Botrychium spp.


Table of Contents


INTRODUCTORY



CITATION:
Fryer, Janet L. 2014. Botrychium spp. 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/plants/fern/botspp/all.html [].

 
Figure 1. Common moonwort. Photo by Abalg, courtesy of Creative Commons. Figure 2. Mountain moonwort. Photo ©2012 Ryan Batten.
Figure 9 and Figure 11 show photos of daisy-leaf and peculiar moonworts, respectively.

SUMMARY:
This section summarizes information on the general ecology and fire ecology of 4 moonwort species: Botrychium lunaria, Botrychium matricariifolium, Botrychium montanum, and Botrychium paradoxum, using the sources available in the scientific literature as of 2014. Scientific literature available on these species was limited as of 2014. Details and documentation of source materials are included in individual sections that follow this summary.

Introductory: The Introductory section discusses the taxonomy of the moonworts discussed in this review, including infrataxa, hybrids, and synonyms.

Distribution and Occurrence: The moonworts discussed in this review are native to North America. All are rare throughout most of or all of their ranges. They occur on a wide variety of sites and in many plant communities, ranging from open, grassy meadows to closed-canopy, old-growth forests.

Botanical and Ecological Characteristics: Moonworts are small, inconspicuous, cryptic species. They have 2 life stages: the gametophyte and sporophyte generations. The entire gametophyte and most of the sporophyte stage is spent below ground; obligatory mycorrhizal associations support moonwort development and growth. Moonworts regenerate from spores, gemmae, and possibly by sprouting. They occur in a wide range of light conditions and successional stages.

Fire Effects and Management: Limited research suggests that moonworts require several years to recover after fire. Mechanisms of postfire regeneration may include sprouting of dormant sporophytes and establishing on burned sites from spores. Occurrence of moonworts in early (≥postfire year 10) to old-growth successional stages suggests moonworts are adapted to a wide variety of fire regimes. Further research is needed on the fire ecology of moonworts.

Management Considerations: Moonworts are rare and have protection status in most states and provinces in which they occur. Herbivores utilize moonworts, but moonworts are probably not important forage due to their small stature and rarity. Their cryptic habit and mostly belowground life cycle make them difficult to survey. Methods for detecting belowground plants are available.

FEIS ABBREVIATION:
BOTSPP
BOTLUN
BOTMAT
BOTMON
BOTPAR

COMMON NAMES:
For the genus Botrychium:
moonwort
grapefern

For Botrychium lunaria:
common moonwort
moonwort grapefern

For Botrychium matricariifolium:
daisy-leaf moonwort
camomile grapefern
matricary grapefern

For Botrychium montanum:
mountain moonwort
western goblin

For Botrychium paradoxum:
peculiar moonwort
paradox moonwort

TAXONOMY:
Moonworts are classified in the genus Botrychium Sw. (Ophioglossaceae) [21,35,63,70]. Within the genus, moonworts are placed in subgenus Botrychium, while grapeferns are placed in the subgenus Sceptridium [17]. This review provides detailed information on 4 moonwort species. Their scientific names are:

Botrychium lunaria (L.) Sw., common moonwort [12,14,21,29,35,63,70,77]
Botrychium matricariifolium (A. Braun ex Dowell) A. Braun ex Koch, daisy-leaf moonwort [22,35,58,63,70]
Botrychium montanum W.H. Wagner, mountain moonwort [21,35,69,70]
Botrychium paradoxum W.H. Wagner, peculiar moonwort [21,35,70]

This review uses common names for plant species. See Appendix B for scientific names of plant species mentioned in this review.

In the broad sense, the Botrychium lunaria complex consists of several closely aligned moonwort species [15,19,67]. These taxa are: common moonwort, giant moonwort, Tunux moonwort, and dainty moonwort. Stensvold [67] and Farrar [19] consider dainty moonwort a variety of common moonwort. Their genetic studies found 3 distinct genotypes of common moonwort (proposed names in parentheses) [67]:

1) a genotype endemic to the western United States and Canada (Botrychium lunaria var. crenulatum (W.H. Wagner) Stensvold, dainty moonwort)
2) a genotype endemic to Greenland, Iceland, and Norway (Botrychium lunaria var. melzeri Stensvold, Melzeri's moonwort)
3) a genotype with a circumpolar distribution (Botrychium lunaria var. lunaria, common moonwort)

Hybrids: Moonworts may hybridize, although resulting plants are usually sterile [27,67]. Common moonwort hybridizes with pointed moonwort [67], spathulate botrychium [67,78], and Mingan moonwort [67]. Mingan moonwort apparently resulted from common moonwort × pale botrychium hybridization [78], and Waterton moonwort from peculiar moonwort × western moonwort hybridization [17,21,78]. Waterton moonwort is known only from western Alberta, in Waterton Lakes National Park [21,39].

SYNONYMS:
For Botrychium lunaria:
Botrychium lunaria (L.) Sw. forma gracile (Schur.) Aschers. & Graebn.
Botrychium lunaria (L.) Sw. forma lunaria [63]
Botrychium lunaria (L.) Sw. var. onondagense (Underw.) House [14]

For Botrychium matricariifolium:
Botrychium lunaria (L.) Swartz var. matricariifolium Döll [21]
Botrychium matricariifolium (A. Braun ex Dowell) A. Braun ex W.D.J. Koch var. matricariifolium [63]

For Botrychium montanum:
none

For Botrychium paradoxum:
none

LIFE FORM:
Fern or fern ally

DISTRIBUTION AND OCCURRENCE

SPECIES: Botrychium spp.
GENERAL DISTRIBUTION:
The 4 moonworts covered in this review are native to North America [21]. All are rare throughout most of or all of their ranges [35]. Their distributions are as follows:

Common moonwort: Common moonwort occurs in North America, South America, Eurasia, Australia, New Zealand, and the Pacific islands [21,30,77]. In North America, it occurs from arctic Alaska [11] east to the southern tip of Greenland and south to California, New Mexico, the Dakotas, and Pennsylvania [12,21,70,77]. It is rare throughout most of its North American distribution, but it has scattered pockets of abundance in the Rocky Mountains [35].

Figure 3. Common moonwort's distribution in the United States and Canada. Map courtesy of USDA, NRCS. 2014. The PLANTS Database. National Plant Data Team, Greensboro, NC. (2014, February 7).

States and provinces [70]:
United States: AK, AZ, CA, CO, ID, ME, MI, MN, MT, ND, NH, NM, NV, NY, OR, PA, SD, UT, VT, WA, WI, WY
Canada: AB, BC, LB, MB, NF, NS, NT, NU, ON, QC, SK, YT


Daisy-leaf moonwort: Daisy-leaf moonwort occurs in eastern North America, Eurasia [21,42,59], Patagonia [22], Australia, and New Zealand [42]. In North America, it occurs from Manitoba east to Labrador and south to the Dakotas, Tennessee, and North Carolina [35,70]. It is the most common moonwort in the Great Lakes and Northeast regions [19], but it is rare on the western edges of its distribution [35].

Figure 4. Daisy-leaf moonwort's distribution in the United States and Canada. Map courtesy of USDA, NRCS. 2014. The PLANTS Database. National Plant Data Team, Greensboro, NC. (2014, February 7).

States and provinces [70]:
United States: CT, DC, DE, IA, IL, IN, KY, MA, MD, ME, MI, MN, NC, ND, NH, NJ, NY, OH, PA, RI, SD, TN, VA, VT, WI, WV
Canada: LB, MB, NB, NF, NS, ON, PE, QC

Mountain moonwort: Mountain moonwort has a scattered distribution in the western United States [21,35,70]. It also occurs in British Columbia [35,70] and Alaska. Except in Washington, it is rare throughout its distribution [35].

Figure 5. Mountain moonwort's distribution in the United States and Canada. Map courtesy of USDA, NRCS. 2014. The PLANTS Database. National Plant Data Team, Greensboro, NC. (2014, February 7).

States and provinces [35,70]:
United States: AK, CA, ID, MT, OR, WA
Canada: BC

Peculiar moonwort: Peculiar moonwort has a scattered distribution in the western United States and southwestern Canada [21,35]. It is rare throughout its distribution [35].

Figure 6. Peculiar moonwort's distribution in the United States and Canada. Map courtesy of USDA, NRCS. 2014. The PLANTS Database. National Plant Data Team, Greensboro, NC. (2014, February 7).

States and provinces [19,70]:
United States: CA, OR, ID, MT, UT, WA
Canada: AB, BC, SK


SITE CHARACTERISTICS AND PLANT COMMUNITIES:
Because moonworts are easily overlooked and rare, little is known of their habitat preferences. They tend to grow in open areas including beaches, subalpine and alpine meadows [62], and wood edges [19,77]. They are uncommon in arid environments [62]. In the United States, most moonwort species grow at high elevations [21]. In the Southern Rocky Mountains, they are most common above 9,000 feet (2,700 m) [62]. Soils supporting moonworts are moist [19] to well-drained and often gravelly or rocky [62]. Moonworts generally favor acidic to neutral soil (pH 4.8-7.2) [65]. However, some species, including common moonwort, tolerate or prefer basic soils [9,19,49]. Moonworts are commonly associated with recent (15-30 years) natural and human-caused disturbances, growing on avalanche chutes, talus slopes, and deglaciated areas; along roadsides, airstrips, and ski runs; and in fields, pastures, and areas of mining activity [67]. Farrar [17] reported that in general, moonworts do not have precise habitat requirements. Associated mycorrhizae (see General Botanical Characteristics) apparently mediate many interactions of moonworts species with their environments [17].

