Collinsonia Verticillata Is a Mid-Sized Forest Herb
HABITAT CHARACTERIZATION AND INVENTORY OF COLLINSONIA VERTICILLATA
IN SHAWNEE STATE FOREST AND STATE PARK, SCIOTO COUNTY, OHIO
A thesis presented to
the faculty of
the College of Arts and Sciences of Ohio University
In partial fulfillment
of the requirements for the degree
Master of Science
Jennifer L. Gray
June 2006 This thesis entitled
HABITAT CHARACTERIZATION AND INVENTORY OF COLLINSONIA VERTICILLATA
IN SHAWNEE STATE FOREST AND STATE PARK, SCIOTO COUNTY, OHIO
by
JENNIFER L. GRAY
has been approved for
the Program of Environmental Studies
and the College of Arts and Sciences by
Philip D. Cantino
Professor of Environmental and Plant Biology
Benjamin M. Ogles
Dean, College of Arts and Sciences GRAY, JENNIFER L., M.S., June 2006. Environmental Studies.
HABITAT CHARACTERIZATION AND INVENTORY OF COLLINSONIA VERTICILLATA
IN SHAWNEE STATE FOREST AND STATE PARK, SCIOTO COUNTY, OHIO (69 pp.)
Director of thesis: Philip D. Cantino
This study seeks to answer the following questions: (1) What is the current status of the
Collinsonia verticillata populations in Ohio? (2) What site and vegetation characteristics are typical of the Ohio habitat of Collinsonia verticillata? Previously known populations were ground-checked to determine current population status. Plots were established within selected populations in order to record vegetation and environmental data.
The Ohio populations of Collinsonia verticillata are stable, and four new sites were discovered over the course of the study. The habitat sites are on steep north to northeast-facing lower slopes on silt-loam soils with low pH. Associated tree species are white oak, sugar maple, and red maple. Viburnum dentatum may be an associated indicator species. Possible threats to the populations include invasive species, logging, and soil disturbance. Frequent monitoring and removal of invasive plants are needed. Location of additional populations in the future is likely.
Approved:
Philip D. Cantino
Professor of Environmental and Plant Biology
4 Table of Contents
Page
Abstract...... 3
List of Tables ...... 5
List of Figures...... 6
Introduction...... 7
Research Questions...... 13
Methods ...... 13 Management and Inventory ...... 13 Plot Establishment ...... 14 Data Collection ...... 16
Analyses...... 17 Herb and Tree Data...... 17 Soil Analysis...... 18 Environmental Factors...... 18
Results...... 18 Inventory...... 18 Herb and Tree Data...... 25 Characterization of the Tree Layer ...... 27 The Herb Layer...... 35 Soil and Environmental Analyses...... 37
Other Observations ...... 41
Discussion...... 47
Conclusions...... 50
Literature Cited ...... 51
Appendix A: Data Sheets...... 54
Appendix B: Soil Analysis Protocols ...... 57
Appendix C: List of All Species Found in Study Plots...... 59
Appendix D: Tree Survey, List of Tree Species in Each Plot...... 61
Appendix E: Quad Survey, Herb Species and Cover Data ...... 66
5 List of Tables
Table Page
1. Status of Collinsonia verticillata populations in Shawnee State Forest ...... 19
2. Plot species totals...... 26
3. Comparison of all plots...... 27
4. Summary of all tree data by plot...... 31
5. Summary of all tree data by species...... 32
6. Summary of canopy tree data by species...... 33
7. Analysis of understory species...... 34
8. Top five species recorded for each plot in the herb layer survey...... 36
9. pH and clay percentage parameters of soils...... 38
10. Properties of soil samples ...... 39
11. Plot aspect and slope...... 41
6 List of Figures
Figure Page
1. Collinsonia verticillata in bloom...... 7
2. U.S. county distribution of Collinsonia verticillata...... 9
3. Extent of Shawnee State Forest and Shawnee State Park...... 10
4. Physiographic map of Ohio...... 11
5. Locations of Collinsonia verticillata within Shawnee State Forest...... 12
6. Plot layout for this study...... 14
7. Locations of plots established for tree and herb sampling...... 15
8. Healthy population of Collinsonia verticillata showing size distribution ...... 20
9. Standing dead trees as a result of a forest fire...... 21
10. Vigorous but yellowed Collinsonia verticillata...... 22
11. Healthy blooming Collinsonia verticillata ...... 23
12. Location of newly discovered populations of Collinsonia verticillata...... 25
13. GPS data point locations as an overlay on soil survey...... 40
14. Collinsonia verticillata growing with multiple flower spikes ...... 42
15. Close up of the flower of Collinsonia verticillata ...... 43
16. Deer browsed Collinsonia verticillata ...... 44
17. Developing nutlets of Collinsonia verticillata...... 45
18. Mature and developing nutlets of Collinsonia verticillata ...... 46
7 Introduction
Collinsonia verticillata Baldwin ex Elliott, commonly called early stoneroot or whorled horse-balm, is a rare, mid-sized forest herb in the mint family (Lamiaceae). Alternate (older) names of Collinsonia verticillata are Hypogon verticillata and Micheliella verticillata. This plant has 2-3 pairs of opposite leaves that are closely crowded, appearing whorled, and have a thick appearance and texture. Collinsonia verticillata blooms from May to July, producing an unbranched inflorescence with whorls of 3-6 pinkish flowers having four stamens each (Figure
1), and can reach as much as 5 dm in height. As the name “stoneroot” implies, Collinsonia verticillata has a very coarse, woody rhizome. It reproduces both from seed and by sprouting from these spreading rhizomes.
Figure 1. Collinsonia verticillata in bloom. Shawnee State Forest, Scioto Co., OH, 5-24-05.
8 The much more common Collinsonia canadensis is easily differentiated from C.
verticillata when in flower, but less easily when only vegetation is present. Collinsonia
canadensis has leaves that are clearly opposite (not appearing whorled), faintly lemon-scented,
and thinner in appearance and texture when compared with those of C. verticillata. Collinsonia canadensis blooms much later, from August to mid-October, and has a branching, yellow- flowered inflorescence with two stamens per flower. Collinsonia canadensis has been used as a
medicinal plant for many years, and it is possible that C. verticillata has been used in this way as
well.
Collinsonia verticillata has been located in less than 60 counties in the eastern U.S. and is globally ranked G2/G3, indicating imperiled or rare status. Most of the populations are found in
Georgia, South Carolina, and Tennessee, but populations have also been found in two counties each of southern Virginia and Kentucky, and one county each of North Carolina, Alabama and
Ohio (Figure 2). Collinsonia verticillata is considered rare or “of concern” in all of these states except Alabama and Tennessee, but it is not included on the U.S Fish and Wildlife Service list of threatened or endangered species. The state conservation status for those states that do include the plant on their lists is as follows: Ohio, Kentucky, and Virginia all list Collinsonia verticillata as
S1, critically imperiled or endangered. North Carolina lists C. verticillata as S2, imperiled.
Georgia lists the plant as S3, vulnerable, and South Carolina lists it as SC, or ‘of state concern’.
The rarity of C. verticillata has meant that it has been the subject of very few projects.
Fone (1987, 1989) and Fone and Wyatt (1989) studied seed production in C. verticillata, finding that seed production was greater with greater plant size, but was limited by low pollination rates.
Peirson (2003) discussed the plant in his study of the systematics of the genus Collinsonia, finding C. verticillata to be a morphologically distinct species. Very little has been written about the ecology and habitat characteristics of this species.
9
Figure 2. U.S. county distribution of C. verticillata, based on combined data from Peirson (2003), the Center for Plant Conservation (2006), University of North Carolina herbarium (2004), USDA NRCS Plants Database (2006), the University of Tennessee Herbarium (2006), and the South Carolina Plant Atlas (2006).
Ohio’s Populations The Ohio populations of Collinsonia verticillata are located within the Shawnee State
Park and Shawnee State Forest in the western portion of Scioto County (Figure 3). Shawnee State
Forest is located in the unglaciated portion of Ohio on the Allegheny Plateau and, more specifically, on the Shawnee-Mississippian Plateau (Zone 15 in Figure 4), an area characterized by Mississippian-age shales, sandstones, and siltstones (Brockman, 1998) and steep, deeply dissected terrain. The Shawnee-Mississippian Plateau occupies a fairly small portion of the state, and its geologic characteristics are similarly restricted, as are these characteristics’ presumed influence on the habitat type. Collinsonia verticillata was first discovered in this area in 1963 by
10 E. Lucy Braun, who located four populations within one mile of each other: one small patch, one of unrecorded size, and two “large discontinuous patches” (Braun, 1964).
Ohio’s Natural Heritage Database, operated by the Ohio Department of Natural
Resources Division of Natural Areas and Preserves, currently has on record 16 separate
populations, all located within a few miles of each other (Figure 5). Populations on record range
in size from ~12 to ~ 10,000 individuals, though many of the records had not been updated for 20
or even 30 years, and the current condition of the recorded populations was unknown at the outset
of my study. The Shawnee State Forest does not currently have a management plan for these
populations.
Figure 3. Extent of Shawnee State Forest and Shawnee State Park in Scioto County and eastern Adams County Ohio. Shawnee State Forest shape file courtesy of Mark Ervin, ODNR Division of Forestry.
11
Figure 4. Physiographic map of Ohio (Brockman, 1998). Shawnee State Forest, in Scioto County and eastern Adams County, is in the unglaciated portion of Ohio on the Shawnee- Mississippian Plateau (zone 15).
12
Figure 5. Locations of Collinsonia verticillata within Shawnee State Forest, Scioto County Ohio, just S and SW of St. Rte. 125. Map produced by ODNR-DNAP, Natural Heritage Database from records held as of 4/2005.
13 Research Questions
There are two primary research questions of this study: (1) What is the current status of the known Collinsonia verticillata populations in Ohio? (2) What site and vegetation characteristics are typical of the Ohio habitat of Collinsonia verticillata? The aim of this study is to update current population status through ground checking current ODNR-DNAP records, and to investigate environmental factors in order to determine those factors that characterize the
‘typical’ habitat of C. verticillata. In addition, recommendations will be made for possible management strategies, as there are currently none in place.
