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Management Recommendations
for
Mingan moonwort (Botrychium minganense Victorin)

v. 2.0

by

Laura L. Potash

December 1998


TABLE OF CONTENTS

EXECUTIVE SUMMARY 1
I. NATURAL HISTORY 3
A. Taxonomic/Nomenclatural History 3
B. Species Description 3
1. Morphology 3
2. Reproductive Biology 4
3. Ecology 4
C. Range, Known Sites 5
D. Habitat Characteristics and Species Abundance 6
II. CURRENT SPECIES SITUATION 6
A. Why Species is Listed under Survey and Manage Standards and Guidelines 6
B. Major Habitat and Viability Considerations 7
C. Threats to the Species 7
D. Distribution Relative to Land Allocations 8
III. MANAGEMENT GOALS AND OBJECTIVES 9
A. Management Goals for the Taxon 9
B. Specific Objectives 9
IV. HABITAT MANAGEMENT 9
A. Lessons from History 9
B. Identification of Habitat Areas for Management 9
C. Management Within Habitat Areas 9
D. Other Management Issues and Considerations 11
V. RESEARCH, INVENTORY, AND MONITORING NEEDS 11
A. Data Gaps and Information Needs 11
B. Research Questions 11
C. Monitoring Needs and Recommendations 12
VI. REFERENCES 13

EXECUTIVE SUMMARY

Species: Botrychium minganense Victorin (Mingan moonwort)

Taxonomic Group: Vascular Plants

ROD Components: 1, 2

Other Management Status: Region 6, Regional Forester's Sensitive Species List (USDA, Forest Service 1991) and BLM's Bureau Assessment List. It is considered sensitive in Oregon and ranked as S2, which is a species that is imperiled because of rarity or because it is vulnerable to extinction or extirpation: typically 6 to 20 occurrences (Oregon Natural Heritage Program 1995). In Washington, Botrychium minganense was previously listed as sensitive but has recently been moved to Review Group 2, which are those taxa with unresolved taxonomic questions (Washington Natural Heritage Program 1997).

Range: On federally owned lands within the range of the northern spotted owl, it occurs on the Gifford Pinchot, Mt. Baker-Snoqualmie, Wenatchee, and Okanogan National Forests (Table 1) in Chelan, Whatcom, Snohomish, Kittitas, Skamania, Lewis, Okanogan, King, and Pierce counties. In Oregon it has been documented on Mt. Hood National Forest and Willamette National Forest (Table 1) in Hood River and Linn counties. There has recently been a documented sighting of Botrychium minganense on the Bureau of Land Management's Eugene District in Oregon. There are currently 2 known sites of Botrychium minganense in California, located in Fresno and Tehama counties (Skinner and Pavlik 1994; California Natural Diversity Database 1998). The California sites for Botrychium minganense are not within the analysis area for the Northwest Forest Plan.

Specific Habitat: Most populations in Oregon and east of the Cascades in Washington are associated with riparian zones and old-growth western red cedar (Thuja plicata) in dense shade, sparse understory, an alluvium substrate, and often a duff layer of Thuja branchlets. However, on the Gifford Pinchot and Mt. Baker-Snoqualmie forests, all known sites are located in subalpine meadows, ski slopes, mossy boulder fields under bigleaf maple, road cuts, shrublands, and alder thickets.

Threats: Potential threats to Botrychium minganense may include exotic weed invasion, trampling by recreational users or livestock, soil compaction, fire, livestock grazing, burial by surface deposition, and timber harvest.

Management Recommendations: Viability of Botrychium minganense is expected to be secure in Washington State under current ROD land allocations. The following measures are only recommended for populations within the analysis area in Oregon and California .

  • Maintain existing hydrologic regime.
  • Provide conditions to maintain fungal diversity to support presumed mycorrhizal relationships between Botrychium minganense and western red cedar.
  • Maintain deep shade.
  • Maintain microclimatic conditions in old-growth stands.
  • Avoid excessive siltation or deposition of soil.
  • Avoid disturbance of duff layer where the species occurs.
  • Avoid actions that would contribute towards establishment of competing exotic vegetation and manually remove noxious weeds.
  • Avoid impacts caused by livestock such as trampling or grazing.

