Botrychium simplex E. Hitchcock (little grapefern) - Colorado Natural ...

Botrychium simplex E. Hitchcock 

(little grapefern) 

A Technical Conservation Assessment 

Prepared for the USDA Forest Service, 

Rocky Mountain Region, 

Species Conservation Project 

May 22, 2006 

David G. Anderson 

Colorado Natural Heritage Program 

Colorado State University 

Fort Collins, CO 

Peer Review Administered by 

Center for Plant Conservation

Anderson, D.G. (2006, May 22). Botrychium simplex E. Hitchcock (little grapefern): a technical conservation 

assessment. [Online]. USDA Forest Service, Rocky Mountain Region. Available: http://www.fs.fed.us/r2/ 

projects/scp/assessments/botrychiumsimplex.pdf [date of access]. 

ACKNOWLEDGMENTS 

The helpfulness and generosity of many experts, particularly Beth Burkhart, Reed Crook, Don Farrar, Cindy 

Johnson-Groh, Annette Miller, Peter Root, Dave Steinmann, Florence Wagner, Jennifer Whipple, and Jennifer 

Winther, are gratefully acknowledged. Their interest in the project, valuable insight, depth of experience, and time 

spent answering questions were extremely valuable and crucial to the project. Herbarium specimen label data were 

provided by Margaret Bolick (NEB); Aleisha Cordell (SJNM); Nan Lederer (COLO); Ron Hartman, Ernie Nelson, 

and Joy Handley (RM); Deborah Lewis (ISC); Steve Rolfsmeier (High Plains Herbarium); Sylvia Kelso (COCO); 

and Catherine Kleier. Jason McNees at NatureServe assisted with heritage data acquisition. Thanks also to Janet 

Coles, Greg Hayward, Greg Karow, Gary Patton, Jim Maxwell, Andy Kratz, Beth Burkhart, Steve Popovich, John 

Proctor, and Joy Bartlett for assisting with questions and project management. Beth Burkhart provided photographs 

by Katherine Zacharkevics and others with the Black Hills National Forest for this assessment. Mary Olivas, Jane 

Nusbaum, Carmen Morales, and Barbara Brayfield provided crucial financial oversight. Shannon Gilpin and Ryan 

Neeper assisted with literature acquisition. Thanks also to my family (Jen, Cleome, and Melia) for their support during 

the synthesis of this document, and to my mother for contributing the Botrychium simplex haiku. 

AUTHOR’S BIOGRAPHY 

David G. Anderson is a botanist with the Colorado Natural Heritage Program (CNHP). Mr. Anderson’s work at 

CNHP includes inventory and mapping of rare plants throughout Colorado, mapping weeds, maintaining and updating 

CNHP’s database, and writing reports on the rare plants of Colorado. He has worked with CNHP since 1999. Much of 

Mr. Anderson’s prior experience comes from five years of fieldwork studying the flora and ecosystem processes of the 

Alaskan and Canadian Arctic. Mr. Anderson also served in the Peace Corps as a science teacher in the Solomon Islands 

from 1996 to 1998. Mr. Anderson received his B.A. in Environmental, Populational, and Organismic Biology from the 

University of Colorado, Boulder (1991) and his M.S. in Botany from the University of Washington, Seattle (1996). 

COVER PHOTO CREDIT 

Botrychium simplex (little grapefern). (Left) photo of the first illustration of B. simplex, published with the 

description of the species (Hitchcock 1823). Photo by the author. (Right) photo by Katherine Zacharkevics, Black 

Hills National Forest Botanist, North Hills District of a plant collected at Dugout Gulch, Wyoming in 2005. 

One reluctant leaf, 

A moonwort sprouts in the fen 

Risking its secrets 

—Jean Anderson 

2

SUMMARY OF KEY COMPONENTS FOR CONSERVATION OF 

BOTRYCHIUM SIMPLEX 

Status 

Botrychium simplex E. Hitchcock (little grapefern) is known from 50 locations in Region 2, 17 of which have 

not been seen within the last 20 years. The population size in Region 2 is unknown, but the estimated total population 

from locations where plant counts have been made is 500 to 600 plants. Other occurrences are known to support 

significant populations of this species, but these have not been counted. Botrychium simplex is not designated as a 

sensitive species in USDA Forest Service (USFS) Region 2, but it is considered a sensitive species in USFS Region 

1 and in the Washington portion of USFS Region 6, and it is considered important for biodiversity analysis in USFS 

Region 4. NatureServe ranks B. simplex as globally secure (G5). Within Region 2, it is ranked imperiled (S2) in 

Colorado and Wyoming, and unrankable (SU) in South Dakota. It has no rank in Nebraska, but it probably warrants a 

rank of critically imperiled (S1) based on the one known occurrence in that state. Botrychium simplex is not listed as 

threatened or endangered under the Federal Endangered Species Act. 

Primary Threats 

Observations and quantitative data suggest several threats to the persistence of Botrychium simplex. The primary 

threats are ski area development and maintenance, road construction and maintenance, timber harvest, recreation, 

fire, grazing, effects of small population size, woody plant encroachment, exotic species invasion, succession, global 

climate change, and pollution. 

Primary Conservation Elements, Management Implications and Considerations 

The responsibility of maintaining viable populations of Botrychium simplex within Region 2 falls largely 

on the USFS because most occurrences and suitable habitat are on National Forest System land. Forty-one of the 

50 known occurrences of this species in Region 2 are on National Forest System land, and 24 of these have been 

observed since 1999. Eight occurrences are known from national parks in the states of Region 2: two occurrences are 

in Rocky Mountain National Park within Region 2 and six are in Yellowstone National Park outside Region 2. Two 

additional occurrences are under unknown management, and two are on public lands managed by the Bureau of Land 

Management. The City of Denver, State of Colorado, and The Nature Conservancy also each have one occurrence. 

Seventeen occurrences have not been seen in more than 20 years, and it is important to determine if B. simplex is 

extant at these locations. Additional inventories are needed to better understand the full range and distribution of 

this species. 

Restoring populations of Botrychium simplex is probably precluded by the difficulty in propagating this 

species. Research is needed to investigate the underground life history, ecology, reproductive biology, the role of 

mycorrhizae, and the role of disturbance in the autecology of B. simplex so that conservation efforts on its behalf 

can be most effective. 

The major conservation element essential to ensuring viable populations of Botrychium simplex in Region 2 

is promoting the processes that create and maintain the early- to mid-seral or other suitable habitats required by B. 

simplex. Unfortunately, these processes are poorly understood. Because new data are just now becoming available and 

our current knowledge of B. simplex in Region 2 is incomplete, it is difficult to formulate conservation strategies at 

present. More complete knowledge of the species’ distribution will permit the identification of areas most suitable for 

conservation management in Region 2. New surveys are needed to better understand how the subspecies of B. simplex 

differ in habitat affinities and autecology in Region 2. Demographic studies designed to determine the impacts of 

grazing, succession, fire, and exotic species on population viability are also high priorities for research on B. simplex 

in Region 2. 

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TABLE OF CONTENTS 

ACKNOWLEDGMENTS ..............................................................................................................................................2 

AUTHOR’S BIOGRAPHY............................................................................................................................................2 

COVER PHOTO CREDIT .............................................................................................................................................2 

SUMMARY OF KEY COMPONENTS FOR CONSERVATION OF BOTRYCHIUM SIMPLEX ...............................3 

Status..........................................................................................................................................................................3 

Primary Threats..........................................................................................................................................................3 

Primary Conservation Elements, Management Implications and Considerations.....................................................3 

LIST OF TABLES AND FIGURES ...............................................................................................................................6 

INTRODUCTION ..........................................................................................................................................................7 

Goal of Assessment....................................................................................................................................................7 

Scope of Assessment..................................................................................................................................................7 

Treatment of Uncertainty in Assessment ...................................................................................................................7 

Treatment of This Document as a Web Publication...................................................................................................8 

Peer Review of This Document .................................................................................................................................8 

MANAGEMENT STATUS AND NATURAL HISTORY .............................................................................................8 

Management Status ....................................................................................................................................................8 

Existing Regulatory Mechanisms, Management Plans, and Conservation Strategies...............................................8 

Adequacy of current laws and regulations ............................................................................................................8 

Adequacy of current enforcement of laws and regulations...................................................................................9 

Biology and Ecology..................................................................................................................................................9 

Classification and description................................................................................................................................9 

Taxonomic status ..............................................................................................................................................9 

Description .....................................................................................................................................................13 

Sources for keys photographs, illustrations, and descriptions........................................................................16 

Distribution and abundance.................................................................................................................................19 

Population trend ..................................................................................................................................................34 

Habitat .................................................................................................................................................................35 

Habitat descriptions and characterization.......................................................................................................35 

Region 2 habitat descriptions .........................................................................................................................35 

Varietal differences in habitat.........................................................................................................................36 

Elevation, slope, and aspect............................................................................................................................37 

Soil..................................................................................................................................................................37 

Moisture..........................................................................................................................................................37 

Disturbance as a habitat attribute ...................................................................................................................38 

Fire..................................................................................................................................................................38 

Meadows.........................................................................................................................................................38 

Mycorrhizae as a habitat attribute ..................................................................................................................39 

Reproductive biology and autecology.................................................................................................................39 

Reproduction ..................................................................................................................................................39 

Phenology.......................................................................................................................................................40 

Fertility ...........................................................................................................................................................40 

Dispersal.........................................................................................................................................................41 

Cryptic phases ................................................................................................................................................41 

Mycorrhizae....................................................................................................................................................41 

Hybridization..................................................................................................................................................42 

Demography ........................................................................................................................................................43 

Community ecology ............................................................................................................................................45 

CONSERVATION.........................................................................................................................................................50 

Threats......................................................................................................................................................................50 

Influence of management activities and natural disturbances on individuals and habitat ..................................51 

Ski area development and maintenance .........................................................................................................51 

Road and trail construction and maintenance.................................................................................................51 

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Timber harvest................................................................................................................................................51 

Recreation.......................................................................................................................................................52 

Fire..................................................................................................................................................................52 

Grazing ...........................................................................................................................................................52 

Effects of small population size......................................................................................................................52 

Woody plant encroachment ............................................................................................................................53 

Exotic species .................................................................................................................................................53 

Global climate change ....................................................................................................................................53 

Pollution .........................................................................................................................................................54 

Over-utilization...............................................................................................................................................54 

Conservation Status of Botrychium simplex in Region 2.........................................................................................54 

Is distribution or abundance declining in all or part of its range in Region 2? ...................................................54 

Do habitats vary in their capacity to support this species? .................................................................................54 

Vulnerability due to life history and ecology ......................................................................................................55 

Evidence of populations in Region 2 at risk .......................................................................................................55 

Management of Botrychium simplex in Region 2....................................................................................................55 

Implications and potential conservation elements ..............................................................................................55 

Tools and practices ..............................................................................................................................................56 

Species inventory............................................................................................................................................56 

Habitat inventory.................................................................................................................................................57 

Population monitoring.........................................................................................................................................58 

Beneficial management actions...........................................................................................................................59 

Restoration ..........................................................................................................................................................60 

Information Needs and Research Priorities .............................................................................................................60 

Additional research and data resources...............................................................................................................62 

DEFINITIONS..............................................................................................................................................................63 

REFERENCES .............................................................................................................................................................65 

EDITORS: Janet Coles, Beth Burkhart, and Kathy Roche, USDA Forest Service, Rocky Mountain Region 

5

Tables: 

LIST OF TABLES AND FIGURES 

Table 1. Classification of Botrychium simplex after USDA Natural Resources Conservation Service........... 9 

Table 2. A partial list of synonyms for Botrychium simplex. ......................................................................... 11 

Table 3. Summary information for the varieties of Botrychium simplex....................................................... 12 

Table 4. Genetic variability in eastern diploid species of Botrychium subgenus Botrychium....................... 12 

Table 5. Hypothetical parents for polyploid species of Botrychium subgenus Botrychium. ......................... 13 

Table 6. Diagnostic characteristics for the determination of the “eastern” and “western” types of 

Botrychium simplex........................................................................................................................................ 15 

Table 7. Known distribution of Botrychium simplex in the western hemisphere. ......................................... 21 

Table 8. Land ownership / management status summary for the 50 known occurrences of Botrychium 

simplex within the administrative boundary of USDA Forest Service Region 2........................................... 23 

Table 9. Land ownership / management status summary for the 50 known occurrences of Botrychium 

simplex within the administrative boundary of USDA Forest Service Region 2. ......................................... 32 

Table 10. Wetland Indicator Status for Botrychium simplex.......................................................................... 38 

Table 11. Associated species reported with Botrychium simplex in USDA Forest Service Region 2. .......... 46 

Figures: 

Figure 1. The first illustration of Botrychium simplex. .................................................................................. 10 

Figure 2. Botrychium simplex sporophytes at Dugout Gulch, Wyoming, Black Hills National Forest after 

emergence, and before separation of the trophophore and sporophore. ........................................................ 15 

Figure 3. Botrychium simplex var. compositum (“western” simplex) collected on Boreas Pass, Colorado... 16 

Figure 4. Botrychium simplex var. tenebrosum from Park County, Wyoming. ............................................. 17 

Figure 5. Botrychium simplex var. simplex (“eastern” simplex) from East Inlet, Rocky Mountain National 

Park, Colorado. .............................................................................................................................................. 17 

Figure 6. Illustration of Botrychium simplex. ................................................................................................ 18 

Figure 7. Illustration of Botrychium tenebrosum........................................................................................... 19 

Figure 8. Global distribution map of Botrychium simplex............................................................................. 20 

Figure 9. Distribution of Botrychium simplex in North America, color-coded by S rank. ............................ 21 

Figure 10. The distribution of Botrychium simplex in the states of USDA Forest Service Region 2............ 33 

Figure 11. Habitat of Botrychium simplex at Redbank Spring, South Dakota on the Black Hills National 

Forest.............................................................................................................................................................. 36 

Figure 12. Lifecycle diagram for Botrychium simplex, illustrating the alternation of generations. .............. 44 

Figure 13. Hypothetical life cycle graph for Botrychium simplex................................................................. 45 

6

INTRODUCTION 

This assessment is one of many being produced 

to support the Species Conservation Project for the 

Rocky Mountain Region (Region 2), USDA Forest 

Service (USFS). Botrychium simplex is the focus 

of an assessment because of its rarity, degree of 

imperilment, and concern for its viability in Region 2. 

It is not currently listed as sensitive by Region 2 or by 

the Bureau of Land Management (BLM) in Colorado or 

Wyoming. Botrychium species have been the focus of 

increasing interest by the USFS and other federal and 

state agencies due to their rarity, difficulty in detection, 

and highly variable populations (Johnson-Groh and 

Farrar 2003). 

This assessment addresses the biology, ecology, 

conservation status, and management of Botrychium 

simplex throughout its range in Region 2. The broad 

nature of the assessment leads to some constraints on 

the specificity of information for particular locales. 

This introduction defines the goal of the assessment, 

outlines its scope, and describes the process used in 

its production. 

Goal of Assessment 

Species conservation assessments produced as 

part of the Species Conservation Project are designed 

to provide forest managers, research biologists, and 

the public with a thorough discussion of the biology, 

ecology, conservation status, and management of 

certain species based on available scientific knowledge. 

The assessment goals limit the scope of the work to 

critical summaries of scientific knowledge, discussion 

of broad implications of that knowledge, and outlines 

of information needs. The assessment does not seek 

to develop specific management recommendations. 

Rather, it provides the ecological backgrounds upon 

which management must be based and focuses on the 

consequences of changes in the environment that result 

from management (i.e., management implications). 

Furthermore, it cites management recommendations 

proposed elsewhere and examines the success of those 

recommendations that have been implemented. 

Scope of Assessment 

This assessment examines the biology, ecology, 

conservation status, and management of Botrychium 

simplex with specific reference to the geographic and 

ecological characteristics of Region 2. Although some 

of the literature on the species may originate from field 

investigations outside the region, this document places 

7 

that literature in the ecological and social context of the 

central Rocky Mountains. Similarly, this assessment 

is concerned with reproductive behavior, population 

dynamics, and other characteristics of B. simplex in the 

context of the current environment rather than under 

historical conditions. The evolutionary environment of 

the species is considered in conducting the synthesis, 

but placed in a current context. 

In producing the assessment, I reviewed refereed 

literature, non-refereed publications, research reports, 

and data accumulated by resource management 

agencies and other investigators. Because basic research 

has not been conducted on many facets of the biology 

of Botrychium simplex, literature on its congeners was 

used to make inferences in many cases. The refereed 

and non-refereed literature on the genus Botrychium 

and its included species is more extensive and includes 

other endemic or rare species. All known publications 

on B. simplex are referenced in this assessment, and 

many of the experts on this species were consulted 

during its synthesis. All available specimens of B. 

simplex in Region 2 were viewed to verify occurrences 

and to incorporate specimen label data. Specimens 

were searched for at COLO (University of Colorado 

Herbarium), CS (CSU Herbarium), RM (Rocky 

Mountain Herbarium), KHD (Kalmbach Herbarium, 

Denver Botanic Gardens), SJNM (San Juan College 

Herbarium), CC (Carter Herbarium), GREE (University 

of Northern Colorado Herbarium), NMCR (New Mexico 

State University Range Science Herbarium), and UNM 

(University of New Mexico Herbarium). The assessment 

emphasizes peer-reviewed literature because this is the 

accepted standard in science. Non-refereed publications 

or reports were regarded with greater skepticism, but 

they were used in the assessment because there is 

very little refereed literature that specifically treats B. 

simplex in Region 2. Unpublished data (e.g., Natural 

Heritage Program records, reports to state and federal 

agencies, specimen labels) were important in estimating 

the geographic distribution of this species. These data 

required special attention because of the diversity of 

persons and methods used in collection. 

Treatment of Uncertainty in 

Assessment 

Science is a rigorous, systematic approach to 

obtaining knowledge. Competing ideas regarding how 

the world works are measured against observations. 

Because our descriptions of the world are always 

incomplete and our observations are limited, science 

focuses on approaches for dealing with uncertainty. 

A commonly accepted approach to science is based

on a progression of critical experiments to develop 

strong inference (Platt 1964). It is difficult to conduct 

experiments that produce clean results in the ecological 

sciences. Often, observations, inference, good thinking, 

and models must be relied on to guide our understanding 

of ecological relations. Confronting uncertainty then is 

not prescriptive. In this assessment, the strength of 

evidence for particular ideas is noted, and alternative 

explanations are described when appropriate. 

Treatment of This Document as a Web 

Publication 

To facilitate use of species assessments in the 

Species Conservation Project, assessments are being 

published on the Region 2 World Wide Web site. Placing 

the documents on the Web makes them available to 

agency biologists and the public more rapidly than 

publishing them as reports. More important, Web 

publication will facilitate revision of the assessments, 

which will be accomplished based on guidelines 

established by Region 2. 

Peer Review of This Document 

Assessments developed for the Species 

Conservation Project have been peer reviewed before 

their release on the Web. This assessment was reviewed 

through a process administered by the Center for Plant 

Conservation, employing two recognized experts 

on this or related taxa. Peer review was designed to 

improve the quality of communication and to increase 

the rigor of the assessment. 

MANAGEMENT STATUS AND 

NATURAL HISTORY 

Management Status 

Botrychium simplex is not currently listed as a 

sensitive species in Region 2 (USDA Forest Service 

2003). Its merits as a sensitive species were evaluated in 

2001 (Ode 2001) and 2002 (Burkhart 2002), but it was 

determined that insufficient information is available to 

determine whether B. simplex meets the requirements 

for sensitive status (Warren 2003). Botrychium simplex 

is considered a sensitive species in Region 1 and in the 

Washington portion of Region 6, and it is considered 

important for biodiversity analysis in Region 4 (Zika et 

al. 1995). 

Botrychium simplex is not included on the BLM 

State Sensitive Species lists in Colorado (Bureau 

8 

of Land Management Colorado 2000) or Wyoming 

(Bureau of Land Management Wyoming 2002), nor is it 

listed as threatened, endangered, or candidate under the 

federal Endangered Species Act (16 USC 1531-1536, 

1538-1540). This species is not listed as endangered or 

vulnerable by the International Union for Conservation 

of Nature and Natural Resources (1978). NatureServe 

considers B. simplex to be globally secure (G5). 

Botrychium simplex is a widely distributed, 

circumboreal species, but there are concerns for its 

viability in many portions of its range. It is classified 

as endangered in Sweden (Nilsson 1981), and it is 

considered endangered in Illinois, Indiana, Maryland, 

and Ohio (USDA Natural Resources Conservation 

Service 2002). In North America, it has been 

documented from 35 states and 10 provinces as well 

as from Greenland. Botrychium simplex has been 

extirpated in three states. For detailed information 

regarding its range-wide status, see the Distribution and 

abundance section. 

Botrychium simplex is known from 50 locations 

in Region 2; 17 of these have not been revisited in more 

than 20 years, and five have not been revisited for at 

least 50 years. In the states of Region 2, B. simplex is 

ranked imperiled (S2) in Colorado and in Wyoming, 

where it has not been tracked by the Wyoming Natural 

Diversity Database since 1986. This species has been 

reported from Nebraska, and while it has not yet been 

assigned a state rank, it will probably receive a rank of 

S1. Botrychium simplex is considered unrankable (SU) 

in South Dakota and it is not known from Kansas. For 

explanations of NatureServe’s ranking system, see the 

Definitions section. 

Existing Regulatory Mechanisms, 

Management Plans, and Conservation 

Strategies 

Adequacy of current laws and regulations 

Botrychium simplex has no legal protection that 

would prevent the destruction of habitat or individuals 

on state and private land in Region 2, or on federal land 

not managed by the USFS. Because it is not listed as 

a sensitive species in Region 2, the USFS does not 

explicitly consider it for special management. As a 

Botrychium species, however, it is documented during 

Biological Evaluations where it may receive some 

special consideration.

Adequacy of current enforcement of laws and 

regulations 

There are no known cases in which an occurrence 

of Botrychium simplex was extirpated due to human 

activities or due to the failure to enforce existing 

regulations in Region 2. It is not known if federal, state, 

or other laws could have prevented the extirpation of 

B. simplex in Connecticut, Maryland, Virginia, and 

possibly Ohio. Thus, it cannot be determined if current 

regulations or their enforcement are adequate for the 

species’ protection. 

Biology and Ecology 

Classification and description 

Botrychium simplex is a member of the adder’s 

tongue family (Ophioglossaceae). Members of the 

Ophioglossaceae are eusporangiate and share a suite 

of characters that are less derived than those of most 

other Pteridophytes (Gifford and Foster 1989). In 

North America, the Ophioglossaceae as circumscribed 

by Wagner and Wagner (1993) is composed of three 

genera: Ophioglossum, Cheiroglossa, and Botrychium. 

Botrychium (grapeferns) is the most diverse of these 

genera with 50 to 60 species worldwide (Wagner and 

Wagner 1993). 

The genus Botrychium contains three subgenera: 

Osmundopteris, Sceptridium, and Botrychium 

(Wagner and Wagner 1993). Subgenus Botrychium 

(the moonworts) is the most diverse of the three, 

with approximately 25 to 30 species. Members of 

this subgenus share many morphological traits, and 

9 

subtle morphological differences make it difficult to 

identify species in the field. Based on nuclear and 

chloroplast DNA, recent phylogenetic research on 

the Ophioglossaceae has shown that the members 

of subgenus Botrychium comprise a monophyletic 

group (Hauk et al. 2003). The diversity of subgenus 

Botrychium in North America was not recognized until 

the 1980s when Drs. Herb and Florence Wagner began 

work in earnest on Botrychium. Table 1 is a summary of 

the classification of B. simplex. 

Taxonomic status 

The taxonomic status of Botrychium simplex 

has been in turmoil for more than 100 years, and there 

remains much uncertainty regarding the appropriate 

circumscription of this taxon. Reverend Edward 

Hitchcock first described B. simplex from plants found 

in Massachusetts (Hitchcock 1823). Botrychium simplex 

was the second species after B. lunaria to be described 

in subgenus Botrychium. Hitchcock’s description 

includes a hand-painted plate of B. simplex (Figure 1) 

and the following introduction: “This species grows, not 

very abundantly, in Conway, Massachusetts. It was first 

noticed, two years since, and with some doubt, referred 

to B. lunaria of Swartz and Wildenow. But upon a 

suggestion of Dr. Torrey that it might be a new species, 

I have several times re-examined it during the two past 

summers, and feel so confident that it is specifically 

distinct from any described Botrychium, that I take the 

liberty to propose for it the name ‘B. simplex.’” 

At least seven varieties of Botrychium simplex 

have been described (Table 2). The description of 

varietal taxa began with A.A. Eaton’s (1899) account 

Table 1. Classification of Botrychium simplex after USDA Natural Resources Conservation Service (2002), with 

sources (not necessarily the original source) of particular portions cited below. 

