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1 September 2016 The Vascular Flora of the University of the South, Sewanee, Tennessee
Jonathan P. Evans, Callie A. Oldfield, Mary P. Priestley, Yolande M. Gottfried, L. Dwayne Estes, Alfire Sidik, George S. Ramseur
Author Affiliations +

Sewanee: The University of the South, located in Franklin County, Tennessee, is a 5,263 ha site that encompasses a variety of plateau and cove habitats on the southern Cumberland Plateau in southeastern Tennessee. The vascular flora of the University of the South was documented from 1948–2015 and comprises 1,118 species and lesser taxa in 553 genera and 150 families. We documented 229 exotic taxa, 20.5% of the flora. This flora contains six state records, 74 Franklin County records, and two potential new species. Eighteen taxa are listed as protected either at the state or federal level, including the federally listed endangered Clematis morefieldii and state listed endangered Diamorpha smallii, Silphium brachiatum, and Symphyotrichum pratense. When compared to the five other published floras for the southern Cumberland Plateau in Tennessee, the University of the South flora is the most diverse, capturing 69% of the total taxa at the species level for the region. The high diversity of plant species in the University of the South flora reflects the broad range of habitats that can be found within the campus and contiguous natural areas. This flora demonstrates that herbaria at small liberal arts colleges can play an important role in the documentation and promotion of plant biodiversity within their region.

INTRODUCTION The campus and contiguous lands owned by Sewanee: The University of the South (UoS) (referred to locally as the “Sewanee Domain”) are located on the Cumberland Plateau in southeastern Tennessee, a region of high botanical diversity within the United States (Kartesz 2015). The area around UoS has been the focal point for several published floras that have contributed to our understanding of this diversity (Clark 1966, Wofford et al. 1979, Clements and Wofford 1991, Fleming and Wofford 2004, Beck and Van Horn 2007). A number of factors contribute to the potential for the UoS to capture a large percentage of the region's botanical diversity: (a) much of the land has been under continuous ownership since 1875, with management for education being a primary landowner objective; (b) there is a spectrum of habitats associated with a variety of historical land uses; and (c) the landscape encompasses a broad range of ecological communities associated with major variation in geology, hydrology, and topography.

The goals of this study were (a) to document the vascular flora of the UoS, (b) document rare or threatened plants, (c) document exotic plants, (d) document new state and Franklin county records, (e) list characteristic habitats for each taxon, and (f) compare floristic similarity of five Tennessee Cumberland Plateau floras. This study represents 50 yr of accumulated botanical study and collecting at this institution and is the first published record of the UoS flora.

Study Area

The land owned by the University of the South is located in southeast Tennessee. The majority of the land is in Franklin County, but a small portion reaches into Marion County (Figure 1). The UoS, a 5,263 ha area, is located between between 35°07′13″N and 35°13′52″N latitudes and between 85°51′24″W and 85°59′27″W longitudes. It is approximately 132 km southeast of Nashville and 58 km northwest of Chattanooga. The UoS occupies the Sewanee, Monteagle, and Sinking Cove USGS topographic sheet quadrangles and is part of the Appalachian Plateau Province. The landscape of the UoS consists of both upland (tableland) and cove environments, spanning an elevational range from 289 to 612 m. The UoS currently consists of an academic campus (155 ha) with adjacent commercial and residential areas (317 ha) that are embedded within and surrounded by diverse natural lands (4,791 ha). There are 15 reservoirs (67.5 ha), and the UoS is crossed by 9 km of major Tennessee Valley Authority (TVA) powerline rights-of-way (two lines, established in 1942 and 1951) that are kept free of woody vegetation. The continuum that exists on the UoS between the human-built environment and the surrounding natural environment contributes to the diversity of plant species.

Figure 1. 

Map of the University of the South lands with location within Tennessee. Plateau forest type is shown in dark gray, and cove forest type is shown in lighter gray. Streams and water bodies are shown in light gray. Paved roads and county lines overlay the map in black.


Land Use and Disturbance History

The lands that make up the present-day UoS have been affected by both prehistoric and historic human management. For 10,000 yr, Native Americans (the last of which were the Cherokees) used the Cumberland Plateau as seasonal hunting grounds and increased the level of fire on the landscape through their burning practices (Miller 2004). Settlers of European descent arrived in the Sewanee area in the early 1800s and began using land for grazing animals and resource extraction including coal and timber (Miller 2004). They also continued the practice of burning in the forests. By the time the University of the South was founded in 1857, Sewanee was a small town and much of the upland forest had been heavily logged to generate income (Cheston 1953). The arrival of the railroad on the mountain in 1852 promoted later economic activity associated with the new University and also resulted in fire disturbances.

In response to concerns about forest condition, in 1900 the University commissioned a forest assessment and plan for the UoS by Gifford Pinchot's Bureau of Forestry in the United States Department of Agriculture (USDA). This assessment concluded that fire, overgrazing, and overlumbering were suppressing regeneration and causing the forest to decrease in value (Foley 1903). During this time, the Department of Forestry was created at the University (Cheston 1953), and a forest manager was hired in the early 20th century to survey the forest and implement regulated timber extraction plans (Miller 2004). Fire suppression began in the 1900s, and by the 1950s, fires no longer represented an important ecological disturbance regime at Sewanee. Water impoundments were initially created for the purpose of fighting fires and later for providing drinking water and recreation areas. A network of firelanes was established in the 1930s and expanded in the 1950s, serving the dual purpose of fire control and recreational access across the upland area of the UoS. Pinus strobus was planted along firelanes to provide aerial visualization of compartment boundaries (Smith et al. 2010) and is now considered invasive as second-generation trees have spread from seed beyond original plantings. Pinus strobus, Pinus taeda, and other species were also planted in scattered experimental plantations by the University in conjunction with United States Forest Service (USFS) silvicultural scientists, who maintained a research station at the University from 1956–1985.

By the 1960s and continuing to the present day, the University significantly decreased logging activities in favor of preserving its forests and promoting recreation and aesthetics. Two old-growth forests, Dick Cove and Shakerag Hollow, have been set aside as a result of their biological diversity. In 2008, the University most recently acquired an adjacent 1,200 ha tract in Lost Cove that is currently managed as a natural area (Smith and Williamson 2008).

A University farm established in 1899 included 6 ha of land managed for pasture and vegetable production (Smith et al. 2010). The practice of running free-range livestock (cattle, pigs) in the forest that predated the University ended by the 1940s. The University farm fenced off large tracts of forest (400 ha) for pasture and running cattle. The original farm operation ceased in the 1960s, with farm fields being allocated to other purposes such as an equestrian center and sports fields. Some were maintained as open fields, and others were abandoned. A new University farm program was established in 2012 and currently occupies 1 ha of land used for agricultural purposes.

As a result of severe hunting pressure during the 1930s, white-tailed deer (Odocoileus virginianus) were largely eliminated from the Sewanee area. Deer were reintroduced in the 1950s, and a predator trapping program was established, targeting bobcats and feral dogs. By the late 1990s, deer densities had increased to levels impacting forest regeneration on the UoS (Reid et al. 2008).

Castanea dentata, once abundant on upland ridges and cove habitats with sandstone colluvium, was decimated by chestnut blight epidemics of the 1930s (Myers et al. 2004), so there are no reproductive individuals present on the UoS. Similarly, the dogwood anthracnose blight caused high mortality in Cornus florida populations in both cove and upland forest communities (Hiers and Evans 1997).


Typical of the southern Cumberland Plateau in Tennessee, the UoS has two major forest types draped across the plateau topography: an upland forest type and adjacent cove forest type (Hinkle 1989). The xeric upland forest is composed of a mixture of oak species (Quercus alba, Quercus coccinea, Quercus prinus, Quercus stellata, Quercus velutina) and hickory species (Carya glabra, Carya pallida, Carya tomentosa) in the canopy along with Acer rubrum, Nyssa sylvatica, and Oxydendrum arboreum (Reid et al. 2008). The understory of the upland forest is composed of a variety of woody shrubs including Kalmia latifolia, Rhododendron spp., and Vaccinium spp., along with a large number of grasses, sedges, and fall-blooming composites (Clements and Wofford 1991). The composition of the adjacent cove forest varies as a function of slope and aspect, but is generally characterized as having a diverse, mixed mesophytic canopy (Aesculus flava, Carya ovata, Juglans nigra, Liriodendron tulipifera, Tilia americana) with an oak (Quercus spp.) component and an Acer saccharum dominated understory and midstory (McGee 1986, Hinkle 1989, Reid et al. 2008). The cove forest understory has a diverse assemblage of herbaceous species (Duffy and Meier 1992).

The upland and cove forest communities differ ecologically. Upland plateau forest dynamics are driven by limited soil resource availability (Hinkle 1989), whereas cove forest dynamics are controlled to a greater degree by limited light availability (Martin 1992). Floristic studies from nearby locations on Cumberland Plateau in southern Tennessee (Wofford et al. 1979: Savage Gulf, Grundy County; Clements and Wofford 1991: Wolf Cove, Franklin County) have determined that less than 25% of the vascular plant species associated with the upland forest is also found in the cove forest. These studies also reveal that despite this dissimilarity, the plateau forest is just as diverse as the cove forest with upland forest species representing 48% of the total flora in both study areas.


The Cumberland Plateau represents the southern extension of the Appalachian Plateau that extends from West Virginia and Kentucky through Tennessee, terminating in Alabama. It is considered the westernmost part of the southern Appalachian region (Southern Appalachian Man and the Biosphere [SAMAB] 1996), bounded by the Ridge and Valley Province to the east and by the Interior Lowland Plateau to the west. The UoS is located on the southern portion of the Cumberland Plateau in Tennessee, which is characterized as having a flat to gently rolling surface underlain by highly resistant Pennsylvanian sandstones and shales (surface: Warren Point Sandstone, Sewanee Conglomerate and Whitwell Shale; Upper Slope: Signal Point Shales and Raccoon Mountain Formations). Where drainages have breached the sandstone, the less resistant Mississippian limestones underneath have eroded to form extensive steep-sided coves (Midslope: Pennington Formation; Lower Slope: Bangor Limestone and Hartselle Formation; Bottomlands: Monteagle Limestone; Fenneman 1938, Knoll et al. 2015).


The soils of this region reflect their underlying substrate and this, along with topographic position, is responsible for the large compositional differences between the forest of the plateau surface (hereafter referred to as plateau forest) and that of the coves (hereafter referred to as cove forest). Soils on the plateau are derived from sedimentary rocks and are acidic and well drained. Soils include Hartsells, Albertville, Lonewood, Ramsey, Hector, and Gilpin. Soils in the bottomlands of the cove are moderately fertile, deep, and moderately well drained. These soils include Sewanee, Ealy, and Clifty. Upper slopes of the cove developed from sandstone, siltstone, and shale. Soils on lower slopes developed from limestone. Slope soils include Grimsley, Jefferson, Bouldin, and Allen (Smalley 1982).

The soils of the plateau surface, being derived from the underlying sandstone, have a very high sand content. This condition makes these soils nutrient poor (low ion exchange capacity), drought prone, and highly acidic with little buffering capacity (Francis and Loftus 1977, Mays et al. 1991). Most of the fine root matter within the plateau forest soil is located in the upper 5 cm of the organic layer. This carpet-like mat of roots suggests that soil resource input (water and nutrients) is mostly coming from above through precipitation and litter turnover. In a study examining the potential impact of increased acid precipitation on cation-poor forest systems, Kelly (1988) observed an annual net loss of base cations (principally Ca2+ and Mg2+) over a five yr period from two forested plateau watersheds (Cross Creek, Franklin County, and Camp Branch, Bledsoe County) near UoS.


The climate of UoS is summarized using the last three decades of weather data taken on the uplands of the Cumberland Plateau by the University. The highest average temperatures occur in July (28.07°C) and August (28.08°C). The lowest average temperature occurs in January (−1.42°C). December and March have the highest precipitation, at 15.43 and 13.92 cm, respectively. Precipitation is lowest in May (11.61 cm) and August (11.33 cm). The average annual precipitation is 155.13 cm.

MATERIALS AND METHODS The vascular flora of the UoS has been studied informally throughout its history, and collections have been made as early as 1948. Some collections were made starting in 1958 as a result of a Plant Systematics course taught at the University. A formal inventory of the UoS flora began in 1995 with the creation of the Sewanee Herbarium (Index Herbariorum Code: UOS). Thorough descriptions of location and habitat are associated with each specimen, and species associations with major habitats have been described using UoS permanent vegetation plots. Standard references used in the identification of species include Radford et al. (1968), Cronquist (1980), Holmgren and Holmgren (1998), Wofford and Chester (2002), Jones (2005), and Weakley (2006). Nomenclature conforms to the Fifth Checklist of Tennessee Vascular Plants (Chester et al. 2009); exceptions were determined by D. Estes. The complete vouchers are stored in the Herbarium of Sewanee: The University of the South (UOS).

Presence of taxa within Franklin County was determined using the Biota of North America (BONAP) taxonomic data center (Kartesz 2015). Native and exotic taxa status was determined using USDA PLANTS database nativity data (USDA, NRCS 2016). When taxa were designated as “Native and Introduced” or “Cultivated, or not in the U.S.” we conformed to the status as listed in the Fifth Checklist of Tennessee Vascular Plants (Chester et al. 2009). Invasive exotic taxa were determined and ranked using the Tennessee Exotic Pest Plant Council list (TNEPPC; 2009). All exotic taxa listed in the appendix were also grouped into the following categories: Invasive, Introduced, and Persistent Planting. Invasive species were defined as all that were on the TNEPPC list (2009). Persistent Plantings were species that appeared to be planted, but had survived for 15 or more years in a natural habitat without tending. Introduced taxa are those that do not fit into the other categories. Not included in the flora are species that were deliberately planted and currently maintained as a defined planting. State and federal status of protected taxa were determined using the Tennessee Natural Heritage Program Rare Plant List (Crabtree 2014).

Floristic Comparison using Sørensen's Coefficient of Community

The flora of the UoS was compared to five others within the Cumberland Plateau in Tennessee to determine similarity. These include (with proximity to UoS noted): Fiery Gizzard (19 km; Clark 1966), Savage Gulf (39 km; Wofford et al. 1979), Wolf Cove (20 km; Clements and Wofford 1991), Fall Creek Falls State Park (74 km; Fleming and Wofford 2004), and Prentice Cooper State Forest and Wildlife Management Area (45 km; Beck and Van Horn 2007). We compared the similarity of these communities using Sørensen's coefficient of community (CC) repeating the methods in Beck and Van Horn (2007). This calculation did not incorporate area or other physical factors that might influence the results.


Survey of the Flora

A total of 3,135 collections were made on the UoS, representing 1,118 species and lesser taxa. The flora of the UoS represent 553 genera, and 150 families (Table 1). The four largest families are Asteraceae (151 taxa), Poaceae (98 taxa), Cyperaceae (76 taxa), and Fabaceae (56 taxa). These four families make up 34.3% of the total flora of the UoS. The largest genera are Carex (53 taxa), Viola (17 taxa), Solidago (15 taxa), Dichanthelium (12 taxa), and Symphyotrichum (12 taxa).

Table 1. 

Taxonomic summary of the flora of the University of the South, Franklin County, Tennessee. Exotic taxa include those that are introduced, invasive, exotic to the UoS, and persistent plantings.


