goodyera repens
Abrahamson, Ilana L. 2013. Goodyera repens. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: [].


lesser rattlesnake plantain
creeping rattlesnake plantain
dwarf rattlesnake plantain
northern rattlesnake plantain

The scientific name of lesser rattlesnake plantain is Goodyera repens (L.) R. Br. (Orichidaceae) [28,31,47].

Goodyera repens var. ophiodes [30,66,84].


Photo used by permission of David McAdoo and the North Carolina Native Plant Society

In December of 2012 an extensive search was done to locate information on lesser rattlesnake plantain (see FEIS's list of source literature). The following paragraphs provide details from the available information.


SPECIES: Goodyera repens
North American distribution of lesser rattlesnake plantain. Map courtesy of the Flora of North America Association. 2013, 29 April.

Lesser rattlesnake plantain is native to North America [28] and Eurasia [87]. This review describes lesser rattlesnake plantain as it occurs in North America. Lesser rattlesnake plantain is transcontinental in Canada and occurs in continuous and discontinuous populations in the United States. Its northern range occurs in Alaska, Yukon, and Newfoundland. In the eastern United States, its southern range reaches North Carolina and Tennessee; in the West, it occurs in disjunct populations in Arizona, New Mexico, and Colorado [28].

States and provinces [78]:
United States: AK, AZ, CO, CT, ID, MA, MD, ME, MI, MN, MT, NC, NH, NM, NY, OH, PA, SD, TN, VA, VT, WI, WV, WY
Canada: AB, BC, LB, MB, NB, NF, NS, NT, ON, PE, QC, SK, YT

Site Characteristics:
Lesser rattlesnake plantain is associated with undisturbed, late-successional and old-growth forests. In much of its range, lesser rattlesnake plantain occurs in shady, moist, coniferous or mixed forests [2,11,17,28,31,40,53,57,69] on mossy or humus-covered ground [28,80], and can sometimes occur in bogs or swamps [28]. However, it also occurs in xeric conditions in the southern Appalachian Mountains including high, rocky, exposed cliffs and slopes of the Carolina hemlock (Tsuga caroliniana) community [77]; xeric heath balds and forest heathlands [56,86]; and relatively xeric eastern hemlock (Tsuga canadensis) forest [79,85].

Lesser rattlesnake plantain occurs at elevations ranging from sea level to about 9,500 feet (0-2,900 m) [28]. This varies with geographic location. For example, lesser rattlesnake plantain occurs at high elevations in disjunct populations in Colorado (8,065-8,258 feet (2,460-2,515 m)) [27] and New Mexico (7,450 feet (2,270 m)) [45] and occurs at low elevation in eastern Canada (450 feet (135 m)) [39].

Soils: Lesser rattlesnake plantain can tolerate a range in soil moisture regimes, from wet sites in the northern part of its range to dry sites in the southern Appalachians. In its northern range, lesser rattlesnake plantain typically grows on sites that have a thick organic layer or moss cover [2,43,63,69,76,80]. Soils on these sites tend to have a high water holding capacity or moderate drainage [34,53,69,83]. In Alaska, lesser rattlesnake plantain sometimes occurs on sites with permafrost [83]. In xeric coniferous heathlands of the southern Appalachian Mountains, lesser rattlesnake plantain occurs on dry, nutrient poor, leached, acidic sites and is classified as an acidophile [56,86]. Lesser rattlesnake plantain is considered an indicator species in serpentine woodlands of the southern Appalachian region [79].

Plant Communities:
Lesser rattlesnake plantain occurs in many vegetation types including late-successional coniferous forests [80,81,83], mixed hardwood forests [12,74], somewhat dry forests [31], heath balds and forest heaths [86], and bogs and swamps [28]. It is commonly associated with white spruce (Picea glauca) in boreal forests [22,49,53,76,80,81,83], and eastern hemlock in eastern montane forests [13,21,56,61,68,77,79,85].

Lesser rattlesnake plantain is an indicator of mature vegetation throughout its range. On the Tanana River, Alaska, lesser rattlesnake plantain is an indicator of the mature floodplain successional stage dominated by white spruce (150-200 years old) [32]. In southwestern Manitoba, lesser rattlesnake plantain was most common in mature northern whitecedar (Thuja occidentalis)-balsam fir (Abies balsamea)-black spruce (Picea mariana)-white spruce communities [37]. In boreal mixedwood stands in northern Ontario, lesser rattlesnake plantain was an indicator of unharvested forest [73]. In mature boreal mixedwood forest stands of western Canada, lesser rattlesnake plantain was an indicator of conifer forest patches (mostly dominated by white spruce) rather than broadleaf forest patches (mostly quaking aspen (Populus tremuloides)) (P=0.003) [82].

See Table 1 for additional descriptions of plant communities and site characteristics in areas where lesser rattlesnake plantain typically occurs or has been observed. See the Fire Regime Table for a list of plant communities in which lesser rattlesnake plantain may occur and information on the fire regimes associated with those communities.


SPECIES: Goodyera repens
Botanical description:
Lesser rattlesnake plantain is a perennial orchid arising from short rhizomes that are shallowly rooted in mosses and the upper organic layer [2,31,63]. It has erect stems and typically grows 4 to 6 inches (10-15 cm) tall but can reach 8 inches (20 cm) [31]. The mostly basal leaves are typically ovate or ovate-lanceolate [28,31]. Seven to 36 small (3-5 mm long) white or pale green flowers occur close together on one-sided racemes [28,31]. Like other species in this genus, lesser rattlesnake plantain has perfect, bilateral flowers [31]. The fruit is a capsule that contains numerous tiny and lightweight seeds [2,5].