The following sub-questions will help in providing a thorough answer to the primary research questions:
Inventory and Ecological Information How healthy are the known populations? What are the environmental characteristics of the growing sites? Comparative Factors Are the site characteristics of the current populations’ locations similar? Management How is Collinsonia verticillata being managed by other groups or organizations? What are possible management strategies for the Ohio populations?
Methods
Management and Inventory
I attempted to collect management information from agencies and organizations in the
southeast U.S. that have C. verticillata on their properties, in order to determine what
management strategies, if any, they currently use. I received no response, and I suspect this is
because there are no management plans in effect that specifically deal with the health and
perpetuation of Collinsonia verticillata. Because of the lack of response, this topic will not be covered further in the results section.
14 The ODNR DNAP Natural Heritage Database records were updated through field- checking known locations and counting or visually estimating the number of individual plants
(ramets) at each site, also noting age distribution and site condition. An attempt was made to visit all 16 sites, though not all 16 were accessible (see results). Each updated site was marked with
GPS to accurately record locations for future reference.
Plot Establishment
While inventorying each site, a count of individuals was taken for the entire population of
Collinsonia verticillata. Following this, plots (Figure 6) were established within some populations for collection of herb layer and tree data. Herb data were collected along a diagonal transect across the plot, while tree data were collected throughout the entire plot area.
Figure 6. Plot layout for this study. Circles indicate soil sample locations. Fourteen 0.25m2 herb layer sampling quads were spaced along the diagonal transect, with one every two meters.
15 A total of eight 20m x 20m plots were established for this study (Figure 7) within DNAP sites
004, 008, 009, 010, 016 and two newly discovered sites. Plot locations were determined by proximity to current C. verticillata populations. Where area was sufficient and access was possible, plots were located either fully within large populations or fully including small populations and the areas surrounding them. Two plots (2 and 3) were established within one large population (ODNR Site 009) where the population size allowed for a distance of 20+ meters between plots. Once established, one corner of each plot was recorded with a Garmin GPS II
PLUS handheld GPS unit, and each corner was permanently marked with a steel bar painted yellow at the top. Directionality of the plot axes was recorded as well.
Figure 7. Locations of plots established for tree and herb sampling of Collinsonia verticillata habitat.
16 Data Collection
Within each plot, a 28.28m transect was placed on the diagonal from upslope to downslope for the collection of herb data. Data were collected using a 0.25 m2 (50x50 cm) quad
placed at 2-meter intervals. The first quad was placed at the 1.5 m mark (covering the ground
space from 1.5 to 2m) on the upslope side of the tape. Subsequent quads followed at 2m intervals:
3.5m, 5.5m, 7.5m, etc., continuing to 27.5m for a total of 14 quads per plot. Species and % cover
were recorded for all species using the following scheme: 0=overhanging, 1= <1%, 2= 1-5%,
3=5-25%, 4=25-50%, 5= 50% or greater. Unknown plants were given a cover value, and a
specimen was taken from outside of the study quad and pressed for later identification.
Nomenclature follows Gleason and Cronquist (1991) for all recorded species except Fraxinus
biltmoreana, taken from Yatskievych and Turner (1990), and Carex cumberlandensis from Naczi
et al (2001). Data collection was completed from mid-May to mid-July of 2005.
Species and DBH (diameter breast height) measurements were recorded for all mature trees, and for saplings over 1.3 meters in height and with 2.5 cm or greater DBH. Multiple stems arising from a common root system were recorded separately if they branched below 1.3 m.
Standing dead trees were not measured.
Soil samples were taken in two corners of each plot, one from the highest elevation and one from the lowest elevation in the plot. The slope and aspect of each plot were also recorded.
Photos were taken of each plot, and photo directionality and location were noted on the data sheet. Specialized data forms were produced for recording data specific to this study (Appendix
A).
17 Analyses
Herb and Tree Data
Analysis provided a means to discern patterns that characterize the habitat of C. verticillata. First, the number of herb species and tree species for each plot were summed separately and then added together to create a total species count and species list for each plot.
This information was used to determine a total species count for all plots combined. Species richness and average species per plot were also calculated.
Cover data for herb species were totaled by species for each 400m2 plot and for all plots combined. This provided a value of total possible cover value between 0 and 560 (14 quads x 5 x
8 plots) overall for each herb species. The number of plots and number of quads in which each herb layer species occurred was also recorded.
Woody species data were totaled for each 400 m2 (0.04 ha) plot, creating a stem count for each woody species with stems over 2.5 cm in diameter. Stem counts for tree species were used to calculate mature tree density (DEN) in trees and saplings per hectare, and relative density
(RDEN), calculated as ((DENSPEC/DENSUM)*100). DBH measurements were used to calculate the
basal area (BA), or dominance, of each tree species in m2/ha. Using the BA calculations, relative dominance, or relative basal area (RBA; %) was calculated. Following McCarthy et al (2001), a relative importance value (RIV) was calculated for each tree species based on the average of the sum of relative density (RDEN; %), and relative dominance (RBA; %). These calculations were also done separately for all tree species with DBH of 10.0 cm or greater to characterize the forest canopy, and for all tree species with DBH of 2.5-9.99 cm to characterize the understory, as well as summarized by plot.
Species lists, including all herb layer and tree survey species, for all plots were compared to determine the number of species in common for each possible pair of plots. Sorensen similarity
(or the Bray-Curtis coefficient) was then calculated for all plot comparisons using the following
18 equation: 2w/ (A+B), in which A and B are species totals for the two plots being compared, and w=the number of species the two plots have in common (McCune and Grace, 2002). The result of this equation multiplied by 100 provides a value of percent similarity between the vegetation of the two plots.
Soil Analysis
Soil samples were analyzed for texture and pH. Texture analysis was done using the hydrometer method. Measurements of pH were taken by the glass electrode method using an
Oakton pH/mv/°C Meter (pH 10 Series) calibrated with buffer solutions for pH 7 and 4. Analysis protocols (Appendix B) were taken from McCarthy (1997).
Environmental Factors
Environmental characteristics (elevation, slope aspect, pH) of the plots were compared in order to determine if there were noticeable similarities between plots. Species and environmental factors that are found to be similar between most or all of the plots will be considered to be characteristics that in part typify the habitat of Collinsonia verticillata in Scioto County.
Results
Inventory
Of the sixteen previously known Collinsonia verticillata sites, eleven were successfully located and inventoried (Table 1). The sites that were not inventoried (005-007, 011 and 013) were either rendered inaccessible due to extensive forest damage sustained during a severe ice storm in February of 2003 or were not located at the specified map location and coordinates. The condition of Collinsonia verticillata on updated sites was, for the most part, healthy, and most populations were the same size as previously recorded. All of the stable or increased populations
19 had a wide range of different sized and (presumably) aged individuals, from small, sterile
(non-flowering) plants to much larger, fully flowering and seed-producing individuals (Figure 8).
Table 1. Status of Collinsonia verticillata populations in Shawnee State Forest, Scioto County Ohio. Sight records of author and ODNR Natural Heritage records.
Current Previous Occurrence Number of Compared to Inventory Inventory Associated Number Updated ramets Previous # Condition Date Date Plot # 1 Y > 50 no prev. info. widely scattered 5/25/05 6/13/73 N/A 2 Y ~ 60 no prev. info. clustered 5/25/05 1981 N/A clustered–open 3 Y 500-?* indeterminable areas 7/7/05 5/14/85 N/A 4 Y 1000-5000 consistent scattered clusters 7/6/05 5/16/85 6 5 N - - - - 5/8/91 N/A 6 N - - - - 5/16/85 N/A 7 N - - - - 5/16/85 N/A 8 Y < 1000 consistent scattered clusters 6/22/05 5/19/85 4 9 Y 5000-10,000 consistent scattered clusters 6/15/05 6/4/03 02 and 03 10 Y 5000-10,000 consistent scattered 7/7/05 5/24/85 7 11 N - - - N/A 12 Y 1 decrease solitary 7/19/05 5/26/85 N/A 13 N - - - 5/25/85 N/A 14 Y 50-100 increase clustered 6/22/05 5/8/91 N/A closely clustered 15 Y 8 decrease (~4sq ft) 5/20/05 5/17/98 N/A 16 Y 500-1000 consistent scattered 5/24/05 5/12/04 1 *See text below for explanation.
20
Figure 8. Healthy population of Collinsonia verticillata showing size distribution. Shawnee State Forest, Scioto County, OH. 5-13-05.
The majority of the updated sites were damaged by the 2003 ice storm. The most
damaged areas were characterized by very heavy undergrowth and were hard to navigate due to
the presence of large fallen trees and limbs. Even in these areas, the presence of at least the large
flowering individuals of C. verticillata was evident due to the showy spikes, which rose above other herbs, and the distinctive look of the plant. It is possible that counts were underestimated, however, because smaller plants may have been hidden under the debris left by the ice storm.
Of particular note are sites 003 and 016. An especially hot, natural fire burned site 003 a
few years ago (Kevin Bradbury, Shawnee State Park Manager, personal communication) and
killed nearly all of the mature trees. Standing dead trees remain, and a heavy undergrowth of ~ 3-
10 ft seedlings and saplings covers the area (Figure 9). The Collinsonia plants survived this fire
and are growing vigorously, though yellowed and having thicker leaves than normal, in the open
21 areas closest to the road (Figure 10). The dense undergrowth made it impossible to determine the full extent of the population, but 500+ individuals were visible in the more open areas.
Figure 9. Standing dead trees as a result of a forest fire. ODNR-DNAP Heritage Database C. verticillata site 003, Shawnee State Forest, Scioto County, OH. 7-07-05.
22
Figure 10. Vigorous but yellowed Collinsonia verticillata behind sprouting tulip poplar and oak, ODNR-DNAP Heritage Database site 003, Shawnee State Forest, Scioto County OH. 7- 07-05.
Site 016 is located in an area that is managed by ODNR, Division of Forestry using controlled burns. On this site, the mature trees were not damaged by the fire, but smaller saplings and seedlings seem to have been killed. As on site 003, the Collinsonia verticillata does not seem to have been affected by the fire, and remains healthy (Figure 11). The example of these two sites suggests that C. verticillata is hardy enough to survive both low intensity managed fires and much more intense natural fires.
