National Park Service

U.S. Department of the Interior

Natural Resource Stewardship and Science

Plant Community Composition and Structure 

Monitoring for Wind Cave National Park

2011-2015 Summary Report

Natural Resource Report NPS/NGPN/NRR—2016/1236
ON THE COVER

Long-term monitoring plot PCM-0028 at Wind Cave National Park, 2015

Photograph courtesy of the National Park Service.
Plant Community Composition and Structure 

Monitoring for Wind Cave National Park

2011-2015 Summary Report

Natural Resource Report NPS/NGPN/NRR—2016/1236

Isabel W. Ashton

Christopher J. Davis

National Park Service

Northern Great Plains Inventory & Monitoring Network

231 East St. Joseph Street

Rapid City, SD 57701

June 2016 

U.S. Department of the Interior

National Park Service

Natural Resource Stewardship and Science 

Fort Collins, Colorado
ii

The National Park Service, Natural Resource Stewardship and Science office in Fort Collins, 

Colorado, publishes a range of reports that address natural resource topics. These reports are of 

interest and applicability to a broad audience in the National Park Service and others in natural 

resource management, including scientists, conservation and environmental constituencies, and the 

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information is scientifically credible, technically accurate, appropriately written for the intended 

audience, and designed and published in a professional manner. Data in this report were collected 

and analyzed using methods based on established, peer-reviewed protocols and were analyzed and 

interpreted within the guidelines of the protocols.

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This report is available in digital format from the Northern Great Plains Inventory & Monitoring 

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report in a format optimized for screen readers, please email irma@nps.gov.

Please cite this publication as:

Ashton, I. W. and C. J. Davis. 2016. Plant community composition and structure monitoring for 

Wind Cave National Park: 2011-2015 summary report. Natural Resource Report NPS/NGPN/NRR—

2016/1236. National Park Service, Fort Collins, Colorado.

NPS 108/133334, June 2016 
iii

Contents 

Page

Figures...................................................................................................................................................iv

Tables.....................................................................................................................................................v

Executive Summary..............................................................................................................................vi

Acknowledgments...............................................................................................................................viii

Introduction............................................................................................................................................1

Methods..................................................................................................................................................2

NGPN and NGPFire Monitoring Plots 2011-2015 ........................................................................2

Other Monitoring Plots (1997-2015)..............................................................................................5

Data Management and Analysis.....................................................................................................6

Reporting on Natural Resource Condition .....................................................................................7

Results and Discussion ..........................................................................................................................9

Status & Trends in Vegetation Community Composition and Structure of WICA .......................9

Species Richness, Diversity, and Evenness.............................................................................13

Comparing Forest and Grassland Plots ...................................................................................16

The Influence of Climate and Fire on Plant Community Structure and Diversity .......................19

Rare Plants....................................................................................................................................22

Conclusion ...........................................................................................................................................23

Literature Cited ....................................................................................................................................25

Appendix A: List of plant species found at WICA 1998-2015............................................................28
iv

Figures 

Page

Figure 1. Map of Wind Cave National Park (WICA) plant community monitoring plots, 

1998-2015 ..............................................................................................................................................3

Figure 2. Long-term monitoring plot layout used for sampling vegetation in Wind Cave 

National Park. ........................................................................................................................................4

Figure 3. The Northern Great Plains Inventory & Monitoring vegetation crew used pointintercept (left panel and center) and quadrats (right panel) to document plant diversity 

and abundance........................................................................................................................................4

Figure 4. Average cover by lifeform of native (green) and exotic (red) plants recorded in 

monitoring plots in Wind Cave National Park (1998-2015)..................................................................9

Figure 5. The average absolute cover of the 10 most common native (green) and exotic 

(red) plants recorded at Wind Cave National Park in 1998-2015........................................................10

Figure 6. Trends in the relative cover of exotic plants and annual bromes in Wind Cave 

National Park from 1998-2015 ............................................................................................................11

Figure 7. A map of long-term monitoring plots and the relative cover of exotic plants.....................12

Figure 8. The relationship between native species richness and the relative cover of 

annual bromes......................................................................................................................................13

Figure 9. A photograph of long-term monitoring plot WICA_PCM_0007 and 

WICA_PCM_0018 which has a low and high diversity of native plant species, 

respectively. .........................................................................................................................................14

Figure 10. Trends in native species richness and evenness in Wind Cave National 

Park,1998-2015....................................................................................................................................15

Figure 11. Comparison of forest and grassland plots in Wind Cave National Park. Values 

are means ± one standard error. ...........................................................................................................17

Figure 12. Examples of dense forest plots (left), moderately dense forest plots (center) 

and grassland plots (right)....................................................................................................................18

Figure 13. Map comparing exotic species relative cover in forest and grassland plots in 

Wind Cave National Park. Plot marker size is proportional to the cover of exotic species 

observed in each plot, as described in the legend. ...............................................................................18

Figure 14. The total annual precipitation anomaly from 1998-2015 for Wind Cave 

National Park .......................................................................................................................................19

Figure 15. Map of recent fire history at Wind Cave National Park. ...................................................20

Figure 16. Native species richness and percent cover of annual bromes in plots with 

different fire histories...........................................................................................................................21

Figure 17. Examples of two rare species found in plant community monitoring plots at 

Wind Cave National Park ....................................................................................................................22
v

Tables 

Page

Table ES-1. Natural resource condition summary table for plant communities in Wind 

Cave National Park ..............................................................................................................................vii

Table 1. Exotic species surveyed for at Wind Cave National Park as part of the early 

detection and rapid response program within the Northern Great Plains Network. ..............................5

Table 2 Definitions of state and global species conservation status ranks*..........................................6

Table 3. Key to the symbols used in the Natural Resource Condition Table........................................8

Table 4. Average plant species richness in monitoring plots at Wind Cave National Park 

from 1998 to 2015................................................................................................................................13

Table 5. Rare species occurrence in Wind Cave National Park from 1998-2015...............................22

Table 6. Natural resource condition summary table for plant communities in Wind Cave 

National Park .......................................................................................................................................24
vi

Executive Summary

The Northern Great Plains Inventory & Monitoring Program and Fire Effects Program have been 

monitoring vegetation in Wind Cave National Park for over 18 years. While methods have changed 

slightly, this report summarizes data from 97 locations from 1998-2015. Below, we list the questions 

we asked and provide a summarized answer, for more details see the Results and Discussion section.

We conclude with a Natural Resource Condition Table (Table ES-1) that summarizes the current 

status and trends in a few key vegetation metrics. 

1. What is the current status of plant community composition and structure of WICA 

vegetation (species richness, cover, and diversity) and how has this changed from 1998-

2015? 

The ponderosa pine woodlands and mixed-grass prairie within the boundaries of WICA are

characterized by moderate to high native species richness. The closed canopy forest generally 

maintains lower species richness than the open grassland areas. We found no significant trends in 

native diversity or evenness from 1998 to 2015. Invasive annual brome grasses are an emerging 

threat in WICA, and keeping them in low abundance will help maintain native plant diversity. Other 

exotics, such as Kentucky bluegrass are more prevalent, but there is no evidence of an increasing 

trend in their cover. 

2. What are the differences and similarities in forested and grassland condition and trends?

We examined the status and trends in native species richness, exotic species cover, and litter cover 

between forest and grasslands plots in WICA. We found that grassland plots had higher native 

richness and higher cover of annual brome grasses but there was not a difference in the relative cover 

of exotic plant species. Despite some differences in species composition, trends over time appear 

consistent in grasslands and forested plots. This suggests that both ecotypes are responding similarly 

to climate and other external drivers. 

3. How do trends in vegetation correlate with climate and fire history?

The large variability in the climate of WICA has made it difficult to discern strong patterns linking 

temperature, precipitation, and plant community structure (e.g. exotic cover, species richness). A 

longer time series of vegetation data will make it easier to elucidate trends in the future. 

WICA has been using prescribed fire as a management tool since the 1970s. There was no difference 

in native diversity among plots that had burned recently and those that had not burned for over 25 

years. However, the sites which burned recently have a lower cover of annual brome grasses than 

sites that have not burned in over 20 years. Ongoing adaptive management programs and research 

will provide better guidance to the park on whether prescribed burns are an effective tool to reduce 

the cover of annual bromes. 
vii

4. What, if any, rare plants were found in WICA long-term monitoring plots? 

We identified 2 rare plant species in WICA between 1998 and 2015- nylon hedgehog cactus

(Echinocereus viridiflorus) and a moonwort species (Botrychium sp.). These plants are found in such 

low abundance and in such few plots, it will take many years to determine any trends in rare plant 

cover. We recommend a more targeted survey of rare plants be completed when funds are available. 

Table ES-1. Natural resource condition summary table for plant communities in Wind Cave National Park.

Current values are based on data from 2011-2015 and trends are based on data from 1998-2015. 

Indicator of 

Condition

Specific 

Measures

Current

Value 

(mean ± 

SE)

Reference 

Condition 

and Data 

Source

Condition 

Status/Trend

Rationale for Resource 

Condition

Upland Plant 

Community 

Structure and 

Composition

Native species 

richness (1m2

quadrats) 

11.1 ± 0.4 

species

8-18 species 

The ponderosa pine woodland 

and mixed-grass prairie plant 

communities within the 

boundaries of WICA are 

characterized by moderate to 

high native species richness that 

is within the natural range of 

variability (Symstad and Jonas 

2014). Native diversity and plant 

cover is higher in grassland plots 

than forested plots. Native 

evenness is high indicating one 

species is not dominating the 

community and it not changed 

significantly since monitoring 

began in 1998.

Evenness

(pointintercept 

transects)

0.77 ± 

0.006 

To be 

determined

Exotic Plant 

Early 

Detection and 

Management

Relative cover 

of exotic 

species 

20.9 ± 

1.4% < 10 % cover

Many areas of WICA have a high 

cover of exotic species. Kentucky 

bluegrass is the most 

widespread and abundant 

invasive plant species. There is 

no evidence of an increasing or 

decreasing trend in total exotic 

cover or Kentucky bluegrass 

cover. While still in low 

abundance, annual brome cover 

has shown an increasing trend 

since 1998.

