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The Vascular Flora of an Old-Growth Mixed Mesophytic Forest in Southeastern Kentucky

Posted on: Sunday, 26 February 2006, 03:02 CST

By McEwan, Ryan W; Paratley, Robert D; Muller, Robert N; Riccardi, Cynthia L

MCEWAN, R. W. (Department of Environmental and Plant Biology, Ohio University, Athens, OH 45701), R. D. PARATLEY (Department of Forestry, University of Kentucky, Lexington, KY 40546), R. N. MULLER (Santa Barbara Botanical Garden, Santa Barbara, CA 93105), AND C. L. RICCARDI (U.S. Forest Service, Pacific Northwest Research Station, Seattle, WA 98103). The vascular flora of Big Everidge Hollow, an old-growth mixed mesophytic forest. J. Torrey Bot. Soc. 132: 618- 627. 2005.-Because the goal of natural areas management is often the preservation of biodiversity, documenting botanical species richness is critically important. We conducted a series of botanical surveys in Big Everidge Hollow, a 52 ha watershed containing old-growth forest, on the Cumberland Plateau of eastern Kentucky. We contrasted our findings with a floristic survey that included parts of our study area, conducted approximately 20 years prior. Our research, from 1999 through 2001, yielded 263 species from 176 genera and 82 families, including 19 species that were new records from the site. These new species may have been overlooked in the previous study because of their scarcity or highly cryptic taxonomy, or they may have established in the years between studies. Of the 263 species recorded in our surveys, only one is considered non-native. This remarkable absence of exotic species indicates the high levels of "ecological integrity" inherent the study site and suggests an increasingly vital role for old-growth forests as reference ecosystems.

Key words: old-growth forest, flora, species richness, Cumberland Plateau, Lilley Cornett Woods.

As natural areas management is increasingly driven by the need to preserve biodiversity, maintaining plant species richness becomes critically important (Wilson 1988, U.S. Forest Service 1997, Wiser et al. 1998, Collins et al. 2001, Mace et al. 2003). Recent research has documented the loss of plant species richness due to timber harvesting (Meier et al. 1995, Jules 1998), and the loss of species over time in forested preserves (Davison and Forman 1982, Rooney and Dress 1997). These studies, among others, indicate the necessity of creating sitespecific floras and documenting shifts in species richness over time (Palmer et al. 1995). These efforts may be particularly important in the floristically rich central Appalachian region of eastern North America where management for biodiversity has been emphasized (Wofford 1989, KBTF 1995, SAMAB 1996, Dobson et al. 1997, Collins et al. 2001).

Old-growth forests are considered repositories of species richness and may provide refugia for species susceptible to loss (Meier et al. 1995, Rooney and Dress 1997, Jules 1998). Old-growth eastern forests often have a disturbance regime characterized by "gap-dynamics" (Runkle 1982) and these gaps may facilitate floristic diversity. Tree-fall gaps create a spatially heterogeneous light environment at the forest floor (Bazzaz and Wayne 1994) providing an array of habitats for plant species. The coarse woody debris that results from these tree-falls (Muller 2003) provides a heterogeneous substrate that enhances plant species richness (Bratton 1976, Thompson 1980). Forest harvesting can eliminate plant species by direct mechanical damage, by suspending gap-dynamics, and by altering growing conditions at the forest floor (Meier et al. 1995, Jules 1998, Matlack 2005). Because of potential species loss following anthropogenic disturbance, old-growth forests are often used as "reference" ecosystems and are considered ecological and floristic benchmarks (Meier et al. 1995, Davis 1996).

FIG. 1. Topographic map of Big Everidge Hollow, an old-growth forest watershed in southeastern Kentucky. Symbols on map indicate overstory plot locations designated in Fagus grandifolia ([white circle]), mixed mesophytic ([white triangle up]) and Quercus montana- Acer rubrum ([white square]) communities. Map was redrawn from Muller (1982).

