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Herbaceous Plants and Grasses in a Mountain Longleaf Pine Forest Undergoing Restoration: A Survey and Comparative Study
Martin L. Cipollini, Joshua Culberson, Cade Strippelhoff, Thomas Baldvins, and Kalia Miller

Southeastern Naturalist, Volume 11, Issue 4 (2012): 637–668

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2012 SOUTHEASTERN NATURALIST 11(4):637–668 Herbaceous Plants and Grasses in a Mountain Longleaf Pine Forest Undergoing Restoration: A Survey and Comparative Study Martin L. Cipollini1,*, Joshua Culberson2, Cade Strippelhoff 3, Thomas Baldvins4, and Kalia Miller1 Abstract - The Berry College Longleaf Pine Management Area consists of old-growth fire-suppressed mountain Pinus palustris (Longleaf Pine) stands embedded within an encroaching matrix of mixed pines and hardwoods. Since 2001, portions of this area have been subjected to restoration efforts involving logging followed by burning, foliar herbicide application, and planting, as well as burning and hardwood control using herbicides in unlogged old-growth stands. To document the herbaceous plants and grasses of this site and to begin to address questions concerning the short-term impacts of management practices on these species, flowering specimens were systematically collected in managed and unmanaged stands in 2008 and 2009. We recorded 201 species in 35 families, including 70 species of Asteraceae, 35 species of Poaceae, 17 species of Fabaceae, and 10 grass-like species other than Poaceae (Cyperaceae, Iridaceae, and Juncaceae). Native herbaceous plants most commonly found included: Houstonia caerulea, Hypoxis hirsuta, Solidago odora, Oxalis stricta, Coreopsis major, Hypericum hypericoides, Lespedeza procumbens, Hieracium venosum, and Packera paupercula. While only 14 species were found in unmanaged old growth, 127 were found in managed old growth, and 167 in logged areas. Fire suppressed old-growth mountain Longleaf Pine forests are generally virtually devoid of understory plant diversity; these results suggest that reduction in canopy density and leaf litter can substantially recover herbaceous and grass species diversity. However, the extent to which understory diversity can be fully recovered in any specific site remains in question, particularly if local propagule sources have vanished during the period of fire suppression. A comparison with historical species lists at our site, and with other mountain Longleaf Pine forests in various stages of fire maintenance, is presented to help define characteristic understory species for mountain Longleaf Pine forests. Introduction In 2001, Berry College established a project to begin restoring its relict mountain Pinus palustris Mill. (Longleaf Pine) forests on Lavender Mountain, Floyd County, GA. The mountain Longleaf Pine forests of the Berry College campus represent an ecologically significant landscape type about which there is a paucity of knowledge (Stowe et al. 2002). Most information on Longleaf Pine forests comes from sandhills or coastal plain landscape types, i.e., wiregrass country. Eleusine indica (L.) Gaertn. (Wiregrass), Gopherus polyphemus Daudin, 1Department of Biology, Berry College, Mount Berry, GA 30149. 2575 Oglethorpe Mountain Road, Jasper, GA 30143. 3Environmental Safety and Occupational Health, Georgia Tech Research Institute, Atlanta, GA 30332. 4Joseph W. Jones Ecological Research Center at Ichauway, Newton, GA 39870. *Corresponding author - mcipollini@berry.edu. 638 Southeastern Naturalist Vol. 11, No. 4 (Gopher Tortoise), and many of the scrub oaks considered characteristic of the better-known Longleaf Pine ecosystems are far outside the range of Berry College’s stands. The college campus has a number of very old trees, some in excess of 250 years old. There are very few tracts of old Longleaf Pine left in the south, and almost none left within “mountain” or “montane” Longleaf Pine areas of northwestern Georgia and northeastern Alabama (Varner and Kush 2004, Varner et al. 2000). As with most remaining old-growth Longleaf Pine forests, stands at Berry had been fire-suppressed for a long time—perhaps as long as 60 or 70 years (M.L. Cipollini, pers. observ.). Prior to recent restoration efforts, native herbaceous plant and grass cover had been virtually completely lost, other than perhaps patches of plants in scattered refugia or as dormant seeds in the soil. Nevertheless, much of the steep hillsides of Lavender Mountain had never been plowed, which increases the likelihood that understory species characteristic of fire-maintained mountain Longleaf Pine forests might be recovered via management practices. Andrews (1917) and Jones (1940) both conducted surveys of the plant communities on the Berry Campus. These historical surveys as well as surveys conducted recently at fire-managed and fire-suppressed mountain Longleaf Pine sites in Georgia and Alabama (Carter and Londo 2006, Varner et al. 2000, Womack and Carter 2011) can serve as sources for determining potential or “target” plant communities of restored mountain Longleaf Pine forests. Mountain Longleaf Pine forests occupied a relatively small part of the historic range of Longleaf Pine, and are confined to middle to high elevations of northwest Georgia and northeast Alabama (Varner 1999, Varner et al. 2003; see also Harper 1905). Due to their occurrence within relatively developed, agricultural, and fire-suppressed landscapes (in comparison with the more well-studied Longleaf Pine forests of the southeast coastal plain), mountain Longleaf Pine habitat loss may be even more dramatic than that seen in coastal systems (Brockway et al. 2005, Varner 1999). Means (1996) reported that less than 4000 ha of mountain Longleaf Pine forest remained (0.0001% of the range) as of 1995. The loss of these forests has prompted many landowners and organizations to begin to establish Longleaf Pine restoration projects. Fire is particularly important to this ecosystem partly because Longleaf Pines are relatively poor dispersers, and seed germination and early growth is enhanced by bare soil conditions and high light intensity (Landers et al. 1995, Platt et al. 1988). Along with Longleaf Pine, many plant species in these habitats are expected to display firedependent life histories and to require open conditions that minimize competition with non-fire-tolerant species, facilitate seed germination, and maintain required high light and low litter levels (Peet and Allard 1993). By restoring these forests using mechanisms that open the canopy, remove encroaching hardwoods, and lower litter levels, land managers have the potential to restore the plant community to one similar in makeup to its pre-fire-suppression condition. Management typically consists of strategies such as prescribed burning, herbicide injection of competing trees, and direct removal of timber, which may or may not be accompanied by planting. Generally, these have proven effective in re-establishing 2012 M.L. Cipollini, J. Culberson, C. Strippelhoff , T. Baldvins, and K. Miller 639 Longleaf Pine as the dominant tree (e.g., Kush et al. 2004). The effects of Longleaf Pine restoration on other aspects of the biological community are not well studied in mountain Longleaf Pine forests. Key questions of interest to this study concern the effect management practices have on the understory plant community and what is the target understory plant community for mountain Longleaf Pine restoration. One of the goals of the Berry College Longleaf Pine Project (Cipollini 2005) is the restoration of the historical understory plant community to Lavender Mountain. The main purpose of this project was to initiate a baseline study of the understory plant community in areas undergoing restoration, with reference to two unmanaged (fire-suppressed) old-growth areas (UM sites). This study represents the first major initiative to document herbaceous plants and grasses in this community and may assist others in developing lists of species expected to be found in similar habitats undergoing restoration. This study has implications for mountain Longleaf Pine management projects, especially those undertaken in areas where the understory plant community has been more-or-less eliminated (i.e., where local refugia and/or seed banks do not exist). By comparing our findings with historical records and with data derived from other sites designated as “mountain Longleaf Pine” forest, we hope to generate a draft list of species expected to be found in restored areas. Field Site Description Study stands were in the Berry College Longleaf Pine Management Area (BCLPMA; 34.34481°N, -85.20717°W) on Lavender Mountain, Floyd County, GA, an area of ≈140 ha containing ≈120 ha of relict Longleaf Pine stands (Cipollini 2005). The BCLPMA is centrally located within a Georgia Department of Natural Resources Wildlife Management area on the wooded ≈10,500-ha Berry College campus. This large campus is adjacent to other large protected forest areas (Rocky Mountain Recreational Area and Chattahootchee National Forest, both to the north of campus). Old-growth Longleaf Pines in the BCLPMA range from about 75–250 years old, and fires have been suppressed on Lavender Mountain since the college was founded in 1902. Prior to restoration efforts, relict Longleaf Pine stands had sustained hardwood encroachment, and excessive litter and woody fuel build-up (Cipollini et al. 2005, Currie et al. 2006, Huber et al. 2006). Pinus echinata Mill. (Shortleaf Pine) share dominance with Longleaf Pines in areas of greatest Longleaf