23
Figure 11. Healthy blooming Collinsonia verticillata at ODNR DNAP Heritage database site 016, controlled burn management area, Shawnee State Forest, Scioto County, OH. 5-24-05.
Only two of the updated sites (012 and 015) were found to have reduced numbers. For site 012, only one plant was located on a site previously listed as having “hundreds.” The specified area was easily located by using both sight and GPS to find specified coordinates.
Though the area was damaged by the 2003 ice storm, the forest was still easily navigable, and an extensive search produced only one individual. The extreme decrease in numbers, along with inconsistencies in the location description and coordinate location, leads me to believe that the map location, description, and coordinates may have been incorrectly recorded, as no other
24 apparent disturbance to the area existed that would explain the greatly reduced numbers.
Site 015 was previously recorded as a very small population of ~12 plants, and currently, only eight remain. The area was heavily damaged by the February 2003 ice storm and lies on the edge of an aging white pine plantation in which all but a few of the pines have died and fallen.
The area has a heavy cover in most places of 3-5ft tall seedlings and saplings and, in more open
areas, a heavy low cover of Japanese honeysuckle (Lonicera japonica) and poison ivy
(Toxicodendron radicans). Immediately next to the small patch of Collinsonia verticillata, there is a large multiflora rose (Rosa multiflora) bush that threatens to overtake the few remaining plants.
In addition to updating the previously known locations of C. verticillata, four new locations were discovered (Figure 12). N1 was a rather large population of ~100 plants, N2 had only 8, N3 had ~ 25, and N4 had ~20. These areas were located not through an extensive search, but merely by walking the existing trails in Shawnee State Park and exploring the vegetation for a short distance on either side of the trail. The relative ease with which these populations were located gives some weight to the idea that other populations of C. verticillata may exist in other less accessible parts of Shawnee State Forest or surrounding areas.
25
Figure 12. Location of newly discovered populations of Collinsonia verticillata within Shawnee State Forest, Scioto County, OH.
Herb and Tree Data
The total number of species recorded in the eight study plots was 156 (Appendix C),
including 33 species of trees, 13 shrub species, and 111 species of ‘herbs’ (including forbs, ferns,
grasses, sedges, and vines). The number of species per plot (species richness) ranged from 50 to
67, showing minimal variation in species richness between plots. The average number of species
recorded per plot was 58 (Table 2).
Comparisons of plot species lists to determine species in common, and calculation of
Sorenson Similarity (Table 3), illustrate the minor variation between plots. Percent species overlap ranged from 37.0% to 74.0%. When Sorenson Similarity values were compared, percent similarities ranged from 38.5% to 68.5%. The two plots that were most similar were plots 2 and
26 3. This high degree of similarity is expected, as these are the two plots that were established within the same large population of C. verticillata. The two plots with the least similarity were plots 2 and 5, though percent species overlap of plot 2 with plot 5 was 40%, and overlap of plot 5 with plot 2 was 37%. The Sorenson similarity value for the comparison of plots 2 and 5 was
38.5%, still a fairly high degree of similarity.
Table 2. Plot species totals. Total Plot Species 1 67 Total species in all plots: 156 2 50 Average species per plot: 58 3 58 4 59 5 54 6 60 7 59 8 57
27 Table 3. Comparison of all plots using species presence or absence to calculate species overlap and Sorenson Similarity (or Bray-Curtis Coefficient).
# Of # of # Species in % % Sorenson % Plots Species Plot Species Common overlap overlap Similarity Compared Plot A (A) B (B) (w) A B ((2*w)/(A+B))*100 3-Feb 2 50 3 58 37 74.00 63.79 68.52 4-Mar 3 58 4 59 38 65.52 64.41 64.96 7-Apr 4 59 7 59 35 59.32 59.32 59.32 8-Mar 3 58 8 57 33 56.90 57.89 57.39 8-Apr 4 59 8 57 33 55.93 57.89 56.9 4-Feb 2 50 4 59 31 62.00 52.54 56.88 7-Mar 3 58 7 59 33 56.90 55.93 56.41 6-Jan 1 66 6 60 35 53.03 58.33 55.56 6-Apr 4 59 6 60 33 55.93 55.00 55.46 8-May 5 54 8 57 30 55.56 52.63 54.05 7-Jun 6 60 7 59 32 53.33 54.24 53.78 3-Jan 1 66 3 58 33 50.00 56.90 53.23 6-Feb 2 50 6 60 29 58.00 48.33 52.73 4-Jan 1 66 4 59 32 48.48 54.24 51.2 7-Jan 1 66 7 59 32 48.48 54.24 51.2 6-Mar 3 58 6 60 30 51.72 50.00 50.85 8-Feb 2 50 8 57 27 54.00 47.37 50.47 5-Mar 3 58 5 54 28 48.28 51.85 50 8-Jul 7 59 8 57 29 49.15 50.88 50 5-Apr 4 59 5 54 28 47.46 51.85 49.56 7-Feb 2 50 7 59 27 54.00 45.76 49.54 2-Jan 1 66 2 50 27 40.91 54.00 46.55 8-Jun 6 60 8 57 27 45.00 47.37 46.15 7-May 5 54 7 59 26 48.15 44.07 46.02 8-Jan 1 66 8 57 27 40.91 47.37 43.9 5-Jan 1 66 5 54 26 39.39 48.15 43.33 6-May 5 54 6 60 23 42.59 38.33 40.35 5-Feb 2 50 5 54 20 40.00 37.04 38.46
Characterization of the Tree Layer
Summary By Plot
Each plot was first totaled individually (Appendix D) and those data were used to create the following summary. In all of the eight (0.04 ha each) plots combined, 33 species of trees were recorded (Table 4). The number of species per plot ranged from 6 to 14, and the average number
28 of species per plot was 10. Stem counts per plot ranged from 22 to 45 with a total of 245 stems in all plots combined, and an average per plot stem count of about 31. Overall, tree density (DEN) ranged from 550 to 1125 stems/ha with an average density of 765.63 stems/ha. Basal area (BA) ranged from 20.68 to 41.38 m2/ha with an average BA of 26.37 m2/ha. When understory and
canopy trees were separated, 111 of the 245 total stems were classified as understory, and the
remaining 134 were classified as canopy. Canopy trees were designated as those having a DBH of
10.0 cm or greater. All trees having a DBH of 9.99 cm or less were designated as understory. On
average, the stem count of the understory was 13.88 stems/plot and the average density of
understory species was 346.88 stems/ha with an average BA of only 0.99 m2/ha. The canopy, on average, had a stem count of 16.75 stems/plot, an average stem density of 418.75 stems/ha, and an average BA of 25.38 m2/ha.
Species Analysis
When the 33 recorded tree species are examined (Table 5), the most commonly recorded species was sugar maple (Acer saccharum), which was recorded 76 times, making up 31%
(RDEN=31.02) of the total recorded trees. The second most common species of tree was red maple (Acer rubrum), recorded 46 times, with RDEN of 18.8% of the total recorded trees.
Following this was white oak (Quercus alba) with RDEN of 10.6%, black gum (Nyssa sylvatica) with RDEN of 6%, slippery elm (Ulmus rubra) with RDEN of 3.7%, and ironwood (Carpinus caroliniana) with RDEN of 3.3%. Each remaining species made up only 2% or less of the total recorded trees (RDEN< 2).
When relative importance value (RIV) and relative basal area (RBA) were examined,
however, the tree with the greatest RBA (31.38%) was white oak, though white oak was second
in relative importance (RIV= 21) to sugar maple (RIV=23), the second most dominant tree based
on RBA, with 14.99% of the total recorded BA, just less than half that of white oak. As stated
29 above, however, this species had the greatest RIV. These two species together accounted for over 46% of the total sample area BA. The remainder of the BA was composed of many species, each making up less than 10% of the total BA in the study area.
Analysis of all tree data together gives the impression of a maple-dominated forest. The differences in dominance represented by the density and BA data suggest that the forest composition cannot be accurately characterized by analyzing all the tree data at once. To better understand the forest structure and composition, two other analyses were done, one including only trees designated as “canopy” and one including all other tree species, designated as
“understory.”
Canopy trees were designated as those having a DBH greater than 10.0 cm, and (Table
6). The canopy was composed of 24 species, 18 of which were only found as canopy species with
DBH greater than 10.0cm. There were a total of 134 individuals that met the canopy criteria. The
canopy in the study area was dominated by white oak (RIV = 26), having a total BA of 66.19
m2/ha and a RBA of 32.60%, in other words, making up 32.60% of the total BA in the canopy.
Sugar maple was the second most dominant species (RIV = 20.41) with a total BA of 28.34 m2/ha
and a RBA of 13.96%. Though the density (DEN = 650 stems/ha) of white oak was less than that
of sugar maple (DEN = 900 stems/ha) the much higher BA and RBA totals for white oak indicate
that this is the dominant canopy species, existing as fewer large individuals with many smaller
sugar maples. All other species made up 10% or less of the total BA (RBA<10%), and only tulip
poplar and red maple had RIVs of greater than 4.5 with RIVs of 8.88 and 7.43 respectively.
The understory (Table 7) was composed of 15 species of trees, only five of which were also included on the canopy species list. The understory of the study area was dominated by sugar maple. This species had the greatest density (DEN = 1000 stems/ha), RDEN (36%), and RBA
(41%) of all the understory species. Sugar maple was followed by red maple, which had a density of 750 stems/ha, a RDEN of 27%, and a RBA of 26.5%. These two species combined made up
30 63% of the understory density, and 67.9% of the total BA of the understory. The RIV of sugar maple was 38.68 and that of red maple was 26.78. Black gum, ironwood, and slippery elm were the three species with the next highest RIV values with 6.86, 6.03, and 5.96 respectively. Black gum had a RDEN of 6.31% and RBA of 7.41%. Ironwood had a RDEN of 7.21% and RBA of
4.86%. Slippery elm had a RDEN of 6.31% and RBA of 5.62%. Each of the ten other species accounted for less than 3% of both the BA and DEN of the understory.