Annual brome 

cover 

3.3 ± 0.4% < 10 % cover
viii

Acknowledgments 

We thank the authors of the NGPN Plant Community Monitoring Protocol, particularly A. Symstad, 

for outstanding guidance on data collection and reporting. Thank you to the current and past staff at 

WICA, particularly G. Schroeder and B. Burkhart, for providing logistical support and performing 

safety checks. The NGPN and NGPFire vegetation field crews collected all the data included in this 

report. We thank D. Swanson for providing data on fire history and A. Weaver for helping with the 

maps for this report. Comments from C. Wienk and T. Richardson improved this report. 
1

Introduction 

The Black Hills is a 1.5 million ha refuge of ponderosa pine (Pinus ponderosa) forest surrounded by 

the mixed-grass prairies of western South Dakota and eastern Wyoming. The Black Hills is a unique 

ecosystem composed of species from the western Rocky Mountains, eastern deciduous forests, 

northern boreal forests, and the surrounding Great Plains (Larson and Johnson 2007).Wind Cave 

National Park (WICA) is located in the southern Black Hills with a purpose to protect the unique 

Wind Cave resources and preserve and enhance the mixed-grass prairie and native wildlife, while 

providing for the enjoyment of the public. The 33,851 acres (13,700 ha) of WICA is a mosaic of 

ponderosa pine forest and mixed-grass prairie, with approximately 30% covered by ponderosa pine 

forests. While some areas have high exotic species cover, the native plant communities within the 

upland areas of WICA are considered to be in good condition (Komp et al. 2011). The Northern 

Great Plains Inventory & Monitoring Program (NGPN) began vegetation monitoring efforts in 

WICA in 2011 (Ashton et al. 2012). Vegetation monitoring protocols and plot locations were chosen 

to represent the entire park (based on the 2010 boundary) and to coordinate efforts with the Northern 

Great Plains Fire Ecology Program (NGPFire). NGPFire had been monitoring upland vegetation in 

WICA since 1997. There is a separate effort to monitor streambank condition because riparian 

communities are an important resource for the park (Burkhart and Kovacs 2013). Additional upland 

plots were added to represent the 2011 Addition to the park property. In this report, we use the data 

from 2011-2015 to assess the current condition of park vegetation and the data from 1998-2015 are 

used to look at longer-term trends. 

Using 18 years of plant community monitoring data in WICA, we explore the following questions: 

1. What is the current status of plant community composition and structure of WICA vegetation 

(species richness, cover, and diversity) and how has this changed from 1998-2015? 

2. What are the differences and similarities in forested and grassland condition and trends?

3. How do trends in condition correlate with climate, fire history, and grazing?

4. What, if any, rare plants were found in WICA long-term monitoring plots? 
2

Methods 

Two different methods and protocols have been used to monitor long-term vegetation plots at WICA

since 1997: the NGPN monitoring protocol (Symstad et al. 2012b, a) and the Fire Monitoring

Handbook (NPS 2003). Below, we briefly describe both methods, but focus on the NGPN monitoring 

protocol which is the current standard and was used to collect most of the data in this report. For 

more detail on either method, please see the protocol publications (cited above). 

NGPN and NGPFire Monitoring Plots 2011-2015

NGPN and NGPFire implemented a survey to monitor plant community structure and composition in 

WICA using a spatially balanced probability design (Generalized Random Tessellation Stratified 

[GRTS]; Stevens and Olsen 2003, 2004). Using a GRTS design, NGPN selected 35 randomly located 

sites within the park to become Plant Community Monitoring plots (PCM plots; Figure1). In 2011, 

the park acquired new property and we selected 10 randomly located sites within the addition using 

the above procedure. The NGPN visits 14 PCM plots within the original boundary every year using a 

rotating sampling scheme where 7 sites were visited in the previous year and 7 sites are new visits. 

After 5 years (2011-2015), most of the 35 PCM plots were visited at least twice between mid-June 

and mid- July. We had time to visit only 2 PCM plots within the new addition since 2012. When a 

PCM plot fell within an active burn unit, NGPFire added additional visits based on a 1, 2, 5, and 10 

year sampling schedule. NGPFire also established and monitored a number of new sites using the 

same GRTS sampling schema focused in active burn units (Fire FPCM plots). From 2011-2015, 25

FPCM plots were established. 

At each of the grassland sites we visited, we recorded plant species cover and frequency in a 

rectangular, 50 m x 20 m (0.1 ha), permanent plot (Figure 2). Data on ground cover, herb-layer 

height ≤ 2 m, and plant cover were collected on two 50 m transects (the long sides of the plot) using 

a point-intercept method (Figure 3). Species richness data from the point-intercept method were 

supplemented in the 20 NGPN plots with species presence data collected in five sets of nested square 

quadrats (0.01 m2

, 0.1 m2, 1 m2, and 10 m2) located systematically along each transect (Figure 2). 

When woody species were present, tree regeneration and tall shrub density data were collected within 

a 10 m radius subplot centered in the larger 50 m x 20 m plot (Figure 2). Trees with diameter at 

breast height (DBH) > 15 cm, located within the entire 0.1 ha plot, were mapped and tagged. For 

each tree, the species, DBH, status, and condition (e.g., leaf-discoloration, insect-damaged, etc.) were 

recorded. Dead and downed woody fuel load data were collected on two perpendicular, 100 ft (30.49 

m) transects with midpoints at the center of the plot (Figure 2), following Brown’s Line methods 

(Brown 1974, Brown et al. 1982). For this report, we do not provide results on the fuels and forest 

structure data that were collected on an annual basis because we have a limited sample and because 

we completed a more large forest survey using 90 plots in 2012 (Ashton et al. 2013) and plan to 

repeat the survey in 2017.
3

Figure 1. Map of Wind Cave National Park (WICA) plant community monitoring plots, 1998-2015. Thirtyseven PCM plots (red) were established by the Northern Great Plains Inventory & Monitoring Program 

(NGPN) and 25 (blue) FPCM plots were established by the Fire Effects Program (NGPFire) between 

2011 and 2015. An additional 35 FMH plots (green) were monitored from 1997-2011 by NGPFire. 
4

Figure 2. Long-term monitoring plot layout used for sampling vegetation in Wind Cave National Park.

Figure 3. The Northern Great Plains Inventory & Monitoring vegetation crew used point-intercept (left 

panel and center) and quadrats (right panel) to document plant diversity and abundance. 

At all PCM plots, but not the FPCM plots, we also surveyed the area for common disturbances and 

target species of interest to the park. Common disturbances included such things as prairie dog 

towns, rodent mounds, animal trails, and fire. For all plots, the type and severity of the disturbances 

were recorded. We also surveyed the area for exotic species that have the potential to spread into the 
5

park and cause significant ecological impacts (Table 1). These species were chosen in collaboration 

with the Midwest Invasive Plant Network, the Exotic Plant Management Team, park managers, and 

local weed experts. For each target species that was present at a site, an abundance class was given 

on a scale from 1-5 where 1 = one individual, 2 = few individuals, 3 = cover of 1-5%, 4 = cover of 5-

25%, and 5 = cover > 25% of the plot. The information gathered from this procedure is critical for 

early detection and rapid response to such threats. 

Table 1. Exotic species surveyed for at Wind Cave National Park as part of the early detection and rapid 

response program within the Northern Great Plains Network. 

Scientific Name Common Name Habitat

Alliaria petiolata garlic mustard Riparian

Polygonum cuspidatum; P. sachalinense; P. x bohemicum knotweeds Riparian

Pueraria montana var. lobata kudzu Riparian

Iris pseudacorus yellow iris Riparian

Ailanthus altissima tree of heaven Riparian

Lepidium latifolium perennial pepperweed Riparian

Arundo donax giant reed Riparian

Rhamnus cathartica common buckthorn Riparian

Heracleum mantegazzianum giant hogweed Riparian

Centaurea solstitialis yellow star thistle Upland

Hieracium aurantiacum; H. caespitosum orange and meadow hawkweed Upland

Isatis tinctoria Dyer's woad Upland

Taeniatherum caput-medusae medusahead Upland

Chondrilla juncea rush skeletonweed Upland

Gypsophila paniculata baby's breath Upland

Centaurea virgata; C.diffusa knapweeds Upland

Linaria dalmatica; L. vulgaris toadflax Upland

Euphorbia myrsinites & E. cyparissias myrtle spurge Upland

Dipsacus fullonum & D. laciniatus common teasel Upland

Salvia aethiopis Mediterranean sage Upland

Ventenata dubia African wiregrass Upland

Other Monitoring Plots (1997-2015)

In 1997, NGPFire began monitoring plots within WICA to evaluate the effectiveness of prescribed 

burns. Starting in 1998, data collection followed the NPS National Fire Ecology Program protocols 

(NPS 2003): in grassland plots vegetation cover and height data were collected using a pointintercept method, with 100 points evenly distributed along a single 30 m transect. In forested sites, 

plots are 0.1 ha (20 x 50 m) in size and point-intercept data was collected along the two 50 m sides.

For each live tree with a DBH > 15 cm located within the 0.1 ha plot, the species and DBH were 

recorded. The densities of smaller trees (2.54 cm ≤ DBH ≤ 15 cm) were measured within a subset of 

the plot area. NGPFire plot locations were located randomly within major vegetation types within 

areas planned for prescribed burning (burn units) in the near future. The data were not collected using 

these protocols in 1997, so this year is excluded from analyses. Hereafter, we refer to these 35 plots 
6

as Fire Monitoring Handbook (FMH) plots. These FMH plots are being retired after 10 years of 

monitoring is completed (e.g. the rebar will be removed). 