Information Needs: What is the genetic basis for defining Botrychium minganense as a species - should the taxon be split into 4 different species or otherwise reclassified based on genetic analyses? Are periodic floods part of the natural disturbance regime at these sites? Is there a correlation between the previous year's weather and population numbers? Are Thuja plicata trees necessary for this species to thrive within the range of the northern spotted owl? What role do mammals play in grazing and spore dispersal? What components of an old-growth forest stand are most critical for providing habitat for Botrychium minganense? For example, is it the low light levels, the soil moisture content, the soil microorganisms, or another factor that is the key component - or is it some interaction among these variables?

I. NATURAL HISTORY

A. Taxonomic/ Nomenclatural History

Botrychium minganense Victorin was originally described in 1927 in the Proceedings and Transcripts of the Royal Society of Canada (Victorin 1927), and is treated as such by Wagner and Wagner (1993) in Flora of North America. In the past this species was considered a variety of Botrychium lunaria, with the synonym Botrychium lunaria (Linnaeus) Swartz var. minganense (Victorin) Dole.

Based on current work in the Pacific Northwest, Wagner (1994) suggests that what is being called Mingan moonwort may in fact be 4 distinct species. If Botrychium minganense Victorin remains as a single taxon, then it is not as rare as some other members of the genus (e.g., Botrychium ascendens). However, if the taxon is split into 4 "new" species then at least one or more of the new species would be very rare indeed. Because the taxonomy of this genus is in such a state of uncertainty, these management recommendations will treat Botrychium minganense Victorin as one species at this time (as it is listed in Table C-3 of the ROD); however, if future changes in taxonomy become accepted by the federal land management agencies, then corresponding revisions in these management recommendations will be necessary. Current genetic research on Botrychium minganense (Linda Swartz, personal communication) may help resolve some of these taxonomic questions.

B. Species Description

1. Morphology

The above-ground or visible parts of this species consist of a single upright stem arising from the ground and terminating in a cluster of tiny ball-like structures that resemble a bunch of grapes (hence the often used common name for the genus of "grapefern"). These globular structures (the sporangia) contain the spores necessary for sexual reproduction. Branching off from the main stem is the sterile, fern-like leaf blade (the trophophore). In the Botrychium genus, the portion of the stem below this juncture of the sterile blade with the main stem is referred to as the common stalk, and the portion with the sporangia is called the fertile stalk (sporophore). At the base of the common stalk, but just below the ground, Botrychium species have several layers of leaf primordia that are the preformed buds of plants that will emerge in future years.

Botrychium minganense is a small, herbaceous, perennial fern. The sterile blade (trophophore) is dull green in color, narrowly oblong to linear in overall outline, about 10cm long x 2.5 cm wide. The sterile blade is once-pinnate, with up to 10 pairs of pinnae (segments). In general the segments are well developed, cuneate to flabellate in shape, and spaced separately from each other along the rachis. The margins of the segments are entire to shallowly crenate. The lowest segments are narrowly fan-shaped. The description uses many qualifying adjectives because this genus is notorious for morphological variation.

Botrychium minganense can be confused with Botrychium spathulatum but the latter species has a more leathery textured, deltate-shaped sterile blade, and is not suspected within the range of the northern spotted owl. In eastern Washington, Botrychium minganense is often difficult to distinguish from Botrychium crenulatum because of wide variations in leaf morphology, but the latter species is more yellow-green in color and the veins are more pronounced on the leaf segments (Swartz, personal communication). Within the range of the spotted owl, Botrychium minganense may often be confused with Botrychium lunaria. The main difference is that the leaf segments in B. lunaria are much more broadly fan-shaped and the pairs are arranged closely enough to overlap.

2. Reproductive Biology

Botrychium species are terrestrial ferns that reproduce by means of microscopic spores (Lellinger 1985). When the spore germinates, it develops into a tiny underground structure (gametophyte) that produces the gametes (egg and sperm). When the sperm is mature it is released from one part of the gametophyte and swims via a thin film of water to the egg. The fertilized zygote then develops roots, stem, and the above-ground structure seen as the fern (sporophyte). The sporophyte produces the spores by the thousands in round sacs (sporangia) borne in clusters at the top of the fertile stalk.