The picture we are left with, after analysis of the separate divisions of the tree data, is a forest canopy heavily dominated by mid-sized white oaks, mid-sized sugar maples, and only occasional other species. The understory was dominated by small sugar and red maples, a few black gum, ironwood, and slippery elm, and a diverse array of other small trees. Overall, the composition of this forest is typical of an Appalachian hardwood forest. White oak, the dominant canopy species, was not found at all in the understory, however, while the most dominant understory species, sugar maple, was found commonly as a lower DBH canopy species. This seems to indicate a possible future shift in forest canopy composition.
31
Table 4. Summary of Tree Data by plot, all trees > 2.5cm DBH. Number of species per plot, stem count, density (DEN), and basal area (BA).
PLOT Total Understory (2.5-9.99 cm DBH) Canopy (10+ cm DBH) # of stem DEN BA stem DEN BA stem DEN BA species count (stems/ha) (m2/ha) count (stems/ha) (m2/ha) count (stems/ha) (m2/ha)
1 14 45 1125 20.68 27 675 1.83 18 450 18.86 2 8 22 550 22.45 9 225 0.67 13 325 21.78 3 13 31 775 25.61 17 425 0.87 14 350 24.74 4 8 23 575 23.31 6 150 0.47 17 425 22.84 5 13 34 850 25.17 14 350 1.33 20 500 23.84 6 10 28 700 23.62 13 325 0.82 15 375 22.80 7 6 22 550 28.72 4 100 0.41 18 450 28.30 8 9 40 1000 41.38 21 525 1.50 19 475 39.88
Totals 33 245 6125 210.94 111 2775 7.91 134 3350 203.03
Avg. 10.13 30.63 765.63 26.37 13.88 346.88 0.99 16.75 418.75 25.38 SD 2.90 8.62 215.45 6.52 7.72 192.93 0.51 2.49 62.32 6.44
32 Table 5. Summary of tree layer survey by species for all plots combined. Stem count, density (DEN), relative density (RDEN), basal area (BA), relative basal area (RBA), and relative importance value (RIV; [RDEN+RBA]/2). Sorted by RIV in descending order.
# of stem DEN BA SPECIES Plots count (stems/ha) RDEN (m2/ha) RBA RIV Acer saccharum 8 76 1900 31.02 31.61 14.99 23.00 Quercus alba 7 26 650 10.61 66.19 31.38 21.00 Acer rubrum 7 46 1150 18.78 8.04 3.81 11.29 Liriodendron tulipifera 5 11 275 4.49 19.39 9.19 6.84 Nyssa sylvatica 5 15 375 6.12 6.63 3.14 4.63 Pinus strobus 1 2 50 0.82 14.56 6.90 3.86 Quercus velutina 2 3 75 1.22 11.97 5.67 3.45 Quercus prinus 2 4 100 1.63 10.13 4.80 3.22 Ulmus rubra 3 9 225 3.67 1.89 0.90 2.29 Quercus sp. 1 1 25 0.41 8.76 4.15 2.28 Carya laciniosa 3 4 100 1.63 4.85 2.30 1.97 Tilia americana 2 4 100 1.63 4.79 2.27 1.95 Carpinus caroliniana 4 8 200 3.27 0.38 0.18 1.72 Carya ovata 4 4 100 1.63 3.18 1.51 1.57 Fraxinus americana 3 3 75 1.22 2.71 1.28 1.25 Quercus coccinea 1 1 25 0.41 3.41 1.62 1.01 Carya sp. 2 2 50 0.82 2.38 1.13 0.97 Quercus rubra 1 1 25 0.41 2.07 0.98 0.70 Ostrya virginiana 2 3 75 1.22 0.23 0.11 0.67 Euonymus sp.(cultivar) 1 3 75 1.22 0.22 0.10 0.66 Cercis canadensis 3 3 75 1.22 0.15 0.07 0.65 Carya cordiformis 1 1 25 0.41 1.83 0.87 0.64 Juglans nigra 1 1 25 0.41 1.73 0.82 0.61 Platanus occidentalis 1 1 25 0.41 1.34 0.63 0.52 Cornus florida 2 2 50 0.82 0.42 0.20 0.51 Fraxinus sp. 1 2 50 0.82 0.08 0.04 0.43 Hamamelis virginiana 2 2 50 0.82 0.07 0.03 0.43 Magnolia tripetala 1 2 50 0.82 0.07 0.03 0.43 Carya tomentosa 1 1 25 0.41 0.76 0.36 0.38 Fagus grandifolia 1 1 25 0.41 0.73 0.35 0.38 Oxydendrum arboreum 1 1 25 0.41 0.27 0.13 0.27 Amelanchier arborea 1 1 25 0.41 0.08 0.04 0.22 Lindera benzoin 1 1 25 0.41 0.01 0.01 0.21 Totals 245 6125 210.94
33 Table 6. Summary of canopy trees by species, all plots combined. All trees DBH 10.0 cm or greater. Stem count, density (DEN), relative density (RDEN), basal area (BA), relative basal area (RBA), and relative importance value (RIV; [RDEN+RBA]/2). Ranked in descending order by RIV.
stem DEN BA SPECIES count (stems/ha) RDEN (m2/ha) RBA RIV Quercus alba 26 650 19.40 66.19 32.60 26.00 Acer saccharum 36 900 26.87 28.34 13.96 20.41 Liriodendron tulipifera 11 275 8.21 19.39 9.55 8.88 Acer rubrum 16 400 11.94 5.94 2.92 7.43 Nyssa sylvatica 8 200 5.97 6.05 2.98 4.47 Pinus strobus 2 50 1.49 14.56 7.17 4.33 Quercus velutina 3 75 2.24 11.97 5.89 4.07 Quercus prinus 4 100 2.99 10.13 4.99 3.99 Carya laciniosa 4 100 2.99 4.85 2.39 2.69 Tilia americana 4 100 2.99 4.79 2.36 2.67 Quercus sp. 1 25 0.75 8.76 4.32 2.53 Carya ovata 4 100 2.99 3.18 1.57 2.28 Fraxinus americana 2 50 1.49 2.56 1.26 1.38 Carya sp. 2 50 1.49 2.38 1.17 1.33 Quercus coccinea 1 25 0.75 3.41 1.68 1.21 Ulmus rubra 2 50 1.49 1.45 0.71 1.10 Quercus rubra 1 25 0.75 2.07 1.02 0.88 Carya cordiformis 1 25 0.75 1.83 0.90 0.82 Juglans nigra 1 25 0.75 1.73 0.85 0.80 Platanus occidentalis 1 25 0.75 1.34 0.66 0.70 Carya tomentosa 1 25 0.75 0.76 0.38 0.56 Fagus grandifolia 1 25 0.75 0.73 0.36 0.55 Cornus florida 1 25 0.75 0.36 0.18 0.46 Oxydendrum arboreum 1 25 0.75 0.27 0.13 0.44 Totals 134 3350 203.03
34 Table 7. Summary of understory species all plots combined. All trees DBH 2.5 – 9.99 cm. Stem count, density (DEN), relative density (RDEN), basal area (BA), relative basal area (RBA), and relative importance value (RIV; [RDEN+RBA]/2). Ranked in descending order by RIV.
stem DEN Sum BA SPECIES count (stems/ha) RDEN (m2/ha) RBA RIV Acer saccharum 40 1000 36.04 3.27 41.32 38.68 Acer rubrum 30 750 27.03 2.10 26.54 26.78 Nyssa sylvatica 7 175 6.31 0.59 7.41 6.86 Carpinus caroliniana 8 200 7.21 0.38 4.86 6.03 Ulmus rubra 7 175 6.31 0.44 5.62 5.96 Ostrya virginiana 3 75 2.70 0.23 2.85 2.78 Euonymus sp.(cultivar) 3 75 2.70 0.22 2.73 2.72 Cercis canadensis 3 75 2.70 0.15 1.89 2.29 Fraxinus americana 1 25 0.90 0.15 1.92 1.41 Fraxinus sp. 2 50 1.80 0.08 0.98 1.39 Hamamelis virginiana 2 50 1.80 0.07 0.91 1.35 Magnolia tripetala 2 50 1.80 0.07 0.91 1.35 Amelanchier arborea 1 25 0.90 0.08 1.05 0.98 Cornus florida 1 25 0.90 0.06 0.81 0.85 Lindera benzoin 1 25 0.90 0.01 0.18 0.54 Totals 111 2775 7.91
35 The Herb Layer
The herb layer survey yielded 145 species comprising 70 forbs, 26 trees, 11 vines, 10 shrubs, 10 ferns, 10 grasses, and 8 sedges (Appendix E). Of the 145 species, only seven were recorded in all eight plots: Acer rubrum, Acer saccharum, Anemonella thalictroides, Galium triflorum, Parthenocissus quinquefolia, Viburnum dentatum, and Viola sororia.
Collinsonia verticillata grew in all eight of the plots but, due to the transect-quad survey method used, was not recorded in one of the plots because it did not occur in any of the quads.
The herb layer was too thick in many of the areas to attempt recording all species, even if they did not occur in the sampling quads, so only the species found within or overhanging the sampling quads were recorded.
The species with the greatest amount of cover in the herb layer survey quads was
Parthenocissus quinquefolia with a total cover value of 111. Parthenocissus quinquefolia was
also the species that was recorded in the greatest number of quads: 48 out of the 112 total.
Tiarella cordifolia had the second highest cover, with a total of 98. Tiarella cordifolia was
recorded in only four of the eight plots, but it was recorded in the second greatest number of
quads (42 out of 112), indicating that in the areas where it occurs, Tiarella cordifolia makes up a
fairly large portion of the herb layer. Being a small herb, this means that there are large numbers
of small individuals making up a large portion of the cover. Laportea canadensis had the third
highest cover, with a total of 80, but was only recorded in 4 of the plots and 29 of the 112 survey
quads. Laportea canadensis presents the opposite situation from Tiarella cordifolia. Being a
rather large (~1-2 feet high) herb, the high cover value was more a result of fewer large
individuals with a great amount of cover each. This illustrates the difficulty of assessing herb
layer species by sum cover with combined plot data, as high cover values have resulted from
locally widespread, small plants; locally widely-spaced large plants; and generally widespread
mid-sized herbs, as in the case of Parthenocissus quinquefolia.
