Botrychium hesperium - (Maxon & Clausen) W.H. Wagner & Lellinger
Western Moonwort
Taxonomic Status: Accepted
Related ITIS Name(s): Botrychium hesperium (Maxon & Clausen) W.H. Wagner & Lellinger (TSN 501021)
French Common Names: botryche de l'Ouest
Unique Identifier: ELEMENT_GLOBAL.2.157840
Element Code: PPOPH010Q0
Informal Taxonomy: Plants, Vascular - Ferns and relatives
 
Kingdom Phylum Class Order Family Genus
Plantae Filicinophyta Ophioglossopsida Ophioglossales Ophioglossaceae Botrychium
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Concept Reference
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Concept Reference: Kartesz, J.T. 1994. A synonymized checklist of the vascular flora of the United States, Canada, and Greenland. 2nd edition. 2 vols. Timber Press, Portland, OR.
Concept Reference Code: B94KAR01HQUS
Name Used in Concept Reference: Botrychium hesperium
Taxonomic Comments: Genus experts (Drs. Florence Wagner and Don Farrar) plan to separate the eastern plants of B. hesperium from the western (primarily Rocky Mountains) plants and call the midwestern plants B. "michiganense". This split seems to be widely accepted, although the formal publication has not yet been made. This record refers to B. hesperium in the broad sense, including B. "michiganense" plants.
Conservation Status
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NatureServe Status

Global Status: G4
Global Status Last Reviewed: 02Jun2008
Global Status Last Changed: 02Jun2008
Rounded Global Status: G4 - Apparently Secure
Reasons: Known from over 120 occurrences thus far; additional occurrences continue to be discovered in many parts of the range. Range includes much of the Rocky Mountains, from the San Francisco Peaks in Arizona through BC and Yukon to the Wrangell-St. Elias Mountains in Alaska, as well as many Pacific Northwest and upper Midwest states and adjacent Canadian provinces, east to Michigan, Ontario, and Quebec and south to South Dakota and Wyoming. Occurrences are very often small and isolated, with counts of aboveground sporophytes rarely exceeding 100; however, additional gametophytes and juvenile sporophytes belowground may add to the size of many occurrences and provide some buffer from environmental stochasticity. Appears to strongly favor open habitats within a forested matrix; threats include succession on the one hand (and suppression of natural disturbances such as fire) and human activities associated with anthropogenically-disturbed habtiats on the other (e.g. potential herbiciding of roadside populations).
Nation: United States
National Status: N3
Nation: Canada
National Status: N3N4 (03Apr2014)

U.S. & Canada State/Province Status
United States Arizona (S1), Colorado (S3), Michigan (S2), Minnesota (SNR), Montana (S3), Oregon (S1), Utah (S1), Washington (S1), Wyoming (S1)
Canada Alberta (SU), British Columbia (S3S4), Ontario (S1), Saskatchewan (S1), Yukon Territory (S1)

Other Statuses

NatureServe Global Conservation Status Factors

Range Extent Comments: Populations are generally highly disjunct, in that most known populations are scattered and often separated by many miles (Anderson and Cariveau 2004). Farrar (2005) believes that B. hesperium s.s. ranges from northern Arizona in the San Francisco Peaks northward through the Rocky Mountains (including the Blue and Wallowa Mountains of Oregon) to southern British Columbia and Alberta, Yukon Territory and southeastern Alaska in the Wrangell-St. Elias Mountains. Furthermore, he notes that B. "michiganense" has been confirmed from Michigan, Minnesota and Ontario in the Lake Superior region, the Black Hills region of South Dakota and eastern Wyoming, northwestern Montana, eastern Washington, Waterton Lakes in Alberta, the Cypress Hills in Saskatchewan, and possible additional locations in the northern Rocky Mountains not yet re-identified. Subnations reporting the presence of one or both taxa include AZ, CO, ID, MI, MN, MT, OR, UT, WA, WY, AB, BC, ON, QC, SK, and YT; Anderson and Cariveau (2004) also report that B. "michiganense" specimens are known from SD, and Farrar includes AK in the range of B. hesperium s.s. and ND, SD, MB, and WI in the range of B. "michiganense". Total range extent (without considering any one particular area to be "disjunct") is 3,000,000+ square km.