Kingdom Plantae (Plants) 

Subkingdom Tracheobionta (Vascular Plants) 

Division Pteridophyta (Ferns) 

Class Filicopsida 

1 Wagner and Wagner 1993 

2 Clausen 1938 

3 Hauk 1995 

Order Ophioglossales 

Family Ophioglossaceae (Adder’s Tongue Family) 

Genus Botrychium (Grapeferns) 

Subgenus Botrychium (Moonworts) 1 

Section Lunariae 2 , Simplex 3 

Species Botrychium simplex E. Hitchcock 1

Figure 1. The first illustration of Botrychium simplex (from Hitchcock 1823). 

of B. tenebrosum. Clute (1901) demoted this taxon to 

varietal status under B. matricariifolium. Clausen (1938) 

recognized the similarity of this taxon to B. simplex and 

included tenebrosum as a variety of B. simplex. Other 

varieties that have been described for B. simplex include 

var. cordatum (Wherry 1937), var. compositum and var. 

typicum (Clausen 1938), and var. laxifolium (Clausen 

1938, Fernald 1949). The plants from Colorado and 

Wyoming observed by Clausen 1938 were placed in var. 

typicum (including unnamed specimens from Gilpin 

and El Paso counties, where the former is probably the 

10 

1919 collection of Hazel Schmoll) and var. compositum 

(I.M. Johnson #2415 on Pikes Peak, Colorado and C.C. 

Parry #306 from Yellowstone Lake, Wyoming). Plants 

not falling into one of the other varieties are referred to 

as var. simplex. 

The varieties of Botrychium simplex are often 

morphologically distinct. During the second half of 

the 20 th century, Drs. Herb and Florence Wagner began 

to question the taxonomic validity of these varieties. 

Wagner and Wagner (1983) noted that “low, dark,

Table 2. A partial list of synonyms for Botrychium simplex. 

Source Taxon 

1 = Kartesz (1999) 

2 Botrychium simplex E. Hitchcock 

2 Botrychium simplex var. cordatum (Fr.) Wherry 

2,3 Botrychium simplex var. laxifolium (R.T. Clausen) Fernald 

2 Botrychium simplex var. laxifolium R.T. Clausen 

2,3 Botrychium simplex var. tenebrosum (A.A. Eaton) R.T. Clausen 

2,3 Botrychium simplex var. typicum R.T. Clausen 

1,3 Botrychium simplex var. compositum (Lasch) Milde 

1,3 Botrychium tenebrosum A.A. Eaton 

1 Botrychium simplex ssp. typicum R.T. Clausen 

2 Botrychium virginicum var. simplex (Hitchc.) A.Gray 

2 Botrychium lunaria var. simplex (Hitchc.) Watt 

2 Botrychium kannenbergii forma compositum Lasch 

2 = The Plant Names Project (1999) 

3 = USDA-Natural Resources Conservation Service (2002) 

acidic forest floors yield var. laxifolium, shaded bog 

edges var. tenebrosum, dry upland fields var. typicum, 

and moist low meadows var. compositum. All of these 

are connected with the others in intermediate habitats 

and there is little consistency.” Thus they suggested that 

B. simplex is an extremely plastic species that exhibits 

a phenotypic response to different habitats, as Paris 

et al. (1989) also suggested. Wagner and Wagner’s 

deconstruction of varietal concepts in B. simplex is 

culminated in their treatment in the Flora of North 

America, where they write “The many environmental 

forms and juvenile stages of B. simplex have resulted 

in the naming of numerous, mostly taxonomically 

worthless, infraspecific taxa” (Wagner and Wagner 

1993, p. 101). Instead, they offer an alternate concept 

of B. simplex in which “eastern” and “western” forms 

are described. The Wagner’s’ western plants conform 

roughly to var. compositum while their eastern plants 

fall into var. simplex. However, var. compositum 

may not properly apply to populations of the Rocky 

Mountain Cordillera where a varietal name was 

not applied (Cronquist et al. 1972). The following 

paragraphs provide detailed descriptions of these 

taxa. Lellinger (1985, p. 112) noted that “eastern, 

western, and Colorado forms exist in this species, [and] 

additional study is needed.” The “Colorado” form of B. 

simplex noted by Lellinger is probably B. minganense, 

which can be mistaken for B. simplex (Root personal 

communication 2003). 

Ongoing genetic research by Dr. Don Farrar and 

Dr. Warren Hauk suggests another concept of Botrychium 

simplex. Significant intraspecific variation in plastid 

11 

DNA sequences occurs in B. simplex, indicating the 

need for more study to evaluate the possibility that B. 

simplex (as currently circumscribed) includes multiple 

taxa (Hauk 1995, Farrar 1998). Current taxonomy 

recognizes four varieties of B. simplex: var. simplex, 

var. tenebrosum, var. compositum, and var. fontanum 

(Table 3; Farrar personal communication 2003, Farrar 

2005). Var. compositum includes plants from Minnesota 

west to Oregon and Washington (Farrar 2001, Farrar 

personal communication 2003). Most plants found in 

Colorado appear to be var. compositum. Var. simplex is 

found in the Black Hills of South Dakota and has been 

found once in Colorado (Farrar personal communication 

2003, Farrar 2005). At least five Wyoming specimens 

are labeled var. tenebrosum; this taxon is recognized by 

Hartman and Nelson (2001) for Wyoming plants. Thus 

it appears that three of the currently recognized varieties 

(var. compositum, var. simplex, and var. tenebrosum) 

may occur in Region 2. However, most Colorado 

and Wyoming specimens have not been evaluated by 

Farrar to assess their taxonomic status (Farrar personal 


Farrar (personal communication 2003, 2005) 

described the fourth variety, var. fontanum, for 

genetically distinct plants from southern California, 

southern Nevada, northeastern Oregon, and 

southeastern Washington. Var. fontanum is typically 

found in calcareous fens and seeps (Farrar 2005). The 

presence of this variety in Colorado or elsewhere in 

Region 2 has not been confirmed, but calcareous fens 

and seeps in Region 2 need to be searched for this taxon 

(Farrar personal communication 2003).

Table 3. Summary information for the varieties of Botrychium simplex recognized by Farrar (2005). 

Variety Diagnostics Range Habitat and notes 

simplex Plants with mostly undivided basal pinnae 

and trophophore stalk equal to or exceeding 

length of the common stalk. Larger plants 

have pinnae that become progressively 

more dissected, with lower pinnae more 

elongated. 

tenebrosum Slender plants with undivided basal pinnae 

and very short trophophore and sporophore 

stalks but relatively long common stalks. 

compositum Plants with secondarily divided basal 

pinnae and trophophore stalk equal to or 

exceeding length of the common stalk. 

fontanum Robust plants with basal pinnae divided 

or not, thick and fleshy with broad, bluish 

green pinnae and terminal pinnae with 

broadly rounded apices. 

Isozyme analysis demonstrates a close relationship 

of most western plants to var. compositum (Farrar 

2001). Although Farrar and Wendel (1996) and Farrar 

(1998) note that the genetic distance between three 

varieties of Botrychium simplex approaches that of full 

species, current genetic evidence suggests that the range 

of variability in B. simplex falls within that of a single 

species (Table 4). As noted by Wagner and Wagner 

(1983), plants in different ecological situations (e.g., 

seasonally dry meadows, saturated fens) have marked 

morphological differences, but they also have marked 

Northeastern US, west through the 

Great Lakes Region, with disjunct 

reports in Nebraska and the Black Hills, 

apparently also in Colorado (Douglass 

#62-25) 

Northeastern US and Great Lakes 

Region west to Iowa and Minnesota; 

reported in Wyoming (Hartman and 

Nelson 2001) 

Mountains of the western US 

(including Colorado) and Canada 

California, Nevada, NE Oregon, SE 

Washington 

12 

meadow, woodland 

forest, swamp margin, 

dune 

meadow, roadside 

Calcareous fens and 

hardwater seeps. 

Possibly in CO; further 

surveys are needed. 

genetic differences (Farrar 2001). Given the current 

uncertainties regarding the proper circumscription of B. 

simplex, it is likely that future investigation will result 

in further changes and refinements that may include the 

description of other varietal taxa or even new species. 

Botrychium simplex is closely related to two rare 

and narrowly endemic moonworts, B. mormo and B. 

pumicola (Hauk 1995, Hauk et al. 2003). Botrychium 

mormo was described in 1981 after three decades of 

study (Wagner and Wagner 1981). It is a tiny moonwort 

Table 4. Genetic variability in eastern diploid species of Botrychium subgenus Botrychium from Farrar (1998). These 

results support the genetic and taxonomic uniqueness of the varieties simplex, tenebrosum, and compositum. Most 

moonworts show little heterozygosity (number of alleles per locus), and do not have a high percentage of polymorphic 

loci. However, B. simplex sensu lato shows very high percentage of polymorphic loci relative to other eastern 

moonworts. When the varieties are analyzed separately they fall into a more normal range, suggesting that they may 

even warrant treatment as full species. 

Species 

Mean sample size 

per locus 

Mean number of alleles 

per locus Percentage of loci polymorphic 

B. lunaria 41.1 1.1 11.1 

B. campestre 96.9 1.2 16.7 

B. pallidum 20.6 1.1 5.6 

B. simplex sensu lato 96.2 1.7 61.1 

var. simplex 27 1.1 5.6 

var. tenebrosum 27.8 1.1 11.1 

var. compositum 15.8 1.1 5.6 

B. mormo 48.8 1.1 5.6 

B. lanceolatum ssp. lanceolatum 24.6 1 0 

B. lanceolatum ssp. angustifolium 29.2 1.1 5.6

that is found in rich woods of Michigan, Wisconsin, and 

Minnesota (Wagner and Wagner 1981). Botrychium 

simplex is the most similar species to B. mormo, and the 

two can be difficult to distinguish (Chadde and Kudray 

2001a). These species do not occur together in Region 

2. Botrychium mormo is genetically distinct from all 

varieties of B. simplex (Farrar and Wendel 1996). 

Botrychium pumicola is known only from volcanic 

substrates and frost pockets in Oregon (Hopkins et al. 

2001). The morphological distinctness of B. pumicola 

from B. simplex has been long recognized. Botrychium 

pumicola was described in 1900 (Coville 1900) and has 

now been shown to be clearly genetically distinct from 

B. simplex (Farrar 2000). Using isozyme data (Hauk 

and Haufler 1999) and rbcL data (Hauk et al. 2003), B. 

pumicola appears to be the most closely related to B. 

simplex of the species analyzed. Botrychium pumicola 

was also the closest relative in a combined analysis of 

rbcL, trnL-F and morphological data, with B. montanum 

the next closest relative (Hauk et al. 2003). Botrychium 

mormo was not included in this phylogenetic study. 

Botrychium pumicola is one of a handful of 

moonwort species that have been observed to produce 

gemmae (Camacho 1996, Camacho and Liston 2001), 

which are minute vegetative propagules abscised at 

maturity from the parent plant (Farrar and Johnson- 

Groh 1990). Camacho (1996) found 0 to 10 gemmae 

per plant in B. pumicola. 

Botrychium campestre also produces gemmae 

and was the first moonwort species in which they 

were documented (Farrar and Johnson-Groh 1986, 

Farrar and Johnson-Groh 1990, Johnson-Groh et al. 

2002). Subsequent research has found them on other 

diploid species, including B. pumicola. Botrychium 

gallicomontanum, a rare allotetraploid species for 

which B. campestre and B. simplex are the putative 

parent species, is also known to reproduce with gemmae 

(Farrar and Johnson-Groh 1991). The production 

of gemmae as vegetative propagules by these three 

species, all of which are found in relatively xeric sites, 

suggests that it is an adaptation for reproduction in dry 

sites (Camacho 1996). Farrar and Johnson-Groh (1986) 

13 

examined B. simplex for gemmae, but none were found. 

Gemmae may confer a lesser advantage to B. simplex, 

which is typically found in habitats that are at least 

seasonally wet. 

Botrychium pumicola is unusual in that its spores 

are dispersed in loose groups of four spores also called 

tetrads (Wagner 1998). Despite the close relationship 

between B. simplex and B. pumicola, this phenomenon 

has not been documented in B. simplex. 

Botrychium simplex is a putative parent species for 

several polyploid nothospecies in subgenus Botrychium 

(Table 5; Wagner 1993, Farrar and Wendel 1996). 

Description 

Botrychium subgenus Botrychium sporophytes 

are simple plants recognized by their small size and 

distinctive leaf and spore structures. Members of this 

subgenus are usually less than 15 cm in height. They 

possess a trophophore, or sterile leaf-like structure 

that is often heavily lobed or segmented, but rarely 

truly pinnate (Wagner and Wagner 1993). Members 

of the subgenus Botrychium usually only produce one 

leaf each year and in some years produce no leaves 

(Johnson-Groh 1998). On the same stalk sits a fertile 

sporophore that is often taller than the trophophore. The 

sporophore contains 20 to 100 grape-like sporangia, 

each containing possibly thousands of spores (Farrar 

and Johnson-Groh 1986, Wagner 1998). 

Botrychium species can be difficult to identify due 

to their subtle diagnostic characters, frequent occurrence 

with other Botrychium species, and morphological 

variability (Paris et al. 1989). Because they are such 

simple plants, there are few morphological characters 

that can be used to distinguish species; identification 

is often based on very subtle characters (Hauk and 

Haufler 1999). Identification is facilitated by the use 

of dichotomous keys (see Weber and Wittmann 2001a 

and Weber and Wittmann 2001b); however, these do 

not guarantee a positive identification, and it is often 

necessary to get verification by a Botrychium expert. 

Table 5. Hypothetical parents for polyploid species of Botrychium subgenus Botrychium (after Hauk 1995, from 

Wagner 1993 and Farrar and Wendel 1996). 

Polyploid species Hypothetical parents 

B. hesperium B. lanceolatum x B. simplex 

B. pseudopinnatum B. pinnatum x B. simplex 

B. gallicomontanum B. campestre x B. simplex

Botrychium simplex is challenging to identify 

with confidence since it is highly variable, small, and 

cryptic. Farrar (2001) notes that “B. simplex is by far 

the most variable of diploid moonworts,” and Wagner 

and Wagner (1983) wrote that “Only a few botrychiums 

have such astonishing variability as B. simplex.” They 

also noted that “no species approaches B. simplex in the 

extent of its variability. Var. compositum stands in vivid 

contrast to var. tenebrosum.” 

Within Region 2, both the eastern and western 

forms described by Wagner and Wagner (1993) are 

present, and it appears that all three of the varieties 

recognized by Wagner and Wagner (1993) occur here. 

Wagner and Wagner (1993) provide a summary of the 

diagnostic characters of Botrychium simplex; these are 

summarized in Table 6. A comparison of the varieties 

recognized by Farrar (2005) is included in Table 3. 

Botrychium simplex is a small perennial fern, 

seldom exceeding 8 cm tall (Lorain 1990). It has 

small roots (0.5 to 1 mm in diameter) and a highly 

variable trophophore, which is 1 to 7 cm long, oblong 

to long-elliptic, 0.3 to 2 cm wide, truncate to round 

at the base, round at the apex, pinnate or sometimes 

nearly simple, entire, round pinna apices, and entire 

or crenulate margins (Lellinger 1985, Farrar 2005). 

Useful field marks for B. simplex include its diminutive 

size, succulent stem, single compound leaf that is often 

clasping the sporophore, its unbranched fertile frond, 

and the tendency for the trophophore and sporophore 

to connect at ground level (Figure 2; Farrar 2005). It 

is highly variable and only distinguished absolutely 

from other grapeferns by its larger spores, which 

are unusually large for a diploid species, ranging in 

diameter from 0.035 to 0.050 mm (Rook 2002, Farrar 

2005). Occasional plants are found that have sporangia 

dotting the margins of the trophophore (Figure 3). 

These are called supernumerary sporangia and are seen 

infrequently in all moonwort species (Farrar 2005). 

Botrychium simplex is diploid with 45 chromosomes 

(2n = 90) (Wagner 1993, Wagner and Wagner 1993). 

Detailed descriptions for both “eastern” and 

“western” Botrychium simplex appear in Wagner and 

Wagner 1993. Several field characteristics are useful for 

distinguishing these two types of B. simplex. Western B. 

simplex has a sporophore that is longer relative to the 

trophophore, lacks a common stalk, and has fan-shaped 

pinnae. See Table 6 for a comparison of diagnostic 

characteristics between these types, and Figure 3, 

Figure 4, and Figure 5 for specimens representing 

these types. 

14 

Botrychium simplex var. compositum is roughly 

equivalent to the “western” B. simplex of Wagner and 

Wagner 1993 (Figure 3). It is usually has a three-parted 

leaf as shown in Gray (1908), Hitchcock and Cronquist 

(1969), and Cronquist et al. (1972). The shape of 

the trophophore is distinctive in having three main 

branches (Lorain 1990). The sporophore diverges at or 

just above ground level (Welsh et al. 1993, Farrar 2005) 

or sometimes below ground level (as noted on some 

specimens collected by Peter Root). 

Botrychium simplex var. tenebrosum occurs in 

eastern North America (Wagner and Wagner 1993) and 

elsewhere, including Wyoming (Figure 4; Hartman and 

Nelson 2001). It was thought by Wagner and Wagner 

(1993) and others to be a persistent juvenile, but current 

genetic evidence suggests otherwise. Var. tenebrosum is 

distinguished from var. simplex (Figure 5) by its smaller 

size, slender stature, and its simple and rudimentary 

trophophore attached near the top of an exaggerated 

common stalk. Wagner and Wagner (1993) describe 

a western equivalent to B. simplex var. tenebrosum, 

with a lower attachment of the trophophore, which is 

longer and more herbaceous in texture. The spores of 

var. tenebrosum are larger than those of var. simplex 

(Eaton 1899). Clausen (1938) includes photographs 

of specimens of B. simplex var. tenebrosum. Var. 

tenebrosum has been mistaken for B. simplex var. 

simplex and B. matricariifolium. 

Botrychium simplex is frequently confused with 

other species of Botrychium in Region 2 and elsewhere. 

Within Region 2, B. simplex specimens have been 

misidentified as B. lunaria and B. hesperium. In several 

cases, one plant on an herbarium sheet containing 

several plants has been annotated as B. lunaria or in one 

case B. paradoxum. Many medium or large specimens 

of B. simplex have flabellate pinnae that strongly 

resemble B. lunaria (Wagner and Wagner 1981). In 

Oregon, well-formed plants can be mistaken for B. 

pumicola, and immature plants are easily confused 

with B. minganense and B. lunaria (Zika et al. 1995). 

A key point in separating B. simplex from simplex-like 

minganense (B. “colorado”) is that in B. simplex, the 

sporophore separates from the stipe at or just below the 

soil surface (Figure 2; Root personal communication 

2003). Before its circumscription, B. montanum was 

known as B. simplex in Montana (Vanderhorst 1997). 

Botrychium simplex also looks like B. mormo (Wagner 

1998). In Iowa, Michigan, and Nebraska, B. simplex has 

been found associated with B. campestre, with which 

it is the putative parent species of B. gallicomontanum 

(Farrar and Johnson-Groh 1991, Farrar personal 

communication 2003).

Table 6. Diagnostic characteristics presented in Wagner and Wagner (1993) for the determination of the “eastern” and “western” types 

of Botrychium simplex. 

Characteristic B. simplex sensu lato “eastern” B. simplex “western” B. simplex 

Sporophore 1-pinnate; one to eight times the length of the 

trophophore 

Figure 2. Botrychium simplex sporophytes at Dugout Gulch, Wyoming, Black Hills National Forest (BOSI-12) after emergence, and 

before separation of the trophophore and sporophore. Over 200 individuals were seen at this location in 2004. Photo by Katherine 

Zacharkevics, provided by Beth Burkhart. 

15 

One to four times the length of the 

trophophore; arises from common 

stalk below middle to near the top, 

well above the leaf sheath 

Three to eight times the length 

of the trophophore, arises 

directly from the top of the leaf 

sheath 

Common stalk Absent or well developed Well developed Much reduced or absent 

Trophophore Stalk 0 to 3cm, 0 to 1.5 times the length of the 

trophophore rachis, blade dull to bright green, 

linear to ovate-oblong to fully triangular with 

ternately arranged pinnae 

Pinnae Up to seven pairs of pinnae or well-developed 

lobes, spreading to ascending, closely or widely 

separated, distance between 1st and 2nd pinnae 

pairs is frequently greater than between 2nd 

and 3rd pairs, basal pinna pair larger and more 

complex than adjacent pair, cuneate to fanshaped, 

strongly asymmetric, venation pinnate 

or like ribs of fan, with midrib 

Habitat Dry fields, marshes, bogs, swamps, roadside 

ditches 

Nonternate or if subternate, the 

lateral pinnae are smaller than 

the central pinnae and simple 

to merely lobed, tip undivided, 

texture papery to herbaceous 

Lateral pinnae are smaller than 

central pinnae, simple to merely 

lobed or rarely pinnate, pinnae 

adnate to rachis, rounded and 

ovate to spatulate, segment sides at 

angles mostly less than 90 degrees 

Ternate with three equal 

segments, or rarely non-ternate 

but resembling a single segment 

of a ternate blade, tip divided 

usually into three parts, texture 

thin and herbaceous 

Pinnae usually strongly 

contracted at the base to stalked, 

angular to fan-shaped, segment 

sides at angles mostly more than 

90 degrees as in Botrychium 

lunaria 

Often upland fields Along marshy margins and in 

meadows

Figure 3. Botrychium simplex var. compositum (“western” simplex) collected on Boreas Pass, Colorado. 

The gametophytes of Botrychium species remain 

poorly understood. They are achlorophyllous and 

are wholly dependent on mycorrhizal fungi for their 

water, mineral nutrients, and carbohydrates (Campbell 

1922, Bower 1926, Scagel et al. 1966, Gifford and 

Foster 1989, Schmid and Oberwinkler 1994). The 

gametophytes of other Botrychium species have been 

cultured and studied (Campbell 1911, Whittier 1972, 

Whittier 1973, Whittier 1981, Whittier 1984, Melan 

and Whittier 1989, Thomas and Whittier 1993). 

The gametophyte of Botrychium simplex was 

studied in detail by Campbell (1922). Like all other 

Botrychium species studied, the gametophyte of B. 

simplex has a dorsal ridge that bears the gameteproducing 

structures. Antheridia are borne at the top 

of the dorsal ridge and archegonia on either side of 

the dorsal ridge. The gametophytes of B. simplex 

are monoecious, with both male and female gameteproducing 

structures. The gametophyte is up to 5 

mm long and is usually obovoid or club-shaped. The 

rhizoids (root-like structures) form on the lower surface. 

Botrychium simplex gametophytes have been found 

with young sporophytes attached (Clausen 1938). 

16 

Sources for keys photographs, illustrations, and 

descriptions 

There are numerous sources of keys, photographs, 

illustrations, and descriptions that are of great value in 

identification of Botrychium simplex. The best source 

currently available for use in Region 2 is Farrar (2005), 

which includes keys, descriptions, photographs, and a 

discussion of the diagnosis, distribution, and habitats 

of all western North American moonworts. This source 

includes details for each of the currently recognized 

varieties of B. simplex (vars. simplex, tenebrosum, 

compositum, and fontanum), but unfortunately it is 

unpublished and is not yet widely available. Root 

(2003) is another useful but unpublished source of 

information on moonworts in Colorado that includes 

keys, silhouettes, and diagnostic information. Wagner 

and Wagner (1993) include a description of B. simplex. 

Lellinger (1985) includes a good description, key, 

and photograph. Internet sources including Wisconsin 

State Herbarium (2003) and Rook (2002) contain 

photographs, habitat information, and links to other 

sources. Wagner and Wagner (1983) include a figure 

showing the varieties of B. simplex.

Figure 4. Botrychium simplex var. tenebrosum from Park County, Wyoming (Kirkpatrick #5317). 

Figure 5. Botrychium simplex var. simplex (“eastern” simplex) from East Inlet, Rocky Mountain National Park, 

Colorado (Douglass #62-25). 

17

There are also several sources of illustrations 

of Botrychium simplex. Hitchcock (1823) includes 

the first illustration of B. simplex (Figure 1). Huxley 

(1972) contains an illustration of B. simplex in Europe. 

Hitchcock et al. (1969) and Cronquist et al. (1972) 

include an illustration of “western” B. simplex (var. 

compositum). This illustration is also included in 

Lackschewitz (1991). Another illustration of “western” 

B. simplex is found in Dorn and Dorn (1972). Gray (1908) 

includes an illustration of var. compositum. Chadde and 

Kudray (2001a) provide an illustration of “eastern 

simplex.” A description and illustration are included in 

Britton and Brown (1913) for both B. simplex (Figure 

6) and B. tenebrosum (Figure 7). Polunin (1959) 

includes an illustration of var. tenebrosum. Weber and 

Wittmann (2001a, 2001b) provide a brief description, 

an illustration, and a key for Botrychium in Colorado. 

Campbell (1922) includes detailed descriptions and 

numerous detailed illustrations of the gametophyte and 

embryo of B. simplex. 

Figure 6. Illustration of Botrychium simplex (Britton and Brown 1913). 

18 

Silhouettes can be helpful in identifying 

moonworts. Farrar (2001) includes the silhouettes 

of many moonwort species, including Botrychium 

simplex. Mantas and Wirt (1995) also include 

silhouettes, illustrations, and a description of B. simplex 

in Montana. 

Because of its wide distribution, many floras 

and field guides describe Botrychium simplex (e.g., 

Coulter and Nelson 1909, Rydberg 1922, Davis 1952, 

Harrington 1954, Peck 1961, Gleason and Cronquist 

1963, Munz and Keck 1968, Huxley 1972, Great Plains 

Flora Association 1986, Lackschewitz 1991, Welsh et 

al. 1993). Lorain (1990) includes a key for moonworts 

found in the Idaho Panhandle and a description of B. 

simplex (var. compositum). Full technical descriptions 

for B. simplex and its varieties are in Clausen (1938). 

