Calcareous Glade Monitoring at Chickamauga and Chattanooga National Military Park Final Report—Third Sampling of Baseline Transects Established in 1993

National Park Service U.S. Department of the Interior

Natural Resource Stewardship and Science Calcareous Glade Monitoring at Chickamauga and Chattanooga National Military Park Final Report—Third Sampling of Baseline Transects Established in 1993

Natural Resource Report NPS/CUPN/NRR—2021/2239

The production of this document cost $53,732, including costs associated with data collection, processing, analysis, and subsequent authoring, editing, and publication.

ON THE COVER Leavenworthia exigua var. exigua, an endemic special concern species characteristic of cedar glades at Chickamauga and Chattanooga National Military Park. Photo by Tom Govus.

Calcareous Glade Monitoring at Chickamauga and Chattanooga National Military Park Final Report—Third Sampling of Baseline Transects Established in 1993

Natural Resource Report NPS/CUPN/NRR—2021/2239

Thomas E. Govus1 and Bill J. Moore2

1Independent Botanist/Vegetation Ecologist 3711 Big Creek Road Elijay, Georgia 30536

2National Park Service Ecologist/Data Manager Cumberland Piedmont Network P.O. Box 8 Mammoth Cave, Kentucky 42259

March 2021

U.S. Department of the Interior National Park Service Natural Resource Stewardship and Science Fort Collins, Colorado

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 public.

The Natural Resource Report Series is used to disseminate comprehensive information and analysis about natural resources and related topics concerning lands managed by the National Park Service. The series supports the advancement of science, informed decision-making, and the achievement of the National Park Service mission. The series also provides a forum for presenting more lengthy results that may not be accepted by publications with page limitations.

All manuscripts in the series receive the appropriate level of peer review to ensure that the information is scientifically credible, technically accurate, appropriately written for the intended audience, and designed and published in a professional manner.

This report received peer review by qualified subject-matter experts with the appropriate background and expertise needed to evaluate the technical and scientific aspects of this study. Peer review was overseen by a peer review manager.

Views, statements, findings, conclusions, recommendations, and data in this report do not necessarily reflect views and policies of the National Park Service, U.S. Department of the Interior. Mention of trade names or commercial products does not constitute endorsement or recommendation for use by the U.S. Government.

This report is available from the Cumberland Piedmont Network website and the Natural Resource Publications Management website. If you have difficulty accessing information in this publication, particularly if using assistive technology, please email [email protected].

Please cite this publication as:

Govus, T. E., and B. J. Moore. 2021. Calcareous glade monitoring at Chickamauga and Chattanooga National Military Park: Final report—third sampling of baseline transects established in 1993. Natural Resource Report NPS/CUPN/NRR—2021/2239. National Park Service, Fort Collins, Colorado. https://doi.org/10.36967/nrr-2284881.

NPS 301/175135, March 2021 ii

Contents Page

Figures...... v

Tables ...... v

Appendices ...... vii

Executive Summary ...... ix

Acknowledgments ...... xi

Introduction ...... 1

Methods ...... 3

Glade Species Lists...... 3

Baseline Monitoring ...... 3

Woody Cover...... 4

Ground Cover ...... 5

Herb Density ...... 5

Follow up Taxonomic Issues Relating to the 1993 Data ...... 6

Photomonitoring procedures ...... 7

Results of the Third Round of Sampling ...... 9

Change in Large Woody Cover Greater Than or Equal to 1 meter (3.3 ft) ...... 10

Change in Plant Density Summary Results ...... 16

Discussions and Conclusions ...... 21

Woody Cover...... 21

Plant Density ...... 22

Exotic Species ...... 24

Management ...... 25

Future Sampling ...... 26

Literature Cited ...... 29

iii

Figures

Page

Figure 1. General location of glades sampled for woody cover; some were further divided by Sutter et al. (1994) into separate glades for monitoring purposes (i.e., glade 3A and 3B) ...... 10

Figure 2. Glade means for percent woody cover for eight of the highest cover species. Red line is mean across glades ...... 13

Figure 3. General location of glades sampled for plant density measurements...... 17

Figure 4. Sum total change in characteristic glade species densities over three sampling intervals at Chickamauga and Chattanooga National Military Park ...... 23

Figure 5. A baseline photograph of glade 24A showing degraded condition, particularly encroachment by Chinese privet (Ligustrum sinense)...... 26

Tables

Page

Table 1. Baseline length, segment (i.e., interval in which monitoring transects were randomly placed), number of monitoring transects, and combined monitoring transect length...... 4

Table 2. Percent change in total woody cover summed across species from 1993 to 2006- 2008 and 1993 to 2019 in monitored glades on Chickamauga and Chattanooga National Military Park...... 12

Table 3. Glade minimum and maximum percent woody cover for eight of the highest cover species by sampling session...... 13

Table 4. 2019 Percent cover values for select high cover species greater than or equal to 1 meter (3.3 ft) showing the glade mean value for each sample glade...... 14

Table 5. Difference in percent woody cover, comparing 1993 and 2019 values ...... 15

Table 6. Characteristic glade species 1993 density (individuals per square meter) and subsequent change in density based on 2019 survey results ...... 19

Table 7. Plant density values (plants/m²) of characteristic glade species averaged across sampled glades along with minimum and maximum glade averages (parentheses) ...... 20

Appendices

Page

Appendix A. Compiled Woody Cover Data for all Glades Sampled ...... 31

Appendix B. Compiled Herbaceous Plant Density Data for All Glades Sampled ...... 45

Appendix C. Compiled Small Woody Plant Density Data for All Glades Sampled (Woody Plants less than 0.5 meters) ...... 51

vii

Executive Summary

Calcareous glades, sometimes referred to as cedar glades, are characterized by limestone and dolomite outcrops having very thin rocky soils and with extreme environmental conditions. These rare habitats are unique, highly specialized communities that support many rare endemic or near endemic species in the southeastern United States. In 1993, The Nature Conservancy established several baseline and monitoring transects within a subset of glades on Chickamauga and Chattanooga National Military Park for the purpose of tracking changes in these biologically significant habitats through time. Woody cover data were resampled in nine glades during the 2006 field season and two glades during the 2008 field season. Plant density data were also resampled in five glades during the 2008 field season. All woody cover and plant density data collected during the combined 2006 and 2008 seasons were collected again most recently in 2019. The conclusion of the researchers following the 2006-2008 effort was that there was a decline in the natural integrity of the glade communities most notably reflected in an increase in woody cover and declining populations of several herbaceous species characteristic of glade communities. Results from the 2019 effort suggest a continued decline. Specifically, Eastern red cedar (Juniperus virginiana) has exhibited a consistent increase in cover through time across a majority of the monitored glades. Between 1993 and 2019 this species has exhibited statistically significant, double digit increases in percent cover in eight of the 11 monitored glades. Though not nearly as dramatic as the increases observed by red cedar an additional 11 woody species have also exhibited statistically significant increases in percent woody cover in one or more monitored glades. As a result, a fundamental feature of the glades (i.e., their openness) is being lost, making it more difficult to distinguish the glades from the forest/woodland complex surrounding them. Additionally, glade endemic or near endemic species have shown statistically significant declines from 1993 to 2019, while others went undetected in some glades in 2019. Nashville breadroot (Pediomelum subacaule) and Gattinger's prairie-clover (Dalea gattingeri) exhibited consistent (and statistically significant) declines in the majority of glades where they were monitored. In most cases 2019 observed densities were less than half the densities observed in 1993 for these species. Other glade endemic or near endemic species, glade larkspur (Delphinium carolinianum ssp calciphilum) and Tennessee gladecress (Leavenworthia exigua var. exigua) had extremely low densities or went undetected in 2019 surveys.

Exotics are also a concern. Chinese privet (Ligustrum sinense) increased substantially in glades 8A and 24A, and three glades (8A, 11, and 24A) now have total woody cover values at or near 10%. Chinese privet appears to be the only invasive species that poses a serious threat to the habitat integrity at the moment, but a new non-native species, waxy-leaf privet (Ligustrum quihoui) was detected on one transect in glade 6. To our knowledge, this occurrence represents a state record for Georgia and is of concern because it has been reported to be invasive in diabase glade communities in North Carolina. The removal of privet from the center and edges of these glades remains an important management priority.

We are also supportive of park’s plans to conduct a prescribed fire in the Viniard-Alexander Prescribed Burn Unit, which will impact glades 13, 14 and 24A, as well as other smaller glades and glade-like habitats in the vicinity. While it is hoped that fire will impact the encroachment of woody

vegetation it is questionable whether fire alone will be enough to restore the previously open nature to these glades. If this proves to be the case, removal of individual red cedars should also be considered. This management action has been implemented successfully on other national park units as part of their glade management and restoration activities.

Acknowledgments

We would like to thank the National Park Service for assistance with this effort. In particular, Teresa Leibfreid, Cumberland Piedmont Inventory and Monitoring Network ecologist deserves appreciation for helping with the many issues that came up during field work, data processing and statistical analysis. Likewise, Jim Szyjkowski, at Chickamauga and Chattanooga National Military Park helped to make sure assistance was available for field work. Kaylee Moore, Zack Irick and Trent Deason all helped with field work and their assistance was much appreciated. Ellen Cheng, Quantitative Ecologist, with the NPS Southeast Region Inventory and Monitoring, provided critical help with data analysis.

Finally, gratitude is extended to the researchers at The Nature Conservancy who established the initial baselines in 1993 and collected the original data that made all of this important work possible.

Introduction

Calcareous glades (also referred to as cedar glades) are a highly significant component of biodiversity and a rare plant association type for the Southeastern United States. The center of their distribution is the Central Basin and Highland Rim of Tennessee, but occurrences are also known from southern Kentucky, northern Alabama and northwestern Georgia. The flora of these distinctive, limestone and dolomite bedrock barrens are characterized by a high number of endemic and rare plant species compared to their relatively small size and extent. Few examples are protected, and many examples have been degraded or destroyed through development.

Sutter et al. (1994) identified 23 glades on the Chickamauga Battlefield unit in Walker County, Georgia. While Chickamauga and Chattanooga National Military Park (CHCH) is comprised of many units spanning three counties and two states, this is the only unit of the park with calcareous glades. In 1993, The Nature Conservancy established a system of monitoring, utilizing a more or less permanent baseline with perpendicular monitoring transects, on a subset of these glades for the purpose of tracking changes in these biologically significant habitats through time. Selection of the monitored glades was made by the investigators in 1993, as well as staff from the Georgia Natural Heritage Program and Chickamauga and Chattanooga NMP. These selections were subjectively based on “. . . size, presence of rare plant species, species diversity, and level of disturbance.” The methods and results of this effort are described by Sutter et al. (1994) in a report to the National Park Service. At the time of this initial report, it was recommended that resampling be conducted approximately every five years. The second round of sampling did not occur until 2006–2008, some 14 years later.

During the summer of 2005, National Park Service employees began an effort to relocate the established monitoring sites and create an ArcView GIS database of locations of marked starting points and end points for these baselines. This effort was documented by an internal report to the National Park Service by Rutledge and Diggs (2006). Of the total 12 baselines that were established by Sutter et al. (1994), 11 had the tags for the starting and ending points in good enough condition to be relocated and resampled accurately. During the summer of 2006 the National Park Service awarded a grant to NatureServe to resample, following the original monitoring protocol, analyze the collected data, and provide management suggestions based on the results. The results of that effort were reported by Govus and Lyons (2009) and Sutter et al. (2011).

A third round of sampling occurred from May–June of 2019, some 25 years after the initial study. Permanent monuments (metal conduit) were installed at the start, mid-points (where a directional change was made) and endpoints of the baselines. These are marked with engraved metal tags that identify the glade and tag tree number. It is hoped, these will aid future investigators in re-locating the baselines.

Methods

Readers of the original monitoring report (Sutter et al. 1994) and later reports might be confused as to how many glades were actually identified in 1993, and how many were selected for monitoring. Per Sutter et al. (1994) species lists were developed “for each of the twenty-three glades… baseline monitoring data was collected for nine selected glades… and herbaceous vegetation data was collected for a subset of six of the nine glades.” A permanent baseline transect (referred to simply as baseline in this report) was established in each of the nine glades along the primary longitudinal axis of the glade, off of which monitoring transects were established. However, upon further inspection by the initial investigators, three of the nine glades selected for monitoring were determined to be more or less “glade complexes”, necessitating multiple baselines to adequately sample. These additional baselines were annotated with a letter to differentiate them. Specifically, glade 3 became 3A and 3B, 8 became 8A and 8B and 24 became 24A and 24B. Also, Sutter et al. (1994) analyzed and reported these data as separate glades in effect increasing the number of monitored glades from nine to twelve. This referencing of twelve monitored glades was continued in subsequent reports (Sutter et al. 2011) and the authors of this report continue to report the number of monitored glades as twelve as well.

Unfortunately, during the 2005 effort of attempting to relocate the original 12 baselines, it was discovered glade number 24B had apparently lost a number of tag trees due to pine beetle damage and along with its unusual layout, it was felt that this baseline could not be located carefully enough for it to be included in a resampling effort. Thus 12 baselines were monitored in 1994, but only 11 in 2006-2008 and 2019.

Sutter et al. (1994) collected multiple types of data with the intent of providing baseline monitoring data to park managers to assess change within these calcareous glades. Sources of change include natural succession, fire exclusion and other human impacts. Per Sutter et al. (1994), these data, “…can be used to determine management activities, develop protection actions, and gain a greater understanding of these significant resources.” The multiple types of data collected in 1993 included glade species lists and various types of data collected from established baselines, each of which are briefly described below.

Glade Species Lists Glade specific species lists were developed for each identified glade over the course of the initial field season (Sutter et al. 1994). The stated purpose of these lists was to allow glades to be compared “…for their diversity, rare species, and species composition.” Glades were initially numbered 1–24 (23 was skipped). Glade 3, 8, and 24 were subsequently split over the course of that initial season into an A and B. Species lists for 3A, 3B, 8A and 8B were maintained separately. A single combined list was maintained for 24. Collection of these species lists has not occurred since that initial effort in 1993.

Baseline Monitoring For all other types of measurements, a baseline was established along the longest axis of the glade openings and marked with metal tags on nearby trees. The diameter breast height (DBH) of tagged 3

trees, along with distance and compass bearing between tagged trees and in some cases distance and compass bearing from the tagged tree to the baseline were recorded. Tag trees were established at or very near the beginning and end of each baseline and at midpoints in instances where the baseline bearing needed to “bend” in order to remain within the glade along its longest axis. Hand drawn maps were also provided to further assist in the relocation of the baselines. Monitoring transects were established off the baseline randomly within consecutive 10- to 20-meter (32.8- to 65.6-feet [ft]) segments, depending on overall glade length (Table 1).

Monitoring transects were established at a right angle to the baseline (using a sighting compass). These monitoring transects extended past the edge of the glade to the closest tree within the woodlands and a nail was used to mark the endpoints. All of this information was organized into an Appendix of the initial report (Sutter et al. 1994) to aid in relocation of the baselines.

Table 1. Baseline length, segment (i.e., interval in which monitoring transects were randomly placed), number of monitoring transects, and combined monitoring transect length [m—meters].