Pine density, although Pinus taeda L. (Loblolly Pine), Pinus virginiana Mill. (Virginia Pine), and most of the hardwoods had apparently invaded during the period of fire suppression. Starting in 2001, a series of management activities were initiated in this area as part of a long-term project to restore the Longleaf Pine ecosystem (see Table 1 for management details). For this study, a stand was defined as an area with a uniform management history since 2001 centered within former or existing areas of high Longleaf Pine 640 Southeastern Naturalist Vol. 11, No. 4 density. This study encompassed 16 stands, ranging in intensity of management. Eight stands (2.1–4.3 ha each; 26.7 ha total) had been heavily managed (HM), having been logged to mitigate infestations of mature Loblolly Pines and Shortleaf Pines with Dendroctonus frontalis Zimmerman (Southern Pine Beetle). Longleaf Pines and most hardwoods were also removed at the time of logging. Following logging, HM stands were treated with foliar and injection applications of herbicides targeted to residual hardwoods, Rubus spp., and invasive plants, were planted with Longleaf Pine seedlings, and were burned several times. Six stands (2.1–8.3 ha each; 28.6 ha total) were mature, formerly fire-suppressed Longleaf Pine stands that were similar to HM stands prior to management, containing a mixture of mature pines and encroaching hardwoods. These stands were intermediately managed (IM), having had most hardwoods injected with herbicides and having been burned several times. Two stands (5.7 ha each; 11.4 ha total), similar in condition to managed stands prior to management, were completely unmanaged (UM). All stands were embedded within a matrix of infrequently burned mature mixed pine and hardwood forest, so HM and IM stands constituted “islands” within a matrix whose forest composition and structure was similar to UM stands. An attempt was made to select stands for study so that area, slope position, aspect, and other characteristics of the stands did not confound differences in management history. Stands were between 200 m and 400 m in elevation on south- and southwest-facing slopes of 1–45% grade. The rocky, acidic, well-drained soils are dominated by fine sandy loams, stony fine sandy Table 1. Summary of major recent management activities in the areas surveyed in this study, organized by general management practice. Management Site name Management practicesA began Area (ha) HM sites: Logged pine areas, with foliar and injection herbiciding of hardwoods, burning and planting of Longleaf Pine seedlings since 2001. Clear Cut CC (1), HS (4), FH (2), B (2), P (2) 2003 3.19 SAVE Middle SC (1), HS (4), B (1), P (2) 2001 5.00 SAVE West SC (1), HS (3), P (3) 2001 2.11 Seed Tree Cut CC (1), HS (2), FH (1), B (2), P (3) 2005 4.26 Selective Cut 1 and 2 East CC (1), HS (4), FH (2), B (4), P (3) 2003 6.07 Selective Cut 1 and 2 West SC (1), HS (4), FH (2), B (3), P (2) 2003 6.07 IM sites: Mature Longleaf Pine areas, with injection herbiciding of hardwoods and one or more prescribed burns since 2001 Stands A and B HS (2), B (3) 2001 7.09 Stands C, D, and E HS (2), B (2) 2003 15.47 SAVE East HS (2), B (1) 2005 3.89 UM sites: Fire-suppressed mature Longleaf Pine areas Stands F and H No active management N/A 11.44 ACC = clear cut logging, with goal of removing most canopy trees ≥ 30 cm in diameter; SC = selective cut logging, with goal of removing all pines and most hardwoods <30 cm in diameter; B = nongrowing season prescribed burn; HS = wide-spaced injection of ArsenalAC with goal of reducing 50% of hardwoods <30 cm in diameter with each application; FH = targeted foliar application of Garlon 4 or Garlon 3A herbicides to residual hardwoods, Rubus spp., and invasive species. 2012 M.L. Cipollini, J. Culberson, C. Strippelhoff , T. Baldvins, and K. Miller 641 loams, and gravelly silt-loams (Soil Survey Staff, USDA NRCS 2011). The two managed classes (HM and IM) were represented by similar total areas, whereas the unmanaged class (UM) was represented by an area of about half that size. We chose to survey a smaller area of unmanaged stands because previous informal surveys as well as formal studies of herbaceous plant and grass cover (e.g., Currie et al. 2006) had revealed extremely low abundance and diversity of herbs and grasses in stands similar to the 2 we surveyed. Unmanaged old-growth Longleaf Pine stands within the BCLPMA and on other areas on Lavender Mountain are uniformly devoid of understory herbs and grasses, being heavily suppressed by deep shade and decades of litter accumulation. Understory plants there are generally woody shrubs (e.g., Vaccinium spp.) and vines (e.g., Smilax spp.). Methods Quantitative vegetation characteristics In spring and summer 2009, 45 points were randomly located in each of 12 of the 16 stands (5 HM, 5 IM, and the 2 UM stands). From each point, the number of pines, hardwoods, and dead trees >5 m in height were counted using the Bitterlich (1948) variable plot method, using a 1-m-long angle gauge with a 1.4-cm-wide sight. To estimate basal area (m²/ha) of trees in the area surrounding each point, counts of trees appearing wider than the sight were divided by 2 (Barbour et al. 1999). Within a 1-m2 quadrat established at each point, the percent cover of each of the following variables was visually estimated: woody plants 1–5 m tall, woody plants <1 m tall, grasses, herbaceous plants, and litter. Litter depth was taken at the center point, and litter volume (L/m2) was calculated as a product of cover and depth. Vegetation data were analyzed using multiple analysis of variance (MANOVA; IBM®, SPSS Statistics 19, Chicago, IL) to determine differences among management classes. Plant specimen collection and identification Each of the 16 stands was visited by 4 surveyors for one 3-hr period (total = 12 person hours) each month from June–October 2008, and again in April and May of 2009. During each survey, the researchers systematically walked through the entire stand searching for plants in flower. When a specimen was encountered, the latitude and longitude coordinates were recorded and the specimen was tentatively identified at least to genus in the field using field guides such as Newcomb (1977). Voucher specimens were taken for each plant thus identified, duplicate specimens were combined, and the best specimen of each species collected was mounted for verification of identification and added to Berry College’s herbarium. Using these specimens, tentative identifications were cross-referenced with Radford et al. (1968), various other guides (e.g., Dean et al. 1973, Duncan and Foote 1975, Horn et al. 2005, Knobel 1980), and a species checklist for the flora of Floyd County, GA (R. Ware and T. Ware, Georgia Botanical Society, Rome, GA, unpubl. data; hereafter “Wares’ Flora 642 Southeastern Naturalist Vol. 11, No. 4 of Floyd County”). The PLANTS on-line database (USDA, NRCS 2011) was utilized to further cross-check each specimen and to assign accepted species names and plant codes. Species were categorized according to sun- vs. shadetolerance, life history (perennial, biennial, annual), soil-moisture preferences, preference for disturbed sites (i.e., weedy vs. non-weedy species), and native status using the reference materials listed above, as well as Weakley (2011) and University of Texas (2011). We first used these data to describe species richness and species characteristics in areas of the BCLPMA with contrasting recent management histories (Table 1). To provide a rough estimate of relative abundance of each species in each management class (HM, IM, and UM), the total number of records of each species found in each management class was tabulated. This approach permitted a comprehensive species list to be developed for the three main management classes, and provided a first approximation of the relative importance of different species. Comparison with historical data and with other sites Our comprehensive species list was compared with lists of plants considered diagnostic of upland Longleaf Pine habitats on Lavender Mountain when it was still under fire maintenance (Andrews 1917) and for upland habitats soon after fire-suppression efforts began (Jones 1940). Our list was also compared with lists of diagnostic species for recently fire-maintained mountain Longleaf Pine areas of Fort McClellan, Calhoun County, AL (Maceina et al. 2000, Varner et al. 2000) and for fire-suppressed mountain Longleaf Pine areas of Thunder Scout Reservation, Upson County, GA (Carter and Londo 2006) and Horseblock Mountain, Talledega National Forest, Cleburne County, AL (Womack and Carter 2011). Species were grouped according to their presence and absence in each of these data sets. This comparison of species lists served to help identify species that are likely to be encountered within Longleaf Pine stands in the BCLPMA or in mountain Longleaf Pine habitats in general, but may have not been recorded in our current study. Results Quantitative vegetative characteristics Multiple analysis of variance revealed significant differences among the management classes for all vegetative variables except woody plants 0–5 m tall (Table 2). In general, there was greater grass and herbaceous plant cover and lower litter volume and cover of living trees >5 m tall as management intensity increased. In comparison with IM and UM stands, HM stands contained about half the basal area of hardwoods, one fourth the basal area of pines, 4 to 15 times greater grass cover, 2 to 3 times the herbaceous plant cover, and 50% to 30% lower litter volume, respectively. IM stands had greater basal area of dead trees >5 m in comparison with both HM and UM stands (about 300% and 350% greater, respectively). 