Vegetative Survey of Martha’s Meadow, an Open
Limestone Habitat in Northwestern Georgia
Martin L. Cipollini, Maya Strahl, Nicole Soper Gorden, Patricia Tomlinson, and Richard Ware
Southeastern Naturalist, Volume 12, Issue 2 (2013): 317–338
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2013 SOUTHEASTERN NATURALIST 12(2):317–338
Vegetative Survey of Martha’s Meadow, an Open
Limestone Habitat in Northwestern Georgia
Martin L. Cipollini1,*, Maya Strahl2, Nicole Soper Gorden3, Patricia Tomlinson4,
and Richard Ware5
Abstract - The vegetation of Martha’s Meadow (Berry College, Floyd County, GA) is
distinct from that of surrounding flatwood areas and appears to be floristically similar
to a limestone glade community. Limestone (cedar) glades of the southeastern United
States are characterized by high species richness and diversity, calcareous soil, and up
to 26 endemic or near-endemic indicator species. To describe the vegetation of Martha’s
Meadow and to determine its affinity to other limestone-based communities, a comprehensive
species survey was augmented by quantitative surveys in May, July, and October
of 2006. A total of 203 species in 56 families were identified, including nine limestoneassociated
species designated as rare in the state of Georgia. Juniperus virginiana, a key
species associated with glade communities, was important in both the overstory and the
understory. Other important overstory species included Pinus taeda, Quercus shumardii,
Q. muehlenbergii, and Ostrya virginiana. The perennial grass Danthonia spicata, the
southeastern sedge Carex cherokeensis, and the herb Verbesina virginica, which is commonly
associated with alkaline soils, were among the most important understory species,
but no well-recognized limestone glade endemics were found. Two invasive grasses
(Festuca subverticillata and Microstegium vimineum) were also among the most important
understory species. Several ordination and clustering methods were used to compare
the community structure of Martha’s Meadow with data sets derived from other open
calcareous and non-calcareous habitats throughout the southeastern and mid-western
United States. Regardless of method, Martha’s Meadow appeared to be most similar to
several well-recognized Georgia limestone glades. While Martha’s Meadow has some
characteristics of edaphically determined limestone glades, it also lacks some characteristic
traits (e.g., key limestone endemic species), making definitive classification tenuous.
Evidence of on-going succession involving woody species suggests that the site might
be best classified as a xeric limestone prairie (barrens) that requires disturbance or active
management to maintain canopy openness and understory diversity.
The terms cedar glade, limestone glade, limestone cedar glade, deep-soil
barren, xeric limestone prairie, and xeric limestone barrens all refer to scattered
areas in the eastern half of the United States that contain thin, calcium-rich soils
and high herbaceous plant cover (Baskin and Baskin 2004, Baskin et al. 2007,
Mann et al. 1999, McClain and Ebinger 2002, McDearman 2004). All are characterized
by sparse to nonexistent woody vegetation cover, high grass cover,
1Department of Biology, Berry College, Mount Berry, GA 30149. 2Cold Spring Harbor
Laboratory, Cold Spring Harbor, NY 11724. 3Department of Biology, University of Massachusetts,
Amherst, MA 01003. 4Department of Biology, Berry College, Mount Berry,
GA 30149 (deceased). 5Georgia Botanical Society, Rome, GA 30165. *Corresponding
author - firstname.lastname@example.org.
318 Southeastern Naturalist Vol. 12, No. 2
and high herbaceous plant diversity. They occur in unglaciated areas, with soils
that tend to be too shallow, dry, and nutrient-poor to allow the development of
extensive root systems, thus excluding most woody species. Vegetation type and
density is correlated with bedrock exposure and soil-depth gradients, giving rise
to patches of vegetation types ranging from treeless, grass-dominated to open
woodland habitats (Quarterman 1950a, b).
True southeastern limestone glades are considered to be mainly edaphic
communities, with their distinctive vegetation characteristics resulting from
dry, high-calcium, shallow soils, and high light levels resulting from a minimal
woody overstory (Baskin and Baskin 1988, 2003, 2004; Baskin et al. 2007). Climate
does not appear to be the main determining factor of species composition;
many species are common and have wide ranges throughout the eastern, southeastern,
and midwestern United States (Baskin and Baskin 2003). Because they
are, by definition, mainly determined by existing soil characteristics, true limestone
glades appear to be restricted to their original boundaries and do not require
disturbances for maintenance of canopy openness (Baskin and Baskin 1988,
2003). The edaphic nature of such glades seems to preclude particular species
from exerting substantial control over community composition. Along with other
types of forest openings, they are important habitats for rare and endemic species
(Copenheaver et al. 2004). Fire may play a role in maintaining some limestone
communities, since some putative limestone “glades” may not be maintained
strictly by edaphic conditions. McDearman (2004), for example, claims that fire
affects and perhaps even determines true limestone glades by selecting against
woody species that may otherwise colonize glade edges. Likewise, Baskin et al.
(2007) argue for a dichotomy between primarily edaphically determined limestone
glades (cedar glades) and primarily disturbance-maintained xeric limestone
prairies (rocky limestone barrens) in areas outside of the central Tennessee basin,
a summary of which can be found in Table II of their paper.
Martha’s Meadow is a small area located on limestone-based soil on the Berry
College campus (Mount Berry, Floyd County, GA). Its open-canopied overstory
contains several woody species commonly found in limestone habitats, including
Juniperus virginiana L. (Eastern Redcedar), Quercus muehlenbergii Engelm.
(Chinkapin Oak), and Q. shumardii Buckl. (Shumard’s Oak). At the time of
this study, the site contained a more or less continuous herbaceous and grassy
understory. Preliminary informal floristic surveys that found several protected
limestone-affiliated species (see Results) led to permanent protection of the site
by the college as a putative limestone glade in 2003. Many similar small, isolated
limestone sites occur throughout northwestern Georgia, northeastern Alabama,
and eastern Tennessee and harbor variable numbers of endangered or rare species
(e.g., sites associated with, but discontinous from, the Coosa Valley Prairies;
GADNR 2012). Questions concerning such small, but botanically important
sites include whether management (fire, mowing, canopy removal, etc.) might
be necessary to maintain plant community characteristics, or whether protection
alone (or management restricted to invasive species control) might be sufficient.
Understanding the relationship of small, isolated sites to larger and better-studied
sites may provide insight into their origin, floristic make-up, and management.
2013 M.L. Cipollini, M. Strahl, N. Soper Gorden, P. Tomlinson, and R. Ware 319
This paper, focusing on the plant community at Martha’s Meadow, represents
a case study of one such site. The purposes of this study were: a) to determine
the species composition of the site, b) to make a quantitative assessment of the
canopy and understory strata, c) to compare its species composition to data sets
from other limestone glade type sites within the southeastern US as a means of
determining its floristic relationship to those sites, and d) to provide baseline data
necessary to guide long-term management for this and similar sites.
Field Site Description
Martha’s Meadow (34.2965°N, 85.1995°W, elevation 180–190 m) is a small
(≈1 ha) natural site located on Berry College’s main campus in Mount Berry, GA.