Glade Baseline Length Segment Length # Monitoring Combined Monitoring Transect (m) (m) Transects Length (m)*

2 119.02 10 12 275.06

3A 54.35 10 5 92.36

3B 84.10 10 9 147.32

6 59.09 10 6 109.97

8A 115.16 10 11 274.01

8B 141.60 15 10 210.95

11 56.35 10 5 85.47

13 60.95 10 10 463.53 (13**) (37.55) (10) (–)

14 99.40 10 10 287.10

21 49.85 10 5 97.10

24A 86.00 10 8 337.89

24B*** 201.20 20 10 368.49

* Calculated from Distance to Glade Edge found in Appendix I, Sutter et al. (1994) ** Glade 13 baseline was established with a disconnected section, resulting in two sections or lengths. The second section is listed in parentheses. *** Sampled in 1993, only

Woody Cover The first and most important type of data involved line intercept measurements for woody cover. These are designed to track the potential encroachment of both native and non-native woody plants that might compete with the sun loving herbaceous characteristic glade species that are of 4

conservation concern. For woody cover measurements, it was interpreted that the original study divided plants into two classes: those 1 meter (3.3 ft) or greater, and those between 0.5–1 meter (1.6– 3.3 ft). Instructions provided in Sutter et al. (1994) were not clear, and later information provided by the lead investigator (Rob Sutter pers. comm.) indicated that these two measurements were actually height classes and that all vegetation within these two classes were measured. As a consequence, the second monitoring effort (Govus and Lyons 2009) under measured vegetation within the lower class (< 1 meter [3.3 ft]), and the results between the initial and first resample were not comparable. An additional error occurred during this most recent monitoring effort (i.e., 2019). The intention was to resample as was sampled in 2008 (using plant height as a criteria) – but a mental error led to the collection of the lower plant class data as plants less than 1 meter (3.3 ft), rather than plants less than 1.0 m but ≥ 0.5 m. So once again, data regarding plants less than 1 meter cannot be compared. Coverage by length along the monitoring transects were measured for each species and summarized as percent cover along the total length of each monitoring transect.

Data on woody cover were originally collected for twelve baselines at nine different glades or glade complexes (2, 3A, 3B, 6, 8A, 8B, 11, 13, 14, 21, 24A and 24B). These data were resampled in 2006– 2008 and 2019 for 11 of the 12 baselines. Field crews in 2006-2008 and 2019 were unable to relocate the tag trees for the 12th baseline (i.e., 24B).

Ground Cover Ground cover characteristics were collected during the initial sampling event (Sutter et al. 1994) to document the condition and composition of the ground cover layer within the glades in terms of general classes of biotic and abiotic components (Sutter et al. 1994). Classes of abiotic substrates and plants were constructed and included categories such as “tufted grasses,” “litter,” “forbs, grasses, sedges,” “soil,” “gravel,” “rock,” “cobble,” “Pleurochaete squarrosa (a moss of limestone glades),” and “other mosses.” The same approach used for woody cover of measuring and recording length along the monitoring transects for each class of ground cover component (≥ 5 centimeters [2 inches {in}]) was used. The goal of these measurements was to detect changes from increased trampling (with an expected increase in soil and gravel and decrease in grass and forbs)—or increased shading (with an expected decrease in grasses and forbs and an increase in litter and soil).

Ground cover data were collected on a subset of the “nine” glades chosen for woody cover monitoring. The six chosen included 3B, 8B, 11, 14, 21 and 24B. These data were not collected during the 2006–08 efforts or the most recent 2019 sampling effort.

Herb Density The third and final type of baseline data collected during the original study was the use of 1-meter (3.3-ft) wide belt transects to collect plant density data for selected glade species. This was accomplished by laying a 1-meter frame along the monitoring transects, effectively creating a 1- meter wide belt transect, segmented into 1-meter frames, along the length of the monitoring transects. Within these frames select herbaceous species were counted and recorded as well as woody species less than 0.5 meter (1.6 ft; Sutter et al. 1994). These data were collected from the same baselines chosen for the collection of ground cover data (i.e., 3B, 8B, 11, 14, 21 and 24B.

During the 2006–2008 monitoring effort, a decision was made to substantially reduce the number of species tallied. As an example, over 40 species were tallied in glade 3B in 1993. The list was reduced based upon glade endemism, abundance during the initial monitoring effort, and ease of identification (Govus and Lyons 2009). Thus, these data were recollected in 2006–08 and 2019, utilizing a much reduced and glade specific list in five glades (i.e., 3B, 8B, 11, 14, and 21). As noted earlier, field crews were unable to relocate the tag trees for baseline 24B.

Additional details regarding previous sampling events can be found in Sutter et al. (1994), Govus and Lyons (2009) and Sutter et al. (2011).

Follow up Taxonomic Issues Relating to the 1993 Data During the course of the initial round of data collection, it became apparent that certain species documented by the original study were incorrectly identified (some of these are difficult entities to properly name). Others have been taxonomically reclassified. In particular, for woody plant coverage, blue ash (Fraxinus quadrangulata) was frequently identified with a high cover value in both height classes (but curiously absent from the plant density measurements for woody species < 0.5 meters [1.6 ft]). This species is easily identified by its square and often winged stems. No evidence of blue ash was seen in any of the glades during sub sequential studies. No observations of this species as being widespread was reported during the inventory and monitoring effort led by NatureServe during three years of field work from 2003–2006 (Govus and White 2006). It is assumed that the early study encountered the more common white ash (Fraxinus americana) and incorrectly identified it as blue ash. For comparison purposes, this study uses their coverage values for blue ash and compares them to the values of white ash (which is fairly frequent in nearly all glades as a small tree or sapling).

Another species that is strongly suspected of being incorrectly identified from the plant density measurements in 1993 is Packera plattensis (= Senecio plattensis, prairie ragwort) which was frequently reported from many of the monitoring transects. No evidence exists for the occurrence of this species in Georgia (Tom Patrick, pers com; Weakley 2019) and it was not observed during this study. It was most likely confused with a more common glade species, balsam ragwort (Packera paupercula var. paupercula—a somewhat smaller concept than Senecio pauperculus) which was documented from the same monitoring transects as the prairie ragwort. Sometimes young individuals of balsam ragwort can have a degree of tomentum along the base of the leaves (typically disappearing with age) which can cause confusion with proper identification. All of the diminutive species of ragwort in 2006-2008 and 2019 sampling efforts are regarded as Packera paupercula var. paupercula.

In addition, it is believed that the chestnut oak (Quercus montana) infrequently reported within the monitoring transects of the first study was really chinquapin oak (Quercus muhlenbergii), a limestone species often found within the glades. These species have similar looking leaves. Despite extensive field work at Chickamauga Battlefield, this investigator has never seen chestnut oak within the glade system. Since it is not impossible for this to occur, and the amount of Quercus montana recorded in 1993 was small, this potential error was ignored and will not have a significant impact on overall woody cover results. A final likely misidentification seems to occur within the genus Celtis. 6

The original study variably identifies these as C. occidentalis, C. laevigata and Celtis sp. Our team strongly believes that all of the Celtis occurring in these glade systems are Celtis tenuifolia, a species specially adapted to the dry and harsh conditions of the glades.

Under the category of change in nomenclature, the species referred to in the initial study as Rhamnus caroliniana is now considered to be Frangula caroliniana (Weakley 2019). It is a frequent and often abundant woody component of limestone glades. In addition, the larkspur associated with these glades was originally identified as Delphinium virescens which was an older and broader concept. Taxonomists now treat this taxon as a finer concept, Delphinium carolinianum ssp. calciphilum (Weakley 2019).

Lastly, and perhaps more problematic for data comparison purposes, Viola palmata (wood violet) was reported exclusively in the early study as a frequently abundant species. Whereas Viola palmata may occur in some glades, the more regularly encountered species is Viola egglestonii (glade violet)—a glade specialist and more distinctively lobed violet. The acaulescent (without leafy stem) violets are a difficult group to sort out and some taxonomists include V. egglestonii within the concept of Viola palmata (V. palmata var. pedatifida) or within Viola septemloba (V. septemloba ssp. egglestonii). This study treats the deeply dissected, glabrous violets within the glades as V. egglestonii and assumes most (if not all) of the violets reported in the original 1993 data were this species.

The current report uses Weakley 2019 as its taxonomic standard, as this is the source of nomenclature for other ongoing inventory and monitoring efforts at Chickamauga and Chattanooga National Military Park. Any discrepancies in describing taxa reported by earlier reports can be resolved by referring to synonymy.

Photomonitoring procedures One suggestion from the second round of sampling the cedar glades at Chickamauga and Chattanooga National Military Park was the need for the development of photomonitoring techniques to quickly capture qualitative images of glade conditions. As a part of this third round of resampling (2019), a procedure was developed and photomonitoring data forms were generated to document the collection of photographic images of all of the glades sampled. This simple process involves using the baseline starting, midpoint and endpoints as photo stations to take images of the glades. In general, if the baseline is within the middle of the glade, photos are taken in the direction of the baseline, but for glades where the baseline is along the edge of a glade (several small glades are sampled this way), the camera is directed towards the center of the glade and the azimuth of the direction is recorded. It is intended that the data recorded regarding photographs taken during this sampling serve as a template for subsequent efforts. A detailed protocol was also developed to name the photos so that it indicates the glade number, tag tree number, distance along baseline and azimuth of the photo. Forty-seven photos of the 11 glades sampled were taken, labeled and are currently archived with the National Park Service. These should provide a valuable quick method of evaluating the habitat quality of the glades. Examples of pristine versus degraded glade photos are provided in the results section of this report.

Results of the Third Round of Sampling

As was the case in 2006-2008, a total of 11 baselines were sampled in 2019. An ArcView shapefile of the baselines is archived by the National Park Service and includes starting and endpoint locations as well as midpoint witness tree locations and tag numbers (when they were a component). Figure 1 shows the general location of sampled glades. Some glades that are in close proximity were divided into “A” and “B” segments by Sutter et al. (1994) and are sampled separately. All 11 glades (2, 3A, 3B, 6, 8A, 8B, 11, 13, 14, 21 and 24A) were sampled for woody cover with line intercept transects. Glades 3B, 8B, 11, 14 and 21 were sampled with belt transects for plant density (Figure 3.). The line intercept transect formed the established edge of the belt transect (refer to previous Methods section). Both the line intercept and belt transect are often referred to as monitoring transect(s) in this report, as they are the same basic line or transect.

An important point should be made regarding the methods used to collect and analyze these data to compare them with the initial study of 1993. It was hoped that some statement could be made with what was defined by The Nature Conservancy as the “glade edge.” Their definition was that “the glade edge was recognized where the cover of woody species was 100% for one full meter of the line intercept” (Sutter et al. 1994). This definition does not make sense as even within fairly open glades there are areas where cover by large arborescent species (especially red cedar) can be greater than a meter. This definition also did not make sense to the field crew(s) implementing the methodology in 1993 as can be seen by the fact there are several transect notes indicating sections of monitoring transects where woody and ground cover data were not collected “due to a non-glade area.” These notes, which include start and ending points along monitoring transects where ground cover, woody cover and belt transect data were not collected, can be found in Appendix 1 of Sutter et al. (1994). No other mention of these “field modifications” in data collection can be found in Sutter et al. (1994). There are no other conditions stipulated in the original report, and in examining previous end points defined as glade edges based on measurements from the baseline, many of these appear arbitrary and one could argue in many cases that the glade edge could be either farther away from the baseline or closer to the baseline. The best indication of glade edge, which seemed more consistent to researchers conducting this study, was the point at which leaf litter became continuous and there was no herbaceous cover. This point always occurred closer to where the woodlands surrounding the glade complex became continuous. It was also noted, during the course of collecting woody cover data, some transects (especially in glade 14) included fairly long sections of woodland habitat that were islands of woody vegetation within the glade matrix. In looking at these variable circumstances, it was decided for the purposes of the second and third resample, it was imperative to attempt to resample the original transects as closely as possible to the original measurements so that the change in woody cover could be accurately compared. This included adhering to the non-collection sections of the monitoring transects recorded by the 1993 field team(s). No attempt was made to redefine the glade edge or change dimensions of the original transects, beyond those noted in 1993.

Figure 1. General location of glades sampled for woody cover; some were further divided by Sutter et al. (1994) into separate glades for monitoring purposes (i.e., glade 3A and 3B). A total of 11 baselines representing 11 glades were sampled.

Change in Large Woody Cover Greater Than or Equal to 1 meter (3.3 ft) The woody cover data (≥ 1 meter [3.3 ft]) are compiled and summarized as average percent cover by species for each glade in tables provided in Appendix I. A recent review by the National Park Service Regional Inventory and Monitoring Program of the data analysis techniques of earlier reports noted that if woody cover is correlated with transect length in a glade, then the average woody cover across transects would differ from the estimated woody cover for that glade (“glade mean,” defined as the proportion of glade expected to be covered by vegetation of a woody species). For example, if shorter transects (perhaps representing glade edges or narrow pinch points) tend to have greater encroachment of woody cover than do longer transects, then averaging woody cover across transects 10

without accounting for transect lengths may yield a higher cover estimate than the glade mean. This overestimate would be due to weighting the woody cover estimates of shorter transects equally with that of longer transects, when the latter actually represent a larger portion of the glade.

In an attempt to more accurately portray glade conditions within this report, all woody cover values for the three sample events are reported as the glade means for a particular woody species in a particular surveyed glade. The woody cover “glade mean” for a species is calculated as the total length of surveyed transect overlapped (covered) by large (≥ 1 meter [3.3 ft]) woody plants of that species, divided by the total length of surveyed transects in the glade. This calculation is equivalent to taking a weighted mean of transect-level percent woody cover, in which the woody cover estimate for each transect is weighted by the length of that transect. Additionally, and perhaps more importantly, because the glade mean represents a statistical sample, 95% confidence intervals for glade means can be bootstrapped from the sampled transect data, providing a level of confidence or statistical significance to the calculation.

Our presentation of woody cover as glade means differs from how woody cover was summarized in earlier reports (Sutter et al. 1994; Govus and Lyons 2009; Sutter et al. 2011). Those earlier studies reported what we will refer to as a “transect mean” which averages the percent woody cover across transects, with all transects equally weighted despite differences in transect length. As a result, percent cover values reported in this report will differ somewhat from values reported in those earlier reports.

Large woody cover data were recorded by species along the monitoring transects. It should be noted however, these data were not recorded in such a way as to account for overlap between species. For instance, absolute woody cover for species A + B could be 50%, but that 50% could mean anywhere from 25% of the transect was covered (because species A and B completely overlapped) to 50% (because species A and B had no overlap). Given this important caveat, there seems an obvious and generally consistent pattern of increase in total woody cover (i.e., summed across all species) within each of the 11 monitored glades (Table 2.). The change in percent total woody cover from 1993 for each glade is summarized in Table 2 where percent total woody cover change is calculated by summing the percent change of individual species between the sampling events (i.e., 1993 to 2006- 2008 and 1993 to 2019). For example, if species A increased by 12% and species B decreased by 5%, this would equate to a 7% increase in total woody cover.

The consistency of the results suggests that the cedar glades may be slowly and steadily closing in and becoming more woodland like.

Red cedar (Juniperus virginiana), which is the highest cover species within the sampled glades by a large margin (Figure 2 and Table 3), exhibited a consistent increase within the majority of glades through time (Figure 2). This clear pattern of consistent increase through time across a majority of monitored glades was not replicated for other high cover species, however.

Loblolly pine (Pinus taeda), the second highest cover value species across the 11 monitored glades in 2019 (Figure 2 and Table 3), had a mean value of 9% cover during both the 1993 and 2006-2008

sampling sessions but increased to 15% mean cover in 2019 (Figure 2). This overall increase was driven by large increases in cover in four of the 11 glades (Figure 2). Yet, it exhibited a decline on at least three glades and was all together absent on four (Figure 2 and Table 3).