New Species, State, and County Records

Two specimens (Helianthus sp. nov. and Sisyrinchium sp. nov.) are thought to be new species and are being evaluated by D. Estes. We documented four native state records, Agrimonia microcarpa, Carex rugosperma, Rubus longii, Viburnum alabamense, and two exotic state records, Arum italicum and Salvia pratensis. The taxa included in the UoS flora represent 38.7% of the Tennessee vascular plant taxa (Chester et al. 2009). This study documented 74 county records for Franklin County based on Biota of North America data (Kartesz 2015). This increases the number of documented species in Franklin County from 1,971 to 2,053. The taxa included in the UoS flora represent of 43.5% of the documented Franklin County species (Kartesz 2015).

Exotic Taxa

The UoS flora contains 229 exotic taxa, making up 20.5% of the total vascular plant flora. Of these, 73 are designated as invasive by the TNEPPC representing 54% of all invasive plant species in the state (2009). In the Appendix, the following species were considered exotic in Tennessee (Chester et al. 2009) despite their native status in USDA Plants Database (USDA, NRCS 2016): Amaranthus spinosus, Chamaesyce prostrata, Dichondra carolinensis, Gaillardia pulchella, Helianthus annuus, Iva annua, Magnolia grandiflora, Oenothera speciosa, Oxalis corniculata, and Prunella vulgaris. We also deviate from the USDA PLANTS database (USDA, NRCS 2016) and Fifth Checklist of Vascular Plants in Tennessee (Chester et al. 2009) in listing seven species as exotic to the UoS: Catalpa speciosa, Halesia tetraptera, Magnolia macrophylla, P. taeda, Quercus phellos, Taxodium distichum, and Tsuga canadensis. These represent naturalized populations that are the offspring of plantings. Pinus strobus was treated as another exception, listing it as both invasive and exotic to the Cumberland Plateau (Little 1971). Eighteen taxa were listed as persistent plantings: Abelia ×grandiflora, Forsythia viridissima, Hyacinthoides non-scripta, Ilex crenata, Lamiastrum galeobdolon, Lysimachia clethroides, Malus prunifolia, Narcissus triandrus, Picea abies, Philadelphus inodorus (native to Tennessee planting), Phlox subulata (native to Tennessee planting), Phyllostachys aurea, Prunus avium, Pyrus communis, Rosa wichuraiana, Viburnum carlesii, Viburnum rhytidophyllum, and Yucca flaccida (native to Tennessee planting). Of these persistent taxa, I. crenata, M. prunifolia, P.s aurea, and V. rhytidophyllum are beginning to invade Tennessee and these populations will continue to be monitored.

Protected Taxa

The UoS lands contain eighteen taxa (1.6% of the flora) that are considered protected at the state or federal level. The most notable of these are Clematis morefieldii, which is federally listed endangered (LE), and Diamorpha smallii, Silphium brachiatum, and S. pratense, which are listed endangered by the state. State threatened species include: Allium burdickii, Juglans cinerea, Liatris cylindracea, Lycopodiella alopecuroides, Phemeranthus teretifolius, Sceptridium jenmanii, Silphium pinnatifidum, and Trichomanes boschianum. Special concern species include: Berberis canadensis, Danthonia epilis, Helianthus eggertii, Lonicera dioica, Panax quinquefolius, and Thermopsis mollis.

Habitats and Characteristic Species

Species were assigned to one or more of 15 habitat types divided between upland (plateau surface) and cove environments (Table 2). Habitats within the upland and cove environments were further differentiated as a function of canopy openness, geology, topography and human disturbance regime. The 10 nonanthropogenic habitat types correspond to the NatureServe Terrestrial Ecological Classification System (NatureServe 2013) in the following way: Plateau Surface–Allegheny-Cumberland Dry Oak Forest and Woodland (CES 202.359), Southern Appalachian Low Elevation Pine Forest (CES 202.332); Sandstone Outcrop and Bluff–Cumberland Sandstone Glade and Barrens (CES 202.337); Plateau Wetland–Central Interior Highlands and Appalachian Sinkhole and Depression Pond (CES 202.018), Cumberland Seepage Forest (CES 202.361); Sandstone Cliff Face and Rockhouse–Cumberland Acidic Cliff and Rockhouse (CES 202.309), Southern Appalachian Spray Cliff (CES 202.288); Upper Slope Dry–Southern Interior Low Plateau Dry-Mesic Oak Forest (CES 202.898); Upper Slope Mesic–South-Central Interior Mesophytic Forest (CES 202.887); Limestone Outcrop and Glades–Central Interior Highlands Calcareous Glades and Barrens (CES 202.691), Southern Interior Calcareous Cliff (CES 202.356); Lower Slope Dry–Southern Ridge and Valley/Cumberland Dry Calcareous Forest (CES 202.457); Floodplain–South Central Interior Large Floodplain (CES 202.705); Lower Slope Mesic–South-Central Interior Mesophytic Forest (CES 202.887). The 5 anthropogenically-modified habitats are Reservoir, Former Agriculture, Lawn, Plateau Open, and Cove Open.

Table 2.  

List of habitats for upland and cove environments associated with the University of the South.


Upland Habitats

Plateau surface. The upland topography gently varies from convex to concave surfaces, and there are subtle changes in vegetation that follow these gradients. Oaks (Q. alba, Q. coccinea, Q. montana, and Q. velutina) and hickories (C. glabra, C. pallida, and C. tomentosa) dominate the canopy along with A. rubrum, N. sylvatica, O. arboreum, and Prunus serotina. There is a distinct understory layer of shrubs and small trees including: Sassafras albidum, Vaccinium corymbosum, Vaccinium stamineum, and Viburnum acerifolium. Common vines include: Parthenocissus quinquefolia, Smilax spp., Toxicodendron radicans, and Vitis vulpina. Common herbaceous species include: Carex digitalis var. digitalis, Chimaphila maculata, Coreopsis major, Dichanthelium acuminatum, Dichanthelium boscii, Ipomaea pandurata, Houstonia caerulea, and Houstonia purpurea. Arundinaria appalachiana, a species associated with this habitat, was recently described as a distinct species (Triplett et al. 2006) and forms large clonal populations along upland drainages.

Sandstone outcrop and bluff. On the plateau surface, there are places where the soil is thin and the underlying sandstone is sometimes exposed. Gradients of soil depth determine openness of canopy. On the UoS, sandstone outcrops are associated with the Sewanee Conglomerate formation. Shallow depressions and fissures in the rock surface maintain a thin layer of soil and support a variety of annual and perennial herbs including four plant species that are endemic on the UoS to these habitats: D. smallii, Minuartia glabra, Opuntia humifusa, and Phemeranthus teretifolius. Sandstone outcrops are generally embedded in a matrix of open woodland of low statured trees dominated by Pinus virginiana, Q. coccinea, Quercus marilandica, Q. montana, and Vaccinium arboreum. On the bluff edge and upland ridges, Q. montana is often the dominant species, sometimes forming open woodlands with an ericaceous understory of heath species including: K. latifolia, V. corymbosum, and Viburnum pallidum. These ridges were once codominated by C. dentata before the chestnut blight (Foley 1903). At one such bluff edge site, a remnant Pinus echinata population can be found comingled with Q. prinus. Cypripedium acaule and Danthonia sericea are characteristic herbs of this habitat.

Plateau wetland. There are a variety of wetland habitats associated with the plateau surface including streams, vernal pools, and maple-gum swamps. Canopy species associated with these habitats include: A. rubrum, L. tulipifera, Liquidambar styraciflua, N. sylvatica, and Q. alba. Species typical of the subcanopy and shrub layer include A. rubrum, Aronia arbutifolia, Ilex opaca, O. arboreum, Rhododendron canescens, Rubus hispidus, Smilax bona-nox, Vaccinium fuscatum, and Viburnum nudum. A variety of ferns, forbs, and sedges are common in the understory, including Carex debilis, Medeola virginiana, Mitchella repens, Osmunda regalis, Osmundastrum cinnamomea, Polygonatum biflorum, and Thelypteris noveboracensis. Bartonia virginica is an uncommon mycoheterotrophic species restricted to the fringes of vernal pools and upland seeps.

Reservoir. Reservoirs represent novel habitats on the Cumberland Plateau since there are no natural permanent bodies of water. Plants species associated with the open aquatic habitats arrived through long-distance dispersal and have accumulated over time. Brasenia schreberi is a common, rooted, floating species found in most reservoirs. Rooted submerged and emergent species associated with reservoir edges include: Callitriche heterophylla, Eleocharis spp., Fimbristylis autumnalis, Iris pseudacorus, Isoetes engelmannii, Juncus spp., Murdannia keisak, Myriophyllum aquaticum, Scirpus georgianus, Potamogeton spp., and Rhynchospora spp.

Former agriculture. In addition to the former agricultural land associated with the University farm, there are several small abandoned farm areas on the upland portion of UoS that are in the process of reverting back to forest. Some of these fields were planted in P. taeda and P. strobus by the University after they were no longer used for agricultural purposes. Pinus strobus plantations still persist in places; however, many of the P. taeda stands have succumbed to pine bark beetle mortality. Pinus strobus has spread into the native forest as a result of wind dispersal from these plantings creating belts of dense understory pine parallel to these roads. The plant communities that have recolonized former farm sites represent novel mixtures of both cove and plateau species, along with introduced species (Lonicera japonica, Microstegium vimineum, and Rosa multiflora) and persistent plantings (P. avium, P. communis, and Y. flaccida). The intensive liming of the soil within farmed areas substantially changed soil properties and may be contributing to the presence of the more calciphilic species typical of the nearby cove habitats such as A. saccharum and Quercus muehlenbergii. Common tree species include A. rubrum, C. florida, Diospyros virginiana, L. tulipifera, L. styraciflua, P. virginiana, and Robinia pseudoacacia. Common herbs include Asplenium platyneuron, Botrypus virginianus, Carex albicans var. albicans, Cynoglossum virginianum, Oxalis dillenii, Polystichum acrostichoides, Pycnanthemum loomisii, and Sanicula canadensis.

Lawn. The University campus and its associated residential areas have maintained open mown lawns, cemeteries, and fields for more than 175 years. These areas have accumulated novel assemblages of grasses and forbs that are dominated by introduced and native species, such as Anthoxanthum odoratum, Cerastium fontanum, Cynodon dactylon, Dichondra carolinensis, Glechoma hederacea, Ranunculus abortivus, Schedonorus arundinaceus, Taraxacum officinale, Trifolium repens, and Viola sororia.

Plateau open. A large group of native and exotic species can be found in edge habitats at the intersection of forest and various land uses such as roads, fields, utility corridors and other clearings in the upland portion of the UoS. Wooded greenways along creeks through the campus and residential areas of the UoS represent another novel assemblage of species dominated by woody exotics including Celastrus orbiculatus, Lespedeza cuneata, Ligustrum sinense, Lonicera fragrantissima, L. japonica, and R. multiflora.

Cove Habitats

Sandstone cliff face and rockhouse. Beneath the bluff, where the sandstone cap of the plateau upland gives way to cove below, there are a number of habitats that include the sandstone cliff face, rockhouse, and spray zones of waterfalls. The sandstone cliff face represents vertical sandstone habitats ranging from dry to wet, including waterfall spray zones. Characteristic species include: Asplenium montanum, Asplenium pinnatifidum, Dentaria multifida, Heuchera spp., Huperzia porophila, Iris cristata, Micranthes careyana, M. repens, Rhododendron maximum, Sedum ternatum, Silene rotundifolia, Stellaria pubera, and Thalictrum spp. Rhododendron maximum is at the southwestern edge of its range (Kartesz 2015) and is represented on UoS by a single population. At the base of these cliffs, where less resistant sedimentary layers have eroded, there exist extensive overhangs, or rockhouses, that Native Americans used for thousands of years as temporary rock shelters (Walck et al 1996). Trichomanes boschianum is a rare fern endemic to this unique habitat.

Upper and lower slope mesic. Upper cove slopes with a north or east facing aspect support what is typically referred to as a “mixed mesophytic” community (Duffy and Meier 1992). This moist cove forest type supports a rich assemblage of characteristic canopy species including A. saccharum, A. flava, C. ovata, Fraxinus biltmoreana, J. nigra, Quercus rubra, and T. americana. Understory trees and shrubs include Asimina triloba, Cercis canadensis, Lindera benzoin, and Staphylea trifolia. The old growth forests in Shakerag Hollow and Dick Cove support a diverse vernal herbaceous flora: Arisaema triphyllum, Caulophyllum thalictroides, Claytonia virginica, Delphinium tricorne, Dicentra cucullaria, Hydrophyllum canadense, Podophyllum peltatum, Polymnia laevigata, Phacelia bipinnatifida, Phlox divaricata, Sanguinaria canadensis, Tradescantia subaspera, Trillium spp., and Viola spp. On the lower slopes, deeper and more basic soils support the highest plant diversity on the UoS. Characteristic canopy species include A. saccharum, Celtis occidentalis, Fagus grandifolia, F. biltmoreana, and Q. muehlenbergii. Understory herb species include: Anenome acutiloba, Epifagus virginiana, Hexastylis arifolia, and Sanicula spp. Understory vines and shrubs include: Isotrema macrophyllum, Bignonia capreolata, Euonymus americanus, L. benzoin, P. quinquefolius, Smilax spp., and Viburnum rufidulum. Large, dense Arundinaria gigantea clones can be found in some locations within this habitat.

Upper and lower slope dry. Upper slopes of the cove with a south or west facing aspect support forest communities characterized by the following woody plants: A. saccharum, C. ovata, F. biltmoreana, L. tulipifera, Q. alba, R. pseudoacacia, Ulmus alata, and V. rufidulum. The herbaceous community includes A. triphyllum, Bromus pubescens, D. boscii, Galium spp., P. acrostichoides, Scleria oligantha, and Smallanthus uvedalius. Upper portions of both dry and mesic slopes often contain sandstone colluvium that host bouldertop communities (P. bipinnatifida, P. biflorum, Polymnia laevigata, Polypodium appalachianum, T. radicans, and Tradescantia subaspera). This unique plant assemblage is associated with the shallow soils that accumulate on top of boulders, and has been shown to serve as a refugia in areas of high deer browse (Comisky et al. 2005). With decreasing elevation in the cove, the influence of limestone substrate on soil characteristics becomes more important and there is a corresponding shift in species composition. On the lower portion of dry slopes, common species in the canopy and midstory include: A. saccharum, Carya carolinae-septentrionalis, C. glabra, Crataegus spp., F. biltmoreana, Ostrya virginiana, Q. alba, Q. coccinea, Rhus aromatica, U. alata, and V. rufidulum. Common vines include: B. capreolata, Campsis radicans, and T. radicans. The following are common herbaceous understory species: Carex oxylepis, H. arifolia, Solidago auriculata, Vernonia flaccidifolia, and Viola spp.

Limestone outcrop and glades. Places within the lower coves that have exposed limestone (cliff faces, outcrops) support a unique flora including Asplenium ruta-muraria, L. dioica, Pellaea atropurpurea, and Polypodium polypodioides. Outcrops can be surrounded by an open woodland or glade community dominated by a rich assemblage of trees and vines: Amelanchier arborea, Berchemia scandens, B. capreolata, C. carolinae-septentrionalis, Celtis tenuifolia, C. canadensis, Fraxinus quadrangulata, Juniperus virginiana, Ostrya virginiana, P. quinquefolia, Q. rubra, Rhamnus caroliniana, R. aromatica, Sideroxylon lycioides, Symphoricarpos orbiculatus, and U. alata. Characteristic herbs include Brachyelytrum erectum, Desmodium spp., D. boscii, Elymus hystrix, Eryngium yuccifolium, Manfreda virginica, Matelea carolinensis, S. oligantha, Ruellia humilis, Salvia lyrata, and Zizia aptera.

