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The Vascular Flora and Phytogeographical Analysis of the Tennessee River Gorge, Hamilton and Marion Counties, Tennessee
Emily Blyveis and Joey Shaw

Southeastern Naturalist, Volume 11, Issue 4 (2012): 599–636

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2012 SOUTHEASTERN NATURALIST 11(4):599–636 The Vascular Flora and Phytogeographical Analysis of the Tennessee River Gorge, Hamilton and Marion Counties, Tennessee Emily Blyveis1,2,* and Joey Shaw1 Abstract - An inventory of the vascular flora of the Tennessee River Gorge (TRG) was conducted between April 2009 and July 2011. The TRG is a 41-km-long river canyon located on the Cumberland Plateau physiographic province within Hamilton and Marion counties, TN. According to prior ecological surveys, the 4970-ha study area contains 12 natural communities. This survey documented 123 families, 392 genera, and 700 species and sub-specific taxa, including 138 county range extensions. Approximately 13% of the flora (92 species) is considered non-native. Fifteen imperiled species were reported, including 3 that are undescribed and 9 not documented in any previous Cumberland Plateau flora. In addition to the floristic component of this work, a phytogeographical analysis was conducted for the TRG and 12 other Cumberland Plateau floras in Tennessee and Kentucky. An important finding of this research was the identification of a biogeographical transition zone, from a higher northern to a higher southern species richness. This biogeographic break was identified in the vicinity of Fentress, Scott, and Morgan counties, TN. Introduction The Tennessee River Gorge (TRG) is a large river canyon located at the southeastern boundary of the Cumberland Plateau physiographic province within Tennessee. It is in a region of secondary-growth eastern deciduous forest approximately 8 km west of Chattanooga. The gorge bisects 41 km of the Cumberland Plateau with escarpments reaching 305 m, thus earning the name “Tennessee’s Grand Canyon” (Luther 1979:13). The east and west borders of the TRG are immediately drawn by the Ridge and Valley physiographic province and the Sequatchie Valley, respectively. The northern half of the gorge is part of the Cumberland Plateau’s Walden Ridge geologic formation. Raccoon Mountain makes up the southern half of the gorge and is considered an extension of Walden Ridge. A survey of the northern half of the gorge was conducted by Beck and Van Horn (2007) in their flora of Prentice Cooper State Forest (Prentice Cooper or PC). The southern half of the TRG and the adjacent ridges of Little Cedar Mountain (LCM), however, have remained understudied areas of scientific and conservation interest. Currently, land ownership of the TRG is divided among multiple entities. The Tennessee Valley Authority manages several thousand hectares in the gorge for hydroelectric power generation, public recreation, and wildlife habitat. The 1Department of Biological and Environmental Sciences, University of Tennessee at Chattanooga, 615 McCallie Avenue, Chattanooga, TN 37403. 2Current address - 3621 Prospect Church Road, Apison, TN 37302. *Corresponding author - emilyblyveis@ 600 Southeastern Naturalist Vol. 11, No. 4 Tennessee River Gorge Trust (TRGT) is a 501(c)(3) non-profit land trust that promotes environmental stewardship through public education outreach programs and land acquisition (TRGT 2011). Currently, the TRGT protects more than 6800 ha of undisturbed natural lands in the gorge. The remaining parcels are owned by multiple private holders. The land is subject to multi-purpose usage; however, the rugged topography has provided an obstacle for significant development as well as a platform for grassroots conservation. In 1983, the Tennessee Nature Conservancy contracted with the Tennessee Natural Heritage Program to conduct an ecological survey of the gorge. This study resulted in the delineation of the natural communities, the documentation of many rare species, and the identification of large tracts of ecologically significant habitat (Bridges et al. 1984). The Cumberland Plateau forests have been best characterized by Hinkle (1978) as oak, oak-hickory, and oak-pine community types with mixed mesophytic elements limited to ravines and escarpment slopes. Using data gathered by Bridges et al. (1984), Carroll (n.d.) produced a detailed map delineating the forest communities of the TRG. Their results exhibit Hinkle’s (1978) description of the forests in this region. Floristic studies of the Cumberland Plateau include Clark (1966), Wofford et al. (1979), Sole et al. (1983), Schmalzer et al. (1985), Clements and Wofford (1991), Allawos (1994), Goodson (2000), Bailey and Coe (2001), Weckman et al. (2003), Fleming and Wofford (2004), McEwan et al. (2005), Beck and Van Horn (2007), and Huskins and Shaw (2010). These floras consist of 3 from Kentucky and 10 from Tennessee, and their findings are summarized in Table 1. Prentice Cooper is the largest study area located in Hamilton, Marion, and Sequatchie counties, TN. It covers 10,300 ha and contains 1072 documented species (Beck and Van Horn 2007). The smallest study site is Big Everidge Hollow in Letcher County, KY, which contains 52 ha and 263 reported species (McEwan et al. 2005). These studies reveal the plateau’s high species richness as compared with other physiographic provinces in the state (TENN 2010, Wofford and Chester 2002). This high species richness is a natural consequence of topographic variation, geology, slope, aspect, and moisture differences on the plateau, which in turn affords a diversity of plant habitats (Braun 1950, Fenneman 1938, Hinkle 1978). While botanical inventories remain an important tool for conservation, they also increase the resolution of individual species’ ranges and may clarify geographical affinities associated with floristic compositions (Allawos 1994, Bailey and Coe 2001, Clements 1989, Murrell 1985, Murrell and Wofford 1987). Phytogeographical distribution analyses can improve our understanding of the origins of historical plant migrations, relict populations, and forest community disjunctions (FNA 1993, Graham 1972, Murrell 1985). Several botanists have conducted some level of a distribution analysis on the flora of east Tennessee. Murrell and Wofford (1987) analyzed the phytogeographical significance of Big Frog Mountain in the Blue Ridge physiographic province of Polk County. On the Cumberland Plateau, Clements (1989) analyzed the geographic distributions of the Wolf Cove flora in Franklin County. Allawos (1994) examined the distributions of the North White Oak Creek Gorge flora located in Fentress and Scott counties. In the same area, Bailey and Coe (2001) evaluated the geographic 2012 E. Blyveis and J. Shaw 601 affinities of the Clear Fork River and New River riparian flora. Shaw and Wofford (2003) investigated the phytogeography of Big South Fork National River and Recreation Area’s woody flora in Scott County, TN, and McCreary County, KY. Each of these investigations revealed broad, central floristic distributions with significant northern components. While these were all sound phytogeographical studies that revealed similar distributional patterns, comparisons between them are dubious for the reasons of incompatible taxonomic nomenclature, differing range interpretations derived by the authors, and differing phytogeographical references employed. No such distribution analyses have been pursued in the recent floristic surveys on the southern plateau’s Walden Ridge section (Beck and Van Horn 2007, Huskins and Shaw 2010), of which the current study i s a part. The objectives of this investigation are to (1) inventory the vascular flora of the TRG and the adjacent LCM, (2) document rare and non-native taxa, (3) determine county records for Hamilton and Marion counties, (4) compare the floristic richness and similarity of the TRG with other Cumberland Plateau floras, and (5) elucidate the phytogeographical affinities of the TRG and Cumberland Plateau flora. Study Area The Tennessee River Gorge covers 10,927 ha of land from its northern escarpment on Walden Ridge to its southern escarpment on Raccoon Mountain. The delineated project area contains 4970 ha, with elevations ranging from 193 m to 583 m. It is located between 35°00'25.200" (35.007°) and 35°07'12.000" (35.12°) N latitudes and 85°21'10.800" (-85.353°) and 85°35'16.800"(-85.588°) W longitudes (Fig. 1). The study area consists of two distinct units: Unit One includes the southern half of the gorge, on Raccoon Mountain, from the river margins to the escarpments and covers 4310 ha. A small eastern portion of it is located in Figure 1. Map of the Tennessee River Gorge study area in Hamilton and Marion counties, TN. 602 Southeastern Naturalist Vol. 11, No. 4 Hamilton County, but the majority extends westward into Marion County. Unit Two is defined as the middle and southern ridges of Little Cedar Mountain (LCM) at the western terminus of the gorge and is surrounded to the east and south by Nickajack Lake. The western margin of Unit Two parallels Anderson Ridge and the Sequatchie Valley. Located entirely within Marion County, LCM contains 660 ha of karst limestone woodlands, bluffs, and barrens. The USGS Gap Analysis Program characterizes the TRG vegetation as predominantly southern dry calcareous forest and south-central interior mesophytic forest (USGS-NBII 2011). General habitat types include riparian areas, Tennessee River floodplains, mesophytic gorge slopes, ravines, Cumberland Plateau uplands, and ruderal sites. Within these broad habitat classifications, several smaller unique habitats are supported, including limestone sinkholes, a hardwood marsh, sandstone and limestone rock outcrops and bluffs, rich mesic coves, and limestone glades. Geography The study area is adjacent to the southern extremity of Walden Ridge, a southeastern sub-region of the Cumberland Plateau physiographic province in Tennessee. The Cumberland Plateau is part of the larger Appalachian Plateau, which extends in a northeastern to southwestern orientation from southern New York to central Alabama (Fenneman 1938, Hinkle 1978, Luther 1977). Fenneman (1938) divided the Appalachian Plateau into seven sections, and the southernmost of these is the Cumberland Plateau, which reaches its northern limits in northeastern Kentucky and its southern extent in central Alabama, spanning a distance of approximately 467 km (Hinkle 1978). Within Tennessee, the plateau covers 6920 km2 (Luther 1977). The Sequatchie Valley bisects the plateau through the southern half of the state, creating a large, straight, and narrow northeast– southwest trending valley. This division by the Sequatchie Valley separates the western Cumberland Plateau proper from the slender, eastern sub-region of Walden Ridge. The Cumberland