Among the eight highest cover species, Virginia pine (Pinus virginiana) is the only species to exhibit a somewhat consistent decline over time in most glades (Figure 2) and is no longer one of the highest cover species based on 2019 results. Curiously, Chinese privet (Ligustrum sinense) exhibited a rather consistent increase in glades where it was present from 1993 to 2006-2008 and just as consistent of a decrease from 2006/08 to 2019 (Figure 2). Remaining species with high ranking cover values in 2019 show wide disparity in trends across glades and through time (Figure 2).

Table 2. Percent change in total woody cover summed across species from 1993 to 2006-2008 and 1993 to 2019 in monitored glades on Chickamauga and Chattanooga National Military Park.

Glade # 2006-2008 2019

2 34.1 72.7

3A 52.2 94.3

3B 17.1 17.7

6 -3.89 45.5

8A 31.5 81.4

8B 20.8 44.6

11 4.75 18.0

13 26.5 47.9

14 33.4 46.1

21 32.1 56.6

24A 13.0 17.2

Standard errors or confidence intervals for the mean % woody cover across glades were not calculated because the eleven sampled glades do not represent a statistical sample from the overall population of glades within Chickamauga and Chattanooga National Military Park (Sutter et al. 1994). Instead minimum and maximum cover values for each species are presented in Table 3 to provide a measure of the variability of individual species cover across the sampled glades. As can be seen from these values the amount of cover that a particular species provides varies widely across glades and sampling periods. For reference purposes, Table 4 contains the 2019 glade by glade cover value for select species, primarily those with high cover values relative to all the woody species recorded.

Figure 2. Glade means for percent woody cover for eight of the highest cover species. Red line is mean across glades. Y axis scales differ among facets.

Table 3. Glade minimum and maximum percent woody cover for eight of the highest cover species by sampling session.

Species 1993 2006/08 2019

Mean Min Max Mean Min Max Mean Min Max

Juniperus virginiana 20.8 10.1 48.6 32.1 19.6 58.9 47.3 22.4 65.1

Pinus taeda 9.12 2.17 20.3 9.46 0 28.1 14.9 0 46.4

Ligustrum sinense 1.20 0 4.54 4.21 0 12.8 3.42 0 11.2

Frangula caroliniana 0.44 0 3.61 2.06 0 5.90 2.80 0 9.57

Cercis canadensis 0.50 0 2.73 2.06 0 8.75 2.43 0 7.42

Ulmus alata 0.31 0 2.29 1.83 0 5.48 2.70 0 7.88

Pinus virginiana 2.41 0 6.13 1.42 0 5.57 1.00 0 3.66

Rhus aromatica 0.15 0 0.71 2.30 0 5.91 2.02 0 7.42

Table 4. 2019 Percent cover values for select high cover species greater than or equal to 1 meter (3.3 ft) showing the glade mean value for each sample glade.

Species Glade 2 Glade 3A Glade 3B Glade 6 Glade 8A Glade 8B Glade 11 Glade 13 Glade 14 Glade 21 Glade 24A

Juniperus virginiana 58.91 59.07 65.12 55.94 49.59 44.40 22.35 39.10 31.83 57.74 36.36

Pinus taeda 33.57 26.11 0 0 46.42 30.03 6.02 9.68 12.28 0 0

Ligustrum sinense 1.35 0 2.22 0 9.28 2.09 11.22 0 0 2.49 8.98

Frangula caroliniana 8.04 9.57 0.65 0.99 1.79 0.45 0 2.85 1.94 4.55 0

Ulmus alata 7.61 4.26 2.41 0 2.33 7.88 2.59 0.76 0.15 0.00 1.74

Cercis canadensis 0 0 1.94 7.42 3.48 0.00 0 2.07 4.68 7.12 0.04

Rhus aromatica 2.08 0.65 0.98 2.48 0.51 0.19 1.64 3.46 2.78 0 7.42

Pinus virginiana 0 0 3.66 3.34 1.15 0 1.44 1.39 0 0 0

Although the surveyed glades are not a statistical sample of CHCH glades, the surveyed transects within each glade can be considered a statistical sample for that glade and 95% confidence intervals can be calculated. Table 5 contains the difference in glade mean percent woody cover, comparing 1993 and 2019 results. A value greater than zero indicates percent woody cover was higher by that amount in 2019 than in 1993. Values in red indicate a statistically significant result (i.e., 95% CI of the difference did not include zero at alpha = 0.05). Results from 2006–08 surveys were not factored into these calculations. Table 5 includes all results (either positive or negative) that were determined to be significant. In other words, if a species did not exhibit a statistically significant increase or decrease in at least one glade it is not included within Table 5 with the notable exception of Pinus species.

Table 5. Difference in percent woody cover, comparing 1993 and 2019 values. A value greater than zero indicates % woody cover was higher in 2019 than in 1993. A red number indicates the difference is statistically significant at alpha = 0.05 (i.e., the 95% CI does not include 0). Table includes all species with a significant change in cover in at least one glade in addition to Pinus species.

Species Glade 2 Glade 3A Glade 3B Glade 6 Glade 8A Glade 8B Glade 11 Glade 13 Glade 14 Glade 21 Glade 24A

Cercis canadensis N/A N/A 1.74 5.87 3.48 N/A N/A 2.01 4.65 4.39 -0.91

Cornus florida 1.90 1.35 N/A N/A N/A N/A N/A N/A N/A N/A N/A

Diospyros virginiana N/A N/A 0.03 N/A 0.22 N/A 1.51 1.09 2.42 0.72 N/A

Frangula caroliniana 7.49 9.57 0.65 -2.62 1.70 0.30 N/A 2.61 1.72 4.55 N/A

Fraxinus americana -1.41 5.51 1.37 1.36 2.01 0.36 2.18 1.13 1.67 N/A 1.94

Juniperus virginiana 36.6 45.5 16.5 31.85 30.89 27.89 7.48 24.6 21.7 45.08 3.51

Ligustrum sinense 0.68 -0.05 2.22 NA 7.68 -0.80 6.68 N/A N/A 2.49 5.56

Pinus echinata 0.62 7.91 N/A N/A 0.30 N/A 0.94 0 N/A N/A 0

Pinus taeda 18.5 22.5 -5.58 -2.17 26.1 9.94 -5.44 5.56 5.92 -8.42 -3.11

Pinus virginiana -0.61 -4.22 -2.47 -1.34 0.48 -0.90 -3.57 -1.36 -0.10 -0.71 -0.70

Quercus N/A N/A N/A 7.27 N/A N/A 2.56 3.99 1.95 N/A 0.63 muehlenbergii

Quercus stellata N/A -0.41 1.37 1.14 0.91 0.50 1.46 1.55 2.71 2.58 0.96

Rhus aromatica 1.81 0.40 0.98 2.48 0.51 -0.52 1.64 3.42 2.78 0 7.10

Ulmus alata 5.32 4.26 2.41 0 2.33 7.48 2.59 0.72 0.24 N/A 1.07

Change in Plant Density Summary Results Figure 3 shows the location of glades sampled for plant density measurements. Per Sutter et al. (1994) six of the nine glades where woody cover data were collected were selected for collection of plant density data. This subset of six glades was chosen based on some combination of glade size, presence of rare species, species diversity and level of disturbance. As noted earlier, field teams in 2006 and 2019 were unable to relocate the tag trees for glade 24B which reduced the overall number of glades sampled for herb density from six to five.

Another important point worth repeating is the list of species monitored for plant density in 1993 was reduced during the 2006-2008 monitoring effort and this reduced list was carried forward to 2019. The reasoning behind the list reduction centered on the fact the 1993 list was too long and included many widely distributed and non-glade specific species. Many low frequency species based on the 1993 results were also excluded. As stated in Govus and Lyons (2009) the criteria for species selection were species

• that are significant glade endemics

• have a fairly substantial population (based on the 1993 data)

• are large enough and/or easy to detect and tally accurately

It is also important to note that this list varied by glade, largely based on the second criteria above. If the species did not have a fairly substantial population in 1993 it was not tallied in 2008 or 2019. However, characteristic glade species (see below) were surveyed for in each of the five sampled glades in 1993, 2006-2008 and 2019.

Figure 3. General location of glades sampled for plant density measurements.

The compiled data for the plant density transects on a glade by glade basis is presented in Appendix II in tabular form. Table 6 (below) shows the change in average plant density of characteristic glade species for the five glades sampled from 1993 to 2019. The values reported here differ in two important ways from Sutter et al. (1994). One is that we are using the glade mean rather than the transect mean (refer to earlier discussion on this point in this report). In addition, in the process of recovering, reviewing and recalculating values, it was discovered that Sutter et al. (1994) oddly did not include the B side of the transects in their calculations (most sampling transects include a long A side and a shorter B side), our analysis/calculations within this report and appendices include the B side transect data. As such 1993 values reported in Table 6 and 7 will vary from those reported in Sutter et al. (1994) and any subsequent report relying on Sutter et al. (1994) as the source.

Nashville breadroot (Pediomelum subacaule), glade violet (Viola egglestonii) and Gattinger's prairie- clover (Dalea gattingeri) exhibited consistent (i.e., across the sampled glades) and statistically significant declines in the glades where they were monitored. One could arguably include pinnate prairie coneflower (Ratibida pinnata) as it exhibited significant declines and extremely low densities in two of three monitored glades. But it also increased, though not significantly in glade 14. Perhaps most alarming is cutleaf prairie dock (Silphium pinnatifidum), which was surveyed for in three glades but went undetected in 2019. Glade larkspur (Delphinium carolinianum ssp. calciphilum) and Tennessee gladecress (Leavenworthia exigua var. exigua) also exhibited extremely low densities in 2019 surveys, including going undetected in some glades. The fact that some of these declines (and absences) were not significant is a function of the extremely low densities already observed for these characteristic glade species in 1993.

At the other end of the spectrum balsam ragwort (Packera paupercula var. paupercula) and glade St. John’s-wort (Hypericum dolabriforme) showed statistically significant increases in three of the five glades where surveys were conducted. Interestingly or confounding is the fact balsam ragwort and St. John’s-wort, exhibited substantive declines between 1993 and 2006-2008 survey efforts (Table 7, Govus and Lyons 2009). Yet, they averaged the highest densities among the characteristic glade species in 2019 (Table 7).

With that said, however, it is worth pointing out once again that glades were subjectively selected for monitoring and as such do not represent a randomized sample from among the glades on Chickamauga and Chattanooga National Military Park. As such statistics pooling across glades should be viewed with this caveat in mind. It is also worth noting there was also generally a great deal of variability amongst the glades sampled within survey periods, as can be seen from the minimum and maximum values also reported in Table 7. As a reminder, a total of five glades were sampled for herb density in both the 2006-2008 and 2019 survey efforts. It would have been six but field crews in 2006-2008 and 2019 were unable to relocate the tag trees and thus reestablish the baseline for glade 24A. For consistency purposes results for glade 24A were excluded from the results reported in Table 7.

Table 6. Characteristic glade species 1993 density (individuals per square meter) and subsequent change in density based on 2019 survey results. The 1993 density is bracketed followed by the 2019 difference in density, unbracketed value. A negative 2019 value indicates density declined by that number of individuals/m2. The 2019 change in density value is red if statistically significant at alpha = 0.05.

Species Glade 3B Glade 8B Glade 11 Glade 14 Glade 21

Pediomelum subacaule N/A* [11.88] [10.37] [15.34] [0.91] -7.31 -6.03 -8.15 0.22

Viola egglestonii [0.92] [2.94] [3.97] [3.84] [1.52] -0.68 -2.38 -3.35 -3.63 -1.36

Dalea gattingeri [2.25] [1.76] [3.46] [2.56] [3.96] -1.50 -0.87 -2.78 -0.71 -3.24

Ratibida pinnata [1.09] N/A* N/A* [0.09] [1.97] -1.08 0.48 -1.86

Ophioglossum engelmannii [2.20] N/A* [1.44] [0.01] N/A* -1.70 -0.62 -0.01**

Silphium pinnatifidum N/A* [0.72] N/A* [0.57] [0.03] -0.72** -0.57** -0.03**

Delphinium carolinianum ssp [1.93] [0.02] N/A* N/A* [0.05] calciphilum -1.17 -0.02** -0.05**

Leavenworthia exigua var. [0.30] N/A* [0.66] [0.003] [0.26] exigua -0.30** -0.55 -0.003** -0.24

Scutellaria leonardii [0.79] [0.12] [0.43] [0.30] [1.24] -0.11 0.35 0.19 0.36 -0.02

Ruellia humilis [0.83] [1.61] [3.29] [2.78] [5.50] -0.46 4.82 4.42 -2.08 -4.86

Packera paupercula var. [0.76] [0.51] [0.81] [1.23] [0.02] paupercula 0.60 8.61 -0.42 5.62 -0.01

Hypericum dolabriforme [1.08] [0.80] [2.64] [1.79] [3.50] 2.86 3.96 8.21 1.01 11.35

* NA indicates species was not detected in both the 1993 and 2019 survey efforts. ** Indicates (emphasizes) species was not detected during 2019 surveys

Table 7. Plant density values (plants/m²) of characteristic glade species averaged across sampled glades along with minimum and maximum glade averages (parenthesis). For each species, n = number of glades included in calculation.

Species 1993 2008 2019

Pediomelum subacaule 9.63 5.37 4.31 (n=4) (0.91–15.34) (1.09–8.11) (1.13–7.19)

Dalea gattingeri (n=5) 2.80 1.04 0.98 (1.76–3.96) (0.59–1.80) (0.68–1.85)

Viola egglestonii (n=5) 2.64 0.21 0.36 (0.92–3.97) (0.08–0.33) (0.16–0.62)

Ophioglossum 1.22 0.38 0.66 engelmannii (n=3) (0.01–2.20) (0.0–0.47) (0.0–0.82)

Ratibida pinnata (n=3) 1.05 0.10 0.23 (0.09–1.97) (0.0–0.21) (0.01–0.57)

Silphium pinnatifidum 0.65 0.04 0.00 (n=3) (0.03–0.72) (0.0–0.08) (0.0)

Leavenworthia exigua var. 0.39 0.32 0.04 exigua (n=4) (0.003–0.66) (0.0–0.64) (0.0–0.11)

Delphinium carolinianum 0.67 0.00 0.25 ssp. calciphilum (n=3) (0.02–1.93) (0.0) (0.0–0.76)

Scutellaria leonardii (n=5) 0.58 0.01 0.73 (0.12–1.24) (0.0–0.05) (0.47–1.22)

Ruellia humilis (n=5) 2.80 1.31 3.17 (0.83–5.50) (0.27–3.80) (0.37–7.71)

Packera paupercula var. 0.71 0.51 3.55 paupercula (n=5) (0.02–1.23) (0.0–2.11) (0.01–9.12)

Hypericum dolabriforme 1.96 0.99 7.44 (n=5) (0.80–3.50) (0.19–1.92) (2.80–14.85)

Discussions and Conclusions

Woody Cover Total woody cover (i.e., summed across all species) of plants greater than or equal to 1 meter (3.3 ft) increased within and across the sampled glades, having increased on 95% of the sampled transects (86 of 91 transects) when comparing 1993 and 2019 results. This would support the conclusion that the increase in woody cover is widely dispersed across the monitored glades. Additionally, 10 of the 11 monitored glades contained at least one woody species which exhibited a statistically significant increase in cover from 1993 to 2019. The only glade to have no statistical increases was 3B. Seven of the remaining 10 monitored glades had three or more woody species exhibiting a statistical increase in cover. However, in looking at change in woody cover by species it is clear the most consistent and substantial change in woody cover resulted from an increase in Eastern red-cedar (Juniperus virginiana) and loblolly pine (Pinus taeda) with eight and six of the 11 sampled glades showing a statistically significant increase respectively. These are the only two species to show statistical increases on more than half of the glades, and they are the only two species to have double-digit increases in cover on any of the glades monitored in 1993 and 2019. Furthermore, cedar and loblolly pine accounted for 60% or more of the increase in total woody cover observed between 1993 and 2019 in nine of the 11 glades. The two exceptions being glade 11 and 24A. Interestingly these are the smallest and second largest monitored glades, based on combined transect length (Table 1).