Floodplain. There is limited bottomland habitat present on the UoS along Lost Cove Creek and at the bottom of Shakerag Hollow on Mud Creek. There is a small portion of floodplain habitat near the big sink that experiences substantial annual flooding. Canopy tree species include Acer negundo, A. saccharum, F. grandifolia, L. tulipifera, and L. styraciflua. Understory woody species include Carpinus caroliniana, Cornus alternifolia, and L. benzoin. Alluvial soils support a characteristic herbaceous community: Arnoglossum reniforme, Diphasiastrum digitatum, Mertensia virginica, Pedicularis canadensis, and Samolus valerandi.

Cove open. Old fields, abandoned homesites, roadsides, and TVA transmission line rights-of-way are included here. Limestone-based soils vary in depth, and slope varies from level to gently sloping. This habitat is dominated by graminoids and forbs. Graminoid species include Carex spp., Chasmanthium sessiliflorum, Elymus virginicus, Muhlenbergia sobolifera, Panicum spp., and Saccharum alopecuroides. Herbs include Allium canadense, Anemone virginiana, Calystegia silvatica, Chenopodium album, Desmodium cuspidatum, Frasera carolinensis, Lespedeza procumbens, Packera glabella, R. humilis, S. auriculata, and V. flaccidifolia. Woody plants include semiwoody Hypericum sphaerocarpum, as well as lianas, Cocculus carolinus and Lonicera sempervirens, and small trees such as Crataegus intricata and Prunus americana.

Two TVA transmission powerline rights-of-way (constructed in 1942 and 1951) extend across the UoS property. With the primary management objective of keeping the lines clear of woody plants, TVA's approach is to maintain an early-successional habitat dominated by woodland grasses, ferns, and low-growing shrubs. A limestone barren type habitat is present in the older right-of-way on a west-facing slope of the plateau. This small (0.5 ha) area is characterized by thin calcareous soil with patches of exposed rock and gravel on level to slightly sloping ground. Dominant perennial grasses include Panicum spp., Schizachyrium scoparium var. scoparium, and Sorghastrum nutans. The many sedge species include Carex glaucodea, C. granularis, C. kraliana, C. meadii, and C. vulpinoidea. Rare herbaceous species, including L. cylindracea, S. pinnatifidum, and S. pratense, are found here, among other forbs such as M. virginica, Ratibida pinnata, and Verbesina alternifolia.

Community Comparisons

The flora of UoS was compared to the species found in five other southern Cumberland Plateau floras: Fiery Gizzard (Clark 1966), Savage Gulf (Wofford et al. 1979), Wolf Cove (Clements and Wofford 1991), Fall Creek Falls (Fleming and Wofford 2004), and Prentice Cooper (Beck and Van Horn 2007). If the intersection of these six floras is interpreted to be a representation of a regional flora for the southern Cumberland Plateau in Tennessee (1,595 species of vascular plants), then UoS flora is the most diverse, capturing 69% of all taxa at the species level. The coefficient of community indicated that it was most similar to Fall Creek Falls (total: 70.6; native-only: 72.9) and Prentice Cooper (total: 66.8; native-only: 70.8) when considering both the total flora and the native only species. The least similar was Fiery Gizzard (total: 55.8; native-only: 61.2). The flora of the UoS contained the highest percentage of exotic species (20.7%), with Prentice Cooper coming in second (17.1%), followed by Fall Creek Falls (13.1%). Wolf Cove (5.3%), Savage Gulf (6.4%), and Fiery Gizzard (6.9%) had fewer exotic species. When taking into account the total species found at each location, we found that the UoS flora contained 84.4% of the Wolf Cove flora, 79.7% of the Fall Creek Falls flora, and 79.8% of the Fiery Gizzard flora, closely followed by the others (Table 3).

Table 3. 

Comparison of Tennessee Cumberland Plateau floras including number of species, number of species on University of the South (UoS), percent of total species on the UoS, number of native and exotic species, and community coefficient of native and exotic species.


CONCLUSION The UoS flora represents a large percentage of the total Tennessee flora (39%; Chester et al. 2009) as well as the flora of Franklin County (44%; Kartesz 2015) and the southern Cumberland Plateau region (69%). At the species level, the UoS flora is the most diverse flora compiled for the south Cumberland Plateau in Tennessee and it contains 18 listed state/federal rare, threatened, and endangered species along with six state records. The high diversity of plant species in the UoS flora reflects the broad range of habitats that can be found within the campus and contiguous natural areas of the University of the South. The UoS property contains 13 of the characteristic ecological systems associated with the Cumberland Plateau region as identified by NatureServe (2013). Augmenting the high diversity of native plant species are a novel assemblage of introduced species that reflect the long history of varied land use within this landscape.

Colleges and universities in this country that own natural areas have a unique opportunity to both protect and study the biodiversity for which they are stewards (Muller and Maehr 2000). While natural area stewardship is now recognized as an important part of the sustainability commitment manifest by academic institutions (Krasny and Delia 2014), it is unusual for American universities or colleges to make conservation of biodiversity a primary objective of land management (Straka 2010). As one of the largest owners of natural lands of any college or university in the country, the University of the South has prioritized the protection of biodiversity and specifically its rich vascular plant flora as part of its ecosystem management philosophy. In addition, having a carefully catalogued living flora in association with an academic campus provides an unparalleled educational opportunity for the study of botany. Finally, this flora demonstrates that herbaria at small liberal arts colleges can play an important role in the documentation and promotion of plant biodiversity within their region (Boyd 2008, Thompson 2008).

ACKNOWLEDGMENTS The authors would like to thank the many former students and staff of the Sewanee Herbarium who have contributed their time and expertise to this project over the years. In particular, we would like to acknowledge the following individuals: Ashley Block, Nathan Bourne, Caitlin Elam, Nick Hollingshead, Jacquelyn Presley, Leighton Reid, Katie Qualls, Emily Sprouse, and Hali Steinman. We acknowledge the Friends of the Sewanee Herbarium and the Department of Biology for financial support to this project. Gratitude is also extended to Al Good and Todd Crabtree.



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ANNOTATED APPENDIX This list of taxa consists of five groups: Pteridophytes, Gymnosperms, Basal Dicots, Monocots, and Eudicots. Taxa are arranged inside of these groups alphabetically by family. Nomenclature follows the Tennessee Flora; exceptions are determined by D. Estes. An asterisk (*) indicates that the taxa is exotic. A double asterisk (**) indicates that it is considered invasive according to Tennessee Exotic Pest Plant Council (2009). A plus sign (+) indicates that it is not native to the southern Cumberland Plateau, but is native to the Southeastern United States. Protected taxa are noted with bold letters; LE = listed endangered, E = endangered, T = threatened, S = special concern, CE = commercially exploited. Characteristic habitats are given as codes: 1 = Sandstone Outcrop and Bluff, 2 = Plateau Surface, 3 = Plateau Wetland, 4 = Former Agriculture, 5 = Reservoir, 6 = Lawn, 7 = Plateau Open, 8 = Sandstone Cliff Face and Rockhouse, 9 = Upper Slope Dry, 10 = Upper Slope Mesic, 11 = Limestone Outcrop and Glades, 12 = Lower Slope Dry, 13 = Lower Slope Mesic, 14 = Floodplain, 15 = Cove Open. Following the habitat codes is the specimen collection number.



Asplenium montanum Willd.; 8.UOS-100.

Asplenium pinnatifidum Nutt.; 8.UOS-6269.

Asplenium platyneuron (L.) B.S.P.; 3, 4, 7, 8, 9, 10, 11, 13, 14.UOS-1366.

Asplenium resiliens Kunze; 11.UOS-3988.

Asplenium rhizophyllum L.; 8, 10, 11.UOS-1651.

Asplenium ruta-muraria L.; 11.UOS-8177.

Asplenium trudelli Wherry (pro sp.) [montanum × pinnatifidum]; 8.UOS-8530.


Woodwardia areolata (L.) T. Moore; 3.UOS-1705.


Dennstaedtia punctilobula (Michx.) T. Moore; 2.UOS-6138.

Pteridium aquilinum (L.) Kuhn; 7.UOS-139.


Dryopteris celsa (Wm. Palmer) Knowlt., Palmer & Pollard ex Small; 14.UOS-8333.

Dryopteris goldiana (Hook. ex Goldie) Gray; 10.UOS-7799.

Dryopteris marginalis (L.) Gray; 10, 14.UOS-1360.

Polystichum acrostichoides (Michx.) Schott; 1, 2, 3, 4, 9, 10, 11, 13, 14, 15.UOS-2210.


Equisetum arvense L.; 7.UOS-2487.


Trichomanes boschianum Sturm; 8.UOS-6265. T.

Trichomanes intricatum Farrar; 13.UOS-8527.


Isoetes engelmannii A. Braun; 5.UOS-8511.


Dendrolycopodium obscurum (L.) A.Haines; 2, 4.UOS-8634.

Diphasiastrum digitatum (Dill. ex A. Braun) Holub; 2, 4, 14.UOS-4845.

Huperzia porophila (Lloyd & Underw.) Holub; 8.UOS-4861.

Lycopodiella alopecuroides (L.) Cranfill; 5, 7.UOS-7688. T.


Onoclea sensibilis L.; 3.UOS-2476.


Botrypus virginianus (L.) Holub; 3, 9, 10, 11, 14.UOS-4473.

Ophioglossum pycnostichum (Fernald) A. Löve & D. Löve; 3, 14.UOS-6251.

Sceptridium biternatum (Sav.) Lyon; 3, 7, 10, 14.UOS-2053.

Sceptridium dissectum (Spreng.) Lyon; 4.UOS-8322.

Sceptridium jenmanii (Underw.) Lyon; 4.UOS-8371. T.


Osmunda claytoniana L.; 3.UOS-8889.

Osmunda regalis L. var. spectabilis (Willd.) A. Gray; 3.UOS-2685.

Osmundastrum cinnamomea L.; 1, 3.UOS-7912.


Pleopeltis polypodioides (L.) Andrews & Windham var. michauxiana (Weath.) Andrews & Windham; 11.UOS-4871.

Polypodium appalachianum Haufler & Windham; 1, 8.UOS-1369.

Polypodium virginianum L.; 10.UOS-4033.


Adiantum pedatum L.; 10.UOS-3989.

Cheilanthes lanosa (Michx.) D.C. Eaton; 11.UOS-123.

Pellaea atropurpurea (L.) Link; 11, 15.UOS-7413.

Vittaria appalachiana Farrar & Mickel; 8.UOS-8367.


Phegopteris hexagonoptera (Michx.) Fée; 9, 10, 13.UOS-2700.

Thelypteris noveboracensis (L.) Nieuwl.; 2, 3.UOS-2735.


Athyrium filix-femina (L.) Roth ssp. asplenioides (Michx.) Hultén; 3.UOS-2681.

Cystopteris bulbifera (L.) Bernh.; 10, 11.UOS-3984.

Cystopteris protrusa (Weatherby) Blasdell; 8, 10, 14.UOS-6342.

Cystopteris tennesseensis Shaver; 8, 11.UOS-8496.

Deparia acrostichoides (Sw.) M. Kato; 14.UOS-8655.

Diplazium pycnocarpon (Spreng.) Broun; 9, 10.UOS-3990.

Woodsia obtusa (Spreng.) Torr.; 13.UOS-8620.



Juniperus virginiana L.; 1, 2, 4, 7, 9, 11, 14.UOS-1394.

+Taxodium distichum (L.) Rich.; 7.UOS-8288.


*Picea abies (L.) Karst.; 7.UOS-8522.

Pinus echinata Mill.; 1.UOS-1401.

+Pinus strobus L.; 2, 3, 4, 7.UOS-1395.

+Pinus taeda L.; 4, 7.UOS-1403.

Pinus virginiana Mill.; 1, 2, 4, 7.UOS-1402.

+Tsuga canadensis (L.) Carr.; 7.UOS-8215.

Basal Dicots


Asimina triloba (L.) Dunal; 4, 9, 10.UOS-3320.


Asarum canadense L.; 10.UOS-4034.

Endodeca serpentaria (L.) Raf.; 2, 9, 10, 11, 12, 13, 14.UOS-7725.

Hexastylis arifolia (Michx.) Small var. ruthii (Ashe) Blomquist; 10, 12, 13, 14.UOS-1570.

Isotrema macrophyllum (Lam.) C.F.Reed; 10, 11.UOS-7803.


Brasenia schreberi J.F. Gmel.; 5.UOS-1993.


Calycanthus floridus L. var. floridus; 2, 3.UOS-4447.


Lindera benzoin (L.) Blume; 9, 10, 13, 14.UOS-546.

Sassafras albidum (Nutt.) Nees; 1, 2, 3, 9, 11.UOS-513.


Liriodendron tulipifera L.; 1, 2, 3, 4, 7, 9, 10, 13, 14.UOS-4484.

Magnolia acuminata (L.) L.; 9, 10, 12, 13, 14.UOS-3322.

+Magnolia grandiflora L.; 2.UOS-4851.

+Magnolia macrophylla Michx.; 2, 3.UOS-4806.


Saururus cernuus L.; 5.UOS-2200.



Camassia scilloides (Raf.) Cory; 14.UOS-8368.

Manfreda virginica (L.) Salisb. ex Rose; 11, 15.UOS-4733.

+Yucca filamentosa L.; 7.UOS-6249.


Alisma subcordatum Raf.; 7.UOS-7709.

Sagittaria australis (J.G. Sm.) Small; 3.UOS-2067.


Allium burdickii (Hanes) A.G. Jones; 10.UOS-7702. T-CE.

Allium canadense L.; 9, 15.UOS-7866.

Allium cernuum Roth; 10, 11.UOS-7717.

**Allium vineale L.; 4, 7.UOS-948.

Nothoscordum bivalve (L.) Britton; 7.UOS-3709.


*Leucojum aestivum L.; 3.UOS-7807.

*Narcissus poeticus L.; 4.UOS-7580.

*Narcissus pseudonarcissus L.; 10.UOS-7565.

*Narcissus triandrus L.; 7.UOS-8888.


Arisaema dracontium (L.) Schott; 9.UOS-8713.

Arisaema triphyllum (L.) Schott; 9, 10, 11, 13, 14.UOS-2313.

*Arum italicum Mill.; 7.UOS-8247.

Landoltia punctata (G. Mey.) D.H. Les & D.J. Crawford; 5.UOS-8616.


*Asparagus officinalis L.; 7.UOS-2059.


Uvularia grandiflora Sm.; 10, 12.UOS-7870.

Uvularia perfoliata L.; 9, 10, 11, 12.UOS-934.

Uvularia sessilifolia L.; 3, 11.UOS-6271.


*Commelina communis L.; 7, 14.UOS-2065.

Commelina virginica L.; 3.UOS-7860.

**Murdannia keisak (Hassk.) Hand.-Maz.; 5.UOS-8883.

Tradescantia ohiensis Raf.; 2.UOS-7926.

Tradescantia subaspera Ker-Gawl.; 9, 10, 14.UOS-2075.


Carex aggregata Mackenzie; 15.UOS-8365.

Carex albicans Willd. ex Spreng. var. albicans; 4, 15.UOS-6608.

Carex albolutescens Schwein.; 3.UOS-8006.

Carex albursina Sheldon; 10.UOS-7822.

Carex amphibola Steud.; 4, 10, 11.UOS-8327.

Carex atlantica Bailey ssp. atlantica; 7.UOS-1695.

Carex basiantha Steud.; 4, 9.UOS-8703.

Carex blanda Dewey; 2, 10, 13, 14.UOS-6606.

Carex careyana Torr. ex Dewey; 10, 14.UOS-8658.

Carex caroliniana Schwein.; 4, 9.UOS-8675.

Carex cephalophora Muhl. ex Willd.; 7, 12.UOS-8005.