2012 M.L. Cipollini, J. Culberson, C. Strippelhoff , T. Baldvins, and K. Miller 643 Herbacous plants and grasses of the BCLPMA Altogether, 607 specimens comprising 201 species were collected among the 16 stands. Of the species identified (Table 3, Appendix I), ≈35% were Asteraceae (70 species), followed by Poaceae with ≈17% (35 species) and Fabaceae with ≈8% (17 species). The 35 Poaceae were dominated by Dichanthelium spp. and Panicum spp. grasses, in particular D. boscii (Poir.) Gould & C.A. Clark (Bosc’s Panic Grass). In fact, the Dichanthelium genus was the most species rich overall (Table 3). Part of our intent was to compare managed and unmanaged areas in species diversity and plant characteristics. Although not based upon equal sample areas (area surveyed in UM stands was approximately one half of the areas surveyed in HM and IM stands), differences among the 3 management classes in plant species diversity were disproportionalty pronounced, with 165, 127, and 14 species recorded in HM, IM, and UM stands, respectively. Examining the most commonly Table 3. Summary of the 35 plant families encountered in this study, including the total number of species found within each family and the proportion of the total species represented by that family. Family Species Proportion Asteraceae 70 34.8% Poaceae 35 17.4% Fabaceae 17 8.5% Lamiaceae 9 4.5% Rubiacea, Oxalidaceae, Polygalaceae 6 each 3.0% each Campanulaceae, Cyperaceae 5 each 2.5% each Clusiaceae 4 2.0% Euphorbiaceae, Iridaceae, Scrophulariaceae, Verbenaceae 3 each 1.5% each Ericaceae, Juncaceae, Melasomataceae, Rosaceae, Violaceae 2 each 1.0% each Apiaceae, Apocynaceae, Brassicaceae, Caryophyllaceae, 1 each 0.5% each Chenopodiaceae, Commelinaceae, Geraniaceae, Liliaceae, Linaceae, Passifloraceae, Phytolaccaceae, Plantaginaceae, Polemoniaceae, Portulacaceae, Ranunculaceae, Solanaceae Table 2. Means and standard deviations for vegetative characteristics of the study areas from which herbaceous plants were collected in this study. Identical letters denote means that did not differ significantly among management classes based upon one-way analyses of variance following significant overall MANOVA results. Management class (n) Vegetative characteristic HM (5) IM (5) UM (2) Hardwood >5 m tall (m2/ha) 2.4 ± 1.8 a 4.4 ± 1.3 b 4.8 ± 1.4 c Pine >5 m tall (m2/ha) 1.5 ± 1.1 a 6.8 ± 1.5 b 5.2 ± 0.0 c Dead tree >5 m tall (m2/ha) 3.1 ± 2.2 a 5.9 ± 1.4 b 4.5 ± 1.0 a Woody plants >1–5 m (% cover) 2.4 ± 3.1 a 1.9 ± 1.5 a 2.8 ± 1.0 a Woody plants 0–1 m (% cover) 5.3 ± 4.5 a 5.4 ± 4.0 a 2.0 ± 2.1 a Grasses (% cover) 30.8 ± 18.1 a 7.2 ± 4.8 b 2.4 ± 2.9 c Herbaceous plants (% cover) 11.6 ± 12.4 a 8.7 ± 4.5 b 4.5 ± 2.2 c Litter (L/m2) 11.9 ± 9.1 a 20.6 ± 4.8 b 32.0 ± 16.2 c 644 Southeastern Naturalist Vol. 11, No. 4 encountered species in each stand type (“top 20”; Table 4) reveals that HM stands held the highest ratio of sun-loving to shade-tolerant plants, a high proportion of perennials, and the highest ratio of dry- to moist-soil-adapted plants. Six legume species were among the most common species in these sites. These stands also showed a high proportion of plants responding to disturbance and 2 alien annuals associated with dry soils. IM stands had a higher proportion of sun-loving plants than did UM stands, as well as a greater proportion of perennials to annuals. Species dominating these areas fell between HM and UM stands in soil-moisture preferences, but many of the perennials were Asteraceae with preferences for dry soil conditions and disturbed sites. The few species found in UM areas had the lowest ratio of sun-loving to shade-tolerant plants, 3 annuals associated with moist soils, the lowest ratio dry- to moist-soil-adapted plants, and a relatively low proportion of plants associated with disturbance. Dichanthelium spp. and Panicum spp. grasses with at least partial preferences for moist, shaded sites dominated this group of species. Comparisons with historical records and other mountain Longleaf Pine sites In comparing our species list with those of previous studies on Lavender Mountain as well as with other sites characterized as mountain Longleaf Pine (Appendix II), we found the following patterns: Species Group 1. There were 74 species found in our site, in other mountain Longleaf Pine sites, or in our site prior to the 1940s. Most of these are sun-loving perennials and annuals with preference for dry soils. Some are shade or partshade tolerant, a few are pioneer and alien species, and nearly all are in Wares’ Flora of Floyd County. Species Group 2. Eighty-three species were recorded in mountain Longleaf Pine sites other than ours and in Wares’ Flora of Floyd County. This group includes a mixture of shade-tolerant native perennials, sun-loving species, weeds, and 5 invasive aliens. These species are a mix of those that generally respond well to open conditions and disturbance, as well as species associated more with undisturbed (fire-suppressed) forests. Some are known to be present on Lavender Mountain from undocumented observations, and some were recorded in our site in the past, but were not collected and recorded during the formal 2008–2009 census (e.g., some Asclepias spp., Galium spp., Oenothera spp., Linum spp., etc.). Species Group 3. Ninety-seven species were found in our site and in Wares’ Flora of Floyd County, but not at our site in the past and not at any other mountain Longleaf Pine site examined. This group is dominated by widespread, sun-loving, and weedy species, including 12 invasive species. This group does, however, include many grass and grass-like species not necessarily associated with disturbed sites. Species Group 4. A group of 28 species was recorded in our site, but not at other mountain Longleaf Pine sites or in Wares’ Flora of Floyd County. Of these species, many are reported as having either US coastal plain beach or sand 2012 M.L. Cipollini, J. Culberson, C. Strippelhoff , T. Baldvins, and K. Miller 645 Table 4. Most frequent species in each of the 3 major management classes, based upon the total number of recorded sightings in each management class. Listed are the top 20 including species tied in frequency (which makes for more than 20 for the IM site and combined lists), sorted in descending rank. Detailed information on these species can be found in Appendixces I and II. HM Stands (top “20”) IM Stands (top “20”) UM Stands (all 14 species) All stands combined (top “20”) Oxalis stricta Houstonia caerulea Hypoxis hirsuta Houstonia caerulea Houstonia caerulea Solidago odora Dichanthelium laxiflorum Hypoxis hirsuta Chaenorhinum minus Hieracium venosum Tradescantia virginica Solidago odora Dichanthelium boscii Hypoxis hirsuta Coreopsis major Oxalis stricta Packera paupercula Oxalis violacea Dichanthelium boscii Chaenorhinum minus Lespedeza procumbens Coreopsis major Dichanthelium commutatum Coreopsis major Hypericum hypericoides Eupatorium rotundifolium Panicum philadelphicum Hypericum hypericoides Piptochaetium avenaceum Hypericum hypericoides Hieracium piloselloides Lespedeza procumbens Pycnanthemum muticum Pipochaetium avenaceum Hieracium venosum Pipochaetium avenaceum Coreopsis major Clitoria mariana Houstonia caerulea Hieracium venosum Eupatorium hyssopifolium Hieracium piloselloides Oxalis dillenii Packera paupercula Lespedeza virginica Lespedeza procumbens Panicum dichotomiflorum Dichanthelium boscii Mimosa strigillosa Sericocarpus linifolius Polygala verticillata Clitoria mariana Strophostyles umbellata Solanum carolinense Solidago odora Hieracium piloselloides Chrysopsis mariana Symphyotrichum dumosum Oxalis violacea Clitoria mariana Symphyotrichum racemosum Pycnanthemum muticum Hypoxis hirsuta Chaenorhinum minus Solanum carolinense Krigia virginica Erechtites hieracifolia Erigeron strigosus Lespedeza frutescens Erigeron strigosus Eupatorium hyssopifolium Silene virginica Hypochaeris radicata Eupatorium rotundifolium Oxalis stricta Hypochaeris radicata Pityopsis graminifolia Lespedeza virginica Sorghastrum nutans Mimosa strigillosa Pityopsis graminifolia Symphyotrichum dumosum 646 Southeastern Naturalist Vol. 11, No. 4 dune affiliations, or northeastern, midwestern, and northwestern US prairie or mountain affiliations. Seven are listed as introduced species. This group contains mostly sun-loving perennials and a few annuals, with a tendency for moist soil preferences. Species Group 5. Eight species were listed in other mountain Longleaf Pine sites or in historical records for Lavender Mountain, but were not found in our site during this study, and were not listed in Wares’ Flora of Floyd County. While some of these species have been recorded in nearby counties and in northern Georgia in general, 2 of them are far out of range and likely are incorrect (Stylosanthes calcicola Small [Everglade Key Pencilflower] and Tephrosia mohrii (Rydb.) Godfrey [Pineland Hoarypea]). The other 6 species represent a small group of sun-loving (mostly) perennials associated with variable site preferences. Discussion This study represents to our knowledge the first study of understory plant diversity in mountain Longleaf Pine forests in northwestern Georgia (other studies having been conducted in middle Georgia or northeastern Alabama). As expected based upon differences in management history, differences existed among the management classes for all quantitative vegetative variables except woody plants 0–5 m tall (Table 