Almost without exception red cedar dwarfed nearly all other increases in cover in the respective glades, ranging from 21 to 45% in the eight glades where it had a statistically significant increase in cover. The fact that Juniperus is at or near 40% average cover in a majority of the sampled glades further demonstrates that these glades are now approaching a woodland condition. This is not to say that other species have not exhibited notable increases. For instance, six species had a statistically significant increase greater than or equal to 5% in at least one glade. These include Eastern redbud (Cercis canadensis) Carolina buckthorn (Frangula caroliniana), white ash (Fraxinus americana), Chinese privet (Ligustrum sinense), fragrant sumac (Rhus aromatica) and winged elm (Ulmus alata). However, it is worth noting that many of these species had larger increases from 1993 to 2006-2008 for some glades followed by declines from 2006-2008 to 2019. In short, the results for species other than red cedar are somewhat mixed and much less compelling of an increase.

Thus, with the exception of red cedar, species level analysis of woody cover data is highly variable across the monitored glades and sampling sessions. While it is true some species have shown moderate increases (~5% or more) over the last two-plus decades species specific increases are not consistent across the monitored glades. Even loblolly pine, which had statistical increases on six of the monitored glades, exhibited declines on the remaining five though two of these were not significant. In short, the majority of woody species have not exhibited a single trajectory of increase over time and have not exhibited a statistically significant increase in cover in a majority of monitored glades. The one monumental exception being eastern red cedar (Juniperus virginiana).

Duncan et al. (2007), in summarizing Quarterman (1989) states that “Although colonizing trees may be a natural component of the glades, they may negatively affect populations of light-demanding species typical of open glade habitat.” The increase in woody cover at Chickamauga and Chattanooga National Military Park has most likely been enhanced by fire exclusion (Govus and White 2006). In studying fire reintroduction into calcareous glade systems, Duncan et al. (2007) found that in all of the glades they studied, ecotonal vegetation appeared to be advancing into the habitat. They further state that, “Historically, fires along the ecotone probably slowed or prevented encroachment, and prescribed fires could provide the same service.”

Plant Density The results of plant density measurements are mixed and at least initially, difficult to interpret. This is particularly true when considering results from the previous sampling event in 2008 (Govus and Lyons 2009) pointed to a very strong relationship between increases in woody cover and decrease in plant density measurements for all glade characteristic species. When examining 2019 data there is possibly a weak negative association between percent total woody cover and herb density, but factors other than the increase in woody cover should also be considered.

National Weather Service (2020) records for the year 2007 report an exceptional drought for the Chattanooga area with an annual rainfall total of 38.6 inches (98 centimeters [cm]; compared to a historical seasonal average of 52.48 inches [133.3 cm] per year). For the prior year leading up to the 1993 sample the rainfall reported for the study area was 55.86 inches (141.9 cm), nearly three inches (7.6 cm) above average. Cedar glades are believed to be maintained primarily by a combination of extreme edaphic conditions associated with the shallow, calcareous, rocky soils as well as extreme climatic variation (alternating wet winter seasons with hot and often droughty summers; Baskin and Baskin 1978). The exceptional drought of 2007 may have had a strong influence on the plant density measurements made in 2008 (Figure 4). Rainfall records for 2018 show a near record high rainfall of 65.9 inches (167.4 cm; 13 inches [33 cm] above average), and an unusually high 11.3 inches (28.7 cm) for the month of March just prior to the most recent sampling event. It is certainly reasonable to conclude that rainfall has some explanatory value in the paucity of glade characteristic species observed in 2008 surveys and the comparative abundance in 2019. However, this arguably makes the statistically significant declines of some species, specifically Pediomelum subacaule, Viola egglestonii and Dalea gattingeri, more dramatic.

Baskin and Baskin (2003) refer to species “characteristic” of cedar glades. The 12 species in Table 7 have all been considered to be characteristic species though some do occur over a wider range of habitats. Baskin and Baskin (2003) went on to evaluate greater than 400 vascular plant species of cedar glades in the southeastern United States. From this list they identified 21 endemic or near endemic taxa to be “the most distinct floristic element of the southeastern glades.” Due to their reduced distribution and lack of competitive advantage outside of glade environments, these endemic or near endemic taxa are deserving of greater conservation and management attention.

Figure 4. Sum total change in characteristic glade species densities over three sampling intervals at Chickamauga and Chattanooga National Military Park. Refer to Table 6 for individual species averages and confidence intervals.

Per Baskin and Baskin (2003) Leavenworthia is the only genus that is restricted to cedar glades and glade-like areas in unglaciated eastern United States. Additionally, Dalea gattingeri, Delphinium carolinianum ssp. calciphilum and Pediomelum subacaule were also identified as glade endemic or near endemics of the Ridge and Valley of northwest Georgia (among other regions). Within our data set, Dalea gattingeri and Pediomelum subacaule exhibited large statistical declines when comparing 1993 and 2019 density values within the monitored glades. Pediomelum declined in all four of the glades where it was monitored, including three statistically significant declines. Dalea declined in all five of the glades where it was monitored, including four statistically significant declines (Table 6). Most certainly a function of the already low densities first observed in 1993, Delphinium carolinianum ssp. calciphilum and Leavenworthia exigua var. exigua did not exhibit statistically significant declines in densities from 1993 to 2019. However, both were not observed in 2019 in glades where they were observed in 1993 with current densities for both species at or near what is arguably the bottom limit of detectability (Table 6).

At the other end of the spectrum, the overall increase in densities of glade characteristic species in 2019 is driven by significant increases in two species, Hypericum dolabriforme and Packera paupercula var. paupercula (Table 6). Hypericum dolabriforme was at one time considered by Baskin and Baskin (1986, 1989) to be endemic to southeastern cedar glades. However, in addition to cedar glades, this species has been found to occur in other rocky, disturbed habitats including

disturbed roadsides in the Interior Low Plateaus (Baskin and Baskin 2003). Thus, its presence or trend is no longer considered to be as indicative of glade quality. Weakley (2019) includes uplands, cobble bars, riverbanks, glades and barrens among the habitats for Packera paupercula var. paupercula. As such it also is not a clear indicator of high-quality limestone cedar glade environments.

It is not immediately obvious as to why certain species like Pediomelum have declined sharply while these other species have increased. Given the increase in woody cover it could be expected that characteristic glade (i.e., heliophytic) species would all have lower densities. However, it does stand to reason that glade endemic or near endemic species would be the first to lose any competitive advantage as habitat quality degraded (Quarterman 1950). It also makes sense that Hypericum and Packera having a wider niche are able to take advantage of the increased rainfall observed in 2019 and increase in abundance in this somewhat modified habitat, but this is conjecture. Sutter et al. (1994) established this study to track changes in habitat quality within the glades of Chickamauga and Chattanooga National Military Park and specific questions like these are beyond that scope.

What we can see is that the continued increase in red cedar and the decline in endemic or near endemic glade species points to a continued decline in glade quality on Chickamauga and Chattanooga National Military Park. As such management actions designed to reverse these trends are encouraged as the current trajectories do not bode well for the continued existence of these unique and ecologically important habitats.

Exotic Species The results of woody cover measurements reveal statistically significant increases in Chinese privet (Ligustrum sinense), a species which Weakley (2019) refers to as “…one of the most noxious of all our weeds.” Specifically, Chinese privet (Ligustrum sinense) increased substantially in glades 8A and 24A, and three glades (8A, 11, and 24A) now have total woody cover values at or near 10%. Cofer et al. (2008) state that ecological models predict that stressful environments such as cedar glades, which have a high rate of endemism, might easily be invaded by exotics. For the glades at Chickamauga and Chattanooga National Military Park, Chinese privet (Ligustrum sinense) appears to be the only invasive species that poses a serious threat to the habitat integrity at the moment. Other exotic species that have appeared in sampling transects include tree-of-heaven (Ailanthus altissima) and black raspberry (Rubus occidentalis) but these are only trace occurrences.

Of particular interest in regard to exotic species occurring within these habitats, waxy-leaf privet (Ligustrum quihoui) was sampled for the first time in one transect in glade 6. To our knowledge, this occurrence represents a state record for Georgia and is of concern because it has been reported to be a threat to diabase glade communities in North Carolina (Alan Weakley, pers com). Known occurrences range from North Carolina, Virginia, Alabama, Florida and Mississippi. It is hoped that this discovery will serve as an “early detection” situation and this small occurrence (a single shrub) can be eliminated and other potential plants may also be located and removed.

Management With such a large increase in coverage by Eastern red cedar (Juniperus virginiana) the time has come to consider removal of individual trees. This has been used as a management tool for maintaining cedar glade habitat at Stones River National Battlefield, combined with a program of NPS chainsaw training and certification (Terri Hogan, pers com). Care must be taken to make sure that removed trees are not tag trees for the baseline, and any elimination of cedar trees must include the entire removal of all the plant materials. As Baskin and Baskin (2000) have stated, “Without removal of invading woody plants by fire or other means, succession in these rocky, calcareous openings is to forest.” Fire is not likely to remove the large cedars that are now well established within the glades. Engaging with glade managers in the region such as Tennessee Department of Environment and Conservation’s Division of Natural Areas is encouraged as they have a history of actively managing glade and barren habitats using fire, manual thinning, and herbicide. The Couchville Cedar Glade near Nashville is a good example of where these management tools have been implemented successfully.

One very positive development in the management of glade habitats at Chickamauga and Chattanooga National Military Park is that a prescribed burn was scheduled for February and March of 2020 (Rob Klein, pers com). The Viniard-Alexander Prescribed Burn Unit which is 61.1 hectares (151 acres [ac]) will impact glades 13, 14 and 24A and several smaller glades and surrounding glade like habitats in the vicinity. Fire is an important component of these systems and in the absence of fire, woody plant encroachment can become a serious problem (Mills 2008). Mills (2008) also found that for the glades he studied, in the absence of fire, a 56% reduction in the number of original native species occurred over a 50-year period. Duncan et al. (2007) found that reintroduction of fire did not harm glade species and helped prevent encroachment of the surrounding forest. They also found that glade fires may not be hot enough to significantly damage roots and seed banks and that fires spread unevenly. Historically it is believed that natural fires along the ecotone have probably slowed or prevented the encroachment of woody vegetation and that prescribed fires could provide the same service. At Chickamauga and Chattanooga National Military Park, where investigators have resampled these glades over two periods, a very strong sense has developed that the glade edges are encroaching notably into the habitat. This prescribed burn should produce a marked improvement for the ecotones of all the glades where fire is able to penetrate. It will be of interest to see if the next round of sampling reflects any changes related to this fire. It is hoped, as conditions allow, that several prescribed burns can be undertaken at Chickamauga and Chattanooga National Military Park. The natural fire return interval for this area is around 15 years (Frost 1998) and fire has been excluded at this park for more than 120 years.

The removal of Chinese privet (Ligustrum sinense) from the center and edges of these glades remains an important management priority. The woody density data for each glade can serve as a guide for which glades to target first. Glades 11 (11.2% coverage by privet), 8A (9.28%) and 24A (8.98%) should be treated first, followed by glades 3B, 8B and 21. It is encouraging that even now, 25 years into the monitoring process, that four of the glades show no signs of infestation by privet. If meaningful work is sought out for conservation groups like the Youth Conservation Corps and others, privet removal would be an excellent opportunity. Since privet resprouts after cutting, the

careful use of painting the cut stems with herbicide is strongly recommended. Figure 5 shows an example of a degraded glade (24A) with a well-established privet population.

Figure 5. A baseline photograph of glade 24A showing degraded condition, particularly encroachment by Chinese privet (Ligustrum sinense).

Future Sampling During the process of collecting monitoring data from this last round of sampling, discussions developed about the amount of time required to complete all of the field work involved. Some fourteen days were required to collect woody cover and plant density data. In particular, the collection of plant density values with continuous belt transects was singled out as a possible area where the sampling protocol might be streamlined. For glades 8B and 14 a total of four days was required to sample the plant density values. These are larger glades with some long transects ranging from over 30 meters (98.4 ft) up to 47 meters (154.2 ft). One possible solution would be to randomly sample half of these longer transects with the goal that no glade should take more than one day to sample. This should be discussed with a biostatistician to make sure no bias is introduced in the sampling, and a “cut off point” for transect length should be identified for guiding the modification from the original methodology.

It should be pointed out that the 11 sampled glades vary widely in size and shape. It seems logical that larger and wider glades would show less pronounced changes in glade quality, while small glades that often are narrow ellipses with short transects might seem to be changing more quickly. The larger glades presumably have shallower soil, much more exposed limestone and less edge effect. The opposite is true for small glades that are narrow and may have deeper soil. Harper (1926) also expresses the opinion that for cedar glades to support endemic and characteristic species they need to be of “substantial width and considerable area.” Likewise, Somers et al. (1986) recognizes that floristics and composition and the rate of succession may vary depending upon glade size. In studying cedar glade habitats in Middle Tennessee, Cofer et al. (2008) noted that Leavenworthia and Pediomelum were the most frequent endemics, and more abundant in large, open glades. Given these observations, it would seem worthwhile for future efforts to investigate relationships between glade size and soil depth to observed vegetational change within the monitored glades in Chickamauga and Chattanooga National Military Park. In addition to quantifying soil depths, an evaluation of historical imagery (if available) could be used to determine previous glade boundaries and size. Both soil depths and previous boundary or size would be extremely helpful in developing future management prescriptions. Future imagery could also potentially be used to evaluate effectiveness of proposed management actions (e.g., prescribed fire, red cedar tree removal).

A final important observation was made during this most recent sampling interval, some of the early spring ephemerals, especially Leavenworthia exigua var. exigua, were nearly done blooming and fruiting and beginning to senesce, making detection of individuals very difficult. This could be the result of shifting climate patterns and shifting blooming times earlier over the 25-year period that this project has been ongoing. During the 2019 field season, plant density sampling began on May 9 (which is based on the original sampling time that The Nature Conservancy used). It is recommended that future monitoring efforts closely watch flowering times and be prepared to adjust the timing of monitoring accordingly. This may be somewhere close to the first day of May.

Lastly, though not a part of this monitoring effort, glade 3A has been observed to support a few individuals of blue wild indigo (Baptisia aberrans), another characteristic glade species (Baskin and Baskin 2003) and S2 (State imperiled) in the state of Georgia (Baptisia australis [Linnaeus)] R. Brown var. aberrans [Larisey] M. Mendenhall). While this beautiful, showy flower was not initially tracked by Sutter et al. (1994), only one plant was noticed during this last sampling period and it appeared weak and less robust due to dense shade. This species has been reported from other glades at Chickamauga and Chattanooga National Military Park (Allison 1991).