Carex cherokeensis Schwein.; 7.UOS-8374.

Carex communis Bailey; 10, 15.UOS-8325.

Carex complanata Torr. & Hook.; 5.UOS-3698.

Carex crinita Lam. var. brevicrinis Fern.; 3, 7.UOS-8022.

Carex cumberlandensis Naczi, Kral & Bryson; 9, 10.UOS-8001.

Carex debilis Michx. var. debilis; 2, 3.UOS-8400.

Carex digitalis Willd. var. digitalis; 2, 10.UOS-8302.

Carex frankii Kunth; 3, 15.UOS-1230.

Carex glaucescens Ell.; 3.UOS-8021.

Carex glaucodea Tuckerman ex Olney; 15.UOS-8649.

Carex granularis Muhl. ex Willd.;14.UOS-6604.

Carex gravida Bailey; 13.UOS-8397.

Carex gynandra Schwein.; 3.UOS-8755.

Carex hirsutella Mackenzie; 2, 4, 11, 12, 15.UOS-8328.

Carex intumescens Rudge; 3.UOS-1691.

Carex jamesii Schwein.; 4, 9, 15.UOS-8644.

Carex joorii Bailey; 3.UOS-4317.

Carex kraliana Naczi & Bryson; 12, 15.UOS-8642.

Carex laxiflora Lam.; 14.UOS-8654.

Carex longii Mackenzie; 1, 7.UOS-8366.

Carex louisianica Bailey; 3.UOS-7826.

Carex lurida Wahlenb.; 3, 10.UOS-2632.

Carex meadii Dewey; 15.UOS-8641.

Carex mitchelliana M.A. Curtis; 3.UOS-8517.

Carex nigromarginata Schwein.; 2, 10.UOS-7999.

Carex oligocarpa Schkuhr ex Willd.; 11, 12, 15.UOS-8329.

Carex oxylepis Torr. & Hook. var. oxylepis; 12, 13, 15.UOS-8355.

Carex pensylvanica Lam.; 1, 9, 12.UOS-8638.

Carex plantaginea Lam.; 10.UOS-1683.

Carex prasina Wahlenb.; 10.UOS-7825.

Carex purpurifera Mackenzie; 9, 10, 11, 14.UOS-7610.

Carex reznicekii Werier; 2, 10.UOS-8659.

Carex rosea Schkuhr ex Willd.; 13, 14.UOS-4464.

Carex rugosperma Mack.; 1.UOS-8637.

Carex seorsa Howe; 10.UOS-7640.

Carex striatula Michx.; 2, 9.UOS-8646.

Carex styloflexa Buckley; 3.UOS-7641.

Carex swanii (Fern.) Mackenzie; 3.UOS-7997.

Carex texensis (Torr.) L.H. Bailey; 14.UOS-8754.

Carex venusta Dewey; 14.UOS-8753.

Carex virescens Muhl. ex Willd.; 2, 3, 4, 7.UOS-4470.

Carex vulpinoidea Michx.; 7, 15.UOS-3054.

Cyperus croceus Vahl; 6.UOS-1759.

Cyperus echinatus (L.) Alph. Wood; 6.UOS-4802.

*Cyperus esculentus L.; 7.UOS-8882.

Cyperus flavescens L.; 7.UOS-3704.

Cyperus squarrosus L.; 1.UOS-8364.

Cyperus strigosus L.; 7.UOS-525.

Dulichium arundinaceum (L.) Britton; 5.UOS-6334.

Eleocharis acicularis (L.) Roem. & Schult.; 5.UOS-7707.

Eleocharis erythropoda Steud.; 9.UOS-8765.

Eleocharis microcarpa Torr. var. filiculmis Torr.; 5.UOS-7533.

Eleocharis obtusa (Willd.) Schult.; 5.UOS-1699.

Eleocharis quadrangulata (Michx.) Roem. & Schult.; 5.UOS-2719.

Eleocharis tenuis (Willd.) Schult. var. verrucosa (Svens.) Svens.; 5, 7.UOS-6601.

Fimbristylis autumnalis (L.) Roem. & Schult.; 5.UOS-7844.

Isolepis carinata Hook. & Arn. ex Torr.; 6.UOS-8714.

Rhynchospora capitellata (Michx.) Vahl; 5.UOS-3261.

Rhynchospora glomerata (L.) Vahl; 5.UOS-6610.

Schoenoplectus purshianus (Fernald) M.T. Strong; 5.UOS-4826.

Scirpus atrovirens Willd.; 3, 5.UOS-3701.

Scirpus cyperinus (L.) Kunth; 5, 7.UOS-3084.

Scirpus georgianus R.M.Harper; 5.UOS-3699.

Scirpus pendulus Muhl.; 5, 15.UOS-8353.

Scleria oligantha Michx.; 4, 7, 9, 11, 15.UOS-7406.


**Dioscorea polystachya Turcz.; 7, 14.UOS-4456.

Dioscorea villosa L.; 2, 7, 9, 10, 11, 12, 13.UOS-7992.


*Hemerocallis fulva (L.) L.; 7, 15.UOS-906.


*Hyacinthoides non-scripta (L.) Chouard ex Rothm.; 7.UOS-8277.

**Muscari neglectum Guss. ex Ten.; 6, 7.UOS-3457.

**Ornithogalum umbellatum L.; 6, 10.UOS-7569.


**Egeria densa Planch.; 5.UOS-1687.

**Najas minor All.; 5.UOS-2111.


Hypoxis hirsuta (L.) Coville; 2, 10, 11.UOS-2470.


Iris cristata Aiton; 8, 10, 14.UOS-7594.

**Iris pseudacorus L.; 5.UOS-7724.

Iris virginica L.; 7.UOS-1709.

Sisyrinchium albidum Raf.; 9.UOS-2575.

Sisyrinchium angustifolium Mill.; 9, 10.UOS-2569.

Sisyrinchium atlanticum E.P. Bicknell; 3, 7, 15.UOS-1582.

Sisyrinchium sp. nov. ; 13.UOS-8879.


Juncus acuminatus Michx.; 5.UOS-4830.

Juncus anthelatus (Wiegand) R.E. Brooks; 7.UOS-8396.

Juncus canadensis J. Gay ex Laharpe; 3.UOS-4313.

Juncus coriaceus Mackenzie; 7.UOS-7663.

Juncus debilis Gray; 3, 7.UOS-4314.

Juncus diffusissimus Buckl.; 3, 7.UOS-4829.

Juncus effusus L.; 5, 7.UOS-886.

Juncus longii Fern.; 5.UOS-8037.

Juncus marginatus Rostk.; 7.UOS-4315.

Juncus repens Michx.; 5.UOS-7422.

Juncus tenuis Willd.; 1, 2, 3, 4, 7.UOS-4316.

Luzula acuminata Raf. var. carolinae (S. Watson) Fern.; 14.UOS-8203.

Luzula bulbosa (Alph. Wood) Smyth & Smyth; 2, 6.UOS-2857.

Luzula echinata (Small) F.J. Herm.; 1, 2, 4, 7, 9, 14.UOS-7905.

Luzula multiflora (Ehrh.) Lej.; 4, 6, 7.UOS-2947.


Erythronium americanum Ker-Gawl.; 10.UOS-6272.

Lilium canadense L.; 3.UOS-913.

Medeola virginiana L.; 2, 3.UOS-6274.

Prosartes lanuginosa (Michx.) D. Don; 9, 10, 13, 14.UOS-6279.

Prosartes maculata (Buckley) A. Gray; 9, 10.UOS-6344.


Amianthium muscitoxicum (Walt.) Gray; 3.UOS-6331.

Chamaelirium luteum (L.) Gray; 3.UOS-2998.

Melanthium parviflorum (Michx.) S. Watson; 10.UOS-4860.

Stenanthium gramineum (Ker-Gawl.) Morong; 3.UOS-7859.

Trillium cuneatum Raf.; 9, 10, 13, 14.UOS-3166.

Trillium grandiflorum (Michx.) Salisb.; 10, 14.UOS-6276.

Trillium sulcatum Patrick; 10.UOS-6275.


Aletris farinosa L.; 3, 7.UOS-6358.


Aplectrum hyemale (Muhl. ex Willd.) Torr.; 10.UOS-8274.

Corallorhiza odontorhiza (Willd.) Poir. var. odontorhiza; 6.UOS-4757.

Corallorhiza wisteriana Conrad; 6.UOS-1659.

Cypripedium acaule Ait.; 1, 2.UOS-3014. CE.

Cypripedium pubescens Willd.; 10.UOS-8275.

Galearis spectabilis (L.) Raf.; 10.UOS-5672.

Goodyera pubescens (Willd.) R. Br.; 2, 10, 13.UOS-4836.

Isotria verticillata Raf.; 2, 3, 10.UOS-2800.

Liparis liliifolia (L.) Rich. ex Ker Gawl.; 4, 11.UOS-8502.

Malaxis unifolia Michx.; 2, 6.UOS-1706.

Platanthera ciliaris (L.) Lindl.; 7.UOS-4867.

Platanthera clavellata (Michx.) Luer; 3.UOS-7414.

Spiranthes cernua (L.) Rich.; 7.UOS-2112.

Spiranthes lacera (Raf.) Raf. var. gracilis (Bigelow) Luer; 7.UOS-2754.

Spiranthes ovalis Lindl. var. erostellata Catling; 13.UOS-8520.

Spiranthes vernalis Engelm. & Gray; 7.UOS-6327.

Tipularia discolor (Pursh) Nutt.; 2, 11.UOS-4783.


Agrostis perennans (Walt.) Tuckerman; 2, 4, 7.UOS-4319.

**Agrostis stolonifera L.; 4.UOS-8326.

Andropogon gerardii Vitman; 7.UOS-3263.

Andropogon glomeratus (Walt.) B.S.P.; 3.UOS-3318.

Andropogon gyrans Ashe; 7.UOS-7862.

Andropogon ternarius Michx.; 7.UOS-3076.

Andropogon virginicus L.; 4, 7.UOS-7956.

*Anthoxanthum odoratum L.; 4, 6, 7.UOS-3174.

Aristida dichotoma Michx. var. dichotoma; 7.UOS-3264.

**Arthraxon hispidus (Thunb.) Makino; 7.UOS-7011.

Arundinaria appalachiana Triplett, Weakley & L.G. Clark; 2.UOS-8027.

Arundinaria gigantea (Walt.) Muhl.; 11, 12, 13.UOS-7941.

Brachyelytrum erectum (Schreb. ex Spreng.) P. Beauv.; 9, 10, 11, 14.UOS-7015.

*Bromus commutatus Schrad.; 7.UOS-8766.

*Bromus hordeaceus L.; 4.UOS-8280.

Bromus pubescens Muhl. ex Willd.; 1, 2, 9, 10, 11, 13, 14, 15.UOS-4485.

*Bromus racemosus L.; 7.UOS-8035.

Calamagrostis cinnoides (Muhl.) W.P.C.Barton; 5.UOS-7693.

Chasmanthium latifolium (Michx.) Yates; 4, 7.UOS-2694.

Chasmanthium laxum (L.) Yates; 2.UOS-4320.

Chasmanthium sessiliflorum (Poir.) Yates; 2, 4, 13, 15.UOS-8238.

Cinna arundinacea L.; 7.UOS-8362.

*Cynodon dactylon (L.) Pers.; 6.UOS-8617.

*Dactylis glomerata L.; 4, 7.UOS-2017.

Danthonia epilis Scribn.; 1, 7.UOS-8009. S.

Danthonia sericea Nutt.; 1.UOS-8010.

Danthonia spicata (L.) P. Beauv. ex Roem. & Schult.; 1, 2, 4, 7, 9, 11, 12.UOS-3172.

Diarrhena americana P. Beauv.; 13.UOS-8471.

Dichanthelium acuminatum (Sw.) Gould & C.A. Clark ssp. fasciculatum (Torr.) Freckmann & Lelong; 1, 2, 3.UOS- 3253.

Dichanthelium acuminatum (Sw.) Gould & C.A. Clark ssp. lindheimeri (Nash) Gould & C.A. Clark; 1, 2, 3, 4, 7.UOS- 8622.

Dichanthelium boscii (Poir.) Gould & C.A. Clark; 1, 2, 3, 4, 7, 9, 10, 11, 12, 13, 14.UOS-4318.

Dichanthelium clandestinum (L.) Gould; 7.UOS-7651.

Dichanthelium commutatum Schult. ssp. ashei (Pearson ex Ashe) Freckmann & Lelong; 2.UOS-8751.

Dichanthelium commutatum (J.A.Schultes) Gould ssp. commutatum; 2, 4, 7.UOS-7644.

Dichanthelium depauperatum (Muhl.) Gould; 7.UOS-8652.

Dichanthelium dichotomum (L.) Gould ssp. dichotomum; 2.UOS-7534.

Dichanthelium laxiflorum (Lam.) Gould; 1, 4, 7, 11.UOS-7646.

Dichanthelium ovale (Elliot) Gould & C.A. Clark ssp. villosissimum (Nash) Freckmann & Lelong; 9.UOS-8475.

Dichanthelium polyanthes (Schult.) Mohlenbr.; 4, 7.UOS-6616.

Dichanthelium scoparium (Lam.) Gould; 7.UOS-7624.

Digitaria ciliaris (Retz.) Koel.; 7.UOS-6597.

*Digitaria ischaemum (Schreb.) Schreb. ex Muhl.; 7.UOS-3262.

*Echinochloa colona (L.) Link; 15.UOS-8363.

*Eleusine indica (L.) Gaertn.; 6.UOS-2722.

Elymus glabriflorus (Vasey) Scribn. & C.R. Ball; 7.UOS-7849.

Elymus hystrix L.; 2, 9, 11, 12, 13, 14.UOS-1950.

Elymus villosus Muhl. ex Willd.; 2.UOS-8361.

Elymus virginicus L. var. virginicus; 1, 6, 15.UOS-8482.

*Eragrostis cilianensis (All.) Vign. ex Janchen; 4.UOS-7961.

Eragrostis hirsuta (Michx.) Nees; 7.UOS-4079.

Eragrostis spectabilis (Pursh) Steud.; 7.UOS-8038.

Festuca subverticillata (Pers.) Alexeev; 14.UOS-8011.

Glyceria striata (Lam.) Hitchc.; 3, 7.UOS-4907.

*Holcus lanatus L.; 7.UOS-2018.

Hordeum pusillum Nutt.; 7.UOS-8111.

Leersia oryzoides (L.) Sw.; 7.UOS-7662.

Leersia virginica Willd.; 3.UOS-7877.

*Lolium multiflorum Lam.; 7.UOS-3055.

Melica mutica Walt.; 2, 4, 9, 11, 14, 15.UOS-6340.

**Microstegium vimineum (Trin.) A. Camus; 2, 3, 4, 9, 13, 14, 15.UOS-2737.

**Miscanthus sinensis Anderss.; 7.UOS-3327.

Muhlenbergia schreberi J.F. Gmel.; 7.UOS-8039.

Muhlenbergia sobolifera (Muhl. ex Willd.) Trin.; 12, 15.UOS-8308.

Muhlenbergia tenuiflora (Willd.) B.S.P.; 10.UOS-6592.

Panicum anceps Michx. ssp. anceps; 2, 15.UOS-3058.

Panicum flexile (Gattinger) Scribn.; 7, 15.UOS-7830.

Panicum philadelphicum Bernh. ex Trin. ssp. gattingeri (Nash) Freckmann & Lelong; 7.UOS-7854.

Panicum rigidulum Bosc ex Nees ssp. rigidulum; 7.UOS-3265.

Panicum verrucosum Muhl.; 7.UOS-7697.