Data Management and Analysis

We used FFI (FEAT/FIREMON Integrated; http://frames.gov/ffi/) as the primary software 

environment for managing our sampling data. FFI is used by a variety of agencies (e.g., NPS, USDA 

Forest Service, U.S. Fish and Wildlife Service), has a national-level support system, and generally 

conforms to the Natural Resource Database Template standards established by the Inventory and 

Monitoring Program. Species scientific names, codes, and common names are from the USDA Plants 

Database (USDA-NRCS 2015). However, nomenclature follows the Integrated Taxonomic 

Information System (ITIS) (http://www.itis.gov). In the few cases where ITIS recognizes a new name 

that was not in the USDA PLANTS database, the new name was used, and a unique plant code was 

assigned. This report uses common names after the first occurrence in the text, but scientific names 

can be found in the Appendix A. Plant life forms (e.g., shrub, forb) were based on definitions from 

the USDA Plants Database (USDA-NRCS 2015). The conservation status ranks of plant species in 

South Dakota is determined by the South Dakota Natural Heritage Program (SDNHP). For the 

purpose of this report, a species was considered rare if its conservation status rank was S1, S2, or S3. 

See Table 2 for a detailed definition of each conservation status rank.

After data for the sites were entered, 100% of records were verified to the original data sheet to 

minimize transcription errors. A further 10% of records were reviewed a second time. After all data 

were entered and verified, automated queries were used to check for errors in the data. When errors 

were caught by the crew or the automated queries, changes were made to the original datasheets 

and/or the FFI database as needed. Summaries were produced using the FFI reporting and query tools 

and statistical summaries, and graphics were generated using R software (version 3.2.2). In most 

cases, linear mixed models were used to analyses for differences across time or treatments, where 

plot was considered a random factor. 

Table 2 Definitions of state and global species conservation status ranks*.

Status 

Rank Category Definition

S1/G1 Critically imperiled Due to extreme rarity (5 or fewer occurrences) or other factor(s) making it 

especially vulnerable to extirpation.

S2/G2 Imperiled Due to rarity resulting from a very restricted range, very few populations (often 20 

or fewer), steep declines, or other factors making it very vulnerable to extirpation.

S3/G3 Vulnerable Due to a restricted range, relatively few populations (often 80 or fewer), recent 

widespread declines, or other factors making it vulnerable to extirpation. 

S4/G4 Apparently secure Uncommon but not rare; some cause for concern due to declines or other factors.

S5/G5 Secure Common, widespread and abundant.

S#S#/

G#G#

Range rank

(e.g. S2S3)

Used to indicate uncertainty about the status of the species or community. 

Ranges cannot skip more than one rank.

*Adapted from NatureServe status assessment table (http://www.natureserve.org/conservationtools/conservation-status-assessment)
7

We measured diversity at the plots in 2 ways: species richness and Pielou’s Index of Evenness. 

Species richness is simply a count of the species recorded in an area. Peilou’s Index of Evenness, J’, 

measures how even abundances are across taxa. It ranges between 0 and 1; values near 0 indicate 

dominance by a single species and values near 1 indicate nearly equal abundance of all species 

present. Plant richness was calculated for each plot using the total number of species intersected 

along the transects. Average height was calculated as the average height per plot using all species 

intersected on the transects.

Climate data from the Wind Cave National Park, SD weather station were downloaded from 

NOAA’s online database (NOAA 2015). A fire history map was compiled for WICA and crossreferenced with plot locations. For each plot visit, we determined the number of years since it burned 

and the number of recorded fires. Plots where no burns were recorded, were excluded from the 

analysis. 

Reporting on Natural Resource Condition

Results were summarized in a Natural Resource Condition Table based on the templates from the 

State of the Park report series (http://www1.nrintra.nps.gov/im/stateoftheparks/index.cfm). The goal 

is to improve park priority setting and to synthesize and communicate complex park condition 

information to the public in a clear and simple way. By focusing on specific indicators, such as exotic 

species cover, it will also be possible and straightforward to revisit the metric in subsequent years. 

The status and trend of each indicator is scored and assigned a corresponding symbol based on the 

key found in Table 3. 

We chose a set of indicators and specific measures that can describe the condition of vegetation in 

the Northern Great Plains and the status of exotic plant invasions. The measures include: native 

species richness, evenness, relative cover of exotic species, and annual brome cover. Reference 

values were based on descriptions of historic condition and variation, past studies, and/or 

management targets. Current park condition was compared to a reference value, and status was 

scored as good condition, warrants moderate concern, or warrants significant concern based on this 

comparison (Table 3). Good condition was applied to values that fell within the range of the 

reference value, and significant concern was applied to conditions that fell outside the bounds of the 

reference value. In some cases, reference conditions can be determined only after we have 

accumulated more years of data. When this is the case, we refer to these as “To be determined” and 

estimate condition based on our professional judgment.
8

Table 3. Key to the symbols used in the Natural Resource Condition Table. The background color 

represents the current status, the arrow summarizes the trend, and the thickness of the outside line 

represents the degree of confidence in the assessment. A symbol that does not contain an arrow 

indicates that there is insufficient information to assess a trend. Based on the State of the Park reports 

(http://www.nps.gov/stateoftheparks/).

Condition Status Trend in Condition

Confidence in 

Assessment

Warrants 

Significant Concern Condition is Improving High

Warrants 

Moderate Concern Condition is Unchanging Medium

Resource is in Good 

Condition Condition is Deteriorating Low
9

Results and Discussion

Status & Trends in Vegetation Community Composition and Structure of WICA 

The plant species list for WICA includes 871 species and we found 424 plant species from 1998-

2015 at WICA (Appendix A). Graminoids, which includes grasses, sedges, and rushes, accounted for 

most of the vegetative cover at WICA, but forbs, shrubs, trees, and vines were also present (Figure

4). We found 73 exotic plant species at WICA and none were the targeted early detection exotics 

(Table 1). Exotic graminoids were particularly abundant (Figure 4). The trees and subshrubs were all 

native species.

Figure 4. Average cover by lifeform of native (green) and exotic (red) plants recorded in monitoring plots 

in Wind Cave National Park (1998-2015). Absolute cover can be greater than 100% because the pointintercept methods records layers of overlapping vegetation. 

Western wheatgrass (Pascopyrum smithii) was the most abundant native grass and averaged close to

20% absolute cover (Figure 5). Other common native grasses include little bluestem (Schizachyrium 

scoparium), threadleaf sedge (Carex filifolia), needle and thread (Hesperostipa comata), and big 

bluestem (Andropogon gerardii). Kentucky bluegrass (Poa pratensis) was the most pervasive exotic 

at WICA. The average percent cover of exotic plant species in WICA fluctuates from year to year

(Figure 6), and there was very little exotic cover in 2008. Currently (2011-2015), the relative cover of 

exotic species at long-term plots in WICA averages 20.9 ± 1.4 % (mean ± one standard error). 

However, there is a great deal of variation across the park and some sites have few exotic species 

while others are heavily invaded (Figure 7). Much of this cover is Kentucky bluegrass, which 

averaged 11.9 ± 1.0 % from 2011-2015 and there is no evidence that it has decreased or increased 

since 1998. Kentucky bluegrass is a cool-season perennial grass that has recently been increasing in 

abundance in and around the Northern Great Plains (DeKeyser et al. 2013) and its presence is often 
10

correlated with declines in native species richness (Miles and Knops 2009). The Fish and Wildlife 

Service has begun an adaptive management program to control Kentucky bluegrass in South Dakota 

and North Dakota refuges (Grant et al. 2009). Their work suggests that Kentucky bluegrass can often 

spread and become a dominant component in mixed-grass prairie that is rested for long periods (e.g. 

neither burned nor grazed) (Grant et al. 2009).

Figure 5. The average absolute cover of the 10 most common native (green) and exotic (red) plants 

recorded at Wind Cave National Park in 1998-2015. Bars represent means ± standard errors.

Another concern in WICA is the spread of invasive cheatgrass (Bromus tectorum) and Japanese 

brome (Bromus japonicus). Together, the two species accounted for an average of 5.2 ± 0.7% cover 

from 2011 to 2015. Cheatgrass and Japanese brome are Eurasian annual grasses that have been a part 

of the NGP landscape for more than a century, but their invasion in the region has accelerated since 

1950 (Schachner et al. 2008). Our data suggest that the cover of annual bromes has been increasing 
11

over time at WICA (Figure 6). This is a concern because the presence of bromes is associated with a 

decrease in native species richness in the Northern Great Plains (F1,551=36.5, P<0.0001; Figure 8). An 

adaptive management program for annual bromes in the NGPN, starting in 2017, should help guide 

management actions in the future so that abundances of annual bromes remain low in WICA.

Figure 6. Trends in the relative cover of exotic plants and annual bromes in Wind Cave National Park

from 1998-2015. Points represent mean ± one standard error and sample size is to the right of the point. 

Years with fewer than three monitoring plots were excluded from the graph. The shaded area highlights 

the period from 2011-2015 when sampling methods were consistent and distribution of plots was more 

even and consistent across years. The dashed line represents the maximum and minimum cover values 

for each year. 
12

Figure 7. A map of long-term monitoring plots and the relative cover of exotic plants. Red dots indicate 

plots with a high cover of exotic plants (>15%), yellow has a moderate cover, and green dots indicate 

plots with low exotic species cover (<9%). 
13

Figure 8. The relationship between native species richness and the relative cover of annual bromes (in 

long-term monitoring plots in National Park units of the Northern Great Plains, 1998-2015.

Species Richness, Diversity, and Evenness 

One of the ways for the NPS to measure effectiveness at the mission to ‘preserve ecological 

integrity’ is to examine trends in native plant diversity and evenness within their boundaries. Average 

species richness has been measured by point-intercept since 1998 and in 1 m2 and 10 m2 quadrats

since 2011 (Table 4). 