Because of its proximity to a former recreational lake (Victory Lake), the area
had been periodically mowed in the past and was likely subject to occasional foot
traffic at various times since the lake was established in the 1920s. Maintenance
and foot traffic ceased when the adjacent lake area drained into a karst sinkhole
and the depression was filled in the late 1990s. The site was identified as being
floristically distinct from surrounding areas, was discovered to harbor several
rare limestone-affiliated species, and was protected as a putative limestone glade
in 2003. Management of the site to maintain plant community composition was
proposed, and the site was mown in the fall of 2004 in an attempt to control what
was interpreted as encroaching woody species. Soils at the site are Townley and
Cunningham silt loams (2–10% slope; Soil Survey Staff, NRCS 2011), which
consist of thin layers of well-drained loam over silty or shaly clays, with about
0.5 to 1 m to paralithic bedrock. Mean annual precipitation at the site is ≈1.3 m,
mean annual air temperature is ≈15 °C, and the frost-free perio d is ≈190 days.
General species survey
A general species survey was conducted from 2002 to 2006. Periodically
throughout each growing season, the area was thoroughly searched for flowering
species not previously recorded, which were added to a running list of species.
Quantitative species surveys
Quantitative surveys were conducted in May, July, and October of 2006 using
a modified version of the North Carolina Vegetation Survey Protocol (Peet et al.
1998). A single 50- x 70-m plot was located at the center of the meadow. The plot
was divided into thirty-five 10- x 10-m modules. At the southeast corner of each
10- x 10-m module, a 1- x 1-m quadrat was established.
Because the area lacked substantial shrub or subcanopy layers, the vegetation
on the site was divided into just two strata: the understory included all plants
less than 1.4 m in height, and the overstory included all trees, shrubs, and vines ≥1.4 m
in height. The understory survey was conducted by visually estimating percent
cover for all species in the 1- x 1-m quadrats during each census period. All
species were assigned either a known scientific name or a descriptive name until
320 Southeastern Naturalist Vol. 12, No. 2
the species could be identified, and herbarium specimens were taken unless the
species was thought to be rare or endangered. Cover estimated as being less than
1.0% was recorded as 0.5% for data-analysis purposes.
For all understory species, mean cover (C; sum of cover values) and frequency
(F; total modules containing the species) values were calculated for each census.
Total C (TC; sum of all cover values), and total F (TF; sum of all frequency
values) values were calculated and used to determine relative cover (RC) and
frequency (RF) values for each species. Importance values (IV) were then calculated
as the sum of RC and RF values for each species.
For the overstory, all individuals were located in summer 2005, their Cartesian
coordinates within each 10- x 10-m module recorded to the nearest 0.01 m,
and circumferences at breast height (CBH) measured to the nearest 0.01 m. For
each species, absolute and relative density (RD), frequency, cover (basal area)
values, and importance values (IV = RD + RF + RC) were calculated. Percent
canopy cover was also estimated in each 1- x 1-m quadrat, and used to estimate
canopy cover across the entire plot.
Species identifications and ecological characteristics were determined by consulting
Brown and Kirkman (1990), Dean et al. (1973), Duncan and Foote (1975),
Flora of North America Editorial Committee (1993+), Georgia Department of
Natural Resources - Wildlife Resources Division (2011), Hitchcock and Chase
(1951), Knobel (1980), Newcomb (1977), Pohl (1968), Preston (1989), Radford
et al. (1968), University of Texas at Austin (2011), University of Wisconsin - Stevens
Point (2011), and Weakley (2011). Final species names and codes followed
the United States Department of Agriculture, Natural Resource Conservation
Service (2011) PLANTS Database. Dried specimens were deposited in the Berry
College Department of Biology’s herbarium.
Comparative data analysis
Comprehensive species lists were obtained from 21 studies of sites characterized
as limestone/cedar glades or barrens in Alabama, Georgia, Illinois,
Kentucky, Tennessee, and West Virginia (see Table 1 for site descriptions and
references). Data sets included data from single intensively surveyed sites as well
as comprehensive lists for habitat types within certain states (e.g., Georgia limestone
glades). The latter data sets help root the resulting ordinations. To further
root the ordinations, four data sets from non-limestone grassy communities in
Georgia, including a mountain Pinus palustris Mill. (Longleaf Pine) forest with
acidic sandstone soils on the Berry College campus, were used as outgroups. The
Community Analysis Package (CAP; Pisces Conservation, Ltd. 2004) program
was used to conduct reciprocal averaging (RA, also known as correspondence
analysis) ordination followed by a divisive cluster analysis (TWINSPAN), based
upon species presence/absence data for each site. Reciprocal averaging ordination
is good for examining patterns in overall species similarity in data sets taken
across environmental gradients, but can be disproportionately influenced by rare
species. For the RA analysis reported here we down-weighted rare species by
selecting that option in CAP (although results did not differ markedly from those
when species were weighted equally). Non-metric multidimensional scaling
2013 M.L. Cipollini, M. Strahl, N. Soper Gorden, P. Tomlinson, and R. Ware 321
ordinations (NMDS) produced qualitatively similar patterns as did RA, but we
report RA results here due to their use in TWINSPAN clustering. TWINSPAN
produces a cluster diagram based upon an RA ordination by roughly dividing the
sites into groups that score high and low on the first axis, and then continuing this
process until all groups are successively divided. We chose a level of division
so that the smallest cluster produced included two sites. At each successive division,
TWINSPAN identifies species strongly associated with the two branches
that were produced, as well as those not strongly associated with either branch.
Finally, as an alternative to TWINSPAN divisive clustering, we conducted an
agglomerative cluster analysis in CAP (Bray-Curtis average linkage method)
using the same 25 datasets. Because such species were not surveyed thoroughly
Table 1. Data sets used in the comparative analysis of Martha’s Meadow with limestone glade type
and non-limestone communities in the southeastern United States.
Code Data set description
AL1 Alabama limestone glades comprehensive listA
AL2 Northern Alabama limestone glades, Moulton ValleyB
AL3 Northern Alabama limestone glades, Little MountainB
AL4 Northern Alabama limestone glades, Tennessee ValleyB
AL5 Northern Alabama limestone glades, Jackson County Mountains B
AL6 Northern Alabama limestone glades, Sequatchi ValleyB
GA1 Georgia limestone glades comprehensive listA
GA2 Georgia, Chickamauga National Battlefield Plot 16 C
GA3 Georgia, Chickamauga National Battlefield Plot 19 C
GA4 Georgia, Chickamauga National Battlefield Plot 21 C
GA5 Georgia, Chickamauga National Battlefield Plot 51 C
GN1 Georgia, Thunder Mountain Longleaf Pine-Post Oak-Black-seeded Needlegrass communityD
GN2 Georgia, Thunder Mountain Longleaf Pine-Turkey Oak-Goat's Rue communityD
GN3 Georgia, Thunder Mountain Mockernut Hickory-Post Oak-Yellow Passion Flower communityD
GN4 Georgia, Berry College Mountain Longleaf Pine Management AreaE
IL1 Southern Illinois limestone glades comprehensive list F
IL2 Southern Illinois limestone barrens comprehensive list F
KY1 Kentucky, Blue Licks Battlefield State Park G
MM Georgia, Martha's Meadow comprehensive species list H
TN1 Tennessee limestone glades comprehensive listA
TN2 Middle Tennessee cedar gladeB
TN3 Middle Tennessee cedar glade, Grindelia lanceolata communityI
WV1 West Virginia limestone barrens comprehensive listJ
WV2 West Virginia cedar glades comprehensive listJ
WV3 West Virginia glade woodlands comprehensive listJ
ABaskin and Baskin 1975; Baskin et al. 1968.