Lesser rattlesnake plantain leaves and inflorescence. Photos used by permission of David McAdoo and the North Carolina Native Plant Society.

Raunkiaer [67] life form:

Seasonal Development:
Lesser rattlesnake plantain is an evergreen perennial with leaves active throughout the winter [29]. The flowering period ranges from early July to mid-September depending on its location. In Montana, flowering begins in late July or early August and extends into mid-September [2]. In northern Michigan, flowering was observed from mid-July to late August with a median bloom date of July 31 [35]. In the mountains of Virginia, lesser rattlesnake plantain flowers in July and August [52]. After the plant blooms, the flowering rosette dies, and buds along the rhizome are released from dormancy [17].

Regeneration processes:
Lesser rattlesnake plantain regenerates sexually through seeds and vegetatively via rhizomes. Rhizomes that arise from seed require 5 years to produce a rosette of evergreen leaves, which produces a flowering stalk and seed capsules after approximately 3 more years [48]. The tiny seeds of lesser rattlesnake plantain are wind dispersed [2,5]. No information was available on lesser rattlesnake plantain soil seed banks in North America at the time of this writing (2013); however, research from old-growth Norway spruce (Picea abies) forests in Norway suggests that lesser rattlesnake plantain does not have a long-lived seed bank [71,72]. Goodyera species often form colonies due to their rhizomatous growth habit [17].

Interbreeding among Goodyera species is possible where mixed-species populations occur. See Kallunki [35] for details. Although lesser rattlesnake plantain is geitonogamous, self-pollination is unlikely to occur in nature because the flowers are slightly protandrous. When experimentally pollinated, lesser rattlesnake plantain collected from northern Michigan was capable of self-fertilization. Bumblebees are the principal pollinators of Goodyera species in North America [35].

Like other terrestrial orchids, lesser rattlesnake plantain requires a mycorrhizal symbiont for seed germination and early seedling development [4,10,24]. The symbiont is usually Rhizoctonia goodyerae-repentis. Lab experiments have demonstrated that lesser rattlesnake plantain can be germinated asymbiotically in the presence of an external carbohydrate source, whereas inoculated lesser rattlesnake plantain does not require a carbohydrate [24,65]. Other lab experiments with seeds collected from Pinnacle Mountain, South Carolina, found that germination rates were greater when seeds were exposed to light (37% germination) compared to those incubated in continuous darkness (25% germination), and light had no effect on seed mortality [55].

Successional status:
Lesser rattlesnake plantain is a shade-tolerant, mid- to late-successional species generally occurring in stands more than 70 years old [22,23,32,33,36,46,69,80,83].

In white spruce-fir and black spruce forests across the North American taiga from Newfoundland to Alaska, lesser rattlesnake plantain was prevalent in mature, undisturbed forest stands (i.e., stands that had no evidence of fire, insect infestation, or other major natural disturbance). Lesser rattlesnake plantain had a frequency of 59% in white spruce-fir stands and 54% in black spruce stands [39].

Chronosequence studies of primary succession on Alaskan floodplains indicate that lesser rattlesnake plantain typically occurs in mature white spruce stands that range from 120 to 300 years old [32,80,81,83]. On the Tanana River in interior Alaska, lesser rattlesnake plantain occurs in 150- to 300-year-old stands after balsam poplar-dominated stands shift to closed white spruce and develop a nearly complete ground cover of feathermosses (Hylocomiaceae) [32,80,81]. One of the studies on the Tanana River floodplain characterized lesser rattlesnake plantain as an indicator of the mature white spruce stage; it was not found in earlier successional stages or the subsequent black spruce stage [32]. Near the Chena River in interior Alaska, lesser rattlesnake plantain had 40% frequency in 120-year-old white spruce stands, but was absent from younger willow and balsam poplar stands and older (220-year-old) white spruce-black spruce and black spruce/Sphagnum stands [83]. Similarly, along the Peace River in British Columbia, lesser rattlesnake plantain is generally more common on sites dominated by white spruce, feather moss (Hylocomium splendens), and Schreber's moss (Pleurozium schreberi) and less common on sites dominated by earlier successional balsam poplar [76].

On boreal sites in eastern Canada, lesser rattlesnake plantain occurred only in mid- to late-successional stands [22,23]. Around Lake Duparquet in northwestern Quebec, lesser rattlesnake plantain appeared in stands dominated by balsam fir, black spruce, and paper birch (Betula papyrifera) 74 to 143 years after fire [22]. A study of mature second-growth stands and undisturbed forest stands at 2 stages of development (senescent and old growth) in the Gaspé Peninsula, Quebec, found that lesser rattlesnake plantain was not present in any early second-growth forests (50-year-old, even-aged balsam fir-white birch stands) but had 6% frequency in senescent forests (70-year-old, even-aged balsam fir stands) and 3.3% frequency in old-growth forests (90-year-old, uneven-aged balsam fir-black spruce and uneven-aged balsam fir-white spruce stands) [23]. In upland plantations and managed stands of jack pine (Pinus banksiana) and black spruce in Ontario, lesser rattlesnake plantain frequency corresponded with the greatest time since disturbance and stand establishment. Measurements taken over 20 years showed that lesser rattlesnake plantain increased from 6% to 11% mean relative frequency in jack pine stands, and from 4% to 17% mean relative frequency in black spruce stands [33].