Escarpment lines the eastern border of Walden Ridge and faces the western perimeter of the Ridge and Valley physiographic province. Raccoon, Lookout, and Sand Mountains are located immediately south of Walden Ridge, and they represent extensions of the plateau spreading in to northern Alabama and Georgia. Geology and soils Geology of the TRG is characterized by rocks of Ordovician to Pennsylvanian age (Hack 1966). On the plateau uplands, the Sewanee Conglomerate Formation is the youngest geologic layer (Avel and Hartman 1979). It contains hard, erosion-resistant conglomerate and sandstone. At a slightly lower elevation, formations from the Gizzard Group comprise sandstone, siltstone, shale, and coal. The escarpments are formed from Pennington Formation shale. Pennington outcrops are unstable and have resulted in 3 major landslides in the TRG in recent times (Carroll n.d.). The lower slopes are characterized by a wide band of Mississippian age Bangor limestone from the Hartselle Formation. Bangor has a high dissolvability, thus producing many solution cavities and sinkholes. The lowest elevations at the base of the slopes are the Tennessee River riparian zones, consisting of flood plains and alluvial fan deposits. Geology of LCM is primarily 2012 E. Blyveis and J. Shaw 603 characterized by Mississippian Bangor limestone (Avel and Hartman 1979, Wilson 1979). LCM exists at a lower elevation than the adjacent gorge escarpments, and the terrain consists of karst limestone rocklands with ledges and outcrops occupying 51% of the surface area (USDA, NRCS 2011b). Abundant solution cavities on LCM provide an exceptional example of limestone dis solution. Elevation differences in the study area offer great contrasts in the geologic parent materials as well as a diversity of soil types (Jackson 1982). Soils on the plateau uplands overlay sandstone bedrock and are typically strongly acidic, very well drained, and low in fertility. The upper slopes contain a moderately deep profile over limestone and shale bedrock. The two major soil types are classified in the Lily-Ramsey Association (USDA, NRCS 2011b). A majority of the middle and base slopes are delineated in the Bouldin-Gilpen Group, characterized by bouldery colluviums of sandstone, limestone, and shale. Silt loam deposits of the Hamblen-Staser Association are sparsely observed along the riparian areas (Jackson 1982). Soils of the Talbott Series cover approximately 49% of the surface area on LCM (USDA, NRCS 2011b). The remaining 51% is categorized as a rockland of Bangor limestone outcrops. The Talbott Series consists of moderately deep soils weathered from limestone. They are well drained, gently sloping to moderately steep, with a silty clay texture. Due to the unique soil situations and geology, LCM supports a considerably different flora from the rest of the gorge, exhibiting a local area of high endemism (Estes and Beck 201 1). Climate The climate of the TRG is characterized by warm, humid summers and cool, mild winters and a growing season ranging from 180 to 220 days (NOAA 2001). Climate data were gathered by the National Oceanic and Atmospheric Administration (NOAA) between the years of 1971 and 2000 from the lower elevation Lovell Field Weather Station in Hamilton County and the higher elevation Monteagle Weather Station in Marion County. The Lovell Field Station is located at 35.03°N latitude and -85.20°W longitude, approximately 27 km east of the study area at an elevation of 204 m. The Monteagle Station is located at 35.22°N latitude and -85.84°W longitude, approximately 14 km northeast of the study area at an elevation of 563 m (SRCC 2011). The lower gorge experiences an average annual mean temperature of 15.5 °C (SRCC 2011); the highest mean monthly temperature is 26.4 °C during July, and the lowest is 4.1 °C in January. The average annual mean temperature on the plateau uplands is 13.3 °C; the highest mean monthly temperature is 23.7 °C in July, and the lowest is 1.5 °C in January. In terms of rainfall, the gorge lowlands receive 138.3 cm annually; the maximum monthly amount of precipitation occurs in March (15.7 cm), and the lowest amount occurs in October (8.3 cm). The plateau uplands receive considerably more precipitation annually than the lowlands. Normal annual precipitation on the uplands is 161.9 cm; the highest monthly amount falls in March (17.6 cm), and the lowest volume falls in August (10.3 cm). The most significant differences are that the upper gorge, on average, is 2.2 °C cooler, and receives approximately 24 cm more precipitation annually than the lower gorge. 604 Southeastern Naturalist Vol. 11, No. 4 Methods Specimen collection Sixty-one total collecting trips were made from March to November of 2009 to 2011. A habitat classification map of the TRG (Carroll n.d.) was consulted to determine locations of the different habitat types. An effort was made to visit each representative habitat in the study area biweekly throughout the growing season. In areas of major collection, GPS coordinates were recorded using a Garmin eTrex Vista Cx (Garmin International, Inc., Olathe, KS), accurate to 5 m. Accession numbers were assigned to each specimen along with locality data and supplementary field notes. Species identifications were made using multiple taxonomic treatments, including Radford et al. (1968), Cronquist (1980), Wofford (1989), Gleason and Cronquist (1991), Wofford and Chester (2002), selected volumes from the Flora of North America series (FNA 2002 and 2003), Jones (2005), and Weakley (in prep.). Species determinations were cross examined with voucher specimens from UCHT and TENN as well as images from the online databases of the University of Tennessee Herbarium at Knoxville and the United States Department of Agriculture Natural Resources Conservation Services PLANTS Database (USDA, NRCS 2011a). County records were determined by county-level distribution maps from TENN (2010). The nomenclature and native status of all taxa reported follows the USDA PLANTS Database (USDA, NRCS 2011a) and A Fifth Checklist of Tennessee Vascular Plants (Chester et al. 2009). The Tennessee Natural Heritage Program Rare Plant List (TDEC 2008) was consulted for rare species designations. Voucher specimens were processed according to standard herbarium protocols, and the collection was deposited at UCHT. Duplicates and triplicates were sent to APSC and TENN. Statistical analysis Quantitative comparisons were generated between the flora of the TRG and 12 other studies on the Cumberland Plateau using Sørenson’s coefficient of community (CC). This calculation of floristic similarity is given by the formula CC = 2c / (s1 + s2), where c is the number of species common to both communities, s1 represents the number of species known from the TRG community, and s2 is the number of species in the community being compared (Smith and Smith 2002). The CC index can fall between zero and one: a value closer to zero indicates dissimilarity between the compared communities, and a value closer to one signifies a greater similarity. The species numbers for all 12 floristic studies used in this calculation were standardized for synonymy by Huskins (2008) in a normalized comparative plant list, which will be further discussed below. Phytogeographical analysis Methodology for the phytogeographical distribution analysis was adapted from Murrell (1987) and Allawos (1994) and was developed in Microsoft Office Excel 2007 (Microsoft Corporation, Redmond, WA) spreadsheets. The MS Excel 2003 version of Huskins’ (2008) “Normalized Comparative Plant List” was provided as an integral resource for this study. The document contains a comparative checklist 2012 E. Blyveis and J. Shaw 605 of presence/absence species data from the 12 Cumberland Plateau floras presented in Table 1 with the exception of the current study and the New River Gorge flora on the Allegheny Plateau. Huskins normalized the checklist by converting each study’s nomenclature to a standardized type used by the USDA PLANTS Database. This was an important step toward making possible a broad and comparative phytogeographical analysis. The current study further developed the Normalized Comparative Plant List to include presence/absence species data for the TRG, as well as a phytogeographical analysis of all species from the 13 Cumberland Plateau floras. One flora (New River Gorge or NRG; Suiter and Evans 1999) located on the Allegheny Plateau in southern West Virginia was added to extend the analysis farther north on this contiguous plateau province and to add a northern flora comparable in size to the Tennessee Cumberland Plateau studies. Distribution maps from the USDA PLANTS Database (USDA, NRCS 2011a) were consulted for accurate species range descriptions. A column was added to the Normalized Comparative Plant List (Huskins 2008) to describe the geographic “center of distribution” of every native species in the list of all 14 floras (Table 1). Five possible centers of distribution were recognized: central, northern, eastern, southern, and western. Each taxon was assigned to one of the five categories based on the approximate center of its distribution. These headings were followed by a more detailed range description. Species with widespread geographic distributions in the eastern US were classified as central species. Taxa with broad ranges to the north that bottlenecked southward were characterized as northern. Similarly, species with broad southern distributions that tapered northward were considered southern taxa. Eastern taxa were those with distributions along the east coast ranging as far west as the Appalachian Mountains. Species with western distributions extended from the west coast to the Midwestern region. The second component of this two-fold analysis was to categorize the “geographical affinities” of the TRG flora into 4 distinct groups: intraneous, extraneous, strict endemic, or introduced. This study defines taxa with intraneous distributions as those for which the TRG study area is located well within the margin of their range, with a buffer width of no less than 5 counties of average county size for the eastern US. The extraneous group consists of taxa for which the study area is situated on the periphery of their range with a buffer width of no more than 5 average-sized counties. The strict endemic species have ranges restricted to the southern Cumberland Plateau sensu Fenneman (1938), and the introduced group includes all non-native species. Results and Discussion Floristic summary In total, 960 specimens were collected from the study area over 3 growing seasons. Additionally, 124 TRG herbarium vouchers from other collectors were examined. From these combined collections, 700 species and sub-specific taxa were identified, representing 392 genera from 123 families of vascular plants (Table 2). The