The limited information available on habitats of common, daisy-leaf, mountain, and peculiar moonworts (as of 2014) is summarized below. See the Fire Regime Table for a list of plant communities in which these moonworts may occur and information on the fire regimes associated with those communities. See Appendix B for scientific names of plant species mentioned in this review.

Common moonwort: Common moonwort is cosmopolitan in habitat [19]. It grows on shores, cliff ledges, scree, and open gravelly slopes; in open fields, meadows, and woodlands [19,49]; and occasionally in forests. It tolerates and often grows on dry soils [21] such as sand dunes, but it is most common on moist, well-drained woodland and meadow soils [19]. It prefers neutral [49] to basic [12,49,63] soils.

In Alaska, common moonwort occurs on grassy slopes [29] and shrubfields. In arctic northeastern Alaska, a common moonwort population grew on a south-facing slope in the organic layer of a sandy loam derived from glaciofluvial materials. The plant community was a shrub-grassland dominated by grayleaf willow, shrubby cinquefoil, northern rough fescue, and arctic brome. Common moonworts grew in the middle of the community, appearing as scattered individuals or in pairs. They were growing within grass clumps beneath grayleaf willows [11]. Northeast of Nome, common moonwort was abundant (40% frequency) in an arctic dwarf birch-bog birch dwarf shrubland [25]. In southeastern Alaska, it grew on the face of Yakutat Glacier [67]. Near Palmer in south-central Alaska, it grew in slender wheatgrass, bluejoint reedgrass, and northern rough fescue-mountain cranberry grassland communities [24].

Near the Alaskan Highway in Yukon, Corell [13] observed "numerous" common moonwort plants growing in a limestone sink above timberline. Krummholz subalpine fir, Sitka clubmoss, fir clubmoss, and alpine clubmoss grew on the same site [13].

Common moonwort grows mostly in open communities in the conterminous United States. On the Pacific coast, it grows on grassy slopes and in fields, moist meadows, heathlands, and in open forests. It occurs from low to high elevations [54]. In the Intermountain West, it grows on wet to moist sites at middle to high elevations [14]. In Idaho, it grows in mountain meadows and western redcedar-western hemlock forests. A survey on the Kootenai National Forest, Idaho, found common moonwort grew exclusively in old-growth western redcedar and western hemlock forests [72]. In Glacier National Park, Montana, it grows in mountain meadows, mossy banks, and bogs. In the 1920s, it was noted as "plentiful" on the moraine of Grinnell Glacier [66]. It is reported in grassy meadows and on wood edges in Utah, from 7,700 to 11,500 feet (2,350-3,500 m) elevation [77]. In Colorado, it grows in mountainous regions at elevations up to 12,000 feet (4,000 m) [26]. In the Rocky Mountains of central Colorado, common moonwort was noted in limber pine-Rocky Mountain Douglas-fir-interior ponderosa pine forests [60]. In the San Francisco Peaks of north-central Arizona, it grew in alpine meadows dominated by Ross' avens and sedges. These communities occur at and above 10,100 feet (3,500 m) [52]. Near timberline at 11,600 feet (3,550 m), common moonwort was found in a krummholz Great Basin bristlecone pine-Engelmann spruce/Ross' avens community [56]. In New England, common moonwort occurs on open slopes [64] and in open woodlands [16].

Daisy-leaf moonwort: This species is cosmopolitan in habitat [19]. It grows on roadsides and in meadows, woodlands, forest edges, open to closed forests, and rich swamps [12,19,41,63]. It tolerates moist to dry soils that range from acidic to neutral pH [19]. Across Canada and the United States, it occurs from sea level to 4,000 feet (1,200 m) [21]. In the Intermountain West, it grows in woodlands at low to middle elevations [28]. It occurs above 4,000 feet (1,200 m) in the Black Hills of South Dakota [19]. It is reported in moist woodlands in the Great Plains [23]. In Minnesota, daisy-leaf moonwort gametophytes, collected in the field and identified in the laboratory, were growing with least moonwort in a white oak-bur oak-northern red oak forest [10]. In Michigan, daisy-leaf moonwort grows in conifer and hardwood-conifer forests. In Gogebic and Ontonagon counties, it was found on mesic to dry sites [44]. On the Keweenaw Peninsula, Michigan, daisy-leaf moonwort was noted in a 100+-year-old white spruce-balsam fir forest with eastern white pine and quaking aspen. The site was mesic, and the canopy was opening due to canopy-gap succession [45]. Daisy-leaf moonwort is reported in grassy meadows in Illinois [58]. In Nova Scotia, it grows in hardwood communities on leaf mold or rich alluvial soils [58]. In the Northeast, it occurs in thickets and woodlands in "subacid" soil [22]. In New Jersey, daisy-leaf moonwort occurred in a pin oak woodland and on the border of a pitch pine barren. The pin oak community soil was moist, rich clay, while soil on the pine barren border was dry sand [41]. In Maryland, a daisy-leaf moonwort population was found in "damp woods" along a tributary of the Patapsco River [75]. A population in Montgomery County, Maryland, grew beneath a dense eastern poison-ivy thicket in a yellow-poplar/flowering dogwood stand [20]. In Great Smoky Mountains National Park, North Carolina, daisy-leaf moonwort grew in mesophytic cove forests of yellow-poplar, red maple, and eastern hemlock. A daisy-leaf moonwort population was noted on north-facing slopes of 5% to 50%, at 2,250 feet (686 m) elevation. Daisy-leaf moonworts were most common in areas without leaf litter and duff [57].

Mountain moonwort: Mountain moonwort grows in dark, moist habitats. This, and its gray-green color, earns it the alternate common name "western goblin" [19]. It grows in conifer forests, usually in or near riparian zones [19,21,49,74], and in moist meadows, seeps, and fens [19,74]. Mountain moonwort has been found on calcareous soils [19]. Across its range, it occurs from 3,000 to 6,500 feet (1,000-2,000 m) elevation [21]. In Washington, it prefers old-growth western redcedar riparian forests and is uncommon in other plant communities. Western redcedar forests with mountain moonwort occupy upper terraces near small to moderate-sized streams. Soils are moist, with high concentrations of organic matter and minerals [19]. Surveyors on the Kootenai National Forest found mountain moonwort exclusively in mature second-growth and old-growth western redcedar and western hemlock forests [72]. In California, mountain moonwort occurred in conifer forests at 5,000 to 6,000 feet (1,500-1,800 m), growing mostly in the southern Cascade Range [69]. Specimens were collected near a stream in Butte and Tehama counties, in a mixed-conifer forest of incense-cedar, white fir, sugar pine, and Pacific ponderosa pine [19,76]. Farrar [19] suggested that litter from the Cupressaceae family, which includes western redcedar and incense-cedar, favors mountain moonwort growth. Additionally, mycorrhizae associated with Cupressaceae are also associated with Botrychium [9,19].

Figure 7. Mountain Moonwort growing in incense-cedar debris and litter in an ephemeral drainage on the El Dorado National Forest, California. Photo ©2012 Belinda Lo, courtesy of Creative Commons.

Peculiar moonwort: This species grows in snowfields, pastures [21], mountain meadows [19,49], woodlands, and conifer forests [49,74], including western redcedar forests [21]. It occurs from 5,000 to 9,000 feet (1,500-3,000 m) elevation across it range [21,74]. Three peculiar moonwort populations have been found in Yosemite National Park, California, all in soils derived from metasedimentary parent materials [19]. On the Kootenai National Forest, peculiar moonwort grew in western redcedar and western hemlock forests. Soils were volcanic loess overlying dense glacial till, but peculiar moonworts were growing in deep litter and humus, not the underlying soil [72]. In Montana, peculiar moonwort grows in northern rough fescue-Idaho fescue meadows [39], dense fireweed stands, and near Rocky Mountain lodgepole pine, Engelmann spruce, and alpine larch trees [74]. Elevational ranges in Montana are from 3,000 to 8,400 feet (900-2,500 m) [39]. In Waterton Lakes National Park, Alberta, peculiar moonwort grew in streamside mountain meadows dominated by northern rough fescue and Idaho fescue [39]. In Willmore Wilderness Park of west-central Alberta, it occurred in subalpine fir-Engelmann spruce/kinnikinnick/boreal wildrye forests. Sites with peculiar moonwort ranged from 4,700 to 7,249 feet (1,433-2,210 m) elevation; soils were mostly calcareous [9].

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Botrychium spp.
GENERAL BOTANICAL CHARACTERISTICS:
Botanical descriptions:
Descriptions in this review cover characteristics that may be relevant to fire ecology and are not meant for identification. Moonworts, general: Moonworts are perennial herbs. They are small, inconspicuous [49,53], and structurally simple [67]. Wagner [75] described moonworts as requiring "persistent searching (meaning on hands and knees)" to find. Once found, moonwort species are difficult to identify. They are considered cryptic [51,78] because their distinguishing morphological characters are few, variable, and often subtle; they are reproductively isolated; and species within the genus have often been misplaced taxonomically [51,67]. Aboveground morphology of individuals and populations may change annually with environmental conditions [78]. "Reliable field determination of moonworts depends on the careful use of technical keys and comparison with silhouette outlines of verified specimens. It is complicated because there is often a high degree of morphological variability between individuals in a population and between populations of the same species, several species may grow together at the same site, and the few diagnostic characters may not be apparent in small plants" [49]. Genetic analysis may be needed to distinguish between moonwort species [78].