36 To better characterize the herb layer, a list was made comparing the top five species in each plot by both cover and number of quad occurrences (Table 8). Using this information, it is apparent that Parthenocissus quinquefolia was the most common species recorded, appearing in the top five species both by number of quads and sum cover in five of the eight survey plots.
Tiarella cordifolia was the second most common species by both measures.
Table 8. Top five species recorded for each plot in the herb layer survey, both by sum of cover and total quads.
Plot Plot Species in top 5 by sum cover Number(s) Species in top 5 by quads Number(s) Acer rubrum 4 Acer rubrum 3,4,6,7 Acer saccharum 7 Acer saccharum 7 Amphicarpaea bracteata 6 Anemonella thalictroides 3 Anemonella thalictroides 3,8 Asarum canadense 8 Asarum canadense 8 Carex woodii 7 Carex woodii 7 Cimicifuga racemosa 7 Cimicifuga racemosa 7 Collinsonia canadensis 2 Collinsonia canadensis 2 Desmodium nudiflorum 6 Dioscorea villosa 2,4 Disporum lanuginosum 7 Fraxinus americana 3 Galium triflorum 1 Galium triflorum 1 Impatiens capensis 5 Impatiens capensis 5 Laportea canadensis 1,2,5,6 Laportea canadensis 1,2,5 Liriodendron tulipifera 1 Lonicera japonica 5 Parthenocissus quinquefolia 1,2,4,6,8 Parthenocissus quinquefolia 1,2,4,6,8 Polemonium reptans 6 Polemonium reptans 6 Polystichum acrostichoides 8 Polystichum acrostichoides 8 Prunus serotina 4 Sassafras albidum 1 Sassafras albidum 1 Tiarella cordifolia 2,3,5,8 Tiarella cordifolia 2,3,5,6 Toxicodendron radicans 2,5 Toxicodendron radicans 3 Uvularia perfoliata 7 Uvularia perfoliata 7 Viburnum acerifolium 1,2,4 Viburnum acerifolium 4 Viburnum dentatum 4,6 Viburnum dentatum 4,6 Viola canadensis 3,5 Viola canadensis 3,5 Viola sororia 6
37 The third most common species by this method of comparison was Laportea canadensis, followed by Viburnum dentatum and Viola canadensis. Species that were in the top five in more
than one plot according to number of quads but not cover are also important to note because they
occur widely but the plants are small. Of these species, Acer rubrum was the most common,
occurring in the top five species in four plots by quads, versus one plot by cover. Anemonella
thalictroides appeared in the top five in two plots when compared by quads versus one by cover.
Of all of the above species that were commonly found in the study plots, the one that stood out most as a habitat indicator associated with Collinsonia verticillata was Viburnum dentatum. In my casual observations, while the other species mentioned above seemed to be widespread in areas not having populations of Collinsonia verticillata, Viburnum dentatum did not seem to be as widespread. My observation of Viburnum dentatum along a trail led twice to the discovery of new locations of Collinsonia verticillata growing underneath the shrubs. This apparent association was not rigorously tested but bears further investigation.
Soil and Environmental Analyses
Based on maps produced by the USDA Soil Conservation Service in the Soil Survey of
Scioto County, the main soil association in the Shawnee State Forest habitat area of Collinsonia verticillata is the Shelocta-Brownsville association, very steep (ScF), which is characterized by fairly deep soils derived from sandstones and siltstones on side slopes in especially steep terrain
(slopes from 40-70%). The Shelocta soils and Brownsville soils are both silt-loams. The ScF
association makes up 29.6% of Scioto County’s total acreage. While this soil type is the most
common in the area, however, it is relatively rare outside of this area. The C. verticillata habitat
area also contains three other soil types: the Berks channery silt loam, (BeC), which lies on upper
slopes and narrow ridges and makes up 2.3% of Scioto County’s acreage; the Shelocta-Wharton-
Latham association, steep (SfE), which is also silt loam on steep side slopes and makes up 4.1%
38 of Scioto County’s acreage; and the Skidmore silt loam (Sk), found in valley bottoms along small streams, which makes up 1.7% of Scioto County’s acreage. The properties of these soils
(Table 9) were compared with the analysis results from samples taken in the study plots.
Table 9. pH and clay percentage parameters of soils in the Collinsonia verticillata habitat area of Shawnee State Forest. Source: USDA SCS Soil Survey of Scioto County, 1989. Soil % clay % Slope Association Soil Type pH Range range Range ScF Shelocta 4.5-5.5 10-25 40-70 Brownsville 3.6-6.0 8-18 40-70 SfE Shelocta 4.5-5.5 10-25 25-40 Wharton 4.0-6.0 15-25 25-40 Latham 3.6-6.5 20-27 25-40 BeC Berks 3.6-6.5 5-23 7-15+ Sk Skidmore 5.6-7.8 17-27 0-3
Based on soil samples taken in the study plots (Table 10), all of the study plots were established on silt loam, except for one, which was on silty clay-loam. The soil sample pH ranged from 3.59 to 5.48, well within the specified ranges of the soil types in the area, and slope ranged from 24% to 44% (Table 11), also well within the parameters of the soil types and terrain. When soil survey data are compared with plot locations, the majority of sites are located on the ScF association, or on the border between ScF, Sk, and SfE (Figure 13), with only two sites falling fully within the SfE association or on the border of SfE and Sk. All sites were located on low to mid elevation slopes, so none occurred on the BeC soil type.
39 Table 10. Properties of soil samples taken in Collinsonia verticillata habitat plots. Sample 1 was from the upslope area of each plot, Sample 2 from the downslope area of each plot.
Soil Sample % Sand % Clay % Silt Type pH
Plot 1-Sample 1 27.53 5.00 67.47 silt loam 4.31
Plot 1-Sample 2 15.01 7.50 77.49 silt loam 5.09
Plot 2-Sample 1 26.80 6.37 66.84 silt loam 4.78
Plot 2-Sample 2 38.53 3.84 57.63 silt loam 5.12
Plot 3-Sample 1 20.14 9.98 69.88 silt loam 4.07
Plot 3-Sample 2 10.01 12.50 77.49 silt loam 4.28
Plot 4-Sample 1 27.60 9.99 62.42 silt loam 4.66
Plot 4-Sample 2 12.51 25.00 62.49 silt loam 4.81
Plot 5-Sample 1 7.53 27.49 64.98 silty clay loam 4.82
Plot 5-Sample 2 35.04 6.25 58.71 silt loam 3.59
Plot 6-Sample 1 12.53 14.99 72.47 silt loam 4.9
Plot 6-Sample 2 12.50 15.00 72.50 silt loam 4.85
Plot 7-Sample 1 10.01 17.50 72.49 silt loam 3.87
Plot 7-Sample 2 7.52 20.00 72.49 silt loam 3.81
Plot 8-Sample 1 10.01 20.00 69.99 silt loam 3.86
Plot 8-Sample 2 20.00 7.50 72.50 silt loam 5.48
40
Figure 13. GPS data point locations as an overlay on soil survey data.
41 Table 11. Plot aspect and slope. Aspect Slope Plot # Slope % (azimuth) Degrees 1 90(E) 18 32.49 2 0(N) 24 44.52 3 352 (NNW) 19 34.43 4 75 (ENE) 24 44.52 5 33 (NNE) 16.5 29.62 6 65 (ENE) 13.5 24.01 7 50 (ENE) 16 28.67 8 70 (ENE) 19.5 35.41
Plot aspect (Table 11) ranged from 352 degrees to 90 degrees, on hillsides facing north- northwest to east, but most commonly on ENE-facing slopes. When all new sites, plots, and
ODNR-DNAP Natural Heritage database records were examined together, out of the now 20 total
Collinsonia verticillata habitat sites, 18 are on slopes within this range of aspects. The remaining two sites are on southeast-facing slopes.
Other Observations
Over the course of my field season, I made some observations regarding the growth habits and environment of Collinsonia verticillata that do not fit into any of the ‘results’ sections of this paper but are important nonetheless.
One of the characteristics listed in descriptions of C. verticillata is a single flower spike
(Kentucky State Nature Preserves Commission, 2005; Gleason & Cronquist, 1991; NatureServe
Explorer, 2006). Over the course of my field season, I observed numerous C. verticillata plants with multiple flower spikes (Figure 14). These plants did not seem to have been damaged in any way, so the multiple spikes did not seem to be a response to injury in these cases. The number of plants I observed with multiple (usually three) flower spikes suggests to me that this is a rather common characteristic of C. verticillata, at least within the Ohio populations.
42
Figure 14. Collinsonia verticillata growing with multiple flower spikes. 5-24-05.
The flowers of the Shawnee populations of C. verticillata were all in the pink to white color range (Figure 15). Flowers are sometimes described as yellow (Kentucky State Nature
Preserves Commission, 2005; Gleason & Cronquist, 1991; NatureServe Explorer, 2006). The reference to yellow flowers may be the result of confusion with, or merely a lack of distinction from, the yellow flower color of the more common Collinsonia canadensis.
43
Figure 15. Close up of the flower of Collinsonia verticillata. 5-13-05.
I observed deer browsing on C. verticillata in only one location, directly along a trail.
The leaves and developing flower spikes were eaten (Figure 16). The plants recovered by growing new leaves but did not develop another flower spike once the original one had been eaten. This seems to indicate that C. verticillata has no trouble recovering vegetatively from deer browsing, but that deer browsing has the potential to reduce the number of flowering and seed- producing plants. The rarity of deer-browsed Collinsonia plants, even in some areas where adjacent species had been browsed, suggests that this species is not desirable to deer.
44
Figure 16. Deer browsed Collinsonia verticillata showing initial damage at left (5-13-05), and vegetative recovery at right on 6-16-05.
In the time I spent in the field, over long days that were spread out over the entire blooming season, I saw no insects on the flowers of Collinsonia verticillata. No pollinators at all were apparent.