On southern boreal mixedwood sites in Michigan, Manitoba, and Saskatchewan, lesser rattlesnake plantain is associated with late-successional, mixed coniferous forest [46,69]. In a study of primary succession across a sand-dune chronosequence bordering northern Lake Michigan, lesser rattlesnake plantain was found in trace amounts (<0.1% cover) on dune ridges that ranged from 175 to 835 years old. It was not present on the earliest successional dunes (25 years old) dominated by beach grass (Ammophila breviligulata), or on 55- to 175-year-old dunes dominated by evergreen shrubs. It was first present after the development of mixed pine forest (225-440 years old) consisting of eastern white pine (Pinus strobus), red pine (Pinus resinosa), white spruce, balsam fir, northern whitecedar, and paper birch [46]. In the mixedwoods of Manitoba and Saskatchewan, lesser rattlesnake plantain occurs after the development of the feathermoss stratum, which is rarely present until stands are more than 70 years old [69].

Lesser rattlesnake plantain is uncommon in Montana, but it sometimes occurs in older forests dominated by Douglas-fir with a well-developed moss layer dominated by feather moss and Schreber's moss [2], or on northerly aspects in late-successional spruce (Picea) or subalpine fir (Abies lasiocarpa) forests [63].

Postfire regeneration strategy [75]:
Rhizomatous herb, rhizome in organic layer

Fire adaptations and plant response to fire:
Fire adaptations: Although no information was available on the immediate effects of fire on lesser rattlesnake plantain at the time of this writing (2013), most literature describes lesser rattlesnake plantain as poorly adapted to fire [2,63,70,88]. Its roots, like those of other Goodyera species, rarely penetrate the mineral soil [17]. Its rhizomes are shallowly rooted in mosses and the upper organic layer [2,63], which makes them susceptible to damage from fire. Habitat and germination/establishment requirements of lesser rattlesnake plantain (i.e., shade, thick organic layer, intact mycorrhizal community) make it unlikely to "respond well to fire" [2]. Rowe [70] characterizes lesser rattlesnake plantain as a fire avoider; these are defined as "shade-tolerant species that slowly reinvade burned areas; late successional, often with symbiotic requirements". Avoiders cannot establish immediately after fire but only after some habitat requirement has been met (e.g., sufficient shade or humus accumulation). These species occur late in succession in areas with long fire-return intervals [70].

Plant response to fire: Lesser rattlesnake plantain is consistently absent from postfire plant communities, suggesting that it is sensitive to fire. In a study on the effects of wildfire on understory plant community development in boreal mixedwood communities near Slave Lake, Alberta, lesser rattlesnake plantain was present in prefire sample plots but absent from postfire and manipulated (burned and sapling/herb layers removed) plots [49]. In northeastern Minnesota, lesser rattlesnake plantain had 20% and 13% frequency in unburned second-growth (about 70 years old), mixed conifer-hardwood stands but was nearly absent from 2 similar stands that were burned by wildfires. On 1 burned site, lesser rattlesnake plantain did not occur 3, 5, or 14 years after fire. On a second burned site, where "little or no soil burn occurred even though it was a hot fire", lesser rattlesnake plantain was not observed 3 or 5 years after fire and had a 3% frequency 11 years after fire [38]. Lesser rattlesnake plantain was absent from postfire communities in jack pine and black spruce plantations in northeastern Minnesota [3] and oak-pine (Quercus-Pinus) forest in North Carolina [25], despite its presence in similar unburned sites. In vegetation that was characterized as "standing virgin forest communities" before fire in northeastern Minnesota, only 1 individual lesser rattlesnake plantain was observed in the first 5 years after fire [62]; no information was given on prefire plant communities or unburned controls.

In the southern Appalachian Mountains in northeastern Tennessee, lesser rattlesnake plantain was observed in some postfire sites but at a lower frequency than in unburned controls. Zimmerman [88] examined the effects of prescribed fire on the herbaceous layer by comparing 6 burned sites with 6 unburned controls. Postfire measurements were taken between 2 and 6 years after fire. Leaf litter, canopy cover, soil moisture, and soil pH were "slightly affected" by the fire. Lesser rattlesnake plantain was absent from some of the 6 burned sites, whereas it was one of the most common herbaceous species in their unburned counterparts. Lesser rattlesnake plantain occurred 20 times in burned plots and 46 times in unburned controls. The research does not indicate whether the plants in burned sites occurred in unburned patches within the burn perimeter, were burned and sprouted after fire, or were newly germinated seedlings. The author suggests that lesser rattlesnake plantain "is an indicator of moist woods and is not adapted to a regular fire regime" [88].

On the Lewis and Clark National Forest in Montana, where lesser rattlesnake plantain is an imperiled species, stand-replacement fires (wildfire and prescribed) and timber harvests (clearcuts and seed tree cuts) may have eliminated or greatly reduced the occurrence of lesser rattlesnake plantain. One population may have been eliminated by the Turkey Fire, and 2 other populations were greatly reduced following stand-replacement wildfires. One of these wildfires left an area of unburned habitat where lesser rattlesnake plantain is still present. No prefire data on lesser rattlesnake plantain occurrence was available, but it likely occurred on these sites before the fire removed the shade and organic layers of the forest floor. Populations of lesser rattlesnake plantain extended up to the boundary of, but not into, a recent clearcut and severely burned forest in 2 river drainages of the Little Belt Mountains, suggesting that lesser rattlesnake plantain cannot withstand clearcutting and/or high severity fire [63].