Moonworts have 2 distinct generations: the gametophyte (n) and sporophyte (2n) [2,48]. The gametophyte spends its life entirely below ground. The gametophyte generation is longest and is little studied. The sporophyte develops and spends most of its life below ground [48]. Its frond emerges above ground at sexual maturity, and the frond is featured in identification keys. Both generations require mycorrhizae for development [34,67]; the gametophyte is entirely dependent on mycorrhizae for water and nutrients [2,34]. Persistence of associated mycorrhizae is likely the most important factor in persistence of moonwort populations (Johnson-Groh 1999 cited in [2]). See Regeneration Processes for further discussion of the moonwort life cycle.

Figure 8. A cutleaf grapefern gametophyte. Ophioglossaceae gametophytes are barely visible without magnification and are rarely microphotographed. Photo © Dr. Steven J. Baskauf, Bioimages.

Moonwort gametophytes grow entirely below ground. They are spherical to oblong, ranging from 0.1 [61] to 3.0 mm [10] in diameter. They lack chlorophyll; young gametophytes are nearly colorless but turn brown with age [10]. Rhizoids cover most of their outer surface [67] (pictured in Figure 8 above).

The mature sporophyte consists of the aboveground frond and the belowground caudex, rhizomes, and roots. The frond stalk is usually divided into 2 segments. The sterile segment (the trophophore) supports leaflets or pinnae, while the upper, fertile segment (the sporophore) supports the sporangia (spore cases) [21,67,79]. The common name "moonwort" refers to the half-moon shape of the leaflets [54]. Moonworts are distinguished by the morphology of their leaflets (for example, simple, pinnate, or dissected) [28,63]. The sporangia are arranged in grapelike clusters [21,28,67]. Moonwort spores are just visible without magnification [62]. The sporophyte has a belowground caudex; fleshy, adventitious roots attach to the caudex [31,67,77]. Moonworts are rhizomatous [67]; rhizomes spread horizontally for 1 to 8 inches (3-20 cm) [34]. Roots are few in number (5-30) and lack root hairs ([34], Bower 1926 cited in [67]).

Moonwort population sizes vary greatly, from fewer than 10 to thousands of sporophyte plants (review by [2]). Population size may appear to fluctuate without discernable causes (U.S. Forest Service 1999 in [46]), although population size cannot be determined accurately without intensive sampling (see Other Management Considerations). Moonworts tend to grow in clumps, often within 0.4 inch (1 cm) of one another [78]. Peculiar moonwort, western moonwort, and their hybrid, Waterton moonwort, grow together in Waterton Lakes National Park [39]. Because moonworts are easily overlooked, they may be more common than surveys to date (2014) indicate [49].

Moonworts are short-lived [39]; they are thought to live for 5 to 10 years [62].

Moonworts manufacture trehalose sugar; that ability is also noted in clubmosses but is unusual among higher plants. Trehalose enables plants to withstand desiccation and rehydrate quickly [80].

Mycorrhizae infect moonwort stems, roots, and rhizoids (Campbell 1908, Bower 1926, Rayner 1927, cited in [67]). These fungal associates are largely unidentified, and some species may be undescribed. Vesicular-arbuscular [72,79] and ectomycorrhorizal [21,49] fungi are associated with moonworts. Some mycorrhizae associated with moonworts have been identified as Glomus spp. ([67,79]).

Common moonwort: Common moonwort's fleshy frond arises from a caudex [31] about 5 mm thick [49]. The frond is long (up to 10 inches (25 cm)) and somewhat leathery [12,54]. Frond leaflets are highly variable in form. Environmental conditions during elongation, such as early-season freezing temperatures, affect the morphology of fully extended leaflets [29]. Electron micrographs showed endophytic fungi infecting common moonwort gametophytes but not sporophytes. The gametophytes were 0.1 to 0.3 mm in diameter, with a few rhizoids [61].

Daisy-leaf moonwort: Daisy-leaf moonwort's frond is relatively long (≥4 inches (10 cm)) and membranous to fleshy [12,21]. The sporophyte has short rhizomes and thick roots. In France, daisy-leaf moonwort fronds emerged when plants averaged about 10 years old. Sporophytes lived another 2 (rarely 3) years after reaching sexual maturity. Over 4 years, an average of 50% of sporophytes were not relocated in surveys (and presumed dead) [48].

Figure 9. Daisy-leaf moonwort frond, showing sporangia-bearing, fertile (upper) and sterile (lower) segments. Photo by Radim Paulič, courtesy of Creative Commons.

Mountain moonwort: Mountain moonwort's frond is somewhat succulent. It may reach 5 inches (12 cm) tall, although it is often shorter [49].

Peculiar moonwort: Peculiar moonwort's frond has 2 spore-bearing segments; unlike other moonworts, this species lacks a sterile frond segment. Mature sporophytes usually range from 3 to 5 inches (7-15 cm) tall in open sun but are shorter in shade [21,49]. Vesicular-arbuscular mycorrhizae [79] and/or ectomycorrhizae [21,49] infect peculiar moonwort's roots and rhizoids. In Waterton Lakes National Park, peculiar moonwort sporophytes lived 6 years or less [39].

Raunkiaer [55] life form:
Chamaephyte
Geophyte

SEASONAL DEVELOPMENT:
Moonworts are ephemeral. The single frond of sexually mature sporophytes emerges in spring or early summer [67], and spores disperse in summer. The frond desiccates by midsummer to late fall [67].

Mature moonwort sporophytes may remain dormant for a year or more before re-emerging [31,39]. Drought and lack of nutrients apparently induce dormancy in moonwort species [76,78], although other as yet unidentified factors may also induce dormancy. In Waterton Lakes National Park, 78% of peculiar moonworts sampled were dormant for 1 year, 19% for 2 years, and 3% for >2 years. Approximately 280 peculiar moonwort sporophytes were located in this 4-year study [39]. Annual dormancy rate averaged 13% for a daisy-leaf moonwort population in France. Daisy-leaf moonwort fronds desiccated in spring, prior to spore dispersal, in drought years [47].

Table 1. Phenology of common, daisy-leaf, mountain, and peculiar moonworts
Species Region Event Period
Common moonwort across range fronds emerge
spring
fronds desiccate late summer [21]
Minnesota fronds emerge May
fronds desiccate mid-August [46]
Northeast spores mature June-August [42]
New England spores mature July-August [64]
Daisy-leaf moonwort across range fronds emerge
spring
fronds desiccate mid-late summer [21]
Great Plains spores mature June-August [23]
Illinois spores mature June-July [58]
Northeast spores mature June-August [22,42]
Canada spores mature June-July [12]
France fronds emerge
late April-early May
spores mature May
spores disperse early-mid-June
fronds senesce mid- late June [47]
Mountain moonwort across range fronds emerge late spring-summer [21]
Canada spores mature June-August [12]
Peculiar moonwort across range spores mature June-August [21]
across range

fronds emerge

spring

fronds desiccate mid- to late summer [49]

REGENERATION PROCESSES:
The life cycle of moonworts is similar to that of ferns in the class Filicopsida, but it is only partially understood for moonworts [62]. Spores (meiospores) produced by the sporophyte germinate below ground. The resulting gametophyte forms a caudex and grows for several years. When mature, the gametophyte produces female archegonia and antheridia with eggs and sperm, respectively [17]. Fertilization occurs below ground [17,67,79]. Sperm require wet soil to swim to eggs [2,39]; free-swimming sperm may fertilize eggs from either their parent or adjacent gametophytes [39]. After fertilization, the resulting sporophyte remains attached to the gametophyte. After several years of development, the sporophyte frond stalk emerges above ground, and the frond leaf expands [39,79]. Length of time required for gametophytes to develop from spores, and for sporophytes to develop and emerge above ground, is little studied. Little is known of the gametophyte portion of the life cycle in general [39].

Figure 10. Moonwort life cycle. Only the mature sporophyte appears above ground. Diagram by Janet Fryer, U.S. Forest Service.

Breeding system: Moonworts are monoecious [2].

Overall genetic diversity is low in moonworts compared to other ferns and fern allies. Soil restricts movement of sperm between moonwort gametophytes, so inbreeding is prevalent. There is a high level of intragametophytic selfing ([27,67], review by [2]). In the intragametophytic selfing breeding system of moonworts, sperm released by belowground antheridia swim less than 1 mm to eggs in archegonia on the same gametophyte [17,67]. Except in rare cases, the resulting sporophytes are genetically nearly identical to their parents [67]. Farrar [17] reported that <1% of moonwort plants tested showed genetic evidence of outcrossing. However, he pointed out that although moonworts have likely had low genetic diversity for thousands to millions of years, their low genetic diversity is apparently not a barrier to population viability [17]. Long-distance spore dispersal and clumping of belowground gametophytes [78] allows for limited genetic exchange.

Metapopulation dynamics are likely important in maintaining moonwort populations, but they are poorly understood for Botrychium [2].

Despite barriers to cross-fertilization, hybridization and polyploidy occur in moonworts, sometimes resulting in new species. For example, daisy-leaf moonwort is a tetraploid that probably resulted from hybridization between diploid lanceleaf moonwort and diploid pale botrychium [27,78].

Spore production and dispersal: Spore output of moonworts is large [2]. A single sporangium contains thousands of spores [67], and a single sporophore may bear >100 sporangia (see Figure 1).

Dispersal occurs during the sporophyte stage [39,79]. Wind and air currents disperse most moonwort spores [62], although water and animals also disperse spores [2,67]. Spores stick to animal hides or feathers and travel through the digestive tracts of herbivores. Viable spores have been collected from feces of deer and small mammals (review by [2]). Wind and animals may disperse spores long distances [34,67]. Birds may have dispersed common moonwort to Australia, New Zealand, and the Pacific islands [15]. However, most spores fall beneath or near the parent plant [67]. Researchers estimate that moonwort spores usually disperse <10 feet (3 m). Rain, frost heaving [62], and erosion [67] work spores into the ground.