BBaskin and Baskin 1995; Baskin et al. 1995.
CThomas Govus, Ellijay, GA, unpubl. data.
DCarter and Londo 2006.
EMartin Cipollini, Mount Berry, GA, unpubl. data.
FHiekens and Robertson 1994.
GBaskin and Baskin 1985a.
IBaskin and Baskin 1996.
322 Southeastern Naturalist Vol. 12, No. 2
in Martha’s Meadow, we did not include bryophytes, lycophytes, pteridophytes,
or lichens that appeared commonly in other species lists.
In August 2005, soil-core samples (10 cm depth) were taken from each of
10 randomly selected 10- x 10-m modules. These samples were combined into
a single bulk sample representing the entire plot. The bulk sample was sent to
Georgia Cooperative Extension Service (Athens, GA) for analysis of available
Ca, K, Mg, Mn, NO3-N, P, and Zn (lbs/acre), as well as pH-CaCl2 extract, pHwater
extract, and lime buffer capacity (LBC = ppm CaCO3/pH unit; a measure
of a soil’s resistance to increases in pH).
General species survey
The general survey identified 56 families, 142 genera, and 203 species within
Martha’s Meadow (Appendix 1). The four families containing the most species
were Poaceae (25 species), Asteraceae (25 species), Fabaceae (20 species), and
Rosaceae (9 species). Ten of these species are considered rare (species of special
concern) in the state of Georgia (Table 2; GA DNR 2011), and nine of these rare
species have known affiliations with calcareous or high pH soils. Twenty-eight
(13.8%) species are not native to North America (USDA, NRCS 2011), and five
of these species are considered invasive in natural habitats in Georgia (Elaeagnus
umbellata Thunb. [Autumn Olive], Lespedeza cuneata [Dum.-Cours.] G.
Don [Sericea Lespedeza], Ligustrum sinense Lour. [Chinese Privet], Lonicera
japonica Thunb. [Japanese Honeysuckle], and Microstegium vimineum (Dum.-
Cours.) G. Don [Nepalese Browntop]; Georgia Exotic Pest Plant Council 2012).
Table 2. Plant species found in Martha’s Meadow and considered rare in the state of Georgia (for
a key to rarity rankings, see GA DNR 2011). All species are affiliated with calcareous or high pH
soils except C. catesbiana ssp. sericata. Authorities, common names, and species codes can be
found in Appendix 1.
Species GA DNR List Rarity rankings
Asclepias hirtella (Pennell) Woodson (Green Milkweed) Special Concern G5, S2
Asclepias viridiflora Raf. (Green Comet Milkweed) Watch List G5, S3
Asclepias viridis Walt (Green Antelopehorn) Watch List G4G5, S3
Calystegia catesbeiana Pursh ssp. sericata (House) Special Concern G3T2T3Q, S2S3
Brummitt (Catesby’s False Bindweed)
Carex meadii Dewey (Mead’s Sedge) Watch List G4G5, SU
Delphinium tricorne Michx. (Dwarf Larkspur) Special Concern G5, S2S3
Prenanthes barbata (Torr. & Gray) Milstead Special Concern G3, S2
Scutellaria parvula Michx. var. missouriensis (Torr.) Special Concern G4Q, S2
Goodman & C.A. Lawson (Leonard’s Skullcap)
Sida elliottii Torr. & Gray (Elliott’s Fanpetals) Special Concern G4G5, S2
Thaspium barbinode (Michx.) Nutt. [=chapmanii] Special Concern GNR, S3
2013 M.L. Cipollini, M. Strahl, N. Soper Gorden, P. Tomlinson, and R. Ware 323
Quantitative understory survey
A total of 141 species were identified in the quantitative understory surveys.
During the spring, summer, and fall surveys, 59, 127, and 72 species were identified,
respectively. The IVs were relatively small and similar due to the high
number of species identified. The herbaceous or grass-like species Danthonia
spicata (L.) Beauv. Ex Roemer & Schultes (Poverty Oatgrass), Carex cherokeensis
Schwein. (Cherokee Sedge), and Verbesina virginica L. (White Crownbeard)
were among the most important species in all three censuses, as were seedlings
of the trees Celtis laevigata Willd. (Sugarberry) and J. virginiana (Table 3).
Pinus taeda L. (Loblolly Pine) seedlings were abundant in the spring, but fell
off slightly during a drought in early summer to 11th in importance in summer
Table 3. Ten most important species from the quantitative understory surveys in 2006. Asterisks
denote introduced (non-native) species, and species are sorted in descending order of overall importance
Danthonia spicata 0.27
Festuca subverticillata 0.22
Verbesina virginica 0.15
Carex cherokeensis 0.12
Salvia lyrata L. (Lyreleaf Sage) 0.09
Viola triloba Schwein. (Three-Lobe Violet) 0.06
Juniperus virginiana 0.06
Celtix laevigata 0.06
Pinus taeda 0.05
Sanicula canadensis L. (Canadian Blacksnakeroot) 0.05
Danthonia spicata 0.21
Carex cherokeensis 0.18
Verbesina virginica 0.16
Festuca subverticillata 0.09
Microstegium vimineum 0.06
Scleria oligantha Michx. (Littlehead Nutrush) 0.06
Salvia lyrata 0.05
Juniperus virginiana 0.05
Ostrya virginiana 0.05
Celtis laevigata 0.04
Danthonia spicata 0.31
Carex sp. 0.16
Verbesina virginica 0.14
Festuca subverticillata 0.13
Carex cherokeensis 0.07
Juniperus virginiana 0.07
Celtis laevigata 0.05
Smilax bona-nox L. (Saw Greenbriar) 0.05
Tridens flavus (L.) A.S. Hitchc. (Purpletop Tridens) 0.04
Microstegium vimineum* 0.04
324 Southeastern Naturalist Vol. 12, No. 2
and fall seasons. A widespread and sometimes invasive native grass Festuca subverticillata
(Pers.) Alexeev (Nodding Fescue) was among the most important in
all three seasons, and the introduced invasive grass M. vimineum was among the
most important in the summer and fall censuses.