As part of an ongoing, long-term ecological analysis (e.g., Muller 1982, McEwan et al. 2000, Muller 2003, McEwan et al. 2005), we conducted a series of botanical surveys of Big Everidge Hollow (BEH), a 52 ha old-growth mixed mesophytic forest watershed in eastern Kentucky. Big Everidge Hollow is part of the Lilley Cornett Woods Appalachian Research Station (LCWARS) and a floristic study of the entire Research Station had been conducted approximately twenty years prior to our analysis (Sole et al. 1983). Our objectives were to 1) describe the flora of the area and 2) detail species additions over the course of the approximately twenty years between studies. We also discuss implications of our floristic work for biodiversity management.

Methods. SITE DESCRIPTION. Big Everidge Hollow is an old-growth mixed mesophytic watershed located on the Cumberland Plateau of eastern Kentucky (37 05' N, 83 00' W, Roxana Quadrangle; Fig. 1). This location is representative of the larger Rugged Eastern section of the Cumberland Plateau and is near the geographic center of the mixed mesophytic forest described by Braun (1950). Regional climate is humid continental with warm summers, cool winters and no distinct dry season (Trewartha 1968). Mean annual temperature and precipitation are 13 C and 113 cm respectively (Hill 1976). Elevation ranges from 320 m to 600 m and mean slope is 50% (Muller 1982). The BEH watershed is generally surrounded by second-growth forest, although an adjacent watershed was contour mined in 1970 (Muller 1982). Big Everidge Hollow is a "U"-shaped watershed that opens to the east and is drained by a perennial stream. There is no evidence of commercial timber cutting in the watershed throughout its history. The only known direct human impacts on the watershed are from 1) removal of some dead trees, principally Castanea dentata stems following the blight, 2) hog grazing in a small section of the north-facing slope and 3) occasional illegal collection of medicinal herbs (e.g., Panax quinquefolius) in some areas of the forest. No significant damage from ice glaze or severe winds has been documented. The role of fire in the pre-history of the stand is unknown; however, no obvious signs of forest fire in the recent past have been found. This stand, therefore, has a disturbance regime characterized by gap-phase dynamics (Romme and Martin 1982).

The highly dissected terrain of the BEH watershed creates marked ecological contrast with minor changes in topographic or edaphic conditions (Martin 1975, Muller 1982, McEwan et al. 2005). Previous analysis of the woody overstory vegetation has distinguished three ecological communities: a lower slope Fag us grandifolia community, a mid-slope community characterized by Acer saccharum and Tilia americana, and an upper slope Quercus montana-Acer rubrum community (Muller 1982). Detailed analyses of vegetation-site relationships can be found in Muller (1982) and McEwan et al. (2005).

In order to document dynamics in the woody overstory vegetation, Muller (1982) installed 80 circular permanent plots, 0.04 ha in size, using a stratified random sampling scheme, throughout BEH. Inventories of woody stems ≥ 2.5 cm dbh were conducted in 1979 and 1989 (Muller 1982 and unpublished data). In order to expand the scope of the long-term project, McEwan et al. (2000) established four nested shrub-layer (10 m^sup 2^) and ground-layer (1 m^sup 2^) subplots in each original overstory plot (320 plots total). See McEwan et al. (2000) for details on plot establishment.

Table 1. Summary of the major taxonomic groups and percent native species (Jones 2005) documented in Big Everidge Hollow, an old- growth forest in southeastern Kentucky.