Area of Occupancy: 126-12,500 4-km2 grid cells
Area of Occupancy Comments:  

Number of Occurrences: 81 - 300
Number of Occurrences Comments: Over 120 extant occurrences are known, although this is almost certainly an underestimate as only about half of the jurisdictions believed to be part of the range (Farrar 2005) have mapped their occurrences so far, and additional occurrences continue to be discovered in many jurisdictions rangewide. At this time there are an additional 22 unranked occurrences and 12 historical occurrences known.

Population Size Comments: Often occurs as one or a few individuals scattered among other Botrychium species; at times occurs in pure stands, sometimes in large numbers. In a detailed analysis of sizes of Colorado populations, Anderson and Cariveau (2004) reported that "known populations in Colorado range in size from 1 individual to somewhere between 50 and 100 individuals, with most reports documenting between 4 and 20 plants." Still, a recent survey estimated the total count of B. hesperium in Summit County, CO alone to be 567 (Anderson and Cariveau 2004). In Oregon, 7 known occurrences are estimated to total less than 100 plants overall (smallest: 1 plant, largest: 40 plants); in Montana, many sites are poorly documented in terms of population size or are small in size, though several sites have been observed with >100 plants (S. Mincemoyer, pers. comm. 2008). Further complicating matters, the number of aboveground sporophytes observed in a given year may be a poor indicator of total occurrence size in this species, because a high proportion of an occurrence typically remains underground as gametophytes and juvenile sporophytes; thus a single emergent sporophyte may indicate the presence of a viable occurrence or a recent colonist (Anderson and Cariveau 2004). Farrar (2005) notes that "Botrychium hesperium is considered common throughout most of its range, but it may be less abundant than previously thought, due to its confusion with similar taxa, especially B. echo in the southern part of its range".

Number of Occurrences with Good Viability/Integrity: Some (13-40)
Viability/Integrity Comments: Just 15 of the 120 mapped and ranked extant occurrences are believed to have good or excellent viability. Many populations of this species are small and/or occur in uncertain habitat, where they may be threatened by such factors as succession or detrimental human activities. Many of the EOs thus far ranked as having good viability occur in western Montana, where several sites have been observed with >100 plants.

Overall Threat Impact: Medium
Overall Threat Impact Comments: Threats to B. hesperium are not well understood. Because this species occurs in both naturally and artificially disturbed sites, threats include natural plant succession as well as the same human activities (recreation, road and trail maintenance activities, selection of grazing areas) that have also apparently resulted in suitable habitat (especially when populations are small). Anderson and Cariveau (2004) point out that habitat created by anthropogenic disturbance has not yet been proven to support viable populations in the long-term; it is possible that human-created habitats may become inhospitable later due to processes such as microbial or fungal succession. Agriculture and forestry activities may also threaten this species in some areas. In Colorado, in rough order of decreasing priority, threats are listed as habitat loss, recreation, succession, overgrazing, effects of small population size, sedimentation, timber harvest, exotic species invasion, global climate change, and pollution (Anderson and Cariveau 2004). These authors suggest that minimizing soil disturbance may be important to the species; they note that off-road vehicle use (both motorized and non-motorized) represents a significant threat, and that the use of livestock grazing to enhance habitat is risky.Threats in the Wallowa Mountains of Oregon include fire suppression, pack animal grazing, wood-cutting, and recreation-associated activities (Anderson and Cariveau 2004). In Montana, many populations occur on roadsides or other similarly open or disturbed habitats, leaving them vulnerable to activities such as weed invasion, weed spraying and road maintenance (S. Mincemoyer, pers. comm. 2008).

Short-term Trend: Decline of <30% to increase of 25%
Short-term Trend Comments: Essentially unknown, as high variation in number of emergent stalks among years makes detection of overall trends very difficult (Anderson and Cariveau 2004). Abundance of B. hesperium has been observed to have increased in the vicinity of ski resorts, but the long-term suitability of these habitats is unknown (Anderson and Cariveau 2004). It is possible that fire suppression has decreased availability of suitable habitat, but no data are available to support or refute this idea.There have been no known cases since this species was recognized in which an occurrence was extirpated due to human activities (Anderson and Cariveau 2004).

Long-term Trend: Decline of <30% to increase of 25%
Long-term Trend Comments: In post-settlement times, Kolb and Spribille (2001) (cited in Anderson and Cariveau 2004) hypothesize that the abundance of Botrychium has increased due to increased anthropogenic disturbance (associated with ski runs, roads, clear cuts, trails, mine sites, etc.) However, Botrychium habitat may also have decreased due to fire suppression and grazing of western grasslands and meadows. Unfortuantely, little data are available to address either of these ideas.