Because lesser rattlesnake plantain occurs in a variety of communities, it is subject to many different fire regimes, but most of them have relatively long fire-return intervals. The Fire Regime Table summarizes characteristics of fire regimes for vegetation communities in which lesser rattlesnake plantain may occur. For additional information on fire regimes of communities where lesser rattlesnake plantain occurs, see FEIS reviews of dominant species such as white spruce and black spruce.

Federal legal status:

Other status:
Information on state- and province-level protection status of plants in the United States and Canada is available at NatureServe. As of 1984, lesser rattlesnake plantain was ranked as apparently secure to vulnerable in Colorado, imperiled to vulnerable in Montana and North Carolina, imperiled in Arizona, Pennsylvania, and possibly Virginia, critically imperiled to imperiled in Kentucky and West Virginia, and critically imperiled in Massachusetts and Tennessee. It was possibly extirpated from Connecticut and Maryland [60]. Lesser rattlesnake plantain was ranked as apparently secure or secure in Canada, except in Nova Scotia where it is vulnerable and Prince Edward Island where it is imperiled [60].

Other management:
Lesser rattlesnake plantain is negatively affected by timber harvesting due to its need for shade and undisturbed forest floor. A Montana Field Guide [59] states that “any activities or events that substantially reduce the overstory and/or removes the duff/litter layer, such as thinning and logging operations or severe fires are likely to have adverse impacts” to lesser rattlesnake plantain. Lesser rattlesnake plantain is typically absent in the short to mid-term after all types of timber harvest. It was absent for at least 10 years after harvest in oak-hickory (Quercus-Carya) forests in southwestern Virginia and northeastern West Virginia [12]; it was observed 17 years after thinning and shelterwood harvest in the Lewis and Clark National Forest, Montana [63]; and it was absent from stands less than 70 years old after harvests in eastern boreal, balsam fir forest on the Gaspé Peninsula, Canada [23]. In uncut white spruce/alder communities near Fairbanks, Alaska, lesser rattlesnake plantain had 14% to 21% frequency but was absent during the first 2 years after logging. Of the 50 species present in the plots before logging, only 4 species–including lesser rattlesnake plantain–were absent after logging [26]. In the Lewis and Clark National Forest, Montana, lesser rattlesnake plantain was absent from a seed-tree harvest stand except in 2 small, undisturbed patches within the harvest unit. Abundant lesser rattlesnake plantain populations were observed in adjacent uncut stands. The author states that "it is likely that the loss of shade, organic matter, and moss during the clearcut, seed tree and dozer piling treatments, changed the microclimate on the forest floor to such an extent that lesser rattlesnake plantain could not survive or reproduce after treatment" [63].


SPECIES: Lesser rattlesnake plantain
The following table provides fire regime information that may be relevant to lesser rattlesnake plantain habitats. Follow the links in the table to documents that provide more detailed information on these fire regimes.

Fire regime information on vegetation communities in which lesser rattlesnake plantain may occur. This information is taken from the LANDFIRE Rapid Assessment Vegetation Models [42], which were developed by local experts using available literature, local data, and/or expert estimates. This table summarizes fire regime characteristics for each plant community listed. The PDF file linked from each plant community name describes the model and synthesizes the knowledge available on vegetation composition, structure, and dynamics in that community. Cells are blank where information is not available in the Rapid Assessment Vegetation Model.
Northern and Central Rockies Southwest Great Lakes
Northeast Southern Appalachians
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
Minimum interval
Maximum interval
Southwest Forested
Lodgepole pine (Central Rocky Mountains, infrequent fire) Replacement 82% 300 250 500
Surface or low 18% >1,000 >1,000 >1,000
Ponderosa pine-Douglas-fir (southern Rockies) Replacement 15% 460    
Mixed 43% 160    
Surface or low 43% 160    
Riparian forest with conifers Replacement 100% 435 300 550
Southwest mixed conifer (cool, moist with aspen) Replacement 29% 200 80 200
Mixed 35% 165 35  
Surface or low 36% 160 10  
Southwest mixed conifer (warm, dry with aspen) Replacement 7% 300    
Mixed 13% 150 80 200
Surface or low 80% 25 2 70
Spruce-fir Replacement 96% 210 150  
Mixed 4% >1,000 35 >1,000
Northern and Central Rockies
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
Minimum interval
Maximum interval
Northern and Central Rockies Forested
Douglas-fir (warm mesic interior) Replacement 28% 170 80 400
Mixed 72% 65 50 250
Lodgepole pine, lower subalpine Replacement 73% 170 50 200
Mixed 27% 450 40 500
Ponderosa pine (Black Hills, high elevation) Replacement 12% 300    
Mixed 18% 200    
Surface or low 71% 50    
Ponderosa pine-Douglas-fir Replacement 10% 250   >1,000
Mixed 51% 50 50 130
Surface or low 39% 65 15  
Upper subalpine spruce-fir (Central Rockies) Replacement 100% 300 100 600
Great Lakes
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
Minimum interval
Maximum interval
Great Lakes Forested
Conifer lowland (embedded in fire-resistant ecosystem) Replacement 36% 540 220 >1,000
Mixed 64% 300    
Eastern white pine-eastern hemlock Replacement 54% 370    
Mixed 12% >1,000    
Surface or low 34% 588    
Great Lakes spruce-fir Replacement 100% 85 50 200
Minnesota spruce-fir (adjacent to Lake Superior and Drift and Lake Plain) Replacement 21% 300    
Surface or low 79% 80    
Northern hardwood-eastern hemlock forest (Great Lakes) Replacement 99% >1,000    
Northern hardwood maple-beech-eastern hemlock Replacement 60% >1,000    
Mixed 40% >1,000    
Pine-oak Replacement 19% 357    
Surface or low 81% 85    
Red pine-eastern white pine (less frequent fire) Replacement 30% 166    
Mixed 47% 105    
Surface or low 23% 220    
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
Minimum interval
Maximum interval
Northeast Forested
Beech-maple Replacement 100% >1,000    
Eastern white pine-northern hardwood Replacement 72% 475    
Surface or low 28% >1,000    
Northern hardwoods-eastern hemlock Replacement 50% >1,000    
Surface or low 50% >1,000    
Northeast spruce-fir forest Replacement 100% 265 150 300
Southern Appalachians
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
Minimum interval
Maximum interval
Southern Appalachians Forested
Appalachian oak forest (dry-mesic) Replacement 6% 220    
Mixed 15% 90    
Surface or low 79% 17    
Eastern hemlock-eastern white pine-hardwood Replacement 17% >1,000 500 >1,000
Surface or low 83% 210 100 >1,000
Eastern white pine-northern hardwood Replacement 72% 475    
Surface or low 28% >1,000    
Mixed-mesophytic hardwood Replacement 11% 665    
Mixed 10% 715    
Surface or low 79% 90    
Southern Appalachian high-elevation forest Replacement 59% 525    
Mixed 41% 770    
*Fire Severities—
Replacement: Any fire that causes greater than 75% top removal of a vegetation-fuel type, resulting in general replacement of existing vegetation; may or may not cause a lethal effect on the plants.
Mixed: Any fire burning more than 5% of an area that does not qualify as a replacement, surface, or low-severity fire; includes mosaic and other fires that are intermediate in effects.
Surface or low: Any fire that causes less than 25% upper layer replacement and/or removal in a vegetation-fuel class but burns 5% or more of the area [8,41].