2). In general, there was greater grass and herbaceous plant cover, and lower litter volume and cover of living trees >5 m tall as management intensity increased. The only exception to this pattern was for dead trees, which were retained at higher levels in IM stands due to herbicide-injection practices that left standing dead trees. Although not represented by equal sample areas, managed stands had about 10 times greater species diversity than unmanaged stands, which corresponds with informal surveys in many fire-suppressed old-growth mountain Longleaf Pine stands whose understories are virtually devoid of herbaceous plant and grass cover. Local examples of such depauperate understories can be seen in other areas on Lavender Mountain southwest of the Berry College campus and on Horseleg Mountain (Marshall Forest TNC Preserve, Rome, Floyd County, GA); indeed, most extant old-growth mountain Longleaf Pine stands are in this condition (Varner and Kush 2004). The small group of species found in unmanaged stands might be considered to be the group most tolerant of long-term fire suppression in this system; as expected, many of these plants have preferences for shady or moist conditions. Species in managed stands of the BCLPMA were dominated by perennials with preferences for dry and sunny conditions, and disturbed acidic soils, and included few alien species. None of the alien species are considered particularly invasive, although woody or semi-woody species such as Pueraria lobata (Willd.) Ohwi (Kudzu), Ligustrum sinense Lour. (Chinese Privet), Lonicera japonica Thunb. (Japanese Honeysuckle), and Paulownia tomentosa (Thunb.) Steud. (Royal Pawlonia) are invasive to some degree in managed sites, and have been managed in recent years via targeted herbicide applications. 2012 M.L. Cipollini, J. Culberson, C. Strippelhoff , T. Baldvins, and K. Miller 647 In our comparative study of mountain Longleaf Pine sites, the following conclusions can be made about each species group: Species Group 1. Most of these species are sun-loving perennials adapted to dry sites, but the presence of some shade-tolerant, moisture-loving species in this group may be indicative of the general state of fire suppression currently seen in such systems. Of the 16 stands we surveyed in the BCLPMA, 14 were in some state of management that had substantially opened up the canopy, controlled competing hardwoods and shrubs, and substantially decreased litter layers. Considering this, species group 1 may be most representative of species that might be expected for mountain Longleaf Pine sites in general, at least when they are in some early stage of restoration. Species Group 2. This group includes a mix of shade-tolerant and sun-loving natives, and some weedy non-natives. Excluding the non-natives, the characteristics of this group also suggest inclusion in the baseline list of species considered to be representative of mountain Longleaf Pine stands undergoing restoration. Many of these may have simply gone unrecorded in our 2008–2009 study—either being rare or non-flowering during the time of the study . Species Group 3. About half of the Asteraceae and Poaceae species we recorded are included in this group, as were nearly half of the non-native species. Because this group is dominated by widespread, sun-loving, weedy species, we expect that many members of this group represent transient species likely to have responded to recent management-induced disturbances. We further suspect that some of these species will be unlikely to be maintained in great abundances as succession ensues and regular low-intensity fires become a main controlling factor (prescribed burns at 3- to 5-year intervals are the only planned activities for managed sites). On the other hand, many of the native grass and grass-like species in this group might be fire tolerant or fire regenerating, and might indeed be representative of the target flora. Grass-like species are commonly overlooked or lumped into general groups (e.g., “Andropogon spp.”, “Panicum spp.”) in short-term surveys, so we suspect that these may have been missed in some of the studies we used for our comparisons. Species Group 4. Nearly half of these species were listed as out of range (either approaching Floyd County, GA from the north or northwest, or from the southeastern coastal plain). Many of the species in this group are associated with sunny, moist conditions. It is possible that this group represents a subset of species at the limits of their distributions (or expanding their distribution), with overlap in the sunny, wet, nutrient-poor soils found in some areas within our site. Several of these, including the introduced Chaenorhinum minus (L.) Lange (Dwarf Snapdragon), were among the most commonly encountered species in managed sites, and reflect conditions found in some of the lower-lying areas within the BCLPMA. Species Group 5. While some of these species have been reported in nearby counties and in northern Georgia in general, the 2 species far out of range are unlikely to be found in mountain Longleaf Pine habitats. The other 6 species 648 Southeastern Naturalist Vol. 11, No. 4 represent a small group of sun-loving (mostly) perennials with variable site preferences. As such, this latter group is unlikely representative of mountain Longleaf Pine sites specifically, but might occasionally be found in such sites. Overall, the BCLPMA species list was dominated, as expected, by Asteraceae, Poaceae, and Fabaceae. With 201 confirmed species, it is clear that the understories of these sites are not nearly as species rich as might be found in a typical coastal plain Longleaf Pine forest (cf. Peet and Allard 1993); nevertheless, this number is nearly twice the number of species found in the few comprehensive studies available for comparison. Maceina et al. (2000) and Varner et al. (2000, 2003), for example, recorded only ≈100 herbaceaous plant and grass species in mountain Longleaf Pine habitats maintained by frequent fires. This is about half the number of species we recorded in our managed areas. One reason for this difference is that more extensive area was surveyed for a longer period of time in our study. Our survey could represent a more exhaustive look at grass- and grass-like species, in comparison with similar studies done in mountain Longleaf Pine habitats, which might explain why we recorded more of these types. Many of these species, particularly perennial warm-season bunch grasses, may be fire tolerant and expected of fire-maintained mountain Longleaf Pine forests. On the other hand, many of the plants (particularly those in species group 3, which is dominated by weedy plants associated with soil disturbances) may drop out with continued fire management (herbiciding and man-made physical soil disturbances having ended). Removing subtantial hardwood component either through logging, fire, or herbiciding leads to substantial canopy openness and (in the case of logging operations) soil disturbances. We expect canopy openness to be reduced somewhat as Longleaf Pines grow and add canopy, and substantial soil disturbances to be minimal in the future. Ultimately, we expect HM stands to end up with approximately the same canopy structure as IM stands. This could lead to loss of some species dependent upon disturbances other than periodic fires, or on full sun exposure. If this happens, the species number in a dynamically stable fire-maintained condition could then approximate that found by other researchers. That said, over 200 species, including woody species, were recently recorded in the understory of a frequently burned mountain Longleaf Pine forest on the Talladega National Forest, AL (A. Dotson and R. Carter, Jacksonville State University, Jacksonville, AL, unpubl. data). Our combined species list (Appendix II) may be used by land managers and others interested in understanding, managing, or restoring understory plant communities in mountain Longleaf Pine forests. Species not found, but expected to be found, within a similar system undergoing restoration might be targeted for intensive restoration projects such as direct seeding or other planting efforts. Projects of this sort are increasing in frequency within the historic mountain Longleaf Pine range. There is no doubt that restoration efforts involving canopy reduction, targeted herbicide use, and prescribed fire can bring about a dramatic return of many of the species expected to be present in formerly fire-maintained systems. The 2012 M.L. Cipollini, J. Culberson, C. Strippelhoff , T. Baldvins, and K. Miller 649 recovery of an intact understory plant community is, however, dependent upon a local seed source or soil seed bank. It is fortunate in this respect that most of our study areas in the BCLPMA had been unplowed and had not been subjected to intensive logging during their long period of fire suppression. Dormant seeds in the soil or local refugia (e.g., road banks, steep disturbed hillsides, occasional small burn areas, nearby commercial logging areas) have likely been the sources of the dramatic understory response to management within the BCLPMA. The BCLPMA site may further serve as a source for the introduction of species to other restoration sites. Controlling invasive species, recovering “lost” species, and maintaining a prescribed fire regime will likely be necessary to maintain this level of plant diversity. Effects on the biota beyond the plant community are likewise expected. For example, a study conducted in these sites at about the same time frame as our studies (J. Kronenberger et al., Berry College, Mount Berry, GA, 2009 unpubl. data) demonstrated a dramatically higher bird abundance and species diversity within managed sites in the BCLPMA, changes which are likely influenced heavily by the changes described here in the vegetative community. Potential negative effects of returning fire to remnant fire-suppressed Longleaf Pine stands must be considered, since burning conditions must be carefully selected in order to avoid post-fire mortality to adult trees via smoldering duff (cf. Cipollini et al. 2008). Conclusions Fire suppression in old-growth mountain Longleaf Pine forests is associated with a dramatic decrease in understory plant diversity, leading to a handful of species adapted to the shadier, less xeric, undisturbed conditions and/or large litter volume on the forest floor. Reduction of canopy, prescribed fire, and hardwood herbicide applications not only help to regenerate mountain Longleaf Pines, but can also dramatically increase diversity of herbaceous plant and grass or grass-like species, particularly those adapted to dry, sunny, and periodically disturbed or burned conditions. Management practices that extensively open canopies may result in the highest overall understory plant species diversity in the early stages of restoration, but loss of some short-lived weedy species might be expected as canopies close following Longleaf Pine establishment or natural regeneration. Canopy closure can be a problem encountered following restoration efforts involving the planting of Longleaf Pine seedlings within a short time frame (even-age stands). Less-intensive management practices such as prescribed burning and targeted herbicide application to non-fire tolerant trees can promote substantial ground-cover restoration while leaving standing trees. Our comparative species list provides baseline date to land managers involved in regional mountain Longleaf Pine restoration efforts involving understory regeneration. Old-growth mountain Longleaf Pine habitats with open canopies and substantial diverse understory plant communities are very rare, so prescribed fire and other management practices must be continued to maintain diverse 650 Southeastern Naturalist Vol. 11, No. 4 communities where such management is possible. Such effects are not restricted to Longleaf Pine habitats, as multiple forest types have been subjected to the negative effects of fire-suppression and mesophication over the last century (Nowacki and Abrams 2008). Acknowledgments We would like to thank the Berry College Student Work Office, Council for Student Scholarship Committee, School of Mathematics and Natural Sciences (Dean D.B. Conn), and the National Fish and Wildlife Foundation/Southern Company Longleaf Legacy program for funding; the Biology Department for providing equipmen; President S. Briggs, the Educational Land Management program, and the Forestry and Land Management office (Director W. Yeomans) for allowing us to work in the BCLPMA and for logistical support (especially with prescribed burning); R. Armstrong, E. Lane, C. McDaniel, and the Spring 2009 Forest Ecology class for helping to collect and identify plant specimens; R. and T. Ware for sharing their Flora of Floyd County data file; and 2 anonymous reviewers for comments on an earlier version of this manuscript. Literature Cited Andrews, F.F. 1917. Agency of fire in propagation of Longleaf Pines. Botanical Gazette 64:497–508. Barbour, M.G., J.H. Burk, W.D. Pitts, F.S. Gilliam, and M.W. Schwartz. 1999. Terrestrial Plant Ecology. Benjamin/Cummings, Menlo Park, CA. 649 pp. Bitterlich, W. 1948. Die winkelzahlprobe. 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Landers, J.L., D.H. van Lear, and W.D. Boyer. 1995. The Longleaf Pine forests of the Southeast: Requiem or renaissance? Journal of Forestry 93:39–44. Maceina, E.C., J.S. Kush, and R.S. Meldahl. 2000. Vegetational survey of a montane Longleaf Pine community at Fort McClellan, Alabama. Castanea 65:147–154. Means, B.D. 1996. Longleaf Pine forest, going, going, …. Pp. 210–229, In M.B. Davis (Ed.). Eastern Old-Growth Forests: Prospects for Rediscovery and Recovery. Island Press, Washington, DC. 400 pp. Newcomb, L. 1977. Newcomb’s Wildflower Guide. Little, Brown, and Company, New York, NY. 490 pp. Nowacki, G.J., and M.D. Abrams. 2008. The demise of fire and “mesophication” of forests in the Eastern United States. BioScience 58:123–138. Peet, R.K., and D.J. Allard. 1993. Longleaf Pine vegetation of the southern Atlantic and eastern Gulf Coast regions. Proceedings of the Tall Timbers Fire Ecology Conference 18:45–81. Platt, W.J., G.W. Evans, and M.M. Davis. 1988. Effects of fire season on flowering of forbs and shrubs in Longleaf Pine forests. Oecologia 76:353–363. Radford, A.E., H.E. Ahles, and C.R. Bell. 1968. Manual of the Vascular Flora of the Carolinas. University of North Carolina Press, Chapel Hill, NC. 1245 pp. Soil Survey Staff, USDA, NRCS. 2011. Web soil survey. Available online at http//websoilsurvey. nrcs.usda.gov/app. Accessed 15 August 2011 Stowe, J.P., Jr., J.M. Varner III, and J.P. McGuire. 2002. Montane Longleaf pinelands … a disappearing treasure. Tipularia 17:8–15. United States Department of Agriculture, Natural Resource Conservation Service (USDA, NRCS). 2011. The PLANTS database. National Plant Data Team, Greensboro, North Carolina. Available online at http://plants.usda.gov. Accessed continuously. University of Texas. 2011. Native plant database. Lady Bird Johnson Wildflower Center, Austin, Texas. Available online at http://www.wildflower.org/plants/. Accessed continuously. Varner, J.M., III. 1999. Longleaf Pine forests … in the mountains? Alabama’s Treasured Forests, Fall 1999:30–31. Varner, J.M., III, and J.S. Kush. 2004. Remnant old-growth Longleaf Pine (Pinus palustris Mill.) savannas and forests of the southeastern USA: Status and threats. Natural Areas Journal 24:141–149. 652 Southeastern Naturalist Vol. 11, No. 4 Varner, J.M., III, J.S. Kush, and R.S. Meldahl. 2000. The mountain Longleaf Pine resources of Fort McClellan, Alabama: Final report on their status, ecology, and management needs. Longleaf Pine Stand Dynamics Laboratory, Auburn University School of Forestry and Wildlife Sciences, Auburn, AL. 83 pp. Varner, J.M., III, J.S. Kush, and R.S. Meldahl. 2003. Structural characteristics of frequently burned old-growth Longleaf Pine stands in the mountains of Alabama. Castanea 68:211–221. Weakley, A.S. 2011. Flora of the Southern and Mid-Atlantic States. May 2011 Edition. UNC Herbarium, North Carolina Botanical Garden, University of North Carolina at Chapel Hill, Chapel Hill, NC. Available online at http://www.herbarium.unc.edu/ flora.htm. Accessed 12 September 2011. Womack, B., and R. Carter. 2011. Landscape-scale forest classification in the Horseblock Mountain Region of the Talladega National Forest, Alabama. Natural Areas Journal 31:51–64. 2012 M.L. Cipollini, J. Culberson, C. Strippelhoff , T. Baldvins, and K. Miller 653 Appendix 1. List of plant species identified within the Berry College Longleaf Pine Management Area from June 2008 through May 2009. Code = PLANTS codes from USDA, NRCS (2011). Family Species Common name Code Apiaceae Sanicula canadensis L. Canadian Blacksnakeroot SACA15 Apocynaceae Apocynum cannabinum L. Indianhemp APCA Asteraceae Achillea millefolium L. Common Yarrow ACMI2 Asteraceae Ageratina altissima (L.) King & H. Rob. White Snakeroot AGALA var. altissima Asteraceae Ageratina aromatica (L.) Spach Lesser Snakeroot AGAR4 Asteraceae Ambrosia artemisiifolia L. Annual Ragweed AMAR2 Asteraceae Antennaria parlinii Fernald ssp. parlinii Parlin's Pussytoes ANPAP Asteraceae Antennaria plantaginifolia (L.) Richardson Woman's Tobacco ANPL Asteraceae Arnoglossum atriplicifolium (L.) H. Rob. Pale Indian Plantain ARAT Asteraceae Chrysopsis mariana (L.) Elliot. Maryland Goldenaster CHMA14 Asteraceae Cirsium arvense (L.) Scop Canada Thistle CIAR4 Asteraceae Cirsium discolor (Muhl. ex Willd.) Spreng. Field Thistle CIDI Asteraceae Cirsium vulgare (Savi) Ten. Bull Thistle CIVU Asteraceae Conyza canadensis (L.) Cronquist var. Canadian Horseweed COCAC3 canadensis Asteraceae Coreopsis major Walter Greater Tickseed COMA6 Asteraceae Elephantopus tomentosus L. Devil’s Grandmother ELTO2 Asteraceae Erechtites hieracifolia (L.) Raf. ex DC. American Burnweed ERHI2 Asteraceae Erigeron annuus Pers. Eastern Daisy Fleabane ERAN Asteraceae Erigeron philadelphicus L. Philadelphia Fleabane ERPH Asteraceae Erigeron strigosus Muhl. ex Willd. Prairie Fleabane ERST3 Asteraceae Eupatorium album L. White Thoroughwort EUAL2 Asteraceae Eupatorium capillifolium (Lam.) Small Dog Fennel EUCA5 Asteraceae Eupatorium hyssopifolium L. Hyssopleaf Thoroughwort EUHY Asteraceae Eupatorium perfoliatum L. Common Boneset EUPE3 Asteraceae Eupatorium rotundifolium L. Roundleaf Thoroughwort EURO4 Asteraceae Eutrochium purpureum (L.) E.E. Lamont Sweetscented Joe Pye EUPU10 Weed Asteraceae Gamochaeta purpurea (L.) Cabrera Spoonleaf Purple GAPU3 Everlasting Asteraceae Helenium amarum (Raf.) H. Rock var. Yellowdicks HEAMA amarum Asteraceae Helianthus divaricatus L. Woodland Sunflower HEDI2 Asteraceae Helianthus mollis Lam. Ashy Sunflower HEMO2 Asteraceae Helianthus angustifolius L. Swamp Sunflower HEAN2 Asteraceae Heterotheca camporum (Greene) Shinners Lemonyellow False HECA16 Goldenaster Asteraceae Heterotheca subaxillaris (Lam.) Britton & Camphorweed HESU3 Rusby Asteraceae Hieracium gronovii L. Queendevil HIGR3 Asteraceae Hieracium piloselloides Vill. Tall Hawkweed HIPI2 Asteraceae Hieracium venosum L. Rattlesnakeweed HIVE