Intrinsic Vulnerability Comments: This species' tendency to grow in small, somewhat isolated populations with highly variable numbers of individuals may make populations susceptible to local extirpation; however, many populations may be buffered by the presence of underground gametophytes and juvenile sporophytes (Anderson and Cariveau 2004). This species is also very difficult to propagate, which probably precludes restoration efforts (Anderson and Cariveau 2004).

Environmental Specificity Comments: Depends on disturbance to maintain suitable habitat; current sites are in most cases destined to become unsuitable as a result of natural succession (Anderson and Cariveau 2004). In addition, Botrychium species rely upon mycorrhizae in both the sporophytic and gametophytic stages. The ubiquity and low host specificity of AM fungi suggest that mycorrhizae may not be a limiting factor in the distribution of B. hesperium, but changes in the mycoflora during succession may affect habitat quality (Anderson and Cariveau 2004).

Other NatureServe Conservation Status Information

Distribution
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Global Range: Populations are generally highly disjunct, in that most known populations are scattered and often separated by many miles (Anderson and Cariveau 2004). Farrar (2005) believes that B. hesperium s.s. ranges from northern Arizona in the San Francisco Peaks northward through the Rocky Mountains (including the Blue and Wallowa Mountains of Oregon) to southern British Columbia and Alberta, Yukon Territory and southeastern Alaska in the Wrangell-St. Elias Mountains. Furthermore, he notes that B. "michiganense" has been confirmed from Michigan, Minnesota and Ontario in the Lake Superior region, the Black Hills region of South Dakota and eastern Wyoming, northwestern Montana, eastern Washington, Waterton Lakes in Alberta, the Cypress Hills in Saskatchewan, and possible additional locations in the northern Rocky Mountains not yet re-identified. Subnations reporting the presence of one or both taxa include AZ, CO, ID, MI, MN, MT, OR, UT, WA, WY, AB, BC, ON, QC, SK, and YT; Anderson and Cariveau (2004) also report that B. "michiganense" specimens are known from SD, and Farrar includes AK in the range of B. hesperium s.s. and ND, SD, MB, and WI in the range of B. "michiganense". Total range extent (without considering any one particular area to be "disjunct") is 3,000,000+ square km.

U.S. States and Canadian Provinces
Color legend for Distribution Map

U.S. & Canada State/Province Distribution
United States AZ, CO, MI, MN, MT, OR, UT, WA, WY
Canada AB, BC, ON, SK, YT

Range Map
No map available.


U.S. Distribution by County Help
State County Name (FIPS Code)
CO Archuleta (08007), Boulder (08013), Clear Creek (08019), Conejos (08021), Eagle (08037), Gilpin (08047), Grand (08049), Huerfano (08055), Jackson (08057), Lake (08065), Larimer (08069), Mineral (08079), Pitkin (08097), Rio Grande (08105), Saguache (08109), San Juan (08111), Summit (08117), Teller (08119)
MI Alger (26003), Alpena (26007)*, Chippewa (26033)
OR Umatilla (41059), Union (41061), Wallowa (41063)
WA Chelan (53007), Ferry (53019), Pend Oreille (53051), Snohomish (53061), Stevens (53065)
* Extirpated/possibly extirpated
U.S. Distribution by Watershed Help
Watershed Region Help Watershed Name (Watershed Code)
04 Betsy-Chocolay (04020201)+, Tahquamenon (04020202)+, Carp-Pine (04070002)+, Thunder Bay (04070006)+*
10 North Platte Headwaters (10180001)+, Upper Laramie (10180010)+, Upper South Platte (10190002)+, Clear (10190004)+, St. Vrain (10190005)+, Big Thompson (10190006)+*, Cache La Poudre (10190007)+
11 Arkansas Headwaters (11020001)+, Fountain (11020003)+, Huerfano (11020006)+
13 Rio Grande headwaters (13010001)+, Alamosa-Trinchera (13010002)+*, Saguache (13010004)+, Conejos (13010005)+
14 Colorado headwaters (14010001)+, Blue (14010002)+, Eagle (14010003)+, Roaring Fork (14010004)+, Upper San Juan (14080101)+, Animas (14080104)+
17 Pend Oreille (17010216)+, Franklin D. Roosevelt Lake (17020001)+, Kettle (17020002)+, Colville (17020003)+, Sanpoil (17020004)+, Lake Chelan (17020009)+, Upper Grande Ronde (17060104)+, Wallowa (17060105)+, Sauk (17110006)+
+ Natural heritage record(s) exist for this watershed
* Extirpated/possibly extirpated
Ecology & Life History
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Basic Description: A small, erect, perennial fern. B. hesperium is similar to other Botrychium spp. but is a duller green color than B. echo, and has ovate to oblong lower pinnae. Leaves appear in midspring and die in early fall. Spores are produced in July.