largest documented family was Asteraceae with 94 species representing 13.4% of the flora. This family was followed by Poaceae with 51 species (7.3%), Cyperaceae with 48 (6.9%), Fabaceae with 35 (5%), Rosaceae with 27 606 Southeastern Naturalist Vol. 11, No. 4 Table 1. Floras of the Allegheny and Cumberland Plateaus in West Virginia, Kentucky, and Tennessee, as well as area, results of Sørenson’s coefficient of community (CC), phytogeographical summary, and relevant species data. Percentages have been rounded to the nearest whole number where appropriate, and they refer to the total species number of the flora listed at the far left column. All species numbers were normalized for synonymy by Huskins and Shaw (2010) in concurrence with USDA PLANTS Database. Study sites: NRG = New River Gorge (Suiter and Evans 1999), PK = Pilot Knob (Weckman et al. 2003), LCW = Lilley Cornett Woods (Sole et al. 1983), BEH = Big Everidge Hollow (McEwan et al. 2005),NWOCG = North White Oak Creek Gorge (Allawos 1994), CF/NR = Clear Fork/New River (Goodson and Bailey 2001), OB = Obed (Schmalzer et al. 1985), FCF = Fall Creek Falls (Fleming and Wofford 2004), SG = Savage Gulf (Wofford et al. 1979), NCCG = North Chickamauga Creek Gorge State Natural Area (Huskins and Shaw 2010), FG = Fiery Gizzard (Clark 1966), PC = Prentice Cooper State Forest (Beck and Van Horn 2007), WC = Wolf Cove (Clements and Wofford 1991), TRG = Tennessee River Gorge (this study). Species in common Area with Study site (ha) CC value TRG TRG> TRG< Central Northern Eastern Southern Western Introduced Totals NRG 25,123 N/A N/A N/A N/A 666 (74%) 96 (11%) 1 (0.1%) 17 (2%) 2 (0.2%) 118 (13%) 900 PK 262 0.5308 319 381 184 419 (83%) 18 (4%) 0 9 (2%) 0 55 (11%) 502 LCW 220 0.4905 298 402 217 403 (78%) 30 (6%) 0 17 (3%) 2 (0.2%) 61 (12%) 515 BEH 52 0.3780 182 518 81 237 (90%) 15 (6%) 0 9 (3%) 0 1 (0.4%) 263 NWOCG 5407 0.5106 312 388 210 414 (79%) 35 (7%) 3 (0.6%) 27 (5%) 1 (0.2%) 41 (8%) 522 CF/NR 1896 0.5183 333 367 252 465 (80%) 32 (6%) 2 (0.3%) 38 (7%) 1 (0.2%) 44 (8%) 585 OB 4000 0.5516 393 307 332 581 (80%) 35 (5%) 1 (0.1%) 46 (6%) 0 60 (8%) 725 FCF 8900 0.6257 494 206 385 670 (76%) 44 (5%) 2 (0.2%) 59 (7%) 0 103 (12%) 879 SG 4047 0.5615 386 314 289 545 (81%) 39 (6%) 0 46 (7%) 1 (0.1%) 42 (6%) 675 NCCG 2862 0.6043 394 306 210 448 (74%) 29 (5%) 1 (0.2%) 49 (8%) 0 74 (12%) 604 FG 3626 0.5644 366 334 231 480 (80%) 37 (6%) 0 40 (7%) 2 (0.3%) 37 (6%) 597 PC 10,300 0.6682 592 108 481 774 (72%) 33 (3%) 1 (0.09%) 87 (8%) 0 176 (16%) 1072 WC 1000 0.5385 343 357 231 468 (82%) 28 (5%) 0 47 (8%) 0 28 (5%) 574 TRG 4970 N/A N/A N/A N/A 514 (73%) 24 (3%) 0 70 (10%) 0 92 (13%) 700 2012 E. Blyveis and J. Shaw 607 (3.9%), and Lamiaceae with 26 (3.7%). The largest genera were Carex with 31 species, followed by Quercus (14), Viola (12), and Symphyotrichum (10). In all, 138 county range extensions were documented, which implies that 19.7% of the TRG flora was not known to occur in the respective counties from which they were collected. There were 15 county records for Hamilton County and 123 for Marion County. The percentages of each taxonomic division in the flora are shown in Table 2, and these proportions are similar to that of other regional floras. Rare species The TRG contains fifteen imperiled taxa, comprising 2.1% of the total vascular flora. Twelve of these species are state or federally listed, and 3 are undescribed (Table 3). Found only in the calcareous habitats of LCM, the undescribed taxa Table 2. Floristic summary of the TRG. Values in parentheses represent the proportions of Liliopsida and Magnoliopsida to the division Magnoliophyta. % of total Division Families Genera Species species composition Equisetophyta 1 1 1 0.10% Pteridophyta 9 18 29 4.10% Coniferophyta 3 4 7 1.00% Magnoliophyta 110 369 663 94.70% (Liliopsida) (15) (72) (150) (21.40%) (Magnoliopsida) (95) (297) (513) (73.30%) Total 123 392 700 100% Table 3. Rare species of the TRG study area with state statuses as well as state and global ranks where applicable. A double dagger denotes a taxon that has not been documented in any previous Cumberland Plateau flora. Scutellaria montana is also federally listed as threatened. Rank Scientific name Common name State status State Global Arabis patens Sull.‡ Spreading Rockcress Endangered S1 G3 Castanea dentata (Marsh.) American Chestnut Special concern S2S3 G4 Borkh. Clematis sp. nov.‡ Clematis species Undescribed taxon Cotinus obovatus Raf.‡ American Smoketree Special concern S2 G4 Helianthus sp. nov.‡ Sunflower species Undescribed taxon Hydrastis canadensis L. Goldenseal Commercially exploited S3 G4 Liatris cylindracea Michx.‡ Ontario Blazing Star Threatened S2 G5 Lonicera dioica L.‡ Limber Honeysuckle Special concern S2 G5 Onosmodium bejariense DC. Soft-hair Marbleseed Endangered S1 G4 ex A. DC. var. hispidissimum (Mack.) B.L. Turn.‡ Panax quinquefolius L. American Ginseng Commercially exploited S3S4 G3G4 Phemeranthus mengesii (W. Menges' Flameflower Threatened S2 G3 Wolf) Kiger Polymnia johnbeckii D. Estes‡ John Beck’s Leafcup Proposed endangered S1 G1 Scutellaria montana Chapm. Large-flowered Skullcap Threatened S2 G3 Sisyrinchium sp. nov.‡ Blue-eyed grass species Undescribed taxon Viola tripartita Ell. Threepart Violet Special concern S2S3 G5T3 608 Southeastern Naturalist Vol. 11, No. 4 are considered rare species as they presumably have low population numbers and narrow ranges as a result of strict habitat requirements. The Natural Heritage Program of the Tennessee Department of Environment and Conservation assigns a non-legal state rank and status to rare taxa based on the number of occurrences throughout the state. These S-ranks range from S1 to S4. A rank of S1 indicates critical endangerment and vulnerability to extirpation from Tennessee, S2 is assigned to taxa with 6 to 20 occurrences in the state, S3 signifies species with 21 to 100 occurrences in Tennessee, and S4 designates taxa that are abundant within the state yet rare in parts of their range (TDEC 2008). Three taxa in the TRG hold an S1 classification, 5 are S2 ranked, 1 has an S3 rank, 3 have split classifications, and 3 are not yet ranked on the Tennessee Natural Heritage Program Rare Plant List due to their undescribed status. Interestingly, 9 of the imperiled species reported in this study were not documented in any other flora on the Cumberland Plateau, and they all occur on LCM. Introduced species In total, 92 non-native species were documented, accounting for 13.1% of the flora. The families that contributed the greatest numbers of introduced taxa included Fabaceae (13 species), Poaceae (12), and Asteraceae (6). The TRG contained 33 species reported on the Tennessee Exotic Pest Plant Council (TNEPPC 2011) list. There were 13 rank-one species (known to be invasive), 14 rank-two species (spread easily in native plant communities), and 6 rank-three species (possess invasive characteristics, but not considered to spread as easily into native plant communities). The rank-one taxa that are most abundant in the TRG included: Ligustrum sinense Lour. (Chinese Privet), Lonicera japonica Thunb. (Japanese Honeysuckle), Microstegium vimineum (Trin.) A. Camus (Nepalese Browntop), and Pueraria montana (Lour.) Merr. var. lobata (Willd.) Maesen and S. Almeida (Kudzu). Floristic comparison Species richness of the TRG was compared to 8 other regional floras using a species-area curve generated specifically for the Cumberland Plateau within Tennessee by Huskins and Shaw (2010). Using this curve, a richness measure of approximately 760 taxa was predicted for a study area the size of the TRG. In comparison to the 3 other floras that were below the Huskins and Shaw (2010) regression line (NCCG, FG, and SG), the TRG was within a range of expected richness values. Edge effects and land-use history are factors that could account for a lower than expected species richness. With respect to the TRG and Prentice Cooper as a contiguous flora, the Tennessee River Gorge region represents a remarkably species rich geologic feature of the Cumberland Plateau in Tennessee, supporting a total of 1180 species and lesser taxa. The Sørenson’s coefficient of community was used to generate the similarity indices that are reported in Table 1. Prentice Cooper was the site shown to be most floristically similar to the TRG, with a CC value of 0.66. Fall Creek Falls was the next most similar with a value of 0.62, and then North Chickamauga Creek Gorge with a value of 0.60. Floras indicated as the most dissimilar to the 2012 E. Blyveis and J. Shaw 609 TRG included Big Everidge Hollow (0.37), Lilley Cornett Woods (0.49), and North White Oak Creek Gorge (0.51). Big Everidge and Lilley Cornett were areas much smaller than the remaining 10 study sites, and the North White Oak flora was shown to be species poor. Such factors may explain the low CC value computed for these sites. The TRG is most floristically similar to and shares the greatest number of species (592) in common with Prentice Cooper (Table 1). This finding was anticipated since the two sites are only separated by the Tennessee River. Interestingly, PC had the greatest number of species (481) that were not likewise found within the TRG study area. While this discrepancy could be interpreted as dissimilarity, it is more likely attributed to Prentice Cooper’s exceptionally large study area (10,300 ha), which contained greater habitat diversity of 18 general habitat types and almost twice the species richness of the TRG. Sørenson’s coefficient of community does not account for the correlation between area and richness. As a result, this calculation is capable of producing a low CC value for two sites that are similar in floristic composition but greatly different in area. Fall Creek Falls and North Chickamauga Creek Gorge had the second and third highest species numbers in common with the TRG (494 and 394, respectively). These relatively high values may be explained by the presence of a gorge system and predominance of gorge vegetation communities throughout both sites. Phytogeographical analysis The geographic range of every native taxon in 13 Cumberland Plateau floras and 1 Allegheny Plateau flora was examined in an attempt to better understand the phytogeography of the Cumberland Plateau. The results are summarized in Table 1. Taxa with broad central distributions, ranging between Canada and Florida, dominate the Cumberland Plateau, accounting for 90% (Big Everidge Hollow) to 72% (Prentice Cooper) of the species composition. Southern species made up the second largest composition of the native flora on the plateau, with 9 of the 14 study sites having a larger southern than northern component. The TRG flora had the highest percentage of southern taxa (10%), and Pilot Knob had the lowest (2%). Conversely, the New River Gorge (NRG) contained the greatest percentage of northern