*Paspalum dilatatum Poir.; 7.UOS-2740.

Paspalum laeve Michx.; 7.UOS-8033.

Paspalum pubiflorum Rupr. ex Fourn.; 7, 15.UOS-8317.

*Phleum pratense L.; 4.UOS-8605.

*Phyllostachys aurea Carrière ex A. Rivière & C. Rivière; 7.UOS-8323.

Piptochaetium avenaceum (L.) Parodi; 2, 11.UOS-4474.

*Poa annua L.; 7.UOS-7656.

Poa autumnalis Muhl. ex Ell.; 4, 10.UOS-8013.

*Poa compressa L.; 1, 4.UOS-8014.

*Poa pratensis L.; 4, 7.UOS-8015.

Poa sylvestris Gray; 7, 13, 14, 15.UOS-6614.

*Poa trivialis L.; 4.UOS-8241.

Saccharum alopecuroides (L.) Nutt.; 7, 15.UOS-3328.

**Schedonorus arundinaceus (Schreb.) Dumort.; 6, 7.UOS-8240.

Schizachyrium scoparium (Michx.) Nash var. scoparium; 7, 15.UOS-8000.

**Setaria faberi Herrm.; 4.UOS-8603.

Setaria parviflora (Poir.) Kerguélen; 4, 7.UOS-7657.

**Setaria pumila (Poir.) Roemer & J.A.Schultes; 4, 7.UOS-930.

**Setaria viridis (L.) P. Beauv. var. viridis; 7.UOS-2799.

Sorghastrum nutans (L.) Nash; 7, 15.UOS-2723.

**Sorghum halepense (L.) Pers.; 7.UOS-2662.

Sphenopholis nitida (Biehler) Scribn.; 4, 7.UOS-8016.

Sporobolus clandestinus (Biehler) Hitchc.; 11.UOS-8357.

Sporobolus compositus (Poir.) Merr. var. drummondii (Trin.) Kartesz & Gandhi; 15.UOS-8345.

*Sporobolus indicus (L.) R. Br.; 7.UOS-7850.

Sporobolus vaginiflorus (Torr. ex Gray) Wood var. vaginiflorus; 7.UOS-7851.

Tridens flavus (L.) Hitchc.; 7, 13, 15.UOS-3254.

*Vulpia myuros (L.) C.C. Gmel.; 4.UOS-8242.

Vulpia octoflora (Walt.) Rydb. var. glauca (Nutt.) Fern.; 9.UOS-8647.


*Eichhornia crassipes (Mart.) Solms; 5.UOS-7743.


Potamogeton diversifolius Raf.; 5.UOS-2699.

Potamogeton nodosus Poir.; 5.UOS-1696.


*Convallaria majalis L. var. majalis; 7.UOS-3452.

Maianthemum racemosum (L.) Link; 2, 3, 10, 13, 14.UOS-1965.

Polygonatum biflorum (Walt.) Ell.; 2, 3, 10, 11, 12, 13.UOS-2248.


Smilax bona-nox L.; 1, 2, 3, 7, 9, 11, 12, 13, 14.UOS-2027.

Smilax glauca Walt.; 1, 2, 9, 13, 14.UOS-2621.

Smilax hugeri (Small) J.B.S. Norton ex Pennell; 10, 11, 13.UOS-4858.

Smilax rotundifolia L.; 1, 2, 3, 10, 11, 13.UOS-630.

Smilax tamnoides L.; 10, 13, 14.UOS-8167.


Sparganium americanum Nutt.; 5.UOS-4827.


Typha latifolia L.; 5.UOS-2671.


Xyris torta Sm.; 3.UOS-4810.



Ruellia caroliniensis (J.F. Gmel.) Steud.; 7, 11, 15.UOS-7921.

Ruellia humilis Nutt.; 6, 15.UOS-4737.


Sambucus canadensis L.; 3, 7, 10, 13.UOS-950.

Viburnum acerifolium L.; 1, 2, 10.UOS-900.

Viburnum alabamense (McAtee) Sorrie; 3.UOS-4498.

*Viburnum carlesii Hemsl.; 7.UOS-8533.

Viburnum cassinoides L.; 2.UOS-7619.

Viburnum nudum L.; 3.UOS-4084.

Viburnum recognitum Fernald; 3.UOS-4811.

*Viburnum rhytidophyllum Hemsl.; 7.UOS-8358.

Viburnum rufidulum Raf.; 2, 9, 11, 12, 13, 14.UOS-3748.


Liquidambar styraciflua L.; 2, 3, 4, 7, 13, 14.UOS-819.


+Amaranthus spinosus L.; 7.UOS-4843.

*Chenopodium album L. var. album; 15.UOS-8344.

*Dysphania ambrosioides (L.) Mosyakin & Clemants; 7.UOS-2026.


Rhus aromatica Ait.; 9, 11, 12, 13.UOS-860.

Rhus copallinum L.; 1, 4, 7.UOS-1948.

Rhus glabra L.; 7, 15.UOS-8515.

Toxicodendron radicans (L.) Kuntze; 1, 2, 3, 4, 7, 8, 9, 10, 11, 12, 13, 14.UOS-4475.

Toxicodendron vernix (L.) Kuntze; 3.UOS-4081.


Chaerophyllum tainturieri Hook.; 7.UOS-2217.

**Conium maculatum L.; 7.UOS-3171.

Cryptotaenia canadensis (L.) DC.; 10.UOS-4505.

**Daucus carota L.; 4, 7.UOS-924.

Erigenia bulbosa (Michx.) Nutt.; 10.UOS-7563.

Eryngium prostratum Nutt. ex DC.; 5.UOS-7417.

Eryngium yuccifolium Michx.; 7, 11.UOS-4911.

*Foeniculum vulgare Mill.; 7.UOS-8886.

Osmorhiza claytonii (Michx.) C.B. Clarke; 10.UOS-3703.

Osmorhiza longistylis (Torr.) DC.; 2, 10, 13.UOS-4798.

Oxypolis rigidior (L.) Raf.; 3.UOS-4075.

Sanicula canadensis L.; 2, 3, 9, 10, 13.UOS-4506.

Sanicula odorata (Raf.) K.M. Pryer & L.R. Phillippe; 10, 13.UOS-4489.

Sanicula smallii E.P. Bicknell; 15.UOS-7809.

Sanicula trifoliata E.P. Bicknell; 10.UOS-7625.

Taenidia integerrima (L.) Drude; 11, 12, 15.UOS-7673.

Thaspium barbinode (Michx.) Nutt.; 9, 10, 11, 13, 14, 15.UOS-6518.

Thaspium chapmanii (Coult. & Rose) Small; 11.UOS-8253.

Thaspium trifoliatum (L.) A. Gray var. trifoliatum; 9.UOS-550.

*Torilis arvensis (Huds.) Link; 7.UOS-2654.

Zizia aptera (Gray) Fern.; 12, 15.UOS-7674.

Zizia aurea (L.) W.D.J. Koch; 7.UOS-1960.


Amsonia tabernaemontana Walt. var. tabernaemontana; 15.UOS-7425.

Apocynum cannabinum L.; 7.UOS-1233.

Asclepias amplexicaulis Sm.; 7.UOS-4799.

Asclepias exaltata L.; 7.UOS-4797.

Asclepias quadrifolia Jacq.; 2, 9.UOS-850.

Asclepias syriaca L.; 4, 7.UOS-912.

Asclepias tuberosa L. ssp. tuberosa; 7, 11.UOS-1491.

Asclepias variegata L.; 2, 7.UOS-896.

Asclepias verticillata L.; 11, 15.UOS-4742.

Gonolobus suberosus auct. non (L.) R. Br.; 14.UOS-8319.

Matelea carolinensis (Jacq.) Woods.; 11, 15.UOS-6247.

**Vinca major L.; 6.UOS-8524.

**Vinca minor L.; 7.UOS-4759.


Ilex ambigua (Michx.) Torr.; 2.UOS-4529.

*Ilex crenata Thunb.; 2.UOS-8516.

Ilex longipes Chapman ex Trel.; 9.UOS-8158.

Ilex montana Torr. & Gray ex Gray; 2.UOS-7971.

Ilex opaca Ait.; 2, 3, 4, 14.UOS-2228.

Ilex verticillata (L.) Gray; 2.UOS-4528.


Aralia racemosa L.; 10.UOS-3978.

Aralia spinosa L.; 7.UOS-2684.

**Hedera helix L.; 7, 15.UOS-4760.

*Hydrocotyle sibthorpioides Lam.; 6.UOS-8321.

Panax quinquefolius L.; 9, 10.UOS-3175. S-CE.


Achillea millefolium L. var. occidentalis DC.; 4, 7.UOS-952.

Ageratina altissima (L.) King & H. Rob. var. altissima; 7, 10, 14, 15.UOS-2047.

Ambrosia artemisiifolia L.; 7.UOS-482.

Ambrosia trifida L.; 7.UOS-2006.

Antennaria plantaginifolia (L.) Richardson; 1, 2.UOS-2326.

Antennaria solitaria Rydb.; 2.UOS-7571.

*Anthemis arvensis L.; 7, 15.UOS-2646.

*Arctium minus Bernh.; 7.UOS-1625.

Arnoglossum atriplicifolium (L.) H. Rob.; 2.UOS-8352.

Arnoglossum reniforme (Hook.) H. Rob.; 10, 14.UOS-8213.

*Artemisia vulgaris L.; 7.UOS-2032.

Bidens aristosa (Michx.) Britton; 7.UOS-2044.

Bidens bipinnata L.; 7.UOS-2691.

Bidens frondosa L.; 7, 15.UOS-2045.

Bidens polylepis S.F. Blake; 15.UOS-6337.

Brickellia eupatorioides (L.) Shinners; 13, 14.UOS-8041.

**Carduus nutans L.; 4, 7.UOS-7939.

Chrysopsis mariana (L.) Ell.; 7.UOS-2176.

**Cichorium intybus L.; 7.UOS-4864.

Cirsium altissimum (L.) Hill; 9, 15.UOS-3982.

Cirsium discolor (Muhl. ex Willd.) Spreng.; 7.UOS-4818.

**Cirsium vulgare (Savi) Ten.; 4, 7.UOS-8348.

Conoclinium coelestinum (L.) DC.; 3, 7.UOS-2033.

Conyza canadensis (L.) Cronquist var. canadensis; 7.UOS-2025.

Conyza canadensis (L.) Cronquist var. pusilla (Nutt.) Cronquist; 6.UOS-6360.

Coreopsis lanceolata L.; 4, 6, 7.UOS-2152.

Coreopsis major Walt.; 1, 2, 12.UOS-951.

Coreopsis tinctoria Nutt.; 7.UOS-3326.

Coreopsis tripteris L.; 15.UOS-4747.

Doellingeria infirma (Michx.) Greene; 9, 10.UOS-8263.

Doellingeria umbellata (Mill.) Nees; 3.UOS-7405.

Elephantopus carolinianus Raeusch.; 4, 10, 13.UOS-3979.

Elephantopus tomentosus L.; 4, 7.UOS-1221.

Erechtites hieraciifolia (L.) Raf. ex DC.; 7.UOS-4086.

Erigeron annuus (L.) Pers.; 3, 7.UOS-2339.

Erigeron philadelphicus L.; 6.UOS-2625.

Erigeron pulchellus Michx.; 6, 7, 9.UOS-2312.

Erigeron strigosus Muhl. ex Willd. var. strigosus; 4, 7.UOS-2405.

Eupatorium album L. var. album; 7.UOS-2159.

Eupatorium altissimum L.; 7.UOS-7691.

Eupatorium capillifolium (Lam.) Small; 7.UOS-7954.

Eupatorium hyssopifolium L.; 7.UOS-1212.

Eupatorium perfoliatum L.; 7.UOS-2029.

Eupatorium pilosum Walt.; 7.UOS-7758.

Eupatorium rotundifolium L. var. ovatum (Bigelow) Torr.; 7.UOS-2157.

Eupatorium rotundifolium L. var. rotundifolium; 7.UOS-2021.

Eupatorium serotinum Michx.; 2, 7.UOS-2022.

Eupatorium sessilifolium L.; 11.UOS-2705.

Eurybia divaricata (L.) Nesom; 10, 15.UOS-7012.

Eurybia hemispherica (Alexander) Nesom; 7, 15.UOS-6590.

Eurybia surculosa (Michx.) Nesom; 1, 7.UOS-2821.

Eutrochium fistulosum (Barratt) Lamont; 7.UOS-2035.

Eutrochium purpureum (L.) Lamont; 7, 9, 10.UOS-4819.

Fleischmannia incarnata (Walter) King & H. Rob.; 12, 15.UOS-8046.

+Gaillardia pulchella Foug.; 7.UOS-7637.

*Galinsoga quadriradiata Cav.; 6, 15.UOS-910.

Gamochaeta argyrinea G. L. Nesom; 7.UOS-7937.

Helenium amarum (Raf.) H. Rock; 7.UOS-8350.

Helenium flexuosum Raf.; 7, 15.UOS-3268.

+Helianthus annuus L.; 7.UOS-8885.

Helianthus divaricatus L.; 7.UOS-3841.

Helianthus eggertii Small; 12.UOS-8291. S.

Helianthus hirsutus Raf.; 12.UOS-8880.

Helianthus microcephalus Torr. & Gray; 7.UOS-2074.

Helianthus sp. nov. ; 15.UOS-8064.

Helianthus strumosus L.; 7.UOS-4541.

Helianthus tuberosus L.; 7.UOS-488.

Heliopsis helianthoides (L.) Sweet; 12, 14, 15.UOS-4753.

Heterotheca camporum (Greene) Shinners var. glandulissima Semple; 7.UOS-4543.

Hieracium gronovii L.; 2, 7.UOS-7960.

Hieracium venosum L.; 2, 7.UOS-7788.

*Hypochaeris radicata L.; 4.UOS-8281.

Ionactis linariifolia (L.) Greene; 1, 2, 7.UOS-4823.

+Iva annua L.; 7.UOS-7720.

Krigia biflora (Walt.) Blake; 4, 7.UOS-4476.

Krigia caespitosa (Raf.) Chambers; 4.UOS-7631.

Krigia dandelion (L.) Nutt.; 6.UOS-7918.

Krigia virginica (L.) Willd.; 1, 7, 9.UOS-4446.

Lactuca canadensis L.; 2, 7.UOS-6335.

Lactuca floridana (L.) Gaertn.; 7, 13.UOS-1947.

*Lactuca saligna L.; 7, 15.UOS-4544.

*Lapsana communis L.; 3.UOS-4801.

**Leucanthemum vulgare Lam.; 7.UOS-2255.

Liatris cylindracea Michx.; 15.UOS-4732. T.

Liatris microcephala (Small) K. Schum.; 1.UOS-2039.

Liatris scariosa (L.) Willd.; 1.UOS-4820.

Liatris squarrulosa Michx.; 4, 7.UOS-2051.

Mikania scandens (L.) Willd.; 7.UOS-8360.

Packera anonyma (Alph. Wood) W.A. Weber & A. Löve; 1, 7.UOS-855.

Packera glabella (Poir.) C. Jeffrey; 7.UOS-2189.

Packera obovata (Muhl. ex Willd.) W.A. Weber & A. Löve; 15.UOS-3528.

Parthenium integrifolium L. var. integrifolium; 7.UOS-2656.

Pityopsis aspera (Shuttlw. ex Small) Small var. adenolepis (Fernald) Semple & F.D. Bowers; 7.UOS-3732.

Pityopsis graminifolia (Michx.) Nutt. var. latifolia (Fernald) Semple & F.D. Bowers; 7.UOS-7767.