Table 4. Average plant species richness in monitoring plots at Wind Cave National Park from 1998 to 

2015. Values represent means ± one standard error. 

Category

Point-intercept

(1998-2011; n=98)

1 m2 quadrats

(2011-2015, n=69)

10 m2 quadrats

(2011-2015, n=69)

Species richness 21.6 ± 0.6 12.7 ± 0.4 23.6 ± 0.6 

Native species richness 18.5 ± 0.5 11.1 ± 0.4 20.4 ± 0.6

Exotic species richness 3.3 ± 0.2 1.7 ± 0.1 3.2 ± 0.2

Graminoid species richness 11.3 ± 0.3 5.4 ± 0.2 8.0 ± 0.2

Forb species richness 8.3 ± 0.4 6.5 ± 0.3 13.7 ± 0.5

While there was some variation across the park, the plots we visited in WICA tended to have a 

moderate to high number of native plants compared to other mixed-grass prairies. Species richness in 

the mixed-grass prairie is determined by numerous factors including fire regime, grazing, prairie dog 

disturbance, and weather fluctuations (Symstad and Jonas 2011). While it is difficult to define a 

reference condition for species richness, which naturally varies considerably across both space and 
14

time, the natural range of variation over long-time periods may be a good starting point (Symstad and 

Jonas 2014). Long-term records of species diversity in mixed-grass prairie in a moderately grazed 

site in Montana ranged between 8 and 18 species per square meter (10-90th percentile range) between 

1933-1945 (Symstad and Jonas 2014). Since there are no long-term records of mixed-grass prairie 

diversity within the Black Hills, we are relying on these data as a reference condition. Future work is 

needed to develop a robust reference condition for plant communities in the Black Hills. Compared 

to the nearby mixed-grass prairie, WICA is within the natural range (Table 4). The lowest diversity 

of native plants was found in closed canopy ponderosa pine forests, such as plot WICA_PCM_007 

where there were only an average of 5 native species per square meter (Figure 9). Grassland sites 

tended to be more diverse and WICA_PCM_0018 and WICA_PCM_0026 had an average of 16.9 

and 16.8 species per m2

, respectively (Figure 9). 

Figure 9. A photograph of long-term monitoring plot WICA_PCM_0007 and WICA_PCM_0018 which has 

a low and high diversity of native plant species, respectively. 

We did not find any trends in species richness or evenness (Figure 10). Native diversity in 1m2

quadrats was fairly similar from 2011-2015; it ranged from a low in 2012 of 9.4 ± 0.8 (a drought 

year) to a high of 12.5 ± 1.1 in 2014 (a wet year). We have a longer record of native richness from 

point-intercept data (Figure 10; top). From 1998-2015, we recorded an average between 12.6 and 

21.6 native species. Annual average evenness of native species ranged from 0.65 to 0.81 during this 

time period, indicating the plots were not strongly dominated by a single species (Figure 10: bottom).

There is a great deal of variation in species richness and evenness among sites within WICA (dashed 

lines in Figure 10 represent the maximum and minimum values) which makes long-term trends in 

these metrics difficult to detect.

Disturbance from grazing, fire, and humans affects plant community structure and composition in 

mixed-grass prairie. We estimated the approximate area affected by natural and human disturbances 

at each site we visited in 2011-2015 by surveying the area for ~ 5 minutes at the end of the plot visit. 

The most common disturbance was from fires and evidence of recent fire was found at 56 plot visits. 

There was also evidence of animal use including trails, wallows, and grazing. We found no 

correlation with total disturbance, small or large animal disturbance and native richness or exotic 

cover. As more monitoring data are collected in future years, we may be able to better explore the 

statistical relationship between these metrics and disturbance. 
15

Figure 10. Trends in native species richness and evenness in Wind Cave National Park,1998-2015. Data 

are means ± one standard error. The dashed line indicates the maximum and minimum values for each 

year. 
16

Comparing Forest and Grassland Plots

Wind Cave NP is located at the transition zone between the mixed-grass prairie of the Northern Great 

Plains and ponderosa pine forests of the Black Hills. Approximately 30% of the land area of WICA

is dominated by ponderosa pine forest (Cogan et al. 1999). Fire is an important component of 

ponderosa pine and mixed-grass prairie ecosystems, and promotes healthy function of both systems 

while limiting encroachment of ponderosa pine into prairie habitat (Brown and Sieg 1999). Recent 

work has shown that changing climate may alter the complex interactions between ponderosa pine 

forests, climate, and fire, causing shifts in the extent of ponderosa pine forests in the Black Hills 

(King et al. 2013; King et al. 2015).

Randomly located NPGN vegetation monitoring plots in WICA are comprised of 71% grassland 

plots and 29% forested plots, which agrees with previous estimates of plant community composition 

in the park. Plots were considered forested if the mean diameter at breast height (DBH) is greater 

than 15 m2

/ha at an individual plot. We compared forested plots to grassland plots to determine 

whether exotic species abundance and other plot attributes differed between these ecotypes (Figure 

11).

Originally, we suspected that forested plots harbored a greater abundance of exotic species when 

compared with adjacent grasslands (Figures 12 & 13), but exotic species relative cover was not 

significantly different between forest and grassland plots (Figure 11A; F1,54=2.8, P=0.10). Exotic 

species cover did change over time in both plot types (F1,54=15.8, P<0.01).

Native species richness was greater in grassland plots (Figure 11B; F1,54=7.9, P<0.01), and changed 

over time in both plot types (F1,54=6.0, P=0.02). Relative cover of annual brome grasses (B. tectorum 

and B. japonicus) was greater in grassland plots (Figure 11C; F1,54=7.9, P<0.01) and varied over time 

in both plot types (F1,54=5.0, P=0.02); and litter cover was greater in forest plots in 2011, but by 2015 

litter cover was the same in both plot types after a 16% increase in grassland plots (Figure 11D;

F1,54=9.2, P<0.01). 
17

Figure 11. Comparison of forest and grassland plots in Wind Cave National Park. Values are means ± 

one standard error.
18

Figure 12. Examples of dense forest plots (left), moderately dense forest plots (center) and grassland 

plots (right).

Figure 13. Map comparing exotic species relative cover in forest and grassland plots in Wind Cave 

National Park. Plot marker size is proportional to the cover of exotic species observed in each plot, as 

described in the legend.
19

The Influence of Climate and Fire on Plant Community Structure and Diversity

Climate

The Northern Great Plains has a continental climate, with hot summers and very cold winters. The 

30-year normal temperatures at a nearby weather station, Wind Cave NP ranged from average 

minimum monthly temperatures in December of 13.9° F to maximum monthly July temperatures of 

85.1° F (based on 1980-2010). The 30-year normal annual precipitation totals 19.65 inches. Annual 

precipitation at WICA in 1998-2015 was variable and ranged between 9.6 and 28.8 inches, in 2008

and 1998, respectively. There were dry years in the early to mid-2000s and in 2012 (Figure 10). The 

last few years have been much wetter than average. The native vegetation is adapted to this variation, 

and productivity responds strongly to decreases in spring and summer precipitation (Yang et al. 

1998, Smart et al. 2007). Species richness and diversity in regional grasslands are also sensitive to 

temperature and precipitation fluctuation, but the response is complex and less predictable (Jonas et 

al. 2015). We found no statistically significant relationships between climate and vegetation, 

although there was a tendency for plant height to increase with precipitation and native species 

richness to decline with hotter maximum temperatures. Continued monitoring and a longer time 

series of vegetation data and climate will allow us to better understand how WICA vegetation 

responds to climate. 

Figure 14. The total annual precipitation anomaly from 1998-2015 for Wind Cave National Park. Positive 

values (blue) represent years wetter than and negative values (red) are drier than the 1981-2010 normal. 

The anomaly is measured in inches and based on data from a nearby weather station.
20

Fire History

Historically, smaller fires maintained a relatively stable composition of grassland and forest in the

Black Hills (Shepperd and Battaglia 2002, Brown and Cook 2006). During the period of 1500 to 

early 1900, the average number of years between wide-spread fires in the area was estimated at 10-12

years (Brown and Sieg 1999). Natural fires were suppressed for most of the last century, but the use 

of prescribed burning in Northern Great Plains parks to mitigate the effects of the absence of natural 

fires has increased over time since its start at Wind Cave NP in 1973 (Wienk et al. 2010). As of 2015, 

there is a mosaic of recently burned and unburned areas WICA (Figure 15).

Figure 15. Map of recent fire history at Wind Cave National Park.
21

Figure 16. Native species richness and percent cover of annual bromes in plots with different fire 

histories. 

The effects of specific prescribed burns on vegetation and more details about fires at WICA can be 

found in past NGPFire annual reports (see http://www.nps.gov/ngpfire/docs.htm). Here, we were 

interested in determining the relationship between fire history and vegetation. We compared two 

vegetation metrics: the native species richness and the relative cover of annual bromes with the 

length of time between the site visit and the last time a site had been burned (years since fire). For 

example, a site that burned in the spring and then was visited in the summer would be 0 years since 

fire. 

We found no relationship between native richness and time since a plot had burned (Figure 16;

F1,177=0.3, P=0.57). We also found no relationship between the time since a plot had burned and the 

relative cover of exotic plants (F1,177=2.1, P=0.15). There is a significant relationship between annual 

brome and fire (Figure 16; F1,177=19.0, P<0.0001). While there was variation in the data, the sites 

which have not burned in over 20 years have a higher cover of annual bromes than sites that had

burned more recently. This suggests that burning may assist in reducing annual brome cover. 

However, the increasing trend in annual brome abundance across the park (Figure 6) despite fairly 

frequent prescribed burns suggests that burning alone may not be sufficient. The best approach will 

likely include burning and targeted herbicides. Ongoing research on this topic and an upcoming 

adaptive management initiative for annual brome control in NGPN parks should provide more data 

and guidance to help with these management decisions. 
22

Rare Plants 

While locating rare species was not the focus of this study, two plant species of conservation concern 

were observed within vegetation monitoring plots at WICA. Nylon hedgehog cactus (Echinocereus 

viridiflorus S3) was observed in 10 plots, in very low abundances (Table 5). A moonwort species 

(Botrychium sp.) was observed in a single location. This moonwort individual could not be identified 

to the species level, but all moonwort species that occur in South Dakota are rare (an example of a 

moonwort is shown in Figure 12).