species (11%), while Prentice Cooper had the lowest (3%). The southern and northern geographic affinities of all 14 floras are illustrated in Figure 2. Study sites are arranged on the graph from left to right as northernmost location on the plateau to southernmost, respectively. As might be expected, the northernmost areas contain more northern than southern taxa, and the southernmost areas support more southern than northern taxa. The important finding of this research was the identification of a biogeographical transition zone, from a higher northern to a higher southern species richness. This biogeographic break was identified between the North White Oak Creek Gorge (NWOCG) in Fentress and Scott counties, TN, and the Clear Fork and New River in Fentress, Morgan, and Scott counties, TN at approximately 36.29°N latitude and -84.71°W longitude. It will be interesting to do similar analyses with floras of other physiographic provinces to see if this break is a function of latit ude. 610 Southeastern Naturalist Vol. 11, No. 4 The findings of the phytogeographical analysis for the Cumberland Plateau show a moderate, but evident, increasing southern affinity from the Clear Fork/New River site to the TRG (Fig. 2). From NWOCG in northeast Tennessee to the NRG in southern West Virginia, a clear northern affinity was observed. When interpreting the phytogeographical significance of the Cumberland Plateau, Estill and Cruzan (2001) suggest that one must consider both biogeographical history and contemporary ecology. As early as Braun’s (1950) studies of the Mixed Mesophytic Forest Region, it was observed that the vegetation of the Cumberland Plateau is coextensive with that of the unglaciated Appalachian Plateau. Several authors have suggested that the Blue Ridge and Cumberland Plateau physiographic provinces provided a pathway for southward migration of northern species during Pleistocene glaciations (Cain 1930, Davis 1983, Delcourt 1980, Watts 1970). Many northern taxa have since retreated to the Allegheny Plateau and farther north due to climatic warming during the Quaternary Period; however, northern elements persist on the Cumberland Plateau, particularly in coves, ravines, and gorges (Braun 1950, Caplenor 1979, Quarterman et al. 1972). To what extent northern affinities diminish and southern elements increase on the plateau had not been clarified. After 45 years of floristic research, however, the phytogeographical patterns of the Cumberland Plateau are becoming better understood. In addition to historical biogeography, land-use history and contemporary ecology also have influenced the affinities of the plateau flora. The southern Cumberland Plateau in Tennessee has experienced continual timber harvesting and hardwood-to-pine conversion during the past 60 years (McGrath et al. 2004). Agricultural land uses and urban sprawl have reduced the once characteristic mature hardwood forests of the plateau. These land-altering practices have been more widespread on the southern Cumberland Plateau, presumably because of better accessibility when compared to areas of higher topographic relief to the north. In addition to the effects of human disturbance, the narrowing southern Figure 2. Geographical affinities of 14 Cumberland and Allegheny Plateau floras. 2012 E. Blyveis and J. Shaw 611 plateau is bounded by physiographic transitions with the Ridge and Valley and the Interior Low Plateaus, bringing about changes in edaphic and climatic conditions (Braun 1950, Cathey 1990, Griffith et al. 1997, Omernik 1987). According to Cathey (1990), a transverse gradient of plant hardiness zones exists in northerncentral Tennessee. The area north of Blount, Loudon, and Cumberland counties experiences average annual minimum temperatures of -26 to -18 °C, while the area immediately south reaches minimum temperatures of -18 to -15 °C. This climatic gradient overlaps with the phytogeographical transition zone observed in the vicinity of the NWOCG and Clear Fork/New River floras (Fig. 2). To further elucidate distributional characteristics of the TRG flora, each taxon was assigned to 1 of 4 major categories: intraneous, extraneous, strict endemic, or introduced. All but the introduced taxa were further examined to characterize their centers of distribution. The results of this dual geographic distribution analysis for the TRG are reported in Table 4. A majority of the flora (514 taxa, 73.4%) consists of species with central distributions, with the TRG study area being intraneous to their ranges. Southern intraneous taxa made up 6.6%, and southern extraneous species constituted a larger element than northern intraneous species (2.6% and 2.1%, respectively). Only 9 taxa (1.3%) with northern extraneous distributions to the TRG were documented, and 6 taxa (0.8%) were strict endemics to the southern Cumberland Plateau in southeast Tennessee, northwest Georgia, and northeast Alabama. The TRG supports a larger composition of southern species, with Piedmont and Atlantic-Gulf Coastal Plains distributions, than northern species typical of the Mixed Mesophytic and Northern Hardwood Forest formations. With the study Table 4. Geographic distribution characteristics of the TRG flora. Distributional categories are adapted from Murrell (1987) and Allawos (1994). # of Category Sample taxon Geographic range taxa % Northern 24 3.4% Intraneous Carex pensylvanica Lam. Manitoba, Quebec, and Maine, 15 2.1% (Pennsylvania Sedge) s. to n. Ga., n. Miss., and Ark., w. to Iowa and N.D. Extraneous Houstonia canadensis Willd. N.D., Mich., N.Y., and Maine, 9 1.3% Ex Roem. & Schult. s. to Va., Tenn., and n. Ga., (Canadian Summer Bluet) w. to Missouri Central 514 73.4% Intraneous Quercus alba L. Minn., Ontario, Quebec, and 514 73.4% (White Oak) Maine, s. to Fla., Miss., and La., w. to e. Tex. and Neb. Southern 70 10.0% Intraneous Euphorbia mercurialina Michx. S. Ky. and Va., e. to N.C. and 46 6.6% (Mercury Spurge) n. Ga., s. to Fla. and n. Miss. Extraneous Collinsonia tuberosa Michx. S. Tenn. and N.C., s. to Ga., 18 2.6% (Deepwoods Horsebalm) Miss., and s. Louisiana Endemic Scutellaria montana S.e. Tenn. and n.w. Ga. 6 0.8% Introduced Ailanthus altissima (Mill) Swingle China and Taiwan 92 13.1% (Tree of Heaven) 612 Southeastern Naturalist Vol. 11, No. 4 area demarcated by the Ridge and Valley and the Sequatchie Valley, it is possible that these warmer, drier lowlands influence the affinities of the flora by providing conduits for native southeastern Coastal Plain taxa (e.g., Collinsonia tuberosa Michx. [Deepwoods Horsebalm], Cotinus obovatus Raf. [American Smoketree], and Ilex longipes Chapm. ex Trel. [Georgia Holly]). Of course, one must also consider the suitability of geologic substrate as a factor influencing plant species’ dispersal abilities. The southern affinity is further shown by the greater presence of extraneous southern taxa than extraneous northern taxa. These extraneous species are significant not only for the purpose of elucidating geographical affiliations but also for the broader purpose of monitoring species ranges with respect to climate change. If current climate trends continue, a recession of northern taxa would be anticipated as well as an encroachment of southern taxa through the southern plateau. In its present state, it is suggested that the southern Cumberland Plateau, from northern-central Tennessee and southward, maintains a more typically southern flora influenced by biogeographical history, recent land-use history, and contemporary ecological and environmental factors. Conclusions The TRG represents a floristically diverse and distinct feature of the Cumberland Plateau. Despite previous studies of the southern Cumberland Plateau in Tennessee, the need for this survey was reflected by the documentation of 138 county range extensions for Hamilton and Marion counties. Even with the exhaustive floristic inventory of Prentice Cooper on the northern half of the gorge, 108 species were documented in the TRG that were not found in Prentice Cooper. The TRG contains 15 imperiled taxa, including 9 that were not documented in any other floristic study on the plateau and 3 that are undescribed and potentially endemic to the karst woodlands of the southern Cumberland Plateau’s Little Cedar Mountain and adjacent Sequatchie Valley. While the plateau has historically been regarded as a physiographic refuge for characteristically northern taxa, it is apparent from this cohesive phytogeographical analysis that the central and southern Cumberland Plateau in Tennessee maintains a more typically southern floristic element. The TRG comprises the most distinct southern affinity due in part to its southernmost position on the plateau in Tennessee and its adjacency to the Ridge and Valley and Sequatchie Valley provinces, which are probable distribution pathways for native southeastern taxa of the Atlantic–Gulf Coastal Plains. One major concern within the TRG is the expansion of introduced species. The relatively high number (13.1%) of non-native taxa reflects the magnitude of human disturbance in the TRG. Efforts to eradicate the more aggressive species should be a conservation priority for the local community and for groups like the Tennessee River Gorge Trust who work to preserve the natural condition of the gorge. Unfortunately, issues of natural resource management are complicated by the fact that the Tennessee River Gorge is not administered by one agency but rather falls into the hands of many stakeholders with different land-use practices. While the Tennessee River Gorge Trust and Tennessee Vally Authority currently protect much of the land in the gorge, continued acquisition by the two organizations will be paramount in the preservation of this distinct natural feature. 2012 E. Blyveis and J. Shaw 613 Little Cedar Mountain represents a hotspot of endemism and is the most botanically unique portion of the study area. The 9 rare taxa not documented in any other Cumberland Plateau flora exist on these small mountain ridges. In addition, several other interesting taxa were present on LCM but absent throughout the rest of the study site (e.g., Cheilanthes alabamensis [Buckley] Kunze [Alabama Lipfern], Ophioglossum engelmannii Prantl [Limestone Adderstongue], and Opuntia humifusa [Raf.] Raf. [Prickley Pear Cactus]). The Tennessee Valley Authority administers multiple large tracts on LCM, including the entire southernmost ridge, which has been designated as a habitat protection area/small wild area. From the findings of this study, southern LCM is worthy of its conservation status. Based on the combined results from the flora of Prentice Cooper on the northern half of the gorge and the current study, the Tennessee River Gorge region represents a very rich area with plant species found nowhere else on the Cumberland Plateau. The gorge is a valuable natural resource to the local community, and it is a cultural and natural landmark of the Cumberland Plateau. Acknowledgments The authors wish to thank the two anonymous reviewers for their helpful suggestions to improve the manuscript. Thanks to Mr. Jim Brown of the Tennessee River Gorge Trust, Dr. Patricia Cox of the Tennessee Valley Authority, and Mr. Andrew Carroll of UTC. Gratitude is also extended to Dr. Dwayne Estes of Austin Peay State University and to several field assistants who helped with the collection of speci mens for this project. Literature Cited Allawos, J.G. 1994. The vascular flora of North White Oak Creek Gorge, Scott and Fentress counties, Tennessee. M.Sc. Thesis. University of Tennessee, Knoxville, TN. Avel, A.P., and G.S. Hartman. 