1. Achuff, Peter L.; La Roi, George H. 1977. Picea-Abies forests in the highlands of northern Alberta. Vegetatio. 33(2/3): 127-146. [83873]
2. Achuff, Peter; Schassberger, Lisa. 1991. Status review of Goodyera repens, USDA Forest Service, Region 1, Lewis and Clark National Forest, Montana. Report to the Lewis and Clark National Forest. Helena, MT: Montana Natural Heritage Program. 52 p. [86815]
3. Ahlgren, Clifford E. 1960. Some effects of fire on reproduction and growth of vegetation in northeastern Minnesota. Ecology. 41(3): 431-445. [207]
4. Arditti, Joseph. 1967. Factors affecting the germination of orchid seeds. Botanical Review. 33(1): 1-97. [86852]
5. Arditti, Joseph; Ghani, Abdul Karim Abdul. 2000. Tansley review no. 110. numerical and physical properties of orchid seeds and their biological implications. New Phytologist. 145(3): 367-421. [86847]
6. Bailey, L. H., Jr. 1882. Limits of Michigan plants. Botanical Gazette. 7(8/9): 105-108. [86830]
7. Barclay-Estrup, P.; Duralia, T. E.; Harris, A. G. 1991. Flowering sequence of the orchid genus Goodyera in Thunder Bay, Alaska. Rhodora. 93(874): 141-147. [15379]
8. Barrett, S.; Havlina, D.; Jones, J.; Hann, W.; Frame, C.; Hamilton, D.; Schon, K.; Demeo, T.; Hutter, L.; Menakis, J. 2010. Interagency Fire Regime Condition Class Guidebook. Version 3.0, [Online]. In: Interagency Fire Regime Condition Class (FRCC). U.S. Department of Agriculture, Forest Service; U.S. Department of the Interior; The Nature Conservancy (Producers). Available: [2013, May 13]. [85876]
9. Barringer, Kerry; Clemants, Steven E. 2003. The vascular flora of Black Rock Forest, Cornwall, New York. Journal of the Torrey Botanical Society. 130(4): 292-308. [86841]
10. Baskin, Carol C.; Baskin, Jerry M. 2001. Seeds: ecology, biogeography, and evolution of dormancy and germination. San Diego, CA: Academic Press. 666 p. [60775]
11. Beaudry, Leisbet; Coupe, Ray; Delong, Craig; Pojar, Jim. 1999. Plant indicator guide for northern British Columbia: boreal, sub-boreal, and subalpine biogeoclimatic zones (BWBS, SBS, SBPS, and northern ESSF). Victoria, BC: British Columbia Ministry of Forests, Forestry Division Sciences Branch. 134 p. [70419]
12. Belote, Russell Travis. 2008. Diversity, invasibility, and stability of Appalachian forests across an experimental disturbance gradient. Blacksburg, VA: Virginia Polytechnic Institute and State University. 152 p. Dissertation. [86816]
13. Boyd, Ty B. 2011. Impacts of the loss of hemlock canopy on southern Appalachian herbaceous communities. West Lafayette, IN: Purdue University. 70 p. Thesis. [86828]
14. Breitung, August J. 1957. Annotated catalogue of the vascular flora of Saskatchewan. The American Midland Naturalist. 58(1): 1-72. [86832]
15. Burn, C. R.; Friele, P. A. 1989. Geomorphology, vegetation succession, soil characteristics and permafrost in retrogressive thaw slumps near Mayo, Yukon Territory. Arctic. 42(1): 31-40. [69761]
16. Carleton, T. J.; Maycock, P. F. 1980. Vegetation of the boreal forests south of James Bay: non-centered component analysis of the vascular flora. Ecology. 61(5): 1199-1212. [14734]
17. Case, Frederick W., Jr. 1988. The jewel orchids of North America. American Orchid Society Bulletin. 57(7): 758-765. [62674]
18. Catling, Paul M.; Kostiuk, Brenda. 2011. Some wild Canadian orchids benefit from woodland hiking trails - and the implications. The Canadian Field-Naturalist. 125(2): 105-115. [86831]
19. Clark, Julie Bennett. 2012. The vascular flora of Breaks Interstate Park, Pike County, Kentucky, and Dickenson County, Virginia. Lexington, KY: University of Kentucky. 87 p. Thesis. [86836]
20. Clarkson, Roy B. 1966. The vascular flora of the Monongahela National Forest, West Virginia. Castanea. 31(1): 1-119. [86813]
21. Cooperrider, Tom S.; Thorne, Robert F. 1964. The flora of Giles County, Virginia. II. Castanea. 29(1): 46-70. [80547]
22. De Grandpre, Louis; Gagnon, Daniel; Bergeron, Yves. 1993. Changes in the understory of Canadian southern boreal forest after fire. Journal of Vegetation Science. 4(6): 803-810. [23019]
23. Desponts, Mireille; Brunet, Genevieve; Belanger, Louis; Bouchard, Mathieu. 2004. The eastern boreal forest old-growth balsam fir forest: a distinct ecosystem. Canadian Journal of Botany. 82(6): 830-849. [50267]