Table 5. Rare species occurrence in Wind Cave National Park from 1998-2015. Status ranks are based 

on South Dakota Natural Heritage Program designations. Plot count is the number of unique plots a 

species was recorded in across all years. Mean cover is the average cover of that species across all 

years.

Species Common Name State Rank Global 

Rank

Plot 

Count

Mean Cover 

(%)

Echinocerus viridiflorus nylon hedgehog cactus S3 G5 10 <0.01

Botrychium sp. moonwort species S1-S3 G1-G5 1 0.00

Figure 17. Examples of two rare species found in plant community monitoring plots at Wind Cave 

National Park. Left: Nylon hedgehog cactus (Echinocereus viridiflorus S3). Right: an example of a 

moonwort species (Botrychium lineare S1).

There have been some vegetation mapping and survey projects completed at Wind Cave and the 

surrounding Black Hills (Cogan et al. 1999, Marriott 2000), but we recommend a survey focused on 

locating rare plant populations at WICA be completed when funds are available. A full rare plant 

survey will be more likely to accurately quantify the status of rare plants at WICA and identify rare 

plant populations in areas of the park with no NGPN monitoring plots. 
23

Conclusion

The Northern Great Plains Inventory & Monitoring Program and Fire Effects Program have been 

monitoring vegetation in Wind Cave National Park for over 18 years. While methods have changed 

slightly, this report summarizes data from 97 locations from 1998-2015. Below, we list the questions 

we asked and provide a summarized answer, for more details see the Results and Discussion section.

We conclude with a Natural Resource Condition Table (Table 6) that summarizes the current status 

and trends in a few key vegetation metrics. 

1. What is the current status of plant community composition and structure of WICA

vegetation (species richness, cover, and diversity) and how has this changed from 1998-

2015? 

The ponderosa pine woodlands and mixed-grass prairie within the boundaries of WICA are

characterized by moderate to high native species richness. The closed canopy forest generally 

maintains lower species richness than the open grassland areas. We found no significant trends in 

native diversity or evenness from 1998 to 2015. Annual bromes are an emerging threat in WICA, and 

keeping them in low abundance will help maintain native plant diversity. Other exotics, such as 

Kentucky bluegrass are more prevalent, but there is no evidence of an increasing trend in their cover. 

2. What are the differences and similarities in forested and grassland condition and trends?

We examined the status and trends in native species richness, exotic species cover, and litter cover 

between forest and grasslands plots in WICA. We found that grassland plots had higher native 

richness and higher cover of annual brome grasses but there was not a difference in the relative cover 

of exotic plant species. Despite some differences in species composition, trends over time appear 

consistent in grasslands and forested plots. This suggests that both ecotypes are responding similarly 

to climate and other external drivers. 

3. How do trends in vegetation correlate with climate and fire history?

The large variability in the climate of WICA has made it difficult to discern strong patterns linking 

temperature, precipitation, and plant community structure (e.g. exotic cover, species richness). A 

longer time series of vegetation data will make it easier to elucidate trends in the future. WICA has 

been using prescribed burning as a management tool since the 1970s. There was no difference in 

native diversity among plots that had burned recently and those that had not burned for over 25 years. 

However, the sites which burned recently have a lower cover of annual brome grasses than sites that 

have not burned in over 20 years. Ongoing adaptive management programs and research will provide 

better guidance to the park on whether prescribed burns should be used to reduce the cover of annual 

bromes. 

4. What, if any, rare plants were found in WICA long-term monitoring plots? 

We identified 2 rare plant species in WICA between 1998 and 2015- nylon hedgehog cactus and a 

moonwort species (Botrychium sp.). These plants are found in such low abundance and in such few 
24

plots, it will take many years to determine any trends in rare plant cover. We recommend a more 

targeted survey of rare plants be completed when funds are available. 

Table 6. Natural resource condition summary table for plant communities in Wind Cave National Park.

Current values are based on data from 2011-2015 and trends are based on data from 1998-2015. 

Indicator of 

Condition

Specific 

Measures

Current

Value 

(mean ± 

SE)

Reference 

Condition 

and Data 

Source

Condition 

Status/Trend

Rationale for Resource 

Condition

Upland Plant 

Community 

Structure and 

Composition

Native species 

richness (1m2

quadrats) 

11.1 ± 0.4

species 8-18 species 

The ponderosa pine woodland

and mixed-grass prairie plant 

communities within the 

boundaries of WICA are 

characterized by moderate to 

high native species richness that 

is within the natural range of 

variability (Symstad and Jonas 

2014). Native diversity and plant 

cover is higher in grassland 

plots than forested plots. Native 

evenness is high indicating one 

species is not dominating the 

community and it not changed 

significantly since monitoring 

began in 1998.

Evenness

(pointintercept 

transects)

0.77 ± 

0.006

To be 

determined

Exotic Plant 

Early 

Detection and 

Management

Relative cover 

of exotic 

species 

20.9 ± 

1.4% < 10 % cover

Many areas of WICA have a 

high cover of exotic species. 

Kentucky bluegrass is the most 

widespread and abundant 

invasive plant species. There is 

no evidence of an increasing or 

decreasing trend in total exotic 

cover or Kentucky bluegrass 

cover. While still in low 

abundance, annual brome cover 

has shown an increasing trend 

since 1998.

Annual brome 

cover 

3.3 ± 0.4% < 10 % cover
25

Literature Cited 

Ashton, I., M. Prowatzke, M. Bynum, T. Shepherd, S. K. Wilson, and K. Paintner-Green. 2012. 

Wind Cave National Park plant community composition and structure monitoring: 2011 annual 

report. Natural Resource Technical Report NPS/NGPN/NRTR—2012/534. National Park 

Service, Fort Collins, Colorado.

Ashton, I., M. Prowatzke, S. K. Wilson, P. Graeve, and D. Swanson. 2013. Forest structure and fuel 

loads at Wind Cave National Park: 2012 status report. Natural Resource Technical Report 

NPS/NGPN/NRTR—2013/776. National Park Service, Fort Collins, Colorado.

Brown, J. K. 1974. Handbook for inventorying downed material. General Technical Report INT-16. 

USDA Forest Service, Intermountain Forest and Range Experiment Station, Ogden, UT.

Brown, J. K., R. D. Oberhue, and C. M. Johnston. 1982. Inventorying surface fuels and biomass in 

the Interior West. General Technical Report INT-129. USDA Forest Service, Intermountain 

Forest and Range Experiment Station, Ogden, UT.

Brown, P. and C. Sieg. 1999. Historical variability in fire at the ponderosa pine - Northern Great 

Plains prairie ecotone, southeastern Black Hills, South Dakota. Ecoscience 6:539-547.

Brown, P. M. and B. Cook. 2006. Early settlement forest structure in Black Hills ponderosa pine 

forests. Forest Ecology and Management 223:284-290.

Burkhart, B. A. and K. L. Kovacs. 2013. Condition assessment of streambanks and streamside 

vegetation on perennial streams in Wind Cave National Park: 2009-2012. Natural Resource 

Technical Report NPS/NGPN/NRTR—2013/758. National Park Service, Fort Collins, Colorado.

Cogan, D., H. Marriott, J. Von Loh, and M. J. Pucherelli. 1999. USGS-NPS Vegetation Mapping 

Program Wind Cave National Park, South Dakota. USDI Bureau of Reclamation. Denver, CO.

DeKeyser, E. S., M. Meehan, G. Clambey, and K. Krabbenhoft. 2013. Cool season invasive grasses 

in Northern Great Plains natural areas. Natural Areas Journal 33:81-90.

Grant, T. A., B. Flanders-Wanner, T. L. Shaffer, R. K. Murphy, and G. A. Knutsen. 2009. An 

emerging crisis across northern prairie refuges: prevalence of invasive plants and a plan for 

adaptive management. Ecological Restoration 27:58-65.

Jonas, J. L., D. A. Buhl, and A. J. Symstad. 2015. Impacts of weather on long-term patterns of plant 

richness and diversity vary with location and management. Ecology 96:2417-2432.

Komp, M. R., K. J. Stark, A. J. Nadeau, S. Amberg, E. Iverson, L. Danzinger, L. Danielson, and B. 

Drazkowski. 2011. Wind Cave National Park: Natural resource condition assessment. Natural 

Resource Report NPS/WICA/NRR—2011/478. National Park Service, Fort Collins, Colorado.
26

Larson, G. E. and J. R. Johnson. 2007. Plants of the Black Hills and Bear Lodge Mountains, 2nd 

edition. South Dakota State University, Brookings, SD.

Marriott, H. 2000. The Black Hills community inventory. Nature Conservancy.

Miles, E. K. and J. M. Knops. 2009. Shifting dominance from native C4 to non‐native C3 grasses: 

relationships to community diversity. Oikos 118:1844-1853.

NOAA. 2015. Climate Data Online at http://www.ncdc.noaa.gov/cdo-web/. National Oceanic and 

Atmospheric Administration. National Centers for Environmental Information.

NPS. 2003. Fire monitoring handbook. National Park Service. Fire Management Program Center, 

National Interagency Fire Center, Boise, ID.

Schachner, L. J., R. N. Mack, and S. J. Novak. 2008. Bromus tectorum (Poaceae) in midcontinental 

United States: population genetic analysis of an ongoing invasion. American journal of botany 

95:1584-1595.

Shepperd, W. D. and M. A. Battaglia. 2002. Ecology, silviculture, and management of Black Hills 

ponderosa pine.in USDA, editor. Rocky Mountain Research Station General Technical Report 

RMS-GTR-97.

Smart, A. J., B. H. Dunn, P. S. Johnson, L. Xu, and R. N. Gates. 2007. Using weather data to explain 

herbage yield on three Great Plains plant communities. Rangeland Ecology & Management 

60:146-153.

Stevens, D. L. and A. R. Olsen. 2003. Variance estimation for spatially balanced samples of 

environmental resources. Environmetrics 14:593-610.