Much remains a mystery about the reproductive biology of this entire genus because traditional ways of observing genetic and reproductive traits of a species, through reciprocal transplants or common garden experiments, have been difficult to conduct with this genus, presumably because of their delicately attuned mycorrhizal relationships (Wagner and Wagner 1983). The growth rate is so slow that ordinarily only a single leaf is produced per year - primordia for several years are contained within the bud, and only one primordium matures each season. In Botrychium minganense, asexual reproduction is sometimes accomplished via gemmae, which are spherical units that are produced on the underground stem and are capable of developing into sporophytes (Farrar and Johnson-Groh 1990; Francisco Camacho, personal communication).

Another aspect of the biology of this genus with management implications is that several species apparently undergo periods of dormancy, where the plant will not emerge for one to several years and then reemerge in the exact same location (Gehring and Potash 1995; Montgomery 1990).

3. Ecology

As mentioned above, Botrychium species are thought to depend on mycorrhizal relationships with other species in order to thrive. Underground associations between the fern roots and fungi are apparently fragile but essential, and presumably are established during the gametophyte phase (Zika 1992; Camacho, personal communication). Certain species of fungi may be necessary for establishment and survival. It may be the fungal symbiont that is most affected by changes in canopy coverage, summer temperature, and soil moisture (Zika 1992). Gehring and Potash (1995) found that the distribution of several Botrychium species was independent of the distribution of western redcedar in a monitoring study near Mt. Baker in Washington, but it has been suggested that there may be a mycorrhizal association between Thuja plicata and Botrychium species (USDA, Forest Service and USDI, Bureau of Land Management 1994).

Dr. Warren Wagner suggests that spore ingestion by animals may be important as a dispersal mechanism and has observed many Botrychium species that have been grazed by deer (personal communication). On the Mt. Baker-Snoqualmie NF, evidence of grazing by slugs on Botrychium spp. has been observed.

C. Range, Known Sites

Botrychium minganense is a North American species that is wide ranging across Canada from coast to coast. In the United States the Flora of North America (Morin 1993) shows it occurring in Alaska, Washington, Oregon, Idaho, Montana, Utah, California, Nevada, Wyoming, Arizona, Colorado, North Dakota, Minnesota, Wisconsin, Michigan, New Hampshire, New York, Vermont, and Maine.

On federally owned lands in Washington within the range of the northern spotted owl, it occurs on the Gifford Pinchot, Mt. Baker-Snoqualmie, Wenatchee, and Okanogan National Forests (Table 1), in Chelan, Whatcom, Snohomish, Kittitas, Skamania, Lewis, Okanogan, King, and Pierce counties.

In Oregon, it has been documented on Mt. Hood National Forest and Willamette National Forest (Table 1) in Hood River and Linn counties. One site of Botrychium minganense has recently been located on the Eugene District of BLM. There are no documented sightings of Botrychium minganense within the range of the northern spotted owl in California (Skinner and Pavlik 1994; California Natural Diversity Database 1998).

Table 1. Federal land ownership and number of populations of Botrychium minganense
within the range of the northern spotted owl.

Ownership

Number of Records

Willamette National Forest

1

Mt. Hood National Forest

5

Gifford Pinchot National Forest

3

Mt. Baker-Snoqualmie National Forest

18

Wenatchee National Forest

41

Okanogan National Forest

9

Eugene District BLM

1

D. Habitat Characteristics and Species Abundance

It has been reported that Botrychium minganense "exhibits wide ecological amplitude, occurring in a wide range of habitats, particularly east of the Cascades, where it occurs in open shrubland and barren slopes. However, in the area under consideration [within the range of the northern spotted owl] it typically occurs in older forest stands." (USDA, Forest Service and USDI, Bureau of Land Management 1994). Many populations in Oregon and east of the Cascades in Washington seem to fit this description, as summarized by Zika (1992) in a report for Mt. Hood National Forest. The colonies are associated with riparian zones and old-growth western red cedar (Thuja plicata) in dense shade, sparse understory, on alluvium substrate and often a duff layer of Thuja branchlets. On the Wenatchee National Forest most records are from old-growth stands in river bottoms in the western hemlock/devil's club plant association or the western hemlock/wild ginger association (Steve Rust, personal communication). Soils on the Wenatchee sites are often saturated in the spring (in some instances individuals emerge under running snowmelt), but tend to dry out later in the growing season. Plants do not occur in soils wet enough to support skunkcabbage, but grow adjacent to these areas.