Pluchea camphorata (L.) DC.; 3.UOS-2181.

Polymnia canadensis L.; 7, 9, 13.UOS-949.

Polymnia laevigata Beadle; 7, 9, 10, 11, 13.UOS-7016.

Prenanthes altissima L.; 10.UOS-4825.

Prenanthes serpentaria Pursh; 2, 7, 13.UOS-4083.

Pseudognaphalium obtusifolium (L.) Hilliard & Burtt; 7.UOS-2030.

Pyrrhopappus carolinianus (Walt.) DC.; 7, 15.UOS-8267.

Ratibida pinnata (Vent.) Barnh.; 7, 15.UOS-4734.

Rudbeckia fulgida Aiton var. fulgida; 12.UOS-8264.

Rudbeckia hirta L. var. hirta; 7.UOS-2072.

Rudbeckia hirta L. var. pulcherrima Farw.; 4.UOS-3682.

Rudbeckia laciniata L. var. laciniata; 14.UOS-8183.

Rudbeckia triloba L. var. triloba; 7.UOS-653.

*Senecio vulgaris L.; 7.UOS-8531.

Sericocarpus asteroides (L.) Britton, Sterns & Poggenb.; 7.UOS-8266.

Sericocarpus linifolius (L.) B.S.P.; 7.UOS-1208.

Silphium asteriscus L. var. astericus; 7.UOS-2665.

Silphium asteriscus L. var. dentatum; 7.UOS-8660.

Silphium brachiatum Gattinger; 12.UOS-8176. E.

Silphium pinnatifidum Ell.; 15.UOS-4736. T.

Smallanthus uvedalia (L.) Mack. Ex Small; 9, 15.UOS-4743.

Solidago altissima L. ssp. altissima; 4, 7.UOS-7955.

Solidago arguta Aiton var. caroliniana A. Gray; 7.UOS-6582.

Solidago auriculata Shuttlw. ex Blake; 11, 12, 15.UOS-8047.

Solidago caesia L.; 2, 10, 12, 13.UOS-4815.

Solidago curtisii Torr. & A. Gray var. flaccidifolia (Small) R.E.Cook & Semple; 10, 15.UOS-8628.

Solidago erecta Pursh; 7.UOS-7632.

Solidago flexicaulis L.; 9, 10.UOS-7804.

Solidago hispida Muhl. ex Willd.; 7.UOS-8301.

Solidago nemoralis Ait.; 7.UOS-1219.

Solidago odora Ait.; 2.UOS-4816.

Solidago patula Muhl. ex Willd.; 7.UOS-7721.

Solidago rigida L. ssp. glabrata (E.L. Braun) Heard & Semple; 12, 15.UOS-8484.

Solidago rugosa Mill. aspera (Aiton) Cronquist ssp. aspera (Aiton) Cronquist; 7.UOS-6764.

Solidago speciosa Nutt. speciosa ssp. speciosa; 10.UOS-4817.

Solidago ulmifolia Muhl. ex Willd.; 9, 10, 11, 12.UOS-7728.

*Sonchus asper (L.) Hill; 7.UOS-2645.

Symphyotrichum concolor (L.) Nesom; 1.UOS-6520.

Symphyotrichum cordifolium (L.) Nesom; 10, 12.UOS-1225.

Symphyotrichum dumosum (L.) Nesom; 7.UOS-6523.

Symphyotrichum laeve (L.) A. Löve & D. Löve var. laeve; 11.UOS-8356.

Symphyotrichum lanceolatum (Willd.) Nesom var. latifolium (Semple & Chmielewski) G.L. Nesom; 7.UOS-7699.

Symphyotrichum lateriflorum (L.) A. Löve & D. Löve; 3, 14.UOS-6763.

Symphyotrichum ontarionis (Wieg.) Nesom; 7.UOS-7698.

Symphyotrichum patens (Ait.) Nesom var. patens; 7, 11, 13.UOS-1211.

Symphyotrichum pilosum (Willd.) Nesom; 7.UOS-1226.

Symphyotrichum pratense (Raf.) Nesom; 15.UOS-7993. E.

Symphyotrichum shortii (Lindl.) Nesom; 9, 12, 15.UOS-4824.

Symphyotrichum undulatum (L.) Nesom; 7.UOS-7783.

*Tanacetum vulgare L.; 7.UOS-7813.

*Taraxacum officinale F.H. Wigg.; 6, 7.UOS-3966.

**Tragopogon dubius Scop.; 7.UOS-8468.

Verbesina alternifolia (L.) Britton ex Kearney; 15.UOS-7687.

Verbesina virginica L.; 7, 11, 15.UOS-3981.

Vernonia flaccidifolia Small; 11, 12, 15.UOS-8157.

Vernonia gigantea (Walter) Trel.; 7.UOS-2034.

Xanthium strumarium L.; 7.UOS-8756.


Impatiens capensis Meerb.; 3, 7.UOS-2048.

Impatiens pallida Nutt.; 10, 15.UOS-7930.


**Berberis bealei Fortune; 7.UOS-2729.

Berberis canadensis Mill.; 11.UOS-8656. S.

**Berberis thunbergii DC.; 2, 7.UOS-6270.

Caulophyllum thalictroides (L.) Michx.; 10, 13.UOS-3190.

Jeffersonia diphylla (L.) Pers.; 13.UOS-8185.

Podophyllum peltatum L.; 9, 10, 14.UOS-2345.


Alnus serrulata (Ait.) Willd.; 5.UOS-2179.

Betula nigra L.; 7.UOS-4077.

Carpinus caroliniana Walt.; 13, 14.UOS-8168.

Corylus americana Walt.; 10.UOS-2184.

Ostrya virginiana (Mill.) K. Koch; 11, 12, 13.UOS-2706.


Bignonia capreolata L.; 9, 11, 12, 13, 14.UOS-2702.

Campsis radicans (L.) Seem. ex Bureau; 7, 9, 12.UOS-1681.

+Catalpa speciosa (Warder) Warder ex Engelm.; 4, 7.UOS-8467.


**Buglossoides arvensis (L.) I.M. Johnst.; 6.UOS-8476.

Cynoglossum virginianum L.; 4, 9, 10, 12, 13, 14.UOS-2474.

Hydrophyllum appendiculatum Michx.; 10.UOS-2631.

Hydrophyllum canadense L.; 7, 10.UOS-874.

Hydrophyllum macrophyllum Nutt.; 10.UOS-4805.

Lithospermum canescens (Michx.) Lehm.; 9, 10.UOS-8174.

Lithospermum tuberosum Rugel ex DC.; 11, 12, 13.UOS-4452.

Mertensia virginica (L.) Pers. ex Link; 14.UOS-8099.

Myosotis macrosperma Engelm.; 7, 9.UOS-2359.

**Myosotis scorpioides L.; 7.UOS-2361.

Nemophila aphylla (L.) Brummitt; 7, 13.UOS-8100.

Phacelia bipinnatifida Michx.; 10, 14.UOS-1637.


**Alliaria petiolata (Bieb.) Cavara & Grande; 7, 15.UOS-4443.

*Arabidopsis thaliana (L.) Heynh.; 6.UOS-2394.

*Barbarea verna (Mill.) Asch.; 7.UOS-2422.

*Barbarea vulgaris W.T. Aiton; 7.UOS-7893.

Boechera canadensis (L.) Al-Shehbaz; 11.UOS-2703.

Boechera laevigata (Muhl. ex Willd.) Al-Shehbaz; 11, 12.UOS-7598.

*Brassica rapa L.; 7, 15.UOS-8526.

*Capsella bursa-pastoris (L.) Medik.; 7.UOS-2373.

*Cardamine hirsuta L.; 3, 6, 7, 8.UOS-2463.

Cardamine parviflora L. var. arenicola (Britton) O.E. Schulz; 9.UOS-6346.

Cardamine pensylvanica Muhl. ex Willd.; 14.UOS-7601.

Dentaria diphylla Michx.; 9, 10, 14.UOS-3973.

Dentaria laciniata Muhl. ex Willd.; 10, 12.UOS-5668.

Dentaria multifida Muhl. ex Ell.; 10.UOS-1568.

*Draba verna L.; 6, 7.UOS-2461.

**Hesperis matronalis L.; 2.UOS-8169.

**Lepidium campestre (L.) W.T. Aiton; 7.UOS-7891.

Lepidium virginicum L.; 7.UOS-943.

*Lunaria annua L.; 7, 15.UOS-7863.

**Rorippa nasturtium-aquaticum (L.) Hayek; 7.UOS-2638.

Sibara virginica (L.) Rollins; 6.UOS-3951.

*Sisymbrium officinale (L.) Scop.; 7, 15.UOS-947.

*Thlaspi arvense L.; 7.UOS-8249.


Opuntia humifusa (Raf.) Raf.; 1.UOS-3278.


Campanula americana L.; 13, 15.UOS-4764.

Lobelia cardinalis L.; 3.UOS-1528.

Lobelia inflata L.; 7, 15.UOS-2070.

Lobelia puberula Michx.; 7.UOS-2010.

Lobelia spicata Lam.; 4, 7, 15.UOS-6336.

Triodanis perfoliata (L.) Nieuwl. var. perfoliata; 4, 7.UOS-878.


Celtis laevigata Willd.; 7, 10.UOS-2046.

Celtis occidentalis L.; 2, 9, 13.UOS-8721.

Celtis tenuifolia Nutt.; 11, 13.UOS-8210.


Lonicera dioica L.; 11.UOS-8712. S.

**Lonicera fragrantissima Lindl. & Paxton; 7.UOS-4451.

**Lonicera japonica Thunb.; 3, 7, 13, 14.UOS-871.

**Lonicera maackii (Rupr.) Herder; 4.UOS-8320.

**Lonicera morrowii Gray; 7.UOS-3180.

Lonicera sempervirens L.; 7, 10, 15.UOS-925.

Symphoricarpos orbiculatus Moench; 7, 11, 13, 14.UOS-4869.


*Arenaria serpyllifolia L. var. tenuior Mert. & W.D.J. Koch; 7.UOS-2918.

*Arenaria serpyllifolia L. var. serpyllifolia; 4.UOS-7928.

Cerastium brachypodum (Engelm. ex A. Gray) B.L. Rob.; 7.UOS-4065.

*Cerastium fontanum Baumg. ssp. vulgare (Hartman) Greuter & Burdet; 4, 6, 7.UOS-4069.

*Cerastium glomeratum Thuill.; 7.UOS-2634.

*Cerastium semidecandrum L.; 7.UOS-4071.

*Dianthus armeria L.; 4.UOS-956.

Minuartia glabra (Michx.) Mattf.; 1.UOS-2978.

*Saponaria officinalis L.; 4, 7.UOS-4080.

Silene antirrhina L.; 7.UOS-4104.

Silene rotundifolia Nutt.; 8.UOS-870.

Silene stellata (L.) W.T. Aiton; 2, 9.UOS-3179.

Silene virginica L. var. virginica; 2.UOS-3782.

*Silene vulgaris (Moench) Garcke; 7.UOS-8040.

*Stellaria media (L.) Vill.; 7, 10.UOS-2460.

Stellaria pubera Michx.; 8, 9, 10, 13, 14, 15.UOS-4108.


**Celastrus orbiculatus Thunb.; 7.UOS-2028.

**Euonymus alatus (Thunb.) Siebold; 2, 10.UOS-6338.

Euonymus americanus L.; 3, 7, 13.UOS-2627.

Euonymus atropurpureus Jacq.; 10, 14, 15.UOS-8314.

**Euonymus hederaceus Champ. & Benth.; 7, 10.UOS-7426.


Lechea racemulosa Michx.; 7.UOS-7988.


Calystegia catesbeiana Pursh; 7.UOS-3794.

Calystegia silvatica (Kit.) Griseb. ssp. fraterniflora (Mack. & Bush) Brummitt; 15.UOS-1371.

Cuscuta pentagona Engelm.; 15.UOS-8316.

+Dichondra carolinensis Michx.; 6.UOS-8519.

*Ipomoea hederacea Jacq.; 7.UOS-2676.

Ipomoea pandurata (L.) G. Mey.; 2.UOS-2689.

*Ipomoea purpurea (L.) Roth; 7.UOS-2752.


Cornus alternifolia L. f.; 14.UOS-8165.

Cornus amomum Mill.; 3.UOS-5769.

Cornus florida L.; 2, 4, 9, 12, 13, 14.UOS-3763.


Diamorpha smallii Britton ex Small; 1.UOS-2985. E.

*Sedum sarmentosum Bunge; 1.UOS-2640.

Sedum ternatum Michx.; 8, 10, 14.UOS-3815.


Melothria pendula L.; 7.UOS-2186.

Sicyos angulatus L.; 15.UOS-8303.


Diospyros virginiana L.; 1, 2, 4, 7, 9, 11, 12, 13.UOS-2199.


**Elaeagnus umbellata Thunb. var. parviflora (Wall. ex Royle) C.K. Schneid.; 7.UOS-6325.


Chimaphila maculata (L.) Pursh; 1, 2, 3, 9, 13, 14.UOS-818.

Epigaea repens L.; 1, 2.UOS-2479.

Kalmia latifolia L.; 1, 2.UOS-1682.

Lyonia ligustrina (L.) DC. var. ligustrina; 3.UOS-8436.

Monotropa hypopithys L.; 2.UOS-6329.

Monotropa uniflora L.; 10.UOS-4839.

Oxydendrum arboreum (L.) DC.; 1, 2, 3, 4, 9, 14.UOS-3449.

Rhododendron canescens (Michx.) Sweet; 3.UOS-4223.

Rhododendron maximum L.; 8.UOS-4535.

Vaccinium arboreum Marsh.; 1, 2.UOS-865.

Vaccinium corymbosum L.; 1, 2, 3, 4, 9, 11.UOS-1423.

Vaccinium fuscatum Ait.; 3.UOS-8251.

Vaccinium pallidum Ait.; 1, 2, 3.UOS-4237.

Vaccinium stamineum L.; 1, 2, 4.UOS-813.


Acalypha ostryifolia Riddell; 6.UOS-8487.

Acalypha rhomboidea Raf.; 7.UOS-7739.

Chamaesyce nutans (Lag.) Small; 7.UOS-1963.

+Chamaesyce prostrata (Ait.) Small; 7.UOS-2721.

Croton monanthogynus Michx.; 7.UOS-2930.

Croton willdenowii G.L. Webster; 1.UOS-4847.

Euphorbia corollata L.; 7, 9.UOS-1207.

Euphorbia cyathophora Murr.; 7.UOS-4756.

*Euphorbia dentata Michx.; 7, 15.UOS-7738.

*Euphorbia helioscopia L.; 7, 15.UOS-8105.

Euphorbia mercurialina Michx.; 10, 13.UOS-4487.


**Albizia julibrissin Durazz.; 7.UOS-1962.

Amorpha fruticosa L.; 7.UOS-4483.

Amphicarpaeabracteata (L.) Fern.; 2, 9, 10, 11, 14.UOS-2077.

Apios americana Medik.; 7.UOS-6246.

Cercis canadensis L.; 9, 10, 11, 12, 13.UOS-2669.

Chamaecrista fasciculata (Michx.) Greene; 7.UOS-7014.

Chamaecrista nictitans (L.) Moench; 7, 11, 14, 15.UOS-4539.

Cladrastis kentukea (Dum.-Cours.) Rudd; 10.UOS-4467.

Clitoria mariana L.; 2, 11.UOS-1733.

Desmodium cuspidatum (Muhl. ex Willd.) DC. ex Loud.; 15.UOS-4750.

Desmodium glabellum (Michx.) DC.; 7, 13.UOS-7737.