Stevens, D. L. and A. R. Olsen. 2004. Spatially balanced sampling of natural resources. Journal Of 

The American Statistical Association 99:262-278.

Symstad, A. J. and J. L. Jonas. 2011. Incorporating biodiversity into rangeland health: plant species 

richness and diversity in Great Plains grasslands. Rangeland Ecology & Management 64:555-

572.

Symstad, A. J. and J. L. Jonas. 2014. Using natural range of variation to set decision thresholds: a 

case study for Great Plains grasslands. Pages 131-156 in G. R. Gutenspergen, editor. Application 

of threshold concepts in natural resource decision making. Springer, New York.

Symstad, A. J., R.A. Gitzen, C. L. Wienk, M. R. Bynum, D. J. Swanson, A. D. Thorstenson, and K. 

J. Paintner. 2012a. Plant community composition and structure monitoring protocol for the 

Northern Great Plains I&M Network-Standard Operating Procedures: version 1.01. Natural 

Resource Report NPS/NGPN/ NRR-2012/489.1.

Symstad, A. J., R.A. Gitzen, C. L. Wienk, M. R. Bynum, D. J. Swanson, A. D. Thorstenson, and K. 

J. Paintner. 2012b. Plant community composition and structure monitoring protocol for the 
27

Northern Great Plains I&M Network: version 1.01. Natural Resource Report NPS/NGPN/ NRR2012/489.

USDA-NRCS. 2015. The PLANTS Database (http://plants.usda.gov, 28 October 2015). National 

Plant Data Team, Greensboro, NC 27401-4901 USA.

Wienk, C., A. Thorstenson, J. Freeman, and D. Swanson. 2010. Northern Great Plains Fire Ecology 

Program review: 1997-2007. Natural Resource Report NPS/NRDS/NRDS—2010/112. National 

Park Service, Fort Collins, Colorado.

Yang, L., B. K. Wylie, L. L. Tieszen, and B. C. Reed. 1998. An analysis of relationships among 

climate forcing and time-integrated NDVI of grasslands over the US northern and central Great 

Plains. Remote Sensing of Environment 65:25-37.
Appendix A: List of plant species found at WICA 1998-2015

Below is a list of all the plant species found in WICA long-term plant community monitoring plots. 

The species are grouped by plant family. An “X” in the exotic column means that species is not 

native to the park or in the case where only the genera was identified, there are some species within 

that genera that are exotic. Species considered to be rare in South Dakota are marked in the final 

column and the state ranks are provided. The definitions can be found in Table 4 of the report. 

Family Symbol Scientific Name Common Name Exotic Rare

Aceraceae ACNE2 Acer negundo boxelder

Agavaceae YUGL Yucca glauca soapweed yucca

Amaranthaceae

AMBL Amaranthus blitoides mat amaranth X

AMRE Amaranthus retroflexus redroot amaranth

DYAM99 Dysphania ambrosioides Mexican tea X

Anacardiaceae

RHAR4 Rhus aromatica fragrant sumac

RHTR Rhus trilobata skunkbush sumac

TORY Toxicodendron rydbergii western poison ivy

Apiaceae

LOFO Lomatium foeniculaceum desert biscuitroot

MUSIN Musineon wildparsley

MUTE3 Musineon tenuifolium slender wildparsley

Apocynaceae

APAN2 Apocynum androsaemifolium spreading dogbane

APCA Apocynum cannabinum common dogbane

Asclepiadaceae

ASCLE Asclepias milkweed

ASOV Asclepias ovalifolia oval-leaf milkweed

ASPU Asclepias pumila plains milkweed

ASSP Asclepias speciosa showy milkweed

ASST Asclepias stenophylla slimleaf milkweed

ASVE Asclepias verticillata whorled milkweed

ASVI Asclepias viridiflora green comet milkweed

Asteraceae

ACMI2 Achillea millefolium common yarrow

AGGL Agoseris glauca pale agoseris

AMAR2 Ambrosia artemisiifolia annual ragweed

AMPS Ambrosia psilostachya Cuman ragweed

ANMI3 Antennaria microphylla littleleaf pussytoes

ANNE Antennaria neglecta field pussytoes

ANPA4 Antennaria parvifolia small-leaf pussytoes

ANRO2 Antennaria rosea rosy pussytoes

ANTEN Antennaria pussytoes X

ARCA12 Artemisia campestris field sagewort

ARCA13 Artemisia cana silver sagebrush

ARDR4 Artemisia dracunculus tarragon

ARFI2 Artemisia filifolia sand sagebrush

ARFR4 Artemisia frigida fringed sagewort

ARLU Artemisia ludoviciana white sagebrush

ARTEM Artemisia sagebrush X

ASTER Aster aster X

BREU Brickellia eupatorioides false boneset

CEST8 Centaurea stoebe spotted knapweed X
Family Symbol Scientific Name Common Name Exotic Rare

CIAR4 Cirsium arvense Canada thistle X

CIDR2 Cirsium drummondii dwarf thistle

CIFL Cirsium flodmanii Flodman's thistle

CIRSI Cirsium thistle X

CIUN Cirsium undulatum wavyleaf thistle

CIVU Cirsium vulgare bull thistle X

COCA5 Conyza canadensis horseweed

CORA4 Conyza ramosissima dwarf horseweed

CYXA Cyclachaena xanthifolia giant sumpweed

DYPA Dyssodia papposa fetid marigold

ECAN2 Echinacea angustifolia blacksamson echinacea

ERCA4 Erigeron canus hoary fleabane

ERFL Erigeron flagellaris trailing fleabane

ERFO3 Erigeron formosissimus beautiful fleabane

ERGL2 Erigeron glabellus streamside fleabane

ERIGE2 Erigeron fleabane

EROC Erigeron ochroleucus buff fleabane

ERPU2 Erigeron pumilus shaggy fleabane

ERST3 Erigeron strigosus prairie fleabane

ERSU2 Erigeron subtrinervis threenerve fleabane

EUSI13 Eurybia sibirica arctic aster

GNPA Gnaphalium palustre western marsh cudweed

GRSQ Grindelia squarrosa curlycup gumweed

GUSA2 Gutierrezia sarothrae broom snakeweed

HEAN3 Helianthus annuus common sunflower

HELIA3 Helianthus sunflower

HEMA2 Helianthus maximiliani Maximilian sunflower

HEPA19 Helianthus pauciflorus stiff sunflower

HEVI4 Heterotheca villosa hairy false goldenaster

HIAL2 Hieracium albiflorum white hawkweed

HIERA Hieracium hawkweed X

HYFI Hymenopappus filifolius fineleaf hymenopappus

LACTU Lactuca lettuce X

LASE Lactuca serriola prickly lettuce X

LIPU Liatris punctata dotted blazing star

LOAR5 Logfia arvensis field cottonrose X

LYJU Lygodesmia juncea rush skeletonplant

MUOB99 Mulgedium oblongifolium blue lettuce

OLRI Oligoneuron rigidum stiff goldenrod

PACA15 Packera cana woolly groundsel

PAPL12 Packera plattensis prairie groundsel

PSMA11 Pseudognaphalium macounii Macoun's cudweed

RACO3 Ratibida columnifera upright prairie coneflower

SECR Senecio crassulus thickleaf ragwort

SENEC Senecio ragwort X

SOCA6 Solidago canadensis Canada goldenrod

SOLID Solidago goldenrod
Family Symbol Scientific Name Common Name Exotic Rare

SOMI2 Solidago missouriensis Missouri goldenrod

SOMO Solidago mollis velvety goldenrod

SONE Solidago nemoralis gray goldenrod

SOPT4 Solidago ptarmicoides prairie goldenrod

SORI2 Solidago rigida stiff goldenrod

SOSP2 Solidago speciosa showy goldenrod

SYER Symphyotrichum ericoides white heath aster

SYFA Symphyotrichum falcatum white prairie aster

SYLA3 Symphyotrichum laeve smooth blue aster

SYLA6 Symphyotrichum lanceolatum white panicle aster

SYMPH4 Symphyotrichum aster

SYNO2 Symphyotrichum novaeangliae New England aster

SYOB Symphyotrichum 

oblongifolium aromatic aster

TAOF Taraxacum officinale common dandelion X

TEAC Tetraneuris acaulis stemless four-nerve daisy

TOEX2 Townsendia exscapa stemless Townsend daisy

TRDU Tragopogon dubius yellow salsify X

XASP99 Xanthisma spinulosum lacy tansyaster

Betulaceae BEPA Betula papyrifera paper birch

Boraginaceae

CRMI5 Cryptantha minima little cryptantha

CRTH Cryptantha thyrsiflora calcareous cryptantha

CYOF Cynoglossum officinale houndstongue X

HADE Hackelia deflexa nodding stickseed

LACE Lappula cenchrusoides Great Plains stickseed

LAOC3 Lappula occidentalis flatspine stickseed

LASQ Lappula squarrosa European stickseed X

LIIN2 Lithospermum incisum narrowleaf stoneseed

MELA3 Mertensia lanceolata prairie bluebells

MEOB Mertensia oblongifolia oblongleaf bluebells

ONBE Onosmodium bejariense soft-hair marbleseed

Brassicaceae

ALDE Alyssum desertorum desert madwort X

ARABI2 Arabis rockcress

ARHI Arabis hirsuta hairy rockcress

ARPY4 Arabis pycnocarpa creamflower rockcress

BOFE Boechera fendleri Fendler's rockcress

BOHO99 Boechera holboellii Holboell's rockcress

BOPI99 Boechera pinetorum Holboell's rockcress

BRASS2 Brassica mustard X

CABU2 Capsella bursa-pastoris shepherd's purse X

CAMI2 Camelina microcarpa littlepod false flax X

DEPI Descurainia pinnata western tansymustard

DESO2 Descurainia sophia herb sophia X

DRABA Draba draba

DRAU Draba aurea golden draba

DRRE2 Draba reptans Carolina draba

ERAS2 Erysimum asperum western wallflower
Family Symbol Scientific Name Common Name Exotic Rare