The pattern of seed development did not seem to have any typical pattern. I observed flowers that developed no seeds, with the undeveloped ovary lobes still visible in the calyx, as well as flowers developing one to three one-seeded nutlets, with the remaining undeveloped ovary lobes still visible in the calyx. Figure 17 shows developing nutlets of C. verticillata.
45
Figure 17. Developing seeds (or nutlets) of Collinsonia verticillata. 6-15-05.
Nutlets of C. verticillata are rather large (~3mm in diameter) and pubescent, but they do not seem to act as burrs. I observed the mature nutlets hanging from the calyx seeming ready to fall to the ground (Figure 18). If this is the normal mechanism of seed dispersal, it may help to explain the isolated nature of the C. verticillata populations and their apparent lack of expansion over time.
46
Figure 18. Mature and developing seeds (nutlets) of Collinsonia verticillata, with mature seeds hanging from the calyx. 6-15-05.
Over the course of the field season, the only population I observed that seemed seriously
threatened was being affected by a forest cover change that paved the way for invasive plant
species, which now crowd the population. This seems to indicate that any land use or
management practices that cause disturbance and open areas to invasive plants may be potentially
damaging to the populations of Collinsonia verticillata. Though I stated previously that the
Shawnee State Forest populations have survived both natural and prescribed fires, these disturbed
areas may be more open to invasive species.
47 The invasive species that were found most commonly in the C. verticillata habitat areas were multiflora rose (Rosa multiflora) and Japanese honeysuckle (Lonicera japonica).
These species, though a problem, have existed as invasive species in the Shawnee State Forest for many years. More aggressive invasive plant species such as Asian stilt grass (Microstegium vimineum), garlic mustard (Alliaria petiolata), Asian bittersweet (Celastrus orbiculatus), and
Amur honeysuckle (Lonicera maackii) have not been observed in the C. verticillata habitat areas.
The invasive plants currently threatening C. verticillata populations should be removed. In addition, the habitat areas should be monitored often to ensure that invasive species that are already in the area do not become more of a threat to Collinsonia verticillata, and to ensure that any newly arrived invasive species are quickly discovered and eradicated before large populations can become established.
Based on my observations, it also seems that logging in the habitat areas could be damaging by opening areas to invasive species. Furthermore, because the rhizomes of Collinsonia verticillata seem to grow very close to the surface of the soil, any soil disturbance could be potentially damaging to the plants.
Discussion
This study has attempted to describe the areas of Shawnee State Forest and State Park that are inhabited by Collinsonia verticillata, but, as in the case of previous studies of the species, has not provided a complete explanation for why the species is located where it is. Collinsonia verticillata is believed to be native to Ohio, and for this reason, aspects of the habitat are looked to in order to explain the local distribution. Within the Shawnee populations, it seems possible that soil type, aspect, and pH may all contribute to the suitability of habitat for Collinsonia verticillata. In addition, it seems possible that the extremely rocky, steep slopes and stream edges on which Collinsonia verticillata is found in Shawnee are not conducive to other uses, and these
48 areas have therefore remained safe from disturbances that may have obliterated the populations. There may be other aspects of the habitat area, however, that were not examined in this study, which could also provide explanation for the local distribution.
In explaining the overall distribution of Collinsonia verticillata, one must question whether the disjunct Ohio populations are natural or introduced. The stability of the Shawnee populations over the past 20 or more years seems to suggest that the populations are native. It would seem that since the populations expand at such a slow rate, Collinsonia verticillata must have been in the area for quite a long time in order for populations numbering in the thousands to have become established. Also in support of the hypothesis that the Ohio populations are native,
Scioto County is home to other species of plants that occur there at the northernmost edges of their distributions. Examples of these include creeping aster (Eurybia surculosa), cross vine
(Bignonia capreolata), and carolina thistle (Cirsium carolinianum) (McCormack, 2001; ODNR-
DNAP, 2006; USDA-NRCS Plants Database, 2006). The disjunct distributions of southern
Appalachian species in Ohio are often attributed to their habitat areas’ close proximity to the course of the former Teays River (Backs, 2004), a pre-Pleistocene north-flowing river that
“originated in what is now North Carolina and flowed northwest, entering Ohio in Scioto
County” (McCormack, 2001). Southern Appalachian species reaching the northernmost extent of their range in southern Ohio are thought to have migrated north along the Teays River valley or to have been carried north as seeds by the river or its tributaries (Backs, 2004; McCormack, 2001).
McCormack (2001) specifically referred to Collinsonia verticillata as one of the “vegetative renegades” brought to the Shawnee region by the Teays drainage system prior to the Pleistocene glaciation. The Teays River, if an explanation for the Ohio disjunction, may also explain other disjunctions in the species’ distribution, such as the ones in eastern Kentucky, but it seems unlikely that this explanation would fit with the disjunct populations in southern Virginia as the
49 counties in which C. verticillata has been located lie outside of the area drained by the Teays and its tributaries (Janssen, 1953).
In support of an alternative hypothesis, the relatively recent discovery in 1963 of
Collinsonia verticillata in Shawnee State Park, and the populations’ close proximity to human impacted or human constructed areas such as roads, trails, and old home sites, suggest possible introduction to the area. The proximity of the Shawnee C. verticillata populations to human impacted areas may simply be a function of botanists locating plants in the most accessible areas of the park and forest, rather than exploring more remote locations, or may instead be a consequence of human introduction. The Shawnee State Forest is located in the Virginia Military
District, an area ceded to the state of Virginia (formerly including Kentucky) to provide lands as bounty to soldiers who had served in the Revolutionary War. Many Revolutionary War veterans
(or others to whom they sold their claims) moved from the Appalachian southeast to Ohio during the 1790s and early 1800s to lay claim to these lands (Evans, 1903; Bannon, 1920). Collinsonia, as a known medicinal plant, may have been brought along and planted in the Shawnee area.
Brothers (1992) studied plant disjunctions caused by human introduction in the eastern U.S. and found that human-caused plant introductions usually resulted in plant establishment either north or east of the plant’s native habitat. In addition, Brothers observed that disjunctions caused by human introduction and a species’ subsequent escape from cultivation were common in perennial herbs and were most common in northward plant migrations. In addition to the possibility of introduction by early settlers as a medicinal plant, there is the possibility of accidental introduction by humans or animals as a result of development in the area, possibly by workers or animals from one of the many the WPA and CCC camps established within Shawnee State Forest in the 1930s, during the Great Depression.
If additional C. verticillata populations are located in the less accessible areas of the park and forest, it would provide support for the hypothesis that the species is native. If no further
50 populations are located in the future, however, introduction remains a possible explanation for the Ohio disjunction.
Conclusions
The Shawnee populations of Collinsonia verticillata are currently stable, though the full
effects of recent fire, fire management, and the recent severe ice storm may not yet be apparent.
The increased presence of invasive species could pose a threat in the future. Continued, more
frequent monitoring is necessary.
In Scioto County, Collinsonia verticillata typically grows on steep north to northeast-
facing lower slopes, and occasionally on southeast-facing slopes, ranging in slope from 24 to
44%. The habitat sites are on silt loam soil with pH between 3.6 and 5.5, most commonly on the
Shelocta-Brownsville association, and less commonly on Skidmore silt loam and the Shelocta-
Wharton-Latham association. Collinsonia verticillata is typically found in white oak dominated
forests with a diverse understory dominated by sugar maple and red maple, and a rich herb layer.
Viburnum dentatum may be an associated indicator species. It remains unclear whether the
Shawnee populations are truly native or were introduced some time ago and have now become
naturalized.
New populations of Collinsonia verticillata were located, without extensive search, over
the course of this study. It seems quite possible that, by using the above habitat characteristics to
narrow the search area, additional populations will be located.
51 Literature Cited
Backs, J. 2004. Ohio’s Ancient Nile: The Teays River. Ohio State Parks 10. http://www.ohiodnr.com/parks/explore/magazine/sprsum04/teaysriver.htm
Bannon, H.T. 1920. Scioto sketches: an account of discovery and settlement of Scioto County, Ohio. A.C. McClurg and Co., Chicago, Illinois.
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Brockman, C.S. 1998. Physiographic Regions of Ohio. State of Ohio, Department of Natural Resources, Division of Geological Survey. Columbus, Ohio.
Brothers, T.S. 1992. Postsettlement plant migrations in northeastern North America. American Midland Naturalist 128:72-82.
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Fone, A.L. 1987. Effects of seed abortion and components of plant size and reproduction on seed yield in Collinsonia verticillata (Lamiaceae). American Journal of Botany 74:641.
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53 U.S. Department of Agriculture Soil Conservation Service. 1989. Soil survey of Scioto County, Ohio. In cooperation with U.S. Department of Agriculture, Forest Service; the Ohio Department of Natural Resources, Division of Soil and Water Conservation; and the Ohio Agricultural Research and Development Center. Washington D.C.
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54 Appendix A DATA SHEETS
55
56
57 Appendix B SOIL ANALYSIS PROTOCOLS (McCarthy, 1997)
Soil pH (glass electrode method)
Troubleshoot any pH meter problems prior to analysis.
Refill electrodes if necessary (KCl/AgCl) & test meter.
• Standardize pH meter with pH 4.0 and 7.0 buffer solution • Weigh 25 g of air-dried soil, and place into a 100 ml beaker. • Add 25 ml of distilled and de-ionized water to the beaker. • Agitate beaker on an orbital flatbed shaker at 150 OPM for 60 seconds • Let beaker stand for 10 minutes • Insert the pH electrode into the soil-water solution and swirl gently. • Read the pH immediately on a standardized pH meter. • Rinse the electrode with de-ionized water & wipe lightly with a kim-wipe.
Supplies needed: • Buffer solution pH 4.00 (Fisher #SB101-500) • Buffer solution pH 7.00 (Fisher #SB107-500) • de-ionized distilled water (5th floor Porter)
Materials needed: • Corning 350 pH/ion meter w/pH electrode • 100 ml beakers • Orbital flatbed shaker
58 Soil Texture (hydrometer method) If possible, this entire procedure should be done in a constant temp room. (NB: this procedure is not recommended for calcareous soils , saline soils, or soils > 2% organic matter.)
