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. Introduction 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 - mcipollini@berry.edu. 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. Methods 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. HThis study. IBaskin and Baskin 1996. JBartgis 1993. 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. Soil chemistry 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). Results 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 (Barbed Rattlesnakeroot) 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 (Hairyjoint Meadowparsnip) 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 value (IV). Season/species IV Spring 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 Summer 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 Fall 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 (Chalk Maple) 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 (Common Persimmon) 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 chemistry 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. Discussion 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. Parameter Value Ca 10,440 Mg 230 K 140 Mn 70 N-NO3 10 P 10 Zn 10 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. Conclusions 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 origin include: 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 cover, 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. Acknowledgments 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. 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USDA PLANTS 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] (Hairyjoint Meadowparsnip) 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 Milkvine) Asclepiadaceae MAGO Matelea gonocarpos (Walt.) Shinners (Angularfruit Milkvine) 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 Everlasting) 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 (Small’s Ragwort) Asteraceae PRBA Prenanthes barbata (Torr. & Gray) MilsteadA (Barbed Rattlesnakeroot) 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 USDA PLANTS Family code Scientific name, authority, and common name Asteraceae SMUV Smallanthus uvedalius (L.) Mackenzie ex Small (Hairy Leafcup) Asteraceae SOAU2 Solidago auriculata Shuttlw. ex Blake (Eared Goldenrod) Asteraceae SOOL Sonchus oleraceus* L. (Common Sowthistle) Asteraceae SYPI2 Symphyotrichum pilosum (Willd.) Nesom (Hairy White Oldfield Aster) Asteraceae TAOF Taraxacum officinale* G.H. Weber ex Wiggers (Common Dandelion) 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- Not) 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 Pea) 336 Southeastern Naturalist Vol. 12, No. 2 USDA PLANTS Family code Scientific name, authority, and common name Fabaceae CECA4 Cercis canadensis L. (Eastern Redbud) Fabaceae CHFA2 Chamaecrista fasciculata (Michx.) Greene (Partridge Pea) Fabaceae CHNI2 Chamaecrista nictitans (L.) Moench (Sensitive Partridge Pea) 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 Lespedeza) 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 Snoutbean) 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- Eyed Grass) Iridaceae SIAT Sisyrinchium atlanticum Bickn. (Eastern Blue-Eyed Grass) 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 Beefsteakplant) 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 USDA PLANTS 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 Seal) 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 Brome) 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 Woodoats) Poaceae DAGL Dactylis glomerata* L. (Orchardgrass) Poaceae DASP2 Danthonia spicata (L.) Beauv. ex Roemer & Schultes (Poverty Oatgrass) Poaceae DIBO2 Dichanthelium boscii (Poir.) Gould & C.A. Clark (Bosc’s Panicgrass) Poaceae DICO2 Dichanthelium commutatum (J.A. Schultes) Gould (Variable Panicgrass) Poaceae DILA9 Dichanthelium laxiflorum (Lam.) Gould (Openflower Rosette Grass) 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 Browntop) Poaceae PAAN Panicum anceps Michx. (Beaked Panicgrass) Poaceae PAVI2 Panicum virgatum L. (Switchgrass) 338 Southeastern Naturalist Vol. 12, No. 2 USDA PLANTS 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 Creeper) 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 2011). *Exotic species. †Species listed as invasive by Georgia Exotic Pest Plant Council (2012).