1979. Engineering Geology of Hamilton County, Tennessee. Pp. 65–82, In R.J. Floyd, R.C. Milici, and R.L. Wilson (Eds.). Geology of Hamilton County, Tennessee. 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Weckman, and N.R. George. 2003. Checklist of the vascular flora of Pilot Knob State Nature Preserve, Powell County, Kentucky. Journal of the Kentucky Academy of Science 64(1):36–54. Wilson, R.A. 1979. The stratigraphy of exposed rocks in Hamilton County, Tennessee. Pp. 15–37, In R.C. Milici (Ed.). Geology of Hamilton County, Tennessee. Tennessee Division of Geology Bulletin Volume 79. Wofford, B.E. 1989. Guide to the Vascular Flora of the Blue Ridge. University of Georgia Press, Athens, GA. Wofford, B.E., and E.W. Chester. 2002. Trees, Shrubs, and Woody Vines of Tennessee. University of Tennessee Press, Knoxville, TN. Wofford, B.E., T.S. Patrick, L.R. Phillippe, and D.H. Webb. 1979. The vascular flora of Savage Gulf, Tennessee. Sida 8(2):135–151. 2012 E. Blyveis and J. Shaw 617 Appendix 1. Annotated checklist of the flora of the Tennessee River Gorge. The list is organized by division, subdivision, family, genus, and species. The taxonomic nomenclature of these groups follows the USDA PLANTS Database (USDA, NRCS 2011a) and A Fifth Checklist of Tennessee Vascular Plants (Chester et al. 2009). A single asterisk (*) precedes an introduced taxon, two asterisks (**) precede a rare species, and a double dagger (‡) indicates a rare taxon collected from the TRG yet absent from the 12 other Cumberland Plateau studies. Relative abundance, habitat association, and accession number follows each species, and for taxa not collected in this study, the original collector and institution where specimen was deposited are provided. Key to habitat association abbreviations DA = Disturbed areas LCM = Little Cedar Mountain LGS = Lower gorge slopes LMS = Limestone sinkholes MSS = Mesophytic slopes PL = Power line PS = Plateau surface RA = Riverine area SW = Swales UGS = Upper gorge slopes WL = Wetlands Key to relative abundance abbreviations (following Murrell and Wofford 1987) C = Common: characteristic and dominant F = Frequent: generally encountered O = Occasional: well distributed, but not anywhere abundant I = Infrequent: scattered locations throughout S = Scarce: several locations, or scattered small populations R = Rare: one or two locations in small populations VR = Very rare:a single locale, few individuals DIVISION EQUISETOPHYTA EQUISETACEAE Equisetum hyemale L. var. affine (Engelm.) A.A. Eaton; O. RA, SW. 483. DIVISION PTERIDOPHYTA ASPLENIACEAE Asplenium montanum Willd.; O. LGS, PS, UGS. 152. Asplenium platyneuron (L.) Britton, Stearns & Poggenb. var. platyneuron; C. LGS. 49. Asplenium resiliens Kunze; S. LCM. 164. Asplenium rhizophyllum L.; I. UGS. 307. BLECHNACEAE Woodwardia areolata (L.) T. Moore; R. SW. 628. DENNSTAEDTIACEAE Pteridium aquilinum (L.) Kuhn; R. DA. 528. DRYOPTERIDACEAE Athyrium filix-femina (L.) Roth; C. LGS. 557. Athyrium filix-femina (L.) Roth. ssp. asplenioides (Michx.) Hultén; C. PL. 556. Cystopteris fragilis (L.) Bernh. var. tennesseensis (Shaver) McGregor; VR. UGS. 667. 618 Southeastern Naturalist Vol. 11, No. 4 Cystopteris protrusa (Weath.) Blasdell; S. LGS. 175. Diplazium pycnocarpon (Spreng.) Broun; VR. LMS. 80. Dryopteris marginalis (L.) A. Gray; C. LGS, UGS. 52. Onoclea sensibilis L.; R. WL. 485. Polystichum acrostichoides (Michx.) Schott; C. LGS, UGS. 76. Woodsia obtusa (Spreng.) Torr. O. LCM, LGS, UGS. 700. OPHIOGLOSSACEAE Botrychium dissectum Spreng.; S. LGS, MSS. 636. Botrychium virginianum (L.) Sw.; O. LCM, LGS, MSS, SW. 47. Ophioglossum engelmannii Prantl; I. LCM. 623. Ophioglossum vulgatum L.; S. PS, SW. Beck-TENN. OSMUNDACEAE Osmunda cinnamomea L.; S. SW, UGS. 657. Osmunda regalis L. var. spectabilis (Willd.) A. Gray; I. DA, PS. 548. POLYPODIACEAE Pleopeltis polypodioides (L.) Andrews & Windham ssp. michauxiana (Weath.) Andrews & Windham; F. LCM, PS, UGS. 77. PTERIDACEAE Adiantum capillus-veneris L.; R. LCM. 575. Adiantum pedatum L.; O. LCM, LGS, MSS, SW. 50. Cheilanthes alabamensis (Buckley) Kunze; S. LCM. 298. Cheilanthes lanosa (Michx.) D.C. Eaton; I. LCM. 468. Pellaea atropurpurea (L.) Link; I. LCM. 150. THELYPTERIDACEAE Phegopteris hexagonoptera (Michx.) Fée; O. LGS, MSS, UGS. 228. Thelypteris noveboracensis (L.) Nieuwl.; I. PS, SW. 640. DIVISION CONIFEROPHYTA CUPRESSACEAE Juniperus virginiana L.; C. DA, LCM, PS. 144. PINACEAE Pinus echinata Mill.; C. LGS, PS, UGS. 174. Pinus strobus L.; C. DA, PL, PS. 569. Pinus taeda L.; C. DA, PL, PS, UGS. 256. Pinus virginiana Mill.; C. DA, LGS, PL, PS, UGS. 122. Tsuga canadensis (L.) Carriére; I. PS, MSS. 568. TAXODIACEAE Taxodium distichum (L.) Rich.; R. PS, SW. 726. DIVISION MAGNOLIOPHYTA CLASS LILIOPSIDA 2012 E. Blyveis and J. Shaw 619 AGAVACEAE Manfreda virginica (L.) Salisb. ex Rose; I. DA, UGS. 487. Yucca filamentosa L.; O. DA, LCM, RA. 497. ALISMATACEAE Sagittaria australis (J.G. Sm.) Small; I. RA, WL. 692. Sagittaria latifolia Willd.; I. RA, WL. 510. ARACEAE Arisaema dracontium (L.) Schott; R. SW. 698. Arisaema triphyllum (L.) Schott; C. LCM, LGS, MSS, PS, RA, SW, WL. 79. Arisaema triphyllum (L.) Schott ssp. quinatum (Buckley) Huttleston; S. LGS, SW. 624. COMMELINACEAE *Commelina communis L.; F. DA, LGS, LMS, RA. 124. Commelina diffusa Burm. f.; S. RA. Sharp et al.-TENN. *Murdannia keisak (Hassk.) Hand.-Maz.; S. LCM, WL. 516. Tradescantia subaspera Ker Gawl.; F. DA, LGS. 69. CYPERACEAE Carex albicans Willd. ex Spreng. var. albicans; F. LGS, MSS. 588. Carex albicans Willd. ex Spreng. var. emmonsii (Dewey ex Torr.) J. Rettig; C. LGS. 309. Carex amphibola Steud.; C. LCM, LGS, PS, UGS. 620. Carex atlantica L.H. Bailey ssp. atlantica; F. LCM, LGS. 634. Carex austrocaroliniana L.H. Bailey; O. LGS, MSS, UGS. 600. Carex blanda Dewey; C. LCM, LGS, MSS, PS, SW, UGS. Beck & Estes-TENN. Carex caroliniana Schwein.; O. LGS, RA. 218. Carex cherokeensis Schwein.; O. LCM, RA, WL. 216. Carex complanata Torr. & Hook.; F. DA, LGS, PS. Bridges & Somers-UCHT. Carex crinita Lam. var. brevicrinis Fernald; I. LCM, RA, WL. 276. Carex cumberlandensis Naczi, Kral & Bryson; O. MSS. Beck-TENN. Carex digitalis Willd. var. digitalis; C. LCM, LGS, MSS, PS, SW, UGS. Beck-TENN. Carex eburnea Boott; S. LCM, UGS. 585. Carex festucacea Schkuhr ex Willd.; F. LCM, LGS. 217. Carex frankii Kunth; O. RA, WL. 210. Carex granularis Muhl. ex Willd.; F. DA, LCM, LGS, PS. Bridges & Somers-UCHT. Carex grayi Carey; O. LCM, RA, WL. 207. Carex hirsutella Mack.; F. LGS, PS, UGS. Beck & Estes-TENN. Carex jamesii Scwein.; O. LGS. Bridges & Somers-UCHT. Carex lupulina Muhl. ex Willd.; I. LCM, WL. 384. Carex lurida Wahlenb.; I. LCM, RA, WL. 385. Carex meadii Dewey; VR. LCM. Estes-TENN. Carex muehlenbergii Schkuhr ex Willd. var. enervis Boott.; S. LCM. Bridges & Somers-UCHT. Carex nigromarginata Schwein.; C. DA, LCM, LGS, MSS, PS, SW, UGS. 283. Carex pedunculata Muhl. ex Willd.; I. PS, UGS. 282. Carex pensylvanica Lam.; I. LGS, MSS, UGS. 745. Carex platyphylla Carey; I. LGS. Bridges & Somers-UCHT. Carex purpurifera Mack.; F. LGS, MSS. Beck-TENN. Carex rosea Schkuhr ex Willd.; F. LGS, MSS, PS, UGS. Bridges & Somers-UCHT. Carex virescens Muhl. ex Willd.; O. MSS. Beck-TENN. Carex vulpinoidea Michx.; O. LCM, WL. 733. *Cyperus difformis L.; I. RA, WL. 165. 620 Southeastern Naturalist Vol. 11, No. 4 Cyperus echinatus (L.) Alph. Wood; I. RA, WL. Beck-TENN. Cyperus pseudovegetus Steud.; O. RA, WL. Beck-TENN. Cyperus strigosus L.; I. RA, WL. 564. Eleocharis microcarpa Torr.; I. DA, WL. Beck-TENN. Eleocharis obtusa (Willd.) Schult.; S. WL. Bridges & Somers-UCHT. Fimbristylis autumnalis (L.) Roem. & Schult.; I. WL. Bridges & Somers-UCHT. Kyllinga gracillima Miq.; S. LCM, WL. 703. Rhynchospora capitellata (Michx.) Vahl; O. RA, WL. Bridges & Somers-UCHT. Rhynchospora corniculata (Lam.) A. Gray; O. DA, RA, WL. Beck-TENN. Schoenoplectus pungens (Vahl) Palla; I. RA, WL. 554. Schoenoplectus tabernaemontani (C.C. Gmel.) Palla; O. RA, WL. 565. Scirpus atrovirens Willd.; I. RA, WL. Beck-TENN. Scirpus cyperinus (L.) Kunth; O. RA, WL. 197. Scleria ciliata Michx.; R. LGS. Bridges & Somers-UCHT. Scleria oligantha Michx.; S. LCM. 686. Scleria triglomerata Michx.; S. LGS. Bridges & Somers-UCHT. DIOSCOREACEAE *Dioscorea oppositifolia L.; C. DA, LCM, LGS, PS, RA, SW, UGS, WL. 181. Dioscorea villosa L.; C. DA, LCM, LGS, MSS, PS, RA, SW, UGS. 328. IRIDACEAE Iris cristata Aiton; I. LGS, RA. 321. Sisyrinchium angustifolium Mill.; O. LCM, LGS, RA, SW. 348. **‡Sisyrinchium sp. nov.; VR. LCM. Estes-TENN. JUNCACEAE Juncus acuminatus Michx.; O. RA, WL. Beck-TENN. Juncus coriaceus Mack.; F. RA, WL. 317. Juncus effusus L.; F. RA, WL. 209. Juncus marginatus Rostk.; I. RA, WL. Price & Murphy-TENN. Juncus tenuis Willd.; O. LGS, RA, SW, WL. 199. Luzula echinata (Small) F. J. Herm.; O. PS, UGS. 323. Luzula multiflora (Ehrh.) Lej.; C. LGS, MSS, PS, SW, UGS. 157. LEMNACEAE Spirodela polyrrhiza (L.) Schleid.; S. RA. 470. LILIACEAE Aletris farinosa L.; S. DA, PL. 665. Allium canadense L.; I. RA, WL. 644. *Allium vineale L.; O. LCM, LGS, UGS. 350. Chamaelirium luteum (L.) A. Gray; R. PS. 626. Erythronium americanum Ker Gawl.; O. LGS, MSS, SW. 268. *Hemerocallis fulva (L.) L.; S. DA. 351. Hypoxis hirsuta (L.) Coville; O. LCM, WL. 370. Maianthemum racemosum (L.) Link ssp. racemosum; C. LGS, MSS, PS, SW, UGS. 322. Medeola virginiana L.; S. LGS, MSS. Bridges & Somers-UCHT. *Narcissus pseudonarcissus L.; R. DA, RA. 253. Nothoscordum bivalve (L.) Britton; I. LCM, MSS. 582. Polygonatum biflorum (Walter) Elliot; C. LGS, LMS, MSS, PS, SW, UGS. 325. 