Asteraceae Hypochaeris radicata L. Hairy Cat’s Ear HYRA3 Asteraceae Ionactis linariifolius (L.) Greene Flaxleaf Whitetop Aster IOLI2 Asteraceae Krigia biflora (Walter) S.F. Blake Twoflower KRBI Dwarfdandelion Asteraceae Krigia virginica (L.) Willd. Virginia Dwarfdandelion KRVI Asteraceae Lactuca canadensis L. Canada Lettuce LACA Asteraceae Lactuca hirsuta Muhl. ex Nutt Hairy Lettuce LAHI Asteraceae Lactuca serriola L. Prickly Lettuce LASE 654 Southeastern Naturalist Vol. 11, No. 4 Family Species Common name Code Asteraceae Leontodon autumnalis L. Fall Dandelion LEAU2 Asteraceae Liatris pilosa (Aiton) Willd. var. pilosa Shaggy Blazing Star LIPIP Asteraceae Oenothera perennis L. Little Evening Primrose OEPE Asteraceae Packera anonyma (Alph. Wood) W.A. Small's Ragwort PAAN6 Weber & A. Löve Asteraceae Packera obovata (Muhl. ex Willd.) Roundleaf Ragwort PAOB6 W.A. Weber & A. Löve Asteraceae Packera paupercula (Michx.) A. Löve Balsam Groundsel PAPA20 & D. Löve Asteraceae Pityopsis aspera (Shuttlw. ex Small) Pineland Silkgrass PIASA2 Small var. aspera Asteraceae Pityopsis graminifolia (Michx.) Nutt. Narrowleaf Silkgrass PIGRG var. graminifolia Asteraceae Pseudognaphalium helleri (Britton) Heller's Cudweed PSHEH7 Anderb. ssp. helleri Asteraceae Pseudognaphalium obtusifolium (L.) Rabbit-tobacco PSOB3 Hillard & B.L. Burtt Asteraceae Sericocarpus linifolius (L.) Britton, Narrowleaf Whitetop Aster SELI5 Sterns, & Poggenb. Asteraceae Silphium asteriscus L. Starry Rosinweed SIAS2 Asteraceae Solidago bicolor L. White Goldenrod SOBI Asteraceae Solidago nemoralis Aiton Gray Goldenrod SONE Asteraceae Solidago odora Aiton Anisescented Goldenrod SOOD Asteraceae Solidago petiolaris Aiton Downy Ragged Goldenrod SOPE Asteraceae Solidago altissima L. Canada Goldenrod SOAL6 Asteraceae Sonchus asper (L.) Hill Spiny Sowthistle SOAS Asteraceae Sonchus oleraceus L. Common Sowthistle SOOL Asteraceae Symphyotrichum laeve (L.) A. Löve & Smooth Blue Aster SYLAP D. Löve Asteraceae Symphyotrichum lanceolatum (Willd.) White Panicle Aster SYLAI3 G.L. Nesom Asteraceae Symphyotrichum patens (Aiton) G.L. Late Purple Aster SYPA11 Nesom Asteraceae Symphyotrichum racemosum (Elliot) Smooth White Oldfield SYRA5 G.L. Nesom Aster Asteraceae Symphyotrichum dumosum (L.) G.L. Rice Button Aster SYDU2 Nesom Asteraceae Symphyotrichum ontarionis (Wiegand) Bottomland Aster SYON2 G.L. Nesom Asteraceae Symphyotrichum pilosum Willd. Hairy White Oldfield Aster SYPI2 Asteraceae Taraxacum officinale F.H. Wigg. Common Dandelion TAOF Asteraceae Tragopogon lamottei Rouy Jack-go-to-bed-at-noon TRLA30 Asteraceae Verbesina alternifolia (L.) Britton ex Wingstem VEAL Kearney Brassicaceae Cardamine pensylvanica Muhl. ex Pennsylvania Bittercress CAPE3 Willd. Campanulaceae Lobelia nuttallii Schult. Nuttall's Lobelia LONU Campanulaceae Lobelia puberula Michx. Downy Lobelia LOPU Campanulaceae Lobelia siphilitica L. Great Blue Lobelia LOSI Campanulaceae Triodanis biflora (Ruiz & Pav.) Greene Small Venus’ TRBI2 Looking-glass Campanulaceae Triodanis perfoliata (L.) Nieuwl. Clasping Venus’ TRPE4 Looking-glass Caryophyllaceae Silene virginica L. Fire Pink SIVI4 2012 M.L. Cipollini, J. Culberson, C. Strippelhoff , T. Baldvins, and K. Miller 655 Family Species Common name Code Chenopodiaceae Chenopodium album L. Lambsquarters CHAL7 Clusiaceae Hypericum drummondii (Grev. & Hook.) Nits and Lice HYDR Torr. & A. Gray Clusiaceae Hypericum gentianoides (L.) Britton, Orangegrass HYGE Sterns, & Poggenb. Clusiaceae Hypericum hypericoides (L.) Crantz St. Andrew’s Cross HYHYH ssp. hypericoides Clusiaceae Hypericum punctatum Lam. Spotted St. Johnswort HYPU Commelinaceae Tradescantia virginiana L. Virginia Spiderwort TRVI Cyperaceae Carex comosa Boott Longhair Sedge CACO8 Cyperaceae Carex squarrosa L. Squarrose Sedge CASQ2 Cyperaceae Scleria pauciflora Muhl. ex Willd. Fewflower Nutrush SCPA5 Cyperaceae Scleria reticularis Michx. Netted Nutrush SCRE Cyperaceae Scleria triglomerata Michx. Whip Nutrush SCTR Ericaceae Chimaphila maculata (L.) Pursh Striped Prince’s Pine CHMA3 Ericaceae Epigaea repens L. Trailing Arbutus EPRE2 Euphorbiaceae Cnidoscolus urens (L.) Arthur var. Finger Rot CNURS stimulosus (Michx.) Govaerts Euphorbiaceae Euphorbia corollata L. Flowering Spurge EUCO10 Euphorbiaceae Euphorbia pubentissima Michx. False Flowering Spurge EUPU7 Fabaceae Chamaecrista nictitans (L.) Moench Sensitive Partidge Pea CHNI2 Fabaceae Clitoria mariana L. Atlantic Pigeonwings CLMA4 Fabaceae Crotalaria sagittalis L. Arrowhead Rattlebox CRSA4 Fabaceae Lespedeza capitata Michx. Roundhead Lespedeza LECA8 Fabaceae Lespedeza frutescens (L.) Hornem. Shrubby Lespedeza LEFR5 Fabaceae Lespedeza hirta (L.) Hornem. Hairy Lespedeza LEHI2 Fabaceae Lespedeza procumbens Michx. Trailing Lespedeza LEPR Fabaceae Lespedeza repens (L.) W. Bartram Creeping Lespedeza LERE2 Fabaceae Lespedeza violacea (L.) Pers. Violet Lespedeza LEVI6 Fabaceae Lespedeza virginica (L.) Britton Slender Lespedeza LEVI7 Fabaceae Melilotus officinales (L.) Lam. Yellow Sweetclover MEOF Fabaceae Mimosa microphylla Torr. & A. Gray Littleleaf Sensitive Briar MIMI22 Fabaceae Strophostyles umbellata (Muhl. ex Pink Fuzzybean STUM2 Willd.) Britton Fabaceae Tephrosia virginiana L. Virginia Tephrosia TEVI Fabaceae Trifolium campestre Schreb. Field Clover TRCA5 Fabaceae Trifolium dubium Sibth. Suckling Clover TRDU2 Fabaceae Trifolium reflexum L. Buffalo Clover TRRE2 Geraniaceae Geranium carolinianum L. Carolina Geranium GECA5 Iridaceae Sisyrinchium albidum Raf. White Blue-eyed Grass SIAL3 Iridaceae Sisyrinchium angustifolium Mill. Narrowleaf Blue-eyed Grass SIAN3 Iridaceae Sisyrinchium montanum Greene Strict Blue-eyed Grass SIMO2 Juncaceae Juncus canadensis J. Gay ex Laharpe Canadian Rush JUCA3 Juncaceae Juncus effusus L. Common Rush JUEF Lamiaceae Mentha X piperita L. Peppermint MEPI Lamiaceae Monarda fistulosa L. Wild Bergamot MOFI Lamiaceae Prunella vulgaris L. Common Selfheal PRVU Lamiaceae Pycnanthemum muticum (Michx.) Pers. Clustered Mountainmint PYMU Lamiaceae Salvia lyrata L. Lyreleaf Sage SALY2 Lamiaceae Scutellaria elliptica Muhl. ex Spreng Hairy Skullcap SCEL Lamiaceae Scutellaria integrifolia L. Helmet Flower SCIN2 Lamiaceae Scutellaria lateriflora L. Blue Skullcap SCLA2 Lamiaceae Trichostema dichotomum L. Forked Bluecurls TRDI2 Liliaceae Hypoxis hirsuta (L.) Coville Common Goldstar HYHI2 656 Southeastern Naturalist Vol. 11, No. 4 Family Species Common name Code Linaceae Linum virginianum L. Woodland Flax LIVI Melastomataceae Rhexia virginica L. Handsome Harry RHVI Melostomataceae Rhexia mariana L. Maryland Meadowbeauty RHMA Oxalidaceae Oxalis corniculata L. Creeping Woodsorrel OXCO Oxalidaceae Oxalis dillenii Jacq. Slender Yellow Woodsorrel OXDI2 Oxalidaceae Oxalis grandis Small Great Yellow Woodsorrel OXGR Oxalidaceae Oxalis montana Raf. Mountain Woodsorrel OXMO Oxalidaceae Oxalis stricta L. Common Yellow Oxalis OXST Oxalidaceae Oxalis violacea L. Violet Woodsorrel OXVI Passifloraceae Passiflora incarnata L. Purple Passionflower PAIN6 Phytolaccaceae Phytolacca americana L. American Pokeweed PHAM4 Plantaginaceae Plantago aristata Michx. Largebracted Plantain PLAR3 Poaceae Andropogon gerardii Vitman Big Bluestem ANGE Poaceae Andropogon virginicus L. Broomsedge Bluestem ANVI2 Poaceae Andropogon capillipes Nash Chalky Bluestem ANCA4 Poaceae Chasmanthium sessiliflorum (Poir.) Longleaf Woodoats CHSE2 Yates Poaceae Dichanthelium acuminatum Gould & Tapered Rosette Grass DIACF C.A. Clark var. fasciculatum (Torr.) Freckmann Poaceae Dichanthelium boscii (Poir.) Gould & Bosc's Panicgrass DIBO2 C.A. Clark Poaceae Dichanthelium commutatum (Schult.) Variable Panicgrass DICO2 Gould Poaceae Dichanthelium depauperatum (Muhl.) Starved Panicgrass DIDE4 Gould Poaceae Dichanthelium dichotomum (L.) Gould Cypress Panicgrass DIDID var. dichotomum Poaceae Dichanthelium laxiflorum (Lam.) Gould Openflower Rosette Grass DILA19 Poaceae Dichanthelium sabulorum (Lam.) Gould Hemlock Rosette Grass DISAP & C.A. Clark var. patulum (Scribn. & Merr.) Gould & C.A. Clark Poaceae Dichanthelium scoparium (Lam.) Gould Velvet Panicgrass DISC3 Poaceae Dichanthelium sphaerocarpon (Elliot) Roundseed Panicgrass DISPI Gould var. isophyllum (Scribn.) Gould & C.A. Clark Poaceae Dichanthelium villosissimum (Nash) Whitehair Rosette Grass DIVIV Freckmann var. villosissimum Poaceae Eragrostis frankii C.A. Mey. ex Steud Sandbar Lovegrass ERFR Poaceae Eragrostis refracta (Muhl.) Scribn. Coastal Lovegrass ERRE Poaceae Melica mutica Walter Twoflower Melicgrass MEMU Poaceae Panicum amarum Elliot Bitter Panicgrass PAAM2 Poaceae Panicum anceps Michx. Beaked Panicgrass PAAN Poaceae Panicum capillare L. Witchgrass PACA6 Poaceae Panicum dichotomiflorum Michx. Fall Panicgrass PADI Poaceae Panicum flexile (Gattinger) Scribn. Wiry Panicgrass PAFL2 Poaceae Panicum philadelphicum Bernh. ex Trin. Philadelphia Panicgrass PAPH Poaceae Panicum virgatum L. Switchgrass PAVI2 Poaceae Paspalum floridanum Michx. Florida Paspalum PAFL4 Poaceae Piptochaetium avenaceum (L.) Parodi Blackseed Speargrass PIAV Poaceae Poa trivialis L. Rough Bluegrass POTR2 Poaceae Saccharum alopecuroides (L.) Nutt. Silver Plumegrass SAAL21 Poaceae Saccharum baldwinii Spreng. Narrow Plumegrass SABA10 2012 M.L. Cipollini, J. Culberson, C. Strippelhoff , T. Baldvins, and K. Miller 657 Family Species Common name Code Poaceae Saccharum brevibarbe (Michx.) Pers. Sortbeard Plumegrass SABRC3 var. contortum (Elliot) R. Webster Poaceae Schizachyrium scoparium (Michx.) Nash Little Bluestem SCSC Poaceae Setaria viridis (L.) P. Beauv. Green Bristlegrass SEVI4 Poaceae Sorghastrum nutans (L.) Nash Indiangrass