Quantitative overstory survey
In the overstory survey, 123 individuals and 13 species were identified
(Table 4). The five species with the highest importance values were J. virginiana,
P. taeda, Q. shumardii, Q. muehlenbergii, and Ostrya virginiana (P. Mill.)
K. Koch (Hophornbeam). Percent canopy cover, based upon surveys of 1- x 1-m
quadrats, showed high variability with a range from 0 to 270% (average = 88.7%,
standard deviation = 68.6). This pattern reflects a patchy openness associated
with scattered, but large canopy trees.
Comparative data analysis
The ordination data matrix included 826 species among the 25 sites (see
Supplemental Appendix, available online at https://www.eaglehill.us/SENAonline/
suppl-files/s12-2-1141-Cipollini-s1, and, for BioOne subscribers, at http://
dx.doi.org/10.1656/S1141.s1). On the first axis of the RA plot, Martha’s Meadow
grouped with limestone glades in Georgia, West Virginia, and Illinois (Fig. 1).
On the second axis, Martha’s Meadow fell between the four non-limestone sites
in Georgia and four of the limestone sites in Alabama. As a means of interpreting
biological patterns in the RA, we examined the 30 species scoring the highest or
lowest on each of the first two RA axes:
1) Species scoring low on the first axis and associated with limestone sites
in Tennessee and Alabama tended to have western, mid-western, or widespread
distributions. These species tended to be annual/perennial herbs,
Table 4. Summary of the 2005 quantitative overstory survey. See text for variable definitions; species
are sorted in descending order of overall importance value.
Scientific name D F BA RD RF RC IV
J. virginiana 28 18 18.40 0.23 0.30 0.30 0.82
P. taeda 23 12 15.79 0.19 0.20 0.26 0.64
Q. shumardii 8 7 14.70 0.07 0.12 0.24 0.42
O. virginiana 34 3 1.06 0.28 0.05 0.02 0.34
Q. muehlenbergii 7 5 3.95 0.06 0.08 0.06 0.20
C. canadensis 6 4 0.24 0.05 0.07 0.00 0.12
Acer leucoderme Small 7 3 0.50 0.06 0.05 0.01 0.11
Q. stellata 1 1 3.53 0.01 0.02 0.06 0.08
Diospyros virginiana L. 3 3 0.25 0.02 0.05 0.00 0.08
F. americana 2 2 1.70 0.02 0.03 0.03 0.08
Unknown species 2 1 0.56 0.02 0.02 0.01 0.04
Quercus nigra L. (Water Oak) 1 1 0.61 0.01 0.02 0.01 0.03
V. rufidulum 1 1 0.03 0.01 0.02 0.00 0.03
Total 123 61 61.31
2013 M.L. Cipollini, M. Strahl, N. Soper Gorden, P. Tomlinson, and R. Ware 325
and included a number of graminoid species, introduced species (some
apparently from the western US), and some species associated exclusively
with limestone glades.
2) Species scoring high on the first axis, and associated with the four nonlimestone
sites tended to be woody species with southeastern or eastern
distributions that are most often found in dry, open, and acidic sites.
3) Species scoring low on the second axis and associated with Georgia and
Alabama sites (both limestone and non-limestone), tended to be herbs with
eastern and southeastern distributions and preferences for moist open sites.
4) Species scoring high on the second axis and associated with Georgia,
Illinois, and West Virginia limestone sites tended to be perennial/biennial
herbs with eastern and northeastern distributions. A number of these species
are commonly found in barrens, glades, and rocky or limest one sites.
Based upon this interpretation of the RA analysis, Martha’s Meadow could be
said to nest with “limestone” sites on the first axis, and “southern or southeastern”
sites on the second axis.
The TWINSPAN cluster analysis had Martha’s Meadow nested with three
of the Georgia glades in Georgia on a main “limestone” clade containing the
other Georgia glades as well as limestone sites in Tennessee, Alabama, and Kentucky
(Fig. 2). In this analysis, the four non-limestone sites in Georgia grouped
most closely with the two limestone sites from Illinois. Of the 15 species most
characteristic of the main limestone clade, 10 were found in Martha’s Meadow,
and of the top 96 such species, 41 were found in Martha’s Meadow. Limestone
species of concern in the state of Georgia (GA DNR 2011) that were characteristic
of the main limestone clade, yet lacking in Martha’s Meadow included
Dalea gattingeri (Heller) Barneby (Purpletassels), Delphinium carolinianum
Walt. ssp. calciphilum (Carolina Larkspur), Eleocharis compressa Sullivant
(Flatstem Spikerush), Heliotropium tenellum (Nutt.) Torr. (Pasture Heliotrope),
Figure 1. Reciprocal averaging ordination plot showing the first two RA axes. Martha’s
Meadow is represented by “MM”. For other site codes, see Table 1.
326 Southeastern Naturalist Vol. 12, No. 2
Hypericum dolabriforme Vent. (Straggling St. Johnswort), Hypericum sphaerocarpum
Michx. (Roundseed St. Johnswort), Pediomelum subacaule (Torr. &
Gray) Rydb. (Whiterim Scurfpea), and Viola egglestonii Brainerd (Glade Violet).
Species lacking in Martha’s Meadow, but present in the nearest Georgia glade
(Chickamauga National Battlefield plot 16) included 21 species strongly associated
with limestone soils. Of these, five are listed as rare limestone species in
the state of Georgia (D. carolinianum ssp. calciphilum, Erigeron strigosus Muhl.
ex Willd. var. calcicola J . Allison (Prairie Fleabane), H. dolabriforme, P. subacaule,
and V. egglestonii).
The agglomerative cluster analysis showed qualitatively similar results to the
RA and TWINSPAN analyses, with Martha’s Meadow most closely affiliated with
two of the Georgia glades in a clade that included the non-limestone sites in Georgia,
along with limestone sites in West Virginia and Illinois (Fig. 3). The other main
clade included all other limestone sites and the three remaining Georgia glades
(two of which were closely related to the clade containing Martha’s Meadow).
Thus, Martha’s Meadow shows a mix of both biogeographic (southern and southeastern
region) and site-specific (limestone site) species affinities.
Soil test results (Table 5) based upon a single bulked sample show very high
available Ca and Mg, low P and K, circumneutral pH, and a very low LBC, indicative
of limestone-based soils.
The three most commonly occurring families identified in the general survey
of Martha’s Meadow were Asteraceae, Poaceae, and Fabaceae. High species richness
of Asteraceae and Poaceae is associated with xeric limestone prairies, cedar
glades, and deep-soil barrens (Lawless et al. 2004). Fabaceae is usually ranked as
Figure 2. Twinspan cluster plot derived from the RA analysis. Minimum cluster size was
set at two. Martha’s Meadow is represented by “MM”. For other site codes, see Table 1.
2013 M.L. Cipollini, M. Strahl, N. Soper Gorden, P. Tomlinson, and R. Ware 327
the third most common family in xeric limestone prairies and deep soil barrens,
whereas Cyperaceae usually ranks before Fabaceae in true cedar glades.
Juniperus virginiana had the highest importance value in the overstory survey.