ERCA14 Erysimum capitatum sanddune wallflower

ERCH9 Erysimum cheiranthoides wormseed wallflower X

ERIN7 Erysimum inconspicuum shy wallflower

ERYSI Erysimum wallflower X

HEMA3 Hesperis matronalis dames rocket X

LEDE Lepidium densiflorum common pepperweed

LEMO3 Lesquerella montana mountain bladderpod

PHLU99 Physaria ludoviciana foothill bladderpod

PHMO99 Physaria montana mountain bladderpod

SIAL2 Sisymbrium altissimum tall tumblemustard X

THAR5 Thlaspi arvense field pennycress X

TUGL Turritis glabra tower rockcress

Cactaceae

ECVI2 Echinocereus viridiflorus nylon hedgehog cactus S3

ESCOB Escobaria foxtail cactus, beehive cactus

ESMI3 Escobaria missouriensis Missouri foxtail cactus

ESVI2 Escobaria vivipara spinystar

OPFR Opuntia fragilis brittle pricklypear

OPMA2 Opuntia macrorhiza twistspine pricklypear

OPPO Opuntia polyacantha plains pricklypear

OPUNT Opuntia pricklypear

Campanulaceae

CARO2 Campanula rotundifolia bluebell bellflower

TRIOD Triodanis Venus' looking-glass

TRLE3 Triodanis leptocarpa slimpod Venus' looking-glass

TRPE4 Triodanis perfoliata clasping Venus' looking-glass

Caprifoliaceae SYAL Symphoricarpos albus common snowberry

SYOC Symphoricarpos occidentalis western snowberry

Caryophyllaceae

ARENA Arenaria sandwort X

CEAR4 Cerastium arvense field chickweed

CEFO2 Cerastium fontanum common mouse-ear 

chickweed X

PADE4 Paronychia depressa spreading nailwort

PASE Paronychia sessiliflora creeping nailwort

SIAN2 Silene antirrhina sleepy silene

SIDR Silene drummondii Drummond's campion

SILA21 Silene latifolia bladder campion X

SILEN Silene catchfly

SINO Silene noctiflora nightflowering silene X

SIVU Silene vulgaris maidenstears X

Celastraceae CESC Celastrus scandens American bittersweet

Chenopodiaceae

CHAL7 Chenopodium album lambsquarters X

CHBE4 Chenopodium berlandieri pitseed goosefoot

CHDE Chenopodium desiccatum aridland goosefoot

CHENO Chenopodium goosefoot X

CHLE4 Chenopodium leptophyllum narrowleaf goosefoot

CHPR5 Chenopodium pratericola desert goosefoot

CHSI2 Chenopodium simplex mapleleaf goosefoot

MONU Monolepis nuttalliana Nuttall's povertyweed
Family Symbol Scientific Name Common Name Exotic Rare

SATR12 Salsola tragus prickly Russian thistle X

Cistaceae HEBI2 Helianthemum bicknellii hoary frostweed

Commelinaceae TRBR Tradescantia bracteata longbract spiderwort

TROC Tradescantia occidentalis prairie spiderwort

Convolvulaceae

COAR4 Convolvulus arvensis field bindweed X

EVNU Evolvulus nuttallianus shaggy dwarf morning-glory

IPLE Ipomoea leptophylla bush morning-glory

Crassulaceae SELA Sedum lanceolatum spearleaf stonecrop

Cupressaceae JUCO6 Juniperus communis common juniper

JUSC2 Juniperus scopulorum Rocky Mountain juniper

Cyperaceae

CABA3 Carex backii Back's sedge

CABR10 Carex brevior shortbeak sedge

CADU6 Carex duriuscula needleleaf sedge

CAFI Carex filifolia threadleaf sedge

CAIN11 Carex interior inland sedge

CAIN9 Carex inops sun sedge

CANE2 Carex nebrascensis Nebraska sedge

CAPE42 Carex pellita woolly sedge

CAREX Carex sedge

CARI Carex richardsonii Richardson's sedge

CARO5 Carex rossii Ross' sedge

CAST5 Carex stipata awlfruit sedge

ELPA3 Eleocharis palustris common spikerush

Dryopteridaceae

CYFR2 Cystopteris fragilis brittle bladderfern

DRYOP Dryopteris woodfern

GYDR Gymnocarpium dryopteris western oakfern

WOOR Woodsia oregana Oregon cliff fern

WOSC Woodsia scopulina Rocky Mountain cliff fern

Ericaceae ARUV Arctostaphylos uva-ursi kinnikinnick

Euphorbiaceae

EUBR Euphorbia brachycera horned spurge

EUDE4 Euphorbia dentata toothed spurge

EUGL3 Euphorbia glyptosperma ribseed sandmat

EUMA8 Euphorbia marginata snow on the mountain

EUPHO Euphorbia spurge, sandmat X

EUPR3 Euphorbia prostrata prostrate sandmat

EUSE5 Euphorbia serpyllifolia thymeleaf sandmat

EUMA7 Euphorbia maculata spotted sandmat

EUSP Euphorbia spathulata warty spurge

Fabaceae

AMCA6 Amorpha canescens leadplant

ASAG2 Astragalus agrestis purple milkvetch

ASAU4 Astragalus australis Indian milkvetch

ASBI2 Astragalus bisulcatus twogrooved milkvetch

ASCR2 Astragalus crassicarpus groundplum milkvetch

ASDR3 Astragalus drummondii Drummond's milkvetch

ASFL2 Astragalus flexuosus flexile milkvetch

ASGI5 Astragalus gilviflorus plains milkvetch

ASGR3 Astragalus gracilis slender milkvetch
Family Symbol Scientific Name Common Name Exotic Rare

ASLA27 Astragalus laxmannii Laxmann's milkvetch

ASLO4 Astragalus lotiflorus lotus milkvetch

ASMI10 Astragalus missouriensis Missouri milkvetch

ASMU99 Astragalus multiflorus looseflower milkvetch

ASPL2 Astragalus plattensis Platte River milkvetch

ASRA2 Astragalus racemosus cream milkvetch

ASSP6 Astragalus spatulatus tufted milkvetch

ASTRA Astragalus milkvetch

DACA7 Dalea candida white prairie clover

DAEN Dalea enneandra nineanther prairie clover

DALEA Dalea prairie clover

DAPU5 Dalea purpurea purple prairie clover

GLLE3 Glycyrrhiza lepidota American licorice

LAOC2 Lathyrus ochroleucus cream pea

LAPO2 Lathyrus polymorphus manystem pea

MELIL Melilotus sweetclover X

MELU Medicago lupulina black medick X

MEOF Melilotus officinalis yellow sweetclover X

MESA Medicago sativa alfalfa X

OXCA4 Oxytropis campestris field locoweed

OXLA3 Oxytropis lambertii purple locoweed

OXSE Oxytropis sericea white locoweed

OXYTR Oxytropis locoweed

PEAR6 Pediomelum argophyllum silverleaf Indian breadroot

PECU3 Pediomelum cuspidatum largebract Indian breadroot

PEES Pediomelum esculentum large Indian breadroot

PSCU Psoralea cuspidata largebract Indian breadroot

PSLA3 Psoralidium lanceolatum lemon scurfpea

PSTE5 Psoralidium tenuiflorum slimflower scurfpea

THRH Thermopsis rhombifolia golden pea

VIAM Vicia americana American vetch

QUMA2 Quercus macrocarpa bur oak

Fumariaceae COMI2 Corydalis micrantha smallflower fumewort

Geraniaceae GERAN Geranium geranium X

Grossulariaceae

RIAU Ribes aureum golden currant

RICE Ribes cereum wax currant

RIHI Ribes hirtellum hairystem gooseberry

RIOX Ribes oxyacanthoides Canadian gooseberry

Hydrophyllaceae ELNY Ellisia nyctelea Aunt Lucy

Iridaceae SIAN3 Sisyrinchium angustifolium narrowleaf blue-eyed grass

SIMO2 Sisyrinchium montanum strict blue-eyed grass

Juncaceae JUIN2 Juncus interior inland rush

JUNCU Juncus rush X

Lamiaceae

HEDR Hedeoma drummondii Drummond's false pennyroyal

HEHI Hedeoma hispida rough false pennyroyal

MAVU Marrubium vulgare horehound X

MOFI Monarda fistulosa wild bergamot
Family Symbol Scientific Name Common Name Exotic Rare

NECA2 Nepeta cataria catnip X

PRVU Prunella vulgaris common selfheal

SAAZ Salvia azurea azure blue sage

SARE3 Salvia reflexa lanceleaf sage

Liliaceae

ALCE2 Allium cernuum nodding onion

ALLIU Allium onion X

ALTE Allium textile textile onion

CAGU Calochortus gunnisonii Gunnison's mariposa lily

CANU3 Calochortus nuttallii sego lily

LEMO4 Leucocrinum montanum common starlily

MAST4 Maianthemum stellatum starry false lily of the valley

Linaceae LILE3 Linum lewisii Lewis flax

Linaceae LIRI Linum rigidum stiffstem flax

Malvaceae

SPCO Sphaeralcea coccinea scarlet globemallow

SPCOC Sphaeralcea coccinea ssp. 