A. Calibrate each soil hydrometer (ASTM 152H). • Make a Calgon® solution (sodium-hexametaphosphate) by adding 50 g to one liter. • Add 100 ml of Calgon® solution to a glass cylinder; adjust vol up to 1 liter w/dH2O. • Mix thoroughly and allow to stand (ca. 30 min) at a constant temp (20-25 C). • Lower the hydrometer gently into the solution. • Determine the scale reading (RL) at the upper edge of the meniscus on the stem.
B. Second, begin your soil texture analysis. • Weigh 40.0 g of air-dried (105 C, 24 hr) & sieved soil into a 600 ml beaker. (Note: if the soil is a loamy sand or sandy, use 100 g of soil sample.) • Add 100 ml of Calgon® solution and 300 ml dH2O. • Allow the sample to stand overnight. • Transfer the Calgon® -treated sample to a metal dispersing cup. • Mix for 5 min with an electric mixer (milkshake machine). • Transfer the suspension to a cylinder and bring the volume to 1 liter w/dH2O. • Allow the suspension to equilibrate to room temp (20-25 C). • Place a clean hand (or large stopper) tightly over the top of the cylinder and turn over. • Continue with vigorous end-for-end turnovers for several minutes (mix well). • Finish with 2-3 smooth gentle turnovers. • Record the exact clock time when stirring is complete. • Add a drop or two of amyl alcohol to the cylinder if foaming is present on the top surface. • Lower the hydrometer gently into the suspension. • Take a hydrometer reading at 40 sec (R40S). • Remove the hydrometer after the reading, rinse it, wipe dry w/a kim-wipe. • Reinsert the hydrometer gently at 7 h and take another reading (R7H). • Calculate the percent sand, silt, and clay:
Sand (%) = 100 - [(R40S - RL ) * (100/oven-dried soil wt. in g)] Clay (%) = (R7H - RL ) * (100/oven-dried soil wt. in g) Silt (%) = 100 - (sand % + clay %)
Supplies needed: • Calgon® (sodium-hexametaphosphate) (Fisher #04-322B) • Amyl alcohol (50 ml) (usually avail. in soils lab)
Materials needed: • Standard soil hydrometer (ASTM No. 1 152H) • Electric stirrer (milkshake machine) • 1 liter cylinders
59 Appendix C All Species Found In Study Plots
Num Num Scientific Name Plots Scientific Name Plots Acalypha gracilens 1 Clematis virginiana 1 Acer rubrum 8 Collinsonia canadensis 7 Acer saccharum 8 Collinsonia verticillata 7 Adiantum pedatum 4 Conopholis americana 1 Agrimonia rostellata 1 Cornus alternifolia 1 Amelanchier arborea 4 Cornus florida 2 Ampelopsis brevipedunculata 1 Crataegus sp. 2 Amphicarpaea bracteata 6 Cryptotaenia canadensis 4 Anemonella thalictroides 8 Cunila origanoides 1 Aralia racemosa 1 Desmodium glutinosum 1 Arisaema triphyllum 6 Desmodium nudiflorum 5 Aristolochia serpentaria 2 Dioscorea villosa 5 Asarum canadense 6 Disporum lanuginosum 1 Asplenium platyneuron 1 Disporum maculatum 4 Aster cordifolius 1 Erechtites heiracifolia 1 Aster lateriflorus 1 Euonymus atropurpureus 2 Aster schreberi 6 Euonymus sp.(cultivar) 1 Athyrium felix-femina 1 Fagus grandifolia 3 Athyrium thelypterioides 1 Fraxinus americana 7 Botrychium virginianum 3 Fraxinus biltmoreana 1 Brachyeletrum erectum 1 Fraxinus quadrangulata 1 Bromus pubescens 1 Fraxinus sp. 1 Carex cumberlandensis 2 Galium aparine 1 Carex digitalis 2 Galium circaezans 1 Carex glaucoidea 1 Galium concinnum 2 Carex laxiculmis var. Galium triflorum 8 copulata 1 Geranium maculatum 5 Carex laxiflorum 1 Grass sp. 2 Carex sp. 3 Hamamelis virginiana 5 Carex sparganioides 1 Helianthus sp. 3 Carex woodii 2 Hepatica acutiloba 1 Carpinus caroliniana 7 Hydrangea arborescens 3 Carya cordiformis 2 Hydrastis canadensis 1 Carya glabra 1 Impatiens capensis 5 Carya laciniosa 3 Juglans nigra 1 Carya ovata 5 Laportea canadensis 4 Carya sp. 2 Lindera benzoin 6 Carya tomentosa 1 Liriodendron tulipifera 6 Celastrus scandens 2 Lonicera japonica 4 Ceonothus americanus 1 Luzula multiflora 1 Cercis canadensis 4 Lysimachia quadrifolia 1 Cimicifuga racemosa 5 Magnolia tripetala 1 Circaea lutetiana 4 Medeola virginiana 1
60 Num Num Scientific Name Plots Scientific Name Plots Monarda fistulosa 3 Smilax hispida 2 Nyssa sylvatica 6 Smilax rotundifolia 6 Orchis spectabilis 1 Solidago flexicaulis 4 Osmorhiza claytonii 4 Stellaria pubera 1 Osmunda cinnamomea 2 Taraxacum officinale 1 Ostrya virginiana 4 Thelypteris hexagonoptera 4 Oxalis grandis 1 Thelypteris noveboracensis 1 Oxydendrum arboreum 3 Tiarella cordifolia 4 Panicum boscii 1 Tilia americana 3 Panicum latifolium 1 Toxicodendron radicans 6 Panicum sp. 1 Tree sp. 1 Parthenocissus quinquefolia 8 Trillium grandiflorum 6 Pedicularis canadensis 1 Ulmus rubra 7 Phlox divaricata 2 Unknown herb 3 Pilea pumila 4 Unknown seedling 3 Pinus strobus 1 Uvularia perfoliata 6 Platanus occidentalis 1 Uvularia sessilifolia 2 Poa cuspidata 1 Viburnum acerifolium 7 Poa sp. 1 Viburnum dentatum 8 Poa sylvestris 1 Viburnum prunifolium 2 Podophyllum peltatum 5 Viola canadensis 5 Polemonium reptans 3 Viola sororia 8 Polygonum virginianum 2 Vicia caroliniana 1 Polystichum acrostichoides 7 Vitis sp. 1 Potentilla simplex 6 Prenanthes altissima 5 Prunus serotina 7 Pteridium aquilinum 1 Quercus alba 7 Quercus coccinea 1 Quercus prinus 2 Quercus rubra 3 Quercus sp. 1 Quercus velutina 2 Ranunculus recurvatus 1 Rosa carolina 1 Rosa multiflora 2 Rubus pensilvanicus 1 Rubus sp. 1 Sanguinaria canadensis 2 Sanicula canadensis 6 Sassafras albidum 2 Scutellaria incana 3 Scutellaria nervosa 1 Smilacina racemosa 5 Smilax glauca 5 Smilax herbacea 1
61
Appendix D TREE SURVEY List of Tree Species Found in Each Plot Stem count, density (DEN), relative density (RDEN), basal area (BA), relative basal area (RBA), and relative importance value (RIV; [RDEN+RBA]/2). Sum BA PLOT SPECIES stem count DEN (stems/ha) RDEN (m2/ha) RBA RIV
1 Acer rubrum 14 350 31.11 1.92 9.29 20.20 Acer saccharum 4 100 8.89 0.19 0.92 4.90 Carpinus caroliniana 2 50 4.44 0.07 0.35 2.40 Carya laciniosa 1 25 2.22 1.44 6.97 4.60 Cercis canadensis 1 25 2.22 0.06 0.30 1.26 Cornus florida 1 25 2.22 0.06 0.31 1.27 Fraxinus americana 1 25 2.22 0.15 0.73 1.48 Lindera benzoin 1 25 2.22 0.01 0.07 1.15 Liriodendron tulipifera 4 100 8.89 4.42 21.35 15.12 Magnolia tripetala 2 50 4.44 0.07 0.35 2.40 Oxydendrum arboreum 1 25 2.22 0.27 1.30 1.76 Quercus alba 6 150 13.33 7.76 37.54 25.44 Quercus coccinea 1 25 2.22 3.41 16.51 9.37 Ulmus rubra 6 150 13.33 0.83 4.01 8.67 Totals 1125 20.68
62
PLOT SPECIES stem count DEN (stems/ha) RDEN Sum BA (m2/ha) RBA RIV 2 Acer rubrum 6 150 27.27 0.68 3.04 15.16 Acer saccharum 2 50 9.09 0.69 3.06 6.07 Fagus grandifolia 1 25 4.55 0.73 3.26 3.90 Liriodendron tulipifera 1 25 4.55 4.12 18.35 11.45 Nyssa sylvatica 8 200 36.36 2.30 10.26 23.31 Quercus alba 1 25 4.55 5.54 24.66 14.60 Quercus prinus 2 50 9.09 2.16 9.64 9.37 Quercus velutina 1 25 4.55 6.22 27.73 16.14 Totals 550 22.45