This species is associated with essentially all types of glade habitats (McClain and
Ebinger 2002), although is certainly not restricted to them. Because of hardwood
competition, J. virginiana is rarely dominant in other habitat types (Lawson 1990).
This species usually dominates the sparse canopy strata and areas surrounding
cedar glades because it is adapted to the shallow, dry, nutrient-restricted soils associated
with glades, whereas other native trees are less so.
Some other woody species are associated with pockets of deeper soil and on
edge habitats surrounding cedar glades. For example, Baskin and Baskin (1985a)
mention Cercis canadensis L. (Eastern Redbud), Cornus florida L. (Flowering
Figure 3. Agglomerative cluster plot using average linkage and Bray-Curtis similarity
estimates. Martha’s Meadow is represented by “MM”. For other site codes, see Table 1.
Table 5. Test results for a bulked soil sample taken from Martha’s Meadow in 2005. Units for
minerals are kg/ha rounded to the nearest 10 kg/ha. Results for duplicate samples taken from the
bulked soil sample were identical for all tests.
pH (CaCl2) 6.1
pH (water) 6.7
LBC (ppm CaCO3/pH unit) 291
328 Southeastern Naturalist Vol. 12, No. 2
Dogwood), and Viburnum rufidulum Raf. (Rusty Blackhaw) as the most important
woody species occurring within a Kentucky glade and J. virginiana,
Q. muehlenbergii, Q. stellata Wangenh. (Post Oak), Q. velutina Lam. (Black
Oak), C. canadensis, Cornus drummondii C.A. Mey. (Roughleaf Dogwood),
Fraxinus americana L. (White Ash), and V. rufidulum as the most important edge
species. Four of these eight species (C. canadensis, Q. muehlenbergii, Q. stellata,
and F. americana) occur in Martha’s Meadow. Cercis canadensis tends to
inhabit moist, well-drained soils and is often found on limestone bluffs (University
of Texas at Austin 2011). Quercus muehlenbergii is associated with basic,
well-drained or dry limestone-derived soils and limestone outcrops, although it
can also grow in moist areas; in dry limestone soils, it may be a climax species
(Tirmenstein 1991). Quercus stellata and F. americana both tend to be associated
with dry to mesic, acidic soils, although they do show some tolerance of high-Ca
soils (University of Texas at Austin 2011). Quercus shumardii, another important
oak species in Martha’s Meadow, is a more adaptable and widespread oak, and
can be found on a wide variety of (often moist) soil types including calcareousbased
soils (Edwards 1990).
Although small in size, Martha’s Meadow is rich in both woody and herbaceous
native flora, containing over 200 species. The floristic surveys we used to
compare with Martha’s Meadow were located throughout the southeastern United
States, but the size and number of the areas sampled varied, as did the datacollection
methods, making it difficult to compare species richness directly. This
difference in methodology represents a caveat to our comparative site analysis.
Total species numbers in the sites with which we compared Martha’s Meadow
ranged from 54 in a single limestone glade in the Central Basin of Tennessee
(Baskin and Baskin 1996) to 269 in a survey of twenty-two glades in Alabama
(“Alabama limestone glades”; Baskin et al 1995). The long-term and intensive
manner of the data collection in Martha’s Meadow may explain why more species
were identified there than in most studies of single cedar glades (e.g., Baskin and
Baskin 1977, 1985a). Regardless, Martha’s Meadow shows the greatest overall
similarity to several recognized limestone glades in Chickamauga Battlefield
National Park, Walker County, GA, roughly 80 km due north of Berry College
(Allison 1991). Martha’s Meadow is, on the other hand, floristically dissimilar to
an open-canopied mountain Longleaf Pine site located on acidic sandstone soils
located less than 6 km away.
Our data suggest that Martha’s Meadow is floristically close to recognized
limestone or cedar glades, especially those in Georgia. A high species richness
and diversity, the presence of many species with known calcareous soil
affiliations, the characteristic soil (relatively shallow, dry, clayey, circumneutral
pH, high Ca and Mg, low P and N), and a tree canopy dominated by
J. virginiana and other trees commonly associated with limestone sites suggest
that Martha’s Meadow is a limestone glade community. In addition, the
site groups with other limestone glades in Georgia in both the ordination and
the cluster analyses, and Martha’s Meadow contains high abundances of species
commonly present in other glades. Of 55 species of concern in the state
2013 M.L. Cipollini, M. Strahl, N. Soper Gorden, P. Tomlinson, and R. Ware 329
of Georgia associated with limestone glade habitats, eight were recorded at
Martha’s Meadow (GA DNR 2011). The range is 4–12 such species for the
other Georgia glade data sets referenced in this study, so Martha’s Meadow is
not unusual in that regard.
On the other hand, some results and observations make it difficult to argue that
Martha’s Meadow is a true limestone glade. None of the species identified in Martha’s
Meadow are considered indicator species of limestone glades (Baskin and
Baskin 1988). Such species are often referred to as “limestone glade endemics”,
and, while not being entirely confined to limestone glades, are very commonly
found in them. For example, Sporobolus vaginiflorus (Torr. ex A. Gray) Alph.
Wood (Poverty Dropseed), which occurs in species lists of glade flora throughout
the range of the habitat type and has been repeatedly cited as a significant
component of southeastern glade flora, was not recorded in the site. Examples of
Georgia-designated species of concern with limestone affiliations missing from the
site include H. dolabriforme and E. compressa, which were found in all and four
out of five of the other Georgia glades, respectively. The presence of large, mature
specimens of oak and pine trees suggests that the soil is not thin enough to deter
mesophytic species from encroaching in the area. In fact, the amount of canopy
cover present (nearly 90% on average) in Martha’s Meadow may be a key reason
for its lack of endemics. In studies conducted by Baskin and Baskin (1985b, 1988),
high light exposure was proposed to be the key factor influencing the presence of
viable communities of glade endemics. Martha’s Meadow has a fair number of mature
trees, and although most of the reviewed floristic surveys did not specifically
mention quantitative tree cover, it is likely that Martha’s Meadow has a higher tree
cover than most true glade habitats. The species that grouped Martha’s Meadow
with the Georgia glades on the first axis of the RA ordination were mostly common,
wide-ranging species found throughout the southern and, in some cases, the entire
eastern half of North America. Species that separated the site from true limestone
glades in GA tended to be those associated commonly with limestone habitats.
Finally, woody plant seedlings were present throughout the understory, indicating
that the area had been undergoing succession following the 2004 mowing.
Indeed, informal follow-up surveys of woody species in 2009 showed significant
encroachment by P. taeda and J. virginiana seedlings and saplings, suggesting
that frequent fire (or historical mowing) is a key factor in maintaining the relative
openness of the area. As a result of these observations, the area was burned by prescription
in the fall of 2010 and most tree seedlings and saplings were eliminated
from the site.