coccinea scarlet globemallow

Melanthiaceae TOVE2 Toxicoscordion venenosum meadow deathcamas

Montiaceae PHPA29 Phemeranthus parviflorus sunbright

Nyctaginaceae

ABFR2 Abronia fragrans snowball sand verbena

MIHI Mirabilis hirsuta hairy four o'clock

MILI3 Mirabilis linearis narrowleaf four o'clock

Oleaceae FRPE Fraxinus pennsylvanica green ash

Onagraceae

EPBR3 Epilobium brachycarpum tall annual willowherb

OEBI Oenothera biennis common evening primrose

OECE2 Oenothera cespitosa tufted evening primrose

OECO2 Oenothera coronopifolia crownleaf evening primrose

OECU99 Oenothera curtiflora velvetweed

OEEN Oenothera engelmannii Engelmann's evening primrose

OELA Oenothera laciniata cutleaf evening-primrose

OENOT Oenothera evening primrose

OENU Oenothera nuttallii Nuttall's evening primrose

OESE3 Oenothera serrulata yellow sundrops

OESU99 Oenothera suffrutescens scarlet beeblossom

OEVI Oenothera villosa hairy evening primrose

Ophioglossaceae BOTRY Botrychium grapefern S1-S3

Orobanchaceae ORFA Orobanche fasciculata clustered broomrape

Oxalidaceae OXST Oxalis stricta common yellow woodsorrel

Papaveraceae ARPO2 Argemone polyanthemos crested pricklypoppy

Pinaceae PIPO Pinus ponderosa ponderosa pine

Plantaginaceae

PLPA2 Plantago patagonica woolly plantain

SYWY99 Synthyris wyomingensis Wyoming kittentails

Poaceae

ACHY Achnatherum hymenoides Indian ricegrass

ACRO7 Achnatherum robustum sleepygrass

AGROP2 Agropyron wheatgrass X

AGSC5 Agrostis scabra rough bentgrass

AGST2 Agrostis stolonifera creeping bentgrass X

ANGE Andropogon gerardii big bluestem
Family Symbol Scientific Name Common Name Exotic Rare

ARLO16 Aristida longespica slimspike threeawn

AROL Aristida oligantha prairie threeawn

ARPU9 Aristida purpurea purple threeawn

BOCU Bouteloua curtipendula sideoats grama

BODA2 Bouteloua dactyloides buffalograss

BOGR2 Bouteloua gracilis blue grama

BOHI2 Bouteloua hirsuta hairy grama

BRAN Bromus anomalus nodding brome

BRIN2 Bromus inermis smooth brome X

BRJA Bromus japonicus Japanese brome X

BROMU Bromus brome X

BRPO2 Bromus porteri Porter brome

BRTE Bromus tectorum cheatgrass X

BUDA Buchloe dactyloides buffalograss

CALO Calamovilfa longifolia prairie sandreed

DASP2 Danthonia spicata poverty oatgrass

DICL Dichanthelium clandestinum deertongue

DILI2 Dichanthelium linearifolium slimleaf panicgrass

DIOL Dichanthelium oligosanthes Heller's rosette grass

DIWI5 Dichanthelium wilcoxianum fall rosette grass

ECMU2 Echinochloa muricata rough barnyardgrass

ELCA4 Elymus canadensis Canada wildrye

ELEL5 Elymus elymoides squirreltail

ELRE4 Elymus repens quackgrass X

ELTR7 Elymus trachycaulus slender wheatgrass

ELVI3 Elymus virginicus Virginia wildrye

ELYMU Elymus wildrye X

FEID Festuca idahoensis Idaho fescue

FEOV Festuca ovina sheep fescue X

FESA Festuca saximontana Rocky Mountain fescue

HECO26 Hesperostipa comata needle and thread

HESP11 Hesperostipa spartea porcupinegrass

HOJU Hordeum jubatum foxtail barley

HOPU Hordeum pusillum little barley

KOMA Koeleria macrantha prairie Junegrass

MUAS Muhlenbergia asperifolia scratchgrass, alkali muhly

MUCU3 Muhlenbergia cuspidata plains muhly

MUPA99 Muhlenbergia paniculata tumblegrass

MURA Muhlenbergia racemosa marsh muhly

MUSQ3 Munroa squarrosa false buffalograss

NAVI4 Nassella viridula green needlegrass

PACA6 Panicum capillare witchgrass

PANIC Panicum panicgrass X

PASM Pascopyrum smithii western wheatgrass

PAVI2 Panicum virgatum switchgrass

PIMI7 Piptatherum micranthum littleseed ricegrass

POA Poa bluegrass X
Family Symbol Scientific Name Common Name Exotic Rare

POAL2 Poa alpina alpine bluegrass

POCO Poa compressa Canada bluegrass X

PONE Poa nemoralis wood bluegrass

POPA2 Poa palustris fowl bluegrass

POPR Poa pratensis Kentucky bluegrass X

SCPR4 Schedonorus pratensis meadow fescue X

SCPU Schizachne purpurascens false melic

SCSC Schizachyrium scoparium little bluestem

SONU2 Sorghastrum nutans Indiangrass

SPCO16 Sporobolus compositus composite dropseed

SPCR Sporobolus cryptandrus sand dropseed

SPHE Sporobolus heterolepis prairie dropseed

THIN6 Thinopyrum intermedium intermediate wheatgrass X

VUOC Vulpia octoflora sixweeks fescue

Polemoniaceae

COLI2 Collomia linearis tiny trumpet

PHAL3 Phlox alyssifolia alyssumleaf phlox

PHAN4 Phlox andicola prairie phlox

PHHO Phlox hoodii spiny phlox

Polygalaceae

POAL4 Polygala alba white milkwort

POLYG Polygala milkwort

POVE Polygala verticillata whorled milkwort

Polygonaceae

ERAN4 Eriogonum annuum annual buckwheat

ERFL4 Eriogonum flavum alpine golden buckwheat

ERPA9 Eriogonum pauciflorum fewflower buckwheat

FACO Fallopia convolvulus black bindweed X

POAV Polygonum aviculare prostrate knotweed X

PODO4 Polygonum douglasii Douglas' knotweed

PORA3 Polygonum ramosissimum bushy knotweed

Portulacaceae POOL Portulaca oleracea little hogweed X

Primulaceae

ANAR Anagallis arvensis scarlet pimpernel X

ANOC2 Androsace occidentalis western rockjasmine

ANSE4 Androsace septentrionalis pygmyflower rockjasmine

PRPA99 Primula pauciflora darkthroat shootingstar

Ranunculaceae

ANCY Anemone cylindrica candle anemone

ANEMO Anemone Anemone

ANMU Anemone multifida cutleaf anemone

ANPA19 Anemone patens eastern pasqueflower

MYMI2 Myosurus minimus tiny mousetail

Rosaceae

AGST Agrimonia striata roadside agrimony

AMAL2 Amelanchier alnifolia Saskatoon serviceberry

CEMO2 Cercocarpus montanus mountain mahogany

DRAR8 Drymocallis arguta tall cinquefoil

DRFI3 Drymocallis fissa bigflower cinquefoil

FRVI Fragaria virginiana Virginia strawberry

GETR Geum triflorum prairie smoke

PHMO4 Physocarpus monogynus mountain ninebark

POCO13 Potentilla concinna elegant cinquefoil
Family Symbol Scientific Name Common Name Exotic Rare

POHI6 Potentilla hippiana woolly cinquefoil

POPE8 Potentilla pensylvanica Pennsylvania cinquefoil

POTEN Potentilla cinquefoil X

PRAM Prunus americana American plum

PRPU3 Prunus pumila sandcherry

PRVI Prunus virginiana chokecherry

ROAC Rosa acicularis prickly rose

ROAR3 Rosa arkansana prairie rose

ROSA5 Rosa rose

ROWO Rosa woodsii Woods' rose

RUID Rubus idaeus American red raspberry X

RUSA99 Rubus sachalinensis common red raspberry

Rubiaceae GAAP2 Galium aparine stickywilly

GABO2 Galium boreale northern bedstraw

Salicaceae PODE3 Populus deltoides eastern cottonwood

SALIX Salix willow

Santalaceae COUM Comandra umbellata bastard toadflax

Saxifragaceae HERI Heuchera richardsonii Richardson's alumroot

Scrophulariaceae

CASE5 Castilleja sessiliflora Great Plains Indian paintbrush

CASU12 Castilleja sulphurea sulphur Indian paintbrush

COPA3 Collinsia parviflora maiden blue eyed Mary

NUTE Nuttallanthus texanus Texas toadflax

ORLU2 Orthocarpus luteus yellow owl's-clover

PEAL2 Penstemon albidus white penstemon

PEAN4 Penstemon angustifolius broadbeard beardtongue

PEGL3 Penstemon glaber sawsepal penstemon

PEGR5 Penstemon gracilis lilac penstemon

PEGR7 Penstemon grandiflorus large beardtongue

PENST Penstemon beardtongue

VEAR Veronica arvensis corn speedwell X

VETH Verbascum thapsus common mullein X

Selaginellaceae SEDE2 Selaginella densa lesser spikemoss

Smilacaceae SMLA3 Smilax lasioneura Blue Ridge carrionflower

Solanaceae

PHHE5 Physalis heterophylla clammy groundcherry

PHLO4 Physalis longifolia longleaf groundcherry

PHVI5 Physalis virginiana Virginia groundcherry

PHYSA Physalis groundcherry

SOPT3 Solanum ptychanthum West Indian nightshade

SORO Solanum rostratum buffalobur nightshade

SOTR Solanum triflorum cutleaf nightshade

Ulmaceae

CEOC Celtis occidentalis common hackberry

ULAM Ulmus americana American elm

ULMUS Ulmus elm X

ULPU Ulmus pumila Siberian elm X

Urticaceae PAPE5 Parietaria pensylvanica Pennsylvania pellitory

Urticaceae URDI Urtica dioica stinging nettle

Verbenaceae GLBI2 Glandularia bipinnatifida Dakota mock vervain
Family Symbol Scientific Name Common Name Exotic Rare

Verbenaceae VEBR Verbena bracteata bigbract verbena

Verbenaceae VEHA2 Verbena hastata swamp verbena

Verbenaceae VEST Verbena stricta hoary verbena

Violaceae VIAD Viola adunca hookedspur violet

Violaceae VINU2 Viola nuttallii Nuttall's violet, yellow prairie 

violet

Violaceae VIOLA Viola violet X

Violaceae VIPE2 Viola pedatifida prairie violet

Vitaceae PAVI5 Parthenocissus vitacea woodbine
The Department of the Interior protects and manages the nation’s natural resources and cultural heritage; provides scientific 

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affiliated Island Communities.

NPS 108/133334, June 2015
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