24. Downie, D. G. 1940. On the germination and growth of Goodyera repens. Transactions of the Botanical Society of Edinburgh. 33(1): 36-51. [86853]
25. Dumas, Shay; Neufeld, Howard S.; Fisk, Melany C. 2007. Fire in a thermic oak-pine forest in Linville Gorge Wilderness Area, North Carolina: importance of the shrub layer to ecosystem response. Castanea. 72(2): 92-104. [72033]
26. Dyrness, C. T.; Viereck, L. A.; Foote, M. J.; Zasada, J. C. 1988. The effect on vegetation and soil temperature of logging flood-plain white spruce. Res. Pap. PNW-RP-392. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. 45 p. [7471]
27. Flaig, Jeanette H. 2007. A vascular plant inventory of the eastern San Juan Mountains and vicinity in southern Colorado. Laramie, WY: University of Wyoming. 125 p. Thesis. [86840]
28. Flora of North America Editorial Committee, eds. 2013. Flora of North America north of Mexico, [Online]. Flora of North America Association (Producer). Available: [36990]
29. Givnish, T. J. 1990. Leaf mottling: relation to growth form and leaf phenology and possible role as camouflage. Functional Ecology. 4(4): 463-474. [86848]
30. Gleason, Henry A.; Cronquist, Arthur. 1991. Manual of vascular plants of northeastern United States and adjacent Canada. 2nd ed. New York: New York Botanical Garden. 910 p. [20329]
31. Hitchcock, C. Leo; Cronquist, Arthur; Ownbey, Marion. 1969. Vascular plants of the Pacific Northwest. Part 1: Vascular cryptogams, gymnosperms, and monocotyledons. Seattle, WA: University of Washington Press. 914 p. [1169]
32. Hollingsworth, Teresa N.; Lloyd, Andrea H.; Nossov, Dana R.; Ruess, Roger W.; Charlton, Brian A.; Kielland, Knut. 2010. Twenty-five years of vegetation change along a putative successional chronosequence on the Tanana River, Alaska. Canadian Journal of Forest Research. 40(7): 1273-1287. [83520]
33. Hunt, Shelley L.; Gordon, Andrew M.; Morris, Dave M.; Marek, George T. 2003. Understory vegetation in northern Ontario jack pine and black spruce plantations: 20-year successional changes. Canadian Journal of Forest Research. 33(9): 1791-1803. [65102]
34. Jeglum, John K.; He, Fangliang. 1995. Pattern and vegetation--environment relationships in a boreal forested wetland in northeastern Ontario. Canadian Journal of Botany. 73(4): 629-637. [26695]
35. Kallunki, Jacquelyn A. 1981. Reproductive biology of mixed-species populations of Goodyera (Orchidaceae) in northern Michigan. Brittonia. 33(2): 137-155. [62675]
36. Kembel, Steven W.; Dale, Mark R. T. 2006. Within-stand spatial structure and relation of boreal canopy and understorey vegetation. Journal of Vegetation Science. 17(6): 783-790. [86842]
37. Ko Heinrichs, Derrick. 2009. Ecology of northern white-cedar (Thuja occidentalis L.) stands at their northwestern limit of distribution in Manitoba, Canada. Winnepeg, MB: University of Manitoba. 121 p. Thesis. [86812]
38. Krefting, Laurits W.; Ahlgren, Clifford E. 1974. Small mammals and vegetation changes after fire in a mixed conifer-hardwood forest. Ecology. 55(6): 1391-1398. [9874]
39. La Roi, George H. 1967. Ecological studies in the boreal spruce-fir forests of the North American taiga. I. Analysis of the vascular flora. Ecological Monographs. 37(3): 229-253. [8864]
40. La Roi, George H.; Hnatiuk, Roger J. 1980. The Pinus contorta forests of Banff and Jasper National Parks: a study in comparative synecology and syntaxonomy. Ecological Monographs. 50(1): 1-29. [8347]
41. LANDFIRE Rapid Assessment. 2005. Reference condition modeling manual (Version 2.1), [Online]. In: LANDFIRE. Cooperative Agreement 04-CA-11132543-189. Boulder, CO: The Nature Conservancy; U.S. Department of Agriculture, Forest Service; U.S. Department of the Interior (Producers). 72 p. Available: [2007, May 24]. [66741]
42. LANDFIRE Rapid Assessment. 2007. Rapid assessment reference condition models, [Online]. In: LANDFIRE. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Lab; U.S. Geological Survey; The Nature Conservancy (Producers). Available: [2008, April 18] [66533]
43. Larsen, James A. 1980. Boreal communities and ecosystems: the broad view. In: Larsen, James A., ed. The boreal ecosystem. New York: Academic Press: 128-236. [64915]