Taking all factors into consideration, Martha’s Meadow is most akin to a
xeric limestone prairie (rocky limestone barren), based upon the classification
presented by Baskin et al. (2007). According to this system, factors that preclude
Martha’s Meadow from inclusion as a true limestone glade of primarily edaphic
330 Southeastern Naturalist Vol. 12, No. 2
1) moderately deep soils with little exposed rock,
2) Fabaceae more important than Cyperaceae,
3) perennial grasses and herbs more dominant than annuals,
4) no pronounced aspect dominance by spring-flowering plants,
5) cryptogams and cyanobacteria unimportant,
6) high grass cover, low herb cover, and moderate (rather than low) tree
7) dominance of plant species with southern distributions (as opposed to
more northern/western distributions),
8) few endemics/near endemics,
9) evidence that changes in community composition are underway, suggesting
that management will be necessary to maintain openness, and
10) possible origin as an anthropogenic disclimax due to human traffic,
mowing, and possible fire management in the past.
It is possible that Martha’s Meadow has low numbers of characteristic limestone
taxa due to its small size, isolation, and likely anthropogenic impacts due to its
proximity to a former recreational lake and a college campus. Moreover, the presence
of aggressive grasses (e.g., F. subverticillata and M. vimineum) could have
competitively displaced a number of characteristic species formerly present on the
site. These factors would make it difficult for characteristic species to colonize or
re-establish in the site once driven locally extinct. Nevertheless, evidence of ongoing
successional change on a relatively short time span suggests that disturbance
is necessary to maintain glade- or meadow-like conditions at this site. Presence of
at least five plant species listed as invasive in the state of Georgia is also cause for
concern about the native species present there. As such, to maintain the area in its
unique state, future management plans should include prescribed burning and/or
mowing, and possibly selective canopy tree removal to promote open understory
conditions. Questions regarding the nature, timing, season, and frequency of such
actions remain important issues to be addressed. As said at the outset, many superficially
similar limestone habitats are found throughout the region (including others
in Floyd County) and many harbor rare and endangered species. If disturbance is required
to maintain open canopy conditions of these sites, as seems to be the case for
Martha’s Meadow, protection of the sites alone may not be sufficient to ensure the
maintenance of characteristic flora.
This project was funded by a National Science Foundation, Research Experiences for
Undergraduates grant (DEB0354017) and the Berry College student work opportunity
program. The authors thank NatureServe, the National Park Service, and T. Govus for
unpublished data from the Chickamauga glades; and R. Carter for the Thunder Mountain
data sets; A. Huber, K. Currie, C. Worrell, and L. Rogers for helping with field logistics;
A. Good and T. Ware for helping identify plants; J. Barron for reviewing a manuscript
draft; and W. Yeomans and M. Huffman for assistance with management.
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334 Southeastern Naturalist Vol. 12, No. 2
Appendix 1. Plant species identified in Martha's Meadow from 2003 through 2008. In addition
to these species confirmed from flowering specimens, single unidentified species in
the following genera were also recorded: Amorpha, Aster, Carex, Carya, Celtis, Cirsium,
Clematis, Danthonia, Desmodium, Eupatorium, Galactia, Galium, Hypericum, Matelea,
Quercus, Rubus, Sanicula, Sisyrinchium, Smilax, Veronica, Viola, and Vitis.
Family code Scientific name, authority, and common name
Acanthaceae RUHU Ruellia humilis Nutt. (Fringeleaf Wild Petunia)
Aceraceae ACLE Acer leucoderme Small (Chalk Maple)
Agavaceae MAVI5 Manfreda virginica (L.) Salisb. ex Rose (False Aloe)
Anacardiaceae TORA2 Toxicodendron radicans (L.) Kuntze (Eastern Poison Ivy)
Apiaceae CHTA Chaerophyllum tainturieri Hook. (Hairyfruit Chervil)
Apiaceae DACA6 Daucus carota* L. (Queen Anne’s Lace)
Apiaceae SACA15 Sanicula canadensis L. (Canadian Blacksnakeroot)
Apiaceae THBA Thaspium barbinode (Michx.) Nutt.A [=chapmanii]
Apocynaceae APCA Apocynum cannabinum L. (Indianhemp)
Aquifoliaceae ILDE Ilex decidua Walt. (Possumhaw)
Aquifoliaceae ILOP Ilex opaca Ait. (American Holly)
Aristolochiaceae ARSE3 Aristolochia serpentaria L. (Virginia Snakeroot)
Asclepiadaceae ASHI Asclepias hirtella (Pennell) WoodsonA (Green Milkweed)
Asclepiadaceae ASTU Asclepias tuberosa L. (Butterfly Milkweed)
Asclepiadaceae ASVE Asclepias verticillata L. (Whorled Milkweed)
Asclepiadaceae ASVI Asclepias viridiflora Raf.A (Green Comet Milkweed)
Asclepiadaceae ASVI2 Asclepias viridis WaltA (Green Antelopehorn)
Asclepiadaceae MADE3 Matelea decipiens (Alexander) Woods. (Oldfield
Asclepiadaceae MAGO Matelea gonocarpos (Walt.) Shinners (Angularfruit
Asteraceae ACMI2 Achillea millefolium* L. (Common Yarrow)
Asteraceae AMAR2 Ambrosia artemisiifolia* L. (Annual Ragweed)
Asteraceae CIHO2 Cirsium horridulum Michx. (Yellow Thistle)
Asteraceae CIVU Cirsium vulgare* (Savi) Tenore (Bull Thistle)
Asteraceae ERPH Erigeron philadelphicus L. (Philadelphia Fleabane)
Asteraceae EUCA5 Eupatorium capillifolium (Lam.) Small (Dogfennel)
Asteraceae GAPU3 Gamochaeta purpurea (L.) Cabrera (Spoonleaf Purple
Asteraceae HEDI2 Helianthus divaricatus L. (Woodland Sunflower)