SONU2 Poaceae Sorghum halepense (L.) Pers. Johnsongrass SOHA Poaceae Sporobolus indicus (L.) R. Br. var. Smut Grass SPINI2 indicus Polemoniaceae Phlox divaricata L. Wild Blue Phlox PHDI5 Polygalaceae Polygala ambigua Nutt. Whorled Milkwort POAM9 Polygalaceae Polygala curtissii A. Gray Curtiss' Milkwort POCU5 Polygalaceae Polygala nuttallii Torr. & A. Gray Nuttall's Milkwort PONU2 Polygonaceae Polygonum cespitosum Blume, nom. Inq. Oriental Ladysthumb POCE4 Polygonaceae Polygonum persicaria L. Spotted Ladysthumb POPE3 Polygonaceae Rumex acetosella L. Common Sheep Sorrel RUAC3 Portulacaceae Claytonia viriginica L. Virginia Springbeauty CLVI3 Ranunculaceae Ranunculus recurvatus Poir. Blisterwort RARE2 Rosaceae Rubus flagellaris Willd. var. almus Northern Dewberry RUFL L.H. Bailey Rosaceae Waldsteinia fragariodes (Michx.) Tratt. Appalachian Barren WAFR Strawberry Rubiaceae Diodia teres Walter Poorjoe DITE2 Rubiaceae Galium lanceolatum Torr. Lanceleaf Wild Licorice GALA3 Rubiaceae Galium pilosum Aiton Hairy Bedstraw GAPI2 Rubiaceae Houstonia caerulea L. Azure Bluet HOCA4 Rubiaceae Houstonia longifolia Gaertn. Longleaf Summer Bluet HOLO Rubiaceae Houstonia purpurea L. Venus’ Pride HOPU2 Rubiaceae Houstonia serpyllifolia Michx. Thymeleaf Bluet HOSE2 Scrophulariaceae Chaenorhinum minus (L.) Lange Dwarf Snapdragon CHMI Scrophulariaceae Veronica arvensis L. Corn Speedwell VEAR Scrophulariaceae Veronica peregrina L. Neckweed VEPE2 Solanacea Solanum carolinense L. Carolina Horsenettle SOCA3 Verbenaceae Verbena hastata L. Swamp Verbena VEHA2 Verbenaceae Verbena simplex Lehm. Narrowleaf Vervain VESI Verbenaceae Verbena stricta Vent. Hoary Verbena VEST Violaceae Viola bicolor Pursh Field Pansy VIBI Violaceae Viola pedata L. Birdfoot Violet VIPE 658 Southeastern Naturalist Vol. 11, No. 4 Appendix 2. Plant species reported within mountain Longleaf Pine habitats, in comparison with those recorded in this study and in Wares’ Flora of Floyd County, GA. Species ecological characteristics are derived from Radford et al. (1968), University of Texas (2011), USDA, NRCS (2011), and Weakley (2011). H = Harper 19051, LM 1917 = Lavender Mountain 172, LM 40 = Lavender Mountain 19403, FM = Fort McClellan 20004, TM = Thunder Mountain 20065, HM = Horseblock Mountain 20106, LM 09 = Lavender Mountain 2009 (this study), LM UM = Lavender Mountain UM stands, LM IM = Lavender Mountain IM stands “top 20”, LM HM = Lavender Mountain HM stands “top 20”, FC = Floyd County, GA 20067, ST = sun-tolerant (open canopy), ShT = shade-tolerant (closed canopy), P = perennial, B = biennial, A = annual, D = dry soil, M = moist soil, Di = disturbed sites, and I = invasive or alien species. LM LM LM LM LM LM Family Species H 17 40 FM TM HB 09 UM IM HM FC ST ShT P B A D M Di I Species group 1: Species found in our site currently, in our site in the past, or in other mountain Longleaf Pine sites. Apocynaceae Apocynum cannabinum 1 1 1 1 1 1 1 Asteraceae Ageratina aromatic 1 1 1 1 1 1 1 1 Asteraceae Antennaria plantaginifolia 1 1 1 1 1 1 1 Asteraceae Chrysopsis mariana 1 1 1 1 1 1 1 1 Asteraceae Coreopsis major 1 1 1 1 1 1 1 1 1 1 1 1 Asteraceae Elephantopus tomentosus 1 1 1 1 1 1 1 1 Asteraceae Erigeron strigosus 1 1 1 1 1 1 1 1 1 1 1 1 Asteraceae Eupatorium album 1 1 1 1 1 1 1 1 Asteraceae Eupatorium perfoliatum 1 1 1 1 1 1 1 Asteraceae Gamochaeta purpurea 1 1 1 1 1 1 1 1 Asteraceae Helianthus divaricatus 1 1 1 1 1 1 1 1 1 Asteraceae Helianthus mollis 1 1 1 1 1 1 1 1 Asteraceae Hieracium venosum 1 1 1 1 1 1 1 1 1 1 Asteraceae Krigia biflora 1 1 1 1 1 1 1 Asteraceae Krigia virginica 1 1 1 1 1 1 1 1 Asteraceae Liatris pilosa 1 1 1 1 1 1 Asteraceae Packera anonyma 1 1 1 1 1 1 1 1 1 Asteraceae Pityopsis graminifolia 1 1 1 1 1 1 1 1 1 1 Asteraceae Pseudognaphalium obtusifolium 1 1 1 1 1 1 1 1 Asteraceae Sericocarpus linifolius 1 1 1 1 1 1 1 1 1 Asteraceae Silphium asteriscus 1 1 1 1 1 1 1 1 1 Asteraceae Solidago bicolor 1 1 1 1 1 Asteraceae Solidago nemoralis 1 1 1 1 1 1 1 Asteraceae Solidago odora 1 1 1 1 1 1 1 1 1 1 1 1 2012 M.L. Cipollini, J. Culberson, C. Strippelhoff , T. Baldvins, and K. Miller 659 LM LM LM LM LM LM Family Species H 17 40 FM TM HB 09 UM IM HM FC ST ShT P B A D M Di I Asteraceae Symphyotrichum dumosum 1 1 1 1 1 1 1 1 1 1 1 Asteraceae Symphyotrichum laeve 1 1 1 1 1 1 1 1 Asteraceae Symphyotrichum patens 1 1 1 1 1 1 1 1 1 1 Campanulaceae Triodanis perfoliata 1 1 1 1 1 1 1 Clusiaceae Hypericum gentianoides 1 1 1 1 1 1 1 1 Clusiaceae Hypericum hypericoides 1 1 1 1 1 1 1 1 1 1 Clusiaceae Hypericum punctatum 1 1 1 1 1 1 1 1 Cyperaceae Scleria triglomerata 1 1 1 1 1 1 1 Euphorbiaceae Cnidoscolus urens 1 1 1 1 1 Euphorbiaceae Euphorbia corollata 1 1 1 1 1 1 1 1 1 Euphorbiaceae Euphorbia pubentissima 1 1 1 1 1 1 1 Fabaceae Chamaecrista nictitans 1 1 1 1 1 1 1 1 Fabaceae Clitoria mariana 1 1 1 1 1 1 1 1 1 1 1 Fabaceae Lespedeza frutescens 1 1 1 1 1 1 1 1 Fabaceae Lespedeza hirta 1 1 1 1 1 1 1 Fabaceae Lespedeza procumbens 1 1 1 1 1 1 1 1 1 Fabaceae Lespedeza repens 1 1 1 1 1 1 1 Fabaceae Lespedeza virginica 1 1 1 1 1 1 1 1 Fabaceae Mimosa microphylla 1 1 1 1 1 1 1 1 Fabaceae Tephrosia virginiana 1 1 1 1 1 1 1 1 1 1 Fabaceae Trifolium campestre 1 1 1 1 1 1 1 1 1 1 Fabaceae Trifolium dubium 1 1 1 1 1 1 1 1 1 Geraniaceae Geranium carolinianum 1 1 1 1 1 1 1 1 1 Iridaceae Sisyrinchium angustifolium 1 1 1 1 1 1 1 Juncaceae Juncus effusus 1 1 1 1 1 1 Lamiaceae Prunella vulgaris 1 1 1 1 1 1 1 1 1 Lamiaceae Salvia lyrata 1 1 1 1 1 1 1 1 1 Lamiaceae Scutellaria elliptica 1 1 1 1 1 1 1 1 Lamiaceae Scutellaria integrifolia 1 1 1 1 1 1 1 Liliaceae Hypoxis hirsuta 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Oxalidaceae Oxalis corniculata 1 1 1 1 1 1 1 Oxalidaceae Oxalis stricta 1 1 1 1 1 1 1 1 1 1 1 660 Southeastern Naturalist Vol. 11, No. 4 LM LM LM LM LM LM Family Species H 17 40 FM TM HB 09 UM IM HM FC ST ShT P B A D M Di I Passifloraceae Passiflora incarnata 1 1 1 1 1 1 1 1 Poaceae Andropogon gerardii 1 1 1 1 1 1 1 1 Poaceae Andropogon virginicus 1 1 1 1 1 1 1 1 1 1 Poaceae Dichanthelium commutatum 1 1 1 1 1 1 1 1 1 1 Poaceae Dichanthelium sphaerocarpon 1 1 1 1 1 1 1 1 1 Poaceae Dichanthelium villosissimum 1 1 1 1 1 1 Poaceae Melica mutica 1 1 1 1 1 1 Poaceae Panicum virgatum 1 1 1 1 1 1 Poaceae Piptochaetium avenaceum 1 1 1 1 1 1 1 1 1 1 1 Poaceae Schizachyrium scoparium 1 1 1 1 1 1 1 1 1 1 Poaceae Sorghastrum nutans 1 1 1 1 1 1 1 1 1 Polemoniaceae Phlox divaricata 1 1 1 1 1 1 Polygonaceae Rumex acetosella 1 1 1 1 1 1 1 1 1 Pyrolaceae Chimaphila maculata 1 1 1 1 1 1 1 1 Rubiaceae Galium pilosum 1 1 1 1 1 1 1 Rubiaceae Houstonia caerulea 1 1 1 1 1 1 1 1 1 1 1 Rubiaceae Houstonia purpurea 1 1 1 1 1 1 1 1 Violaceae Viola pedata 1 1 1 1 1 1 1 1 1 Total group 1 65 37 64 2 11 59 33 23 5 Species group 2: Species present in other mountain Longleaf Pine sites and in Ware’s Flora of Floyd County. Acanthaceae Ruellia caroliniensis 1 1 1 1 1 1 (J.F. Gmel.) Steud. Apiaceae Angelica venenosa 1 1 1 1 1 1 1 1 (Greenway) Fernald Apiaceae Daucus carota L. 1 1 1 1 1 1 1 Apiaceae Eryngium yuccifolium Michx. 1 1 1 1 1 Aristolochiaceae Hexastylis arifolia (Michx.) 1 1 1 1 1 1 Small Asclepiadaceae Asclepias tuberose L. 1 1 1 1 1 1 1 Asclepiadaceae Asclepias viridis Walter 1 1 1 1 1 1 Asclepiadaceae Asclepias amplexicaulis Sm. 1 1 1 1 1 2012 M.L. Cipollini, J. Culberson, C. Strippelhoff , T. Baldvins, and K. Miller 661 LM LM LM LM LM LM Family Species H 17 40 FM TM HB 09 UM IM HM FC ST ShT P B A D M Di I Asteraceae Brickellia eupatorioides (L.) 1 1 1 1 1 1 Shinners Asteraceae Coreopsis lanceolata L. 1 1 1 1 1 Asteraceae Coreopsis verticillata L. 1 1 1 1 1 Asteraceae Eurybia surculosa (Michx.) 1 1 1 1 1 1 G.L. Nesom Asteraceae Helianthus atrorubens L. 1 1 1 1 1 1 Asteraceae Helianthus hirsutus Raf. 1 1 1 1 1 1 1 Asteraceae Helianthus microcephalus 1 1 1 1 1 1 1 Torr. & A. Gray Asteraceae Liatris spicata (L.) Willd. 1 1 1 1 1 Asteraceae Marshallia obovata (Walter) 1 1 1 1 1 Beadle & F.E. Boynt. Asteraceae Parthenium integrifolium L. 1 1 1 1 1 1 Asteraceae Prenanthes serpentaria 1 1 1 1 1 1 1 Pursh Asteraceae Rudbeckia hirta L. 1 1 1 1 1 1 1 1 1 Asteraceae Sericocarpus asteroides (L.) 1 1 1 1 1 1 Britton, Sterns & Poggenb. Asteraceae Solidago arguta Aiton 1 1 1 1 1 Asteraceae Solidago erecta Pursh 1 1 1 1 1 1 Asteraceae Symphyotrichum lateriflorum 1 1 1 1 1 1 1 (L.) Á. Löve & D. Löve Asteraceae Symphyotrichum undulatum 1 1 1 1 1 1 (L.) G.L. Nesom Asteraceae Vernonia flaccidifolia Small 1 1 1 1 1 1 1 1 Campanulaceae Lobelia spicata Lam. 1 1 1 1 1 1 1 Convolvulaceae Ipomoea pandurata (L.) 1 1 1 1 1 1 G. Mey. Cyperaceae Cyperus retrofractus (L.) 1 1 1 1 1 1 Torr. Cyperaceae Eleocharis obtuse (Willd.) 1 1 1 1 1 1 Schult. 662 Southeastern Naturalist Vol. 11, No. 4 LM LM LM LM LM LM Family Species H 17 40 FM TM HB 09 UM IM HM FC ST ShT P B A D M Di I Dioscoreaceae Dioscorea villosa L. 1 1 1 1 1 1 Fabaceae Baptisia bracteata Muhl. 1 1 1 1 1 ex Elliott Fabaceae Desmodium nudiflorum 1 1 1 1 1 1 Fabaceae Desmodium paniculatum 1 1 1 1 1 1 (L.) DC. Fabaceae Galactia volubilis (L.) Britton 1 1 1 1 1 1 1 Fabaceae Lathyrus hirsutus L. 1 1 1 1 1 1 1 Fabaceae Orbexilum pedunculatum (Mill.) 1 1 1 1 1 1 Rydb. Fabaceae Rhynchosia tomentosa (L.) 1 1 1 1 1 1 Hook. & Arn. Fabaceae Trifolium hybridum L. 1 1 1 1 1 1 1 1 Fabaceae Vicia caroliniana Walter 1 1 1 1 1 Gentianaceae Gentiana villosa L. 1 1 1 1 1 Iridaceae Iris verna L. 1 1 1 1 1 Iridaceae Sisyrinchium atlanticum E.P. 1 1 1 1 1 Bicknell Lamiaceae Monarda clinopodia L. 1 1 1 1 1 1 Lamiaceae Pycnanthemum incanum (L.) 1 1 1 1 1 1 1 Michx. Lamiaceae Salvia urticifolia L. 1 1 1 1 1 Lamiaceae Scutellaria ovate Hill 1 1 1 1 1 Liliaceae Aletris farinose L. 1 1 1 1 1 1 1 Liliaceae Maianthemum racemosum (L.) 1 1 1 1 1 1 Link Liliaceae Polygonatum biflorum (Walter) 1 1 1 1 1 1 Elliott Liliaceae Trillium catesbaei Elliott 1 1 1 1 1 Linaceae Linum medium (Planch.) Britton 1 1 1 1 1 1 1 1 Loganiaceae Spigelia marilandica (L.) L. 1 1 1 1 1 1 Onagraceae Oenothera fruticosa L. 1 1 1 1 1 1 2012 M.L. Cipollini, J. Culberson, C. Strippelhoff , T. Baldvins, and K. Miller 663 LM LM LM LM LM LM Family Species H 17 40 FM TM HB 09 UM IM HM FC ST ShT P B A D M Di I Orchidaceae Malaxis unifolia Michx. 