General Description: Western Moonwort is a small perennial fern with a single erect frond, 3-13 cm high. It is divided into a sterile segment and a fertile segment. The sterile segment has a stalk 0-4 mm long, and a broadly lance-shaped to triangular blade that is pinnately divided with 1-6 pairs of closely adjacent leaflets (pinnae). The basal pinnae are usually partly to wholly pinnately divided and are larger than the lobed or entire-margined upper pinnae. The fertile segment is 2-3 times as long as the sterile segment and 1-3 times pinnately divided into linear segments that bear the spores.
Diagnostic Characteristics: Botrychium subgenus Botrychium is a large group of very similar species, many of which have been recently described. A technical manual should be consulted for positive identification. The deeply pinnately lobed basal pinnae help separate this species from most others in its range. Farrar (2005) provides the following characteristics differentiating B. "michganense" from B. hesperium s.s.: B. "michiganense" differs in its usually unstalked to short-stalked (< 2mm) trophophore and its abrupt transition from the elongated and deeply dissected basal pinnae to the distinctly smaller and scarcely dissected second pinnae pair. Where the two species co-occur (northern Rocky Mountains), B. hesperium var. fenestratum further differs from B. "michiganense" in having pinnae and pinnules overlapping or nearly so. The typical variety of B. hesperium in the southern Rocky Mountains has little to no dissection in pinnae above the first pair and only shallow and narrow dissection in its basal pinnae; however, the typical variety of B. hesperium seldom co-occurs with B. "michiganense".
Palustrine Habitat(s): HERBACEOUS WETLAND, Riparian
Terrestrial Habitat(s): Forest - Conifer, Forest - Mixed, Forest/Woodland, Grassland/herbaceous, Sand/dune, Woodland - Conifer, Woodland - Mixed
Habitat Comments: In the western portion of its range (i.e. excluding "michiganense" material), occurs in the forested montane zone, where the forest is relatively open-canopied and/or within open habitat types, which tend to be subject to periodic disturbance and include subalpine meadows, snow fields, mesic grassy slopes, prairie pothole meadows, edges of lakes, gravel bars, and roadsides. Soils are dry to moist and tend to be coarse and gravelly. Common moonwort associates include B. paradoxum, B. lunaria and B. lanceolatum. Anderson and Cariveau (2004) note that apparently suitable habitat is plentiful within the range but is often not occupied by this species; they hypothesize that this may be due to limitations in successful migration to the site, or the result of other unknown ecological parameters, such as insufficient time since a disturbance event and/or lack of appropriate mycorrhizal symbionts. 1000 - 3500 m. In the eastern portion of its range ("michiganense" material), occurs in sand dune habitats, in moist shrubby jack pine forest in dune valleys, in grassy roadsides and fields, and in open-canopied mesic northern forests and woodlands, sometimes dominated by sugar maple, yellow birch, and hemlock. Occurs as low as 200 m.
Economic Attributes Not yet assessed
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Management Summary
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Stewardship Overview: Requires at least somewhat open habitat, in the past most likely created by natural disturbance events such as fire. Anthropogenically-disturbed areas such as ski runs have been observed to harbor reasonably large numbers of individuals, but habitat created by anthropogenic disturbance has not yet been proven to support viable populations in the long-term (Anderson and Cariveau 2004). Mycorrhizal interactions are also of paramount importance to Botrychium species, and better understanding of this species' mycorrhizal relationships and requirements will lead to better understanding of management techniques that support those relationships (Anderson and Cariveau 2004).
Population/Occurrence Delineation Not yet assessed
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Population/Occurrence Viability
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U.S. Invasive Species Impact Rank (I-Rank) Not yet assessed
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Authors/Contributors
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NatureServe Conservation Status Factors Edition Date: 13Apr1992
NatureServe Conservation Status Factors Author: S. Gottlieb & C. Russell (1992), rev. K. Crowley & M. Penskar (1995), rev. K. Gravuer (2008)
Management Information Edition Date: 02Jun2008
Management Information Edition Author: Gravuer, K.
Element Ecology & Life History Edition Date: 06Oct1994
Element Ecology & Life History Author(s): JM

Botanical data developed by NatureServe and its network of natural heritage programs (see Local Programs), The North Carolina Botanical Garden, and other contributors and cooperators (see Sources).