3 Acer rubrum 12 300 38.71 1.07 4.19 21.45 Acer saccharum 2 50 6.45 2.84 11.10 8.78 Carpinus caroliniana 4 100 12.90 0.19 0.72 6.81 Carya laciniosa 1 25 3.23 1.72 6.72 4.97 Carya ovata 1 25 3.23 0.72 2.83 3.03 Cornus florida 1 25 3.23 0.36 1.40 2.31 Fraxinus americana 1 25 3.23 2.01 7.85 5.54 Hamamelis virginiana 1 25 3.23 0.05 0.19 1.71 Liriodendron tulipifera 1 25 3.23 4.12 16.08 9.65 Nyssa sylvatica 1 25 3.23 0.65 2.54 2.88 Ostrya virginiana 2 50 6.45 0.19 0.75 3.60 Quercus alba 2 50 6.45 3.72 14.52 10.49 Quercus prinus 2 50 6.45 7.96 31.10 18.77 Totals 775 25.61
63
PLOT SPECIES stem count DEN (stems/ha) RDEN Sum BA (m2/ha) RBA RIV 4 Acer rubrum 6 150 26.09 1.63 6.97 16.53 Acer saccharum 5 125 21.74 1.00 4.28 13.01 Carya laciniosa 2 50 8.70 1.69 7.25 7.97 Carya ovata 1 25 4.35 0.75 3.24 3.79 Carya sp. 1 25 4.35 0.83 3.54 3.94 Quercus alba 6 150 26.09 16.58 71.14 48.61 Tilia americana 1 25 4.35 0.81 3.47 3.91 Ulmus rubra 1 25 4.35 0.03 0.12 2.23 Totals 575 23.31
5 Acer rubrum 2 50 5.88 0.26 1.03 3.46 Acer saccharum 18 450 52.94 5.52 21.93 37.43 Carpinus caroliniana 1 25 2.94 0.06 0.22 1.58 Carya ovata 1 25 2.94 0.83 3.28 3.11 Carya sp. 1 25 2.94 1.55 6.16 4.55 Cercis canadensis 1 25 2.94 0.08 0.30 1.62 Euonymus sp.(cultivar) 3 75 8.82 0.22 0.86 4.84 Fraxinus americana 1 25 2.94 0.55 2.18 2.56 Nyssa sylvatica 1 25 2.94 2.91 11.56 7.25 Platanus occidentalis 1 25 2.94 1.34 5.31 4.13 Quercus rubra 1 25 2.94 2.07 8.24 5.59 Quercus sp. 1 25 2.94 8.76 34.81 18.88 Ulmus rubra 2 50 5.88 1.04 4.12 5.00 Totals 850 25.17
64
PLOT SPECIES stem count DEN (stems/ha) RDEN Sum BA (m2/ha) RBA RIV 6 Acer rubrum 5 125 17.86 1.40 5.91 11.88 Acer saccharum 6 150 21.43 6.89 29.17 25.30 Amelanchier arborea 1 25 3.57 0.08 0.35 1.96 Carpinus caroliniana 1 25 3.57 0.07 0.30 1.94 Carya tomentosa 1 25 3.57 0.76 3.23 3.40 Fraxinus sp. 2 50 7.14 0.08 0.33 3.74 Hamamelis virginiana 1 25 3.57 0.02 0.10 1.83 Liriodendron tulipifera 4 100 14.29 4.21 17.80 16.04 Nyssa sylvatica 3 75 10.71 0.20 0.85 5.78 Quercus alba 4 100 14.29 9.92 41.97 28.13 Totals 700 23.62
7 Acer saccharum 13 325 59.09 10.62 36.99 48.04 Carya ovata 1 25 4.55 0.87 3.04 3.80 Cercis canadensis 1 25 4.55 0.01 0.04 2.29 Juglans nigra 1 25 4.55 1.73 6.03 5.29 Quercus alba 4 100 18.18 9.73 33.90 26.04 Quercus velutina 2 50 9.09 5.74 19.99 14.54 Totals 550 28.72
65
PLOT SPECIES stem count DEN (stems/ha) RDEN Sum BA (m2/ha) RBA RIV 8 Acer rubrum 1 25 2.50 1.08 2.60 2.55 Acer saccharum 26 650 65.00 3.86 9.34 37.17 Carya cordiformis 1 25 2.50 1.83 4.41 3.46 Liriodendron tulipifera 1 25 2.50 2.53 6.12 4.31 Nyssa sylvatica 2 50 5.00 0.57 1.38 3.19 Ostrya virginiana 1 25 2.50 0.03 0.08 1.29 Pinus strobus 2 50 5.00 14.56 35.18 20.09 Quercus alba 3 75 7.50 12.94 31.27 19.39 Tilia americana 3 75 7.50 3.98 9.62 8.56 Totals 1000 41.38
66 Appendix E QUAD SURVEY HERB SPECIES AND COVER DATA
Num Sum Scientific Name Num Plots Quads Cover Type Parthenocissus quinquefolia 8 48 111 v Tiarella cordifolia 4 42 98 f Laportea canadensis 4 29 80 f Toxicodendron radicans 6 28 77 f Viburnum dentatum 8 35 73 sh Acer rubrum 8 44 69 t Polystichum acrostichoides 7 26 61 fe Viola canadensis 5 24 57 f Viola sororia 8 33 55 f Viburnum acerifolium 7 20 54 sh Collinsonia canadensis 7 19 52 f Anemonella thalictroides 8 37 50 f Desmodium nudiflorum 5 18 49 f Aster schreberi 6 17 48 f Galium triflorum 8 26 45 f Prunus serotina 7 18 45 t Uvularia perfoliata 6 23 40 f Cimicifuga racemosa 5 18 40 f Amphicarpaea bracteata 6 17 39 f Collinsonia verticillata 7 20 37 f Impatiens capensis 5 17 37 f Geranium maculatum 5 18 33 f Asarum canadense 6 17 33 f Smilacena racemosa 5 16 33 f Dioscorea villosa 5 18 31 v Acer saccharum 8 21 29 t Potentilla simplex 6 17 29 f Trillium grandiflorum 6 13 29 f Lonicera japonica 4 14 28 v Disporum maculatum 4 13 28 f Prenanthes altissima 5 14 27 f Polemonium reptans 3 13 26 f Thelypteris hexagonoptera 4 11 26 fe Carpinus caroliniana 6 21 25 t Pilea pumila 4 16 24 f Arisaema triphyllum 6 12 24 f Adiantum pedatum 4 9 24 fe Fraxinus americana 6 14 23 t Sanicula canadensis 6 10 22 f Lindera benzoin 6 17 21 sh Osmorhiza claytonii 4 9 21 f Carex woodii 2 8 21 s Hydrangea arborescens 3 7 21 sh
67 Num Sum Scientific Name Num Plots Quads Cover Type Circaea lutetiana 4 8 19 f Hamamelis virginiana 5 8 18 t Sassafras albidum 2 7 15 t Carex cumberlandensis 2 7 15 s Disporum lanuginosum 1 5 15 f Ulmus rubra 7 15 14 t Cryptotaenia canadensis 4 9 14 f Podophyllum peltatum 5 9 14 f Smilax rotundifolia 6 10 13 v Carya cordiformis 2 4 13 t Smilax glauca 5 9 12 v Carex sp. 3 5 12 s Solidago flexicaulis 4 5 11 f Helianthus sp. 3 3 11 f Unknown herb 3 5 10 f Euonymus atropurpureus 2 4 10 sh Monarda fistulosa 3 4 10 f Panicum boscii 1 4 9 g Botrychium virginianum 3 5 8 fe Viburnum prunifolium 2 4 8 sh Galium concinnum 2 4 8 f Osmunda cinnamomea 2 4 8 fe Amelanchier arborea 4 7 7 f Scutellaria incana 3 4 7 f Polygonum virginianum 2 3 7 f Athyrium felix-femina 1 3 7 fe Rosa multiflora 2 2 7 sh Carex glaucoidea 1 2 7 s Hydrastis canadensis 1 2 7 f Rubus pensilvanicus 1 2 7 f Liriodendron tulipifera 2 7 6 t Creteagus sp. 2 4 6 t Sanguinaria canadensis 2 4 6 f Uvularia sessilifolia 2 4 6 f Fraxinus quadrangulata 1 3 6 t Carex laxiculmis var. copulata 1 2 6 s Pedicularis canadensis 1 2 6 f Bromus pubescens 1 2 6 g Unknown seedling 3 5 5 f Nyssa sylvatica 3 4 5 t Hepatica acutiloba 1 4 5 f Fraxinus biltmoreana 1 3 5 t Quercus alba 3 3 5 t Carex digitalis 2 3 5 s Celastrus scandens 2 2 5 v Carya glabra 1 2 5 t Oxydendrum arboreum 2 2 5 t
68 Num Sum Scientific Name Num Plots Quads Cover Type Quercus rubra 2 2 5 t Aristolochia serpentaria 2 2 5 f Thelypteris noveboracensis 1 1 5 fe Ampelopsis brevipedunculata 1 4 4 f Smilax hispida 2 2 4 v Carya tomentosa 1 2 4 t Galium circaezans 1 2 4 f Rubus sp. 1 2 4 f Carya ovata 1 1 4 t Cornus alternifolia 1 1 4 sh Carex sparganioides 1 1 4 s Agrimonia rostellata 1 1 4 f Phlox divaricata 2 2 3 f Pteridium aquilinum 1 2 3 fe Clematis virginiana 1 1 3 v Carex laxiflorum 1 1 3 s Aster cordifolius 1 1 3 f Aster lateriflorus 1 1 3 f Lysimachia quadrifolia 1 1 3 f Orchis spectabilis 1 1 3 f Oxalis grandis 1 1 3 f Ranunculus recurvatus 1 1 3 f Scutellaria nervosa 1 1 3 f Panicum latifolium 1 1 3 g Asplenium platyneuron 1 1 3 fe Vitis sp. 1 2 2 v Ostrya virginiana 2 2 2 t Acalypha gracilens 1 2 2 f Tree sp. 1 1 2 t Ceonothus americanus 1 1 2 sh Rosa carolina 1 1 2 sh Aralia racemosa 1 1 2 f Conopholis americana 1 1 2 f Cunila origonoides 1 1 2 f Galium aparine 1 1 2 f Luzula multiflora 1 1 2 f Medeola virginiana 1 1 2 f Taraxacum officinale 1 1 2 f Brachyeletrum erectum 1 1 2 g Grass sp. 1 1 2 g grass sp.-3 1 1 2 g Poa sp. 1 1 2 g Quercus velutina 1 1 1 t Desmodium glutinosum 1 1 1 f Erechtites heiracifolia 1 1 1 f Stellaria pubera 1 1 1 f Panicum sp. 1 1 1 g
69 Num Sum Scientific Name Num Plots Quads Cover Type Poa cuspidata 1 1 1 g Poa sylvestris 1 1 1 g Fagus grandifolia 2 2 0 t Tilia americana 2 2 0 t Athyrium thelipteroides 1 2 0 fe Smilax herbacea 1 1 0 v Cercis canadensis 1 1 0 t Vicia caroliniana 1 1 0 v