1 1 1 1 1 1 1 Poaceae Agrostis hyemalis (Walter) 1 1 1 1 1 1 1 Britton, Sterns, & Poggenb. Poaceae Andropogon ternarius Michx. 1 1 1 1 1 1 1 Poaceae Aristida purpurascens var. 1 1 1 1 virgata (Trin.) Allred Poaceae Danthonia sericea Nutt. 1 1 1 1 1 1 Poaceae Dichanthelium aciculare (Desv. 1 1 1 1 1 ex Poir.) Gould & C.A. Clark Poaceae Dichanthelium angustifolium 1 1 1 1 1 (Elliott) Gould Poaceae Gymnopogon ambiguus 1 1 1 1 1 (Michx.) Britton, Sterns, & Poggenb. Poaceae Poa autumnalis Muhl. ex Elliott 1 1 1 1 1 1 Polemoniaceae Phlox pilosa L. 1 1 1 1 1 1 1 Polygalaceae Polygala verticillata L. 1 1 1 1 1 1 Ranunculaceae Thalictrum thalictroides (L.) 1 1 1 1 1 Eames & B. Boivin Rosaceae Duchesnea indica (Andrews) 1 1 1 1 1 1 1 Focke Rosaceae Fragaria virginiana Duchesne 1 1 1 1 1 Rosaceae Gillenia stipulate (Muhl. ex Willd.) 1 1 1 1 1 Baill. Rosaceae Potentilla canadensis L. 1 1 1 1 1 1 1 1 Rosaceae Potentilla simplex Michx. 1 1 1 1 1 1 1 Rosaceae Rubus flagellaris Willd. 1 1 1 1 1 1 1 Rubiaceae Galium aparine L. 1 1 1 1 1 1 Rubiaceae Galium lanceolatum Torr. 1 1 1 1 1 1 Scrophulariaceae Aureolaria flava (L.) Farw. 1 1 1 1 1 1 1 Scrophulariaceae Aureolaria pectinata (Nutt.) 1 1 1 1 1 Pennell 664 Southeastern Naturalist Vol. 11, No. 4 LM LM LM LM LM LM Family Species H 17 40 FM TM HB 09 UM IM HM FC ST ShT P B A D M Di I Scrophulariaceae Penstemon canescens (Britton) 1 1 1 1 1 1 1 1 Britton Solanacea Physalis virginiana Mill. 1 1 1 1 1 1 1 1 Valerianaceae Valerianella radiate (L.) Dufr. 1 1 1 1 1 1 1 Violaceae Viola affinis Leconte 1 1 1 1 1 Violaceae Viola x palmata L. 1 1 1 1 1 Violaceae Viola sagittata Aiton 1 1 1 1 1 1 Violaceae Viola sororia Willd. 1 1 1 1 1 1 1 Total group 2 63 51 72 4 12 62 49 18 5 Species group 3. Species found in our study and in Ware’s Flora of Floyd County, but not recorded at Lavender Mountain in the past and not at any other mountain Longleaf Pine site. Apiaceae Sanicula canadensis 1 1 1 1 1 1 Asteraceae Achillea millefolium 1 1 1 1 1 1 1 Asteraceae Ageratina altissima var. altissima 1 1 1 1 1 Asteraceae Ambrosia artemisiifolia 1 1 1 1 1 1 1 Asteraceae Arnoglossum atriplicifolium 1 1 1 1 1 1 1 Asteraceae Cirsium discolor 1 1 1 1 1 1 1 1 Asteraceae Cirsium vulgare 1 1 1 1 1 1 1 Asteraceae Conyza canadensis 1 1 1 1 1 1 1 1 Asteraceae Erechtites hieracifolia 1 1 1 1 1 1 1 1 1 Asteraceae Erigeron annuus 1 1 1 1 1 Asteraceae Erigeron philadelphicus 1 1 1 1 1 1 1 Asteraceae Eupatorium capillifolium 1 1 1 1 1 1 1 Asteraceae Eupatorium hyssopifolium 1 1 1 1 1 1 1 1 Asteraceae Eupatorium rotundifolium 1 1 1 1 1 1 Asteraceae Eupatorium purpureum 1 1 1 1 1 1 Asteraceae Helenium amarum 1 1 1 1 1 1 1 Asteraceae Helianthus angustifolius 1 1 1 1 1 1 Asteraceae Heterotheca camporum 1 1 1 1 1 1 Asteraceae Hieracium gronovii 1 1 1 1 1 1 1 Asteraceae Hypochaeris radicata 1 1 1 1 1 1 1 2012 M.L. Cipollini, J. Culberson, C. Strippelhoff , T. Baldvins, and K. Miller 665 LM LM LM LM LM LM Family Species H 17 40 FM TM HB 09 UM IM HM FC ST ShT P B A D M Di I Asteraceae Ionactis linariifolius 1 1 1 1 1 1 Asteraceae Lactuca canadensis 1 1 1 1 1 1 1 Asteraceae Lactuca serriola 1 1 1 1 1 1 1 1 Asteraceae Packera obovata 1 1 1 1 1 1 Asteraceae Packera paupercula 1 1 1 1 1 1 1 Asteraceae Pityopsis aspera 1 1 1 1 1 Asteraceae Pseudognaphalium helleri 1 1 1 1 1 Asteraceae Solidago petiolaris 1 1 1 1 1 1 Asteraceae Solidago altissima 1 1 1 1 1 1 1 Asteraceae Sonchus asper 1 1 1 1 1 1 Asteraceae Sonchus oleraceus 1 1 1 1 1 1 Asteraceae Symphyotrichum ontarionis 1 1 1 1 1 1 Asteraceae Symphyotrichum pilosum 1 1 1 1 1 1 1 Asteraceae Taraxacum officinale 1 1 1 1 1 1 1 1 Asteraceae Verbesina alternifolia 1 1 1 1 1 1 Brassicaceae Cardamine pensylvanica 1 1 1 1 1 1 Campanulaceae Lobelia puberula 1 1 1 1 1 1 1 1 Campanulaceae Lobelia siphilitica 1 1 1 1 1 1 Campanulaceae Triodanis biflora 1 1 1 1 1 1 1 Caryophyllaceae Silene virginica 1 1 1 1 1 1 1 1 Chenopodiaceae Chenopodium album 1 1 1 1 1 1 Clusiaceae Hypericum drummondii 1 1 1 1 1 1 Commelinaceae Tradescantia virginiana 1 1 1 1 1 1 1 Cyperaceae Carex squarrosa 1 1 1 1 1 1 Ericaceae Epigaea repens 1 1 1 1 1 1 1 Fabaceae Crotalaria sagittalis 1 1 1 1 1 1 1 1 1 Fabaceae Lespedeza capitata 1 1 1 1 1 Fabaceae Lespedeza violacea 1 1 1 1 1 1 Fabaceae Melilotus officinales 1 1 1 1 1 1 1 1 1 1 Fabaceae Strophostyles umbellata 1 1 1 1 1 1 1 1 Fabaceae Trifolium reflexum 1 1 1 1 1 1 1 Iridaceae Sisyrinchium albidum 1 1 1 1 1 666 Southeastern Naturalist Vol. 11, No. 4 LM LM LM LM LM LM Family Species H 17 40 FM TM HB 09 UM IM HM FC ST ShT P B A D M Di I Lamiaceae Mentha x piperita 1 1 1 1 1 1 1 1 Lamiaceae Monarda fistulosa 1 1 1 1 1 1 1 1 Lamiaceae Pycnanthemum muticum 1 1 1 1 1 1 Lamiaceae Trichostema dichotomum 1 1 1 1 1 1 1 1 Linaceae Linum virginianum 1 1 1 1 1 1 1 Melastomataceae Rhexia virginica 1 1 1 1 1 Melostomataceae Rhexia mariana 1 1 1 1 1 Oxalidaceae Oxalis dillenii 1 1 1 1 1 1 1 1 Oxalidaceae Oxalis montana 1 1 1 1 1 Oxalidaceae Oxalis violacea 1 1 1 1 1 1 1 1 Phytolaccaceae Phytolacca americana 1 1 1 1 1 1 1 Plantaginaceae Plantago aristata 1 1 1 1 1 1 Poaceae Chasmanthium sessiliflorum 1 1 1 1 1 Poaceae Dichanthelium boscii 1 1 1 1 1 1 1 1 Poaceae Dichanthelium depauperatum 1 1 1 1 1 1 1 Poaceae Dichanthelium dichotomum 1 1 1 1 1 1 1 Poaceae Dichanthelium laxiflorum 1 1 1 1 1 1 1 Poaceae Dichanthelium scoparium 1 1 1 1 1 1 Poaceae Eragrostis frankii 1 1 1 1 1 1 1 Poaceae Eragrostis refracta 1 1 1 1 1 Poaceae Panicum anceps 1 1 1 1 1 1 Poaceae Panicum capillare 1 1 1 1 1 1 1 Poaceae Panicum dichotomiflorum 1 1 1 1 1 1 1 Poaceae Panicum flexile 1 1 1 1 1 1 Poaceae Paspalum floridanum 1 1 1 1 1 Poaceae Saccharum alopecuroides 1 1 1 1 1 1 Poaceae Saccharum baldwinii 1 1 1 1 1 Poaceae Saccharum brevibarbe 1 1 1 1 1 Poaceae Sorghum halepense 1 1 1 1 1 1 1 1 Poaceae Sporobolus indicus 1 1 1 1 1 1 1 1 Polygalaceae Polygala ambigua 1 1 1 1 1 1 1 Polygalaceae Polygala curtissii 1 1 1 1 1 1 1 2012 M.L. Cipollini, J. Culberson, C. Strippelhoff , T. Baldvins, and K. Miller 667 LM LM LM LM LM LM Family Species H 17 40 FM TM HB 09 UM IM HM FC ST ShT P B A D M Di I Polygalaceae Polygonum cespitosum 1 1 1 1 1 1 1 1 Polygalaceae Polygonum persicaria 1 1 1 1 1 1 1 Portulacaceae Claytonia viriginica 1 1 1 1 1 Ranunculaceae Ranunculus recurvatus 1 1 1 1 1 Rosaceae Waldsteinia fragariodes 1 1 1 1 1 1 1 Rubiaceae Diodia teres 1 1 1 1 1 1 Rubiaceae Houstonia longifolia 1 1 1 1 1 1 Scrophulariaceae Veronica arvensis 1 1 1 1 1 1 1 1 Scrophulariaceae Veronica peregrina 1 1 1 1 1 1 1 Solanacea Solanum carolinense 1 1 1 1 1 1 1 Verbenaceae Verbena simplex 1 1 1 1 1 1 1 Verbenaceae Verbena stricta 1 1 1 1 1 1 1 Violaceae Viola bicolor 1 1 1 1 1 1 Total group 3 87 39 66 10 31 64 72 50 12 Species group 4: Species recorded in our study, but not in other mountain Longleaf Pine sites and not in Ware’s Flora of Floyd County. Asteraceae Antennaria parlinii 1 1 1 1 Asteraceae Cirsium arvense 1 1 1 1 1 1 1 Asteraceae Heterotheca subaxillaris 1 1 1 1 1 1 Asteraceae Hieracium piloselloides 1 1 1 1 1 1 1 1 1 Asteraceae Lactuca hirsuta 1 1 1 1 1 1 Asteraceae Leontodon autumnalis 1 1 1 1 1 1 1 Asteraceae Symphyotrichum lanceolatum 1 1 1 1 Asteraceae Symphyotrichum racemosum 1 1 1 1 1 Asteraceae Tragopogon lamottei 1 1 1 1 1 1 1 Campanulaceae Lobelia nuttallii 1 1 1 1 Cyperaceae Carex comosa 1 1 1 1 Cyperaceae Scleria pauciflora 1 1 1 1 Cyperaceae Scleria reticularis 1 1 1 1 1 Iridaceae Sisyrinchium montanum 1 1 1 1 Juncaceae Juncus canadensis 1 1 1 1 Lamiaceae Scutellaria lateriflora 1 1 1 1 1 668 Southeastern Naturalist Vol. 11, No. 4 LM LM LM LM LM LM Family Species H 17 40 FM TM HB 09 UM IM HM FC ST ShT P B A D M Di I Oxalidaceae Oxalis grandis 1 1 1 1 Poaceae Andropogon capillipes 1 1 1 1 Poaceae Dichanthelium acuminatum 1 1 1 1 Poaceae Dichanthelium sabulorum 1 1 1 1 1 Poaceae Panicum amarum 1 1 1 1 Poaceae Panicum philadelphicum 1 1 1 1 1 1 Poaceae Poa trivialis 1 1 1 1 1 1 1 Poaceae Setaria viridis 1 1 1 1 1 1 1 Polygalaceae Polygala nuttallii 1 1 1 1 Rubiaceae Houstonia serpyllifolia 1 1 1 1 Scropulariaceae Chaenorhinum minus 1 1 1 1 1 1 1 1 1 Verbenaceae Verbena hastata 1 1 1 1 Total group 4 26 5 21 2 7 13 24 8 7 Species group 5: Species listed in other mountain Longleaf Pine sites or at Lavender Mountain in the past, but not found in our site currently, and not in Ware’s Flora of Floyd County. Apiaceae Sanicula trifoliata E.P. Bicknell 1 1 1 1 Aristolochiaceae Hexastylis shuttleworthii 1 1 1 1 (Britten & Baker f.) Small Asteraceae Helianthus laevigatus Torr. 1 1 1 1 1 & A. Gray Fabaceae Baptisia tinctoria (L.) R. Br. 1 1 1 1 Fabaceae Chamaecrista fasciculata 1 1 1 1 1 (Michx.) Greene Fabaceae Senna marilandica (L.) Link 1 1 1 1 1 1 Fabaceae Stylosanthes calcicola Small 1 1 1 Fabaceae Tephrosia mohrii (Rydb.) Godfrey 1 1 1 1 Total group 5 8 0 7 0 1 5 3 3 0 1Harper (1905), 2Andrews (1917), 3Jones (1940), 4Maceina et al. (2000), 5Carter and Londo (2006), 6 Womack and Carter (2011), 7Richard Ware, Georgia Botanical Society, Rome, GA, unpubl. data.