Kolb and Spribille (2000) and Farrar (2005) include 

useful tables comparing the diagnostic characteristics of 

western moonwort species including B. simplex. There 

is no type specimen for B. simplex (Zika et al. 1995).

Figure 7. Illustration of Botrychium tenebrosum (Britton and Brown 1913). 

Distribution and abundance 

Members of subgenus Botrychium are distributed 

worldwide, predominantly in temperate and northern 

temperate habitats, but there are also representatives in 

temperate South America, New Zealand, and Australia 

(Clausen 1938, Wagner and Wagner 1993). In general, 

western North America and the Great Lakes region are 

recognized as centers of diversity and abundance for 

this subgenus. 

Botrychium simplex is a circumboreal species, 

known from North America, Greenland, Iceland, 

Europe, Scandinavia, Corsica, and Japan (Figure 

\8; Clausen 1938, Wagner and Wagner 1986, Ollgaard 

1971, Farrar 2001, Anderberg 2003). In the western 

hemisphere, it is known from Greenland across the 

Canadian arctic and subarctic to Alaska, south to 

California, Colorado, Mississippi, and North Carolina 

(Figure 9; Kartesz 1999, Farrar 2005). It is known from 

10 provinces in Canada, 36 U.S. states and the District 

of Columbia. In some parts of its range, B. simplex is 

more common than other Botrychium species, and it is 

ranked G5 (globally secure) accordingly. Along with 

19 

B. matricariifolium, B. simplex is the most common 

moonwort in eastern North America. However, in 

many parts of its range, including Region 2, B. simplex 

is locally quite rare. It has apparently been extirpated 

in Connecticut, Maryland, Virginia, and possibly 

Ohio (Kartesz 1999). See Table 7 for a list of states 

and provinces where B. simplex is found, including 

subnational (S) ranks and special state designations. 

It appears that three of the four currently 

recognized varieties of Botrychium simplex 

(compositum, simplex, and tenebrosum) occur in 

Region 2. Dr. H. Wagner annotated many Colorado 

specimens housed at the University of Colorado 

Herbarium and noted whether they were “western” 

or “eastern” simplex. From Wagner’s annotations, it 

appears that both “eastern” (either var. simplex or an 

undescribed variety) and “western” (probably mostly 

var. compositum) B. simplex have been documented in 

Colorado (Farrar personal communication 2003, Root 

personal communication 2003). However, Wagner 

noted only one specimen (Douglass #62-25; from 

Grand County, Colorado) to be “eastern” simplex, 

and from photographs, Farrar has confirmed that this

Figure 8. Global distribution map of Botrychium simplex (Anderberg 2003). 

is either “eastern” simplex (var. simplex) or the new 

undescribed variety from the Southwest. Farrar may 

employ molecular techniques to confirm the taxonomic 

status of these plants (Farrar personal communication 

2003). This occurrence has not been seen since 

1962, despite attempts by Peter Root to find it (Root 

personal communication 2003). At least five Wyoming 

specimens housed at the Rocky Mountain Herbarium 

are labeled var. tenebrosum, and Wyoming material 

falls into this variety, according to Hartman and Nelson 

(2001). Most Colorado and Wyoming specimens 

have not been re-examined by Farrar in light of the 

contemporary species concept. 

Two erroneous sources of distribution information 

were found in the literature. The distribution map in 

Wagner and Wagner (1993) does not show occurrences 

in the Black Hills, which contain one occurrence in 

Lawrence County, South Dakota and several, newly 

discovered occurrences in Crook County, Wyoming. 

These occurrences are important to note since they add 

continuity to the distribution of Botrychium simplex 

across North America. Wherry (1938) noted that 

20 

Rydberg had misleadingly attributed this species to 

the prairies of Colorado, but the source of Rydberg’s 

erroneous report was not found. Rydberg (1906) and 

Rydberg (1922) do not include the plains of Colorado 

within the distribution of B. simplex. 

Many surveys for Botrychium simplex and other 

moonwort species have been completed in the past 

two decades (e.g., Lorain 1990, Vanderhorst 1997). 

Recent surveys in Region 2 have led to the discovery 

of new moonwort occurrences (e.g., Fertig 2000, Kolb 

and Spribille 2000, Thompson 2000, Buell 2001, 

Steinmann 2001a, Steinmann 2001b, Thompson 2001, 

Abbott 2003, Crook personal communication 2003, 

Farrar personal communication 2003, Fertig 2003, 

Root personal communication 2003, Farrar 2005, 

Burkhart personal communication 2006). The known 

distribution of this species in the states of Region 2 

consists of disjunct clusters of occurrences (Figure 

10, Table 8). This apparent pattern may be an artifact 

of the intensity of searching that has been conducted in 

particular areas rather than an ecologically meaningful 

pattern. For example, extensive searches in Yellowstone

21 

State/Province 

Conservation 

Status 

SX: Presumed 

Extirpated 

SH: Possibly 

Extirpated 

S1: Critically 

Imperiled 

S2: Imperiled 

S3: 

S4: Apparently 

Secure 

S5: 

Vulnerable 

Secure 

Not Ranked/ 

Under Review 

(SNR/SU 

Figure 9. Distribution of Botrychium simplex in North America, color-coded by S rank (from NatureServe 2005). 

Botrychium simplex is now ranked S2 in Colorado. 

Table 7. Known distribution of Botrychium simplex in the western hemisphere (from Ollgaard 1971, Kartesz 1999, 

USDA Natural Resources Conservation Service 2002, and NatureServe 2005). USDA Forest Service Region 2 states 

are in bold. 

Nation State/Province/District S rank State Designation Status 

Canada Alberta S2 

Canada British Columbia S2S3 

Canada New Brunswick S4 

Canada Newfoundland S2 

Canada Northwest Territories SNR 

Canada Nova Scotia S2S3 

Canada Ontario S4? 

Canada Prince Edward Island S1 

Canada Quebec S3S4 

Canada Saskatchewan S1 

Greenland Julianehaab none 

USA California SNR 

USA Colorado S2 

USA Connecticut SH Special Concern Extirpated 

USA Delaware SNR

Table 7 (concluded). 

Nation State/Province/District S rank State Designation Status 

USA District of Columbia SNR 

USA Idaho S2 

USA Illinois S1 Endangered 

USA Indiana S1 Endangered 

USA Iowa S1 Threatened 

USA Maine SNR 

USA Maryland SH Endangered Extirpated 

USA Massachusetts SNR 

USA Michigan SNR 

USA Minnesota S3 Special Concern 

USA Mississippi SNR 

USA Montana SU 

USA Nebraska none 

USA Nevada SNR 

USA New Hampshire SNR 

USA New Jersey SNR 

USA New York SNR Exploitably Vulnerable 

USA North Carolina S1 

USA North Dakota SU 

USA Ohio S1 Endangered Possibly Extirpated 

USA Oregon S4 

USA Pennsylvania S5 

USA Rhode Island S1 

USA South Dakota SNR 

USA Utah S1 

USA Vermont SNR 

USA Virginia S1 Extirpated 

USA Washington S3 Sensitive 

USA West Virginia SNR 

USA Wisconsin unknown 

USA Wyoming S2 

National Park (outside of Region 2) have located 

numerous occurrences, but adjacent areas within 

Region 2 have been less intensively searched. Intensive 

searches on the Black Hills National Forest occurred 

in 2004 and 2005 and resulted in the discovery of 14 

new occurrences, whereas it was previously known 

from only three occurrences. In fact, B. simplex was 

the most commonly encountered moonwort species 

(Burkhart personal communication 2006). On the other 

hand, Hollis Marriott and Walter Fertig conducted 

Botrychium surveys in Wyoming in 1989, 1993, and 

1999 (Fertig 2000). While many occurrences of other 

species were found, these surveys resulted in relatively 

22 

few discoveries of B. simplex plants. Forested habitats 

have not been extensively searched for B. simplex in 

Region 2 (Root personal communication 2003). Also, 

because of the tendency to find B. simplex in wetter, 

shadier habitats than other moonworts in Region 2, this 

species may not be encountered as often and therefore 

be underreported. 

The majority of the known occurrences of 

Botrychium simplex in Region 2 are on National Forest 

System land (Table 9). These are distributed across 11 

national forests, with most occurrences reported from 

the Black Hills, Shoshone, and White River national

Table 8. Summary information for the known occurrences of Botrychium simplex in the states of USDA Forest Service Region 2, including occurrences in Wyoming that are outside 

of the Region 2 administrative boundary. 

Collector / 

Observer Herbarium Notes Habitat 

Unknown Prettyman COLO Wagner 1989: Wet soil of peat bog 

(s.n.) 

“western simplex” 

management Abundance 

Elevation Date first Date last Land ownership/ 

State County Location (ft.) observed observed 

1 Colorado Boulder Caribou Flats 9,750 7/28/1949 7/28/1949 USDA Forest 

Service (USFS) 

Roosevelt 

National Forest 

COLO Verified by Peter Root In the open hillside meadow on the 

south facing slope to the east of the 

trailhead and on the north side of 

the road, along the edges of willows 

and near rocks. 

Unknown David 

Steinmann 

(s.n.) 

10,100 7/27/2001 7/27/2001 USFS 

Roosevelt 


2 Colorado Boulder 4th of July 

Trailhead 

Not reported 

N/A Root personal 

communication 2003; 

note in CNHP files 

from unknown source 

1 Florence 

Wagner 

City of Denver 

Echo Lake Park 

8/4 or 

8/5/1984 

10,600 8/4 or 

8/5/1984 

South of Echo 

Lake 

3 Colorado Clear 

Creek 

On flat, southeast facing, dry, open 

area in bottom of canyon; with 

COLO Originally identified as 

Botrychium hesperium 

“Abundant” M. Aitken and 

E. Vanwie 

(RG30) 

9,020 7/1/1995 7/1/1995 USFS 

Rio Grande 


4 Colorado Conejos Terrace 

Reservoir 

Antennaria spp. Sporulating 

SJNM Rolling tundra with tree islands of 

Picea with many ponds; bare spots 

in grassy matrix on south-facing 

slope 

Unknown S. O’Kane 

(5872), Arnold 

Clifford, Dave 

Jamieson 

11,490 8/8/2001 8/8/2001 USFS 

San Juan National 

Forest 

5 Colorado Conejos Between Fish 

Lake and Blue 

Lake 

Not reported 

GH Wherry (1938), 

Clausen (1938) 

Unknown I.M. Johnston 

(2415) 

Probably USFS 

Pike National 

before 

1937 

6 Colorado El Paso Pikes Peak ~10,940 before 

1937 

Forest 

Near border of lake 

COLO Wagner 1989: 


(orig. labeled 

Botrychium lunaria) 

Unknown H.M. Schmoll 

(11) 

7 Colorado Gilpin Teller Lake 9,600 8/1/1919 8/1/1919 USFS 

Roosevelt 


Not reported 

COLO T. Spribille: one plant 

may be Botrychium 

paradoxum - sent to 

F. Wagner. Specimen 

originally ident. as B. 

lunaria. Annotated by 

P. Root. 

Unknown MM. Douglass 

(62-31) 

8,700 6/29/1962 6/29/1962 National Park 

Service (NPS): 

Rocky Mountain 

National Park 

8 Colorado Grand North Inlet 

Trail

Table 8 (cont.). 


Collector / 

State County Location (ft.) observed observed management Abundance Observer Herbarium Notes Habitat 

9 Colorado Grand East Inlet 8,500 6/27/1962 6/27/1962 NPS 

Unknown M.M. COLO Annotated as 

Near the falls, in wet marshy area of 

Rocky Mountain 

Douglass 

Botrychium simplex spruce-fir forest right beside stream 

National Park 

(62-25) 

by W. Wagner (1989); 

originally ident. as B. 

lunaria; P. Root noted 

that Wagner considers 

this specimen to be 

“eastern simplex.” 

This was confirmed 

by Farrar (personal 

communication 2003) 

Spruce/fir forest and/or subalpine 

meadow 

10 Colorado Gunnison No information 9,200? No data No data No data No data Arnett (2002) RM? Mentioned in Arnett 

(2002), no specimen 

at RM 

RM Sporulating Avalanche meadow 

Unknown M. Arnett 

(7417) 

10,600 8/24/1999 8/24/1999 Bureau of Land 

Management 

(BLM) 

11 Colorado Hinsdale Northeast 

of Gravel 

Mountain 

RM Sporulating Spruce forest to krummholz; along 

gulch 

12,200 8/30/1999 8/30/1999 BLM Unknown M. Arnett 

(7902) 

12 Colorado Hinsdale East fork of 

Alpine Gulch 

RM Sporulating Alpine tundra; basin of an unnamed 

lake 

Unknown M. Arnett 

(6836) 

12,680 8/11/1999 8/11/1999 USFS 

Gunnison 


13 Colorado Hinsdale Upper West 

Fork of 

Mineral Creek 

COLO Sporulating On dry, open northwest slope of tuff 

12 M. Aitken and 

E. Vanwie (RG 

174 & 175) 

9,000 7/27/1995 7/27/1995 USFS 

Rio Grande 


14 Colorado Hinsdale Thirty Mile 

Campground 

Along railroad tracks; open, dry site 

N/A Late in season; many 

dried-up plants 

5 to 10 A. Kolb & T. 

Spribille 

15 Colorado Lake Fremont Pass 11,100 9/12/2000 9/12/2000 USFS 

San Isabel 


COLO? Not reported 

16 Colorado Saguache No information No data No data No data No data No data University 

of Colorado 

Herbarium 

(2003) 

COLO Open, clearcut, subalpine forest 

A. Kolb & T. 

Spribille (8) 

more than 

10 

11,155 809/2000 8/9/2000 USFS 

White River 


17 Colorado Summit Breckenridge 

Ski Area



Collector / 

State County Location (ft.) observed observed management Abundance Observer Herbarium Notes Habitat 

18 Colorado Summit Boreas Pass 11,483 8/9/2000 8/9/2000 USFS 

20 to 50 A. Kolb & T. COLO Open, eroded, grassy slopes at 

White River 

Spribille (11) 

timberline; area of historical burn; 


on gravelly soil among young 

spruce trees; also on roadbeds; 

aspect: west; moisture: dry 

~50 or more Nancy Redner KH Subalpine meadow; downslope 

of subalpine forest of lodgepole 

and spruce-fir, on a 15 to 35 

percent slope; organic and granite 

soils, cobbly, seasonally wet but 

becoming dry; hillside is used by 

elk 

11,400 7/22/1999 7/22/1999 USFS 

White River 


19 Colorado Summit Vail Pass Rest 

Area 

In a narrow road between ski trails, 

in shade 

1 P. Root (1258) KH Three-part trophophore 

partly eaten away, with 

vehicle tracks on the 

plant 

~10,500 7/14/2000 7/14/2000 USFS 

White River 


20 Colorado Summit Copper 

Mountain 

N/A Not reported 

less than 10 A. Kolb & 

T. Spribille 

(2000) 

21 Colorado Summit Keystone ~10,500 2000 2000 USFS 

White River 


COLO Open, eroded, grassy slopes, 

subalpine; area of historical burn; 

red sandstone; west-facing slope 

5 A. Kolb & T. 

Spribille (22) 

Shrine Pass 11,155 8/15/2000 8/15/2000 USFS 

White River 


22 Colorado Summit/ 

Eagle 

Marshy ground; in a marshy area 

among Carex aquatilis 

COCO Baker and Kuntz 

(1983). COCO 

Specimen was verified 

by D. Farrar 

Unknown Darrow and 

Sigstedt (9340) 

(1980); W.L. 

Baker and D. 

Kuntz (1983) 

~9,000 7/02/1980 9/28/1983 State of Colorado 

Dome Rock State 

Wildlife Area 

23 Colorado Teller Mueller Ranch, 

Reservoir #1 

Not reported 

KH Wagner considers all 

but one of the plants 

on this sheet to be 


Unknown J.B. Hartwell, 

C.W.T. 

Penland, and 

G.R. Marriage 

(1251) 

24 Colorado Teller Mount Baldy 10,300 6/28/1942 6/28/1942 USFS 

Pike National 

Forest 

In the sandy floodplain in swales 

under cottonwood and juniper 

along the Niobrara River below 

the narrow canyon area; on river 

floodplain in shaded opening 

between large Juniperus virginiana 

ISC Not found at this 

location in attempts 

to find it in 2004 

by Farrar (personal 


Farrar and 

Johnson-Groh 

(1986), D.R. 

Farrar (86-6- 

“fairly 

common” 

~2,400 Unknown ~6/2/1986 The Nature 

Conservancy 

25 Nebraska Brown Niobrara 

Valley Preserve 

2-2)


Collector / 

Observer Herbarium Notes Habitat 

management Abundance 


(ft.) observed observed 

5,630 6/18/2004 7/8/2004 USFS 

Black Hills 


State County Location 

In thick moss cover, near a faint 

game trail (which has caused 

disturbance on the slope) and a 

young downed spruce; in partial 

shade near a drainage bottom, in 

dry-mesic soil 

N/A Nearing sporulation on 

July 8 

1 C. Skelton, D. 

King, and T. 

Price (BOSI- 

10) 

Custer Lightning 

Creek 

26 South 

Dakota 

Unknown Unknown Unknown from SDNHP In grass with Agoseris glauca 

~4,500 6/20/1953 6/20/1953 USFS 

Black Hills 


Lawrence Steamboat 

Rock 

27 South 

Dakota 

In a large open meadow with high 

forb cover at the edge of an old 

stock pond and near a small stand 

of young aspen and an old roadbed; 

N/A Not yet sporulating 

on June 22, yellowing 

stalk and brown 

sporangia on July 

9; Botrychium 

minganense is also 

found at this location 

14 C. Skelton, 

D. Farrar, T. 

Price, and D. 

King (BOSI-8, 

BOTR-11) 

Lawrence Tinton Road 6,426 6/12/2004 7/9/2004 USFS 

Black Hills 


28 South 

Dakota 

moisture: dry/mesic 

N/A On the toe of a north-northeast 

facing slope above a southeast 

facing draw, with high cover of 

moss and forbs. In partial shade in 

lower slope of the draw. Moisture: 

mesic. The edges of the draw have 

scattered spruce and aspen growing 

in patches. 


King, and T. 

Price (BOSI-9) 

Lawrence Long Draw 6,160 6/17/2004 6/17/2004 USFS 

Black Hills 


29 South 

Dakota 

In an open grass/forb area on the 

edge of a shallow depression; near a 

small ponderosa pine in an area with 

other small ponderosa pine saplings; 

area is grazed; moisture: dry-mesic 

N/A All individuals had 

already sporulated 


King, and T. 

Price (BOSI- 

13) 

Lawrence Meadow Creek 4,600 6/23/2004 6/23/2004 USFS 

Black Hills 


30 South 

Dakota 

At the base of an open slope in a 

grass/forb dominated area; there 

was a skid trail on the slope above 

this occurrence; the slope is fairly 

steep with small shrubs starting to 

emerge 

N/A Possibly two varieties 

of Botrychium simplex 

present; B. pallidum 

also occurs at this 

location 

13 B. Burkhart, 

C. Mayer, 

C. Skelton, 

D. Reyher, 

D. Farrar, 

D. King, R. 

Crook, S. 

Popovich, 

and T. Price 

(BOTR-4) 

6/9/2004 7/13/2004 USFS 

Black Hills 


Lawrence Higgens Gulch 5,480 to 

5,520 

31 South 

Dakota

Date first Date last Land ownership/ 

Collector / 

observed observed management Abundance Observer Herbarium Notes Habitat 

6/10/2004 6/22/2004 USFS 

1 C. Mayer, N/A Sporangia were still In partial shade in an open grassy 

Black Hills 

C. Skelton, 

green on June 10, but meadow; an open old road bed 


D. Farrar, D. 

were yellowing by passes through the meadow where 

King, and T. 

June 22; Botrychium some weeds (Taraxacum sp., 

Price (BOTR- 

pallidum and B. Trifolium) occur 

5) 

michiganense are also 

found at this location 

5/26/2005 6/7/2005 USFS 

26 B. Burkhart, N/A Other Botrychium Open meadow of Black Hills 

Black Hills 

C. Mayer, 

species were seen at Montane Grassland with native 


S. Corey, 

this location but the species composition; area is 

I. Drieling, 

identification is not yet very flat; minor landforms seem 

C. Buckert, 

verified 

important; area has been heavily 

and K. 

grazed; limestone parent material 

Zacharkevics 

(BOTR-31) 


Elevation 

(ft.) 


Lawrence Citadel Rock 5,200 to 

5,240 

32 South 

Dakota 

Pennington Four Corners 6,560 to 

6,600 

33 South 

Dakota 

Open rolling grassland with 

fairly heavy grazing. High forb 

N/A Likely that many 

more individuals 

are in the vicinity. 

Other Botrychium 

species are present at 

this site but isozyme 

analysis is required 

to identify them. This 

is a very rich site for 

Botrychium species. 

~75 B. Burkhart, 

C. Mayer, 

S. Corey, 

I. Drieling, 

C. Buckert, 

and K. 

Zacharkevics 

(BOTR-30) 

6,600 6/8/2005 6/8/2005 USFS 

Black Hills 


Pennington Redbank 

Spring 

34 South 

Dakota 

diversity. Shrubby cinquefoil 

scattered frequently on the hillside. 

Limestone parent material. 

Moisture: mesic. Topographic 

position: lower slope. 

N/A Not yet sporulating On an old mossy skid trail at the 

base of a northeast facing slope; 

spruce dominate the edges of the 

skid trail; in partial shade of a young 

spruce tree upslope of the plant 

1 D. King 

(BOTR-19) 

6,600 7/21/2004 7/21/2004 USFS 

Black Hills 


Pennington Coulsen 

Hughes Draw 

35 South 

Dakota 

In Sibbaldia clumps and bare soil 

between road and willow carr edge 

Unknown P. Root (90-70) KH Western type. 

Trophophores slightly 

concave but basal 

segments strongly 

folded; two parts of 

leaf separated at or just 

below soil surface 

36 Wyoming Carbon Slash Ridge ~9,600 8/4/1990 8/4/1990 USFS 

Medicine Bow 

National Forest



Collector / 


6/30/2003 7/2/2004 USFS 

3 C. Mayer, N/A Botrychium simplex Mossy, grassy area under a thin 

Black Hills 

R. Crook, 

has been found in three cover of bur oak and Pinus 


D. King, D. 

areas at this location; ponderosa; moist to dry-mesic, 

Farrar, M. 

B. michiganense is sandy soil; plants are found among a 

Gabel, K. 

also present at this stand of 1 to 3 foot tall aspen trees; 

Zacharkevics, 

location 

forbs are scattered, grasses small but 

S. Corey, 

nearly continuous in places; bare 

A. Kratz, 

ground is covered occasionally by 

C. Skelton, 

pine needles 

D. Rarrar, 

D. King, S. 

Popovich, 

and T. Price 

(BOTR-1, 

BOTR-2) 

N/A Open sunny roadside meadow with 

Pinus ponderosa at margins 

Elevation 

(ft.) 


4,670 to 

4,680 

37 Wyoming Crook Bearlodge 

Campground 

1 R. Crook, 

D. Farrar, C. 

Mayer, and D. 

King 

38 Wyoming Crook FS Road 830 4,730 6/2/2003 6/2/2003 USFS 

Black Hills 


Near and on the remnants of an 

old road bed; in an open site in 

grass/forb vegetation near a stand 

of medium sized ponderosa pines; 

in a flatter area on the upper slope 

of a north-facing hill; the site is 

N/A In early sporulation 

on June 11; one plant 

sporulating on July 2; 

Botrychium pallidum 

is also found at this 

location 


C. Skelton, 

D. Rarrar, 

D. King, R. 

Crook, and T. 

Price (BOTR- 

6) 

6/11/2004 7/2/2004 USFS 

Black Hills 


39 Wyoming Crook Beaver Creek 4,900 to 

4,940 

currently grazed 

On a steep, north-facing slope 

dominated by native forbs; the 

ground is covered in moss and rocks 

N/A Sporangia are 

immature 

7 D. King, C. 

Skelton, and T. 

Price (BOSI- 

14) 

6/25/2004 6/25/2004 USFS 

Black Hills 


40 Wyoming Crook Warren Peak 6,500 to 

6,600 

where vegetation cover is sparse; 

plants are in an open area with a 

little shade provided by a large 

ponderosa pine; area is grazed


Collector / 


6/21/2004 7/8/2004 USFS 

207 C. Skelton, D. N/A Large variety in stage On a toe slope between two 

Black Hills 

King, T. Price, 

of maturation seen on different drainages; there is a 


and P. Sweanor 

June 21; plants had vague native surface old road bed 

(BOSI-12) 

sporulated on July 8; (currently unused) going along 

both “eastern” and the toe slope where the plants are 

“western” varieties growing; with grasses and small 

were seen; specimens ponderosa pines; area is grazed. 

are being verified by 

D. Farrar 


Elevation 

State County Location (ft.) 

41 Wyoming Crook Dugout Gulch 4,400 to 

4,500 

In an old open unused road bed 

with very little tree cover; plant 

N/A Plant could not be 

found again on July 2 


C. Skelton, 

D. Farrar, 

D. King, R. 

Crook, and T. 