2012 E. Blyveis and J. Shaw 621 Prosartes lanuginosa (Michx.) D. Don; S. LMS. 313. Trillium cuneatum Raf.; C. LCM, LGS, MSS, SW. 1. Uvularia grandiflora Sm.; S. MSS. 314. Uvularia perfoliata L.; C. LCM, LGS, MSS, UGS. 595. Veratrum parviflorum Michx.; R. MSS. 670. ORCHIDACEAE Corallorhiza wisteriana Conrad; I. LCM, MSS. Beck-TENN. Goodyera pubescens (Willd.) R. Br.; C. LCM, LGS, MSS, PS, SW, UGS. 90. Tipularia discolor (Pursh) Nutt.; F. LCM, LGS, MSS, PS, SW, UGS. 115. POACEAE Agrostis perennans (Walter) Tuck.; S. DA, PS. Bridges & Somers-UCHT. Andropogon gyrans Ashe var. gyrans; I. DA, PS. Bridges & Somers-UCHT. Andropogon virginicus L.; O. DA, LGS, PL. 215. Aristida dichotoma Michx. var. dichotoma; F. DA, LCM. Bridges & Somers-UCHT. Aristida oligantha Michx.; O. DA, PS. Bridges & Somers-UCHT. *Arthraxon hispidus (Thunb.) Makino; I. DA, LGS. 235. Arundinaria gigantea (Walter) Muhl. ssp. gigantea; F. LCM, LGS, RA, UGS. 241. Brachyelytrum erectum (Schreb. ex Spreng.) P. Beauv.; F. LCM, LGS. 211. Bromus pubescens Muhl. ex Willd.; S. LGS. Bridges & Somers-UCHT. *Bromus racemosus L.; O. DA, LGS. 273. Chasmanthium latifolium (Michx.) Yates; C. LCM, PS, RA, WL. 149. Chasmanthium sessiliflorum (Poir.) Yates; F. LCM, RA, WL. 170. Cinna arundinacea L.; I. WL. Bridges & Somers-UCHT. Danthonia spicata (L.) P. Beauv. ex Roem. & Schult.; O. LGS. Beck & Estes-TENN. Dichanthelium boscii (Poir.) Gould & C.A. Clark; C. LCM, LGS, MSS, PS, UGS. 155. Dichanthelium clandestinum (L.) Gould; F. SW. Beck-TENN. Dichanthelium commutatum (Schult.) Gould; C. LCM, LGS, PS. Bridges & Somers-UCHT. Dichanthelium dichotomum (L.) Gould var. dichotomum; C. LGS. 204. Dichanthelium laxiflorum (Lam.) Gould; F. DA, LGS, PS, UGS. Bridges & Somers-UCHT. Dichanthelium scoparium (Lam.) Gould; O. DA. Beck-TENN. Dichanthelium sphaerocarpon (Elliot) Gould var. sphaerocarpon; O. LGS, PS, UGS. Bridges & Somers-UCHT. *Digitaria ischaemum (Schreb.) Schreb. ex Muhl.; I. DA, LGS. 196. Digitaria sanguinalis (L.) Scop.; O. DA, RA. 187. *Echinochloa crus-galli (L.) P. Beauv.; R. DA, PS. Bridges & Somers-UCHT. *Eleusine indica (L.) Gaertn.; S. DA, LGS, RA. 202. Elymus virginicus L.; I. RA, WL. 206. Eragrostis spectabilis (Pursh) Steud.; O. LCM, PS. Beck-TENN. Leersia oryzoides (L.) Sw.; S. LGS. Bridges & Somers-UCHT. Leersia virginica Willd.; S. DA. Bridges & Somers-UCHT. *Lolium perenne L. ssp. multiflorum (Lam.) Husnot; O. DA, LGS. 223. Melica mutica Walter; O. LCM, LGS. 158. *Microstegium vimineum (Trin.) A. Camus; C. DA, LGS, PS, RA, WL. 233. Panicum capillare L.; F. PS, RA, SW, UGS. 551. Panicum dichotomiflorum Michx. var. dichotomoflorum; C. PS, SW. 552. Panicum flexile (Gattinger) Scribn.; F. DA, PS, SW, UGS. 553. Panicum rigidulum Bosc ex Nees var. pubescens (Vasey) Lelong; I. LGS. Bridges & Somers- UCHT. Panicum virgatum L.; O. DA, LGS, UGS. 214. 622 Southeastern Naturalist Vol. 11, No. 4 *Paspalum dilatatum Poir.; F. DA, LGS, PS, RA. 198. Paspalum laeve Michx.; F. DA, PL, PS. 224. Piptochaetium avenaceum (L.) Parodi; F. LCM, PS, UGS. 638. Poa cuspidata Nutt.; C. DA, LGS, PS, UGS. 278. Poa sylvestris A. Gray; C. DA, LCM, LGS, PS, UGS. 272. Saccharum alopecuroides (L.) Nutt.; O. PS, SW. 75. *Setaria faberi Herrm.; S. DA. Bridges & Somers-UCHT. Setaria parviflora (Poir.) Kerguélen; O. DA, RA. 200. *Setaria pumila (Poir.) Roem. & Schult. ssp. pumila; O. DA, RA. 201. *Sorghum halepense (L.) Pers.; C. DA, LCM, LGS, PS, UGS. 195. Sphenopholis intermedia (Rydb.) Rydb.; O. DA, RA. 274. Sphenopholis nitida (Biehler) Scribn.; F. LGS, PS. 156. Tridens flavus (L.) Hitchc. var. flavus; C. DA, PS, RA. 244. Zizaniopsis miliacea (Michx.) Döll & Asch.; S. RA. Bridges & Somers-UCHT. POTAMOGETONACEAE Potamogeton diversifolius Raf.; S. RA, WL. Beck-TENN. Potamogeton foliosus Raf.; R. LCM, WL. 383. Potamogeton nodosus Poir.; R. RA. 404. SMILACACEAE Smilax bona-nox L.; F. LGS, PS, UGS. 102. Smilax glauca Walter; C. DA, LCM, LGS, PL, PS, RA, UGS. 58. Smilax rotundifolia L.; C. DA, LCM, LGS, PL, PS, SW, UGS. 59. TYPHACEAE Typha angustifolia L.; O. LCM, RA, WL. 682. DIVISION MAGNOLIOPHYTA CLASS MAGNOLIOPSIDA ACANTHACEAE Justicia americana (L.) Vahl; I. DA, LCM, WL. 381. Ruellia caroliniensis (J.F. Gmel.) Steud.; C. DA, LCM, LGS, PS. 54. ACERACEAE Acer negundo L.; C. DA, LCM, LGS, PS. 39. Acer rubrum L.; C. DA, LCM, LGS, PL, PS, RA, SW, UGS, WL. 60. Acer saccharinum L.; O. RA. 398. Acer saccharum Marsh. var. saccharum; C. DA, LCM, LGS, MSS, PL, PS, RA, SW, UGS. 57. AMARANTHACEAE *Alternanthera philoxeroides (Mart.) Griseb.; R. RA, WL. 679. ANACARDIACEAE **‡Cotinus obovatus Raf.; R. LCM. 306. Rhus aromatica Aiton; F. DA, LCM, LGS, PL, PS, RA. 151. Rhus copallinum L.; C. DA, PL, PS, RA, UGS. 452. Rhus glabra L.; F. DA, PL, PS, UGS. 67. Toxicodendron radicans (L.) Kuntze; C. DA, LCM, LGS, MSS, PL, PS, RA, SW , UGS, WL. 132. 2012 E. Blyveis and J. Shaw 623 ANNONACEAE Asimina triloba (L.) Dunal; C. LCM, LGS, MSS, PS, SW, UGS. 147. APIACEAE Chaerophyllum tainturieri Hook.; F. LGS, LMS, SW. 14. Cicuta maculata L.; I. DA, RA, WL. 93. Cryptotaenia canadensis (L.) DC.; F. LGS, MSS, SW. Beck & Estes-TENN. *Daucus carota L.; C. DA, PL, RA, UGS. 65. Eryngium yuccifolium Michx.; S. PS, SW. 727. Hydrocotyle verticillata Thunb.; I. RA, WL. 92. Osmorhiza claytonii (Michx.) C.B. Clarke; O. LGS, SW. 333. Osmorhiza longistylis (Torr.) DC.; F. LGS, MSS, SW, UGS. 19. Ptilimnium capillaceum (Michx.) Raf.; O. LCM, WL. 371. Sanicula canadensis L.; C. LCM, LGS, MSS, RA, SW, UGS. 382. Sanicula smallii E.P. Bicknell; O. DA, LGS. 354. Thaspium trifoliatum (L.) A. Gray var. aureum (L.) Britton; F. DA, LCM, LGS, RA. 632. *Torilis arvensis (Huds.) Link; F. RA. 424. Zizia aptera (A. Gray) Fernald; F. LGS, PL, RA. Beck-TENN. APOCYNACEAE Amsonia tabernaemontana Walter var. tabernaemontana; S. LCM. 584. *Vinca major L.; R. DA, LGS. 602. *Vinca minor L.; I. DA. 288. AQUIFOLIACEAE Ilex decidua Walter; O. LGS. 462. Ilex longipes Chap. ex Trel.; I. LCM. 749. Ilex opaca Aiton; O. LGS. 142. Ilex verticillata (L.) A. Gray; I. LCM, WL. 378. ARALIACEAE **Panax quinquefolius L.; R. LGS, MSS. 622. ARISTOLOCHIACEAE Aristolochia serpentaria L.; O. LGS, PS, UGS. 649. Aristolochia tomentosa Sims; I. LGS, PS, UGS. 650. Hexastylis arifolia (Michx.) Small var. ruthii (Ashe) Blomquist; C. LGS, MSS, PS, SW, UGS. 9. Hexastylis shuttleworthii (Britten & Baker f.) Small; S. PS. 601. ASCLEPIADACEAE Asclepias quadrifolia Jacq.; O. DA, PS, UGS. 342. Asclepias tuberosa L.; F. DA, LGS, PS, RA. 226. Asclepias variegata L.; F. LGS, LMS, MSS, UGS. 46. Asclepias verticillata L.; VR. LCM. 707. Asclepias viridiflora Raf.; I. LCM. 708. Cynanchum laeve (Michx.) Pers.; I. DA, RA. 472. Matelea carolinensis (Jacq.) Woodson; R. LCM. Bridges & Somers-UCHT. Matelea gonocarpos (Walter) Shinners; S. LCM. 375. ASTERACEAE Achillea millefolium L.; F. PL, PS. 365. 624 Southeastern Naturalist Vol. 11, No. 4 Ageratina altissima (L.) King & H. Rob. var. altissima; C. DA, LGS, UGS. Bridges & Somers- UCHT. Ageratina aromatica (L.) Spach. var. aromatic; O. LGS, MSS. Bridges & Somers-UCHT. Ambrosia artemisiifolia L.; C. DA, LGS, PL, PS, RA, UGS. 524. Antennaria plantaginifolia (L.) Richardson; C. DA, PL, PS, UGS. 279. Antennaria solitaria Rydb.; F. PL, PS, UGS. 280. Arnoglossum atriplicifolium (L.) H. Rob.; O. LCM, WL. 421. Bidens aristosa (Michx.) Britt.; O. PL, PS. 545. Bidens frondosa L.; F. DA, LGS. Pyne-UCHT. Chrysopsis mariana (L.) Elliot; S. RA. 529. *Cichorium intybus L.; R. DA, LCM. 705. Cirsium discolor (Muhl. ex Willd.) Spreng.; O. RA. 526. *Cirsium vulgare (Savi) Ten.; C. DA, PL, PS, RA. 474. Conoclinium coelestinum (L.) DC.; C. DA, LCM, LGS, PS, RA. 512. Conyza canadensis (L.) Cronquist var. canadensis; F. DA, RA. 527. Coreopsis major Walter; C. DA, LGS, PL, PS, RA, UGS. 362. Coreopsis tripteris L.; I. RA, WL. Bridges & Somers-UCHT. Eclipta prostrata (L.) L.; S. RA, WL. Bridges & Somers-UCHT. Elephantopus carolinianus Raeusch.; C. LCM, LGS, PS, UGS. 515. Elephantopus tomentosus L.; C. LCM, LGS, UGS. 460. Erechtites hieraciifolia (L.) Raf. ex DC.; O. DA, LGS, RA. Bridges & Somers-UCHT . Erigeron annuus (L.) Pers.; F. DA, LGS, PL, PS, RA. 42. Erigeron philadelphicus L.; C. DA, LGS, PL, PS, UGS. 10. Erigeron strigosus Muhl. ex Willd. var. strigosus; C. DA, PS, RA. 126. Eupatorium album L. var. album; F. DA, LGS, PL, PS, UGS. 469. Eupatorium capillifolium (Lam.) Small; O. PL, PS, UGS. 543. Eupatorium hyssopifolium L. var. hyssopifolium; O. LGS, PL, PS, SW. 275. Eupatorium perfoliatum L.; I. PS, SW. 534. Eupatorium purpureum L.; C. DA, LGS, MSS, PL, PS, RA, UGS. 73. Eupatorium rotundifolium L. var. ovatum (Bigelow) Torr.; S. PL, PS. 230. Eupatorium rotundifolium L. var. rotundifolium; I. LGS, PS, SW, UGS. 716. Eupatorium serotinum Michx.; O. PS, SW. 538. Eupatorium sessilifolium L.; O. LGS, UGS. Bridges & Somers-UCHT. Eurybia divaricata (L.) G.L. Nesom; F. LCM, LGS, MSS. 520. Eutrochium purpureum (L.) E.E. Lamont var. purpureum; I. PS, SW, UGS. 724. Fleischmannia incarnata (Walter) King & H. Rob.; O. LCM. 194. *Galinsoga quadriradiata Cav.; O. DA, LGS. 696. Helenium amarum (Raf.) H. Rock; F. DA, PL, PS, RA. 125. Helenium flexuosum Raf.; R. PS, SW. 723. Helianthus divaricatus L.; C. DA, LGS. Bridges & Somers-UCHT. Helianthus hirsutus Raf.; C. DA, PS, SW, UGS. 119. Helianthus microcephalus Torr. & A. Gray; C. DA, LGS, PS, RA, UGS. 457. Helianthus tuberosus L.; F. DA, LGS, PL, PS, RA, UGS. 489. **‡Helianthus sp. nov.; VR. LCM. Estes-TENN. Heliopsis helianthoides (L.) Sweet; O. LGS, UGS. 72. Hieracium gronovii L.; C. DA, LGS, PS, UGS. 647. Krigia biflora (Walter) S.F. Blake; C. DA, PS, UGS. 637. Krigia virginica (L.) Willd.; F. DA, PL, PS. 663. Lactuca canadensis L.; O. DA, RA. 193. Lactuca floridana (L.) Gaertn.; O. DA, PL, PS, RA, WL. 499. *Lactuca saligna L.; I. DA, RA. 225. *Leucanthemum vulgare Lam.; S. DA, RA. 41. Liatris aspera Michx.; O. DA, PS, SW, UGS. 536. **‡Liatris cylindracea Michx.; R. LCM. Estes-TENN. Liatris squarrosa (L.) Michx.; R. LCM. 709. 2012 E. Blyveis and J. Shaw 625 Packera anonyma (Alph. Wood) W.A. Weber & A. Löve; I. LCM. Beck-TENN. Packera glabella (Poir.) C. Jeffrey; O. DA, MSS. 448. Packera obovata (Muhl. ex Willd.) W.A. Weber & A. Löve; I. LGS, MSS. 713. Packera paupercula (Michx.) A. Löve & D. Löve; O. PS, UGS. 7. Pityopsis graminifolia (Michx.) Nutt. var. graminifolia; S. DA, PL, PS. 560. Pityopsis graminifolia (Michx.) Nutt. var. latifolia (Fernald) Semple & F.D. Bowers; S. DA, PL, PS. 559. Polymnia canadensis L.; O. LCM, LGS. 300. **‡Polymnia johnbeckii D. Estes; R. LCM. 153. Prenanthes serpentaria Pursh; C. LGS, MSS, RA, UGS. 666. Pseudognaphalium obtusifolium (L.) Hillard & B.L. Burtt ssp. obtusifolium; F. DA, PL, PS. 544. Ratibida pinnata (Vent.) Barnhart; VR. LCM. 685. Rudbeckia fulgida Aiton var. fulgida; C. DA, LGS. Bridges & Somers-UCHT. Rudbeckia hirta L. var. pulcherrima Farw.; O. PL, PS. 363. Sericocarpus linifolius (L.) Britton, Sterns & Poggenb.; F. DA, LGS, PS, UGS. 521. Silphium trifoliatum L. var. latifolium A. Gray; I. LCM. Bridges & Somers-UCHT. Silphium trifoliatum L. var trifoliatum; F. LGS, PS. 409. Smallanthus uvedalius (L.) Mack. ex Small; F. LGS, MSS, UGS. 518. Solidago caesia L.; C. DA, LCM, LGS, PL, PS, RA, SW, UGS. 719. Solidago erecta Pursh; F. PS, RA. 243. Solidago flexicaulis L.; O. DA, LCM, LGS, PL, SW, UGS. 185. Solidago gigantea Aiton; F. DA, LGS, PL, PS. 133. Solidago rugosa Ait. var. aspera (Aiton) Cronquist; C. DA, PL, PS, RA. 138. Solidago sphacelata Raf.; C. DA, PL, PS, SW. 232. Solidago ulmifolia Muhl. Ex Willd.; C. LCM, LGS. 192. Symphyotrichum cordifolium (L.) G.L. Nesom; C. DA, LCM, LGS, PS, SW, UGS. 519. Symphyotrichum divaricatum (Nutt.) G.L. Nesom; C. DA, LGS, PL, PS, RA. 186. Symphyotrichum dumosum (L.) G.L. Nesom var. dumosum; F. DA, PL, PS. 220. Symphyotrichum laeve (L.) A. Löve & D. Löve var. leave; O. DA, LGS, PS, RA. 540. Symphyotrichum novae-angliae (L.) G.L. Nesom; F. DA, PS, SW. 531. Symphyotrichum ontarionis (Wiegand) G.L. Nesom; I. DA, LGS. Bridges & Somers-UCHT. Symphyotrichum patens (Aiton) G.L. Nesom var. patens; C. DA, PS, SW. 189. Symphyotrichum pilosum (Willd.) G.L. Nesom var. pilosum; C. DA, LCM, LGS, PL, PS. 221. Symphyotrichum shortii (Lindl.) G.L. Nesom; F. DA, LGS, PS. Bridges & Somers-UCHT. Symphyotrichum undulatum (L.) G.L. Nesom; I. PS, SW. 219. *Taraxacum officinale F.H. Wigg.; C. DA, PL, PS. 295. Verbesina occidentalis (L.) Walter; F. LGS, MSS, SW. 465. Verbesina virginica L.; I. RA, WL. 717. Vernonia flaccidifolia Small; O. DA, LGS, PL, PS. 117. Vernonia gigantea (Walter) Trel. ssp. gigantea; F. DA, LCM, LGS, PL, RA. 188. BALSAMINACEAE Impatiens capensis Meerb.; C. LGS, LMS, RA, SW, WL. 89. BERBERIDACEAE Caulophyllum thalictroides (L.) Michx.; VR. RA. Sharp et al.