References
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  • Ahlenslager, K. and P. Lesica. 1995. Observations of BOTRYCHIUM WATERTONENSE and its putative parent species, B. HESPERIUM and B. PARADOXUM. Draft manuscript prepared in cooperation with Waterton Lakes National Park, USFWS, and Montana Natural Heritage Program. 13 pp.

  • Ahlenslager, K. and P. Lesica. 1995. Observations of BOTRYCHIUM WATERTONENSE and its putative parent species, B. HESPERIUM and B. PARADOXUM. Draft manuscript prepared in cooperation with Waterton Lakes National Park, USFWS, and Montana Natural Heritage Program. 13 pp.

  • Anderson, D. G. and D. Cariveau 2004. Botrychium hesperium Barneby (western moonwort): A technical conservation assessment. USDA Forest Service, Rocky Mountain Region. Online. Available: http://www.fs.fed.us/r2/projects/scp/assessments/botrychiumhesperium.pdf (Accessed 2008).

  • Argus, G.W., K.M. Pryer, D.J. White and C.J. Keddy (eds.). 1982-1987. Atlas of the Rare Vascular Plants of Ontario.. Botany Division, National Museum of National Sciences, Ottawa.

  • Chadde, S. and G. Kudray. 2001. Conservation assessment for Western Moonwort (Botrychium hesperium). USDA Forest Service, Eastern Region. 33 pp.

  • Douglas, G.W., D. Meidinger, and J. Penny. 2002. Rare Native Vascular Plants of British Columbia, 2nd ed. B.C. Conserv. Data Centre, Terrestrial Inf. Branch, Victoria. 358pp.

  • Douglas, G.W., D. Meidinger, and J. Pojar, editors. 2000. The Illustrated Flora of British Columbia. Volume 5. Dicotyledons (Salicaceae through Zygophyllaceae) and Pteridophytes. British Columbia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests, Victoria.

  • Douglas, G.W., D. Meidinger, and J. Pojar, eds. 2000. Illustrated Flora of British Columbia, Vol. 5, Dicotyledons (Salicaceae through Zygophyllaceae) and Pteridophytes. B.C. Minist. Environ., Lands and Parks, and B.C. Minist. For., Victoria. 389pp.

  • Douglas, G.W., G.B. Straley, and D. Meidinger, eds. 1998. Rare Native Vascular Plants of British Columbia. Conserv. Data Centre, Resour. Inventory Branch, B.C. Minist. Environ., Lands and Parks, Victoria, and B.C. Minist. For., Victoria.

  • Farrar, D. R. 2005b, January last update. Botrychium hesperium species description, map, and photo page. In Farrar, D.R. 2006, June last update. Systematics of moonworts Botrychium subgenus Botrychium. Department of Ecology, Evolution and Organismal Biology, Iowa State University, Ames. Online. Available: http://www.public.iastate.edu/~herbarium/botrychium.html (Accessed 2008)

  • Farrar, D. R. 2005c, January last update. Botrychium michiganense species description, map, and photo page. In Farrar, D.R. 2006, June last update. Systematics of moonworts Botrychium subgenus Botrychium. Department of Ecology, Evolution and Organismal Biology, Iowa State University, Ames. Online. Available: http://www.public.iastate.edu/~herbarium/botrychium.html (Accessed 2008)

  • Farrar, D. R. and S. J. Popovich. 2012. Ophioglossaceae. Pages 24-35 Colorado Flora: Eastern Slope, fourth edition. W.A. Weber and R.C. Wittmann. University Press of Colorado, Boulder, CO.

  • Fertig, W. 1995. More new plant species for Wyoming. Castilleja 14:4-5.

  • Flora of North America Editorial Committee. 1993. Flora of North America North of Mexico. Vol. 2 Pteridophytes and Gymnosperms. Oxford Univ. Press, New York.