Price (BOSI-6) 

42 Wyoming Crook Fawn Creek 4,920 6/11/2004 7/2/2004 USFS 

Black Hills 


is growing in an area covered by 

needle litter; area is grazed 

Coniferous forest, grassy ridge, and 

high basin with internal drainage; 

RM Var. tenebrosum; 

sporulating 

Unknown R.L. Hartman 

(18845) 

8/17/1984 8/17/1984 USFS 

Shoshone National 

Forest 

9,400 to 

10,000 

43 Wyoming Fremont Vicinity of 

Deacon Lake 

grassy slopes 

Grass and moss-rich hummocks 

along banks of small stream in 

marsh at edge of Pinus contorta/ 

Picea engelmannii woods 


sporulating 

Unknown W.Fertig 

(19618) 

8,100 6/25/2001 6/25/2001 USFS 

Bighorn National 

Forest 

44 Wyoming Johnson French Creek 

Swamp 

Among talus boulders with Arnica 

and Carex 


sporulating 

“Rare” R.W. Lichvar 

(3923) 

45 Wyoming Lincoln Corral Creek 9,400 9/17/1980 9/17/1980 USFS 

Bridger-Teton NF 

Mouth of creek 

Cited in Clausen 

(1938) 

GH, UC, 

NY, PH, 

RM, MO 

Unknown C.C. Parry 

(306) 

~7,740 1873 1873 NPS 

Yellowstone 

National Park 

46 Wyoming Park Yellowstone 

Lake, Mouth of 

Pelican Creek 

Riparian areas adjacent to Picea 

engelmannii and Abies lasiocarpa 


sporulating 

forest; moist creek banks 

Unknown R.S. 

Kirkpatrick 

(5317a) 

and R.E.B. 

Kirkpatrick 

47 Wyoming Park Kirwin 9,300 8/13/1984 8/13/1984 USFS 


Forest 

RM Rocky outcrops 


and B.E. 

Nelson 

(20707) 

7/14/1985 7/14/1985 USFS 


Forest 

48 Wyoming Park Hayden Creek 8,600 to 

9,400



Collector / 


7/18/1985 7/18/1985 USFS 

Unknown R.L. Hartman RM Moist north exposure at edge of 


(21051) 

Douglas-fir forest 

Forest 

Elevation 

(ft.) 


6,900 to 

7,800 

49 Wyoming Park Little Bald 

Ridge 

RM Limestone outcrops and adjacent 

slopes 


(21880) 

8/17/1985 8/17/1985 USFS 


Forest 

8,600 to 

9,600 

50 Wyoming Park Ridge 

Northeast 

of Windy 

Mountain 

In moss in montane forest 

RM Determined as 

Botrychium lunaria 

Unknown E.F. Evert 

(4619) 

7,600 8/3/1982 8/3/1982 USFS 


Forest 

51 Wyoming Park Along West 

Blackwater 

Creek Trail 

Not reported 

RM Determined as 

Botrychium lunaria 

Unknown E.F. Evert 

(4594) 

52 Wyoming Park Kitty Creek 7,200 8/1/1982 8/1/1982 USFS 


Forest 

YELLO About 30 meters above Columbia 

Pool; east aspect; 10 percent slope; 

adjacent to hot ground that is too 

hot to support vascular plants; also 

on north aspect; 5 percent slope 

on edge of barren sinter sheet and 

Botrychium lunariawetland west of 

Rustic Geyser 

125 J.J. Whipple 

(4474, 4478) 

7,480 6/27/1995 6/27/1995 NPS 

Yellowstone 

National Park 

53 Wyoming Teton Heart Lake 

Geyser Basin 

YELLO In a meadow near “bobby sox trees” 

(see Definitions for explanation) 

in areas that are less saturated than 

other areas in the meadow 

J.J. Whipple 

(4263, 4264) 

Several 

hundred 

7,240 6/24/1994 6/5/2003 NPS 

Yellowstone 

National Park 

54 Wyoming Teton Lower Geyser 

Basin 

Wetland 

ID is dubious; 

annotated by Don 

Unknown W. Gernon (7) RM, 

YELLO 

~8,000 8/10/1978 8/10/1978 NPS Yellowstone 

National Park 

55 Wyoming Teton Madison 

Plateau 

Despain in 1978 

but consists of one 

misshapen lamina 

(Whipple personal 

communication 2003)



Collector / 

(ft.) observed observed management Abundance Observer Herbarium Notes Habitat 

~8,950 8/20/1979 8/20/1979 NPS 

Unknown T. Caprio (s.n.) YELLO At least one of Not reported 

Yellowstone 

the three plants 

National Park 

on this sheet is 

Botrychium simplex 

(Whipple personal 



56 Wyoming Teton Northwest side 

of Mariposa 

Lake on Two 

Ocean Plat 

Along abandoned road in roadbed 

where asphalt has been removed 

3 J.J. Whipple N/A No collection 

made due to small 

population size; not 

seen again at this site 

despite repeated visits 

7,300 6/7/1994 6/7/1994 NPS 

Yellowstone 

National Park 

57 Wyoming Teton Upper Geyser 

Basin near Old 

Faithful 

RM Var. tenebrosum Open, moist clay soil below white 

limestone boulders at timberline; 

community of Antennaria media/ 

Phlox pulvinata with 60 percent 

vegetative cover; sporulating 

10,000 8/7/1997 8/7/1997 Unknown Unknown W.Fertig 

(17953) 

58 Wyoming Teton West slope 

of Corner 

Mountain 

Herbarium acronyms: 

COCO: Carter Herbarium, Colorado College 

COLO: University of Colorado Museum Herbarium 

GH: Gray Herbarium, New York 

ISC: Ada Hayden Herbarium 

KHD: Kalmbach Herbarium, Denver Botanic Gardens 

MO: Missouri Botanical Garden 

PH: Academy of Natural Sciences, Philadelphia 

RM: Rocky Mountain Herbarium 

SJNM: San Juan College Herbarium, Farmington, NM 

UC: University of California Berkeley 

YELLO: Yellowstone National Park Herbarium

Table 9. Land ownership / management status summary for the 50 known occurrences of Botrychium simplex within 

the administrative boundary of USDA Forest Service Region 2. 

Land Ownership Status Number of Populations Subtotals 

USDA Forest Service 41 

Bighorn National Forest 1 

Black Hills National Forest 16 

Gunnison National Forest 1 

Medicine Bow National Forest 1 

Pike National Forest 2 

Rio Grande National Forest 2 

Roosevelt National Forest 3 

San Isabel National Forest 1 

San Juan National Forest 1 

Shoshone National Forest 7 

White River National Forest 6 

National Park Service 2 

Rocky Mountain National Park 2 

Bureau of Land Management 2 

State of Colorado 1 

Dome Rock State Wildlife Area 1 

City of Denver 1 

Echo Lake Park 1 

The Nature Conservancy 1 

Niobrara Valley Preserve 1 

Unknown 2 

forests. Yellowstone National Park (outside Region 2) 

and Rocky Mountain National Park include six and two 

occurrences, respectively. 

In Colorado, Botrychium simplex is known 

from 24 locations in 14 counties, but data are sparse 

for most occurrences. Almost no data are available 

for occurrences in Gunnison, Jackson, and Saguache 

counties; these locations have not been verified. 

Arnett (2002) reported a collection of B. simplex from 

Gunnison County, but a specimen or other confirmation 

has not been found. A report from Jackson County (“on 

the west side of Cameron Pass”) is mentioned in a 

species abstract draft for B. simplex (Schwab 1992), but 

no specimen or other ancillary information exists for 

this report. Popovich (personal communication 2006) 

and Proctor (personal communication 2006) searched 

suitable habitats near Cameron Pass but did not find 

B. simplex. It now appears that this report arose from a 

1986 observation initially identified as B. simplex, but 

later determined to be B. hesperium (Farrar personal 

communication 2006). Saguache County is included in 

the range for B. simplex by the University of Colorado 

Herbarium (2002), but a corresponding specimen 

32 

for this occurrence has not been found. Two other 

occurrences lacking vouchers are in Clear Creek and 

Lake counties. 

Occurrences in Colorado are patchy, with clusters 

in the vicinity of Pikes Peak, Breckenridge, and 

Granby. Two clusters are in the San Juan Mountains. 

The patchy nature of the Colorado distribution of 

Botrychium simplex is likely to be the result of survey 

intensity because in two cases (around Breckenridge 

(#17 in Table 8) and Granby (#8 and 9 in Table 8)) 

the occurrences were documented during one survey or 

by one person. In areas such as Pikes Peak, numerous 

collectors were involved, but most used known locations 

as the start of their search. 

Three occurrences of Botrychium simplex have 

been documented in the Pikes Peak area in El Paso 

and Teller counties (#6, 23, and 24 in Table 8). The 

occurrence on Pikes Peak that is documented in 

Wherry (1937) was found on the south side of the peak 

(Root personal communication 2003). A moonwort 

survey was conducted on the north side of Pikes 

Peak in 2001; numerous moonworts were found, but

Figure 10. The distribution of Botrychium simplex in the states of USDA Forest Service Region 2. Table 8 is a 

complete summary of the known occurrences in Region 2. 

no B. simplex (Steinmann 2001a, Steinmann personal 


Recent work by Dave Steinmann and Peter Root 

(Steinmann 2001b, Root personal communication 

2003, Steinmann personal communication 2003) 

identified numerous moonwort occurrences in the 

subalpine zone in the Indian Peaks Wilderness Area 

of Colorado. Two occurrences of Botrychium simplex 

are known from Boulder County, one of which was 

discovered in 2001 as a result of this work (#2 in Table 

8). Botrychium simplex was much less commonly 

found than other species including B. lunaria, B. echo, 

B. hesperium, and B. minganense (Steinmann personal 


Annette Kolb, Nancy Redner, Peter Root, Tony 

Spribille, and others conducted extensive surveys for 

Botrychium species at Copper Mountain, Breckenridge, 

33 

and Keystone ski resorts in Summit County, Colorado. 

Their work identified five new occurrences of B. simplex 

in which Kolb and Spribille (2000) estimate a total of 85 

stems (#15, 17, 18, 21, and 22 in Table 8). Thompson 

(2001) presents data on moonwort aggregations in 

Summit County. Aggregations were observed to range 

in size from a single plant to 1.43 acres, with up to 590 

individuals in single species aggregations and as many 

as 1,717 individuals in aggregations containing multiple 

moonwort species. Botrychium simplex was the rarest 

moonwort observed in these surveys relative to B. 

lanceolatum, B. echo, B. minganense, B. hesperium, B. 

lunaria, B. pallidum, and B. pinnatum. 

No population estimates exist for 14 of the 

24 occurrences in Colorado, and based on the 10 

occurrences where there is some indication of 

abundance, the total population for Colorado is between 

100 and 200 individuals (Table 8).

In Wyoming, Botrychium simplex is known from 

23 occurrences in seven counties (Table 8). Wyoming 

Natural Diversity Database has not tracked B. simplex 

since 1986, so they have not maintained information 

for this species (Heidel personal communication 

2003). Six of the Wyoming occurrences are from 

Yellowstone National Park, and another is on the 

Bridger-Teton National Forest in Region 4. The 

other 16 occurrences fall within the administrative 

boundaries of USFS Region 2. Seven occurrences 

are located on the Shoshone National Forest, east of 

Yellowstone National Park; these combined with the 

six on Yellowstone National Park represent the largest 

concentration of occurrences in Wyoming. Walt Fertig 

(2003) discovered B. simplex at French Creek Swamp 

on the Bighorn National Forest in 2001. One occurrence 

is also known from the Medicine Bow National Forest 

(#36 in Table 8). 

In the Black Hills National Forest in Wyoming, 

two occurrences (#37 and 38 in Table 8) of Botrychium 

simplex were discovered in the Bearlodge Mountains 

of Crook County in 2003. Only one plant was found 

at each site. One of these locations, the Bearlodge 

Campground, also contained B. campestre and B. 

“michiganese” and has become notorious for moonwort 

species (Crook personal communication 2003). In 2004 

and 2005, Farrar, his graduate students, and USFS 

botanists conducted intensive surveys of the Black Hills 

National Forest in Wyoming and discovered four more 

occurrences (Burkhart personal communication 2006). 

The largest occurrence of Botrychium simplex in 

Region 2 was one of those discovered in 2004. Located 

at Dugout Gulch in Crook County, this population was 

estimated to contain 207 individuals (Burkhart personal 

communication 2006). Of the 15 Wyoming occurrences 

in Region 2, populations have been estimated for only 6 

and these total 279 individuals. 

Before 2004 Botrychium simplex was known from 

only a single occurrence in South Dakota, a population 

in Lawrence County that had not been seen since 1953 

(#27 in Table 8). This is probably the population 

that is reported for the Black Hills of South Dakota 

by Dorn and Dorn (1972). Ode (2001) mentions the 

existence of another historic occurrence in northeastern 

South Dakota, but no detailed information is available 

regarding this occurrence. As in Wyoming, intensive 

surveys of the Black Hills National Forest in 2004 and 

2005 resulted in the discovery of nine new occurrences 

of B. simplex in South Dakota. The estimated abundance 

for occurrences in South Dakota is 140 individuals. 

34 

In Nebraska, Botrychium simplex is known from a 

single report from Brown County at the Niobrara Valley 

Preserve owned by The Nature Conservancy (Farrar and 

Johnson-Groh 1986, Farrar personal communication 

2003; #25 in Table 8). Here it is described as being 

“fairly common” (Farrar personal communication 2003), 

but there are no data on the size of this occurrence. The 

Ada Hayden Herbarium (ISC) houses a collection from 

this occurrence, but no other reports of B. simplex are 

known from Nebraska (Bolick personal communication 

2003, Farrar personal communication 2003, Lewis 

personal communication 2003, Rolfsmeier personal 

communication 2003). There are no specimens of B. 

simplex at the University of Nebraska State Museum 

Herbarium (NEB) (Bolick personal communication 

2003). Farrar attempted unsuccessfully to find B. 

simplex in the Niobrara Valley Preserve in 2004 (Farrar 

personal communication 2006). 

Botrychium simplex has not been documented 

in Kansas, and it is doubtful that it will be found there 

(Farrar personal communication 2003). 

Population trend 

There are no rigorous quantitative data on 

population trend for Botrychium simplex in Region 

2, and the available data are too sparse to determine 

whether populations have increased or decreased. 

For species such as B. simplex where the proportion 

of dormant plants varies among years, it is difficult 

to accurately monitor population trends (Lesica and 

Steele 1994). 

The recent discovery of many occurrences in 

Region 2 is probably the result of an increase in interest 

in this species and its congeners rather than an increase 

in population. Moonworts are now known to be much 

more widespread and abundant on the Black Hills 

National Forest (Burkhart personal communication 

2006) and in Summit County, Colorado (Thompson 

2001) than previously thought. Additional searches are 

likely to discover more occurrences. 

Many locations of Botrychium simplex in Region 

2 are known only from very old herbarium specimens. 

Because of the uncertain location of these collections, 

it is difficult to determine whether they are extant or 

whether protective management should be implemented 

on their behalf. Seventeen of the 50 known occurrences 

were last visited more than 20 years ago. The failure 

to observe any occurrence in Region 2 more than once 

may herald a decline of this species, but it is probably 

more indicative of the difficulties in finding it.

It is not known how Botrychium simplex responds 

to and recovers from drought. A severe regional drought 

beginning in 2001 and extending into 2003 may have 

reduced populations of B. simplex in Region 2. The 

degree to which B. simplex depends on years that 

are particularly favorable (perhaps with respect to 

precipitation) for successful reproduction is not known. 

There is evidence to suggest that Botrychium 

simplex has declined elsewhere in its range. 

Botrychium simplex is believed to have been extirpated 

from Connecticut, Virginia, Maryland, and possibly 

Ohio (Kartesz 1999, USDA Natural Resources 

Conservation Service 2002). Botrychium simplex is 

apparently increasing within Region 9 (Chadde and 

Kudray 2001b). 

The quality and amount of wetland habitat for 

Botrychium simplex has certainly decreased in portions 

of the states of Region 2, but this decline in habitat 

has been less severe at higher elevations where most 

Region 2 B. simplex occurrences are found. Since 

1986, wetlands have been lost at a rate of 58,500 acres 

per year in the continental United States (Dahl 2000). 

In Colorado alone, an estimated one million acres of 

wetlands (50 percent of the total for the state) were 

lost before 1980 (Dahl 1990). In total, estimated losses 

from all Region 2 states is approximately 39 percent of 

the original wetland acreage (Dahl 1990). Not included 

in these numbers is the loss and degradation of nonjurisdictional 

wetlands (i.e., those wetlands not regulated 

by Section 404 of the Clean Water Act), such as riparian 

areas. It is unknown what percentage of the wetlands 

lost in Region 2 would be considered B. simplex habitat. 

In some cases, anthropogenic disturbance may have 

even created habitat for B. simplex (Thompson 2001). 

This does not, however, suggest that human disturbance 

can be relied upon for the conservation of this species. 

See the following Habitat section for information on 

wetland and other habitats where B. simplex occurs. 

Habitat 

Habitat descriptions and characterization 

Botrychium simplex grows in a variety of 

habitats and conditions and has been noted for its broad 

ecological amplitude (Chadde and Kudray 2001b). 

Hitchcock (1823) originally described B. simplex from 

plants found in “dry hilly pastures,” but it has since 

been found in a wide range of other habitats. Lellinger 

(1985) describes its habitat as “Terrestrial in meadows, 

barrens, and woods, usually in subacid soil.” Wagner 

and Wagner (1993) describe the habitat as “dry fields, 

35 

marshes, bogs, swamps, and roadside ditches.” Rydberg 

(1922) broadly defines the habitat as grassy places and 

open woods. In general, moonworts tend to grow in 

places that are unpromising to botanists (Wagner and 

Wagner 1983, Root personal communication 2003). 

In Michigan, Minnesota, and Wisconsin, 

Botrychium simplex has been reported from both open 

and closed canopy settings. These include rich black 

ash (Fraxinus nigra) and cedar (Thuja occidentalis) 

swamps, jack pine (Pinus banksiana) woods, prairies, 

an open area dominated by reed canary grass (Phalaris 

arundinacea), an open field with non-native grasses, 

stands of northern hardwood forest, and on glacial 

till and outwash. It is also reported from sites that 

have been disturbed by humans, including borrow 

pits, tailings ponds, road shoulders, and old roadbeds 

(Chadde and Kudray 2001b, Burkhart personal 

communication 2006). Botrychium simplex is known 

from prairie habitats in Iowa and Michigan (Johnson- 

Groh 1999). Only three species of moonworts (B. 

simplex, B. campestre, and B. gallicomontanum) are 

restricted to prairie habitats. In Idaho, Davis (1952) 

reports B. simplex from dry woods and meadows 

while Lorain (1990) describes its habitat as the shaded 

understory of western redcedar/oakfern forests. In 

California, it is known from open meadows and damp 

places (Munz and Keck 1968), and it is “always in open 

grassy areas, often more or less marshy” and “in damp 

meadows with sedges (including Carex aurea), grasses, 

and Mimulus primuloides” (Wagner and Devine 1989). 

In Utah, it is known from “moist to somewhat dry 

woods and open slopes at 2,300 to 3,500 m” (Welsh 

et al. 1993). In Montana, B. simplex is known from 

disturbed seral lodgepole pine forests (Mantas and 

Wirt 1995). In Ohio, B. simplex has been documented 

from moist, shaded situations including weedy thickets 

and mesophytic woods (Ohio Department of Natural 

Resources 2003). Herbarium specimens describe other 

habitats, including cold barren knolls in old pastures 

(Vermont); upper lake beach in shade (Ontario); 

cold, damp woods (Maine); and alpine meadows and 

meadow-forest edge (Oregon). 

Region 2 habitat descriptions 

The variability of habitat for Botrychium simplex 

in Region 2 is as great as its morphological variability, 

making it very difficult to characterize its habitat. It is 

found not only in typical moonwort habitat, but also in 

shaded sites and in wetlands, where no other moonworts 

that have been documented in Region 2 are found. Thus, 

the ecological amplitude of B. simplex is probably 

broader than any other moonwort in Region 2. Habitats

documented in herbarium specimens range from “dry, 

open northwest slope of tuff” to “wet soil of peat 

bog” to railroad right-of-way. Farrar (2005) describes 

B. simplex as “a plant of open habitats, occurring in 

pastures, meadows, orchards, prairies, wetlands, fens, 

sand dunes, and in lake and stream edge vegetation,” 

and further states that “most of its habitats are at least 

temporarily wet and some (fens) are permanently 

saturated.” Harrington (1954) reports B. simplex from 

pastures, meadows, and gravelly slopes of open places. 

Schwab (1992) notes that it is usually found in moist 

meadows, gravel slopes, marshy areas, streamsides, 

and open places. Arnett (2002) reports finding B. 

simplex in spruce fir forest and subalpine meadows. 

Cronquist et al. (1972) report it from “moist to rather 

dry meadows in the mountains, apparently not above 

timberline.” Botrychium simplex was found in 2005 in 

open shrublands dominated by Dasiphora floribunda 

(shrubby cinquefoil) in South Dakota (Figure 11). Table 

8 has a summary of habitat descriptions documented for 

Region 2 occurrences. 

Recent surveys have shed light on local variation 

in the kinds of habitats where Botrychium simplex 

is found. In the Black Hills, it was most often found 

on toeslopes adjacent to riparian greenbelts in deep 

36 

soils, and also on abandoned roadbeds (Burkhart 

personal communication 2006, Popovich personal 

communication 2006). In Colorado, it is more often 

reported from wetter sites near streams, but it has also 

been found in seasonally dry sites. Popovich (personal 

communication 2006) speculates that the greater annual 

precipitation in the Black Hills permits B. simplex 

to persist in upland settings (though still typically in 

swales or proximal to a riparian area or floodplain) 

while it is more tightly constrained by water availability 

in Colorado. 

Varietal differences in habitat 

The varieties described for Botrychium simplex 

have significant differences in their habitat affinities. 

The distribution of these varieties in Region 2 may 

account for the diversity of habitats reported for B. 

simplex. For example, B. tenebrosum was described 

from plants growing in deep shade in maple swamps 

(Eaton 1899). Similarly, Wagner and Wagner (1983) 

characterize the habitat for var. tenebrosum as “shaded 

bog edges.” This habitat description does not fit plants 

in Wyoming that fit the physical description of var. 

tenebrosum. Var. typicum (reported in Colorado by 

Harrington (1954)) is known from dry upland fields 

Figure 11. Habitat of Botrychium simplex at Redbank Spring, South Dakota on the Black Hills National Forest 

(BOTR-30). This area was a rich site for Botrychium species in 2005. The shrubs are Dasiphora floribunda. Photo by 

Black Hills National Forest, provided by Beth Burkhart.

(Wagner and Wagner 1983). Wagner and Wagner 

(1983) characterize the habitat for var. compositum as 

moist low meadows. 

Farrar (2005) notes that varietal habitat 

segregation exists and describes general habitat affinities 

for the four varieties he circumscribed. Var. compositum 

is frequently found in meadow and roadside habitats in 

Colorado and is “often the most common, and frequently 

the only species of moonwort present in mountain 

meadow sites throughout the western mountains” 

(Farrar 2005). Farrar (2005) notes that var. compositum 

may become especially common in alluvial meadows 

derived from granitic substrates. Var. simplex is known 

from “meadow woodlands” throughout its range. Var. 

tenebrosum is found in forests, swamp margins, and in 

dune complexes of the northeastern U.S., Greenland, 

and Canada. In northeastern North America it is most 

commonly present in permanently wet, often deeply 

shaded habitats. Var. fontanum occurs in fens and in 

calcareous seeps, but it is not yet known from within 

Region 2. Table 3 is a summary of the distinguishing 

characteristics of these varieties. 

Elevation, slope, and aspect 

Wagner and Wagner (1993) reported a maximum 

elevation of 7,200 ft. for Botrychium simplex. However, 

most occurrences within Region 2 are found at higher 

altitudes. The elevation of the known occurrences of 

B. simplex in Region 2 ranges from approximately 

2,400 ft. along the Niobrara River, where it occurs in a 

floodplain with cottonwood (Populus c.f. deltoides ssp. 

monilifera) and red cedar (Juniperus virginiana) (Farrar 

and Johnson-Groh 1986, Farrar personal communication 

2003), to 12,680 ft. in tundra. Occurrences in Colorado 

are found at higher elevations (8,500 to 12,680 ft.) 

than those in Wyoming (4,670 to 10,000 ft.) and South 

Dakota (approximately 4,500 ft.). 

Botrychium simplex has been documented from 

flat sites as well as from slopes as steep as 35 percent in 

Region 2. It has been found on wasting scree and talus 

slopes and sloping open soil in the alpine (Colorado 

Natural Heritage Program 2006). Most alpine species 

cannot tolerate the chronic disturbance regime in these 

sites. Botrychium simplex is also commonly found on 

flat sites that may or may not be disturbed. 

Botrychium simplex does not appear to favor 

any particular aspect. In two instances, B. simplex 

occurs on south-facing exposures in both alpine and 

arctic habitats. One site is in the alpine of Colorado in 

37 

Conejos County. It has also been found on south-facing 

slopes in Greenland (Ollgaard 1971). The warming 

effects of greater insolation on south-facing slopes are 

well studied (Barbour et al. 1987) and these may allow 

B. simplex to persist at elevations and latitudes outside 

its normal range. This phenomenon has often been 

observed in the alpine and arctic (e.g., Bliss 1987). 