Based on recent analysis of all the sighting reports, there seems to be wide variation in the habitat preference of Botrychium minganense, even within the range of the northern spotted owl. For example, none of the known sites on the Gifford Pinchot or Mt. Baker-Snoqualmie are in old-growth forest. Instead, sites are located in subalpine meadows and ski slopes, mossy talus slopes under bigleaf maple, roadcuts, shrublands, and alder thickets. Most of the recent sightings on the Wenatchee National Forest are also in nonforested habitats. These populations occur in lush herbaceous meadows (Steve Rust, personal communication). Botrychium minganense is considered relatively abundant throughout its range, and it can grow in a wide variety of habitats, including some highly disturbed areas.

II. CURRENT SPECIES SITUATION

A. Why Species is Listed under Survey and Manage Standards and Guidelines.

In Oregon it is vulnerable within the range of the northern spotted owl because of the limited number of sightings and close association with old-growth. Qualified botanists have conducted extensive surveys for Botrychium species in a variety of habitats in Oregon for many years, so the limited number of sightings in old-growth stands does not seem to be an artifact of limited surveys.

In Washington, however, based on analysis of the most current sighting reports, the viability of this species may not be a concern for 2 reasons: In general, populations west of the Washington Cascades do not seem to be closely associated with old-growth. East of the Cascades in Washington, although Botrychium minganense is often associated with old-growth, there are many populations with numerous individuals at each site, and virtually all of them are located in areas with protected land use designations (see section II.D. below). In addition, most of the recent sightings on the Wenatchee National Forest are in non-forested habitats.

B. Major Habitat and Viability Considerations

The major viability considerations for Botrychium minganense are loss of populations due to management actions that change the habitat (canopy coverage, summer temperature, and soil moisture), disturb the presumed mycorrhizal connection between Botrychium species and western red cedar (Thuja plicata), or loss of populations through direct impact caused by trampling, excavation, or burial. Climate change has the potential to affect this species if occupied sites become much warmer and drier, since almost all of the known sites are relatively cool and moist. Based on professional judgement, the relative risk to viability of Botrychium minganense in Oregon is moderate under existing ROD land allocations, and low if the management recommendations in this document are implemented. In Washington State the relative risk to viability of Botrychium minganense is low under existing ROD land allocations without the additional provisions of the "survey and manage" standards and guidelines.

C. Threats to the Species

Potential threats to Botrychium minganense may include:

  • Timber harvest may pose indirect impacts in those portions of the range where Botrychium minganense appears to be closely associated with old-growth, because of significant changes in light regime, hydrology, temperature, and microclimate that occurs when a forested stand is cut down (Fritschen et al. 1971; Chen et al. 1995). Direct impacts from timber harvest would occur if logs are yarded across Botrychium minganense. In May 1996, two of the largest populations on the Mt. Hood National Forest were traded to private landowner who planned to log the sites (J. S. Nugent, personal communication). These populations represented 1/3 of the known sites on the Mt. Hood National Forest.
  • Habitat degradation of native plant communities resulting from exotic weed invasion is a well documented concern and may pose a threat to the habitat of Botrychium minganense.
  • Trampling by recreational users would probably be harmful to this species. Botrychium minganense is a small herbaceous plant that is easily crushed.
  • Soil compaction would presumably have an adverse effect on the underground primordial buds of this species. Over half of the known populations occur on alluvial soil that is somewhat loose in texture.
  • Botrychium minganense may respond poorly to fire; however, the reaction is unknown at the current time.
  • Livestock may have an adverse impact on Botrychium minganense for several reasons. Native species in the Pacific Northwest have not coevolved to be well adapted to large grazing herbivores, and generally do not respond well to this impact. While there is evidence that Botrychium species have been grazed by deer, the impacts from these animals are not equivalent to domestic stock because the latter weigh significantly more and crop plants to ground level. Therefore, in addition to the complete elimination of above-ground growth (by being eaten to ground level), cattle and horse grazing may impact Botrychium minganense due to increased trampling, soil compaction, and introduction of exotic weeds.
  • Burial by surface deposition (resulting from erosion after during construction, flooding, or other events) could directly impact Botrychium minganense because of the small size of this species.

D. Distribution Relative to Land Allocations

As of November 1998 there are 78 known records of Botrychium minganense within the range of the northern spotted owl. Distribution according to land allocation is summarized in Table 2.

Table 2. Land allocation of known sites of Botrychium minganense
within the range of the northern spotted owl.

(Source: Survey and Manage Known Sites Interim Database 6/95 and
Mt. Baker-Snoqualmie NF rare plants database.)