Structure banking: Moonwort spores are stored in the soil [49]. Soil-stored moonwort spores may remain viable for several years; Schneider [62] suggests viability of 5 to 10 years. Johnson-Groh and others [34] suggested that "Botrychium spores likely remain viable for long periods of time". For mountain moonwort, they estimated a density of 6,000 spores/m² for soil samples collected on the Colville National Forest, Washington [34].

With their spores, gametophytes, belowground juvenile sporophytes, and gemmae, moonworts have more reproductive structures than most plant species. Combined, these reproductive resources have been called a "belowground structure bank". For a mountain moonwort population on the Colville National Forest, researchers found structure banks averaged 1.2 aboveground sporophytes/m², 0 belowground sporophytes/m², 738 gametophytes/m², and 0 gemmae/m² [34].

Germination: Moonwort spores probably remain dormant in the soil until soil moisture is sufficient and fungal associates are present [34]. Anecdotal information suggests that moonworts may require open sites with mineral soil for germination and establishment [2]. In Great Smoky National Park, daisy-leaf moonwort was observed only where leaf litter was sparse to lacking [57]. Moonwort spores germinate below ground [73]; they require complete darkness to germinate. Mycorrhizae must infect the gametophyte after only a few cell divisions for development to continue [67].

Figure 11. Peculiar moonwort sporophyte emerging from a gravelly roadside in Alberta. Photo © 2011 Ryan Batten.

Establishment and plant growth: Both gametophytes [2] and sporophytes [67] grow slowly. Sporophytes may take several years for their fronds to emerge above ground [34,67]. Many plants die before their life cycle is complete. In a study of 5 moonwort species including mountain moonwort, overall annual mortality rate of belowground embryonic, belowground juvenile, and emergent adult sporophytes averaged 73% [34].

In Waterton Lakes National Park, peculiar moonwort's ratio of new plant recruitment:surviving plants was significantly higher than ratios of western moonwort or Waterton moonwort (P<0.05). Across 3 years, about 48% of grid-located peculiar moonworts were new plants. However, peculiar moonwort's mortality rate (40%) tended to be higher than mortality rates of the other 2 moonwort species (P=0.06) [39].

Vegetative regeneration: Moonworts may reproduce asexually by producing gemmae [18,49]. These spherical propagules grow on rhizomes [34] or aboveground stems [18] and range from 0.5 to 1.0 mm in diameter. They develop below ground after abscission from the stem. Gemmae mature into sporophytes; their development is similar to that of sexually produced embryos. Limited microscopic examinations (n=12 stems/species) found no gemmae on either common moonworts from Colorado or daisy-leaf moonworts from Michigan [18], although NatureServe [49] reported that common moonworts sometimes produce gemmae. Gemmae have been found on mountain moonworts collected in Idaho [72]. Further examinations are needed to determine the importance of gemmae to moonwort regeneration in general and the moonwort species discussed in this review in particular.

Moonworts have caudices and rhizomes [31,67], and they likely sprout from these organs after top-kill. Further research is needed on the ability of moonworts to sprout after disturbances such as fire or grazing.

SUCCESSIONAL STATUS:
Moonworts are common in full sunlight [2,49,62] but tolerate some shade. They grow on open meadows, slopes, and banks [26,49,62]. Limited observations suggest that in open areas, moonworts may grow in shaded microsites. For example, moonworts have been found growing beneath shrubs [11] and near tall bunchgrasses that provide shade [39].

Moonworts are associated with light to moderate disturbances [39] but also occur in late successional stages [19,72]. A review stated that "moderate disturbance may be a critical part of the autecology of Botrychium species" [2]. However, moonworts are generally found on sites disturbed ≥10 years prior (Johnson-Groh 2003 personal communication in [2]).

Common moonwort: Limited studies and observations suggest common moonwort sporophytes are absent or uncommon in the initial stage of postdisturbance succession. In the Intermountain West, common moonwort is more common in partial shade than in either open meadows or closed forests [14]. In arctic Alaska, common moonwort was rooted in deep organic material over glaciofluvial material. The depth of the organic layer in which common moonwort grew suggested that it did not establish in early succession, although the exact depth was not provided. Common moonwort was not found on young glaciofluvial materials [11]. Researchers in Norway suggested that common moonwort is a late-successional species. It was absent from subalpine grasslands ploughed 3 to 5 years prior; it was found exclusively in unploughed subalpine meadows [6].

Daisy-leaf moonwort: Daisy-leaf moonwort occurs in second-growth forests across its range [21]. It grows on open sites as well as in closed-canopy forests. It grew in a roadside barrow pit in Glacier National Park [38], suggesting that it tolerates disturbed sites. In the southern Appalachian Mountains it is restricted to closed, mature hardwood forests [19]. It was noted on shaded trails in the Carolinas [44].

Mountain moonwort: Mountain moonwort grows in shady woodlands and closed forests [21,69] but also in open meadows [19]. In Washington, it is most common in old-growth western redcedar forests but has also been found in mountain meadows on Mt Baker [19]. On the Kootenai National Forest, Vanderhorst [72] found mountain moonwort in second-growth western redcedar forests ranging from 40 to 210 years old and in old-growth western redcedar forests that were probably >1,000 years old. It was often the only groundlayer species growing in the deeply-shaded litter of old-growth western redcedar stands. Vanderhorst did not find it in clearcuts [72].

Peculiar moonwort: Successional preferences of peculiar moonwort had rarely been studied as of this writing (2014). A peculiar moonwort population was found on a south-facing roadside in Glacier National Park [8], suggesting ability to grow on open, disturbed sites. In Waterton Lakes National Park, peculiar moonwort occurred in meadow portions of Rocky Mountain lodgepole pine forest-mountain meadow mosaics. Rocky Mountain lodgepole pine forests are subject to wildfire and pine beetle attacks, which slows Rocky Mountain lodgepole pine invasion into meadows. Sites with peculiar moonwort had a dense litter layer and bare soil was uncommon, suggesting infrequent disturbances [39].

FIRE EFFECTS AND MANAGEMENT

SPECIES: Botrychium spp.
FIRE EFFECTS:
Immediate fire effect on plant: Immediate effects of fire on moonworts were undocumented as of 2014. Since moonwort gametophytes are below ground, surface or crown fires may not damage them. Gametophytes residing in litter and duff are likely vulnerable to fire, especially ground fire.

Fire during the growing season likely top-kills emergent moonwort sporophytes. Since their caudices and rhizomes are below ground, fire is probably not lethal to sporophytes growing in mineral soil. Fire may not damage dormant moonwort sporophytes. Most moonwort sporophytes are usually dormant during drought [39], so they are likely dormant in extreme fire years.

Postfire regeneration strategy* (adapted from [68]):
Rhizomatous herb, rhizome in soil
Caudex, growing points in soil
Geophyte, growing points in soil
Ground residual colonizer (on site, initial community)
Initial off-site colonizer (off site, initial community)
Secondary colonizer (on- or off-site spore sources)

*These postfire regeneration strategies are possible based on moonwort morphology and methods of regeneration but have not been confirmed in field studies.

Fire adaptations and plant response to fire: Fire adaptations: Although moonwort adaptations to fire were not well studied as of 2014, moonworts have several adaptations that may help them survive fire and establish new plants. Since the entire gametophyte stage and most of the sporophyte stage are spent below ground, moonworts growing in mineral soil are probably protected from fire damage. Sporophytes have belowground caudices and are rhizomatous, so plants in mineral soil probably sprout after fire [34,50]. Presence of a soil spore bank, and spore dispersal aided by wind, water, and animals, likely helps ensure new postfire regeneration. Sporophyte and spore dormancy may delay postfire sprouting until environmental conditions are favorable for growth (see Regeneration Processes). However, these are conjectural adaptations. Studies are needed on the fire ecology of moonworts.

Bare mineral soil favors postfire establishment of clubmoss spores [2]. Because clubmosses are fern allies that, similar to moonworts, have a life cycle that includes belowground gametophyte and aboveground sporophyte stages, Anderson [2] suggested moonworts might also establish from spores after fire prepares a mineral sporebed.

Plant response to fire: To date (2014), only 2 fire studies [9,57] and anecdotal evidence [45] documented postfire responses of the 4 moonworts covered in this review. Moonwort sporophytes growing in mineral soil likely sprout from their caudices and rhizomes after fire. Gametophytes likely establish from spores stored in the soil and dispersed from off site. Postfire reproduction from gemmae may also occur (see Regeneration Processes). A study in Great Smoky Mountain National Park found dormant-season fire had little effect on moonworts [33]. On the Kootenai National Forest, mountain moonworts sprouted after fronds were completely grazed [72], suggesting ability to sprout after top-kill by fire.

Since both gametophytes and sporophytes grow slowly and are always (gametophytes) or mostly (sporophytes) below ground [2], moonworts may not be found in early postfire surveys. Johnson-Groh and others [34] conjectured that after fire, grazing, or logging, "aboveground populations are fairly resilient and rebound following perturbations, although recovery may take several years". However, Anderson [2] speculated that if fire reduces population density, moonwort populations may require ≥80 years to return to prefire levels.

Moonworts occur in fire-adapted ecosystems. In Minnesota, common moonwort grows on sites with a history of fire and logging [46]. Peculiar moonwort is associated with Rocky Mountain lodgepole pine in Montana [74]; Rocky Mountain lodgepole pine forests have a history of mixed-severity fire about every 25 to 75 years and of stand-replacement fire about every 100 to 330 years [4]. See the Fire Regimes section and the Fire Regime Table for further information on the fire regimes of plant communities in which common, daisy-leaf, mountain, and peculiar moonwort may occur.