Soil 

Botrychium species are often found in alkaline, 

calcium-rich soils (Root personal communication 

2003). Occurrences of B. simplex within Region 2 

have been reported in calcareous soils at a variety of 

elevations. Botrychium simplex is also known from 

subacid or acid soils that are high in organic matter in 

Region 2 and elsewhere (Lellinger 1985, Steinmann 

personal communication 2003, Colorado Natural 

Heritage Program 2006). Botrychium simplex has been 

found in acidic soils associated with mines outside 

Region 2 (Chadde and Kudray 2001b). 

Botrychium simplex also occurs on a range of soil 

textures in Region 2, including cobbly, gravelly, sandy, 

and clay loam. It is documented from riparian and 

wetland sites where soils tend to be silty (e.g., marshy 

areas, willow carr edge). In Montana, plants occur 

in open areas with shallow soils, which apparently 

caused stunted growth (Mantas and Wirt 1995). The 

suitability of germination sites varies with microscale 

heterogeneity of soil conditions, which probably causes 

the patchy distribution of Botrychium species (Johnson- 

Groh et al. 2002). 

Moisture 

Habitat descriptions for Region 2 occurrences 

range from “dry slope” and “along railroad tracks 

in dry site” to “in marshy area” and “wetland.” This 

suggests that while habitat for Botrychium simplex 

in Region 2 might be constrained to some extent by 

water availability, the species has a broad tolerance 

of soil moisture conditions. Botrychium simplex is 

often found in wetlands and is a wetland indicator 

species in Region 2 (Table 10; U.S. Fish and Wildlife 

Service 1988), but its fidelity to wetland habitats is 

much lower than that of B. multifidum, with which it 

often grows. Botrychium simplex is more common in 

eastern North America where soils tend to be wetter. 

Although numerous herbarium specimen labels and 

other reports note dry soil conditions, most sites are at 

least seasonally wet. Kolb and Spribille (2000) report 

that sites in Summit County where B. simplex was

Table 10. Wetland Indicator Status for Botrychium simplex (U.S. Fish and Wildlife Service 1988). 

Region Geographic Areas in Region Wetland Indicator Status 

North Plains MT (Eastern), ND, SD, WY (Eastern) FAC 

Central Plains CO(Eastern), NE, KS FAC 

Intermountain CO (Western), NV, UT FACU 

Northwest ID, OR, MT (Western), WA, WY (Western) FACU 

Facultative (FAC): Equally likely to occur in wetlands or non-wetlands (estimated probability 34 to 66 percent). 

Facultative Upland (FACU): Usually occurs in non-wetlands (estimated probability 67 to 99 percent), but occasionally found on wetlands 

(estimated probability 1 to 33 percent). 

found were dry at the time. There are no soil moisture 

data available from which the physiological tolerances 

of B. simplex to desiccation could be determined. 

Disturbance as a habitat attribute 

Although information on the disturbance regime 

in Region 2 occurrences of Botrychium simplex is 

sparse, some observations (within and outside of 

Region 2) suggest that disturbance assists in the creation 

and maintenance of suitable habitat. Some disturbed 

sites in which B. simplex has been found in Region 2 

include scree slopes, subalpine forest clearcuts, “Open 

eroded grassy slopes at timberline in historic burn,” 

ski slopes, roads, abandoned roadbeds, floodplains 

and riparian areas, and railroad right-of-ways (Table 

8). Five of the sites where it was found on the Black 

Hills National Forest in 2004 and 2005 are abandoned, 

unused roadbeds (Burkhart personal communication 

2006). It has also been reported from disturbed sites 

outside of Region 2, including pastures, tailings ponds, 

borrow pits, and road shoulders (Chadde and Kudray 

2001b). There is a general tendency to find Botrychium 

species in sites that were disturbed approximately 10 

years previously (Johnson-Groh and Farrar 2003). 

Buell (2001) noted that Botrychium species were most 

commonly found in Summit County on ski runs that had 

been created more than 30 years previously, and seldom 

on newer ski runs. 

In Yellowstone National Park, Botrychium simplex 

appears to be strongly associated with geothermallyinfluenced 

meadows (Whipple personal communication 

2003). These meadows usually support interesting 

and atypical plant assemblages. They are probably 

maintained by a mild disturbance regime imposed by 

geothermal activity (Whipple personal communication 

2003). They tend to be slightly disturbed but not recently 

disturbed, and competitive species are not common. It is 

interesting to note that B. simplex also shows an affinity 

for hot springs in Greenland, where it was documented 

from a site where the soil is warmed by the spring 

(Ollgaard 1971). This suggests a general affinity for 

38 

geothermal features. The Reproductive biology and 

autecology section of this document provides a more 

detailed treatment of the ecological role of disturbance 

for B. simplex. 

Fire 

Botrychium simplex was documented in forests 

dominated by Pinus contorta (lodgepole pine) and 

P. ponderosa (ponderosa pine) in Region 2. Both are 

fire-adapted species, and fire is a natural and relatively 

frequent part of forests dominated by these species. 

Stand ages of subalpine lodgepole forests south of 

Yellowstone suggest a 200 to 400 year fire interval, but 

at lower elevations the interval may be as short as 50 to 

150 years (Peet 2000). The role of fire in the ecology 

B. simplex is unclear, but there is no evidence that it 

has an affinity for newly burned areas. Although B. 

simplex is associated generally with these forest types, it 

typically grows in streamsides or meadow edge habitats 

where burning may be less frequent or intense. Most 

occurrences of B. simplex known from Region 2 are 

found in or near subalpine forest dominated by Picea 

engelmannii (Engelmann spruce) and Abies lasiocarpa 

(subalpine fir), which have much longer fire-return 

intervals than the aforementioned forest types (Veblen 

personal communication 2003). The Community 

ecology section contains more details on the vegetation 

associated with B. simplex. 

Meadows 

Botrychium simplex is often documented in 

meadows and forest openings in Region 2 and elsewhere. 

Meadows are treeless areas dominated by grasses, 

sedges, and forbs and occur throughout the forested 

zones of the Rocky Mountains (Peet 2000). There is little 

agreement on what ecological processes are responsible 

for the creation and maintenance of meadows. Some 

wet meadows in Rocky Mountain National Park are 

maintained by a combination of saturated soils, high 

snow accumulation, cold air drainage, and fine-textured 

soils (Peet 2000). Wet meadows are extremely variable

in their species composition depending on water 

chemistry and water availability. Subtle hydrologic 

gradients result in varying species dominance in wet 

meadows (Wilson 1969). 

Three occurrences of Botrychium simplex are 

in geothermally influenced meadows in Yellowstone 

National Park. These meadows are maintained by 

a mild disturbance regime imposed by geothermal 

activity (Whipple personal communication 2003). 

Like B. multifidum (subgenus Sceptridium), B. simplex 

shows an affinity for these habitats, but it is absent from 

many B. multifidum locations. The Community ecology 

section of this document describes the plants associated 

with B. simplex. 

Mycorrhizae as a habitat attribute 

Most of the life cycle of Botrychium species 

occurs underground, and scientists understand very 

little about this part of its life cycle. Botrychium 

species rely on mycorrhizal interactions in each of 

their life stages (Campbell 1922, Bower 1926, Scagel 

et al. 1966, Gifford and Foster 1989, Schmid and 

Oberwinkler 1994). Johnson-Groh (1999) hypothesizes 

that mycorrhizae are the most important factor in 

the establishment and persistence of Botrychium 

occurrences. Almost nothing is known about which 

species of mycorrhizal fungi interact with Botrychium 

species, what factors affect the mycorrhizal fungi, 

and what factors affect the interaction between the 

mycorrhizal fungi and Botrychium. The Reproductive 

biology and autecology section of this document 

contains a discussion of mycorrhizal interactions 

with Botrychium. 

Reproductive biology and autecology 

In the Competitive/Stress-Tolerant/Ruderal 

(CSR) model of Grime (2001), characteristics of 

Botrychium species most closely approximate those of 

stress-tolerant ruderals. Like many epiphytes, lichens, 

and bryophytes, they are characterized by small stature, 

slow relative growth rates, and small propagules. A 

distinguishing characteristic of plants in this category is 

that stressful conditions are experienced during growth. 

Botrychium species have high reproductive outputs 

(Wagner 1998), which likens them to other “r” selected 

species using the classification scheme of MacArthur 

and Wilson (1967), equivalent to ruderal species in the 

CSR model. The perennial life history and slow growth 

exhibited by Botrychium species are characteristic of 

stress-tolerant species in the CSR model (Grime 2001). 

39 

Moderate to light disturbance may be a critical 

part of the autecology of Botrychium species, including 

B. simplex (Clausen 1938, Lellinger 1985, Wagner and 

Wagner 1993, Barker and Hauk 2003, Johnson-Groh 

and Farrar 2003). The disturbance regime required by B. 

simplex has not been studied and is not well understood. 

Habitat attributes for most moonwort species suggest 

that they depend on a natural disturbance regime 

imposed by wildfires, floods, landslides, or avalanches 

(Alverson and Zika 1996). While these forces are 

clearly at work in most occurrences of B. simplex, some 

occurrences grow in relatively undisturbed sites (e.g., 

“in moss in montane forest”). The role of disturbance in 

the autecology of B. simplex is less clear than for other 

members of subgenus Botrychium. 

In at least three locations in Region 2, 

Botrychium simplex has been documented from sites 

that burned historically. Reed Crook’s speculations 

(personal communication 2003) on the role of fire 

in the reproductive biology of B. multifidum may 

be relevant for B. simplex as well. There are many 

similarities between the life histories of Botrychium 

species and the club mosses (Lycopodium). Both have 

subterranean, mycoparasitic gametophytes that persist 

for years underground before the sporophytes emerge. 

Lycopodium species have been observed to establish 

after fire, possibly due to the presence of mineral soil at 

the surface into which the spores can fall and establish 

underground. Because the gametophytes require 

years to mature, sporophytes are not observed until 

years after the fire and require about 80 years before 

population levels return to normal (Crook personal 


The wetter habitats where Botrychium simplex is 

found are often dominated by competitive species such 

as Carex aquatilis. Observations suggest that B. simplex 

is not particularly competitive. Whipple (personal 

communication 2003) has noted that it is often found 

with typical competitive wetland species, but that it is 

usually found in disturbed sites where these species 

are not dominant. As noted in the CSR model, highly 

competitive species are not successful in stressed or 

disturbed habitats because they allocate much of their 

available resources to growth, and this is maladaptive in 

stressed or disturbed habitats (Grime 2001). 

Reproduction 

The reproductive biology of ferns is very 

different in many respects from that of the flowering 

plants. Like all Pteridophytes, but unlike angiosperms

and gymnosperms, Botrychium spores develop 

into gametophytes that live independently of the 

sporophyte, and the two often have different ecological 

requirements. The gametophyte is haploid and produces 

male and female sex cells in the antheridia and 

archegonia respectively. Male sex cells are motile and 

must move through a fluid environment to fertilize a 

female egg cell. The subterranean nature of Botrychium 

gametophytes probably restricts many Botrychium 

species to self-fertilization (McCauley et al. 1985, 

Soltis and Soltis 1986). 

Upon fertilization, the diploid phase of the life 

cycle begins, and this produces the sporophyte that 

is the life stage most familiar and visible in ferns, 

particularly in Botrychium since the gametophyte is 

subterranean and difficult to observe. In B. simplex, 

several archegonia may be fertilized at the same time, 

but only one embryo develops per gametophyte. 

Several species of Botrychium reproduce 

asexually via gemmae, which are vegetative propagules 

produced by the sporophyte. These bud-like structures 

are borne on the underground portion of the sporophyte 

among its roots and are abscised at maturity (Farrar 

and Johnson-Groh 1990). Reproduction via gemmae 

omits the haploid phase of the life cycle, and it may be 

an adaptation to dry environments, where it is difficult 

for the male gamete to move through the soil (Farrar 

and Johnson-Groh 1990). Reproduction via gemmae 

has been documented in B. pumicola, a close relative 

of B. simplex (Camacho 1996, Camacho and Liston 

2001), and in B. gallicomontanum, for which B. simplex 

is a putative parent species (Farrar and Johnson-Groh 

1991), but it apparently does not occur in B. simplex 

(Farrar and Johnson-Groh 1986). 

Botrychium simplex typically takes approximately 

five years to produce its first emergent leaf, but it may 

be capable of producing a small, aboveground, fertile 

frond in one year (Campbell 1922). This is in contrast 

to B. matricariifolium, which takes 10 years to develop 

an adult sporophyte (Muller 1993, Zika et al. 1995), and 

B. lunaria, which has a relatively small cotyledon and 

may require up to seven years to produce a sporophore 

(Campbell 1922). When given sucrose supplements, B. 

dissectum plants in culture were observed to sporulate 

in one year, but it is doubtful if this ever occurs in nature 

(Farrar and Johnson-Groh 1990, Whittier 1996). The 

gametophyte and its mycobiont are thought to nourish 

the embryonic sporophyte (Camacho 1996). 

40 

Spores are produced by the sporophyte in 

sporangia that are borne on the sporophore. Members 

of subgenus Sceptridium and Osmundopteris are 

capable of producing a trophophore but no sporophore. 

Moonworts, on the other hand, always produce both 

structures when emergent (Vanderhorst 1997). Spores 

are produced by reduction division (meiosis) and are 

capable of producing a gametophyte. They provide 

the only plausible means of long-distance dispersal for 

Botrychium species. 

Phenology 

Johnson-Groh and Lee (2002) divide the 

aboveground portion of the moonwort life cycle 

into four stages: emergence; separation, when the 

trophophore and sporophore separate from each other 

and open; spore release, or sporulation; and senescence. 

The emergence of Botrychium simplex occurs in the 

first week of June in Yellowstone National Park 

(Whipple personal communication 2003) and in the 

Black Hills (Burkhart personal communication 2006) 

(Figure 2). Separation probably occurs shortly after 

emergence, as all herbarium specimens had already 

separated. As observed by Johnson-Groh and Lee 

(2002) for B. gallicomontanum, the separation stage 

is brief in B. simplex while that of B. mormo is more 

protracted. Herbarium specimens from Colorado and 

Wyoming suggest that B. simplex typically sporulates 

from late June (as noted by Hitchcock 1823) through 

July and August, and sometimes into mid-September. 

Botrychium simplex sporulates as early as late May in 

Ohio (Ohio Department of Natural Resources 2003). 

Most plants have senesced by September to early 

October (Whipple personal communication 2003), but 

some may senesce as early as July (Burkhart personal 

communication 2006). Observations of Black Hills 

National Forest occurrences suggest that phenological 

stages vary considerably in their timing even within 

an occurrence. 

Fertility 

Botrychium species produce as many as 20 to 100 

or more sporangia per sporophore, and each sporophyte 

may produce thousands of spores, possibly the highest 

number of spores per case of all vascular plants 

(Wagner 1998). There has been no rigorous assessment 

of the viability of the spores of B. simplex. Spores of B. 

virginianum germinated on agar showed a 90 percent 

germination rate (Peck et al. 1990).

Dispersal 

Dispersal frequency and distance are not 

known for Botrychium simplex, but some inferences 

can be drawn from studies of Botrychium and other 

fern genera. Researchers have hypothesized that the 

dispersal distances for some Botrychium spores range 

from a few centimeters (Casson et al. 1998, Hoefferle 

1999) up to 3 m (Peck et al. 1990). Dyer (1994) found 

that spore banks were largest in soil samples taken 

immediately below ferns and were considerably smaller 

only 2 m away from spore sources. While most spores 

land close to the parent plant, they occasionally are a 

means of long-distance dispersal because some are 

carried considerable distances by the wind (Briggs and 

Walters 1997). 

In addition to wind dispersal, animals may 

disperse Botrychium spores (Wagner and Wagner 1993, 

Wagner personal communication 2002). The spores 

have thick walls that may help to retain their viability as 

they pass through an animal’s digestive tract (Johnson- 

Groh 1998, Wagner personal communication 2002). It 

has been thought that deer and small mammals may 

disperse the spores of forest species such as B. dissectum 

along trails and roads. After feeding the spores of B. 

virginianum to a vole, J.D. Montgomery was able to 

recover them intact from the vole’s droppings (Root 

personal communication 2003). Elk and other ungulates 

that eat the sporophores of B. simplex could act as 

effective dispersal vectors that would selectively move 

spores to other potentially suitable habitats. Gifford 

and Brandon (1978) suggested that soil movement by 

soil insects, worms, or larger animals may break up 

and disperse gametophytes. Buell (2001) observed that 

distribution patterns of Botrychium in Summit County, 

Colorado often followed swales, heavy equipment 

tracks, and erosion rills, suggesting that surface runoff 

may play a role in the dispersal of spores. 

Recent genetic studies of Botrychium simplex 

suggest, as seen in studies of other ferns cited above, 

that its migration is extremely limited (Farrar 2005). 

Populations of B. simplex in the Sierra Nevada range 

of California showed great variability in allelic 

composition, even among samples taken from single 

2 by 0.5 mile meadow. Differentiation was observed 

even between occurrences 100 m apart that would not 

be possible if unrestricted inter-occurrence migration 

was occurring. Thus, Farrar (2005) concludes that 

occurrences more than a few miles apart are effectively 

isolated and that suitable, uncolonized habitats at these 

distances have a low probability of receiving a sufficient 

number of spores to assure colonization. 

41 

Cryptic phases 

Like other species of Botrychium, B. simplex 

is probably capable of remaining dormant for at least 

one year and perhaps more. Long periods of dormancy 

are well documented for many species in subgenus 

Botrychium (Muller 1993, Kelly 1994, Lesica and 

Ahlenslager 1995, Johnson-Groh 1998, Johnson-Groh 

1999, Johnson-Groh and Farrar 2003). Dormancy is 

relatively brief for B. mormo. Only 24 percent of B. 

mormo plants remaining dormant one year return in the 

following year, and only 4 percent return after a second 

year of dormancy (Johnson-Groh 1998). 

The dominance of subterranean life stages of 

Botrychium species was highlighted by the recent 

work of Johnson-Groh et al. (2002) who observed very 

high ratios of underground (gametophytes and juvenile 

sporophytes) to aboveground structures. 

The importance of spore banks for Botrychium 

simplex is unknown, but recent studies suggest that 

they play a vital role in the survival strategies of some 

ferns (Dyer and Lindsay 1992). Because of the limited 

ability to grow Botrychium spores in culture, it has been 

difficult to observe and quantify spore banks (Johnson- 

Groh and Farrar 2003), but studies of other ferns have 

found diverse spore banks that persist for many years 

(Milberg 1991, Dyer 1994). 

The longevity of the spores of Botrychium simplex 

is unknown. Measurements of non-chlorophyll-bearing 

spores of other fern families have an average viability 

length of 1045 days (Lloyd and Klekowski 1970), and 

the spores of some fern genera may remain viable for 

long periods of time (Miller 1968, Lloyd and Klekowski 

1970, Windham et al. 1986). Spores of other fern genera 

(not Botrychium) have been germinated from 50-yearold 

herbarium specimens (Dyer and Lindsay 1992). 

Mycorrhizae 

Botrychium species rely upon mycorrhizae in 

both the sporophytic (Bower 1926, Gifford and Foster 

1989) and gametophytic (Campbell 1922, Bower 

1926, Scagel et al. 1966, Gifford and Foster 1989, 

Schmid and Oberwinkler 1994) stages. Germination 

can occur without mycorrhizal infection; however, the 

gametophyte will not mature without an arbuscular 

mycorrhizal symbiont (Campbell 1911, Whittier 1972, 

Whittier 1973). Many observations suggest that the 

fungi forming symbioses with Botrychium species are 

zygomycetes that provide resources to gametophytes 

via a “Paris-type” arbuscular mycorrhizal association

(Read et al. 2000). The subterranean, achlorophyllous 

gametophyte of B. lunaria (subgenus Botrychium) 

may live under ground for up to five years (Winther 

personal communication 2002) using carbohydrates 

and minerals gained from the mycorrhizal interaction 

(Schmid and Oberwinkler 1994). Daigobo (1979) 

observed infection of the gametophyte of B. multifidum 

(subgenus Sceptridium) with an endophytic (arbuscular 

mycorrhizal) fungus in all gametophytes examined at all 

developmental stages. Dormancy in Botrychium species 

appears to be related to the health of mycorrhizae 

(Johnson-Groh 1998). 

It is unknown how or if the mycorrhizal 

interaction changes when the gametophyte develops 

into a sporophyte, but it certainly shifts from a parasitic 

relationship to a more mutualistic relationship since the 

sporophyte is chlorophyllous and not wholly dependent 

on a mycobiont for carbohydrates. Botrychium 

sporophytes have reduced, non-proliferous roots that 

lack hairs (Wagner and Wagner 1993), and they depend 

upon mycorrhizae (Bower 1926, Foster and Gifford 

1989). Observations of root samples of B. multifidum 

and B. virginianum showed 100 percent infection, with 

all roots having the same degree of infection (Kempema 

et al. 2003). 

Arbuscular (also referred to in the literature as 

vesicular-arbuscular) mycorrhizae are a known fungal 

symbiont with Botrychium species (Berch and Kendrick 

1982, Schmid and Oberwinkler 1994). Johnson-Groh 

(1999) hypothesizes that the most important factor 

in the establishment and persistence of Botrychium 

occurrences is the presence of mycorrhizae. Little 

is known about the nature of this interaction. Farrar 

(1998) noted that mycorrhizal fungi are low in species 

diversity, ubiquitous in disturbed and undisturbed 

sites, and generalists in what plant species they infect 

(Smith and Read 1997). Recent studies have measured 

high species diversity of arbuscular mycorrhizal (AM) 

fungi in a single hectare (Bever et al. 2001). A single 

plant root was observed to host up to 49 species of 

AM fungi (Vandenkoornhuyse et al. 2002). These 

observations, coupled with the ubiquity and low host 

specificity of AM fungi, suggest that mycorrhizae may 

not be a limiting factor in the distribution of B. simplex. 

Future studies will be able to identify fungal symbionts 

of Botrychium species through experimentation with 

gametophytes in axenic culture (Read et al. 2000). 

Mycorrhizae can affect the composition of a plant 

community by shifting the intensity of competitive 

interactions (Read 1998, Van Der Heijden et al. 1998). 

Marler et al. (1999) found that the exotic spotted 

42 

knapweed (Centaurea maculosa) had more intense 

competitive effects on Festuca idahoensis (Idaho 

fescue) when grown together in the presence of 

mycorrhizal fungi. With their tight association with 

mycorrhizae, similar work with Botrychium species is 

needed to understand the potential for mycorrhizaemediated 

interspecific competition. 

Hybridization 

Hybrids between Botrychium species are rare 

(Wagner and Wagner 1993, Wagner 1998). At least 

10 records of sterile hybrid combinations have been 

documented (Wagner 1980, Wagner et al. 1984, Wagner 

et al. 1985, Wagner and Wagner 1988, Wagner 1991, 

Wagner 1993, Ahlenslager and Lesica 1996). There 

are no records of hybrids in Region 2, but it is possible 

that hybrids may exist where B. simplex occurs with B. 

lunaria in several locations in Region 2. Two sterile 

hybrids have been documented between B. simplex 

and other Botrychium species. Botrychium lunaria 

(Wagner and Wagner 1988) and B. matricariifolium 

(Wagner 1980, Wagner 1991) are both capable of 

hybridizing with B. simplex. The resulting offspring 

have intermediate characteristics and sterile, abortive 

spores (Wagner 1993). 

Evidence from cytological and molecular 

research on many moonwort species strongly suggests 

an allopolyploid origin through historic hybridization 

events (Wagner 1993, Hauk and Haufler 1999, 

Wagner and Grant 2002). Three polyploid moonworts, 

Botrychium hesperium, B. pseudopinnatum, and B. 

gallicomontanum, are believed to have arisen through 

ancient hybridization events involving B. simplex. As 

such, these species are referred to as “nothospecies.” 

Table 5 describes the purported parentage of these 

nothospecies. Other moonworts also have a suspected 

hybrid parentage, including the recently described B. 

alaskense, which is an allotetraploid of B. lunaria and 

B. lanceolatum (Wagner and Grant 2002). 

The anatomy of the gametophyte (Bower 1926) 

and the difficulty that gametes encounter when traveling 

through dry soil (McCauley et al. 1985, Soltis and Soltis 

1986) serve to limit outcrossing and hybridization. 

Morphological and genetic analyses of genus 

communities have demonstrated that hybridization 

rarely occurs, and most hybrids have abortive spores 

(Wagner and Wagner 1983, Wagner et al. 1984, Wagner 

and Wagner 1986). This demonstrates the presence of 

multiple species in these genus communities rather than 

intraspecific variants.

Demography 

Members of the genus Botrychium appear to 

have naturally low rates of outcrossing, resulting in low 

levels of genetic variability (Farrar 1998, Farrar 2005). 

Botrychium species have coped with this for thousands, 

if not millions, of years, and it is therefore not a concern 

in assessing species or population viability. They are not 

subject to inbreeding depression because they do not 

carry a genetic load of deleterious alleles typical among 

outcrossing species (Farrar 2005). 