Land Management Designation

Number of Sites

Adaptive Management Area (AMA)

1

Late-Successional Reserves (LSR)

28

Administratively Withdrawn and AMA

1

Administratively Withdrawn and LSR

6

Administratively Withdrawn

7

Congressionally Withdrawn

15

Managed LSR

1

Matrix

10

Nonfederal Lands

9

TOTAL

78

Populations east of the Cascades in Washington are numerous, with virtually all of them located in areas with protected land use designations. On the Wenatchee National Forest, where the vast majority of the populations are located, all but one site is in an LSR, Wilderness Area, Administratively Withdrawn Area, or Riparian Reserve. The one population without formal protected status is located in an Adaptive Management Area on the Cle Elum Ranger District, but the site was dropped from a proposed timber sale in order to protect Botrychium minganense.

III. MANAGEMENT GOALS AND OBJECTIVES

A. Management Goals for the Taxon

The management goal for Botrychium minganense is to assist in maintaining species viability within the range of the northern spotted owl.

B. Specific Objectives

  • Maintain populations and habitat throughout the species' geographic range.
  • Maintain existing ecological conditions such as hydrologic regime, temperature regime, and light regime. Maintain associated native plant community. These objectives are based on the assumption that the habitat where the majority of populations exist is the optimal habitat for the viability of this species, unless there is evidence to the contrary.
  • Maintain viable populations of Botrychium minganense within occupied habitat, including presumed mycorrhizal relationship with Thuja plicata.

IV. HABITAT MANAGEMENT

A. Lessons from History

Botrychium minganense is often found growing with other Botrychium species (Wagner and Wagner 1983), a fact that may be helpful in conducting surveys. The more common species are often bigger and easier to see but have led workers to scrutinize the area more closely and discover Botrychium minganense.

B. Identification of Habitat Areas for Management

The viability of this species in Washington State is expected to be secure under existing ROD land use allocations, without the additional provisions of the Survey and Manage Standards and Guidelines. These management guidelines recommend protection of individual plants and their associated habitat for all populations on federal land in Oregon and California within the range of the northern spotted owl.

C. Management Within Habitat Areas

The following conservation measures have been identified as critical towards meeting the management goal for the taxon, and should be applied to any known or newly discovered populations within the analysis area in Oregon and California.

  1. Maintain current hydrologic regime. Most of the known sites have saturated soil.
  2. Maintain conditions which enhance fungal diversity. It is thought that there may be a mycorrhizal relationship between Botrychium species and western redcedar (Thuja plicata), so protecting these trees at known sites may be important (USDA Forest Service, USDI Bureau of Land Management 1994).
  3. Maintain deep shade. Most sites are found in the shade of western redcedar, in the interior of old-growth forests, so maintenance of a low light regime may be necessary.
  4. Maintain microclimatic conditions in old-growth stands, such as air temperature, soil temperature, relative humidity, and soil moisture. The modification of forest structure (e.g., canopy removal) has a profound affect on interior microclimates (Chen et al. 1993; Chen et al. 1995) and ground level vegetation (Chen et al. 1992; Frost 1992).
  5. Avoid excessive siltation or deposition of soil. Most sites have bedrock, boulders, or gravel very close to the surface (with a thick covering of moss or duff ). Excess deposition of soil or siltation could be problematic because the plants are very small and could be directly impacted (smothered).
  6. Avoid disturbance of duff layer where the species occurs. Botrychium minganense at most sites is growing in western redcedar (Thuja plicata) duff.
  7. Avoid actions that would contribute towards establishment of competing exotic vegetation, and manually remove noxious weeds. The understory of associated vascular plants at most sites is somewhat depauperate. This is probably a function of low light levels.
  8. Avoid impacts caused by livestock such as trampling or grazing.

The size of the habitat area around known populations where the management recommendations would be implemented may vary depending on environmental conditions at each particular site. The objective is to maintain the necessary microclimatic conditions around known populations of Botrychium minganense in Oregon.