Desmodium glutinosum (Muhl. ex Willd.) Wood; 7, 13.UOS-4808.

Desmodium nudiflorum (L.) DC.; 2.UOS-481.

Desmodium paniculatum (L.) DC.; 7, 11, 12.UOS-7736.

Desmodium pauciflorum (Nutt.) DC.; 7, 10, 12, 14.UOS-7415.

Desmodium perplexum Schub.; 7.UOS-7013.

Desmodium rotundifolium DC.; 4, 7, 11.UOS-2049.

Desmodium viridiflorum (L.) DC.; 7.UOS-8255.

Galactia volubilis (L.) Britton; 2, 11, 13.UOS-8395.

Gleditsia triacanthos L.; 7, 9, 11, 14, 15.UOS-2660.

Gymnocladus dioicus (L.) K. Koch; 9.UOS-8724.

**Kummerowia striata (Thunb.) Schindl.; 15.UOS-8354.

*Lathyrus hirsutus L.; 7.UOS-8469.

*Lathyrus latifolius L.; 7.UOS-894.

**Lespedeza bicolor Turcz.; 7.UOS-465.

**Lespedeza cuneata (Dum. Cours.) G. Don; 3, 4, 7.UOS-2076.

Lespedeza hirta (L.) Hornem.; 4.UOS-2050.

Lespedeza intermedia (S. Watson) Britton; 7.UOS-8523.

Lespedeza procumbens Michx.; 2, 11, 13, 15.UOS-4769.

Lespedeza repens (L.) W. Bartram; 7.UOS-4795.

Lespedeza violacea (L.) Pers.; 2, 7.UOS-6758.

Lespedeza virginica (L.) Britton; 11, 13.UOS-8472.

*Medicago lupulina L.; 4, 7.UOS-2559.

**Melilotus alba Medikus; 7.UOS-1957.

**Melilotus officinalis (L.) Lam.; 7.UOS-1958.

Mimosa microphylla Dry.; 7.UOS-954.

Orbexilum pedunculatum (Mill.) Rydb.; 3.UOS-8439.

Phaseolus polystachios (L.) B.S.P.; 13.UOS-8610.

**Pueraria montana (Lour.) Merr. var. lobata; 7, 15.UOS-1213.

Robinia hispida L. var. hispida; 7.UOS-4916.

Robinia hispida L. var. rosea Pursh; 4, 7.UOS-2489.

Robinia pseudoacacia L.; 1, 2, 4, 7, 9, 10, 12.UOS-2464.

*Securigera varia (L.) Lassen; 7.UOS-2052.

Senna marilandica (L.) Link; 3.UOS-7404.

Stylosanthes biflora (L.) B.S.P.; 4, 7.UOS-2173.

Tephrosia virginiana (L.) Pers.; 7.UOS-1700.

Thermopsis mollis (Michx.) M.A. Curtis ex A. Gray; 2.UOS-2473. S.

*Trifolium campestre Schreb.; 4, 6, 7.UOS-2560.

*Trifolium dubium Sibth.; 7.UOS-2558.

*Trifolium incarnatum L.; 4.UOS-2556.

*Trifolium pratense L.; 4, 6, 7.UOS-884.

*Trifolium repens L.; 6.UOS-2841.

Vicia caroliniana Walt.; 12, 15.UOS-6347.

*Vicia sativa L. ssp. nigra (L.) Ehrh.; 4, 6, 7.UOS-1686.

*Vicia villosa Roth ssp. varia (Host) Corb.; 6, 7, 12.UOS-6250.

**Wisteria sinensis (Sims) DC.; 7.UOS-3564.


Castanea dentata (Marsh.) Borkh.; 2.UOS-2732.

Fagus grandifolia Ehrh.; 2, 13, 14.UOS-2733.

Quercus alba L.; 1, 2, 3, 4, 9, 12, 13, 14.UOS-1152.

Quercus coccinea Münchh.; 1, 2, 3, 4, 7, 9, 12.UOS-7990.

Quercus falcata Michx.; 1, 2, 4, 9.UOS-7900.

Quercus marilandica Münchh.; 1.UOS-3002.

Quercus montana Willd.; 1, 2.UOS-1475.

Quercus muehlenbergii Engelm.; 4, 9, 10, 11, 12, 13, 14.UOS-7608.

+Quercus phellos L.; 7.UOS-7970.

Quercus rubra L.; 9, 10, 11.UOS-7934.

Quercus shumardii Buckl.; 11, 14.UOS-8311.

Quercus stellata Wangenh.; 2, 11.UOS-3601.

Quercus velutina Lam.; 1, 2, 4, 9.UOS-822.


Bartonia virginica (L.) B.S.P.; 3.UOS-6266.

Frasera caroliniensis Walt.; 9, 15.UOS-8260.

Gentiana saponaria L.; 3.UOS-3324.

Gentiana villosa L.; 4, 15.UOS-8335.

Obolaria virginica L.; 9, 10, 11.UOS-1629.

Sabatia angularis (L.) Pursh; 15.UOS-8257.


Geranium carolinianum L.; 4, 6, 7.UOS-2446.

*Geranium dissectum L.; 7.UOS-8248.

Geranium maculatum L.; 9, 10, 12.UOS-1.


**Myriophyllum aquaticum (Vell.) Verdc.; 5.UOS-2718.

Proserpinaca palustris L. var. crebra Fern. & Grisc.; 5.UOS-2697.


Hamamelis virginiana L.; 2.UOS-2709.


*Deutzia scabra Thunb.; 7.UOS-8635.

Hydrangea arborescens L.; 10.UOS-8534.

Hydrangea cinerea Small; 10, 11, 15.UOS-7897.

Philadelphus hirsutus Nutt.; 11.UOS-6352.

+Philadelphus inodorus L.; 7.UOS-705.


Hypericum canadense L.; 7.UOS-8892.

Hypericum crux-andreae (L.) Crantz; 2, 7.UOS-2756.

Hypericum frondosum Michx.; 11.UOS-7727.

Hypericum gentianoides (L.) B.S.P.; 1, 4.UOS-1144.

Hypericum hypericoides (L.) Crantz; 7.UOS-8279.

Hypericum mutilum L.; 7.UOS-2759.

Hypericum nudiflorum Michx. ex Willd.; 7.UOS-6332.

Hypericum prolificum L.; 7.UOS-1949.

Hypericum punctatum Lam.; 1, 15.UOS-8225.

Hypericum sphaerocarpum Michx.; 15.UOS-6362.

Hypericum stragulum P. Adams & N. Robson; 2, 4, 7.UOS-1169.


Itea virginica L.; 3.UOS-6354.


Carya carolinae-septentrionalis (Ashe) Engl. & Graebn.; 10, 11, 12.UOS-8336.

Carya cordiformis (Wangenh.) K. Koch; 14.UOS-8315.

Carya glabra (Mill.) Sweet; 2, 9, 12, 13.UOS-8318.

Carya ovalis (Wangenh.) Sarg.; 9.UOS-8890.

Carya ovata (Mill.) K. Koch; 9, 10, 13.UOS-8312.

Carya pallida (Ashe) Engl. & Graebn.; 1, 2, 9.UOS-8024.

Carya tomentosa (Lam. ex Poir.) Nutt.; 1, 2.UOS-8604.

Juglans cinerea L.; 14.UOS-8237. T.

Juglans nigra L.; 4, 9, 10, 13, 14.UOS-4781.


Agastache nepetoides (L.) Kuntze; 14, 15.UOS-8347.

*Ajuga reptans L.; 7.UOS-7819.

Blephilia hirsuta (Pursh) Benth.; 13, 14.UOS-8211.

*Calamintha nepeta (L.) Savi; 4.UOS-7957.

Callicarpa americana L.; 9, 12.UOS-3983.

Collinsonia canadensis L.; 7.UOS-6587.

**Glechoma hederacea L.; 6, 7.UOS-2905.

Hedeoma pulegioides (L.) Pers.; 9, 15.UOS-8307.

*Lamiastrum galeobdolon (L.) Ehrend. & Polatschek; 7.UOS-8535.

*Lamium amplexicaule L.; 6.UOS-2612.

*Lamium purpureum L.; 6, 7.UOS-2616.

Lycopus virginicus L.; 4, 7.UOS-2175.

**Mentha piperita L. (pro sp.); 7.UOS-6359.

*Mentha rotundifolia (L.) Huds. (pro sp.); 7.UOS-3177.

Monarda fistulosa L.; 7, 11.UOS-4803.

*Mosla dianthera (Buch.-Ham. ex Roxb.) Maxim.; 4, 7, 15.UOS-4087.

*Perilla frutescens (L.) Britton; 7.UOS-2020.

Physostegia virginiana (L.) Benth. ssp. praemorsa (Shinners) Cantino; 15.UOS-7407.

+Prunella vulgaris L.; 6, 7, 15.UOS-946.

Pycnanthemum loomisii Nutt.; 2, 4, 15.UOS-8306.

Pycnanthemum muticum (Michx.) Pers.; 7.UOS-8096.

Pycnanthemum tenuifolium Schrad.; 9.UOS-8262.

Salvia lyrata L.; 6, 11, 13.UOS-1442.

*Salvia pratensis L.; 7.UOS-8538.

Salvia urticifolia L.; 9, 12.UOS-8126.

Scutellaria elliptica Muhl. ex Spreng. var. hirsuta (Short & Peter) Fern.; 2, 4, 11.UOS-2714.

Scutellaria integrifolia L.; 1.UOS-4891.

Scutellaria ovata Hill; 7, 9, 15.UOS-8387.

Scutellaria pseudoserrata Epling; 9, 10, 15.UOS-4491.

Trichostema dichotomum L.; 15.UOS-3271.

Trichostema setaceum Houtt.; 7.UOS-6261.


Utricularia gibba L.; 5.UOS-7705.


Linum medium (Planch.) Britton var. texanum (Planch.) Fernald; 7.UOS-8884.

Linum striatum Walt.; 7.UOS-2716.


Lindernia dubia (L.) Pennell var. anagallidea (Michx.) Cooperr.; 5.UOS-7630.


*Abelia grandiflora (Rovelli ex André) Rehd. [chinensis × uniflora]; 7.UOS-8494.


Spigelia marilandica (L.) L.; 8, 9, 11.UOS-6348.


Cuphea viscosissima Jacq.; 15.UOS-8304.


**Hibiscus syriacus L.; 7, 14.UOS-1853.

Tilia americana L. var. americana; 9, 13.UOS-462.

Tilia americana L. var. heterophylla (Vent.) Loud.; 10, 13.UOS-7903.


Rhexia mariana L. var. mariana; 7.UOS-2667.

Rhexia virginica L.; 3.UOS-2757.


Cocculus carolinus (L.) DC.; 12, 15.UOS-7991.

Menispermum canadense L.; 9, 10, 11, 12, 14.UOS-8332.


*Nymphoides peltata (Gmel.) Kuntze; 5.UOS-8373.


Mollugo verticillata L.; 7.UOS-7976.


Claytonia caroliniana Michx.; 10.UOS-4440.

Claytonia virginica L.; 6, 10, 14.UOS-1449.

Phemeranthus teretifolius (Pursh) Raf.; 1.UOS-7627. T.


**Broussonetia papyrifera (L.) L'Hér. ex Vent.; 7.UOS-8749.

*Fatoua villosa (Thunb.) Nakai; 6.UOS-8390.

*Morus alba L.; 7.UOS-1030.

Morus rubra L.; 10.UOS-8222.


*Anagallis arvensis L.; 7.UOS-8752.

*Lysimachia clethroides Duby; 7.UOS-5722.

**Lysimachia nummularia L.; 3, 7.UOS-2147.

Lysimachia quadrifolia L.; 2, 3.UOS-864.


Nyssa sylvatica Marsh. var. sylvatica; 1, 2, 3, 4, 9, 10, 14.UOS-4472.


Chionanthus virginicus L.; 9, 12.UOS-3880.

Forestiera ligustrina (Michx.) Poir.; 11.UOS-8196.

*Forsythia viridissima Lindl.; 7.UOS-8525.

Fraxinus biltmoreana Beadle; 2, 4, 9, 10, 11, 12, 13, 14.UOS-7021.

Fraxinus quadrangulata Michx.; 9, 10, 11, 14.UOS-6353.

**Ligustrum obtusifolium Sieb. & Zucc.; 7, 15.UOS-7816.

**Ligustrum sinense Lour.; 3, 4, 7, 15.UOS-893.


Circaea lutetiana L. ssp. canadensis (L.) Asch. & Magnus; 3, 10, 11, 13.UOS-2712.

Epilobium coloratum Biehler; 7, 10.UOS-3985.

Gaura filipes Spach; 15.UOS-4739.

Ludwigia alternifolia L.; 7, 15.UOS-2037.

Ludwigia palustris (L.) Ell.; 5.UOS-2698.

Oenothera biennis L.; 7, 15.UOS-496.

Oenothera laciniata Hill; 7.UOS-2644.

+Oenothera speciosa Nutt.; 7.UOS-1455.


Agalinis gattingeri (Small) Small; 7.UOS-1209.

Agalinis purpurea (L.) Pennell; 4, 7.UOS-2724.

Agalinis tenuifolia (Vahl) Raf.; 4, 7.UOS-4714.

Aureolaria pectinata (Nutt.) Pennell; 7.UOS-4542.

Aureolaria virginica (L.) Pennell; 7, 11, 15.UOS-944.

Conopholis americana (L.) Wallr.; 2, 9, 10, 11, 12.UOS-1626.

Epifagus virginiana (L.) W. Bartram; 13, 14.UOS-8195.

Melampyrum lineare Desr. var. latifolium Bart.; 2.UOS-7894.

Orobanche uniflora L.; 9.UOS-4448.

Pedicularis canadensis L.; 10, 14, 15.UOS-3787.


+Oxalis corniculata L.; 6.UOS-8503.

Oxalis dillenii Jacq.; 4, 6, 7, 13.UOS-7904.

Oxalis illinoensis Schwegm.; 10, 13.UOS-4449.

Oxalis violacea L.; 2, 9, 10, 11, 12.UOS-4442.


Dicentra cucullaria (L.) Bernh.; 10.UOS-2306.

Sanguinaria canadensis L.; 10, 13, 14.UOS-7564.

Stylophorum diphyllum (Michx.) Nutt.; 10.UOS-7573.


Parnassia asarifolia Vent.; 3.UOS-2738.


Passiflora incarnata L.; 4.UOS-492.

Passiflora lutea L.; 10, 13.UOS-4032.


**Paulownia tomentosa (Thunb.) Sieb. & Zucc. ex Steud.; 7, 9.UOS-3719.


Penthorum sedoides L.; 3.UOS-7421.


*Mazus pumilus (Burm. f.) Steenis; 6, 7.UOS-2884.

Mimulus alatus Ait.; 15.UOS-1210.

Mimulus ringens L.; 3.UOS-1997.

Phryma leptostachya L.; 9, 13.UOS-3183.


Phyllanthus caroliniensis Walt.; 7.UOS-7986.


Phytolacca americana L.; 1, 2, 7.UOS-942.


Callitriche heterophylla Pursh; 5.UOS-8470.

*Chaenorhinum minus (L.) Lange; 15.UOS-8539.

Chelone glabra L.; 3.UOS-4082.

*Cymbalaria muralis G. Gaertn., B. Mey. & Scherb.; 6.UOS-8891.

Gratiola neglecta Torr.; 3.UOS-7672.

Gratiola virginiana L.; 3.UOS-4828.

Mecardonia acuminata (Walt.) Small; 7.UOS-7639.

Penstemon brevisepalus Pennell; 7.UOS-8726.