Asteraceae HYRA3 Hypochaeris radicata* L. (Hairy Cat’s Ear)
Asteraceae LEVU Leucanthemum vulgare* Lam. (Oxeye Daisy)
Asteraceae PAAN6 Packera anonyma (Wood) W.A. Weber & A. Löve
Asteraceae PRBA Prenanthes barbata (Torr. & Gray) MilsteadA (Barbed
Asteraceae RUFU2 Rudbeckia fulgida Ait. (Orange Coneflower)
Asteraceae RUHI2 Rudbeckia hirta L. (Blackeyed Susan)
Asteraceae SIAS2 Silphium asteriscus L. (Starry Rosinweed)
2013 M.L. Cipollini, M. Strahl, N. Soper Gorden, P. Tomlinson, and R. Ware 335
Family code Scientific name, authority, and common name
Asteraceae SMUV Smallanthus uvedalius (L.) Mackenzie ex Small (Hairy
Asteraceae SOAU2 Solidago auriculata Shuttlw. ex Blake (Eared Goldenrod)
Asteraceae SOOL Sonchus oleraceus* L. (Common Sowthistle)
Asteraceae SYPI2 Symphyotrichum pilosum (Willd.) Nesom (Hairy White
Asteraceae TAOF Taraxacum officinale* G.H. Weber ex Wiggers (Common
Asteraceae VEVI3 Verbesina virginica L. (White Crownbeard)
Asteraceae YOJA Youngia japonica* (L.) DC. (Oriental False Hawksbeard)
Betulaceae OSVI Ostrya virginiana (P. Mill.) K. Koch (Hophornbeam)
Bignoniaceae BICA Bignonia capreolata L. (Crossvine)
Bignoniaceae CARA2 Campsis radicans (L.) Seem. ex Bureau (Trumpet Creeper)
Boraginaceae LICA12 Lithospermum canescens (Michx.) Lehm. (Hoary Puccoon)
Boraginaceae MYMA Myosotis macrosperma Englm. (Largeseed Forget-Me-
Brassicaceae CAHI3 Cardamine hirsuta* L. (Hairy Bittercress)
Caprifoliaceae LOJA Lonicera japonica*† Thunb. (Japanese Honeysuckle)
Caprifoliaceae SYOR Symphoricarpos orbiculatus Moench. (Coralberry)
Caprifoliaceae VIRU Viburnum rufidulum Raf. (Rusty Blackhaw)
Caryophyllaceae STME2 Stellaria media* (L.) Vill. (Common Chickweed)
Celastraceae EUAM9 Euonymus americanus L. (Bursting-Heart)
Clusiaceae HYHY Hypericum hypericoides (L.) Crantz (St. Andrew’s Cross)
Clusiaceae HYMU Hypericum mutilum L. (Dwarf St. Johnswort)
Clusiaceae HYPU Hypericum punctatum Lam. (Spotted St. Johnswort)
Convolvulaceae CACAS3 Calystegia catesbeiana Pursh ssp. sericata (House)
BrummittB (Catesby’s False Bindweed)
Convolvulaceae IPPA Ipomoea pandurata (L.) G.F.W. Mey. (Man of the Earth)
Cornaceae COFL2 Cornus florida L. (Flowering Dogwood)
Cupressaceae JUVI Juniperus virginiana L. (Eastern Redcedar)
Cyperaceae CAAM8 Carex amphibola Steud. (Eastern Narrowleaf Sedge)
Cyperaceae CACH3 Carex cherokeensis Schwein. (Cherokee Sedge)
Cyperaceae CACO9 Carex complanata Torr. & Hook (Hirsute Sedge)
Cyperaceae CAGR8 Carex gracilescens Steud. (Slender Looseflower Sedge)
Cyperaceae CAME2 Carex meadii DeweyA (Mead’s Sedge)
Cyperaceae CAOX Carex oxylepis Torr. & Hook. (Sharpscale Sedge)
Cyperaceae SCOL2 Scleria oligantha Michx. (Littlehead Nutrush)
Ebenaceae DIVI5 Diospyros virginiana L. (Common Persimmon)
Elaeagnaceae ELPU2 Elaeagnus pungens* Thunb. (Thorny Olive)
Elaeagnaceae ELUM Elaeagnus umbellata*† Thunb. (Autumn Olive)
Euphorbiaceae EUCO10 Euphorbia corollata L. (Flowering Spurge)
Euphorbiaceae TRUR2 Tragia urticifolia Michx. (Nettleleaf Noseburn)
Fabaceae CEVI2 Centrosema virginianum (L.) Benth. (Spurred Butterfly
336 Southeastern Naturalist Vol. 12, No. 2
Family code Scientific name, authority, and common name
Fabaceae CECA4 Cercis canadensis L. (Eastern Redbud)
Fabaceae CHFA2 Chamaecrista fasciculata (Michx.) Greene (Partridge
Fabaceae CHNI2 Chamaecrista nictitans (L.) Moench (Sensitive Partridge
Fabaceae CLMA4 Clitoria mariana L. (Atlantic Pigeonwings)
Fabaceae DERO3 Desmodium rotundifolium DC. (Prostrate Ticktrefoil)
Fabaceae GLTR Gleditsia triacanthos L. (Honeylocust)
Fabaceae LECU Lespedeza cuneata*† (Dum.-Cours.) G. Don (Sericea
Fabaceae LEHI2 Lespedeza hirta (L.) Hornem.(Hairy Lespedeza)
Fabaceae LEPR Lespedeza procumbens Michx. (Trailing Lespedeza)
Fabaceae MELU Medicago lupulina* L. (Black Medick)
Fabaceae MIMI22 Mimosa microphylla Dry. (Littleleaf Sensitive-Briar)
Fabaceae RHTO3 Rhynchosia tomentosa (L.) Hook. & Arn. (Twining
Fabaceae SEMA11 Senna marilandica (L.) Link (Maryland Senna)
Fabaceae TRDU2 Trifolium dubium* Sibthorp (Suckling Clover)
Fabaceae TRRE3 Trifolium repens* L. (White Clover)
Fabaceae VICA2 Vicia caroliniana Walt. (Carolina Vetch)
Fagaceae QUMU Quercus muehlenbergii Engelm. (Chinkapin Oak)
Fagaceae QUNI Quercus nigra L. (Water Oak)
Fagaceae QUPA5 Quercus pagoda Raf. (Cherrybark Oak)
Fagaceae QUSH Quercus shumardii Buckl. (Shumard’s Oak)
Fagaceae QUST Quercus stellata Wangenh. (Post Oak)
Gentianaceae GEVI5 Gentiana villosa L. (Striped Gentian)
Gentianaceae SAAN Sabatia angularis (L.) Pursh (Rosepink)
Hamamelidaceae LIST2 Liquidambar styraciflua L. (Sweetgum)
Iridaceae SIAL3 Sisyrinchium albidum Raf. (White Blue-Eyed Grass)
Iridaceae SIAN3 Sisyrinchium angustifolium P. Mill. (Narrowleaf Blue-
Iridaceae SIAT Sisyrinchium atlanticum Bickn. (Eastern Blue-Eyed
Juglandaceae CAAL27 Carya alba Nutt. ex Ell. (Mockernut Hickory)
Juglandaceae CACA38 Carya carolinae-septentrionalis (Ashe) Engl. & Graebn.
(Southern Shagbark Hickory)
Juglandaceae CAOV2 Carya ovata (P. Mill.) K. Koch (Shagbark Hickory)
Lamiaceae MOFI Monarda fistulosa L. (Wild Bergamot)
Lamiaceae MODI4 Mosla dianthera* (Buch.-Ham. ex Roxb.) Maxim. (Miniature
Lamiaceae PRVU Prunella vulgaris L. (Common Selfheal)
Lamiaceae PYIN Pycnanthemum incanum (L.) Michx. (Hoary Mountainmint)
Lamiaceae SAAZ Salvia azurea Michx. ex Lam. (Azure Blue Sage)
Lamiaceae SALY2 Salvia lyrata L. (Lyreleaf Sage)
Lamiaceae SAUR Salvia urticifolia L. (Nettleleaf Sage)
2013 M.L. Cipollini, M. Strahl, N. Soper Gorden, P. Tomlinson, and R. Ware 337
Family code Scientific name, authority, and common name
Lamiaceae SCPAM Scutellaria parvula Michx. var. missouriensis (Torr.)