-UCHT. Jeffersonia diphylla (L.) Pers.; S. LCM, MSS. 576. *Nandina domestica Thunb.; S. LCM. 162. Podophyllum peltatum L.; C. LGS, MSS, SW. 292. BETULACEAE Alnus serrulata (Aiton) Willd.; O. RA. 357. 626 Southeastern Naturalist Vol. 11, No. 4 Carpinus caroliniana Walter; O. LGS, RA. 148. Ostrya virginiana (Mill.) K. Koch; O. LCM, LGS, WL. 379. BIGNONIACEAE Bignonia capreolata L.; C. DA, LCM, LGS, MSS, PS, RA, SW, UGS, WL. 5. Campsis radicans (L.) Seem. ex Bureau; C. DA, LCM, LGS, PL, PS, RA. 78. BORAGINACEAE *Buglossoides arvensis (L.) I.M. Johnston; C. DA, LGS, PS, RA. 4. Cynoglossum virginianum L.; F. LCM, LGS, MSS. 627. Lithospermum canescens (Michx.) Lehm.; I. LCM. 583. Lithospermum latifolium Michx.; R. LCM. 740. Mertensia virginica (L.) Pers. ex Link; VR. LCM, MSS. 605. Myosotis macrosperma Engelm.; O. LGS, MSS. 616. Onosmodium bejariense DC. ex A. DC. var. bejariense B.L. Turner; VR. LCM. Estes et al.-TENN. **‡Onosmodium bejariense DC. ex A. DC. var. hispidissimum (Mack.) B.L. Turner; VR. LCM. Beck-TENN. BRASSICACEAE Arabis laevigata (Muhl. ex Willd.) Poir. var. laevigata; S. LCM. 563. **‡Arabis patens Sull.; R. LCM. Estes-TENN. Cardamine angustata O.E. Schulz; I. LGS, SW. 310. Cardamine concatenata (Michx.) Sw.; O. LCM. 263. Cardamine dissecta (Leavenworth) Al-Shehbaz; O. LGS, MSS, SW. 249. *Cardamine hirsuta L.; I. MSS, SW, UGS. 254. *Draba verna L.; O. DA, RA. 271. Lepidium virginicum L.; I. RA, WL. 680. *Thlaspi alliaceum L.; R. DA, PS. 594. CACTACEAE Opuntia humifusa (Raf.) Raf.; I. LCM. 154. CALYCANTHACEAE Calycanthus floridus L. var. floridus; C. LGS, LMS, SW, UGS. 2. CAMPANULACEAE Campanula divaricata Michx.; O. PS, UGS. 143. Campanulastrum americanum (L.) Small; F. LCM, LGS, WL. 416. Lobelia cardinalis L.; C. DA, LCM, LGS, RA, WL. 130. Lobelia inflata L.; C. DA, LCM, LGS, PS, RA. 109. Lobelia puberula Michx.; F. LCM, PL. 237. Lobelia spicata Lam.; C. DA, LCM, WL. 423. Triodanis perfoliata (L.) Nieuwl.; F. DA, RA. 38. CAPRIFOLIACEAE **‡Lonicera dioica L.; VR. LCM. Pyne-UCHT. *Lonicera fragrantissima Lindl. & Paxton; I. DA, RA. 258. *Lonicera japonica Thunb.; C. DA, LCM, LGS, PL, PS, RA, WL. 17. *Lonicera maackii (Rupr.) Herder; C. DA, LCM, LGS, PL, PS, RA, WL. 182. Lonicera sempervirens L.; VR. LCM. 757. Sambucus nigra L. ssp. canadensis (L.) R. Bolli; C. DA, LGS, PL, UGS. 167. Sambucus racemosa L. var. racemosa; O. DA, UGS. 66. 2012 E. Blyveis and J. Shaw 627 Symphoricarpos orbiculatus Moench; I. LCM, WL. 389. Viburnum acerifolium L.; C. LCM, LGS, MSS, PS, SW, UGS. 70. Viburnum rufidulum Raf.; F. DA, PL, PS, UGS. 227. CARYOPHYLACEAE *Cerastium fontanum Baumg. ssp. vulgare (Hartm.) Greuter & Burdet; F. DA. 110. *Dianthus armeria L.; S. LCM. 690. Silene stellata (L.) W.T. Aiton; I. DA, LGS. 136. Silene virginica L.; I. LCM. 319. *Stellaria media (L.) Vill. ssp. media; F. DA, PL. 262. Stellaria pubera Michx.; F. DA, PS. 287. CELASTRACEAE *Celastrus orbiculatus Thunb.; S. DA. Beck-TENN. Celastrus scandens L.; O. DA, LGS, MSS. 431. Euonymus americanus L.; C. LCM, LGS, LMS, MSS, PS, SW, UGS, WL. 440. Euonymus atropurpureus Jacq.; S. LCM. Bridges & Somers-UCHT. *Euonymus fortunei (Turcz.) Hand.-Maz.; O. LGS, MSS. 612. CHENOPODIACEAE *Chenopodium ambrosioides L.; I. DA, RA. Bridges & Somers-UCHT. CISTACEAE Lechea racemulosa Michx.; S. DA, LCM. Beck-TENN. CLUSIACEAE Hypericum densiflorum Pursh; S. LCM. 710. Hypericum denticulatum Walter; O. LGS, UGS. 461. Hypericum frondosum Michx.; F. LCM, WL. 240. Hypericum gentianoides (L.) Britton, Sterns & Poggenb.; I. PL, PS. 651. Hypericum hypericoides (L.) Crantz ssp. hypericoides; C. LGS, MSS, SW, UGS. 111. Hypericum hypericoides (L.) Crantz ssp. multicaule (Michx. ex Willd.) Robson; C. DA, PL, PS, UGS. 550. Hypericum punctatum Lam.; F. DA, LCM, MSS, PS, RA, UGS. 121. CONVOLVULACEAE Ipomoea pandurata (L.) G. Mey.; O. DA, RA. 412. CORNACEAE Cornus amomum Mill.; F. LCM, RA, WL. 374. Cornus florida L.; C. DA, LCM, LGS, MSS, PL, PS, RA, SW, UGS. 180. Cornus foemina Mill.; I. RA. Beck-TENN. Nyssa sylvatica Marsh.; C. DA, LGS, MSS, PS, SW, UGS. 64. CRASSULACEAE Sedum pulchellum Michx.; O. DA, LCM. 24. Sedum ternatum Michx.; C. LCM, LGS, LMS. 88. CUSCUTACEAE Cuscuta gronovii Willd. ex Schult.; S. DA, LCM. 711. 628 Southeastern Naturalist Vol. 11, No. 4 EBENACEAE Diospyros virginiana L.; C. LCM, LGS, MSS, RA, UGS. 35. ELAEAGNACEAE *Elaeagnus pungens Thunb.; F. DA, LCM, RA. 229. *Elaeagnus umbellata Thunb.; O. DA, RA. 36. ERICACEAE Epigaea repens L.; O. MSS, PS, UGS. 284. Kalmia latifolia L.; O. PS, UGS. 281. Oxydendrum arboreum (L.) DC.; C. DA, LGS, MSS, PL, PS, RA, SW, UGS. 51. Rhododendron catawbiense Michx.; I. MSS, PS, UGS. 562. Rhododendron cumberlandense E. L. Braun; S. PS, UGS. 658. Vaccinium arboreum Marsh.; C. LGS, PS, UGS. 103. Vaccinium corymbosum L.; C. PS, UGS. 71. Vaccinium pallidum Aiton; O. PS, UGS. Beck-TENN. Vaccinium stamineum L.; F. PS, UGS. 702. EUPHORBIACEAE Acalypha gracilens A. Gray; O. DA, RA. 503. Chamaesyce nutans (Lag.) Small; C. DA, LGS, PL. 112. Croton monanthogynus Michx.; F. DA. 127. Euphorbia corollata L.; C. DA, LCM, LGS, PL, PS, UGS. 85. Euphorbia dentata Michx.; S. LGS. Beck-TENN. Euphorbia mercurialina Michx.; C. DA, LGS, PL, RA. 341. FABACEAE *Albizia julibrissin Durazz.; C. DA, PL, RA. 99. Amorpha fruticosa L.; F. DA, LCM, RA. 161. Amphicarpaea bracteata (L.) Fernald; C. LCM, LGS, MSS, RA. 128. Apios americana Medik.; C. LCM, LGS, RA. 473. Cercis canadensis L.; C. DA, LCM, LGS, MSS, PS, RA, SW, UGS. 62. Chamaecrista fasciculata (Michx.) Greene var. fasciculata; F. DA, RA. 99. Chamaecrista nictitans (L.) Moench ssp. nictitans var. nictitans; F. RA. 507. Cladrastis kentukea (Dum. Cours.) Rudd; VR. RA. Bridges & Somers-UCHT. Clitoria mariana L.; F. DA, PS, RA. 120. Desmanthus illinoensis (Michx.) MacMill. ex B.L. Rob. & Fernald; O. DA, LGS. 179. Desmodium nudiflorum (L.) DC.; C. LCM, LGS, LMS, PS, RA, UGS. 45. Desmodium nuttallii (Schindl.) B.G. Schub.; F. DA, LGS. 105. Desmodium pauciflorum (Nutt.) DC.; F. LCM, LGS, RA. 129. Desmodium rotundifolium DC.; C. DA, LCM, LGS, RA, UGS. 101. Gleditsia triacanthos L.; I. RA. 34. *Lathyrus latifolius L.; F. DA, RA. 100. *Lespedeza cuneata (Dum. Cours.) G. Don; C. DA, LCM, RA. 303. Lespedeza virginica (L.) Britton; O. LCM. Beck-TENN. *Medicago lupulina L.; F. DA, PL, PS. 12. *Melilotus officinalis (L.) Lam.; C. DA, PL. 123. Mimosa microphylla Dryand.; O. DA, PL. 492. Orbexilum pedunculatum (Mill.) Rydb. var. pedunculatum; I. DA, PS. 630. Phaseolus polystachyios (L.) Britton, Sterns & Poggenb.; O. DA. Beck-TENN. *Pueraria montana (Lour.) Merr. var. lobata (Willd.) Maesen & S. Almeida; C. DA, PL, RA. 413. 2012 E. Blyveis and J. Shaw 629 Rhynchosia tomentosa (L.) Hook. & Arn.; O. UGS. 466. Robinia pseudoacacia L.; O. DA, RA. 32. *Securigera varia (L.) Lassen; F. DA, RA. 37. Senna marilandica (L.) Link; S. DA, RA. 513. *Trifolium campestre Schreb.; S. DA. Beck & Estes-TENN. *Trifolium incarnatum L.; R. DA, LCM. 619. *Trifolium pratense L.; C. DA, LCM, PL, PS. 8. *Trifolium repens L.; C. DA, PL, PS. 29. Vicia caroliniana Walter; O. DA, LGS. 597. *Wisteria floribunda (Willd.) DC.; O. RA. 756. *Wisteria sinensis (Sims) DC.; F. DA, LGS. 84. FAGACEAE **Castanea dentata (Marsh.) Borkh.; VR. MSS. 78. Fagus grandifolia Ehrh.; C. LCM, LGS, MSS, PS, RA, SW, UGS. 11. Quercus alba L.; C. DA, LCM, LGS, LMS, MSS, PS, RA, SW, UGS. 231. Quercus coccinea Münchh.; C. DA, LGS, MSS, PS. 430. Quercus falcata Michx.; C. DA, LCM, LGS, PL, SW, UGS. 141. Quercus marilandica Münchh.; F. DA, LGS, PS. 674. Quercus michauxii Nutt.; O. DA, RA. 486. Quercus muehlenbergii Engelm.; I. LCM. Beck & Estes-TENN. Quercus nigra L.; O. DA, RA, WL. 505. Quercus pagoda Raf.; F. PS, UGS. 393. Quercus phellos L.; I. DA, LGS. 441. Quercus prinus L.; C. DA, LCM, LGS, MSS, PL, PS, SW, UGS. 140. Quercus rubra L.; C. LGS, PS, RA, UGS. 447. Quercus shumardii Buckley; I. LCM. Pyne & Bowden-TENN. Quercus stellata Wangenh.; O. DA, PS, UGS. 464. Quercus velutina Lam.; C. DA, LGS, PS, UGS. 429. FUMARIACEAE Corydalis flavula (Raf.) DC.; I. LCM. 577. GENTIANACEAE Frasera caroliniensis Walter; O. DA. Beck & Estes-TENN. Obolaria virginica L.; I. MSS, UGS. 570. Sabatia angularis (L.) Pursh; I. LGS. Beck-TENN. GERANIACEAE *Geranium dissectum L.; O. DA. 289. Geranium maculatum L.; C. LGS, MSS, RA. 293. *Geranium molle L.; F. DA. 324. GROSSULARIACEAE Itea virginica L.; F. DA, LGS, PS, RA, UGS. 356. HALORAGACEAE *Myriophyllum spicatum L.; F. DA, RA, WL. 482. HAMAMELIDACEAE Hamamelis virginiana L.; C. LGS, RA, SW. 330. Liquidamber styraciflua L.; C. DA, LCM, LGS, LMS, PS, RA, UGS. 23. 630 Southeastern Naturalist Vol. 11, No. 4 HIPPOCASTANACEAE Aesculus flava Aiton; O. LMS, MSS. 83. HYDRANGEACEAE Hydrangea cinerea Small; C. LGS, SW, UGS. 68. Philadelphus hirsutus Nutt.; F. LCM, MSS. 574. HYDROPHYLLACEAE Hydrophyllum macrophyllum Nutt.; F. LGS. 296. Nemophila aphylla (L.) Brummitt; O. LCM. 586. Phacelia bipinnatifida Michx.; C. DA, LCM, LGS, UGS. 336. Phacelia dubia (L.) Trel. var. dubia; I. DA, LCM. 608. Phacelia dubia (L.) Trel. var. interior Fernald; VR. LCM. Estes-TENN. JUGLANDACEAE Carya alba (L.) Nutt.; C. LCM, LGS, PS, SW, UGS. 439. Carya cordiformis (Wang.) K. Koch; C. LGS, RA. 547. Carya glabra (Mill.) Sweet; C. LGS, PS, SW, UGS. 426. Carya ovata (Mill.) K. Koch; C. LCM, LGS, PS, SW, UGS. 423. Carya pallida (Ashe) Engl. & Graebn.; F. LGS, UGS. Pyne & Bowen-TENN. Juglans nigra L.; C. RA. 405. LAMIACEAE Collinsonia canadensis L.; C. LGS, UGS. 184. Collinsonia tuberosa Michx.; C. LGS, PL, PS, UGS. 238. *Glechoma hederacea L.; C. DA, RA, WL. 643. *Lamium amplexicaule L.; C. DA, PS. 270. *Lamium purpureum L.; C. DA, PS. 257. Lycopus virginicus L.; F. LCM, LGS, UGS. 514. Monarda clinopodia L.; C. DA, LGS, UGS. 641. Monarda fistulosa L.; C. DA, PL, UGS. 364. *Mosla dianthera (Buch.