Future efforts should be made to ascertain the status of this species at Chickamauga and Chattanooga National Military Park, as there are only 22 extant occurrences (NatureServe 2020). The potential decline or loss of special concern species such as this is a reminder of how important monitoring and evaluating the environment in these sensitive habitats is. Other noteworthy glade plants occur in some of the glades but were not tallied as part of this monitoring effort. These include prairie dropseed (Sporobolus heterolepis), which is rare throughout the southeastern United States (Weakley 2019), as well as a taxon believed now to best be treated as Lithospermum bejariense (western marbleseed; Weakley 2019) a species regarded as important in glade conservation. The treatment and

understanding of Lithospermum has changed substantially in recent years and clarifying the status of this taxa at Chickamauga and Chattanooga National Military Park should be a part of future survey efforts.

Literature Cited

Allison, J. R. 1991. Packet of information on Georgia’s cedar glades. Unpublished memorandum. Social Circle, Georgia. Georgia Department of Natural Resources. 6 p.

Baskin, J. M., and C. C. Baskin. 1978. Plant ecology of cedar glades in the Big Barren Region of Kentucky. Rhodora 80:545-557.

Baskin, J. M., and C. C. Baskin. 1986. Distribution and geographical/evolutionary relationships of cedar glade endemics in the southeastern United States. ASB Bulletin, 33:138–154.

Baskin, J. M., and C. C. Baskin. 1989. Cedar glade endemics in Tennessee, and a review of their autecology. Journal Tennessee Academy of Science 64:63-74.

Baskin J. M., and C. C. Baskin. 2000. Vegetation of Limestone and Dolomite Glades in the Ozarks and Midwest Regions of the United States. Annals of the Missouri Botanical Garden, Vol. 87, No. 2, pp. 286–294.

Baskin, J. M. and C. C. Baskin. 2003. The vascular flora of cedar glades of the southeastern United States and its phytogeographical relationships. Journal of the Torrey Botanical Society 130(2):101–118.

Cofer M.S, J.L. Walck and S.N. Hidayati, 2008. Species Richness and Exotic Species Invasion in Middle Tennessee Cedar Glades in Relation to Abiotic and Biotic Factors. The Journal of the Torrey Botanical Society, Vol. 135, No. 4, pp.540–553.

Duncan, R. S., C. B. Anderson, H. N. Sellers and E. E. Robbins. 2007. The Effect of Fire Reintroduction on Endemic and Rare Plants of a Southeastern Glade Ecosystem. Restoration Ecology, Vol. 16, No. 1, pp. 39–49.

Frost, C. 1998. Presettlement fire frequency regimes of the United States: A first approximation. In: Proceedings 20th Tall Timbers Fire Ecology Conference; Fire in Ecosystem Management: Shifting the Paradigm from Suppression to Prescription; 1996 May 7–10; Boise, ID. Tallahassee, FL: Tall Timbers Research Station: 70–81.

Govus, T. E., and R. D. White, Jr. 2006. Vascular plant inventory and plant community classification for Chickamauga and Chattanooga National Military Park. NatureServe Technical Report. Durham, North Carolina. 292 p.

Govus, T. E. and R. Lyons. 2009. Chickamauga and Chattanooga National Military Park Calcareous Glade Monitoring A Resampling of Baseline Transects Established in 1993. Durham, North Carolina: NatureServe.

Harper, R. M. 1926. The Cedar Glades of Middle Tennessee. Ecology, Vol. 7, No. 1, pp. 48–54.

Hogan, T. 2019. Personal communication. Terri Hogan, Invasive Plant Program Manager, Landscape Restoration & Adaptation, National Park Service, Fort Collins, Colorado.

Klein, R. 2019. Personal communication. Fire Ecologist. Appalachian/Piedmont/Coastal Fire Management Zone, Great Smoky Mountains National Park. Gatlinburg, Tennessee.

Mills, J. E. 2008. Fifty years of change in Wisconsin cedar glades. American Midland Naturalist, 150:214–224.

National Weather Service. 2020. NOWData - NOAA Online Weather Data. National Oceanic and Atmospheric Administration, National Weather Service Forecast Office, Morristown, TN. Available from (https://w2.weather.gov/climate/xmacis.php?wfo=mrx) (accessed January 2020).

NatureServe. 2020. NatureServe Explorer [web application]. NatureServe, Arlington, Virginia. Available https://explorer.natureserve.org/. (Accessed: July 02, 2020).

Patrick, T. 2008. Personal communication. Botanist, Nongame Conservation Section, Georgia Dept of Natural Resources. Social Circle, Georgia.

Quarterman, E. 1950. Ecology of Cedar Glades. I. Distribution of Glade Flora in Tennessee. Bulletin of the Torrey Botanical Club, Vol. 77, No. 1.

Quarterman, E. 1989. Structure and dynamics of the limestone cedar glade community of Tennessee. Journal Tennessee Academy of Science 64(3):155–158

Rutledge J., and T. Diggs. 2006. Cedar glade monitoring project progress report. Internal report to the National Park Service, Cumberland Piedmont Network, Mammoth Cave National Park, Mammoth Cave, Kentucky. 5 p.

Somers, P., L. Smith, P. Hamel, and E. L. Bridges. 1986. Preliminary analyses of plant communities and seasonal changes in cedar glades of Middle Tennessee. ASB Bulletin Vol. 33, No. 4.

Sutter, R. D., S. Benjamin, S. Rollins, G. Livingstone and N. Rudd. 1994. Baseline monitoring of calcareous glades at Chickamauga and Chattanooga National Military Park. Report to the National Park Service. The Nature Conservancy, Southeast Regional Office. Durham, North Carolina. 96 p.

Sutter, R. D., T. Govus, R. L. Smyth, C. Nordman, M. Pyne. 2011. Monitoring Change in a Central U.S. Calcareous Glade: Resampling Transects Established in 1993. Natural Areas Journal 31 (2):163–172.

Weakley, A. S. 2019. Flora of the Southern and Mid-Atlantic States. Working Draft of 9 May 2019. Unpublished manuscript. University of North Carolina Herbarium. Chapel Hill, North Carolina.

Weakley, A. S. 2019. Personal communication. Director of the UNC Herbarium (NCU) and Adjunct Associate Professor. North Carolina Botanical Garden, University of North Carolina, Chapel Hill, North Carolina. 30

Appendix A. Compiled Woody Cover Data for all Glades Sampled

List of Tables Page

Table A-1. Glade 2 ...... 32

Table A-2. Glade 3A ...... 33

Table A-3. Glade 3B ...... 34

Table A-4. Glade 6 ...... 35

Table A-5. Glade 8A ...... 36

Table A-6. Glade 8B ...... 37

Table A-7. Glade 11...... 38

Table A-8. Glade 13 ...... 38

Table A-9. Glade 14 ...... 39

Table A-10 Glade 21 ...... 41

Table A-11. Glade 24A ...... 42

Table A-1. Glade 2—Average percent cover values by woody species greater than 1 meter (3.3 ft)—1993–2019, N = 12 transects (Frequency, Glade Mean Percent Cover, Confidence Intervals [CI]).

Species 1993 1993 1993 2006 2006 2006 2019 2019 2019 Freq. % Cover 95% CI Freq. % Cover 95% CI Freq. % Cover 95% CI (mean) (mean) (mean)

Juniperus virginiana 12 22.3 16.6–27.9 11 34.9 24.8–44.7 12 58.91 51.6–65.9

Pinus taeda 10 15.1 9.1–21.6 10 21.1 12.3–33.0 12 33.57 24.1–45.2

Frangula caroliniana 4 0.55 0.13–1.08 6 5.02 1.32–9.95 7 8.04 3.7–12.5

Ulmus alata 7 2.29 0.76–4.19 6 3.77 1.46–6.42 12 7.61 4.9–10.4

Rhus aromatica 1 0.27 0.00–0.85 3 1.36 0.00–3.30 3 2.08 0.00–4.47

Cornus florida 0 0.00 0.00–0.00 1 0.04 0.00–0.14 7 1.90 0.79–3.29

Fraxinus americana 8 2.95 1.38–4.37 5 6.94 0.56–19.18 3 1.54 0.00–3.68

Ligustrum sinense 3 0.67 0.00–1.35 6 4.16 1.30-7.13 5 1.35 0.25–2.49

Rhus copallina 0 0.00 0.00–0.00 0 0.00 0.00–0.00 1 0.67 0.00–2.04

Pinus echinata 0 0.00 0.00–0.00 1 0.65 0.00–2.02 1 0.62 0.00–1.93

Quercus shumardii 0 0.00 0.00–0.00 0 0.00 0.00–0.00 1 0.59 0.00–1.77

Prunus serotina 0 0.00 0.00–0.00 0 0.00 0.00–0.00 1 0.25 0.00–0.84

Liquidambar styracifula 0 0.00 0.00–0.00 0 0.00 0.00–0.00 2 0.23 0.00–0.78

Celtus tenuifolia 0 0.00 0.00–0.00 0 0.00 0.00–0.00 1 0.07 0.00–0.25

Carya sp. 0 0.00 0.00–0.00 1 0.68 0.00–2.11 0 0.00 0.00–0.00

Pinus virginiana 2 0.61 0.00–1.53 0 0.00 0.00–0.00 0 0.00 0.00–0.00

Quercus velutina 0 0.00 0.00–0.00 1 0.19 0.00–0.57 0 0.00 0.00–0.00

Table A-2. Glade 3A—Average percent cover values by woody species greater than 1 meter (3.3 ft)—1993–2019, N = 9 transects (Frequency, Glade Mean Percent Cover, Confidence Intervals [CI]).

Species 1993 1993 1993 2006 2006 2006 2019 2019 2019 Freq. % Cover 95% CI Freq. % Cover 95% CI Freq. % Cover 95% CI (mean) (mean) (mean)

Juniperus virginiana 3 13.6 4.41–21.4 5 41.6 32.4–46.6 5 59.1 41.2–68.6

Pinus taeda 2 3.59 0.00–9.22 3 12.14 0.68–24.8 3 26.1 4.4–45.1

Frangula caroliniana 0 0.00 0.00–0.00 2 5.90 0.00–12.1 2 9.57 0.0–19.6

Pinus echinata 0 0.00 0.00–0.00 0 0.00 0.00–0.00 2 7.91 0.0–17.0

Fraxinus americana 0 0.00 0.00–0.00 3 2.62 0.23–6.01 3 5.51 0.75–10.3

Ulmus alata 0 0.00 0.00–0.00 2 5.48 0.00–11.34 1 4.26 0.00–9.28

Quercus stellata 2 2.97 0.00–10.5 2 2.35 0.00–9.40 2 2.55 0.00–9.03

Quercus phellos 0 0.00 0.00–0.00 0 0.00 0.00–0.00 1 1.98 0.00–5.28

Cornus florida 0 0.00 0.00–0.00 0 0.00 0.00–0.00 1 1.35 0.00–4.79

Rhus aromatica 2 0.25 0.00–0.51 3 2.40 0.84–4.12 1 0.65 0.00–1.73

Ligustrum sinense 1 0.05 0.00–0.27 0 0.00 0.00–0.00 0 0.00 0.00–0.00

Pinus virginiana 2 4.22 0.00–10.0 2 4.41 0.00–10.85 0 0.00 0.00–0.00

Table A-3. Glade 3B—Average percent cover values by woody species greater than 1 meter (3.3 ft)—1993–2019, N = 9 transects (Frequency, Glade Mean Percent Cover, Confidence Intervals [CI]).

Species 1993 1993 1993 2006 2006 2006 2019 2019 2019 Freq. % Cover 95% CI Freq. % Cover 95% CI Freq. % Cover 95% CI (mean) (mean) (mean)

Juniperus virginiana 9 48.6 36.7–59.3 9 58.9 48.0–69.7 9 65.12 50.1–78.4

Pinus virginiana 4 6.13 1.30–12.0 1 2.10 0.00–5.36 1 3.66 0.00–9.32

Ulmus alata 0 0.00 0.00–0.00 2 1.97 0.00–4.52 1 2.41 0.00–6.13

Ligustrum sinense 0 0.00 0.00–0.00 2 2.50 0.00–6.08 2 2.22 0.00–6.61

Cercis canadensis 1 0.20 0.00–0.65 2 1.00 0.00–2.66 1 1.94 0.00–6.20

Quercus stellata 0 0.00 0.00–0.00 2 2.00 0.00–5.35 2 1.37 0.00–3.93

Fraxinus americana 0 0.00 0.00–0.00 0 0.00 0.00–0.00 1 1.36 0.00–4.40

Rhus aromatica 0 0.00 0.00–0.00 2 0.68 0.00–1.85 2 0.98 0.00–2.46

Frangula caroliniana 0 0.00 0.00–0.00 5 3.62 0.96–7.97 3 0.65 0.00–1.12

Diospyros virginiana 0 0.00 0.00–0.00 1 0.67 0.00–1.70 1 0.03 0.00–0.12

Carya carolinae- 1 0.67 0.00–2.20 0 0.00 0.00–0.00 0 0.00 0.00–0.00 septentrionalis

Carya sp. 0 0.00 0.00–0.00 1 1.14 0.00–3.78 0 0.00 0.00–0.00

Celtus tenuifolia 1 0.55 0.00–1.40 1 2.10 0.00–6.68 0 0.00 0.00–0.00

Pinus taeda 4 5.58 0.37–12.86 1 2.56 0.00–8.47 0 0.00 0.00–0.00

Quercus phellos 1 0.34 0.00–1.08 0 0.00 0.00–0.00 0 0.00 0.00–0.00

Table A-4. Glade 6—Average percent cover values by woody species greater than 1 meter (3.3 ft)—1993–2019, N = 6 transects (Frequency, Glade Mean Percent Cover, Confidence Intervals [CI]).

Species 1993 1993 1993 2006 2006 2006 2019 2019 2019 Freq. % Cover 95% CI Freq. % Cover 95% CI Freq. % Cover 95% CI (mean) (mean) (mean)

Juniperus virginiana 6 24.1 9.57–42.2 5 19.6 6.73–35.7 6 55.94 37.0–78.4

Cercis canadensis 3 1.55 0.35–2.87 2 4.09 0.00–11.1 3 7.42 1.26–16.3

Quercus muehlenbergii 0 0.00 0.00–0.00 0 0.00 0.00–0.00 2 7.27 0.00–21.7

Quercus alba 1 1.19 0.00–3.33 1 2.66 0.00–7.45 1 3.35 0.00–9.37

Pinus virginiana 2 4.68 0.00–12.28 1 5.57 0.00–15.65 1 3.34 0.00–9.38

Quercus pagoda 0 0.00 0.00–0.00 0 0.00 0.00–0.00 1 3.15 0.00–9.83

Quercus stellata 2 1.55 0.00–3.30 0 0.00 0.00–0.00 2 2.68 0.00–5.45

Rhus aromatica 0 0.00 0.00–0.00 3 2.76 0.38–6.22 3 2.48 0.09–6.06

Fraxinus americana 0 0.00 0.00–0.00 0 0.00 0.00–0.00 1 1.36 0.00–4.26

Ligustrum quihoi 0 0.00 0.00–0.00 0 0.00 0.00–0.00 1 1.01 0.00–3.37

Frangula caroliniana 4 3.61 0.39–9.03 3 3.46 0.46–6.59 1 0.99 0.00–3.42

Quercus marilandica 0 0.00 0.00–0.00 0 0.00 0.00–0.00 1 0.84 0.00–2.88

Symphoricarpos orbiculatus 0 0.00 0.00–0.00 0 0.00 0.00–0.00 1 0.26 0.00–0.82

Celtus tenuifolia 1 0.79 0.00–2.47 0 0.00 0.00–0.00 0 0.00 0.00–0.00

Juglans nigra 2 2.09 0.00–4.39 2 2.21 0.00–5.17 0 0.00 0.00–0.00

Pinus taeda 2 2.17 0.00–5.10 0 0.00 0.00–0.00 0 0.00 0.00–0.00

Quercus shumardii 1 2.86 0.00–8.91 0 0.00 0.00–0.00 0 0.00 0.00–0.00

Ulmus alata 0 0.00 0.00–0.00 1 0.36 0.00–1.13 0 0.00 0.00–0.00

Table A-5. Glade 8A—Average percent cover values by woody species greater than 1 meter (3.3 ft)—1993–2019, N = 11 transects (Frequency, Glade Mean Percent Cover, Confidence Intervals [CI]).