  • Flora of North America Editorial Committee. 1993a. Flora of North America north of Mexico. Vol. 2. Pteridophytes and gymnosperms. Oxford Univ. Press, New York. xvi + 475 pp.

  • Harms, V.L., P.A. Ryan and J.A. Haraldson. 1992. The rare and endangered vascular plants of Saskatchewan. Prepared for the Saskatchewan Natural History Society. Unpubl.

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  • Kartesz, J.T. 1994. A synonymized checklist of the vascular flora of the United States, Canada, and Greenland. 2nd edition. 2 vols. Timber Press, Portland, OR.

  • Lellinger, D. B. 1981. Notes on North American ferns. American Fern Journal 71:90-94.

  • Lesica, P. and K. Ahlenslager. 1994. Demographic monitoring of three species of BOTRYCHIUM (Ophioglossaceae) in Waterton Lakes Park, Alberta: 1993 progress report. Unpublished report to the U.S. Fish and Wildlife Service. Montana Natural Heritage Program, Helena, Montana. 19 pp.

  • Lesica, P. and K. Ahlenslager. 1994. Demographic monitoring of three species of BOTRYCHIUM (Ophioglossaceae) in Waterton Lakes Park, Alberta: 1993 progress report. Unpublished report to the U.S. Fish and Wildlife Service. Montana Natural Heritage Program, Helena, Montana. 19 pp.

  • Lesica, P. and K. Ahlenslager. 1995. Demography and life history of three sympatric species of BOTRYCHIUM subg. BOTRYCHIUM in Waterton Lakes National Park, Alberta, Canada. Draft manuscript prepared in cooperation with Waterton National Park, U.S. Fish and Wildlife Service, and Montana Natural Heritage Program. 22 pp.

  • Lesica, P. and K. Ahlenslager. 1995. Demography and life history of three sympatric species of Botrychium subq. Botrychium in Waterton Lakes National Prak, Alberta, Canada. Herbarium, Division of Biological Sciences, University of Montana, Missoula, MT. In cooperation with Waterton Lakes National Parks, USF&W Service, and MIHP.

  • Mantas, M. and R. S. Wirt. 1995. Moonworts of western Montana (BOTRYCHIUM subgenus BOTRYCHIUM). Flathead National Forest. 103 pp.

  • Mantas, M. and R. S. Wirt. 1995. Moonworts of western Montana (BOTRYCHIUM subgenus BOTRYCHIUM). Flathead National Forest. 103 pp.

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  • Tera Environmental Consultants. 2013. EXCEL spreadsheet of various rare plant observations.

  • Wagner Jr., W. H. and F. S. Wagner. 1983. Two moonworts of the Rocky Mountains; Botrychium hesperium and a new species formerly confused with it. American Fern Journal 73:53-62.

  • Wagner, Jr., W. H. and F. S. Wagner. 1981. New species of moonworts, BOTRYCHIUM subg. BOTRYCHIUM (Ophioglossaceae), from North America. Amer. Fern J. 71:20-30.

  • Wagner, Jr., W. H., F. S. Wagner, C. Haufler and J. K. Emerson. 1984. A new nothospecies of moonwort (Ophioglossaceae, BOTRYCHIUM). Canadian Journal of Botany 62:629-634.

  • Wagner, Jr., W. H., F. S. Wagner, C. Haufler and J. K. Emerson. 1984. A new nothospecies of moonwort (Ophioglossaceae, BOTRYCHIUM). Canadian Journal of Botany 62:629-634.

  • Wagner, W. and F. Wagner. 1986. Three new species of moonworts (BOTRYCHIUM subgenus BOTRYCHIUM) endemic in western North America. American Fern Journal 76(2):3347.

  • Wagner, W.H. Jr. and F.S. Wagner. 1986. Three new species of moonworts (Botrychium subg. Botrychium) endemic in western North America. American Fern Journal 76(2):33-47.

  • Wagner, W.H., Jr., and F. Wagner. 1981. New species of moonworts, Botrychium subg. Botrychium (Ophioglossaceae), from North America. American Fern J. 71(1):20, 26.

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  • Wagner, W.H., and F.S. Wagner. 1990. Moonworts (Botrychium subg. Botrychium) of the upper Great Lakes region, USA and Canada, with descriptions of two new species. Contr. Univ. Mich. Herb. 17:313-325

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