Favorable conditions for postfire moonwort regeneration include presence of appropriate mycorrhizae, appropriate soil moisture and chemistry, and possibly, mineral soil [17]. Which fungal species and what concentrations of soil water and nutrients are required were unknown as of 2014. Fire severity and effects to litter, duff, and fungal communities are likely important factors in postfire responses of moonworts [72]. Although associated mycorrhizae are critical to survival of moonwort populations, little is known of either the biology of these mycorrhizae or of moonwort-mycorrhizal interactions. The effects of fire on associated mycorrhizae, and mycorrhizal responses to fire, may be critical to postfire survivorship and growth of moonworts [2]. Studies are need on the basic biology and fire ecology of moonwort species and their associated mycorrhizae.

In a pilot study in Great Smoky Mountains National Park, dormant-season (early spring) prescribed fire or raking increased daisy-leaf moonwort density over prefire densities. The Park had 2 known daisy-leaf moonwort populations, 1 with about 42 emerged sporophytes and other with about 114, depending on the year. The populations grew in second-growth yellow-poplar-red maple-eastern hemlock forests, and disturbed leaf litter appeared to favor the daisy-leaf moonwort populations. The larger population was selected for study; it was on a north-facing slope of 5% to 50% at 2,250 feet (686 m) elevation. Daisy-leaf moonwort sporophytes in the larger population were mapped along 2 transects the year prior to treatments (1997). Three 7- × 10-m² plots (1 prescribed fire,1 raked, and 1 control) were established in 1998; treatments were enacted in March 1998. In posttreatment year 1 (1999), daisy-leaf moonwort density increased 35% on the prescribed fire plot and 33% on the raked plot compared to prefire densities (see Figure 11). Total increase on all plots was from 23 emerged daisy-leaf moonworts sporophytes in 1997 to 144 in 1999. The untreated control plot showed no increase during that time. The author concluded that reducing leaf litter favors daisy-leaf moonwort, and that prescribed burning or raking in spring—while daisy-leaf moonwort is still dormant—are effective management tools for maintaining daisy-leaf moonwort populations [57]. Further published studies are needed to confirm trends suggested by this pilot study.

Figure 12. Daisy-leaf moonwort numbers before and after prescribed fire or raking in Great Smoky Mountains National Park [57]
Number of daisy-leaved moonworts before the 1998 spring fire.
Number of daisy-leaved moonworts after the 1998 spring fire.
Number of daisy-leaved moonworts before 1998 spring raking.
Number of daisy-leaved moonworts after 1998 spring raking.

On the Keweenam Peninsula, Michigan, daisy-leaf moonwort was noted in a 100+-year-old white spruce-balsam fir forest that "showed evidence" of previous fire [45].

Fire apparently had little effect on a peculiar moonwort population in west-central Alberta. Ten years after a wildfire in Willmore Wilderness Park, peculiar moonwort occurred on both burned and unburned plots. The difference in cover was not significant. The plant community was a subalpine fir-Engelmann spruce/kinnikinnick/boreal wildrye forest [9].

Postfire responses of other moonwort species may help understand postfire responses of the 4 moonwort species covered in this review. In Iowa and Minnesota, prescribed spring fires in prairie and maple-beech forests had little effect on Iowa moonwort and Frenchman's Bluff moonwort populations [33]. For most study years, there was no significant difference in either the number or size of Frenchman's Bluff moonwort sporophytes on burned vs. unburned plots in Minnesota. However, the population was slow to recover after fires during or soon following a drought year. The researchers suggested that dormant-season fires do not harm moonwort species [32,33] but the combination of fire and drought may kill "a substantial number" of moonworts (abstract [32]).

Limited postfire surveys in forests have found moonworts in unburned fire refugia or in burns that are several decades old. On the Kootenai National Forest, Mingan moonwort plants were found in old-growth western redcedar stands that the wildfires of 1910 missed. They were also found in a second-growth western redcedar stand, next to a fire-scarred western redcedar stump [72]. Anderson [2] stated that for leathery grapefern, "there is no evidence that it has an affinity for recently burned areas". Five years after the 1988 wildfires in Yellowstone National Park, Wyoming, he found leathery grapeferns in a fire-refuge site along Obsidian Creek but not on adjacent burned sites. The leathery grapeferns were growing beneath mature Engelmann spruces [2].

Pre- and postfire surveys in South Dakota found a leathery grapefern population that might have been declining or going locally extinct, but fire was probably not the cause. On the Black Hills National Forest, a leathery grapefern population was located in Bucher Gulch in 1994, but surveyors found no leathery grapefern plants in Butcher Gulch in annual surveys from 1995 through 2001 (Crook 2003 personal communication in [2]). The site was burned in 2002 by a low-severity fire that left a mosaic of unburned patches. A 2003 survey failed to locate any leathery grapefern plants [2]. Without previous annual surveys, wildfire might have been implicated in this apparent population decline. Since no belowground surveys were conducted (see Other Management Considerations), true population numbers could not be estimated.

FUELS AND FIRE REGIMES:
Fuels: Moonwort species are too small, ephemeral, and rare to have much impact on fuel loads.

Fire regimes: Moonworts are apparently adapted to a wide range of successional stages and occur in plant communities with a wide range of fire regimes, from short-interval surface to very long-interval, stand-replacement fires. In the Northern Rocky Mountains, for example, common, mountain, and peculiar moonwort grow in mountain grasslands. These grasslands historically experienced fire about every 25 to 100 years, with intervals on the short end most common [3]. Common moonwort grows in Rocky Mountain Douglas-fir forests, which historically experienced both surface and mixed-severity fires. Fire-return intervals for these forests ranged from about 25 to 100 years [4,5]. On the Kootenai National Forest, mountain moonwort grows in western redcedar and western hemlock forests that range from 40 to >1,000 years old [72]; this suggests adaptation to a wide range of postfire successional stages. Common, mountain, and peculiar moonwort occur in fir-mountain hemlock forests on the Pacific coast. Historically, these forests had mostly stand-replacement fires, with fire-return intervals ranging from about 125 to 600 years [1].

See the Fire Regime Table for further information on fire regimes of vegetation communities in which common, daisy-leaf, mountain, and peculiar moonwort may occur.

FIRE MANAGEMENT CONSIDERATIONS:
As of 2014, little research had been conducted on the response of moonworts to fire. Their occurrence in recently disturbed sites, in midsuccession, and in old growth, as well as open meadows (see Successional Status), suggests moonworts are adapted to a wide range of light conditions and postfire successional stages. Multiple structures for regeneration—most below ground—suggest that moonworts can regenerate successfully after fire. Limited studies show moonwort species can recover from fire and that dormant-season fires have little effect on moonwort populations. Further studies are needed on the fire ecology of moonworts.

MANAGEMENT CONSIDERATIONS

SPECIES: Botrychium spp.
FEDERAL LEGAL STATUS:
Neither common, daisy-leaf, mountain, nor peculiar moonwort have federal legal protection [71].

OTHER STATUS:
Common, daisy-leaf, mountain, and peculiar moonwort are all rare throughout most of or all of their ranges (see General Distribution). All have protection status in various states and provinces. Information on state- and province-level protection status of plants in the United States and Canada is available at NatureServe.

IMPORTANCE TO WILDLIFE AND LIVESTOCK:
Because moonworts are small and ephemeral, herbivores may graze them only incidentally. However, herbivore use of palatable moonwort species may become concentrated in seasons favorable to moonwort emergence and growth. As of 2014, little information was available regarding the relative palatability of moonwort species, and no information was available on their nutritional value.

On the Kootenai National Forest, mountain moonworts showed "considerable levels" of herbivory. Mountain moonworts sprouted after fronds were completely grazed. Cooccurring peculiar moonworts were apparently not grazed [72].

Rocky Mountain bighorn sheep grazed peculiar moonwort in Willmore Wilderness Park. On a scale from 1 (primary forage) to 5 (not grazed), peculiar moonwort utilization was ranked at 2 [9]. In Waterton Lakes National Park, rodents did not graze peculiar moonwort, although they apparently grazed cooccurring Waterton moonwort and western moonwort [39].

Cover value: Moonworts are too small to provide cover for vertebrates.

VALUE FOR REHABILITATION OF DISTURBED SITES:
No information is available on this topic.

OTHER USES:
Moonworts are difficult to cultivate. They cannot survive and grow unless their associated mycorrhizae are present in the soil [73]. Hitchcock and others [28] advised that moonworts "should be left alone by all fortunate enough to discover them".

OTHER MANAGEMENT CONSIDERATIONS:
Surveying and monitoring moonwort populations are difficult because most of the moonwort life cycle is spent below ground, mature plants are often dormant, and moonworts are small and inconspicuous even after they emerge. Accurate estimates of population sizes and trends are unlikely without mapping or marking individual sporophytes, so intensive sampling is probably inevitable when such data are needed [39,40]. Mason and Farrar [43] provide methods to census dormant sporophytes and gametophytes.

APPENDIXES

SPECIES: Botrychium spp.

APPENDIX A: FIRE REGIME TABLE
The following table provides fire regime information that may be relevant to common, daisy-leaf, mountain, and peculiar moonwort habitats. Follow the links in the table to documents that provide more detailed information on these fire regimes.