Chen et al. (1995) quantified distances of edge influence within forests for several microclimatic variables and assessed the influence of edge effects in relation to aspect, time of day, microorganisms, litter, and woody debris. They described gradients from a clear-cut edge to the interior for air temperature, soil temperature, relative humidity, and soil moisture. Edge effects generally ranged from 180-240 m (540-720 ft.) into the forest for air temperature, 60-120 m (180-360 ft.) for soil temperature, >240 m (720 ft.) for relative humidity, and 0-90 m (0-270 ft.) for soil moisture. These distances provide the land manager with the approximate area needed to maintain the appropriate habitat conditions for Botrychium minganense. Consider that the research cited was derived relative to a clear-cut and that various site conditions, activities, and harvest levels will have different influences on microclimatic conditions. For example, managed areas that have a natural break in the topography such as ridge line could be smaller than those areas where the population is adjacent to disturbed locations such as roads or clear-cuts.

D. Other Management Issues and Considerations

Direct impacts, such as trampling by off-trail users, is primarily a concern from June through October when the plant is above ground. These impacts are not expected to be a concern during the winter months.

V. RESEARCH, INVENTORY, AND MONITORING NEEDS

The objective of this section is to identify opportunities for additional information which could contribute to more effective species management. The content of this section has not been prioritized or reviewed as to how important the particular items are for species management. While the inventory, research, and monitoring identified below are not required, these recommendations should be addressed by a regional coordinating body at the Northwest Forest Plan level.

A. Data Gaps and Information Needs

How can we identify high likelihood habitats for this species in order to prioritize inventory efforts or to ensure habitat conservation? This is a difficult question, aptly put by Zika et al. (1995): "There is no good explanation for why Botrychium [spp.] are common in one site and apparently absent from 20-50 seemingly similar sites. Long distance spore dispersal suggests how they can move around between disjunct sites, but it does not explain what it is that characterizes a good site".

Literature searches should be conducted to learn about any research focusing on the relationship between Botrychium minganense and Thuja plicata.

B. Research Questions

Most sites are flat benches associated with old river or stream terraces where soils are alluvial in origin. Are periodic floods part of the natural disturbance regime at these sites and, if so, what effect this disturbance has on Botrychium minganense?

Is there a correlation between the previous year's weather (e.g., dry winter/wet spring) and high or low population counts during the summer?

What is the nature of the relationship between this species and its fungal symbiont? What are the factors that may impact this mycorrhizal relationship? Are Thuja plicata trees necessary for this species to thrive within the range of the northern spotted owl?

What is the genetic basis for defining Botrychium minganense as a species? Should the taxon be split into 4 different species or otherwise reclassified based on genetic analyses?

What role do mammals play in grazing and spore dispersal? Does small mammal grazing have the same effect on Botrychium minganense as deer/elk grazing? What is the effect of livestock grazing on Botrychium minganense?

What components of an old-growth forest stand are most critical for providing habitat for Botrychium minganense? For example, is it the low light levels, the soil moisture content, the soil microorganisms, or another factor that is the key component - or is it some interaction between these variables?

C. Monitoring Needs and Recommendations

Monitoring for this species is needed over a relatively long time span (at least 5 years), not only to mitigate for year by year variation in environmental conditions but because tagged Botrychium species are known to remain below ground, and then reappear after a lapse of one to several years (Montgomery 1990; Gehring and Potash 1995; Zika et al. 1995).

VI. REFERENCES

California Natural Diversity Database. 1998. Special Plants List. California Department of Fish and Game. Sacramento, CA.

Camacho, Francisco. Personal communication. Corvallis, OR; University of Oregon; PhD candidate. 1996.

Chen, J., J. F. Franklin, and T. A. Spies. 1992. Vegetation responses to edge environments in old-growth Douglas-fir forests. Ecological Applications 2:387-396.

Chen, J., J. F. Franklin, and T. A. Spies. 1993. Contrasting microclimates among clear-cut, edge, and interior old-growth Douglas-fir forest. Agricultural and Forest Meteorology 63:219-237.

Chen, J., J. F. Franklin, and T. A. Spies. 1995. Growing season microclimatic gradients from clear-cut edges into old-growth Douglas-fir forests. Ecological Applications 5(1):74-86.

Fritschen, L. J., C. H. Driver, C. Avery, J. Buffo, R. Edmonds, R. Kinerson, P. Schiess. 1971. Dispersion of air tracers into and within a forested area: 3 Res. and Dev. Tech. Rpt. ECOM-68-G8-3. U.S. Army Elect. Command, Atmospheric Science Lab, Fort Huachuca, AZ

Frost, E. J. 1992. The effects of forest clear-cut edges on the structure and composition of old-growth mixed conifer stands in the western Klamath Mountains. M.S. Thesis, Humbolt State University, CA

Gehring, J. and L. Potash. 1995. 1994 Survey of Nooksack Botrychium. Unpublished monitoring results. Mt. Baker-Snoqualmie National Forest, Mountlake Terrace, WA

Idaho Conservation Data Center. 1994. Unpublished database records, Idaho Conservation Data Center, Idaho Dept. of Fish and Game, Boise.