As in other Botrychium species, the gametophyte 

of B. simplex appears to be designed for self-fertilization 

since the antheridia are positioned just above the 

archegonia (Campbell 1922). Water moving through the 

soil is likely to bring the male sex cells to the archegonia 

on the same plant (Bower 1926). Hauk and Haufler 

(1999) found genetic evidence confirming that there 

are extremely high levels of inbreeding in B. simplex, 

and this has been observed in other Botrychium species 

(Soltis and Soltis 1986, Swartz and Brunsfeld 2002, 

Farrar 2005). Moonwort species consistently show very 

low intraspecific allelic variability when compared with 

other ferns and seed plants (Farrar 1998). Watano and 

Sahashi (1992) observed that heterozygous genotypes 

were very rare in four species in subgenus Sceptridium 

due to high rates of inbreeding. McCauley et al. (1985) 

found B. dissectum (subgenus Sceptridium) to have 

an outcrossing rate of less than 5 percent. Camacho 

and Liston (2001) found high genetic diversity in B. 

pumicola, suggesting that outcrossing is occurring 

or that B. pumicola has some other mechanism for 

maintaining high genetic diversity. The rare presence 

of interspecific hybrids in natural settings indicates 

the ability for cross-fertilization hybridization to occur 

(Wagner 1980, Wagner et al. 1984, Wagner et al. 1985, 

Wagner and Wagner 1988, Wagner 1991, Ahlenslager 

and Lesica 1996). 

Given the breeding biology of Botrychium 

simplex, it is possible that they may not be sensitive to the 

effects of inbreeding depression since opportunities for 

sexual recombination are rare. The spatial distribution 

of B. simplex occurrences suggests that gene flow (via 

spore or, less frequently, gamete movement) between 

occurrences is probably very low. 

The life span of the Botrychium simplex 

sporophyte has not been measured. Johnson-Groh 

and Farrar (2003) estimate that moonworts live 

approximately 10 years. The aboveground longevity 

of B. campestre is approximately four years while B. 

43 

mormo rarely live longer than two years (Johnson-Groh 

1998). Botrychium australe plants live an average of 

11.2 years (Kelly 1994), but B. dissectum (subgenus 

Sceptridium) individuals can live at least a few decades 

(Montgomery 1990, Kelly 1994), and B. multifidum 

(subgenus Sceptridium) may live for as long as 100 

years (Stevenson 1975). 

Botrychium gametophytes are reported to persist 

underground for up to five years (Winther personal 

communication 2002) and grow very slowly from 

an embryo into an adult gametophyte that produces 

gametes (Wagner 1998). The longevity and the 

fate of the gametophyte after the production of a 

sporophyte have not been reported (Vanderhorst 1997). 

Sporophytes also may live heterotrophically under 

ground for several years before producing aboveground 

structures (Kelly 1994). Upon emergence above ground, 

the sporophytes begin spore production on their fertile 

lamina (sporophore). Figure 12 is a diagrammatic 

representation of the life cycle of B. simplex, and 

Figure 13 presents a lifecycle graph (after Caswell 

2001). The Reproductive biology and autecology 

section of this document provides a discussion of the 

life cycle of B. simplex. 

No population habitat viability analysis (PHVA) 

has been done for Botrychium simplex. The only 

Botrychium species for which a PHVA has been 

conducted is B. mormo (Berlin et al. 1998). Although 

genetic studies confirm its close relationship with B. 

simplex, B. mormo differs in many significant ways 

from B. simplex and most other moonworts as well. 

Nonetheless, some of the conclusions drawn from 

the model are relevant to most members of the genus. 

Three factors have the most influence in the model: the 

number of viable spores set per sporophyte, the nature 

and extent of a spore bank, and spore germination rate. 

Unfortunately, these are the factors about which the 

least is known. 

Impacts to Botrychium simplex occurrences 

resulting from environmental stochasticity are possible 

(Johnson-Groh et al. 1998). Environmental stochasticity 

includes variation in fecundity and survival as 

a consequence of environmental conditions and 

catastrophic local events. It may lead to local extinction 

(Lande 1998, Oostermeijer et al. 2003). Environmental 

stochasticity can operate at many scales and may affect 

a species across part or all of its range. Maintaining the 

largest occurrences possible is most likely to reduce 

potential negative consequences. Extinction probability 

models show that belowground stages buffer local

Zygote 

Sexual 

reproduction 

RECOMBINATION 

Antheridum 

Egg 

 

 

SPOROPHYTE 

SPOROPHYTE 

GENERATION (2N) 

GAMETOPHYTE 

GENERATION (1N) 

Archegonium 

Antherizoid 

44 

Asexual 

reproduction 

Archesporial cells in 

sporangium 

Spore mother cell 

SEGREGATION 

(MEIOSIS) 

GAMETOPHYTE 

Meiospores 

Figure 12. Lifecycle diagram for Botrychium simplex (after Lellinger 1985), illustrating the alternation of 

generations. 

Botrychium occurrences against extinction (Johnson- 

Groh et al. 1998), but recovery may not be possible 

after major disturbances. 

Botrychium occurrences appear to have a 

metapopulation structure that may be an important 

component of their viability. Johnson-Groh and Farrar 

(2003) provide an overview of the metapopulation 

structure of subgenus Botrychium. Using metapopulation 

classes defined by Hanski and Simberloff (1997), they 

note that Botrychium populations do not conform 

to a single metapopulation class. Rather, complex 

spatial interactions in multiple metapopulation classes 

probably best characterize the structure of Botrychium 

populations. Botrychium simplex and other species of 

Botrychium may depend on a shifting mosaic of suitable 

habitats for their long-term persistence (Clausen 1938, 

Chadde and Kudray 2001b), as does Pedicularis 

furbishiae (Pickett and Thompson 1978, Menges and 

Gawler 1986). If this is the case, then spores are the 

only means by which B. simplex could migrate to new 

locations. Because most Botrychium species are early to 

mid-seral species, they may be expected to drop out as 

succession proceeds to conditions unsuitable to them. 

Succession is of particular concern for meadow species, 

including B. simplex (Johnson-Groh and Farrar 2003). 

Evidence of limited dispersal of B. simplex suggests that 

the probability of successful long-distance dispersal is 

low (Farrar 2005). 

Demographic studies of Botrychium simplex 

are lacking, and basic parameters circumscribing its 

life-history characteristics are unknown. Sporophytes 

of Botrychium species can remain dormant for one

spore 

C 

A 

gametophyte 

D 

100,000 

spores/ 

sporophyte 

B 

45 

juvenile/ 

dormant 

sporophyte 

E 

F 

G 

adult 

sporophyte 

Figure 13. Hypothetical life cycle graph (after Caswell 2001) for Botrychium simplex. Transition probabilities are not 

known and are difficult to quantify since important stages of the lifecycle occur underground (A-G). See Johnson- 

Groh et al. (1998) for the best information currently available regarding these parameters for subgenus Botrychium. 

The number of years needed for a juvenile sporophyte to reach adulthood and emerge from the ground is not known. 

Spore production is estimated from Wagner (1998). No transition probabilities are known for B. simplex. 

or more years, and most of the population at a given 

site probably resides underground (Johnson-Groh et 

al. 2002). Johnson-Groh et al. (2002) found that the 

average ratio of belowground to aboveground plants 

for moonwort species was 332:1. Botrychium mormo, 

the closest relative of B. simplex studied, had among 

the lowest ratios observed, at 65:1. An average of 728 

gametophytes per square meter was documented for B. 

mormo. In an earlier study, Bierhorst (1958) found 20 

to 50 gametophytes of B. dissectum per square foot, 

with relatively few mature gametophytes with attached 

juvenile sporophytes. These observations indicate 

that long-term studies are needed to assess the true 

population size (Johnson-Groh and Farrar 2003), and 

that the observation of a single emergent sporophyte 

may indicate the presence of a viable occurrence 

(Casson et al. 1998), or an early stage of colonization. 

Studying how plants become established is 

problematic because important events in the life cycle 

of Botrychium occur underground. The requirement of 

darkness for spore germination (Whittier 1973) is not 

surprising, given the gametophyte’s need to establish 

a mycorrhizal symbiosis within a few cell divisions 

(Campbell 1911). The mechanism by which spores get 

under ground is not known, but they somehow get from 

the soil surface to a depth of 1 to 2 inches. Water and 

frost action (freezing and thawing), and possibly fire, 

are probably involved (Crook personal communication 

2003, Root personal communication 2003). From 

predictions based on observations of survivorship, 

Johnson-Groh et al. (2002) estimate that 95 percent of 

Botrychium spores are unsuccessful. 

Community ecology 

Botrychium simplex is known from a range of 

plant communities in Region 2, and the list of associated 

species documented with it is long (Table 11). Kolb and 

Spribille (2000) present associated species in plot data 

from Shrine Pass (White River National Forest) and 

Boreas Pass (White River and Pike national forests), 

both in Colorado.

Table 11. Associated species reported with Botrychium simplex in USDA Forest Service Region 2. 

Associated Species Colorado Nebraska South Dakota Wyoming Exotic? 

Abies lasiocarpa X X 

Achillea millefolium X X 

Aconitum columbianum X 

Agoseris glauca X 

Agrestis thurberiana X 

Agrimonia sp. X 

Allium sp. X 

Amelanchier alnifolia X 

Amelanchier sp. X 

Androsace septentrionalis X 

Anemone sp. X 

Antennaria media X 

Antennaria neglecta X 

Antennaria sp. X X X 

Aquilegia sp. X 

Arctostaphylos uva-ursi X X 

Arnica c.f. fulgens X 

Arnica cordifolia X 

Arnica sp. X 

Artemisia ludoviciana X 

Artemisia sp. X 

Astragalus sp. X 

Betula papyrifera X 

Botrychium campestre X X 

Botrychium echo X 

Botrychium hesperium X 

Botrychium lanceolatum X 

Botrychium lunaria X X 

Botrychium michiganense X 

Botrychium minganense X X 

Botrychium multifidum X X 

Botrychium pinnatum X X 

Campanula rotundifolia X X 

Carex aquatilis X X 

Carex aurea X 

Carex parryana X 

Carex rossii X 

Carex sp. X X 

Castilleja sulphurea X 

Cerastium arvense X X 

Cirsium eatonii X 

Cirsium sp. X 

Clematis tenuiloba X 

46



Collinsia parviflora X 

Collomia linearis X 

Comandra umbellata X 

Corylus cornuta X X 

Crataegus chrysocarpa X 

Crepis runcinata X 

Cryptogramma sp. X 

Cynoglossum officinale X X X 

Dasiphora floribunda X 

Delphinium bicolor X X 

Deschampsia caespitosa X X 

Dodecatheon pulchellum X X 

Elymus trachycaulus X 

Erigeron spp. X 

Festuca spp. X 

Fragaria sp. X X 

Fragaria virginiana X X 

Gallium boreale X X 

Gallium sp. X 

Geranium richardsonii X X 

Geum sp. X 

Geum triflorum X X 

Halenia deflexa X 

Heterotheca depressa X 

Heterotheca pumila X 

Hieracium umbellatum X 

Iris missouriensis X 

Juncus balticus X 

Juniperus communis X X 

Juniperus virginiana X 

Koeleria macrantha X 

Leucanthemum vulgare X X 

Lewisia pygmaea X 

Linum perenne X X 

Lithophragma parviflora X X 

Lupinus sp. X 

Mahonia repens X X 

Maianthemum stellatum X X 

Mertensia lanceolata X X 

Mimulus guttatus X 

Monarda fistulosa X X 

Muhlenbergia cuspidata X 

Myosotis verna X 

47



Orthilia secunda X 

Oxalis dillenii X 

Oxalis sp. X 

Oxytropis campestris X 

Panicum acuminatum X 

Pentaphylloides floribunda X 

Phleum pratense X X 

Phlox hoodii X 

Phlox pulvinata X 

Picea engelmannii X X 

Picea glauca X 

Pinus contorta X X 

Pinus ponderosa X X 

Poa bulbosa X 

Poa compressa X 

Poa nemoralis X 

Poa pratensis X X 

Poa sp. X X 

Polygala senega X 

Polygonum viviparum X 

Polytrichum piliferum X 

Populus c.f. deltoides ssp. monilifera X 

Populus tremuloides X 

Potentilla sp. X X 

Prunella vulgaris X 

Pseudotsuga menziesii X 

Pyrola chlorantha X 

Quercus macrocarpa X 

Ranunculus sp. X 

Rhacomitrium canescens X 

Ribes sp. X 

Rosa sp. X 

Rubus idaeus X 

Salix bebbiana X 

Salix planifolia X 

Sanicula marilandica X 

Sedum lanceolatum X 

Sedum sp. X 

Selaginella densa X X 

Senecio canus X 

Senecio sp. X 

Shepherdia argentea X 

Shepherdia canadensis X 

48



Sisyrinchium idahoense X 

Solidago multiradiata var. scopulorum X 

Solidago sp. X X 

Spiraea betulifolia X 

Sporobolus heterolepis X 

Stipa richardsonii X 

Symphoricarpos albus X 

Symphoricarpos sp. X X 

Taraxacum officinale X X 

Taraxacum sp. X X 

Thalictrum alpinum X 

Thalictrum dasycarpum X 

Thalictrum sp. X 

Thermopsis rhombifolia X 

Trifolium repens X X X 

Urtica dioica X 

Vaccinium caespitosum X 

Vaccinium myrtillus X 

Vaccinium sp. X X 

Viola adunca X 

Viola sp. X 

Zigadenus sp. X 

Zigadenus venosus X 

Some species and species groups commonly 

associated with Botrychium simplex in Region 2 reflect 

its habitat affinities. Coniferous trees are common 

associates with B. simplex, particularly in Region 

2. Conifers reported with B. simplex include Picea 

engelmannii, (Engelmann Spruce) Picea glauca (white 

spruce), Pseudotsuga menziesii (Douglas fir), Pinus 

ponderosa (ponderosa pine), P. contorta (lodgepole 

pine), and Abies lasiocarpa (subalpine fir). Sedges 

(Carex species) are also frequently documented with 

B. simplex range-wide and with many occurrences in 

Region 2. The sedge most commonly associated with 

B. simplex in Region 2 is C. aquatilis. Strawberry 

species (Fragaria vesca and F. virginiana) are common 

associates of many Botrychium species including B. 

simplex (Root personal communication 2003). 

Rigorous work on the community ecology of 

Botrychium simplex is lacking. Using phytosociological 

methods, Kolb and Spribille (2000) described the 

community in which Botrychium species were 

found in Summit County, Colorado as “Festuco 

– Heterothecetum pumilae,” (the Fescue- Dwarf 

Aster Community), named for the dominant plants 

49 

(Festuca brachyphylla and Heterotheca pumila) in the 

community. This community is characterized by ruderal 

taxa, including Fragaria virginiana. Botrychium 

simplex is not noted as a characteristic species of this 

community type, but occurrences of B. simplex were 

found in this community type at Shrine Pass and Boreas 

Pass. Detailed community description and plot data can 

be found in Kolb and Spribille (2000). 

Species in subgenus Botrychium often occur 

together in genus communities (Wagner and Wagner 

1983). “Genus community” is a term coined by Wagner 

and Wagner (1983) to describe the peculiar tendency 

for two or more Botrychium species to be found 

in close association with one another. Botrychium 

simplex has been documented with numerous other 

moonwort species in Region 2 and elsewhere. Wagner 

and Wagner (1983) document the co-occurrence 

of B. simplex with nine other moonwort species in 

genus communities (B. minganense, B. crenulatum, 

B. lunaria, B. pinnatum, B. lanceolatum, B. echo, B. 

hesperium, B. matricariifolium, and B. paradoxum). 

Botrychium simplex has also been documented with 

B. ascendens (Wagner and Wagner 1986, Vanderhorst

1997), B. pedunculosum (Wagner and Wagner 1986), 

B. mormo (Wagner and Wagner 1981, Chadde and 

Kudray 2001a), B. multifidum (Wagner and Devine 

1989), and B. campestre and B. gallicomontanum 

(Farrar and Johnson-Groh 1986). Ollgaard (1971) 

reports the presence of B. lanceolatum, B. lunaria, and 

B. multifidum with B. simplex in Greenland. 

Within Region 2, Botrychium simplex is also 

frequently documented in genus communities. 

Colorado element occurrence records note the presence 

of B. echo, B. hesperium, B. lanceolatum, B. lunaria, 

and B. minganense with B. simplex (Colorado Natural 

Heritage Program 2006). Botrychium simplex has 

been found with B. campestre in Brown County, 

Nebraska (Farrar and Johnson-Groh 1986, Farrar 

personal communication 2003). Both B. campestre 

and B. ‘michiganense’ have been documented from the 

Bearlodge Campground in Crook County, Wyoming, 

but they have not been found in close association with 

B. simplex at this location. Botrychium simplex has 

been found with B. pallidum at three locations on the 

Black Hills National Forest, and with other unidentified 

taxa at one location (Burkhart personal communication 

2006). Outside Region 2, B. simplex has been found 

with B. multifidum at two sites in Yellowstone National 

Park (Whipple personal communication 2003), and 

these species are often found together (Crook personal 

communication 2003, Farrar 2005). 

The coexistence of species of Botrychium in genus 

communities is interesting from a community ecology 

standpoint. If the members of genus communities 

occupy the same niche, then they coexist in violation 

of Gause’s competitive exclusion principle (Krebs 

1972). Because water, nutrient, and some carbohydrate 

uptake are mediated by mycorrhizae, it is possible that 

even if genus community members depend on the 

same resources, coexisting plants are not engaged in 

direct interspecific competition. Competition may be 

for access to the mycorrhizae if it is occurring at all. 

No research has been done on Botrychium species with 

respect to these issues. Wagner and Wagner (1983) offer 

an interesting discussion of this issue from a population 

biology standpoint. 

Moonworts clearly tolerate some degree of 

grazing, but there is little information on which to base 

management decisions regarding livestock and wildlife 

grazing (Johnson-Groh and Farrar 2003). Montgomery 

(1990) found that even repeated removal of the leaf 

of Botrychium dissectum (subgenus Sceptridium) 

for three years did not kill the plants, and on this he 

commented (p. 178) “It is certainly remarkable that 

50 

these plants persist.” Observations of B. minganense 

and B. montanum by George Wooten on the Okanogan 

National Forest in Oregon may offer some insights 

into possible impacts of grazing on B. simplex. These 

two species were monitored in a trampled spring area, 

and while they appeared to benefit from disturbance 

in lightly trampled areas, they were destroyed in 

heavily trampled areas. Moderately trampled areas 

were not affected (Roche 2004). The sensitivity of B. 

simplex to grazing is not known, but some observations 

suggest that it tolerates a moderate level of grazing. It 

is documented from pastures in numerous locations 

outside of Region 2. 

There is some speculation that spores of at least 

some Botrychium species are dispersed by mammals, 

based primarily on observations of herbivory on the 

sporophores of B. mormo (Casson et al. 1998, Wagner 

personal communication 2002). Although dispersal by 

animals has not been demonstrated for B. simplex, it is 

possible that elk, deer, and other grazing ungulates act 

as dispersal agents by eating the ripe sporophores in the 

fall. This would be advantageous to B. simplex, since 

elk and deer use B. simplex habitats and would tend 

to deposit the spores in sites suitable for germination. 

Small mammals may also function as short-distance 

spore dispersal agents. The large, thick walled spores 

of Botrychium species may be an adaptation to dispersal 

by herbivores (Wagner 1998). 

Plants on Vail Pass in the White River National 

Forest have been subjected to small mammal predation 

(Colorado Natural Heritage Program 2006). The plant 

found at Copper Mountain also appeared to have been 

partially eaten or damaged, but it also had been driven 

over by a vehicle (Root personal communication 2003). 

There are no reports of parasitism or disease in the 

literature for any Botrychium species in Region 2. 

CONSERVATION 

Threats 

Observations suggest that there are several threats 

to the persistence of Botrychium simplex. In Ohio, 

soil compaction, drying of habitat, and removal of 

vegetation are threats to B. simplex (Ohio Department 

of Natural Resources 2003). Chadde and Kudray 

(2001b) noted that major threats to B. simplex in 

Wisconsin, Michigan, and Minnesota include exotic 

earthworms, exotic plants, succession to closed 

canopy forest, major disturbance (ranked medium for 

degree of impact), and canopy thinning and minor 

disturbance (ranked low for degree of impact). Lorain

(1990) noted no anthropogenic threats to B. simplex in 

Idaho but included windthrow or trampling by large 

game animals as potential natural threats. Threats to 

Botrychium occurrences in the Wallowa Mountains 

of Oregon include natural succession, recreational 

use, and impacts from pack animals and domestic 

livestock (Alverson and Zika 1996). Berlin et al. (1998) 

described and ranked numerous threats to B. mormo. 

These were sorted into four general categories: exotics, 

forestry practices, development and land use, and other. 

While many of these threats are specific to B. mormo, 

some are relevant for B. simplex as well: earthworms, 

clearcutting, salvage sales, road right-of-way 

management, lakeshore development, recreation site 

development, anthropogenic alteration of hydrology, 

human population increase, natural windthrow, and 

flooding. While these factors pose potential threats for 

B. simplex, the nature and magnitude of the threat is 

different for B. simplex due to ecological and geographic 

differences between these species. 

In approximate order of decreasing priority, 

threats to Botrychium simplex in Region 2 include ski 

area development and maintenance, road construction 

and maintenance, timber harvest, recreation, fire, 

grazing, effects of small population size, woody plant 

encroachment, exotic species invasion, succession, 

global climate change, and pollution. More complete 

information on the biology and ecology of this species 

may elucidate other threats specific to Region 2. 

Assessment of threats to this species will be an important 

component of future inventory and monitoring work. 

See the sections below for specific discussion of the 

known threats to B. simplex. 

Influence of management activities and natural 

disturbances on individuals and habitat 

Ski area development and maintenance 

Construction of facilities to support recreational 

skiing presents potential threats to specific moonwort 

occurrence, primarily those on the White River National 

Forest. Because of a lack of baseline data, it is not know 

to what extent the creation of ski runs and ski areas 

has benefited or negatively impacted occurrences of 

Botrychium species, including B. simplex. For species 

that can occur in forested sites such as B. simplex and 

B. multifidum, the potential negative impacts from ski 

run creation are greater than for most other moonwort 

species that are found almost exclusively in open sites. 

Creation of ski runs and facilities for recreational 

skiers has altered habitats in the mountains of Region 

51 

2, and it is not known if occurrences in ski runs remain 

viable for long periods. Their viability could easily 

be compromised by management activities needed to 

maintain ski runs. Snow-making activities can change 

the hydrology of an area by increasing surface runoff and 

snow depth. The impacts of these factors to Botrychium 

simplex, if any, are not known. Construction of a ski 

hut near the Vail Pass occurrences presents a potential 

threat due to disturbance associated with construction 

and increased use of the hut. Because the presence of 

B. simplex was known before construction, the hut was 

located in a site where impacts to the occurrence would 

be reduced. Summer use of the hut could still result in 

trampling of individuals. 

Road and trail construction and maintenance 

Road construction and maintenance threaten 

known occurrences of Botrychium simplex in Region 

2. New road construction and road widening or 

modification can directly impact occurrences of B. 

simplex in the area. Potential secondary effects of roads 

include changes in hydrology, an increase in erosion, 

and the introduction of weed species. 

One occurrence at Copper Mountain in Colorado 

was reported in an infrequently used road where renewed 

use, repair, and/or maintenance may compromise its 

viability. Four other occurrences in Region 2 are known 

to be within or adjacent to active roads in Region 2, and 

others may be as well. In 2004 and 2005, five additional 

occurrences were discovered within or adjacent to old 

unused roadbeds in the Black Hills National Forest. The 

threats to these occurrences appear minimal at present, 

but they could increase if the roads were reopened. One 

Botrychium simplex occurrence is located along active 

railroad tracks where similar threats may exist. 

Timber harvest 

Direct impacts to Botrychium species from 

timber harvest include soil compaction, disruption 

of the organic surface horizon, changes in light, and 

loss of soil nutrients and moisture (Johnson-Groh and 

Farrar 2003). Sedimentation is another documented 

threat. Secondary impacts of timber harvest such as 

road building or other activities may also represent 

potential threats to B. simplex occurrences. It might be 

expected that occurrences of B. simplex within a timber 

harvest area would be negatively impacted. Outside of 

ski areas on the White River National Forest, there are 

no occurrences of B. simplex in Region 2 known to be 

currently or historically impacted by timber harvest.

Recreation 

Recreational use of Botrychium simplex habitat 

presents a direct threat to individuals, which may 

be killed or damaged. Off-road vehicle use (both 

motorized and non-motorized) represents the greatest 

recreational threat to B. simplex. Off-road vehicle 

use has the potential to be highly detrimental to B. 

simplex habitat by disturbing soil and plant community 

integrity, altering hydrology and hydrologic processes, 

increasing erosion, and enhancing the spread of 

invasive plant species. Off-road use of motorized 

vehicles has affected the Caribou Flats area of Boulder 

County, Colorado where one occurrence has been 

documented. Off-road vehicle use could threaten 

occurrences in unused roadbeds on the Black Hills 

National Forest if these areas begin to be used as trails. 

Use of mountain bikes and “mountainboards” (similar 

to snowboards but equipped with wheels for use on 

ski slopes in the summer) has the potential to impact 

individuals on ski slopes. 