44. Lausi, D.; Nimis, P. L. 1985. Quantitative phytogeography of the Yukon Territory (NW Canada) on a chorological-phytosociological basis. Vegetatio. 59(1/3): 9-20. [86833]
45. Legler, Ben S. 2010. A floristic inventory of Vermejo Park Ranch, New Mexico and Colorado. Laramie, WY: University of Wyoming. 177 p. Thesis. [86839]
46. Lichter, John. 1998. Primary succession and forest development on coastal Lake Michigan sand dunes. Ecological Monographs. 68(4): 487-510. [29313]
47. Lichvar, Robert W.; Kartesz, John T. 2012. North American Digital Flora: National wetland plant list, version 3.0, [Online]. Hanover, NH: U.S. Army Corps of Engineers, Engineer Research and Development Center, Cold Regions Research and Engineering Laboratory; Chapel Hill, NC: The Biota of North America Program (Producers). Available: [2013, April 19]. [84896]
48. Luer, Carlyle A. 1975. Goodyera repens (Linnaeus) R. Brown in Aiton, Hort. Kew. ed. 2. 5:198. 1813. In: The native orchids of the United States and Canada excluding Florida. New York: The New York Botanical Garden: 146. [86895]
49. Lypowy, Jaime Nicole. 2009. The effect of forest fire on understory plant community development: a comparison of pre- and post-fire vegetation in the boreal mixedwood forest. Edmonton, AB: University of Alberta. 168 p. Thesis. [86335]
50. MacHattie, L. B.; McCormack, R. J. 1961. Forest microclimate: a topographic study in Ontario. Journal of Ecology. 49(2): 301-323. [86838]
51. MacQuarrie, Kate; Lacroix, Christian. 2003. The upland hardwood component of Prince Edward Island's remnant Acadian forest: determination of depth of edge and patterns of exotic invasion. Canadian Journal of Botany. 81(11): 1113-1128. [47131]
52. Massey, A. B. 1953. Orchids in Virginia. Castanea. 18: 107-115. [86835]
53. Maycock, P. F.; Curtis, J. T. 1960. The phytosociology of boreal conifer-hardwood forests of the Great Lakes region. Ecological Monographs. 30(1): 1-36. [62820]
54. Maycock, Paul F. 1961. The spruce-fir forests of the Keweenaw Peninsula, northern Michigan. Ecology. 42(2): 357-365. [62688]
55. McKinley, Tonya Cherie. 1995. Factors affecting seed germination of the terrestrial orchid Goodyera repens var. ophioides Fernald and the in vitro culture of orchidaceous mycorrhizae. Clemson, SC: Clemson University. 132 p. Dissertation. [86855]
56. McLeod, Donald Evans. 1988. Vegetation patterns, floristics, and environmental relationships in the Black and Craggy Mountains of North Carolina. Chapel Hill, NC: University of North Carolina. 222 p. Dissertation. [60570]
57. Meades, W. J.; Moores, L. 1989. Forest site classification manual: A field guide to the Damman forest types of Newfoundland. Forest Resources Development Agreement FRDA Report 003. St. Johns, NF: Environment Canada. 295 p. [49220]
58. Mitchell, Richard S.; Tucker, Gordon C. 1994. Flora of an unusually diverse virgin and old-growth forest area in the southern Adirondacks of New York. Bulletin of the Torrey Botanical Club. 121(1): 76-83. [86843]
59. Montana Natural Heritage Program. 2012. Northern rattlesnake-plantain - Goodyera repens. In: Montana field guide, [Online]. Helena, MT: Montana Natural Heritage Program (Producer). Available: [2013, April 29]. [86877]
60. NatureServe. 2013. NatureServe Explorer: An online encyclopedia of life, [Online]. Version 7.1. Arlington, VA: NatureServe (Producer). Available [69873]
61. Newell, Claire L.; Peet, Robert K. 1998. Vegetation of Linville Gorge Wilderness, North Carolina. Castanea. 63(3): 275-322. [71985]
62. Ohmann, Lewis F.; Grigal, David F. 1977. Some individual plant biomass values from northeastern Minnesota. Res. Note NC-227. St. Paul, MN: U.S. Department of Agriculture, Forest Service, North Central Forest Experiment Station. 2 p. [8151]
63. Phillips, H. W. 1995. Conservation strategy, northern rattlesnake-plantain (Goodyera repens), Lewis and Clark National Forest. Unpublished report on file at: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory, Missoula, MT. 6 p. [+ maps]. [86857]
64. Poindexter, Derick B.; Murrell, Zack E. 2008. Vascular flora of Mount Jefferson State Natural Area and environs, Ashe county, North Carolina. Castanea. 73(4): 283-327. [86814]
65. Purves, S.; Hadley G. 1976. The physiology of symbiosis in Goodyera repens. New Phytologist. 77(3): 689-696. [86854]