Common moonwort:

Fire regime information on vegetation communities in which common moonwort may occur. This information is taken from the LANDFIRE Rapid Assessment Vegetation Models [37], 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.
Pacific Northwest California Southwest Great Basin
Northern and Central Rockies Northern Great Plains Great Lakes Northeast
Pacific Northwest
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)
Minimum interval
(years)
Maximum interval
(years)
Pacific Northwest Grassland
Alpine and subalpine meadows and grasslands Replacement 68% 350 200 500
Mixed 32% 750 500 >1,000
Idaho fescue grasslands Replacement 76% 40    
Mixed 24% 125    
Pacific Northwest Woodland
Ponderosa pine Replacement 5% 200    
Mixed 17% 60    
Surface or low 78% 13    
Subalpine woodland Replacement 21% 300 200 400
Mixed 79% 80 35 120
Pacific Northwest Forested
California mixed evergreen (northern California and southern Oregon) Replacement 6% 150 100 200
Mixed 29% 33 15 50
Surface or low 64% 15 5 30
Douglas-fir (Willamette Valley foothills) Replacement 18% 150 100 400
Mixed 29% 90 40 150
Surface or low 53% 50 20 80
Douglas-fir-western hemlock (dry mesic) Replacement 25% 300 250 500
Mixed 75% 100 50 150
Douglas-fir-western hemlock (wet mesic) Replacement 71% 400    
Mixed 29% >1,000    
Lodgepole pine (pumice soils) Replacement 78% 125 65 200
Mixed 22% 450 45 85
Mixed conifer (eastside mesic) Replacement 35% 200    
Mixed 47% 150    
Surface or low 18% 400    
Mixed conifer (southwestern Oregon) Replacement 4% 400    
Mixed 29% 50    
Surface or low 67% 22    
Mountain hemlock Replacement 93% 750 500 >1,000
Mixed 7% >1,000    
Oregon coastal tanoak Replacement 10% 250    
Mixed 90% 28 15 40
Pacific silver fir (low elevation) Replacement 46% 350 100 800
Mixed 54% 300 100 400
Pacific silver fir (high elevation) Replacement 69% 500    
Mixed 31% >1,000    
Red fir Replacement 20% 400 150 400
Mixed 80% 100 80 130
Sitka spruce-western hemlock Replacement 100% 700 300 >1,000
Spruce-fir Replacement 84% 135 80 270
Mixed 16% 700 285 >1,000
Subalpine fir Replacement 81% 185 150 300
Mixed 19% 800 500 >1,000
California
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)
Minimum interval
(years)
Maximum interval
(years)
California Grassland
Alpine meadows and barrens Replacement 100% 200 200 400
Wet mountain meadow-lodgepole pine (subalpine) Replacement 21% 100    
Mixed 10% 200    
Surface or low 69% 30    
California Woodland
Ponderosa pine Replacement 5% 200    
Mixed 17% 60    
Surface or low 78% 13    
California Forested
Aspen with conifer Replacement 24% 155 50 300
Mixed 15% 240    
Surface or low 61% 60    
California mixed evergreen Replacement 10% 140 65 700
Mixed 58% 25 10 33
Surface or low 32% 45 7  
Coast redwood Replacement 2% ≥1,000    
Surface or low 98% 20    
Jeffrey pine Replacement 9% 250    
Mixed 17% 130    
Surface or low 74% 30    
Interior white fir (northeastern California) Replacement 47% 145    
Mixed 32% 210    
Surface or low 21% 325    
Mixed conifer (north slopes) Replacement 5% 250    
Mixed 7% 200    
Surface or low 88% 15 10 40
Mixed conifer (south slopes) Replacement 4% 200    
Mixed 16% 50    
Surface or low 80% 10    
Red fir-western white pine Replacement 16% 250    
Mixed 65% 60 25 80
Surface or low 19% 200    
Red fir-white fir Replacement 13% 200 125 500
Mixed 36% 70    
Surface or low 51% 50 15 50
Sierra Nevada lodgepole pine (cold wet upper montane) Replacement 23% 150 37 764
Mixed 70% 50    
Surface or low 7% 500    
Southwest
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)
Minimum interval
(years)
Maximum interval
(years)
Southwest Grassland
Montane and subalpine grasslands Replacement 55% 18 10 100
Surface or low 45% 22    
Montane and subalpine grasslands with shrubs or trees Replacement 30% 70 10 100
Surface or low 70% 30    
Southwest Woodland
Bristlecone-limber pine (Southwest) Replacement 67% 500    
Surface or low 33% >1,000    
Riparian deciduous woodland Replacement 50% 110 15 200
Mixed 20% 275 25  
Surface or low 30% 180 10  
Southwest Forested
Aspen, stable without conifers Replacement 81% 150 50 300
Surface or low 19% 650 600 >1,000
Aspen with spruce-fir Replacement 38% 75 40 90
Mixed 38% 75 40  
Surface or low 23% 125 30 250
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
Great Basin
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)
Minimum interval
(years)
Maximum interval
(years)
Great Basin Grassland
Mountain meadow (mesic to dry) Replacement 66% 31 15 45
Mixed 34% 59 30 90
Great Basin Forested
Aspen with conifer (low to midelevations) Replacement 53% 61 20  
Mixed 24% 137 10  
Surface or low 23% 143 10  
Aspen with conifer (high elevations) Replacement 47% 76 40  
Mixed 18% 196 10  
Surface or low 35% 100 10  
Aspen-cottonwood, stable aspen without conifers Replacement 31% 96 50 300
Surface or low 69% 44 20 60
Aspen, stable without conifers Replacement 81% 150 50 300
Surface or low 19% 650 600 >1,000
Aspen with spruce-fir Replacement 38% 75 40 90
Mixed 38% 75 40  
Surface or low 23% 125 30 250
Douglas-fir (interior, warm mesic) Replacement 28% 170 80 400
Mixed 72% 65 50 250
Ponderosa pine-Douglas-fir Replacement 10% 250   >1,000
Mixed 51% 50 50 130
Surface or low 39% 65 15  
Spruce-fir-pine (subalpine) Replacement 98% 217 75 300
Mixed 2% >1,000    
Northern and Central Rockies
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)
Minimum interval
(years)
Maximum interval
(years)
Northern and Central Rockies Grassland
Mountain grassland Replacement 60% 20 10  
Mixed 40% 30    
Northern and Central Rockies Forested
Douglas-fir (cold) Replacement 31% 145 75 250
Mixed 69% 65 35 150
Douglas-fir (warm mesic interior) Replacement 28% 170 80 400
Mixed 72% 65 50 250
Grand fir-Douglas-fir-western larch mix Replacement 29% 150 100 200
Mixed 71% 60 3 75
Grand fir-lodgepole pine-western larch-Douglas-fir Replacement 31% 220 50 250
Mixed 69% 100 35 150
Lodgepole pine, lower subalpine Replacement 73% 170 50 200
Mixed 27% 450 40 500
Lower subalpine (Wyoming and Central Rockies) Replacement 100% 175 30 300
Mixed-conifer-upland western redcedar-western hemlock Replacement 67% 225 150 300
Mixed 33% 450 35 500
Ponderosa pine (Black Hills, high elevation) Replacement 12% 300    
Mixed 18% 200    
Surface or low 71% 50    
Ponderosa pine (Northern and Central Rockies) Replacement 4% 300 100 >1,000
Mixed 19% 60 50 200
Surface or low 77% 15 3 30
Ponderosa pine-Douglas-fir Replacement 10% 250   >1,000
Mixed 51% 50 50 130
Surface or low 39% 65 15  
Western larch-lodgepole pine-Douglas-fir Replacement 33% 200 50 250
Mixed 67% 100 20 140
Whitebark pine-lodgepole pine (upper subalpine, Northern and Central Rockies) Replacement 38% 360    
Mixed 62% 225    
Upper subalpine spruce-fir (Central Rockies) Replacement 100% 300 100 600
Western redcedar Replacement 87% 385 75 >1,000
Mixed 13% >1,000 25  
Northern Great Plains
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)
Minimum interval
(years)
Maximum interval
(years)
Northern Plains Woodland
Great Plains floodplain Replacement 100% 500    
Northern Great Plains wooded draws and ravines Replacement 38% 45 30 100
Mixed 18% 94    
Surface or low 43% 40 10  
Great Lakes
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)
Minimum interval
(years)
Maximum interval
(years)
Great Lakes Woodland
Great Lakes pine barrens Replacement 8% 41 10 80
Mixed 9% 36 10 80
Surface or low 83% 4 1 20
Northern oak savanna Replacement 4% 110 50 500
Mixed 9% 50 15 150
Surface or low 87% 5 1 20
Great Lakes Forested
Eastern white pine-eastern hemlock Replacement 54% 370    
Mixed 12% >1,000    
Surface or low 34% 588    
Great Lakes floodplain forest Mixed 7% 833    
Surface or low 93% 61    
Great Lakes pine forest, eastern white pine-eastern hemlock (frequent fire) Replacement 52% 260    
Mixed 12% >1,000    
Surface or low 35% 385    
Great Lakes spruce-fir Replacement 100% 85 50 200
Maple-basswood Replacement 33% >1,000    
Surface or low 67% 500    
Maple-basswood mesic hardwood forest (Great Lakes) Replacement 100% >1,000 >1,000 >1,000
Maple-basswood-oak-aspen Replacement 4% 769    
Mixed 7% 476    
Surface or low 89% 35    
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    
Oak-hickory Replacement 13% 66 1  
Mixed 11% 77 5  
Surface or low 76% 11 2 25
Red pine-eastern white pine (less frequent fire) Replacement 30% 166    
Mixed 47% 105    
Surface or low 23% 220    
Northeast
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)
Minimum interval
(years)
Maximum interval
(years)
Northeast Woodland
Eastern woodland mosaic Replacement 2% 200 100 300
Mixed 9% 40 20 60
Surface or low 89% 4 1 7
Pine barrens Replacement 10% 78    
Mixed 25% 32    
Surface or low 65% 12    
Rocky outcrop pine (Northeast) Replacement 16% 128    
Mixed 32% 65    
Surface or low 52% 40    
Northeast Forested
Beech-maple Replacement 100% >1,000    
Eastern white pine-northern hardwood Replacement 72% 475    
Surface or low 28% >1,000    
Northern hardwoods (Northeast) Replacement 39% >1,000    
Mixed 61% 650    
Northern hardwoods-eastern hemlock Replacement 50% >1,000    
Surface or low 50% >1,000    
Northern hardwoods-spruce Replacement 100% >1,000 400 >1,000
Northeast spruce-fir forest Replacement 100% 265 150 300
*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 [7,36].