-Ham. ex Roxb.) Maxim.; O. MSS, UGS. 748. *Perilla frutescens (L.) Britton; F. LGS, MSS, SW. Bridges & Somers-UCHT. Physostegia virginiana (L.) Benth. ssp. praemorsa (Shinners) Cantino; I. LCM. Beck-TENN. Prunella vulgaris L.; C. DA, LGS, PL. 82. Pycnanthemum muticum (Michx.) Pers.; O. PS. Beck-TENN. Pycnanthemum pycnanthemoides (Leavenworth) Fernald var. pycnanthemoides; F. PS, SW, UGS. 456. Pycnanthemum tenuifolium Schrad.; F. MSS, PS. 432. Salvia lyrata L.; C. DA, LCM, LGS, RA. 13. Salvia urticifolia L.; C. DA, LCM, LGS. Scutellaria elliptica Muhl. var. hirsuta (Short & Peter) Fernald; F. DA, LCM, LGS, UGS. 352. **Scutellaria montana Chapm.; I. LGS, UGS. 305. Scutellaria nervosa Pursh; R. LCM. Estes-TENN. Scutellaria ovata Hill; O. LCM, LGS. 684. Scutellaria pseudoserrata Epling; F. LGS, UGS. 653. Stachys cordata Riddell; C. DA, LGS, PS, UGS. 108. Stachys tenuifolia Willd.; F. LGS, RA. 97. Teucrium canadense L.; O. WL. Beck-TENN. Trichostema brachiatum L.; I. RA. 455. 2012 E. Blyveis and J. Shaw 631 LAURACEAE Lindera benzoin (L.) Blume; C. LCM, LGS, MSS, PS, SW, UGS. 106. Sassafras albidum (Nutt.) Nees; C. LCM, LGS, MSS, PS, RA, SW. 61. LINACEAE Linum striatum Walter; S. LCM. Beck-TENN. LOGANIACEAE Spigelia marilandica (L.) L.; F. LCM, LGS, WL. 304. LYTHRACEAE Rotala ramosoir (L.) Koehne; I. RA. Beck-TENN. MAGNOLIACEAE Liriodendron tulipifera L.; C. LGS, MSS, PS, RA, SW, UGS. 104. MALVACEAE Hibiscus moscheutos L.; O. LCM, WL. 415. Sida spinosa L.; I. DA, RA. 495. MENISPERMACEAE Cocculus carolinus (L.) DC.; F. DA, RA, WL. 500. Menispermum canadense L.; O. DA, LCM, WL. 376. MONOTROPACEAE Monotropa hypopithys L.; S. UGS. 395. Monotropa uniflora L.; S. SW, UGS. 177. MORACEAE Morus rubra L.; F. LGS, RA. 427. OLEACEAE *Forsythia viridissima Lindl.; I. DA, LGS. 252. Fraxinus americana L.; C. LGS, RA, UGS. 53. Fraxinus quadrangulata Michx.; S. LCM. 344. *Ligustrum sinense Lour.; C. DA, LCM, LGS, PS, RA, SW, UGS. 27. ONAGRACEAE Circaea lutetiana L. ssp. canadensis (L.) Asch. & Magnus; C. DA, LGS, LMS. 43. Gaura filipes Spach; O. DA, LGS. 20. Ludwigia alternifolia L.; I. PS, SW. 725. Ludwigia decurrens Walter; O. RA, WL. 496. Ludwigia palustris (L.) Elliot; I. RA, WL. Webb-TENN. Oenothera biennis L.; O. DA, RA. 502. Oenothera fruticosa L. ssp. fruticosa; I. LGS. Beck & Estes-TENN. Oenothera laciniata Hill; O. DA, RA. 494. OROBANCHACEAE Conopholis americana (L.) Wallr.; F. LCM, LGS, UGS. 326. 632 Southeastern Naturalist Vol. 11, No. 4 OXALIDACEAE Oxalis stricta L.; C. DA, LGS, SW. 131. Oxalis violacea L.; O. LGS, MSS. 318. PAPAVERACEAE Sanguinaria canadensis L.; S. LCM, MSS. 618. PASSIFLORACEAE Passiflora incarnata L.; I. DA, LGS. 113. Passiflora lutea L.; O. DA, LGS, RA. 478. PHYTOLACCACEAE Phytolacca americana L.; C. DA, LGS, PS, UGS. 107. PLANTAGINACEAE Plantago aristata Michx.; S. PL, PS. 675. *Plantago lanceolata L.; C. DA, PL, PS. 661. Plantago rugelii Decne.; F. DA, PS. 87. Plantago virginica L.; C. DA, PS. 28. PLATANACEAE Platanus occidentalis L.; C. DA, LGS, RA. 55. POLEMONIACEAE Phlox amoena Sims; C. DA, LGS, UGS. 21. Phlox amplifolia Britton; I. MSS, UGS. 693. Phlox carolina L.; O. DA, UGS. 656. Phlox divaricata L.; C. DA, LGS, SW, UGS. 6. POLYGALACEAE Polygala verticillata L. var. verticillata; I. DA, RA. Bridges & Somers-UCHT. POLYGONACEAE Polygonum hydropiperiodes Michx.; C. DA, RA. 425. *Polygonum persicaria L.; C. DA, LGS. 135. Polygonum punctatum Elliot var. punctatum; F. RA. 98. Polygonum scandens L. var. scandens; I. RA, WL. Bridges & Somers-UCHT. Polygonum virginianum L.; C. DA, LGS. 134. Rumex altissimus Alph. Wood; O. SW. Beck-TENN. PORTULACACEAE Claytonia virginica L.; I. LGS, MSS. 617. **Phemeranthus mengesii (W. Wolf) Kiger; R. PL, PS. 642. *Portulaca oleracea L.; I. DA, RA. 511. PRIMULACEAE Dodecatheon meadia L.; O. LCM. 580. Lysimachia tonsa (Alph. Wood) Alph. Wood ex Pax & R. Knuth; C. DA, LGS, UGS. 360. 2012 E. Blyveis and J. Shaw 633 PYROLACEAE Chimaphila maculata (L.) Pursh; C. LGS, LMS, SW, UGS. 48. RANUNCULACEAE Actaea pachypoda Elliot; O. LMS, MSS. 81. Actaea racemosa L. var. racemosa; O. MSS. 671. Anemone virginiana L.; I. LCM, LGS. Bridges & Somers-UCHT. *Clematis terniflora DC. var. terniflora; F. DA, RA. 137. **‡Clematis sp. nov.; VR. LCM. Estes-TENN. Delphinium tricorne Michx.; O. LCM. 320. Hepatica nobilis Schreb. var. acuta (Pursh) Steyerm.; F. LGS, MSS, UGS. 248. **Hydrastis canadensis L.; VR. MSS. Bridges & Somers-UCHT. Ranunculus abortivus L.; C. LGS, MSS, SW, UGS. 339. *Ranunculus ficaria L.; VR. RA. Harris-UCHT. Ranunculus hispidus Michx. var. hispidus; F. LGS, MSS. 596. Ranunculus micranthus Nutt.; O. LCM. 587. Ranunculus recurvatus Poir.; F. LGS, MSS, UGS. 338. Thalictrum clavatum DC.; I. LGS, MSS. 699. Thalictrum revolutum DC.; O. DA, LGS, MSS. 459. Thalictrum thalictroides (L.) Eames & B. Boivin; C. LCM, LGS, MSS. 3. RHAMNACEAE Berchemia scandens (Hill) K. Koch; F. LCM, LGS. 302. Ceanothus americanus L.; I. DA, PL, UGS. 683. Frangula caroliniana (Walter) A. Gray; F. DA, LGS, PL, PS, UGS. 367. ROSACEAE Agrimonia gryposepala Wallr.; I. DA. 491. Agrimonia pubescens Wallr.; C. DA, LGS, PS. 436. Agrimonia rostellata Wallr.; C. DA, LGS. 443. Amelanchier arborea (Michx. f.) Fernald; O. LGS, UGS. 63. Aruncus dioicus (Walter) Fernald; S. MSS. Bridges & Somers-UCHT. *Duchesnea indica (Andrews) Focke; F. DA. 290. Geum canadense Jacq.; C. DA, LGS, RA. 95. Geum virginianum L.; C. LCM, LGS. Beck-TENN. Gillenia stipulata (Muhl. ex Willd.) Baill.; F. LGS, PS, UGS. 467. Malus angustifolia (Aiton) Michx. var. angustifolia; O. DA, LGS. Beck-TENN. Physocarpus opulifolius (L.) Maxim., orth. cons.; F. DA, LGS. Beck & Estes-TENN. Potentilla canadensis L.; F. PS, UGS. 329. Potentilla simplex Michx.; C. DA, LGS. Bridges & Somers-UCHT. Prunus americana Marsh.; C. DA, LCM, LGS, LMS, RA, UGS, WL. 56. Prunus angustifolia Marsh.; I. DA. 490. Prunus mexicana S. Watson; I. LCM, WL. 377. *Prunus persica (L.) Batsch; O. LGS. 454. Prunus serotina Ehrh.; C. DA, LCM, LGS, PS, SW, UGS, WL. 40. *Pyrus calleryana Decne.; O. DA, RA. 480. *Rhodotypos scandens (Thunb.) Makino; I. DA, LGS. 603. Rosa carolina L.; O. UGS. Beck & Estes-TENN. *Rosa multiflora Thunb.; F. DA, RA. 511. Rubus argutus Link; C. DA, LGS. 16. *Rubus bifrons Vest ex Tratt.; O. DA. Beck-TENN. 634 Southeastern Naturalist Vol. 11, No. 4 Rubus flagellaris Willd.; C. DA, LCM. 694. Rubus occidentalis L.; F. DA, PL. 479. *Rubus phoenicolasius Maxim.; F. DA, PL. 477. RUBIACEAE Cephalanthus occidentalis L.; O. RA, WL. 408. Diodia teres Walter; C. DA. 567. Galium aparine L.; C. LGS, MSS, SW. 331. Galium circaezans Michx.; F. DA, LGS. 438. *Galium parisiense L.; I. DA, RA. 166. Galium tinctorium (L.) Scop.; O. RA, WL. 681. Galium triflorum Michx.; F. DA, MSS. 444. Houstonia canadensis Willd. ex Roem. & Schult.; C. LCM, LGS, PL, UGS. 327. Houstonia purpurea L. var. calycosa A. Gray; O. DA. Beck-TENN. Houstonia purpurea L. var. purpurea; F. DA, PS, UGS. 26. Houstonia pusilla Schoepf; S. UGS. 267. Mitchella repens L.; C. LCM, SW, UGS, WL. 388. RUTACEAE *Poncirus trifoliata (L.) Raf.; O. LCM. 301. SALICACEAE Populus deltoides Bartram ex Marsh.; I. DA, RA. 488. Salix caroliniana Michx.; O. RA, WL. 481. Salix nigra Marsh.; O. RA. 406. SAPOTACEAE Sideroxylon lycioides L.; O. LCM, WL. 380. SAURURACEAE Saururus cernuus L.; O. RA, WL. 91. SAXIFRAGACEAE Astilbe biternata (Vent.) Britton; O. LGS, MSS. 74. Heuchera americana L.; F. DA, LGS, MSS, SW, UGS. 44. Heuchera villosa Michx. var. villosa; S. LGS, LMS. 312. Saxifraga careyana A. Gray; I. LGS, MSS. 672. SCROPHULARIACEAE Agalinis purpurea (L.) Pennell; O. LCM. Beck-TENN. Aureolaria pectinata (Nutt.) Pennell; I. PS, SW, UGS. 730. Aureolaria virginica (L.) Pennell; O. UGS. 116. *Chaenorhinum minus (L.) Lange; O. DA, UGS. Beck-TENN. Chelone lyonii Pursh; I. LGS, SW. 169. Lindernia dubia (L.) Pennell var. dubia; O. RA, WL. Bridges & Somers-UCHT. Mimulus alatus Aiton; O. RA, WL. 484. Nuttallanthus canadensis (L.) D.L. Sutton; S. PL, PS. 639. *Paulownia tomentosa (Thunb.) Siebold & Zucc. ex Steud.; O. DA, LGS, PL, PS, RA. 1 14. Penstemon canescens (Britton) Britton; C. DA, LGS, PL, UGS. 25. *Verbascum blattaria L.; O. DA, RA. 31. *Verbascum thapsus L.; F. DA, LCM, RA. 506. 2012 E. Blyveis and J. Shaw 635 *Veronica hederifolia L.; F. DA. 260. *Veronica persica Poir.; F. DA. 255. SIMAROUBACEAE *Ailanthus altissima (Mill) Swingle; C. DA, LCM, LGS, PL, PS, RA. 33. SOLANACEAE Solanum carolinense L.; C. DA, PL. 368. Solanum ptychanthum Dunal; F. DA, LGS, UGS. 168. STAPHYLEACEAE Staphylea trifolia L.; O. LCM, RA. 525. STYRACACEAE Halesia tetraptera Ellis; F. LGS, MSS, RA. 399. THYMELAEACEAE Dirca palustris L.; O. LGS, MSS. 613. TILIACEAE Tilia americana L. var. americana; F. LCM, LGS, RA, WL. 86. Tilia americana L var. heterophylla (Vent.) Loudon; O. DA, LGS, RA. 30. ULMACEAE Celtis laevigata Willd.; I. LCM, WL. 369. Celtis occidentalis L.; F. DA, PL, PS, UGS. 247. Ulmus alata Michx.; C. DA, LCM, LGS, PL, PS, RA, SW, UGS, WL. 118. Ulmus americana L.; O. LCM, LGS, RA, WL. 410. Ulmus rubra Muhl.; F. DA, LCM, LGS, RA. 15. URTICACEAE Boehmeria cylindrica (L.) Sw.; F. LGS, RA. 94. Laportea canadensis (L.) Weddell; O. LGS, LMS, SW. 191. Pilea pumila (L.) A. Gray; C. LGS, MSS, RA, SW. 178. VALERIANACEAE Valerianella radiata (L.) Dufr.; O. LCM. 668. VERBENACEAE Callicarpa americana L.; F. LCM, LGS, UGS. 173. Phryma leptostachya L.; F. LCM. 704. Phyla lanceolata (Michx.) Greene; I. DA, RA. 471. *Verbena brasiliensis Vell.; O. DA. 139. Verbena simplex Lehm.; O. DA. 355. VIOLACEAE Hybanthus concolor (T.F. Forst.) Spreng.; I. LCM. 712. *Viola arvensis Murray; O. DA, LGS, MSS, UGS. 291. Viola bicolor Pursh; C. DA, LGS, PL, UGS. 261. Viola canadensis L.; F. LCM, LMS, UGS. 308. 636 Southeastern Naturalist Vol. 11, No. 4 Viola cucullata Aiton; S. LGS. 269. Viola hastata Michx.; F. LGS, MSS, SW, UGS. 311. Viola hirsutula Brainerd; O. LGS, MSS. 611. Viola x palmata L. (pro. sp.) [brittoniana or pedatifida x affinis or sororia]; O. LGS, UGS. 610. Viola pedata L.; I. DA, PS. 593. Viola pubescens Aiton var. scabriuscula Schwein. ex Torr. & A. Gray; I. LCM. 604. Viola rostrata Pursh; F. LGS, MSS, PS, UGS. 285. Viola sororia Willd.; O. LCM, LGS, MSS. 316. **Viola tripartita Ell.; S. LGS, MSS. 591. VISCACEAE Phoradendron leucarpum (Raf.) Reveal & M. C. Johnst.; O. DA, LGS, RA. 297. VITACEAE Ampelopsis cordata Michx.; F. DA, LCM, WL. 422. Parthenocissus quinquefolia (L.) Planch.; C. DA, LCM, LGS, LMS, PL, PS, RA, SW , UGS. 18. Vitis aestivalis Michx. var. aestivalis; C. DA, LGS, PL, UGS. 458. Vitis labrusca L.; F. DA, RA, UGS. 96. Vitis rotundifolia Michx.; C. DA, LCM, LGS, MSS, PS, SW, UGS. 437. Vitis vulpina L.; S. DA, LGS, UGS. 428.