66. Radford, Albert E.; Ahles, Harry E.; Bell, C. Ritchie. 1968. Manual of the vascular flora of the Carolinas. Chapel Hill, NC: The University of North Carolina Press. 1183 p. [7606]
67. Raunkiaer, C. 1934. The life forms of plants and statistical plant geography. Oxford: Clarendon Press. 632 p. [2843]
68. Rogers, Robert S. 1980. Hemlock stands from Wisconsin to Nova Scotia: transitions in understory composition along a floristic gradient. Ecology. 61(1): 178-193. [62813]
69. Rowe, J. S. 1956. Uses of undergrowth plant species in forestry. Ecology. 37(3): 461-473. [8862]
70. Rowe, J. S. 1983. Concepts of fire effects on plant individuals and species. In: Wein, Ross W.; MacLean, David A., eds. The role of fire in northern circumpolar ecosystems. SCOPE 18. New York: John Wiley & Sons: 135-154. [2038]
71. Rydgren, K.; Okland, R. H.; Hestmark, G. 2004. Disturbance severity and community resilience in a boreal forest. Ecology. 85(7): 1906-1915. [86849]
72. Rydgren, Knut; Hestmark, Geir. 1997. The soil propagule bank in a boreal old-growth spruce forest: changes with depth and relationship to aboveground vegetation. Canadian Journal of Botany. 75(1): 121-128. [27760]
73. Spalvieri, Cristina. 2009. Understory species response to partial harvesting in boreal riparian buffers. Thunder Bay, ON: Lakehead University. 110 p. Thesis. [86818]
74. Stearns, Forest. 1951. The composition of the sugar maple-hemlock-yellow birch association in northern Wisconsin. Ecology. 32(2): 245-265. [86837]
75. Stickney, Peter F. 1989. Seral origin of species comprising secondary plant succession in northern Rocky Mountain forests. FEIS workshop: Postfire regeneration. Unpublished draft on file at: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory, Missoula, MT. 10 p. [20090]
76. Timoney, Kevin P.; Peterson, George; Wein, Ross. 1997. Vegetation development of boreal riparian plant communities after fire, flooding, and logging, Peace River, Canada. Forest Ecology and Management. 93(1-2): 101-120. [27427]
77. Tucker, G. E. 1972. The vascular flora of Bluff Mountain, Ashe County, North Carolina. Castanea. 37(1): 2-26. [73963]
78. U.S. Department of Agriculture, Natural Resources Conservation Service. 2013. PLANTS Database, [Online]. Available: [34262]
79. Ulrey, Christopher Joseph. 2002. The relationship between soil fertility and the forests of the southern Appalachian region. Raleigh, NC: North Carolina State University. 234 p. Dissertation. [86819]
80. Van Cleve, K.; Viereck, L. A.; Dyrness, C. T. 1996. State factor control of soils and forest succession along the Tanana River in interior Alaska, U.S.A. Arctic and Alpine Research. 28(3): 388-400. [65672]
81. Van Cleve, Keith; Viereck, Leslie A. 1981. Forest succession in relation to nutrient cycling in the boreal forest of Alaska. In: Fire and succession in conifer forests of North America. New York: Springer-Verlag: 185-211. [50633]
82. Varela, Virginia Chavez. 2010. Patterns and causes of variation in understory plant diversity and composition in mature boreal mixedwood forest stands of western Canada. Edmonton, AB: University of Alberta. 163 p. Dissertation. [86820]
83. Viereck, Leslie A. 1970. Forest succession and soil development adjacent to the Chena River in interior Alaska. Arctic and Alpine Research. 2(1): 1-26. [12466]
84. Voss, Edward G. 1972. Michigan flora. Part I: Gymnosperms and monocots. Bulletin 55. Bloomfield Hills, MI: Cranbrook Institute of Science; Ann Arbor, MI: University of Michigan Herbarium. 488 p. [11471]
85. Whittaker, R. H. 1956. Vegetation of the Great Smoky Mountains. Ecological Monographs. 26(1): 2-80. [86822]
86. Whittaker, R. H. 1963. Net production of heath balds and forest heaths in the Great Smoky Mountains. Ecology. 44(1): 176-182. [64578]
87. Wu, Z. Y.; Raven, P. H.; Hong, D. Y., eds. 2013. Flora of China, [Online]. Volumes 1-25. Beijing: Science Press; St. Louis, MO: Missouri Botanical Garden Press. In: eFloras. St. Louis, MO: Missouri Botanical Garden; Cambridge, MA: Harvard University Herbaria (Producers). Available: and [72954]
88. Zimmerman, Michael Lee. 2006. The effects of prescribed fire on the herbaceous layer in the southern Appalachian Mountains. Johnson City, TN: East Tennessee State University. 111 p. Thesis. [86868]

FEIS Home Page