Species 1993 1993 1993 2006 2006 2006 2019 2019 2019 Freq. % Cover 95% CI Freq. % Cover 95% CI Freq. % Cover 95% CI (mean) (mean) (mean)

Juniperus virginiana 9 18.7 9.63–27.9 10 25.1 15.2–34.69 10 49.6 40.9–57.6

Pinus taeda 11 20.3 13.3–31.5 10 28.1 17.9–40.78 11 46.4 37.3–57.1

Ligustrum sinense 5 1.59 0.35–3.31 6 8.62 3.19–15.68 8 9.28 4.32–14.8

Cercis canadensis 0 0.00 0.00–0.00 1 0.26 0.00–0.84 4 3.48 0.41–7.57

Fraxinus americana 1 0.51 0.00–1.61 2 1.10 0.00–3.13 3 2.52 0.00–6.38

Ulmus alata 0 0.00 0.00–0.00 2 1.58 0.00–4.23 6 2.33 0.80–4.13

Quercus falcata 2 0.38 0.00–0.95 0 0.00 0.00–0.00 3 2.27 0.00–5.43

Quercus phellos 1 0.17 0.00–0.50 1 0.61 0.00–1.82 2 1.89 0.00–4.24

Frangula caroliniana 1 0.09 0.00–0.26 1 0.42 0.00–1.27 3 1.79 0.00–3.93

Quercus marilandica 3 1.13 0.00–3.01 3 1.96 0.00–4.26 1 1.38 0.00–4.09

Pinus virginiana 2 0.66 0.00–1.92 1 1.70 0.00–5.96 2 1.15 0.00–3.77

Celtus tenuifolia 0 0.00 0.00-0.00 3 0.57 0.00–1.45 2 0.93 0.00–2.26

Quercus stellata 0 0.00 0.00–0.00 2 0.28 0.00–0.68 3 0.91 0.00–2.29

Rhus aromatica 0 0.00 0.00–0.00 2 1.20 0.00–3.07 1 0.51 0.00–1.56

Pinus echinata 0 0.00 0.00–0.00 1 0.85 0.00–2.68 2 0.30 0.00–0.76

Diospyros virginiana 0 0.00 0.00–0.00 1 0.05 0.00–0.15 1 0.22 0.00–0.69

Morus rubra 0 0.00 0.00–0.00 1 0.11 0.00–0.37 0 0.00 0.00–0.00

Prunus serotina 0 0.00 0.00–0.00 1 0.74 0.00–2.25 0 0.00 0.00–0.00

Table A-5 (continued). Glade 8A—Average percent cover values by woody species greater than 1 meter (3.3 ft)—1993–2019, N = 11 transects (Frequency, Glade Mean Percent Cover, Confidence Intervals [CI]).

Species 1993 1993 1993 2006 2006 2006 2019 2019 2019 Freq. % Cover 95% CI Freq. % Cover 95% CI Freq. % Cover 95% CI (mean) (mean) (mean)

Quercus velutina 0 0.00 0.00–0.00 2 1.88 0.00–5.13 0 0.00 0.00–0.00

Rubus arguta 0 0.00 0.00–0.00 1 0.22 0.00–0.67 0 0.00 0.00–0.00

Table A-6. Glade 8B—Average percent cover values by woody species greater than 1 meter (3.3 ft)—1993–2019, N = 10 transects (Frequency, Glade Mean Percent Cover, Confidence Intervals [CI]).

Species 1993 1993 1993 2008 2008 2008 2019 2019 2019 Freq. % Cover 95% CI Freq. % Cover 95% CI Freq. % Cover 95% CI (mean) (mean) (mean)

Juniperus virginiana 10 16.5 9.2–24.7 10 32.6 21.9–43.4 10 44.4 36.6–52.6

Pinus taeda 9 20.1 12.2–27.2 8 19.2 7.92–30.2 8 30.1 17.3–41.9

Ulmus alata 2 0.43 0.00–1.09 3 4.34 0.00–8.95 5 7.90 1.85–14.4

Ligustrum sinense 5 2.89 0.83–5.02 3 4.60 0.00–9.02 2 2.09 0.00–5.61

Quercus falcata 0 0.00 0.00–0.00 0 0.00 0.00–0.00 1 0.94 0.00–3.02

Quercus stellata 0 0.00 0.00–0.00 0 0.00 0.00–0.00 2 0.50 0.00–1.50

Frangula caroliniana 1 0.14 0.00–0.41 0 0.00 0.00–0.00 1 0.45 0.00–1.56

Fraxinus americana 0 0.00 0.00–0.00 1 0.45 0.00–1.55 3 0.36 0.00–0.82

Rhus aromatica 1 0.71 0.00–2.05 1 0.30 0.00–0.85 1 0.19 0.00–0.65

Celtus tenuifolia 1 0.57 0.00–1.64 2 0.99 0.00–2.39 0 0.00 0.00–0.00

Pinus virginiana 2 0.90 0.00–2.09 1 0.61 0.00–2.11 0 0.00 0.00–0.00

Table A-7. Glade 11—Average percent cover values by woody species greater than 1 meter (3.3 ft)—1993–2019, N = 5 transects (Frequency, Glade Mean Percent Cover, Confidence Intervals [CI]).

Species 1993 1993 1993 2006 2006 2006 2019 2019 2019 Freq. % Cover 95% CI Freq. % Cover 95% CI Freq. % Cover 95% CI (mean) (mean) (mean)

Juniperus virginiana 5 14.9 9.17–23.0 5 25.0 14.6–39.2 5 22.4 15.4–34.3

Ligustrum sinense 4 4.54 0.43–8.6 4 12.0 2.84–27.6 5 11.2 8.31–15.7

Pinus taeda 4 11.5 2.99–18.0 2 2.15 0.00–5.71 2 6.02 0.00–15.8

Quercus muehlenbergii 1 0.13 0.00–0.42 0 0.00 0.00–0.00 1 2.69 0.00–8.72

Ulmus alata 0 0.00 0.00–0.00 0 0.00 0.00–0.00 1 2.59 0.00–8.93

Fraxinus americana 0 0.00 0.00–0.00 0 0.00 0.00–0.00 3 2.18 0.71–3.71

Rhus aromatica 0 0.00 0.00–0.00 0 0.00 0.00–0.00 2 1.64 0.00–2.91

Diospyros virginiana 0 0.00 0.00–0.00 0 0.00 0.00–0.00 1 1.51 0.00–4.90

Quercus stellata 0 0.00 0.00–0.00 0 0.00 0.00–0.00 1 1.46 0.00–3.26

Pinus virginiana 2 5.01 0.00–11.0 0 0.00 0.00–0.00 1 1.44 0.00–5.02

Pinus echinata 0 0.00 0.00–0.00 1 1.64 0.00–3.65 1 0.94 0.00–2.09

Table A-8. Glade 13—Average percent cover values by woody species greater than 1 meter (3.3 ft)—1993–2019, N = 10 transects (Frequency, Glade Mean Percent Cover, Confidence Intervals [CI]).

Species 1993 1993 1993 2006 2006 2006 2019 2019 2019 Freq. % Cover 95% CI Freq. % Cover 95% CI Freq. % Cover 95% CI (mean) (mean) (mean)

Juniperus virginiana 9 14.5 9.28–20.2 10 28.5 21.6–34.6 10 39.1 29.6–46.5

Pinus taeda 6 4.12 1.93–6.40 8 6.90 3.27–11.6 7 9.68 5.31–14.5

Quercus muehlenbergii 2 1.95 0.00–5.36 5 4.36 1.24–7.50 5 5.93 2.14–9.31

Table A-8 (continued). Glade 13—Average percent cover values by woody species greater than 1 meter (3.3 ft)—1993–2019, N = 10 transects (Frequency, Glade Mean Percent Cover, Confidence Intervals [CI]).

Species 1993 1993 1993 2006 2006 2006 2019 2019 2019 Freq. % Cover 95% CI Freq. % Cover 95% CI Freq. % Cover 95% CI (mean) (mean) (mean)

Quercus stellata 2 2.18 0.00–6.71 4 3.11 0.06–8.45 5 3.72 0.49–8.63

Rhus aromatica 1 0.04 0.00–0.11 3 1.20 0.00–2.26 7 3.46 1.29–5.44

Frangula caroliniana 1 0.24 0.00–0.77 8 1.47 0.71–2.43 7 2.85 1.17–5.35

Cercis canadensis 1 0.05 0.00–0.16 2 1.35 0.00–3.30 4 2.07 0.42–3.64

Fraxinus americana 2 0.88 0.00–1.95 3 2.28 0.00–3.95 2 2.00 0.00–4.27

Juglans nigra 0 0.00 0.00–0.00 1 0.49 0.00–1.59 3 1.70 0.00–3.79

Diospyros virginiana 3 0.47 0.00–0.98 4 1.18 0.21–2.47 4 1.56 0.37–3.11

Pinus virginiana 6 2.74 0.93–4.88 4 1.18 0.18–2.65 2 1.39 0.00–3.72

Quercus nigra 1 0.83 0.00–2.62 1 1.13 0.00–3.58 1 0.94 0.00–2.98

Quercus shumardii 0 0.00 0.00–0.00 1 0.14 0.00–0.51 2 0.91 0.00–2.76

Ulmus alata 1 0.04 0.00–0.13 2 0.47 0.00–0.94 4 0.76 0.06–1.54

Sideroxylon lycioides 1 0.51 0.00–1.48 1 0.68 0.00–1.99 1 0.47 0.00–1.37

Carya carolinae- 0 0.00 0.00–0.00 0 0.00 0.00–0.00 1 0.28 0.00–0.81 septentrionalis

Morus rubra 0 0.00 0.00–0.00 0 0.00 0.00–0.00 1 0.09 0.00–0.31

Celtus tenuifolia 1 0.24 0.00–0.67 1 0.37 0.00–1.03 2 0.06 0.00–0.14

Carya glabra 1 0.12 0.00–0.39 0 0.00 0.00–0.00 0 0.00 0.00–0.00

Carya pallida 1 0.10 0.00–0.27 1 0.29 0.00–0.81 0 0.00 0.00–0.00

Carya sp. 0 0.00 0.00–0.00 1 0.18 0.00–0.56 0 0.00 0.00–0.00

Pinus echinata 0 0.00 0.00–0.00 1 0.02 0.00–0.05 0 0.00 0.00–0.00

Table A-9. Glade 14—Average percent cover values by woody species greater than 1 meter (3.3 ft)—1993–2019, N = 10 transects (Frequency, Glade Mean Percent Cover, Confidence Intervals [CI]).

Species 1993 1993 1993 2006 2006 2006 2019 2019 2019 Freq. % Cover 95% CI Freq. % Cover 95% CI Freq. % Cover 95% CI (mean) (mean) (mean)

Juniperus virginiana 7 10.1 3.85–17.6 10 22.6 14.2–30.9 10 31.8 24.1–41.5

Pinus taeda 5 6.36 0.84–16.4 7 12.1 3.59–25.9 8 12.3 4.88–24.5

Cercis canadensis 1 0.03 0.00–0.11 3 4.94 0.00–15.0 3 4.68 0.00–12.6

Quercus stellata 1 0.42 0.00–1.61 4 2.08 0.22–5.06 4 3.14 0.79–6.10

Diospyros virginiana 4 0.58 0.07–1.32 5 1.94 0.53–3.86 5 3.00 1.15–5.03

Rhus aromatica 0 0.00 0.00–0.00 6 4.26 1.31–8.11 6 2.78 1.56–4.04

Quercus muehlenbergii 0 0.00 0.00–0.00 2 0.38 0.00–0.92 3 1.95 0.00–4.89

Frangula caroliniana 2 0.22 0.00–0.52 4 2.81 0.56–4.96 4 1.94 0.61–3.14

Fraxinus americana 1 0.10 0.00–0.40 2 0.37 0.00–1.11 5 1.78 0.41–3.39

Quercus shumardii 1 0.25 0.00–0.84 0 0.00 0.00–0.00 2 0.95 0.00–2.61

Ulmus alata 0 0.00 0.00–0.00 0 0.00 0.00–0.00 1 0.24 0.00–0.72

Pinus virginiana 1 0.10 0.00–0.30 0 0.00 0.00–0.00 0 0.00 0.00–0.00

Quercus alba 1 0.19 0.00–0.59 0 0.00 0.00–0.00 0 0.00 0.00–0.00

Quercus marilandica 1 0.14 0.00–0.40 0 0.00 0.00–0.00 0 0.00 0.00–0.00

Quercus rubra 0 0.00 0.00–0.00 1 0.47 0.00–1.61 0 0.00 0.00–0.00

Table A-10 Glade 21—Average percent cover values by woody species greater than 1 meter (3.3 ft)—1993–2019, N = 5 transects (Frequency, Glade Mean Percent Cover, Confidence Intervals [CI]).

Species 1993 1993 1993 2008 2008 2008 2019 2019 2019 Freq. % Cover 95% CI Freq. % Cover 95% CI Freq. % Cover 95% CI (mean) (mean) (mean)

Juniperus virginiana 4 12.7 3.46–22.5 5 36.8 17.6–58.5 5 57.7 38.8–78.1

Cercis canadensis 1 2.73 0.00–6.97 2 8.75 0.00–19.1 1 7.12 0.00–18.2

Frangula caroliniana 0 0.00 0.00–0.00 0 0.00 0.00–0.00 2 4.55 0.00–7.42

Chionanthus virginicus 0 0.00 0.00–0.00 0 0.00 0.00–0.00 1 3.33 0.00–9.85

Quercus stellata 0 0.00 0.00–0.00 1 0.55 0.00–1.45 2 2.58 0.00–7.15

Ligustrum sinense 0 0.00 0.00–0.00 1 1.59 0.00–4.20 2 2.49 0.00–5.73

Quercus marilandica 0 0.00 0.00–0.00 1 3.24 0.00–8.59 1 1.72 0.00–4.56

Celtus tenuifolia 0 0.00 0.00–0.00 1 0.77 0.00–2.28 1 0.84 0.00–2.50

Diospyros virginiana 0 0.00 0.00–0.00 0 0.00 0.00–0.00 1 0.72 0.00–3.00

Quercus falcata 0 0.00 0.00–0.00 0 0.00 0.00–0.00 1 0.42 0.00–1.07

Amelanchier arborea 0 0.00 0.00–0.00 1 0.10 0.00–0.31 0 0.00 0.00–0.00

Pinus taeda 3 8.42 0.41–13.28 0 0.00 0.00–0.00 0 0.00 0.00–0.00

Pinus virginiana 1 0.71 0.00–1.81 0 0.00 0.00–0.00 0 0.00 0.00–0.00

Quercus shumardii 1 0.43 0.00–1.15 0 0.00 0.00–0.00 0 0.00 0.00–0.00

Rhus aromatica 0 0.00 0.00–0.00 2 5.20 0.00–11.23 0 0.00 0.00–0.00

Table A-11. Glade 24A—Average percent cover values by woody species greater than 1 meter (3.3 ft)—1993–2019, N = 8 transects (Frequency, Glade Mean Percent Cover, Confidence Intervals [CI]).