Daisy-leaf moonwort:

Fire regime information on vegetation communities in which daisy-leaf moonwort may occur. This information is taken from the LANDFIRE Rapid Assessment Vegetation Models [37], 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 Great Plains Great Lakes Northeast Southern Appalachians
Northern Great Plains
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)
Minimum interval
(years)
Maximum interval
(years)
Northern Plains Woodland
Great Plains floodplain Replacement 100% 500    
Northern Great Plains wooded draws and ravines Replacement 38% 45 30 100
Mixed 18% 94    
Surface or low 43% 40 10  
Great Lakes
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)
Minimum interval
(years)
Maximum interval
(years)
Great Lakes Woodland
Great Lakes pine barrens Replacement 8% 41 10 80
Mixed 9% 36 10 80
Surface or low 83% 4 1 20
Northern oak savanna Replacement 4% 110 50 500
Mixed 9% 50 15 150
Surface or low 87% 5 1 20
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 floodplain forest Mixed 7% 833    
Surface or low 93% 61    
Great Lakes pine forest, eastern white pine-eastern hemlock (frequent fire) Replacement 52% 260    
Mixed 12% >1,000    
Surface or low 35% 385    
Great Lakes spruce-fir Replacement 100% 85 50 200
Maple-basswood Replacement 33% >1,000    
Surface or low 67% 500    
Maple-basswood mesic hardwood forest (Great Lakes) Replacement 100% >1,000 >1,000 >1,000
Maple-basswood-oak-aspen Replacement 4% 769    
Mixed 7% 476    
Surface or low 89% 35    
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    
Oak-hickory Replacement 13% 66 1  
Mixed 11% 77 5  
Surface or low 76% 11 2 25
Red pine-eastern white pine (less frequent fire) Replacement 30% 166    
Mixed 47% 105    
Surface or low 23% 220    
Northeast
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)
Minimum interval
(years)
Maximum interval
(years)
Northeast Woodland
Eastern woodland mosaic Replacement 2% 200 100 300
Mixed 9% 40 20 60
Surface or low 89% 4 1 7
Pine barrens Replacement 10% 78    
Mixed 25% 32    
Surface or low 65% 12    
Northeast Forested
Appalachian oak forest (dry-mesic) Replacement 2% 625 500 >1,000
Mixed 6% 250 200 500
Surface or low 92% 15 7 26
Beech-maple Replacement 100% >1,000    
Eastern white pine-northern hardwood Replacement 72% 475    
Surface or low 28% >1,000    
Northern hardwoods (Northeast) Replacement 39% >1,000    
Mixed 61% 650    
Northern hardwoods-eastern hemlock Replacement 50% >1,000    
Surface or low 50% >1,000    
Northern hardwoods-spruce Replacement 100% >1,000 400 >1,000
Northeast spruce-fir forest Replacement 100% 265 150 300
Southeastern red spruce-Fraser fir Replacement 100% 500 300 >1,000
Southern Appalachians
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)
Minimum interval
(years)
Maximum interval
(years)
Southern Appalachians Woodland
Oak-ash woodland Replacement 23% 119    
Mixed 28% 95    
Surface or low 49% 55    
Southern Appalachians Forested
Appalachian oak forest (dry-mesic) Replacement 6% 220    
Mixed 15% 90    
Surface or low 79% 17    
Bottomland hardwood forest Replacement 25% 435 200 >1,000
Mixed 24% 455 150 500
Surface or low 51% 210 50 250
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 [7,36].


Mountain moonwort:

Fire regime information on vegetation communities in which moonwort may occur. This information is taken from the LANDFIRE Rapid Assessment Vegetation Models [37], 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.
Pacific Northwest California Northern and Central Rockies
Pacific Northwest
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)
Minimum interval
(years)
Maximum interval
(years)
Pacific Northwest Grassland
Alpine and subalpine meadows and grasslands Replacement 68% 350 200 500
Mixed 32% 750 500 >1,000
Idaho fescue grasslands Replacement 76% 40    
Mixed 24% 125    
Pacific Northwest Woodland
Subalpine woodland Replacement 21% 300 200 400
Mixed 79% 80 35 120
Pacific Northwest Forested
California mixed evergreen (northern California and southern Oregon) Replacement 6% 150 100 200
Mixed 29% 33 15 50
Surface or low 64% 15 5 30
Douglas-fir (Willamette Valley foothills) Replacement 18% 150 100 400
Mixed 29% 90 40 150
Surface or low 53% 50 20 80
Douglas-fir-western hemlock (dry mesic) Replacement 25% 300 250 500
Mixed 75% 100 50 150
Douglas-fir-western hemlock (wet mesic) Replacement 71% 400    
Mixed 29% >1,000    
Mixed conifer (eastside mesic) Replacement 35% 200    
Mixed 47% 150    
Surface or low 18% 400    
Mixed conifer (southwestern Oregon) Replacement 4% 400    
Mixed 29% 50    
Surface or low 67% 22    
Mountain hemlock Replacement 93% 750 500 >1,000
Mixed 7% >1,000    
Ponderosa pine, dry (mesic) Replacement 5% 125    
Mixed 13% 50    
Surface or low 82% 8    
Pacific silver fir (low elevation) Replacement 46% 350 100 800
Mixed 54% 300 100 400
Pacific silver fir (high elevation) Replacement 69% 500    
Mixed 31% >1,000    
Red fir Replacement 20% 400 150 400
Mixed 80% 100 80 130
Sitka spruce-western hemlock Replacement 100% 700 300 >1,000
Spruce-fir Replacement 84% 135 80 270
Mixed 16% 700 285 >1,000
Subalpine fir Replacement 81% 185 150 300
Mixed 19% 800 500 >1,000
California
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)
Minimum interval
(years)
Maximum interval
(years)
California Grassland
Alpine meadows and barrens Replacement 100% 200 200 400
Wet mountain meadow-lodgepole pine (subalpine) Replacement 21% 100    
Mixed 10% 200    
Surface or low 69% 30    
California Forested
Aspen with conifer Replacement 24% 155 50 300
Mixed 15% 240    
Surface or low 61% 60    
California mixed evergreen Replacement 10% 140 65 700
Mixed 58% 25 10 33
Surface or low 32% 45 7  
Coast redwood Replacement 2% ≥1,000    
Surface or low 98% 20    
Jeffrey pine Replacement 9% 250    
Mixed 17% 130    
Surface or low 74% 30    
Interior white fir (northeastern California) Replacement 47% 145    
Mixed 32% 210    
Surface or low 21% 325    
Mixed conifer (north slopes) Replacement 5% 250    
Mixed 7% 200    
Surface or low 88% 15 10 40
Mixed conifer (south slopes) Replacement 4% 200    
Mixed 16% 50    
Surface or low 80% 10    
Red fir-western white pine Replacement 16% 250    
Mixed 65% 60 25 80
Surface or low 19% 200    
Red fir-white fir Replacement 13% 200 125 500
Mixed 36% 70    
Surface or low 51% 50 15 50
Sierra Nevada lodgepole pine (cold wet upper montane) Replacement 23% 150 37 764
Mixed 70% 50    
Surface or low 7% 500    
Sierra Nevada lodgepole pine (dry subalpine) Replacement 11% 250 31 500
Mixed 45% 60 31 350
Surface or low 45% 60 9 350
Northern and Central Rockies
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)
Minimum interval
(years)
Maximum interval
(years)
Northern and Central Rockies Grassland
Mountain grassland Replacement 60% 20 10  
Mixed 40% 30    
Northern and Central Rockies Forested
Douglas-fir (cold) Replacement 31% 145 75 250
Mixed 69% 65 35 150
Douglas-fir (warm mesic interior) Replacement 28% 170 80 400
Mixed 72% 65 50 250
Grand fir-Douglas-fir-western larch mix Replacement 29% 150 100 200
Mixed 71% 60 3 75
Grand fir-lodgepole pine-western larch-Douglas-fir Replacement 31% 220 50 250
Mixed 69% 100 35 150
Lodgepole pine, lower subalpine Replacement 73% 170 50 200
Mixed 27% 450 40 500
Lower subalpine (Wyoming and Central Rockies) Replacement 100% 175 30 300
Mixed-conifer-upland western redcedar-western hemlock Replacement 67% 225 150 300
Mixed 33% 450 35 500
Ponderosa pine-Douglas-fir Replacement 10% 250   >1,000
Mixed 51% 50 50 130
Surface or low 39% 65 15  
Western larch-lodgepole pine-Douglas-fir Replacement 33% 200 50 250
Mixed 67% 100 20 140
Whitebark pine-lodgepole pine (upper subalpine, Northern and Central Rockies) Replacement 38% 360    
Mixed 62% 225    
Upper subalpine spruce-fir (Central Rockies) Replacement 100% 300 100 600
Western redcedar Replacement 87% 385 75 >1,000
Mixed 13% >1,000 25  
*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 [7,36].