Fire 

Fire affects individual plants directly by burning 

their aerial portions, but Botrychium species, including 

B. simplex, appear to suffer no ill consequences from 

this (Johnson-Groh and Farrar 2003). Particularly hot 

fires or fires when the soil is desiccated could result 

in plant mortality, but due to the strong dependence 

of the species on mycorrhizae, removal of leaf tissue 

via burning or other means is probably inconsequential 

to the plant’s survival (Montgomery 1990, Wagner 

and Wagner 1993, Johnson-Groh and Farrar 1996a, 

Johnson-Groh and Farrar 1996b, Johnson-Groh 1999). 

Fires that occur in the summer or fall (when forest fires 

are most common) would preclude any reproductive 

output for that year and might kill spores lying near the 

surface (Root personal communication 2003). 

While fire itself may not have much effect on 

Botrychium simplex, the secondary effects of fire 

can threaten individuals. Sedimentation is one such 

secondary threat. Burial by sediment has resulted in 

the apparent mortality of buried individuals of other 

Botrychium species (Johnson-Groh and Farrar 2003). 

Part of a permanent plot at Frenchman’s Bluff, Minnesota 

was buried by soil excavated by gophers. While this 

resulted in the temporary loss of B. gallicomontanum 

individuals, after 11 years of monitoring at this site 

the population had rebounded (Johnson-Groh 1999, 

Johnson-Groh and Farrar 2003). 

52 

Grazing 

Grazing offers both potential benefits and 

detriments with regard to Botrychium simplex. This 

species is found in pastures outside of Region 2, 

suggesting that it is tolerant of some level of grazing. 

However, it is most often reported from “old pastures,” 

suggesting that B. simplex is less compatible with an 

intensive grazing regime. Elk and other ungulates 

frequent the meadow habitats of B. simplex and may 

occasionally graze it. Grazing can eliminate a season’s 

contribution to the spore bank (Johnson-Groh and Farrar 

2003). Livestock tend to prefer grasses, permitting the 

competitive release of broad-leaved plants, which 

may benefit B. simplex in some situations. However, 

disturbance of the surface by cattle may injure some 

individuals (potentially above and below ground). 

Livestock grazing may indirectly affect B. simplex 

by increasing erosion, altering the plant community 

composition, damaging the soil structure (particularly 

in wet conditions), and introducing invasive plants. 

Therefore, the use of cattle grazing as a management 

tool for the enhancement of habitat is risky for a plant 

as rare as B. simplex. 

Effects of small population size 

Demographic stochasticity is the variation over 

time in vital rates such as recruitment and survival, and 

it is generally only a concern for very small occurrences. 

Because there are limited data on occurrence sizes in 

Region 2, the degree to which many occurrences are 

threatened by the effects of demographic stochasticity 

is unknown. Reported numbers of individuals at most 

occurrences of Botrychium simplex in Region 2 fall 

below the generally accepted minimum effective 

population size of 50 reproductive individuals (Soulé 

1980) needed to buffer against the probability that a 

fluctuation in vital rates will take the occurrence to the 

extinction threshold. Botrychium species have extremely 

low levels of heterozygosity because unmasked 

deleterious alleles have been purged from populations. 

This has led Farrar (2005) to suggest that the lack of 

genetic variability in Botrychium species should not be 

a concern in assessing species or population viability. 

Environmental stochasticity refers to random 

variations in the physical and biological environment. 

For a single occurrence, this includes natural events 

happening at random intervals that cause the deaths of a 

large proportion of individuals in the occurrence. Such 

events may occur very rarely, yet still have a significant

impact on the occurrence (Menges 1991). Maintaining 

multiple occurrences well-distributed throughout 

Region 2 can mitigate the effects of environmental 

stochasticity. Studies of Botrychium simplex and 

other species of Botrychium suggest that there is little 

genetic connectivity among occurrences of B. simplex 

in Region 2, so these occurrences are more vulnerable 

to extirpation by catastrophic events such as fires, 

landslides, and disease. 

Woody plant encroachment 

Encroachment of trees and other woody plants 

is considered a threat to many Botrychium species, 

which are not typically found in shaded or heavily 

forested sites; it is a particular threat to meadow species 

(Johnson-Groh and Farrar 2003). The degree to which 

B. simplex is threatened by succession is less clear since 

it is often documented in forested sites in the shade. 

Exotic species 

Six exotic species have been documented 

with Botrychium simplex in Region 2 (Table 11): 

houndstongue (Cynoglossum officinale), oxeye daisy 

(Leucanthemum vulgare), timothy (Phleum pratense), 

Kentucky bluegrass (Poa pratensis), dandelion 

(Taraxacum officinale), and white clover (Trifolium 

repens). Of these species, it is likely that Kentucky 

bluegrass represents the greatest threat since it can be 

highly invasive and competitive in riparian areas and 

meadows. Its presence in wet sites in Rocky Mountain 

National Park is a concern for land managers (Connor 

personal communication 2003). Oxeye daisy is known 

to occur near B. simplex at one location in South 

Dakota, but it is also spreading in the mountains of 

Colorado. There are no data suggesting a direct impact 

of noxious weeds on Botrychium species (Johnson- 

Groh and Farrar 2003), but their mutual affinity for 

disturbance may cause Botrychium species and their 

habitat to be vulnerable. 

The management of weed species may also be a 

cause for concern with respect to Botrychium species. 

Johnson-Groh (1999) observed the effects of herbicide 

and fire on prairie moonworts (B. simplex, B. campestre, 

and B. gallicomontanum). Plants that had been hit 

directly with the herbicide Roundup were apparently 

killed (Johnson-Groh 1999, Johnson-Groh and Farrar 

2003). Efforts to control noxious weeds will probably 

affect moonwort occurrences in right-of-ways or other 

sites targeted for weed management. At the time this 

report was written, it was not possible to determine if B. 

53 

simplex was more or less affected by herbicide than the 

other moonwort species. 

In the deciduous, hardwood forest habitats 

of Botrychium mormo, which is a close relative of 

B. simplex, invasion of non-native earthworms has 

resulted in dramatic decreases in mycorrhizal fungi 

(Nielsen and Hole 1963, Cothrel et al. 1997, Berlin 

et al. 1998, Gundale 2002). Because B. mormo is an 

obligate mycorrhizal symbiont, this poses a significant 

threat. Most earthworm activity takes place in the O 

horizon (Langmaid 1964) while mycorrhizal activity 

is greatest at the interface of the O and A horizons 

(Smith and Read 1997). The activity of earthworms has 

resulted in the elimination of the duff layer and a shift 

in species composition in B. mormo habitat (Berlin 

et al. 1998, Gundale 2002). Although earthworms 

present a possible threat to B. simplex, no research has 

shown that they are affecting species of Botrychium 

other than B. mormo. Earthworms are a diverse group 

of over 3,500 species worldwide, and the expansion 

of global commerce may be increasing the likelihood 

of exotic earthworm invasions with potential adverse 

effects on soil processes and plant species (Hendrix 

and Bohlen 2002). 

Global climate change 

Global climate change is likely to have wideranging 

effects in the near future for all habitats. 

However, the direction of projected trends is yet 

to be determined, and predictions vary based 

on environmental parameters used in predictive 

models. For example, Manabe and Wetherald (1986) 

demonstrate projections based on current atmospheric 

CO 2 trends that suggest average temperatures will 

increase while precipitation will decrease in the West. 

Decreased precipitation could have dire consequences 

for occurrences of Botrychium simplex in Region 2 

if it results in drying of its moist habitats. Giorgi et 

al. (1998) showed that temperature and precipitation 

increased under simulated doubling of atmospheric 

carbon dioxide levels. Either scenario could affect 

the distribution of montane grasslands in Region 2. 

Temperature increase, predicted by both models, could 

cause vegetation zones to climb 350 ft. in elevation 

for every degree F of warming (U.S. Environmental 

Protection Agency 1997). Global climate change can 

be expected to affect occurrences at the southern edge 

of their natural range, pushing them northward and 

upward. With extensive habitat disturbance in the path 

of potential migration, many organisms will not be able 

to move. Preserves may provide short-term protection,

ut these may be ineffectual if organisms are trapped 

without suitable routes for migration. 

Pollution 

Atmospheric nitrogen deposition has become one 

of the most important agents of vegetation change in 

densely populated regions (Köchy and Wilson 2001). 

Nitrogen loading and the associated vegetation change 

have been observed to be greatest near large metropolitan 

areas (Schwartz and Brigham 2003). Thus, measurable 

impacts from nitrogen pollution might be expected in 

many locations in Region 2, especially in Colorado. 

Nitrogen enrichment experiments show universally that 

nitrogen is limited (Gross et al. 2000). This is likely 

to cause a few species to increase in abundance while 

many others decline (Schwartz and Brigham 2003). 

Acid deposition, which has increased markedly in 

Colorado through the 20 th century, may have already 

caused changes to the soil chemistry that threaten 

the viability of Botrychium simplex. High elevation 

watersheds of the Front Range have already reached an 

advanced stage of nitrogen saturation (Burns 2002). 

Over-utilization 

Voucher specimens do assist with taxonomic 

research on Botrychium simplex. However, caution 

should be exercised in collection. Weber and Wittmann 

(2001a, 2001b) recommend not collecting the roots 

because they contain no diagnostic characteristics, and 

collecting them kills the plant. To minimize the risks of 

infection and of removing the apical bud, Johnson-Groh 

and Farrar (2003) recommend cutting the leaf with a 

knife near ground level rather than pinching or pulling 

it with the fingers. Although evidence suggests that leaf 

removal does not have a significant, long-term effect on 

Botrychium species (Johnson-Groh and Farrar 1996a), 

collection of the species in Region 2 is only advisable 

in larger occurrences. Johnson-Groh and Farrar (2003) 

suggest that no collections be made in occurrences of 

fewer than 20 plants. Instead, they recommend that 

photographs be taken and deposited at an herbarium 

in lieu of a specimen. If collections are made of B. 

simplex, Johnson-Groh and Farrar (2003) recommend 

that no more than 10 percent of an occurrence be 

collected. Also, it is important to collect mature 

specimens since immature sporophytes are difficult 

to identify (Ohio Department of Natural Resources 

2003). Where occurrences are already of questionable 

viability, collection of material from them, even if the 

plants survive, is a risky endeavor. For example, in an 

occurrence of three sporophytes, there is almost no 

margin of error, and accidentally removing the apical 

54 

bud could easily contribute to the extirpation of the 

species at this site. This is a difficult issue for some 

Botrychium species, and particularly for B. simplex, 

since collection and verification by an expert is the only 

way to be absolutely sure of proper identification. 

There are no known commercial uses for 

Botrychium simplex. According to Gerard in his 1633 

herbal (p. 407), “moonewort” (referring to B. lunaria) 

“is singular to heale greene and fresh wounds: it staieth 

the bloudy flix. It hath beene used among the alchymistes 

and witches to doe wonders withall.” Botrychium 

species are not widely sold in the herb trade, but they 

are mentioned as ingredients in tinctures and poultices 

for the treatment of external or internal injuries. There 

is potential for over-utilization of Botrychium species 

if their popularity increases in the herb trade. Because 

they cannot be readily cultivated, any commercial use 

would require the harvest of wild plants. 

Conservation Status of Botrychium 

simplex in Region 2 

Is distribution or abundance declining in all or 

part of its range in Region 2? 

There are no data available from which meaningful 

inferences can be made regarding population trend 

or changes in distribution of Botrychium simplex in 

Region 2. Some occurrences that were last documented 

20 or more years ago have been searched for and not 

relocated, but this does not necessarily indicate that they 

are extirpated. Vague locations of older collections and 

the difficulty of Botrychium hunting create uncertainty 

as to whether some occurrences can ever be relocated. 

A precise location was known for the occurrence at Old 

Faithful in Yellowstone National Park; however, this 

occurrence has not been seen since 1994 despite several 

searches at the right time of year (Whipple personal 

communication 2003). Apparently, the only occurrence 

in the states of Region 2 that has been successfully 

revisited is at Lower Geyser Basin in Yellowstone 

National Park, but this occurrences falls outside the 

administrative boundary of Region 2. The Population 

trend section has more information on population status 

in Region 2. 

Do habitats vary in their capacity to support 

this species? 

In the absence of a disturbance regime, ecological 

succession may lead to unsuitable conditions for 

Botrychium species. This may be less of a concern, 

however, for B. simplex than for other moonwort

species in Region 2 since it is often found in shaded, 

forested areas. Variables such as fire regime, amount of 

litter, soil moisture variation, and soil texture may affect 

habitat suitability. Some habitats may be suitable for 

one variety of B. simplex but unsuitable for others. 

Vulnerability due to life history and ecology 

The vulnerability of Botrychium simplex due to 

its life history and ecology remains uncertain since 

these factors are poorly understood. Chadde and 

Kudray (2001b) note that the intrinsic vulnerability 

of B. simplex is low; occurrences are generally 

resilient and/or resistant to change. However, the 

reliance of most Botrychium species on disturbance 

to create and maintain appropriate habitat may make 

them vulnerable to woody plant encroachment and 

interspecific competition. 

The wetland and riparian habitats of Botrychium 

simplex are the most critically threatened habitats in the 

semi-arid West. These sites are often sought after for 

human uses, and some are heavily used. Consequently, 

they are subject to hydrologic alterations that could 

render them unsuitable for B. simplex. 

Evidence of populations in Region 2 at risk 

The rarity and small occurrence sizes of 

Botrychium simplex in Region 2 suggest that it is 

vulnerable to extirpation locally or even regionally, 

and that all of the Region 2 occurrences are at risk. The 

number of individuals documented thus far from the 

50 occurrences in Region 2 falls between 500 and 600 

plants. Of these 50 occurrences, 17 have not been seen 

again in more than 20 years (Table 8), and the current 

status of these occurrences is uncertain. Stochastic 

events and normal environmental variation could 

result in extirpation of many of the small and localized 

occurrences in Region 2. The quality and availability 

of habitat in Region 2 has probably declined due to 

fragmentation, exotic species invasion, hydrologic 

alteration, and edge effects that decrease the quality 

of small patches of natural vegetation. The majority 

of occurrences and potential habitat are known from 

public lands managed by the USFS and the National 

Park Service, where they receive some degree of 

protection. Chadde and Kudray (2001b) note that the 

habitat integrity is good in Region 9, with most habitats 

and sites protected and not commonly impacted by 

management. This may be true in much of Region 2 

as well, but the creation of ski runs on USFS land in 

Region 2 has resulted in extensive habitat modification 

in areas known to be occupied by B. simplex. Because 

55 

the ecology of this species is poorly understood, current 

management may be placing unsustainable demands on 

the species despite good intentions. 

Current data suggest a high degree of imperilment 

for this species in Region 2, due largely to small 

occurrences, but these data are sparse. More occurrences 

are likely to be found in Region 2 by targeted surveys, 

but the known patterns of distribution and abundance 

in Region 2 suggest that large, robust occurrences 

are unlikely to be found. This underscores the need 

to conduct additional surveys for this species and to 

monitor known occurrences rigorously. 

Management of Botrychium simplex in 

Region 2 

Implications and potential conservation 

elements 

The most current data available suggest that 

Botrychium simplex is imperiled in most of Region 2 

due to the low number of small, isolated occurrences. 

The loss of any occurrence is significant and may 

result in the loss of important components of the 

genetic diversity of the species. Within Region 2, most 

reports of B. simplex, as well as most areas of suitable 

habitat, are on National Forest System land. Thus the 

responsibility of maintaining viable populations within 

the administrative boundary of Region 2 falls largely 

on the USFS. Further research is needed, however, 

before meaningful inference can be offered regarding 

restoration policy. 

Desired environmental conditions for Botrychium 

simplex include sufficiently large areas where the 

natural processes on which the species depends can 

occur, permitting it to persist unimpeded by human 

activities and their secondary effects, such as weeds. 

This includes a satisfactory degree of ecological 

connectivity between occurrences and apparently 

suitable but unoccupied habitat. Given the current 

paucity of detailed information on this species in 

Region 2, it is unknown how far this ideal is from 

being achieved. It is possible that most or all of the 

ecosystem processes on which B. simplex depends are 

functioning properly at many or most of the occurrences 

of this species. Again, further research on this species’ 

ecology and distribution will help to develop effective 

approaches to its management and conservation. Until 

a more complete picture of the distribution and ecology 

of this species is obtained, however, priorities lie 

with conserving the known occurrences and locating 

additional occurrences.

Conservation elements essential to maintaining 

viable occurrences of Botrychium simplex in Region 

2 include early- to mid-seral stage or other suitable 

habitats and the maintenance of processes that create 

and support these habitats. Unfortunately, these are 

poorly understood at present. Given the rate at which 

new data are becoming available and the incompleteness 

of our current knowledge of B. simplex in Region 2, 

it is difficult to formulate conservation strategies at 

present. More complete knowledge of its distribution 

will permit the identification of the occurrence in 

Region 2 that are the most suitable for conservation 

management. Surveys are needed to better understand 

how the recognized subspecies of B. simplex differ in 

habitat affinities, distribution, and autecology in Region 

2. Demographic studies designed to determine the 

impacts of grazing, succession, fire, and exotic species 

on population viability are also high priorities for 

research on B. simplex in Region 2. Research is needed 

to investigate the subterranean life history, ecology, 

reproductive biology, role of mycorrhizae, and role 

of disturbance in the autecology of B. simplex so that 

conservation efforts can be most effective. Restoration 

of occurrences of members of Botrychium subgenus 

Botrychium is probably precluded by the difficulties in 

propagating them. 

Tools and practices 

Species inventory 

Botrychium simplex is seldom seen because it 

often grows in wetter sites than other moonworts (Root 

personal communication 2003), and it is cryptic and 

small. This species can be found from approximately 

mid-June through August and possibly into September, 

but it may remain dormant in less favorable (dry) years. 

Given the difficulty in finding moonworts, inventory 

and monitoring studies are difficult, and dormant 

occurrences have been reported to be extirpated 

(Lesica and Steele 1994). Success in finding B. simplex 

requires training, time, and persistence (Wagner 1946, 

Wagner and Wagner 1976). New occurrences continue 

to be found within Region 2 (e.g., Kolb and Spribille 

2000, Fertig 2003) and elsewhere Repeated surveys 

may be needed to identify all the Botrychium species 

present. The Bearlodge Campground in Crook County, 

Wyoming, where three moonwort species have been 

found over 30 years, is a classic example of the need to 

revisit a location to understand its moonwort flora. 

Johnson-Groh and Farrar (2003) offer 

suggestions for conducting surveys for Botrychium 

species. Suggested techniques include systematic 

56 

search methods, marking plants, collection protocols, 

documentation, and other concerns. The protocols they 

defined are based on the assumption that one week can 

be spent searching approximately 25 high priority sites 

per year, for about one hour each. Four or five people 

are recommended to walk transects at each area selected 

so that better coverage is achieved in the time of the 

visit. When plants are discovered, they are marked with 

pin flags and the surrounding area searched intensively 

but carefully on hands and knees. 

These methods have certain limitations that need 

to be considered. Johnson-Groh and Farrar (2003) note 

four limitations: 

v low confidence in distribution and abundance 

due to population variability 

v difficulty in finding the plants 

v predominance of the belowground lifecycle 

stages 

v the occurrence of many species in genus 

communities that complicates identification. 

Popovich (personal communication 2003) implemented 

this protocol and noted that the area searched appeared to 

have been heavily impacted by the intensiveness of the 

search efforts. It can also be cost-prohibitive to have four 

or five surveyors on site to implement these protocols. 

Johnson-Groh and Farrar (2003) also recommend using 

the “timed meander search procedure” described by 

Goff et al. (1982) for Botrychium searches. 

Due to limitations in time and funding, attempts to 

search for rare plants often involve looking for multiple 

species over large areas. While this approach has been 

effective in finding occurrences of many rare plant 

species, it may not be effective for Botrychium simplex 

since it is difficult to find and identify. Searching for 

B. simplex requires one’s full attention, so attempts to 

search for this species are more likely to be successful 

if it is being sought along with other Botrychium 

species rather than during a general sensitive species 

inventory. Having experts (e.g., contractors, agency 

botanists, or others trained and experienced with 

searching for Botrychium species) conduct searches in 

appropriate habitat may be the most effective approach 

to expanding our knowledge of the distribution of this 

species in Region 2. 

Given the broad range of habitats that Botrychium 

simplex occupies, it is difficult to recommend specific

areas where searches are likely to locate new 

occurrences. Botrychium simplex is found over a wide 

elevation range and is erratically distributed, with less 

habitat fidelity than any other Botrychium species in 

Region 2. Searches that focus on areas that contain 

habitat types in which B. simplex has been previously 

found are most likely to yield new discoveries. Given 

its tendency to occur with other moonwort species, sites 

where other moonwort species have been documented 

should also be searched for B. simplex. Known 

locations for B. multifidum, in particular, warrant 

searching for B. simplex since these species are often 

found together (Crook personal communication 2003). 

This can be difficult, however, because B. multifidum is 

best seen in the fall after the leaves of B. simplex have 

already senesced. 

Recent surveys in Region 2 have been successful 

in locating new moonwort occurrences. Searches in the 

Bearlodge Ranger District of Wyoming and neighboring 

South Dakota identified numerous occurrences of 

Botrychium species (Crook personal communication 

2003, Burkhart personal communication 2006, Farrar 

personal communication 2006). The White River and 

Shoshone national forests are now known to support 

many occurrences of B. simplex, but additional surveys 

will likely find more. Despite recent discoveries of 

moonwort occurrences in Summit County, Colorado, 

more inventories are needed to document the 

distribution of B. simplex there since all potential habitat 

was not visited (Kolb and Spribille 2000). Zimmerman 

Lake and other sites in the Cameron Pass area are high 

priority search areas on the Arapaho-Roosevelt National 

Forest (Schwab 1992). 

Searches in apparently suitable habitats in 

Colorado have failed to identify new discoveries of 

Botrychium simplex (Popovich personal communication 

2006). Near the toll booth along the road to Mount 

Evans at least seven species of moonworts have been 

found, and there is abundant riparian habitat in the 

area that appears suitable for B. simplex; however, in 

repeated searches of this area by experts, no B. simplex 

has yet been found. Near the east portal of the Moffat 

Tunnel, there is also suitable habitat that has been 

searched without finding any B. simplex (Popovich 

personal communication 2006). These observations 

contrast sharply with moonwort survey efforts on the 

Black Hills National Forest in 2004 and 2005, where 

B. simplex was the most commonly found Botrychium 

species (Burkhart personal communication 2006). 

Botrychium simplex has a known affinity for 

plains habitats, and it is probably more abundant 

57 

in plains riparian areas than we realize. Given the 

difficulty in finding this species, it has likely just 

evaded discovery. Portions of major rivers on the plains 

of Region 2 that still have an active floodplain (e.g., 

Platte, Republican, Arikaree) should be included in 

inventory plans for B. simplex. 

Habitat inventory 

Aerial photographs, topographic maps, soil maps, 

and geology maps can be used to refine surveys of large 

areas. Use of these tools is most effective for species 

for which we have basic knowledge of its substrate and 

habitat specificity, such as Botrychium simplex, and 

from which distribution patterns and potential search 

areas can be deduced. 

The use of deductive and inductive techniques 

to model the distribution of Botrychium simplex would 

also aid in refining areas targeted in survey efforts in 

Region 2. Species distribution modeling is an effective 

means of determining the extent of suitable habitat 

on USFS lands. Classification and Regression Tree 

(CART) has been used to model the distribution of 

other sensitive plant species in Wyoming (e.g., Fertig 

and Thurston 2003) and Colorado (Decker et al. 2005). 

Combining CART with envelope models such as 

DOMAIN, BIOCLIM, or MaxEnt can help to refine a 

potential distribution map further by adding inference 

on the likelihood of the presence of B. simplex (Thuiller 

et al. 2003, Beauvais et al. 2004). CART Techniques 

for predicting species distributions are reviewed 

extensively by Scott et al. (2002). A problem with the 

models described above is that they do not account 

for ecologically relevant events that occurred in the 

past. Historic land use practices may have extirpated 

occurrences of B. simplex in Region 2, but without 

a geospatially explicit dataset of these practices, this 

possibility cannot be accounted for in the model. 

In general, the best search areas for all Botrychium 

species are places that have been disturbed in the past one 

to three decades (Wagner and Wagner 1976, Buell 2001, 

Johnson-Groh personal communication 2003, Johnson- 

Groh and Farrar 2003, Root personal communication 

2003, Whipple personal communication 2003). Wagner 

and Wagner (1976) included old pastures, pasturewoods, 

second-growth fields, edges of paths, old apple 

orchards, fencerows, floodplains subject to immersion, 

young pine woods, deserted homesteads, and natural 

deer pastures on a list of good areas to search. Wagner 

and Wagner (1976) noted that areas within 10 to 20 feet 

of the edges of woods are particularly lucrative sites 

to search, and specimen label data from Region 2 and

elsewhere support this statement. Wetlands, riparian 

areas, floodplains, hot springs, streamsides, peaty 

sites, wet slopes, avalanche meadows, wet meadows, 

colluvial slopes, and sparsely vegetated sites above 

treeline are all potential search areas for B. simplex. 

The Community ecology and Habitat sections of this 

document contain discussions of other environmental 

correlates and commonly associated species. 

Soil moisture and texture are not useful for 

identifying potential habitat for Botrychium simplex. 