Species 1993 1993 1993 2006 2006 2006 2019 2019 2019 Freq. % Cover 95% CI Freq. % Cover 95% CI Freq. % Cover 95% CI (mean) (mean) (mean)

Juniperus virginiana 8 32.9 26.3–41.6 8 27.9 16.8–39.8 8 36.4 29.4–44.5

Ligustrum sinense 6 3.42 1.56–5.61 8 12.83 7.22–20.3 8 8.98 4.28–15.3

Rhus aromatica 2 0.33 0.00–0.88 7 5.91 2.13–10.2 7 7.42 3.73–11.3

Fraxinus americana 2 1.08 0.00–2.42 2 2.07 0.00–4.50 4 3.02 0.39–6.20

Rubus occidentalis 0 0.00 0.00–0.00 1 0.25 0.00–0.85 1 0.98 0.00–3.29

Quercus stellata 0 0.00 0.00–0.00 2 0.71 0.00–2.10 3 0.96 0.00–1.87

Celtus tenuifolia 2 0.88 0.00–2.38 2 1.42 0.00–3.48 1 0.76 0.00–2.43

Quercus muehlenbergii 1 0.03 0.00–0.09 0 0.00 0.00–0.00 2 0.66 0.00–1.46

Juglans nigra 1 0.18 0.00–0.59 1 0.12 0.00–0.34 1 0.34 0.00–1.13

Ailanthus altissima 0 0.00 0.00–0.00 0 0.00 0.00–0.00 1 0.23 0.00–0.73

Cercis canadensis 1 0.95 0.00–3.17 2 2.32 0.00–6.98 1 0.04 0.00–0.15

Carya sp. 0 0.00 0.00–0.00 1 0.18 0.00–0.59 0 0.00 0.00–0.00

Pinus echinata 0 0.00 0.00–0.00 1 0.96 0.00–3.20 0 0.00 0.00–0.00

Pinus taeda 5 3.11 1.16–5.37 0 0.00 0.00–0.00 0 0.00 0.00–0.00

Pinus virginiana 1 0.70 0.00–2.33 0 0.00 0.00–0.00 0 0.00 0.00–0.00

Rhus coppalina 1 0.04 0.00–0.12 0 0.00 0.00–0.00 0 0.00 0.00–0.00

Rhus glabra 0 0.00 0.00–0.00 1 0.43 0.00–1.38 0 0.00 0.00–0.00

Rubus arguta 0 0.00 0.00–0.00 1 0.70 0.00–2.23 0 0.00 0.00–0.00

Table A-11 (continued). Glade 24A—Average percent cover values by woody species greater than 1 meter (3.3 ft)—1993–2019, N = 8 transects (Frequency, Glade Mean Percent Cover, Confidence Intervals [CI]).

Species 1993 1993 1993 2006 2006 2006 2019 2019 2019 Freq. % Cover 95% CI Freq. % Cover 95% CI Freq. % Cover 95% CI (mean) (mean) (mean)

Smilax rotundifolia 0 0.00 0.00–0.00 1 0.06 0.00–0.19 0 0.00 0.00–0.00

Ulmus alata 1 0.67 0.00–2.16 2 2.18 0.00–4.83 2 1.74 0.00–3.80

Appendix B. Compiled Herbaceous Plant Density Data for All Glades Sampled

List of Tables Page

Table B-1. Plant density measurements for glade 3B, 1993–2019...... 46

Table B-2. Plant density measurements for glade 8B, 1993–2019...... 47

Table B-3. Plant density measurements for glade 11, 1993–2019...... 48

Table B-4. Plant density measurements for glade 14, 1993–2019...... 49

Table B-5. Plant density measurements for glade 21, 1993–2019 ...... 50

Table B-1. Average herbaceous plant density measurements for glade 3B, 1993–2019. (Frequency, Glade Mean Density [m2—square meters], 95% Confidence Intervals [CI]).

Species 1993 1993 1993 1993 2008 2008 2008 2008 2019 2019 2019 2019 Freq. Sum Density 95% CI Freq. Sum Density 95% CI Freq. Sum Density 95% CI (plants/m2) (plants/m2) (plants/m2)

Hypericum dolabriforme 9 159 1.08 0.64–1.66 8 82 0.56 0.24–0.95 9 580 3.94 2.91–4.89

Packera paupercula 9 112 0.76 0.55–1 4 8 0.05 0.01–0.12 9 201 1.36 0.82–2.1

Delphinium carolinianum 7 284 1.93 0.5–4.31 0 NA 0 0–0 7 112 0.76 0.16–1.57 var. calciphilum

Dalea gattingeri 8 332 2.25 1.04–3.79 6 135 0.92 0.11–2.07 6 110 0.75 0.13–1.65

Scutellaria leonardii 7 116 0.79 0.32–1.41 3 7 0.05 0–0.1 7 100 0.68 0.34–1.07

Ophioglossum 1 324 2.2 0–7.38 1 42 0.29 0–0.96 3 73 0.5 0–1.45 engelmannii

Ruellia humilis 6 122 0.83 0.16–1.69 4 92 0.62 0.04–1.74 3 55 0.37 0–1.09

Viola egglestonii 7 135 0.92 0.11–2.07 5 12 0.08 0.02–0.17 7 36 0.24 0.11–0.38

Manfreda virginica 6 25 0.17 0.06–0.32 0 NA 0 0–0 5 16 0.11 0.03–0.21

Lithospermum canescens 6 25 0.17 0.07–0.3 0 NA 0 0–0 2 8 0.05 0–0.14

Ratibida pinnata 7 160 1.09 0.46–1.75 5 12 0.08 0.02–0.15 1 1 0.01 0–0.02

Leavenworthia exigua 3 44 0.3 0–0.76 0 NA 0 0–0 0 NA 0 0–0 var. exigua

Table B-2. Average herbaceous plant density measurements for glade 8B, 1993–2019. (Frequency, Glade Mean Density [m2—square meters], 95% Confidence Intervals [CI]).

Species 1993 1993 1993 1993 2008 2008 2008 2008 2019 2019 2019 2019 Freq. Sum Density 95% CI Freq. Sum Density 95% CI Freq. Sum Density 95% CI (plants/m2) (plants/m2) (plants/m2)

Packera paupercula 9 107 0.51 0.3–0.75 5 17 0.08 0.02–0.16 9 1924 9.12 5.21–13.53

Ruellia humilis 10 339 1.61 1.05–2.28 9 56 0.27 0.17–0.38 10 1356 6.43 3.53–9.16

Hypericum dolabriforme 10 168 0.8 0.33–1.38 9 405 1.92 0.8–2.97 10 1005 4.76 2.97–6.74

Pediomelum subacaule 10 2506 11.88 8.19–16.57 10 1220 5.78 3.94–7.97 10 963 4.57 2.67–6.7

Dalea gattingeri 10 371 1.76 1.31–2.29 10 125 0.59 0.41–0.8 9 188 0.89 0.43–1.48

Viola egglestonii 9 620 2.94 1.37–4.4 8 53 0.25 0.11–0.42 9 118 0.56 0.31–0.87

Scutellaria leonardii 3 25 0.12 0–0.26 0 NA 0 0–0 6 100 0.47 0.03–0.89

Manfreda virginica 10 118 0.56 0.29–0.94 0 NA 0 0–0 7 36 0.17 0.07–0.32

Lithospermum canescens 3 5 0.02 0–0.06 0 NA 0 0–0 1 2 0.01 0–0.03

Delphinium carolinianum 1 5 0.02 0–0.08 0 NA 0 0–0 0 NA 0 0–0 var. calciphilum

Silphium pinnatifidum 8 151 0.72 0.31–1.05 7 16 0.08 0.03–0.13 0 NA 0 0–0

Table B-3. Average herbaceous plant density measurements for glade 11, 1993–2019. (Frequency, Glade Mean Density [m2—square meters], 95% Confidence Intervals [CI]).

Species 1993 1993 1993 1993 2008 2008 2008 2008 2019 2019 2019 2019 Freq. Sum Density 95% CI Freq. Sum Density 95% CI Freq. Sum Density 95% CI (plants/m2) (plants/m2) (plants/m2)

Hypericum dolabriforme 5 226 2.64 1.93–3.61 2 16 0.19 0–0.4 5 927 10.85 4.89–17.94

Ruellia humilis 5 281 3.29 0.74–5.69 5 325 3.8 2.03–4.97 5 659 7.71 3.83–10.47

Pediomelum subacaule 5 886 10.37 5.36–14.04 5 555 6.49 4.37–8.14 5 371 4.34 1.94–6.68

Ophioglossum 4 123 1.44 0.23–2.73 4 40 0.47 0.12–0.8 4 70 0.82 0.16–1.43 engelmannii

Dalea gattingeri 5 296 3.46 1.47–6.41 4 65 0.76 0.21–1.53 4 58 0.68 0.37–1.08

Scutellaria leonardii 4 37 0.43 0.16–0.88 0 NA 0 0–0 3 53 0.62 0.03–1.26

Viola egglestonii 5 339 3.97 0.46–7.73 4 28 0.33 0.11–0.52 5 53 0.62 0.2–0.93

Mononeuria patula 1 54 0.63 0–2.05 2 9 0.11 0–0.3 2 39 0.46 0–1.22

Packera paupercula 5 69 0.81 0.41–1.19 4 28 0.33 0.09–0.55 2 33 0.39 0–0.85

Leavenworthia exigua 2 56 0.66 0–1.15 3 55 0.64 0.17–0.97 2 9 0.11 0–0.2 var. exigua

Manfreda virginica 3 29 0.34 0.04–0.71 0 NA 0 0–0 3 7 0.08 0.01–0.22

Lithospermum canescens 1 1 0.01 0–0.04 0 NA 0 0–0 0 NA 0 0–0

Table B-4. Average herbaceous plant density measurements for glade 14, 1993–2019. (Frequency, Glade Mean Density [m2—square meters], 95% Confidence Intervals [CI]).

Species 1993 1993 1993 1993 2008 2008 2008 2008 2019 2019 2019 2019 Freq. Sum Density 95% CI Freq. Sum Density 95% CI Freq. Sum Density 95% CI (plants/m²) (plants/m²) (plants/m²)

Pediomelum subacaule 10 4404 15.3 10.2–19.7 9 2329 8.11 5.6–10.9 9 2063 7.19 4.48–9.2

Packera paupercula 9 353 1.23 0.73–1.88 8 606 2.11 0.65–3.31 10 1967 6.85 4.86–8.34

Hypericum dolabriforme 10 514 1.79 1.4–2.1 8 466 1.62 1.09–2.22 10 805 2.8 1.85–4.11

Dalea gattingeri 7 734 2.56 1.75–3.16 6 318 1.11 0.55–1.6 6 532 1.85 0.83–2.68

Ruellia humilis 7 797 2.78 1.06–4.4 9 301 1.05 0.41–1.62 7 202 0.7 0.31–1.06

Scutellaria leonardii 6 85 0.3 0.08–0.68 3 5 0.02 0–0.05 9 189 0.66 0.29–1.24

Ratibida pinnata 1 27 0.09 0–0.28 2 61 0.21 0–0.75 5 163 0.57 0.05–1.61

Lithospermum canescens 7 33 0.11 0.04–0.25 0 NA 0 0–0 10 107 0.37 0.23–0.58

Viola egglestonii 10 1103 3.84 1.74–7.19 8 58 0.2 0.08–0.36 7 61 0.21 0.09–0.35

Manfreda virginica 4 8 0.03 0.01–0.05 0 NA 0 0–0 5 9 0.03 0.01–0.05

Asclepias viridis 0 NA 0 0–0 0 NA 0 0–0 1 5 0.02 0–0.05

Leavenworthia exigua 1 1 0 0–0.01 0 NA 0 0–0 0 NA 0 0–0 var. exigua

Ophioglossum 1 4 0.01 0–0.04 0 NA 0 0–0 0 NA 0 0–0 engelmannii

Silphium pinnatifidum 6 164 0.57 0.26–0.86 0 NA 0 0–0 0 NA 0 0–0

Table B-5. Average herbaceous plant density measurements for glade 21, 1993–2019. (Frequency, Glade Mean Density [m2—square meters], 95% Confidence Intervals [CI]).

Species 1993 1993 1993 1993 2008 2008 2008 2008 2019 2019 2019 2019 Freq. Sum Density 95% CI Freq. Sum Density 95% CI Freq. Sum Density 95% CI (plants/m²) (plants/m²) (plants/m²)

Hypericum dolabriforme 5 340 3.5 2.22–4.6 3 64 0.66 0.08–1.16 5 1442 14.8 9.89–22.5

Scutellaria leonardii 5 120 1.24 0.42–1.88 0 NA 0 0–0 5 118 1.22 0.57–1.75

Pediomelum subacaule 3 88 0.91 0.03–1.75 3 106 1.09 0.28–1.76 3 110 1.13 0.41–1.62

Dalea gattingeri 5 385 3.96 3.04–4.9 5 175 1.8 1.25–2.68 4 70 0.72 0.14–1.46

Ruellia humilis 5 534 5.5 4.77–6.86 5 77 0.79 0.44–1.17 4 62 0.64 0.05–1.52

Viola egglestonii 4 148 1.52 0.53–2.17 2 18 0.19 0–0.38 2 16 0.16 0–0.39

Ratibida pinnata 4 191 1.97 0.19–5.64 0 NA 0 0–0 2 11 0.11 0–0.26

Manfreda virginica 5 56 0.58 0.28–0.95 0 NA 0 0–0 3 4 0.04 0.01–0.11

Leavenworthia exigua 2 25 0.26 0–0.47 0 NA 0 0–0 1 2 0.02 0–0.05 var. exigua

Packera paupercula 1 2 0.02 0–0.06 0 NA 0 0–0 1 1 0.01 0–0.03

Delphinium carolinianum 2 5 0.05 0–0.13 0 NA 0 0–0 0 NA 0 0–0 var. calciphilum

Silphium pinnatifidum 2 3 0.03 0–0.08 0 NA 0 0–0 0 NA 0 0–0

Appendix C. Compiled Small Woody Plant Density Data for All Glades Sampled (Woody Plants less than 0.5 meters)

List of Tables Page

Table C-1. Small woody plant density for glade 3B, 1993–2019...... 52

Table C-2. Small woody plant density for glade 8B, 1993–2019...... 53

Table C-3. Small woody plant density for glade 11, 1993–2019 ...... 55

Table C-4. Small woody plant density for glade 14, 1993–2019 ...... 56

Table C-5. Small woody plant density for glade 21, 1993–2019 ...... 57

Table C-1. Average small woody plant (< 0.5 meter [1.6 ft]) density measurements for glade 3B, 1993–2019. (Frequency, Glade Mean Density [m2—square meters], 95% Confidence Intervals [CI]).