Goodman & C.A. LawsonA (Leonard’s Skullcap)
Liliaceae ALCA3 Allium canadense L. (Meadow Garlic)
Liliaceae NOBI2 Nothoscordum bivalve (L.) Britton (Crowpoison)
Liliaceae POBI2 Polygonatum biflorum (Walt.) Ell. (Smooth Solomon’s
Liliaceae SMBO2 Smilax bona-nox L. (Saw Greenbriar)
Liliaceae TRCU Trillium cuneatum Raf. (Little Sweet Betsy)
Malvaceae SIEL Sida elliottii Torr. & GrayA (Elliott’s Fanpetals)
Menispermaceae COCA Cocculus carolinus (L.) DC. (Carolina Coralbead)
Oleaceae FRAM2 Fraxinus americana L. (White Ash)
Oleaceae LISI Ligustrum sinense*† Lour. (Chinese Privet)
Orchidiaceae SPLAG Spiranthes lacera (Raf.) Raf. var. gracilis (Bigelow) Luer
(Northern Slender Lady’s Tresses)
Oxalidaceae OXST Oxalis stricta L. (Common Yellow Oxalis)
Oxalidaceae OXVI Oxalis violacea L. (Violet Woodsorrel)
Passifloraceae PALU2 Passiflora lutea L. (Yellow Passionflower)
Pinaceae PITA Pinus taeda L. (Loblolly Pine)
Plantaginaceae PLVI Plantago virginica L. (Virginia Plantain)
Poaceae ANGL2 Andropogon glomeratus (Walt.) B.S.P. (Bushy Bluestem)
Poaceae ANVI2 Andropogon virginicus L. (Broomsedge Bluestem)
Poaceae ANOD Anthoxanthum odoratum* L. (Sweet Vernalgrass)
Poaceae BRPU6 Bromus pubescens Muhl. ex. Willd. (Hairy Woodland
Poaceae BRRA2 Bromus racemosus* L. (Bald Brome)
Poaceae CHLA5 Chasmanthium latifolium (Michx.) Yates (Indian Woodoats)
Poaceae CHLA6 Chasmanthium laxum (L.) Yates (Slender Woodoats)
Poaceae CHSE2 Chasmanthium sessiliflorum (Poir.) Yates (Longleaf
Poaceae DAGL Dactylis glomerata* L. (Orchardgrass)
Poaceae DASP2 Danthonia spicata (L.) Beauv. ex Roemer & Schultes
Poaceae DIBO2 Dichanthelium boscii (Poir.) Gould & C.A. Clark (Bosc’s
Poaceae DICO2 Dichanthelium commutatum (J.A. Schultes) Gould (Variable
Poaceae DILA9 Dichanthelium laxiflorum (Lam.) Gould (Openflower Rosette
Poaceae ELHY Elymus hystrix L. (Eastern Bottlebrush Grass)
Poaceae ELVI3 Elymus virginicus L. (Virginia Wildrye)
Poaceae FESU3 Festuca subverticillata (Pers.) Alexeev (Nodding Fescue)
Poaceae MIVI Microstegium vimineum*† (Trin.) A. Camus (Nepalese
Poaceae PAAN Panicum anceps Michx. (Beaked Panicgrass)
Poaceae PAVI2 Panicum virgatum L. (Switchgrass)
338 Southeastern Naturalist Vol. 12, No. 2
Family code Scientific name, authority, and common name
Poaceae POAU Poa autumnalis Muhl. ex Ell. (Autumn Bluegrass)
Poaceae SAAL21 Saccharum alopecuroides (L.) Nutt. (Silver Plumegrass)
Poaceae SPOB Sphenopholis obtusata (Michx.) Scribn. (Prairie Wedgescale)
Poaceae TRFL2 Tridens flavus (L.) A.S. Hitchc. (Purpletop Tridens)
Portulacaceae CLVI3 Claytonia virginica L. (Virginia Springbeauty)
Pyrolaceae CHMA3 Chimaphila maculata (L.) Pursch. (Striped Prince’s Pine)
Ranunculaceae ANVI3 Anemone virginiana L. (Tall Thimbleweed)
Ranunculaceae DETR Delphinium tricorneA Michx. (Dwarf Larkspur)
Ranunculaceae RAAB Ranunculus abortivus L. (Littleleaf Buttercup)
Ranunculaceae RABU Ranunculus bulbosus* L. (St. Anthony’s Turnip)
Ranunculaceae RAHI Ranunculus hispidus Michx. (Bristly Buttercup)
Rhamnaceae BESC Berchemia scandens (Hill) K. Koch (Alabama Supplejack)
Rosaceae AGPU Agrimonia pubescens Wallr. (Soft Agrimony)
Rosaceae CRSP Crataegus spathulata Michx. (Littlehip Hawthorn)
Rosaceae DUIN Duchesnea indica* (Andr.) Focke (Indian Strawberry)
Rosaceae GIST5 Gillenia stipulata (Muhl. ex Willd.) Baill. (American Ipecac)
Rosaceae POCA17 Potentilla canadensis L. (Dwarf Cinquefoil)
Rosaceae POSI2 Potentilla simplex Michx. (Common Cinquefoil)
Rosaceae PRSE2 Prunus serotina Ehrh. (Black Cherry)
Rosaceae ROMU Rosa multiflora* Thunb. ex Murr. (Multiflora Rose)
Rubiaceae DIVI3 Diodia virginiana L. (Virginia Buttonweed)
Rubiaceae GAAP2 Galium aparine L. (Stickywilly)
Rubiaceae GACI2 Galium circaezans Michx. (Licorice Bedstraw)
Rubiaceae GAPI2 Galium pilosum Ait. (Hairy Bedstraw)
Rubiaceae SHAR2 Sherardia arvensis* L. (Blue Fieldmadder)
Scrophulariaceae MEAC Mecardonia acuminata (Walt.) Small (Axilflower)
Scrophulariaceae PELA8 Penstemon laevigatus Ait. (Eastern Smooth Beardtongue)
Solanaceae PHHE5 Physalis heterophylla Nees (Clammy Groundcherry)
Solanaceae PHVI5 Physalis virginiana P. Mill. (Virginia Groundcherry)
Solanaceae SOCA3 Solanum carolinense L. (Carolina Horsenettle)
Ulmaceae CELA Celtis laevigata Willd. (Sugarberry)
Valerianaceae VARA Valerianella radiata (L.) Dufr. (Beaked Cornsalad)
Verbenaceae VEBR2 Verbena brasiliensis* Vell. (Brazilian Vervain)
Violaceae VITR2 Viola triloba Schwein. (Three-Lobe Violet)
Vitaceae PAQU2 Parthenocissus quinquefolia (L.) Planch. (Virginia
Vitaceae VIRO3 Vitis rotundifolia Michx. (Muscadine)
ASpecies of special concern in GA associated with limestone substrates (GA DNR 2011).
BSpecies of special concern in GA not associated with limestone substrates (GA DNR
†Species listed as invasive by Georgia Exotic Pest Plant Council (2012).