Lellinger, David B. 1985. A field manual of the ferns and fern-allies of the United States and Canada. Smithsonian Institution Press. Washington, D.C.

Montgomery, J. D. 1990. Survivorship and predation changes in five populations of Botrychium dissectum in eastern Pennsylvania. American Fern Journal 80:173-182.

Morin, N. (Ed.). 1993. Flora of North America North of Mexico. Volume 2, Pteridophytes and Gymnosperms. Oxford University Press, New York.

Nugent, J. S. Personal Communication. Mt. Hood National Forest, Hood River Ranger District. Biological Technician, Plants. 1995.

Oregon Natural Heritage Program. 1995. Rare, threatened, and endangered plants and animals of Oregon. Oregon Natural Heritage report, Portland.

Rust, Steve. Personal Communication. Boise, ID; Idaho Conservation Data Center; Research Plant Ecologist. 1995.

Skinner, M. And B. Pavlik, Eds. 1994. Inventory of rare and endangered vascular plants of California, fifth edition (special publication #1). California Native Plant Society. Sacramento, CA.

Swartz, Linda. Personal Communication. Moscow, ID; University of Idaho, Forest Resources Department; Graduate Student. 1996.

USDA, Forest Service. 1988. Managing Competing and Unwanted Vegetation. Final Environmental Impact Statement. USDA, Forest Service, Portland, Oregon.

USDA, Forest Service and USDI, Bureau of Land Management. 1994. Final Supplemental Environmental Impact Statement on Management of Habitat for Late-Successional and Old-Growth Related Species Within the Range of the Northern Spotted Owl. Appendix J2, Results of Additional Species Analysis. Portland, Oregon.

USDA, Forest Service. 1991. Regional Forester's Sensitive Species List, for Region 6 (Pacific Northwest).

Washington Natural Heritage Program. 1994. Endangered, threatened, and sensitive vascular plants of Washington. Department of Natural Resources. Olympia. 52 p.

Washington Natural Heritage Program. 1997 (in draft). Endangered, threatened, and sensitive vascular plants of Washington. Department of Natural Resources.

Wagner, David H. 1992. Guide to the species of Botrychium in Oregon. USDA Forest Service report for the Mt. Hood National Forest.

Wagner, David H. 1994. Leaf morphology of the Botrychium lunaria group in Washington and Oregon. Unpublished report, Northwest Botanical Institute, Eugene, Oregon.

Victorin. 1927. Proceedings and Transcripts of the Royal Society of Canada, Series 3, 21:331.

Wagner, W. H. Jr. and F. S. Wagner. 1981. New species of moonworts, Botrychium subg. Botrychium (Ophioglossaceae), from North America. American Fern Journal: Vol. 71:1. p. 20-30.

Wagner, W. H. Jr. and F. S. Wagner. 1983. Genus communities as a systematic tool in the study of new world Botrychium (Ophioglossaceae). Taxon 32: 51-63.

Wagner, W. H. Jr. and F. S. Wagner. 1993. Ophioglossaceae. In Flora of North America Editorial Committee, eds. Flora of North America North of Mexico. Vol. 2. Oxford University Press, New York. pp. 85-106.

Wagner, D. H. 1994. Leaf morphology of the Botrychium lunaria group in Washington and Oregon. Unpublished report for Wenatchee National Forest. Northwest Botanical Institute, Eugene, Oregon.

Washington Natural Heritage Program. 1997. Endangered, threatened, and sensitive vascular plants of Washington - with working lists of rare non-vascular species. Washington State Department of Natural Resources, Olympia. 62 pp.

Williams, C. and B. Smith. 1991. Forested Plant Associations of the Wenatchee National Forest. In Draft. Wenatchee National Forest, Leavenworth, WA

Zika, P. F. 1992. Draft management guide for rare Botrychium species (moonworts and grapeferns) for the Mt. Hood National Forest. Oregon Natural Heritage Program unpublished report for the USDA Forest Service, Portland.

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