Species 1993 1993 1993 1993 2008 2008 2008 2008 2019 2019 2019 2019 Freq. Sum Density 95% CI Freq. Sum Density 95% CI Freq. Sum Density 95% CI (plants/m²) (plants/m²) (plants/m²)

Fraxinus americana 0 NA 0 0–0 9 90 0.61 0.33–0.95 9 137 0.93 0.52–1.45

Celtus tenuifolia 8 29 0.2 0.1–0.3 8 48 0.33 0.17–0.47 7 69 0.47 0.16–0.95

Ulmus alata 0 NA 0 0–0 9 61 0.41 0.25–0.58 8 69 0.47 0.27–0.7

Juniperus virginiana 0 NA 0 0–0 8 48 0.33 0.19–0.47 9 68 0.46 0.28–0.67

Ligustrum sinense 2 18 0.12 0–0.3 6 29 0.2 0.08–0.35 9 66 0.45 0.32–0.59

Symphoricarpos 3 22 0.15 0–0.4 7 36 0.24 0.09–0.39 8 50 0.34 0.13–0.56 orbiculatus

Cercis canadensis 5 13 0.09 0–0.2 5 10 0.07 0.02–0.12 7 18 0.12 0.05–0.23

Quercus stellata 0 NA 0 0–0 5 10 0.07 0.02–0.12 5 11 0.07 0.02–0.13

Quercus muehlenbergii 0 NA 0 0–0 2 2 0.01 0–0.03 3 4 0.03 0–0.07

Quercus phellos 0 NA 0 0–0 0 NA 0 0–0 2 3 0.02 0–0.05

Carya carolinae- 0 NA 0 0–0 0 NA 0 0–0 1 1 0.01 0–0.02 septentrionalis

Cornus florida 0 NA 0 0–0 0 NA 0 0–0 1 2 0.01 0–0.04

Frangula caroliniana 6 13 0.09 0–0.1 3 7 0.05 0.01–0.09 1 1 0.01 0–0.02

Pinus virginiana 0 NA 0 0–0 0 NA 0 0–0 1 1 0.01 0–0.02

Prunus serotina 0 NA 0 0–0 2 3 0.02 0–0.04 1 1 0.01 0–0.02

Quercus marilandica 0 NA 0 0–0 0 NA 0 0–0 1 1 0.01 0–0.02

Rhus aromatica 9 59 0.4 0.3–0.5 8 18 0.12 0.08–0.17 2 2 0.01 0–0.03

Acer rubrum 0 NA 0 0–0 0 NA 0 0–0 2 2 0.01 0–0.04

Table C-1 (continued). Average small woody plant (< 0.5 meter [1.6 ft]) density measurements for glade 3B, 1993–2019. (Frequency, Glade Mean Density [m2—square meters], 95% Confidence Intervals [CI]).

Species 1993 1993 1993 1993 2008 2008 2008 2008 2019 2019 2019 2019 Freq. Sum Density 95% CI Freq. Sum Density 95% CI Freq. Sum Density 95% CI (plants/m²) (plants/m²) (plants/m²)

Rosa multiflora 2 7 0.05 0–0.1 0 NA 0 0–0 1 2 0.01 0–0.04

Ulmus rubra 9 40 0.27 0.2–0.3 0 NA 0 0–0 1 1 0.01 0–0.02

Carya sp. 0 NA 0 0–0 5 9 0.06 0.02–0.13 0 NA 0 0–0

Pinus sp. 0 NA 0 0–0 2 3 0.02 0–0.05 0 NA 0 0–0

Quercus alba 0 NA 0 0–0 1 1 0.01 0–0.02 0 NA 0 0–0

Quercus sp. 0 NA 0 0–0 2 3 0.02 0–0.05 0 NA 0 0–0

Rubus sp. 0 NA 0 0–0 1 1 0.01 0–0.02 0 NA 0 0–0

Acer floridanum 0 NA 0 0–0 4 7 0.05 0.01–0.08 0 NA 0 0–0

Albizia julibrissin 0 NA 0 0–0 5 16 0.11 0.02–0.21 0 NA 0 0–0

Table C-2. Average small woody (< 0.5 meter [1.6 ft]) plant density measurements for glade 8B, 1993–2019. (Frequency, Glade Mean Density [m2—square meters], 95% Confidence Intervals [CI]).

Species 1993 1993 1993 1993 2008 2008 2008 2008 2019 2019 2019 2019 Freq. Sum Density 95% CI Freq. Sum Density 95% CI Freq. Sum Density 95% CI (plants/m²) (plants/m²) (plants/m²)

Fraxinus americana 0 NA 0 0–0 10 66 0.31 0.22–0.42 10 373 1.77 1.25–2.4

Ligustrum sinense 10 293 1.39 0.6–2.1 8 94 0.45 0.21–0.67 10 239 1.13 0.87–1.38

Ulmus alata 10 168 0.8 0.5–1 8 67 0.32 0.16–0.51 10 102 0.48 0.29–0.76

Celtus tenuifolia 6 57 0.27 0–0.6 9 67 0.32 0.09–0.65 10 91 0.43 0.28–0.59

Table C-2 (continued). Average small woody (< 0.5 meter [1.6 ft]) plant density measurements for glade 8B, 1993–2019. (Frequency, Glade Mean Density [m2—square meters], 95% Confidence Intervals [CI]).

Species 1993 1993 1993 1993 2008 2008 2008 2008 2019 2019 2019 2019 Freq. Sum Density 95% CI Freq. Sum Density 95% CI Freq. Sum Density 95% CI (plants/m²) (plants/m²) (plants/m²)

Symphoricarpos 4 13 0.06 0–0.1 4 17 0.08 0.01–0.19 6 54 0.26 0.04–0.66 orbiculatus

Quercus stellata 0 NA 0 0–0 0 NA 0 0–0 6 40 0.19 0.04–0.41

Juniperus virginiana 10 53 0.25 0.2–0.4 7 29 0.14 0.08–0.19 9 38 0.18 0.12–0.26

Quercus falcata 0 NA 0 0–0 0 NA 0 0–0 5 22 0.1 0.01–0.25

Frangula caroliniana 1 1 0 0–0 2 2 0.01 0–0.02 3 9 0.04 0–0.11

Rhus aromatica 1 1 0 0–0 3 3 0.01 0–0.03 3 9 0.04 0–0.1

Acer rubrum 0 NA 0 0–0 0 NA 0 0–0 6 9 0.04 0.02–0.06

Quercus phellos 0 NA 0 0–0 0 NA 0 0–0 3 6 0.03 0–0.06

Prunus serotina 0 NA 0 0–0 0 NA 0 0–0 4 4 0.02 0.01–0.03

Quercus muehlenbergii 0 NA 0 0–0 0 NA 0 0–0 3 4 0.02 0–0.04

Cornus florida 0 NA 0 0–0 1 1 0 0–0.02 0 NA 0 0–0

Diospyros virginiana 0 NA 0 0–0 3 4 0.02 0–0.04 0 NA 0 0–0

Pinus sp. 10 249 1.18 0.8–1.7 0 NA 0 0–0 0 NA 0 0–0

Quercus sp. 0 NA 0 0–0 5 21 0.1 0.02–0.23 0 NA 0 0–0

Rhus coppalina 0 NA 0 0–0 1 2 0.01 0–0.03 0 NA 0 0–0

Acer floridanum 0 NA 0 0–0 1 1 0 0–0.01 0 NA 0 0–0

Table C-3. Average small woody (< 0.5 meter [1.6 ft]) plant density measurements for glade 11, 1993–2019. (Frequency, Glade Mean Density [m2—square meters], 95% Confidence Intervals [CI]).

Species 1993 1993 1993 1993 2008 2008 2008 2008 2019 2019 2019 2019 Freq. Sum Density 95% CI Freq. Sum Density 95% CI Freq. Sum Density 95% CI (plants/m²) (plants/m²) (plants/m²)

Ligustrum sinense 5 72 0.84 0.2–1.8 4 45 0.53 0.26–0.86 5 60 0.70 0.34–1.4

Ulmus alata 5 123 1.44 0.9–1.8 4 53 0.62 0.22–1.1 3 23 0.27 0.02–0.74

Fraxinus americana 0 NA 0 0–0 5 14 0.16 0.1–0.26 5 15 0.18 0.1–0.3

Juniperus virginiana 5 26 0.3 0.2–0.5 2 12 0.14 0–0.23 4 13 0.15 0.05–0.25

Celtus tenuifolia 0 NA 0 0–0 3 4 0.05 0.01–0.1 3 8 0.09 0.01–0.29

Symphoricarpos 0 NA 0 0–0 1 4 0.05 0–0.1 1 3 0.04 0–0.08 orbiculatus

Cercis canadensis 0 NA 0 0–0 0 NA 0 0–0 1 1 0.01 0–0.05

Quercus stellata 0 NA 0 0–0 2 32 0.37 0–0.76 1 1 0.01 0–0.05

Diospyros virginiana 0 NA 0 0–0 2 2 0.02 0–0.05 0 NA 0 0–0

Pinus sp. 5 94 1.1 0.6–1.8 0 NA 0 0–0 0 NA 0 0–0

Quercus muehlenbergii 0 NA 0 0–0 1 1 0.01 0–0.05 0 NA 0 0–0

Quercus sp. 0 NA 0 0–0 1 1 0.01 0–0.04 0 NA 0 0–0

Rosa multiflora 1 1 0.01 0–0 0 NA 0 0–0 0 NA 0 0–0

Table C-4. Average small woody (< 0.5 meter [1.6 ft]) plant density measurements for glade 14, 1993–2019. (Frequency, Glade Mean Density [m2—square meters], 95% Confidence Intervals [CI]).

Species 1993 1993 1993 1993 2008 2008 2008 2008 2019 2019 2019 2019 Freq. Sum Density 95% CI Freq. Sum Density 95% CI Freq. Sum Density 95% CI (plants/m²) (plants/m²) (plants/m²)

Fraxinus americana 9 176 0.61 0.4–0.9 10 231 0.8 0.48–1.17 10 518 1.8 0.93–2.85

Juniperus virginiana 10 339 1.18 0.6–1.8 10 63 0.22 0.14–0.3 10 127 0.44 0.23–0.74

Cercis canadensis 9 99 0.34 0.1–0.9 6 25 0.09 0.02–0.19 10 115 0.4 0.25–0.65

Ulmus alata 7 71 0.25 0.2–0.3 9 61 0.21 0.12–0.33 10 105 0.37 0.2–0.59

Frangula caroliniana 4 18 0.06 0–0.1 5 20 0.07 0.01–0.15 7 96 0.33 0.05–0.66

Celtus tenuifolia 5 24 0.08 0–0.2 7 19 0.07 0.03–0.11 7 65 0.23 0.08–0.39

Rhus aromatica 7 98 0.34 0.1–0.6 4 22 0.08 0.01–0.23 8 63 0.22 0.11–0.39

Quercus stellata 0 NA 0 0–0 1 11 0.04 0–0.13 7 45 0.16 0.07–0.25

Quercus muehlenbergii 0 NA 0 0–0 1 1 0 0–0.01 6 39 0.14 0.02–0.43

Pinus taeda 0 NA 0 0–0 0 NA 0 0–0 4 35 0.12 0.01–0.22

Rosa multiflora 2 37 0.13 0–0.3 0 NA 0 0–0 4 31 0.11 0.02–0.23

Symphoricarpos 0 NA 0 0–0 1 9 0.03 0–0.11 4 21 0.07 0.01–0.2 orbiculatus

Diospyros virginiana 0 NA 0 0–0 6 12 0.04 0.02–0.07 5 17 0.06 0.02–0.1

Ligustrum sinense 4 15 0.05 0–0.1 3 6 0.02 0–0.04 7 17 0.06 0.03–0.09

Carya carolinae- 0 NA 0 0–0 0 NA 0 0–0 1 2 0.01 0–0.03 septentrionalis

Carya sp. 0 NA 0 0–0 2 2 0.01 0–0.02 0 NA 0 0–0

Pinus sp. 10 537 1.87 1.3–2.4 0 NA 0 0–0 0 NA 0 0–0

Table C-4 (continued). Average small woody (< 0.5 meter [1.6 ft]) plant density measurements for glade 14, 1993–2019. (Frequency, Glade Mean Density [m2—square meters], 95% Confidence Intervals [CI]).

Species 1993 1993 1993 1993 2008 2008 2008 2008 2019 2019 2019 2019 Freq. Sum Density 95% CI Freq. Sum Density 95% CI Freq. Sum Density 95% CI (plants/m²) (plants/m²) (plants/m²)

Prunus serotina 0 NA 0 0–0 2 3 0.01 0–0.03 0 NA 0 0–0

Quercus nigra 0 NA 0 0–0 1 1 0 0–0.01 0 NA 0 0–0

Quercus sp. 0 NA 0 0–0 4 5 0.02 0–0.04 0 NA 0 0–0

Table C-5. Average small woody (< 0.5 meter [1.6 ft]) plant density measurements for glade 21, 1993–2019. (Frequency, Glade Mean Density [m2—square meters], 95% Confidence Intervals [CI]).

Species 1993 1993 1993 1993 2008 2008 2008 2008 2019 2019 2019 2019 Freq. Sum Density 95% CI Freq. Sum Density 95% CI Freq. Sum Density 95% CI (plants/m²) (plants/m²) (plants/m²)

Ulmus alata 4 14 0.14 0.1–0.3 5 96 0.99 0.73–1.25 5 56 0.58 0.38–0.84

Fraxinus americana 0 NA 0 0–0 4 12 0.12 0.05–0.21 5 55 0.57 0.32–1.04

Ligustrum sinense 4 6 0.06 0–0.1 3 12 0.12 0.01–0.22 5 55 0.57 0.32–0.96

Cercis canadensis 5 210 2.16 1.2–3.1 5 43 0.44 0.26–0.54 5 53 0.55 0.32–0.82

Prunus serotina 0 NA 0 0–0 2 4 0.04 0–0.09 5 41 0.42 0.34–0.47

Celtus tenuifolia 5 15 0.15 0.1–0.2 3 34 0.35 0.03–0.84 4 29 0.3 0.15–0.46

Juniperus virginiana 5 48 0.49 0.4–0.7 5 33 0.34 0.2–0.54 5 23 0.24 0.14–0.34

Symphoricarpos 3 13 0.13 0–0.4 3 25 0.26 0.01–0.48 2 13 0.13 0–0.29 orbiculatus

Rhus aromatica 3 5 0.05 0–0.1 2 23 0.24 0–0.59 3 8 0.08 0.01–0.19

Table C-5 (continued). Average small woody (< 0.5 meter [1.6 ft]) plant density measurements for glade 21, 1993–2019. (Frequency, Glade Mean Density [m2—square meters], 95% Confidence Intervals [CI]).

Species 1993 1993 1993 1993 2008 2008 2008 2008 2019 2019 2019 2019 Freq. Sum Density 95% CI Freq. Sum Density 95% CI Freq. Sum Density 95% CI (plants/m²) (plants/m²) (plants/m²)

Frangula caroliniana 0 NA 0 0–0 1 1 0.01 0–0.04 3 5 0.05 0.01–0.08

Quercus stellata 0 NA 0 0–0 3 9 0.09 0.02–0.13 2 4 0.04 0–0.09

Pinus sp. 5 121 1.25 0.9 0 NA 0 0–0 0 NA 0 0–0 1.7

Quercus phellos 0 NA 0 0 2 2 0.02 0–0.05 0 NA 0 0–0 0

Quercus sp. 0 NA 0 0 1 1 0.01 0–0.03 0 NA 0 0–0 0

Rosa multiflora 2 8 0.08 0 0 NA 0 0–0 0 NA 0 0–0 0.1

The Department of the Interior protects and manages the nation’s natural resources and cultural heritage; provides scientific and other information about those resources; and honors its special responsibilities to American Indians, Alaska Natives, and affiliated Island Communities.

NPS 301/175135, March 2021

National Park Service U.S. Department of the Interior

Natural Resource Stewardship and Science 1201 Oakridge Drive, Suite 150 Fort Collins, CO 80525

EXPERIENCE YOUR AMERICA TM