Penstemon calycosus Small; 15.UOS-4800.

Penstemon canescens (Britton) Britton; 2, 7, 15.UOS-7856.

Penstemon laevigatus Aiton; 7.UOS-873.

Plantago aristata Michx.; 4, 7.UOS-1995.

*Plantago lanceolata L.; 4, 6.UOS-888.

Plantago rugelii Decne.; 6, 15.UOS-2949.

Plantago virginica L.; 7.UOS-2406.

*Veronica arvensis L.; 4, 6, 7.UOS-7906.

*Veronica officinalis L.; 4, 7.UOS-3628.

*Veronica persica Poir.; 6, 7.UOS-2462.

*Veronica serpyllifolia L.; 6, 7.UOS-3627.

Veronicastrum virginicum (L.) Farw.; 14, 15.UOS-1500.


Platanus occidentalis L.; 10.UOS-417.


Phlox amoena Sims; 2.UOS-2471.

Phlox amplifolia Britton; 15.UOS-8209.

Phlox carolina L.; 15.UOS-8252.

Phlox divaricata L. var. laphamii Wood; 9.UOS-2340.

Phlox divaricata L. var. divaricata; 10.UOS-727.

Phlox maculata L. ssp. pyramidalis (Sm.) Wherry; 7.UOS-2686.

Phlox maculata L. ssp. maculata; 7.UOS-493.

Phlox pilosa L. ssp. pilosa; 10.UOS-8109.

+Phlox subulata L.; 7.UOS-8532.


Polygala curtissii Gray; 7.UOS-2174.

Polygala senega L. var. senega; 9.UOS-557.

Polygala senega L. var. latifolia Torr. & A. Gray; 11.UOS-8359.

Polygala verticillata L.; 4.UOS-4812.


**Fallopia japonica (Houtt.) Ronse Decr.; 4.UOS-4716.

Fallopia scandens (L.) Holub; 7.UOS-2060.

**Persicaria longiseta (Bruijn) Kitag.; 7, 10, 15.UOS-6595.

**Persicaria maculosa Gray; 7, 9.UOS-2069.

Persicaria pensylvanica (L.) Small; 3, 7.UOS-7987.

Persicaria punctata (Ell.) Small; 7, 9.UOS-6361.

Persicaria sagittata (L.) Gross.; 7.UOS-1697.

Persicaria virginiana (L.) Gaertn.; 10, 13, 14.UOS-4731.

*Polygonum aviculare L.; 7.UOS-8750.

*Rumex acetosella L.; 4, 6.UOS-6141.

*Rumex conglomeratus Murr.; 7.UOS-4512.

*Rumex crispus L.; 7.UOS-1955.

*Rumex obtusifolius L.; 7.UOS-1964.


Primula meadia (L.) A.R.Mast & Reveal; 13, 15.UOS-1677.


Actaea pachypoda Ell.; 10, 13.UOS-936.

Anemone acutiloba (DC.) G. Lawson; 9, 10, 13, 14.UOS-396.

Anemone quinquefolia L. var. quinquefolia; 3.UOS-7622.

Anemone virginiana L.; 11, 15.UOS-8045.

Clematis catesbyana Pursh; 15.UOS-8305.

Clematis morefieldii Kral; 12.UOS-6248. LE.

**Clematis terniflora DC.; 7.UOS-2750.

Clematis virginiana L.; 3, 7, 10, 12.UOS-1746.

Delphinium tricorne Michx.; 9, 10.UOS-2283.

Hydrastis canadensis L.; 9, 14.UOS-7583. CE.

Ranunculus abortivus L.; 2, 3, 4, 6, 7, 9, 10.UOS-1172.

**Ranunculus bulbosus L.; 6, 7.UOS-8657.

Ranunculus hispidus Michx. var. hispidus; 3, 7, 9, 15.UOS-1187.

*Ranunculus parviflorus L.; 7.UOS-2244.

Ranunculus recurvatus Poir.; 9, 10.UOS-3971.

*Ranunculus sardous Crantz; 6, 7.UOS-2264.

Thalictrum clavatum DC.; 8.UOS-7864.

Thalictrum dioicum L.; 8, 10.UOS-4490.

Thalictrum thalictroides (L.) Eames & B. Boivin; 9, 10, 12, 13, 14.UOS-1595.

Trautvetteria caroliniensis (Walt.) Vail; 3.UOS-7895.

Xanthorhiza simplicissima Marsh.; 3, 8.UOS-2482.


Berchemia scandens (Hill) K. Koch; 11.UOS-8163.

Ceanothus americanus L.; 7.UOS-2647.

Rhamnus caroliniana Walt.; 4, 7, 11, 13.UOS-2619.


Agrimonia microcarpa Wallr.; 10.UOS-4814.

Agrimonia parviflora Ait.; 7.UOS-7711.

Agrimonia pubescens Wallr.; 7, 15.UOS-2009.

Agrimonia rostellata Wallr.; 13, 15.UOS-4730.

Amelanchier arborea (Michx. f.) Fernald; 1, 2.UOS-7591.

Amelanchier laevis Wieg.; 1.UOS-4288.

*Aphanes microcarpa (Boiss. & Reut.) Rothm.; 6.UOS-8639.

Aronia arbutifolia (L.) Pers.; 2, 3.UOS-3317.

Aronia melanocarpa (Michx.) Ell.; 1, 2.UOS-821.

Aruncus dioicus (Walt.) Fern.; 8, 10.UOS-869.

Crataegus berberifolia Torr. & Gray; 9.UOS-8285.

Crataegus calpodendron (Ehrh.) Medik.; 11, 12.UOS-8289.

Crataegus collina Chapman; 10, 11.UOS-8309.

Crataegus crus-galli L.; 11.UOS-8160.

Crataegus intricata Lange; 15.UOS-8044.

Crataegus iracunda Beadle; 7.UOS-4872.

Crataegus macrosperma Ashe; 3.UOS-7617.

Crataegus pruinosa (Wendl. f.) K. Koch; 2, 9, 10.UOS-6267.

*Duchesnea indica (Andrews) Focke; 3, 6, 7, 9.UOS-877.

Fragaria virginiana Duchesne; 2.UOS-3121.

Geum canadense Jacq.; 2, 7, 8, 10, 13, 15.UOS-4906.

Geum vernum (Raf.) Torr. & Gray; 3.UOS-8504.

Geum virginianum L.; 7, 9, 10, 15.UOS-8221.

Malus angustifolia (Ait.) Michx.; 1, 2.UOS-4874.

Malus coronaria (L.) Mill.; 2.UOS-7815.

*Malus prunifolia (Willd.) Borkh.; 7.UOS-8506.

*Malus pumila Mill.; 7.UOS-7808.

Porteranthus stipulatus (Muhl. ex Willd.) Britton; 2, 7.UOS-861.

Porteranthus trifoliatus (L.) Britton; 7.UOS-2637.

Potentilla canadensis L.; 4, 10.UOS-6140.

*Potentilla recta L.; 7.UOS-777.

Potentilla simplex Michx. var. simplex; 2, 3, 7, 12.UOS-7911.

Prunus americana Marsh.; 15.UOS-8299.

Prunus angustifolia Marsh.; 7.UOS-8627.

*Prunus avium (L.) L.; 7.UOS-7570.

Prunus mexicana S. Watson; 9.UOS-8043.

Prunus munsoniana W. Wight & Hedrick; 7.UOS-7855.

Prunus serotina Ehrh.; 1, 2, 3, 4, 9, 10, 12, 13, 14.UOS-2855.

**Pyrus calleryana Decne.; 4.UOS-8398.

*Pyrus communis L.; 4, 7.UOS-8290.

Rosa carolina L. ssp. carolina; 2.UOS-2628.

**Rosa multiflora Thunb. ex Murr.; 3, 4, 7, 10, 15.UOS-889.

Rosa setigera Michx.; 15.UOS-1703.

*Rosa wichuraiana Crép.; 7.UOS-8505.

Rubus allegheniensis Porter; 1.UOS-1489.

Rubus argutus Link; 2, 4, 7.UOS-7910.

Rubus flagellaris Willd.; 2, 3, 8, 9.UOS-4003.

Rubus hispidus L.; 1, 3, 7.UOS-1693.

Rubus longii Fernald; 7.UOS-4293.

Rubus occidentalis L.; 7.UOS-4454.

**Rubus phoenicolasius Maxim.; 7, 8.UOS-922.

**Spiraea japonica L. f.; 7, 14, 15.UOS-898.


Cephalanthus occidentalis L.; 3.UOS-2736.

Diodia teres Walt.; 1.UOS-4538.

Diodia virginiana L.; 6, 7.UOS-2715.

Galium aparine L.; 3, 7, 10, 13.UOS-2876.

Galium circaezans Michx.; 2, 3, 8, 9, 10, 11, 12, 13, 15.UOS-3839.

Galium lanceolatum Torr.; 2.UOS-7898.

*Galium pedemontanum (Bellardi) All.; 4.UOS-8216.

Galium pilosum Ait.; 4, 7.UOS-7977.

Galium tinctorium (L.) Scop.; 7.UOS-7256.

Galium triflorum Michx.; 9, 10, 11, 14.UOS-7902.

Houstonia caerulea L.; 2, 3.UOS-1501.

Houstonia canadensis Willd. ex Roem. & Schult.; 2.UOS-1494.

Houstonia longifolia Gaertn.; 2.UOS-856.

Houstonia purpurea L. var. purpurea; 2, 9, 13, 15.UOS-2635.

Houstonia purpurea L. var. calycosa A. Gray; 2.UOS-3484.

Houstonia pusilla Schoepf; 2.UOS-2478.

Mitchella repens L.; 1, 3, 8, 14.UOS-2636.

*Sherardia arvensis L.; 6.UOS-6341.


Ptelea trifoliata L.; 9, 11.UOS-4516.


**Populus alba L.; 4.UOS-8243.

Populus deltoides Bartr. ex Marsh.; 7.UOS-8349.

Salix caroliniana Michx.; 7.UOS-4831.

Salix humilis Marsh. var. microphylla (Andersson) Fernald; 7.UOS-8131.

Salix nigra Marsh.; 7, 15.UOS-881.


Phoradendron serotinum (Raf.) M.C. Johnst.; 2, 7.UOS-4308.


Acer negundo L.; 7, 9, 14.UOS-2466.

Acer rubrum L. 1.; 1, 2, 3, 4, 10, 11, 13, 14.UOS-833.

Acer saccharinum L.; 7.UOS-4780.

Acer saccharum Marsh. ssp. floridanum (Chapman) Desmarias; 10, 13.UOS-8372.

Acer saccharum Marsh. ssp. saccharum; 9, 10, 11, 12, 13, 14.UOS-392.

Aesculus flava Ait.; 9, 10, 13, 14.UOS-4784.

Aesculus glabra Willd.; 13.UOS-2539.

*Cardiospermum halicacabum L.; 7. UOS-8893.


Sideroxylon lycioides L.; 9, 11, 13.UOS-7718.


Astilbe biternata (Vent.) Britton; 9, 14.UOS-8440.

Heuchera americana L. var. americana; 1, 8.UOS-7817.

Heuchera parviflora Bartl.; 8.UOS-2682.

Heuchera villosa Michx. var. villosa; 11.UOS-7715.

Micranthes careyana (Gray) Small; 8, 10.UOS-4789.

Mitella diphylla L.; 14.UOS-8180.

Tiarella cordifolia L.; 10, 14.UOS-591.


**Buddleja davidii Franch.; 7.UOS-8887.

Scrophularia marilandica L.; 7, 10, 15.UOS-7985.

*Verbascum blattaria L.; 7.UOS-921.

**Verbascum thapsus L.; 7.UOS-2066.


**Ailanthus altissima (Mill.) Swingle; 7.UOS-2659.


*Datura stramonium L.; 7.UOS-945.

Physalis heterophylla Nees; 8, 11.UOS-8062.

Solanum carolinense L.; 4, 7.UOS-4833.

*Solanum physalifolium Rusby; 7.UOS-8633.

Solanum ptychanthum Dunal; 7.UOS-4866.


Staphylea trifolia L.; 9, 10.UOS-6343.


+Halesia tetraptera Ellis; 7.UOS-7701.


Stewartia ovata (Cav.) Weatherby; 2.UOS-4508.


Samolus valerandi L. ssp. parviflorus (Raf.) Hultén; 14.UOS-8188.


Ulmus alata Michx.; 2, 3, 9, 10, 11, 12.UOS-4488.

Ulmus americana L.; 7.UOS-4856.

Ulmus rubra Muhl.; 9, 13.UOS-4469.

Ulmus serotina Sarg.; 9, 11.UOS-7401.


Boehmeria cylindrica (L.) Sw.; 1, 9, 13.UOS-3980.

Laportea canadensis (L.) Weddell; 10.UOS-4859.

Parietaria pensylvanica Muhl. ex Willd.; 7.UOS-7975.

Pilea pumila (L.) Gray; 3, 10, 14.UOS-4074.

*Urtica dioica L. ssp. dioica; 4, 10, 14.UOS-8049.


Valerianellaradiata (L.) Dufr.; 7.UOS-1684.


Phyla lanceolata (Michx.) Greene; 7.UOS-7820.

Verbena simplex Lehm.; 4, 7.UOS-811.

Verbena urticifolia L.; 7.UOS-2007.


Hybanthus concolor (T.F. Forst.) Spreng.; 10.UOS-4438.

Viola bicolor Pursh; 7.UOS-2650.

Viola blanda Willd.; 8, 10.UOS-2480.

Viola canadensis L.; 10.UOS-8121.

Viola cucullata Ait.; 3.UOS-7578.

Viola hastata Michx.; 2, 10.UOS-2320.

Viola hirsutula Brainerd; 2.UOS-8245.

Viola lanceolata L.; 4, 7.UOS-7621.

Viola palmata L.; 2, 4, 9, 10, 11, 15.UOS-3772.

Viola pedata L.; 4, 7.UOS-2483.

Viola primulifolia L.; 3, 10.UOS-6136.

Viola pubescens Aiton var. scabriuscula Schwein. ex Torr. & A. Gray; 2, 10, 13, 15.UOS-7582.

Viola rostrata Pursh; 10.UOS-4268.

Viola sagittata Aiton var. sagittata; 1.UOS-7603.

Viola sororia Willd. var. sororia; 6, 7, 10.UOS-1596.

Viola striata Ait.; 3, 6, 7, 10.UOS-8246.

Viola subsinuata (Greene) Greene; 7.UOS-8282.

Viola tripartita Elliot var. glaberrima (DC.) Harper; 14.UOS-8244.


Ampelopsis arborea (L.) Koehne; 7.UOS-8488.

Parthenocissus quinquefolia (L.) Planch.; 2, 4, 9, 10, 11, 12, 13, 14.UOS-2620.

Vitis aestivalis Michx. var. aestivalis; 2, 7, 8, 10, 11.UOS-7935.

Vitis cinerea (Engelm.) Engelm. ex Millard var. baileyana (Munson) Comeaux; 2, 9, 11, 15.UOS-832.

Vitis rotundifolia Michx.; 7.UOS-5942.

Vitis vulpina L.; 2, 10.UOS-2180.

Jonathan P. Evans, Callie A. Oldfield, Mary P. Priestley, Yolande M. Gottfried, L. Dwayne Estes, Alfire Sidik, and George S. Ramseur "The Vascular Flora of the University of the South, Sewanee, Tennessee," Castanea 81(3), 206-236, (1 September 2016).
Received: 28 January 2016; Accepted: 1 June 2016; Published: 1 September 2016
Cumberland Plateau
floristic inventory
plant communities
rare species
Sewanee, Tennessee
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