Peculiar moonwort:

Fire regime information on vegetation communities in which peculiar moonwort may occur. This information is taken from the LANDFIRE Rapid Assessment Vegetation Models [37], 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.
Pacific Northwest California Great Basin Northern and Central Rockies
Pacific Northwest
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)
Minimum interval
(years)
Maximum interval
(years)
Pacific Northwest Grassland
Alpine and subalpine meadows and grasslands Replacement 68% 350 200 500
Mixed 32% 750 500 >1,000
Bluebunch wheatgrass Replacement 47% 18 5 20
Mixed 53% 16 5 20
Idaho fescue grasslands Replacement 76% 40    
Mixed 24% 125    
Pacific Northwest Woodland
Subalpine woodland Replacement 21% 300 200 400
Mixed 79% 80 35 120
Pacific Northwest Forested
California mixed evergreen (northern California and southern Oregon) Replacement 6% 150 100 200
Mixed 29% 33 15 50
Surface or low 64% 15 5 30
Douglas-fir (Willamette Valley foothills) Replacement 18% 150 100 400
Mixed 29% 90 40 150
Surface or low 53% 50 20 80
Douglas-fir-western hemlock (wet mesic) Replacement 71% 400    
Mixed 29% >1,000    
Mixed conifer (eastside mesic) Replacement 35% 200    
Mixed 47% 150    
Surface or low 18% 400    
Mixed conifer (southwestern Oregon) Replacement 4% 400    
Mixed 29% 50    
Surface or low 67% 22    
Mountain hemlock Replacement 93% 750 500 >1,000
Mixed 7% >1,000    
Pacific silver fir (low elevation) Replacement 46% 350 100 800
Mixed 54% 300 100 400
Pacific silver fir (high elevation) Replacement 69% 500    
Mixed 31% >1,000    
Red fir Replacement 20% 400 150 400
Mixed 80% 100 80 130
Sitka spruce-western hemlock Replacement 100% 700 300 >1,000
Spruce-fir Replacement 84% 135 80 270
Mixed 16% 700 285 >1,000
Subalpine fir Replacement 81% 185 150 300
Mixed 19% 800 500 >1,000
California
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)
Minimum interval
(years)
Maximum interval
(years)
California Grassland
Alpine meadows and barrens Replacement 100% 200 200 400
Wet mountain meadow-lodgepole pine (subalpine) Replacement 21% 100    
Mixed 10% 200    
Surface or low 69% 30    
California Forested
Aspen with conifer Replacement 24% 155 50 300
Mixed 15% 240    
Surface or low 61% 60    
California mixed evergreen Replacement 10% 140 65 700
Mixed 58% 25 10 33
Surface or low 32% 45 7  
Interior white fir (northeastern California) Replacement 47% 145    
Mixed 32% 210    
Surface or low 21% 325    
Mixed conifer (north slopes) Replacement 5% 250    
Mixed 7% 200    
Surface or low 88% 15 10 40
Red fir-western white pine Replacement 16% 250    
Mixed 65% 60 25 80
Surface or low 19% 200    
Red fir-white fir Replacement 13% 200 125 500
Mixed 36% 70    
Surface or low 51% 50 15 50
Sierra Nevada lodgepole pine (cold wet upper montane) Replacement 23% 150 37 764
Mixed 70% 50    
Surface or low 7% 500    
Great Basin
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)
Minimum interval
(years)
Maximum interval
(years)
Great Basin Grassland
Mountain meadow (mesic to dry) Replacement 66% 31 15 45
Mixed 34% 59 30 90
Great Basin Forested
Aspen with conifer (low to midelevations) Replacement 53% 61 20  
Mixed 24% 137 10  
Surface or low 23% 143 10  
Aspen with conifer (high elevations) Replacement 47% 76 40  
Mixed 18% 196 10  
Surface or low 35% 100 10  
Aspen-cottonwood, stable aspen without conifers Replacement 31% 96 50 300
Surface or low 69% 44 20 60
Aspen, stable without conifers Replacement 81% 150 50 300
Surface or low 19% 650 600 >1,000
Aspen with spruce-fir Replacement 38% 75 40 90
Mixed 38% 75 40  
Surface or low 23% 125 30 250
Spruce-fir-pine (subalpine) Replacement 98% 217 75 300
Mixed 2% >1,000    
Northern and Central Rockies
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)
Minimum interval
(years)
Maximum interval
(years)
Northern and Central Rockies Grassland
Mountain grassland Replacement 60% 20 10  
Mixed 40% 30    
Northern and Central Rockies Forested
Douglas-fir (cold) Replacement 31% 145 75 250
Mixed 69% 65 35 150
Grand fir-Douglas-fir-western larch mix Replacement 29% 150 100 200
Mixed 71% 60 3 75
Grand fir-lodgepole pine-western larch-Douglas-fir Replacement 31% 220 50 250
Mixed 69% 100 35 150
Lodgepole pine, lower subalpine Replacement 73% 170 50 200
Mixed 27% 450 40 500
Mixed-conifer-upland western redcedar-western hemlock Replacement 67% 225 150 300
Mixed 33% 450 35 500
Ponderosa pine (Black Hills, low elevation) Replacement 7% 300 200 400
Mixed 21% 100 50 400
Surface or low 71% 30 5 50
Western larch-lodgepole pine-Douglas-fir Replacement 33% 200 50 250
Mixed 67% 100 20 140
Whitebark pine-lodgepole pine (upper subalpine, Northern and Central Rockies) Replacement 38% 360    
Mixed 62% 225    
Upper subalpine spruce-fir (Central Rockies) Replacement 100% 300 100 600
Western redcedar Replacement 87% 385 75 >1,000
Mixed 13% >1,000 25  
*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 [7,36].

APPENDIX B: Common and scientific names of plants
These species are assoicated with common, daisy-leaf-, mountain, and/or peculiar moonwort. Follow the links to FEIS reviews for further information.
Common name Scientific name
Trees
alpine larch Larix lyallii
balsam fir Abies balsamea
beech Fagus spp.
bur oak Quercus macrocarpa
eastern hemlock Tsuga canadensis
eastern white pine Pinus strobus
Engelmann spruce Picea engelmannii
fir Abies spp.
incense-cedar Calocedrus decurrens
interior ponderosa pine Pinus ponderosa var. scopularum
limber pine Pinus flexilis
maple Acer spp.
mountain hemlock Tsuga mertensiana
northern red oak Quercus rubra
Pacific ponderosa pine Pinus ponderosa var. ponderosa
pin oak Quercus palustris
pitch pine Pinus rigida
quaking aspen Populus tremuloides
red maple Acer rubrum
Rocky Mountain Douglas-fir Pseudotsuga menziesii var. glauca
Rocky Mountain lodgepole pine Pinus contorta var. latifolia
subalpine fir Abies lasiocarpa
sugar pine Pinus lambertiana
western redcedar Thuja plicata
white fir Abies concolor
white oak Quercus alba
white spruce Picea glauca
yellow-poplar Liriodendron tulipifera
Shrubs
arctic dwarf birch Betula nana subsp. exilis
bog birch Betula glandulosa
eastern poison-ivy  Toxicodendron radicans
flowering dogwood  Cornus florida
grayleaf willow Salix glauca
kinnikinnick Arctostaphylos uva-ursi
mountain cranberry Vaccinium vitis-idaea
shrubby cinquefoil Dasiphora floribunda
Forbs
fireweed Chamerion angustifolium
Ross's avens Geum rosii var. turbinatum
Graminoids
arctic brome Bromus pumpellianus
bluejoint reedgrass Calamagrostis canadensis
boreal wildrye Leymus innovatus
Idaho fescue Festuca idahoensis
northern rough fescue Festuca altaica
sedges Carex spp.
slender wheatgrass Elymus trachycaulus
Fern allies
alpine clubmoss Lycopodium alpinum
clubmosses Lycopodium spp.
cutleaf grapefern  Botrychium dissectum
dainty moonwort Botrychium crenulatum
fir clubmoss Huperzia selago
Frenchman's Bluff moonwort Botrychium gallicomontanum
giant moonwort  Botrychium yaaxudakeit
grapeferns Botrychium spp., subgenus Sceptridium
Iowa moonwort Botrychium campestre
lanceleaf moonwort Botrychium lanceolatum
least moonwort Botrychium simplex
leathery grapefern Botrychium multifidum
Mingan moonwort Botrychium minganense
moonworts Botrychium spp., subgenus Botrychium
pale botrychium Botrychium pallidum
pointed moonwort Botrychium acuminatum
Sitka clubmoss Lycopodium sitchense
spathulate botrychium Botrychium spathulatum
tailed grapefern Botrychium ascendens
Tunux moonwort Botrychium tunux
Waterton moonwort Botrychium × watertonense
western moonwort Botrychium hesperium

REFERENCES:


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8. Barton, Drake.; Crispin, Sue. 2004. Conservation status of Botrychium lineare (slender moonwort) in Montana. Helena, MT: Montana Natural Heritage Program. 6 p. [87004]
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12. Cody, William J.; Britton, Donald M. 1989. Ferns and fern allies of Canada. Ottawa, ON: Agriculture Canada, Research Branch. 430 p. [13078]
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14. Cronquist, Arthur; Holmgren, Arthur H.; Holmgren, Noel H.; Reveal, James L. 1972. Intermountain flora: Vascular plants of the Intermountain West, U.S.A. Vol. 1. New York: Hafner Publishing Company, Inc. 270 p. [717]
15. Dauphin, Benjamin; Vieu, Julien; Grant, Jason R. 2014. Molecular phylogentics supports widespread cryptic species in moonworts (Botrychium s.s., Ophioglossaceae). American Journal of Botany. 101(1): 128-140. [87728]
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