However, soil nutrient levels may be a useful filter 

tool in the selection of potential search areas. In four 

states outside of Region 2, soils that support moonwort 

communities averaged 916 ppm of calcium and 140 

ppm of magnesium (Hansen and Johnson-Groh 2003). 

The potential value of these and other edaphic factors 

in identifying habitat for B. simplex in Region 2 has not 

been assessed. 

Population monitoring 

There are several monitoring studies of 

Botrychium simplex from which a monitoring protocol 

could be developed for Region 2. Permanent plot 

monitoring of B. simplex in Iowa and Minnesota is 

described in Johnson-Groh and Farrar (1996b) and 

Johnson-Groh (1999). Many techniques that have been 

used to monitor occurrences of other Botrychium species 

(e.g., Montgomery 1990, Muller 1992, Kelly 1994, 

Johnson-Groh and Lee 2002) could also be suitable 

for monitoring B. simplex, depending on the situation 

and the question to be answered by monitoring. The 

Pike-San Isabel National Forest (Carpenter 1996) has 

drafted protocols for monitoring B. lineare, and these 

may be useful for B. simplex. Johnson-Groh and Lee 

(2002) have studied the phenology and demography 

of B. gallicomontanum and B. mormo, using methods 

that could be applied to B. simplex. These methods are 

presented in the Botrychium Inventory and Monitoring 

Technical Guide (Johnson-Groh and Farrar 2003), 

which has been designed for members of subgenus 

Botrychium. Using the protocols described by Johnson- 

Groh and Farrar (2003) would facilitate the comparison 

of monitoring data from sites across the United States. 

The sampling strategy recommended by Johnson- 

Groh and Farrar (2003) involves the deployment of 

permanently-marked 1 square meter plots within a 

moonwort occurrence. The number of plots needed will 

depend on a power analysis completed after at least 

two years’ of data have been recorded. These plots are 

established non-randomly to capture the variation in 

aspect, drainage, canopy cover, and other environmental 

58 

variation present at the site, and they are oriented northsouth. 

The plots may contain one or numerous species 

of moonworts. Within each plot, a numbered tag is 

installed 2.5 cm north of each moonwort. Searching 

for plants within the plot may require litter removal 

because small plants are often unable to emerge through 

litter. All plants are measured from the base of the plant 

to the top of the tropophore. The developmental stage 

of each sporophyte is recorded (e.g., just emerging, 

releasing spores). Disturbances such as herbivory are 

also documented for each plant or plot. Johnson-Groh 

(1999) notes that it can be difficult to be assured that an 

individual that had been marked in a previous year is the 

same individual in subsequent years. See Johnson-Groh 

and Farrar (2003) for details regarding these protocols. 

For species such as Botrychium simplex where 

the proportion of dormant plants may vary among 

years, it is difficult to monitor population trends (Lesica 

and Steele 1994). However, recent advances have 

been made in mark-recapture methods for species that 

exhibit prolonged dormancy or are difficult to find in 

concealing vegetation (Alexander et al. 1997). These 

methods, which have been applied to plants, may be 

useful for estimating B. simplex populations. 

Annual monitoring of selected occurrences of 

Botrychium simplex in Region 2 could provide data 

that will improve our understanding of its ecology 

and population trends. Population trend monitoring 

protocols pertaining to members of subgenus 

Botrychium are defined in Johnson-Groh and Farrar 

(2003). Counting the emergent sporophytes at each 

occurrence every year would provide valuable data on 

the status of the species. In small occurrences, the entire 

population should be counted; in large occurrences, a 

randomized permanent plot technique could be used 

to monitor a representative sample of the occurrence 

(Chadde and Kudray 2001b). Tracking individuals by 

marking or mapping them within each sampling unit 

could provide information on the life span, dormancy, 

recruitment success, and population trends of this 

species (Kolb and Spribille 2000). It is important to note 

that monitoring Botrychium occurrences is difficult due 

to their curious life history attributes and small size, and 

tens of years of data will be needed to draw meaningful 

inferences (Johnson-Groh and Farrar 2003). Monitoring 

occurrences in Region 2, with return intervals of not 

more than 10 years, is a high conservation priority 

(Kolb and Spribille 2000). 

Adding a photopoint and photoplot component 

to this work, following recommendations presented in 

Elzinga et al. (1998) and Hall (2002), will facilitate the

tracking of individuals and add valuable qualitative 

information. These techniques can be done quickly 

in the field, and although they do not provide detailed 

cover or abundance data, they can help to elucidate 

patterns observed in quantitative data. 

Presence/absence monitoring is not suitable 

for Botrychium simplex due to the small occurrence 

sizes in Region 2. Gathering occurrence data can be 

done rapidly and may require only a small amount of 

additional time and effort (Elzinga et al. 1998). 

Beneficial management actions 

Because many additional occurrences of 

Botrychium simplex have likely not been found in 

Region 2, surveys prior to management actions within 

potential habitat will help to identify new occurrences 

and to alleviate anthropogenic threats to this species 

(Lorain 1990). The value of surveys before construction 

is evinced by the discovery of B. simplex on Vail Pass 

at a site where a ski hut was to be built. The pre-project 

survey probably reduced the impacts to this occurrence. 

Identifying large, high quality occurrences through 

surveys and monitoring will also help managers to 

prioritize conservation actions. Developing a better 

understanding of the species’ centers of distribution will 

assist with the development of regional management 

protocols that favor the persistence of B. simplex. 

There is no evidence of any direct impacts to 

occurrences of Botrychium simplex from recreation, 

but impacts on this species are difficult to document, 

as shown by the fact that no occurrence of B. simplex in 

Region 2 has been seen more than once. Occurrences in 

ski areas are most likely to incur the loss of individuals 

due to summer recreational use of ski slopes. Off-road 

motorized vehicle use also has the potential to affect 

B. simplex. Hiking may result in trampling where B. 

simplex occurs near trails or backcountry huts. Signage 

or temporary fences that could be removed in the 

winter could help dissuade recreational use in known 

occurrences of B. simplex. Mitigating recreation impacts 

to B. simplex may be important at some occurrences. 

Preventing the alteration of the hydrology in 

Botrychium simplex occurrences is an important 

management consideration. This is probably most 

important in wetland sites such as seepy or wet 

meadows, streamsides, floodplains, and wet slopes 

(e.g., #1, 9, 13, 23, 25, 29, and 49 in Table 8). 

The observations of Muller (1999) suggest 

that maintaining occasional perturbations by humans 

59 

or animals recreates pioneer habitats that benefit 

Botrychium matricariifolium. Thompson (2001) noted 

that the clearing of ski runs created a great deal of 

suitable habitat for moonworts, which are not found 

under tree canopies. While this may be true, ski runs 

cannot be depended upon for the long-term conservation 

of moonworts since they are managed for the benefit 

of skiers, not moonworts. For example, installing 

pipelines in ski runs for the production of artificial snow 

will have negative impacts on plants in the ski runs. It 

is not known if the autecological parameters needed 

by moonworts (e.g., appropriate mycorrhizal hosts, 

suitable disturbance regime) can persist indefinitely in 

ski runs. Kolb and Spribille (2000) recommend that B. 

simplex occurrences should be protected from grounddisturbing 

activities. More research is needed on the 

practical application of management protocols for the 

maintenance of moonwort occurrences. 

The utility of fire as a habitat management 

tool for Botrychium simplex is not known. Evidence 

suggests that the direct impacts of fire are not 

detrimental to Botrychium (Johnson-Groh and Farrar 

2003). Fire can slow succession to closed canopy and 

create open habitats for B. simplex. It can also alter 

soil characteristics in ways that may favor B. simplex. 

Burning actually appears to have had positive effects 

on B. campestre, B. gallicomontanum, and B. simplex 

occurrences in Iowa, but fire combined with erosion 

and desiccation, both natural results of fire, may be 

deleterious. There were no significant differences 

in plant size in burned and unburned plots at these 

locations, but fires occurring after drought have resulted 

in population decline (Johnson-Groh and Farrar 1996b, 

Johnson-Groh 1999). The role of fire in the autecology 

of Lycopodium suggests that it might also play a role 

in the autecology of B. simplex since they have similar 

life histories. 

Beneficial management actions regarding 

grazing are unclear, but there is evidence to suggest 

that Botrychium simplex may tolerate some grazing. 

Johnson-Groh and Farrar (2003) wrote: “Managers 

must not arbitrarily increase or decrease grazing because 

of the moonworts. Understanding the history of land 

management, including frequency of grazing, number 

of grazing animals, and timing of grazing will allow 

managers to determine appropriate levels of grazing to 

maintain populations. Removing grazing or increasing 

grazing cannot be expected to maintain populations.” 

The Community ecology and Threats sections of this 

document have more information on grazing.

Any management strategies that protect 

occurrences of Botrychium simplex from invasion by 

exotic species will confer the greatest benefits. Because 

herbicides are known to kill this species, their use 

within occurrences of B. simplex should be limited 

to direct application to the target species. Aggressive 

management of noxious weeds before they become 

dominant in B. simplex occurrences could avert costly 

and risky eradication efforts. Where possible, handpulling 

should be the favored method of managing 

weeds within B. simplex occurrences. 

Mitigating threats to occurrences of Botrychium 

simplex from intensive land use practices (i.e., offroad 

vehicle use) will confer benefits to the species. 

Controlling motorized access to habitat and providing 

appropriate signage at access points may decrease 

impacts to B. simplex. 

Populations of Botrychium are inherently 

variable (Johnson-Groh 1999). Many occurrences are 

small, increasing the likelihood of local extirpation. 

Botrychium simplex may depend on a shifting mosaic 

of suitable habitats in appropriate successional 

stages that can be colonized (as described by Pickett 

and Thompson 1978). If this is the case, then the 

metapopulation dynamics of B. simplex become crucial 

to its management and conservation, and underscore the 

need to conserve nearby areas of suitable habitat that 

are not currently inhabited by B. simplex (Chadde and 

Kudray 2001b). 

Restoration 

Growing any Botrychium species in the greenhouse 

or lab is extremely difficult (Whittier 1972, Gifford and 

Brandon 1978). Botrychium simplex is hardy to USDA 

Zone 3 (average minimum annual temperature –40 ºF) 

and it can be propagated from spores with difficulty, but 

it is not generally cultivated (Rook 2002). No spores of 

B. simplex are currently in storage at the National Center 

for Genetic Resource Preservation (Miller personal 

communication 2003). Collection of spores for longterm 

storage may be useful for future restoration work. 

Herbarium specimens and wild populations (within 

limits) may also provide propagules for restoration. 

While Botrychium species are generally very difficult 

to transplant, B. virginianum transplants were relatively 

successful (Wagner and Wagner 1976). Moonworts 

have been transplanted in Summit County, Colorado in 

sites where pipelines passed through occurrences, but 

it is not known if these efforts were successful (Kolb 

60 

and Spribille 2000, Buell 2001). Buell (2001) details 

moonwort transplant protocols that may be useful for 

B. simplex. 

Information Needs and Research 

Priorities 

Kolb and Spribille (2000) offer suggestions for 

research on Botrychium simplex. Vanderhorst (1997) 

also offers suggestions for research and management 

for occurrences on the Kootenai National Forest. 

Research needs for other moonworts are cited by Farrar 

and Johnson-Groh (1986), Berlin et al. (1998), and 

Johnson-Groh (1999), and because of similarities in the 

life history and ecological needs of Botrychium species, 

many of these apply to B. simplex as well. 

Surveys are among the greatest research needs 

for Botrychium simplex in Region 2. As concern and 

awareness of Botrychium species has increased, more 

inventories have focused on finding these species. 

Consequently, there are many recent discoveries of B. 

simplex and other Botrychium species in Region 2. It 

is likely that occurrences of this and other moonwort 

species remain to be discovered, and many are likely 

to be on USFS land. Better habitat data may help to 

evaluate the environmental tolerances of B. simplex 

(Farrar 2001). While current data portray the range of 

habitats in which Botrychium simplex is found in Region 

2, research is needed to develop an understanding of the 

ecosystem processes on which it depends. Some habitat 

attributes are commonly seen in Region 2 occurrences, 

but the relative importance of these attributes to the 

establishment and persistence of B. simplex is unknown. 

Habitat data will also support the development of 

potential habitat maps, which would be a valuable tool 

in locating new occurrences (Kolb and Spribille 2000). 

The occurrence of Botrychium simplex in 

saturated fen habitats is not well documented, and 

research is needed to assess the importance of this 

habitat (Farrar 2001, Root personal communication 

2003). Saturated fen habitats also need to be searched 

for other moonworts, including B. montanum, B. 

minganense, B. crenulatum, and B. pinnatum (Farrar 

2001, Root personal communication 2003). Botrychium 

crenulatum has not been found in Colorado but it may 

grow in fens (Root personal communication 2003). 

Searches of calcareous fens and seeps for var. fontanum 

within Region 2 are also needed. Other potential habitats 

for B. simplex such as avalanche chutes (ecologically 

similar to ski runs) have not been adequately searched.

A replicated demographic monitoring study 

that compares vital rates of Botrychium simplex in 

burned vs. unburned, grazed vs. ungrazed, shaded 

vs. unshaded, and weedy vs. unweedy sites would 

answer many questions about this species. This is a 

high priority for determining appropriate management 

practices with respect to B. simplex. Monitoring of 

occurrences is needed to better understand life history 

characteristics including age, dormancy, growth rates, 

and reproductive rates in Region 2. In particular, 

the subterranean portion of this species’ life cycle 

remains poorly understood, despite the fact that much 

of its lifespan occurs underground. More information 

regarding this species’ reproductive rates and its ability 

to colonize new sites would improve our understanding 

of this species’ response to change, but these remain 

highly speculative. 

There are no data in Region 2 from which 

population trend can be determined. Johnson-Groh 

and Farrar (2003) have developed and successfully 

used standard population trend monitoring protocols 

for Botrychium species. Other sampling designs have 

been used in the study of Botrychium species as well 

(e.g., Lesica and Steele 1994, Berlin et al. 1998, 

Johnson-Groh 1999). The problem with any of these 

methodologies is that it is very difficult to determine the 

total number of plants in any Botrychium occurrence 

due to the high proportion of plants residing under 

ground as gametophytes and juvenile sporophytes, 

and due to the annual variation in the aboveground 

sporophyte population (Lesica and Steele 1994). There 

are apparently no published data on the prevalence of 

dormancy in B. simplex. Due to the inherent difficulties 

in monitoring Botrychium occurrences, determination 

of population trend requires decades of study (Johnson- 

Groh and Farrar 2003). Periodic monitoring is needed 

to assess population trends at significant moonwort 

occurrences (Kolb and Spribille 2000). 

Although current research suggests that 

Botrychium simplex sensu lato is a valid taxon, 

better circumscription of the varietal taxa in Region 

2 is needed to understand their distribution, relative 

abundance, and habitat affinities (Farrar 1998, Farrar 

2001, Farrar personal communication 2003). Recent 

and ongoing research has resulted in a clearer picture 

of the taxonomically relevant variability within B. 

simplex. Analyzing isozymes, as in Farrar (1998) and 

Farrar (2001), and using DNA sequences, as employed 

by Hauk (1995) and Hauk et al. (2003), is one approach 

to sorting out the distribution and habitat affinities of the 

varieties in Region 2. The distribution of varieties of B. 

61 

simplex is very poorly understood but is important from 

a management and conservation perspective. 

Developing a better understanding of the 

ecological requirements of Botrychium simplex is 

important for its conservation and management. 

Research on the autecology of B. simplex, particularly 

its responses to fire, grazing, disturbance, and 

succession, is needed. The species’ response to 

water table fluctuation, inundation, and drought are 

particularly relevant throughout Region 2. The effects 

of herbivores and exotic species on the viability of B. 

simplex occurrences have not been investigated. More 

research is needed to determine the types, intensities, 

and periodicity of disturbance that create and maintain 

suitable habitat for Botrychium species, including B. 

simplex. Habitat manipulation studies (i.e., addition 

of artificial snow to plots, soil scarification) in robust 

moonwort occurrences could help to identify those 

ecological factors with the greatest influence on the 

distribution and persistence of B. simplex (Kolb and 

Spribille 2000). The amount of disturbance that B. 

simplex can tolerate is not known but is important to 

managers. Current observations of its response to both 

natural and anthropogenic disturbance in Region 2 are 

informal, and quantitative data are needed to understand 

the role of disturbance in the life history, establishment, 

and persistence of B. simplex. 

A clearer understanding of the relationship 

between Botrychium simplex and its mycorrhizal 

symbionts is important. An assessment of the effects 

of disturbance type and periodicity, especially for fire, 

and grazing, on B. simplex’s mycorrhizal symbionts will 

assist managers in developing appropriate management 

protocols. Research is also needed to assess the effect 

of different mycorrhizal species and infection levels 

on spore output. Studies of the role of mammals and 

other potential spore dispersal agents will assist with the 

management of B. simplex. 

Metapopulation studies are very difficult to 

conduct for Botrychium species, but it is likely that the 

metapopulation structure is important (Johnson-Groh 

and Farrar 2003). Investigation of migration, extinction, 

and colonization rates could yield valuable data for 

the conservation of B. simplex and other Botrychium 

species. Gene flow among and within populations 

of B. simplex could be investigated by analyzing the 

frequency of molecular-genetic markers. 

Identifying habitat characteristics through 

vegetation plot sampling would add analytical power

to the assessment of Botrychium simplex habitat. Kolb 

and Spribille (2000) used releves, but other quantitative 

methods would apply as well. Estimating cover and/or 

abundance of associated species is needed to begin 

the investigation of interspecific relationships through 

ordination or other statistical techniques. 

Understanding environmental constraints on 

Botrychium simplex could facilitate the conservation of 

this species. Gathering data on edaphic characteristics 

(e.g., moisture, texture, chemistry, particularly pH) 

from the vegetation or demographic monitoring plots 

described above would permit the canonical analysis 

of species-environment relationships. These data would 

facilitate hypothesis generation for further studies of the 

ecology of this species. 

There are many barriers to habitat and population 

restoration for Botrychium simplex. Botrychium 

species are extremely difficult to propagate (Whittier 

1972, Gifford and Brandon 1978, Wagner and Wagner 

1983), and propagating them for reintroduction to 

the wild is probably not feasible. The subterranean 

ecology of these species is crucial to understanding 

their dynamics, yet it is very poorly understood. As 

obligate mycorrhizal hosts, they cannot survive without 

fungal partners, but very little is known about the 

specifics of this relationship. The difficulties in growing 

Botrychium species are presumably the result of their 

delicately attuned mycorrhizal relationships (Wagner 

and Wagner 1983). The mycobionts of B. simplex have 

not been identified. Restoration or maintenance of 

native vegetation will be a crucial part of any restoration 

of B. simplex. 

Buell (2001, page 11) describes a method for 

transplanting Botrychium species that was used by 

Nancy Redner of the USFS. This method was used 

to mitigate pipeline and road construction impacts on 

Botrychium occurrences at the Copper Mountain Ski 

62 

Resort in Colorado. No data on survivorship of the 

transplanted plants are available, but Buell (2001) 

described this method as “reasonably successful.” 

Several moonwort species, including B. echo, were 

transplanted in 2003 to mitigate construction impacts 

along the Guanella Pass Road (ERO Resources 

Corporation 2003). Their methods involved excavating 

a large area of soil around each plant or cluster of plants 

and not allowing the soil clump to break. Plants and 

their soil were transported using spring-form baking 

pans with removable bottoms. Popovich (personal 

communication 2005) estimated that up to 50 percent 

of the transplants survived into 2004. Cody and Britton 

(1989) note that transplanting Botrychium species is 

usually fatal. Because there has not been any longterm 

assessment of the success of transplants, the value 

of this practice for conservation is questionable and 

cannot be relied upon as a mitigation tool. Monitoring 

transplants could determine whether these techniques 

can be successful (Kolb and Spribille 2000). 

Additional research and data resources 

Farrar, Ahlenslager, and Johnson-Groh are 

currently assessing a suite of Botrychium species for 

Washington and Oregon. Johnson-Groh and Farrar 

(2003) are drafting monitoring protocols for Botrychium 

species. Dr. Reed Crook is planning to conduct habitat 

modeling for B. multifidum in the Black Hills (Crook 

personal communication 2003); this will apply to B. 

simplex since they are often found together. 

Dr. Farrar plans to assess specimens from Region 

2 using molecular phylogenetic techniques. This 

will result in a clearer picture of the distribution of 

Botrychium simplex varieties in Region 2. It will also 

determine whether the undescribed variety collected 

in the southwestern United States is also found in 

Colorado. The habitat differences among the varieties 

in Region 2 will be apparent with more inventories.

DEFINITIONS 

Achlorophyllous – A plant lacking chlorophyll and thus dependent on obtaining carbon from a host or symbiont. 

Allopolyploid – A polyploid formed from the union of genetically distinct chromosome sets, usually two different 

species (Allaby 1998). 

Antheridium – The male sex organ of the gametophyte, where male sex cells are produced by mitosis (Allaby 

1998). 

Archegonium – The female sex organ of the gametophyte, where female sex cells are produced by mitosis (Allaby 

1998). 

Bobby sox trees – Lodgepole pine trees that, having died due to flooding, act as wicks that absorb geothermal water. 

As this water evaporates, it leaves silicon dioxide behind, which colors the base of the trunks white (hence the moniker 

“bobby sox trees”) (Whipple personal communication 2003). 

Congener – A member of the same genus. Botrychium simplex is a congener of B. multifidum. 

Conservation Status Rank – The Global (G) Conservation Status (Rank) of a species or ecological community 

is based on the range-wide status of that species or community. The rank is regularly reviewed and updated by 

experts, and takes into account such factors as number and quality/condition of occurrences, population size, range of 

distribution, population trends, protection status, and fragility. A subnational (S) rank is determined based on the same 

criteria applied within a subnation (state or province). The definitions of these ranks, which are not to be interpreted 

as legal designations, are as follows: 

GX Presumed Extinct – Not located despite intensive searches and virtually no likelihood of 

rediscovery 

GH Possibly Extinct – Missing; known only from historical occurrences but still some hope of 

rediscovery 

G1 Critically Imperiled – At high risk of extinction due to extreme rarity (often five or fewer occurrences), 

very steep declines, or other factors. 

G2 Imperiled – At high risk of extinction due to very restricted range, very few populations (often 20 or 

fewer), steep declines, or other factors. 

G3 Vulnerable – At moderate risk of extinction due to a restricted range, relatively few populations (often 

80 or fewer), recent and widespread declines, or other factors. 

G4 Apparently Secure – Uncommon but not rare; some cause for long-term concern due to declines or 

other factors. 

G5 Secure – Common; widespread and abundant. 

Competitive/Stress-tolerant/Ruderal (CSR) model – A model developed by J.P. Grime in 1977 in which plants are 

characterized as Competitive, Stress-tolerant, or Ruderal, based on their allocation of resources. Competitive species 

allocate resources primarily to growth; stress-tolerant species allocate resources primarily to maintenance; ruderal 

species allocate resources primarily to reproduction. A suite of other adaptive patterns also characterize species under 

this model (Barbour et al. 1987). Some species, including Botrychium simplex, show characteristics of more than one 

strategy. 

Ectomycorrhiza – A type of mycorrhiza where the fungal hyphae do not penetrate the cells of the root, but instead 

form a sheath around the root (Allaby 1998). 

Endomycorrhiza – A type of mycorrhiza where the fungal hyphae do penetrate the cells of the root. Arbuscular 

mycorrhizae are a type of endomycorrhizae (Allaby 1998). 

Eusporangiate – A primitive condition in which the cells that give rise to the sporangia are superficial (lie at the 

surface) (Gifford and Foster 1989). 

Gametophyte – The haploid stage in the life cycle of a plant. This stage lives independently of the sporophyte in ferns. 

In Botrychium the gametophyte is subterranean and is parasitic on mycorrhizal fungi (Gifford and Foster 1989). 

63

Gemma – A minute vegetative propagule abscised at maturity from the parent plant (Farrar and Johnson-Groh 

1990). 

Genus community – Several Botrychium species are commonly found growing together in close proximity. This is 

unusual in the plant world, since members of the same plant genus often do not occur together, probably because of 

competitive interactions that would occur between them. The Wagners coined the term “genus community” to describe 

these peculiar assemblages of Botrychiums (Wagner and Wagner 1983). 

Lamina – The leaf blade of a fern. In Botrychium, the lamina is divided into a fertile segment (the sporophore) and a 

sterile segment (the trophophore) (Lellinger 1985). 

Monoecious – Gametophytes with both male and female gamete producing structures. 

Mycobiont – The fungal partner in a mycorrhizal symbiosis. 

Nothospecies – A species of hybrid origin (International Association for Plant Systematics 2001). 

Ruderal – Plants with an adaptive suite of characteristics, including high reproductive rate, that makes them effective 

colonists and well-suited to disturbed habitats (Barbour et al. 1987). 

Sporophore – The fertile, spore-bearing portion of the leaf of Botrychium species (Foster and Gifford 1989). 

Sporophyte – The diploid portion of the life cycle of plants. Haploid spores are produced by meiosis that gives rise 

to gametophytes (Allaby 1998). 

Sympatry – The occurrence of species (or varieties in the case of Botrychium simplex) together in the same area 

(Allaby 1998). 

Trophophore – The vegetative portion of the leaf of Botrychium species (Foster and Gifford 1989). 

64

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