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. Allgemeine Forst-und Holzwirtschaftliche
Zeitang 59:4–5. [In German.]
Brockway, D.G., K.W. Outcalt, D.J. Tomczak, and E.E. Johnson. 2005. Restoring Longleaf
Pine forest ecosystems in the southern US. Pp. 501–519, In J.A. Stanturf and P.
Madsen (Eds.). Restoration of Boreal and Temperate Forests. CRC Press, Boca Raton,
FL. 600 pp.
Carter, R., and A.J. Londo. 2006. Remnant fire-disturbed Montane Longleaf Pine forest
in west-central Georgia. Pp. 475–477, In K.F. Connor (Ed.). Proceedings of the 13th
Biennial Southern Silvicultural Research Conference. General Technical Report SRS
92. US Department of Agriculture, Forest Service, Southern Research Station, Asheville,
NC. 640 pp.
Cipollini, M.L. 2005. The Berry College Longleaf management plan. Available online at
http://facultyweb.berry.edu/mcipollini/management/The%20Berry%20College%20
Longleaf%20Management%20Plan.htm. Accessed 1 January 2010.
Cipollini, M., A. Gaskell, and C. Worrell. 2005. Use of herbicides and prescribed burning
to control hardwoods in the regeneration and restoration of Mountain Longleaf Pine
habitats. Longleaf Alliance Report 8:9–13.
Cipollini, M., C. Francia, C. Kujala, A. Lottes, N. Malloy, M. Mann, E. Swanson, and
N. Wigington. 2008. Effects of restoration prescribed burning on post-fire mortality
in relict Montane Longleaf Pine (Pinus palustris) in northwestern Georgia. Longleaf
Alliance Report 13:71.
Currie, K., J.M. Varner, J. Kush, and M. Cipollini. 2006. A survey of the herbaceous
vegetation found in the Berry Longleaf Pine Management Area. Longleaf Alliance
Report 9:35–39.
Dean, B.E., A. Mason, and J.L. Thomas. 1973. Wildflowers of Alabama and Adjoining
States. The University of Alabama Press, Tuscaloosa, AL. 230 pp.
2012 M.L. Cipollini, J. Culberson, C. Strippelhoff , T. Baldvins, and K. Miller 651
Duncan, W.H., and L.E. Foote. 1975. Wildflowers of the Southeastern United States. The
University of Georgia Press, Athens, GA. 304 pp.
Harper, R.M. 1905. Some noteworthy stations for Pinus palustris. Torreya 5:55–60.
Horn, D., T. Cathcart, T. Hemmerly, and D. Duhl. 2005. Wildflowers of Tennessee, the Ohio
Valley, and the Southern Appalachians. Lone Pine Publishing, Auburn, WA. 496 pp.
Huber, A., A. Gaskell, C. Worrell, and M. Cipollini. 2006. Estimation of carbon storage
in a mountain Longleaf ecosystem in northwestern Georgia. Longleaf Alliance Report
9:44–48.
Jones, H.C. 1940. Plant ecology of the Berry Schools Property, Floyd County, Georgia.
Ph.D. Dissertation. George Peobody College, Nashville, TN. 111 pp.
Knobel, E. 1980. Field Guide to the Grasses, Sedges, and Rushes of the United States.
Edition 2 revised by M.E. Faust. Dover Publications, Inc., New York, NY. 96 pp.
Kush, J.S., R.S. Meldahl, and C. Avery. 2004. A restoration success: Longleaf Pine
seedlings established in a fire-suppressed, old-growth stand. Ecological Restoration
22:6–10.
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.