BOTANICAL SURVEYS. All taxa reported here were documented in a series of botanical surveys that began in the summer of 1999 and continued through the fall of 2001. In the summer of 1999, the original overstory plots were revisited and all stems ≥ 2.5 cm dbh were identified to species. An inventory of herbaceous species was conducted on all 320 ground-layer subplots in April, June, and August of 2000 and again in April, June and August of 2001. During the herb surveys, all herbaceous plants that covered any portion of the 1 m^sup 2^ ground-layer subplot were identified to species. In July of 2001, a survey of woody sterns was conducted on all 640 shrub-layer and ground-layer subplots. During the woody ground- layer survey, all woody plants that covered any of the 1 m^sup 2^ ground-layer plots and were less than 0.5 m tall were identified to species. During the shrub-layer survey, all woody-stems that were rooted within the 10 m^sup 2^ subplot and were ≥ 0.5 meters tall and < 2.5 cm dbh were identified to species. Finally, during each formal sampling and, during approximately 30 other visits to the study site throughout the growing seasons of 2000 and 2001, any species found throughout the watershed that had not previously been noted was identified and recorded.

Voucher specimens were collected of less common species and species that could only be identified using laboratory or herbarium resources. Because the study site is one of the best examples of old- growth mixed mesophytic forest remaining in North America, efforts were made to minimize the number of collections and to only collect material necessary to identify \individual specimens (e.g., collecting the flowering material only). Voucher specimens were deposited into the University of Kentucky Herbarium. Botanical nomenclature follows Jones (2005).

Results and Discussion. Our field surveys yielded 263 species from 176 genera and 82 families (Table 1). All but one of these species were collected in Big Everidge Hollow (BEH); the additional species (Cimicifuga americana, previously unknown at LCWARS) was noticed by chance in an adjacent old-growth section of LCWARS. Plant species richness in BEH is comparable to other areas within the region, but is markedly less than larger sites that include habitats other than mature upland forest, such as old-fields or wetlands (Table 2).

The species richness of BEH was notably smaller than reported for LCWARS as a whole (Sole et al. 1983). This difference is explained by the fact that Sole et al. (1983) conducted a transect study of a larger area including community types and habitats not represented in BEH (e.g., a small pond). Nonetheless, we found 19 species from 11 families that are new records for the site (Table 3). There are at least three possible reasons why these species were not reported in the Sole et al. (1983) survey. First, they may have colonized BEH in the period between the samplings. For example, a single sapling of Diospyros virginiana, which is common in southeastern Kentucky second-growth forests and easily identifiable, was found in our study. It is highly unlikely that this species would have been overlooked in previous studies, and thus, this species likely colonized the site between samplings. A second possible explanation is that Sole et al.'s (1983) collections did not contain material sufficient to identify obscure or taxonomically ambiguous species. For instance, Sanicula trifoliata is distinguishable from the more common Sanicula canadensis only by examination of mature floral or fruiting material under magnification (Radford et al. 1968, Wofford 1989, Gleason and Cronquist 1991). In addition, we report Acer nigrum.', however, it is considered a variety of Acer saccharum by some taxonomists (Radford et al. 1968), and neither morphological (St. Hilaire and Graves 1999) nor genetic analyses (Skepner and Krane 1998) have resolved its status. Lastly, it is possible the new records were present at LCWARS during the Sole et al. (1983) study but sampling was insufficient to detect all rare species. Eleven of the species that were not detected during the initial survey have a limited spatial distribution within BEH (Table 3) and two of these are listed as species of conservation interest in the state. Dryopteris carthusiana and Solidago curtsii occurred as single individuals in our BEH and are listed respectively as Special Concern and Threatened in Kentucky (Jones 2005). Since our list contained only five species that had a special conservation designation for the state of Kentucky (Appendix 1), the fact that two of these were missed in the previous survey strongly suggests that continued monitoring of the watershed is needed.

Table 2. Comparison of Big Everidge Hollow and local and regional floras. Studies are arranged in descending order by total species. Results of current study in bold italics for clarity. Note that cells containing nl are for non-listed values.

Table 3. Previously unreported species from Big Everidge Hollow, an old-growth forest in southeastern Kentucky.

Many exotic plants often found on trail edges and in openings within second-growth stands in the region were absent from BEH (Appendix 1). Weedy species of the Apiaceae, Brassicaceae, Caryophyllaceae, Fabaceae, Lamiaceae, Asteraceae and Polygonaceae that are widespread through the region do not occur within the watershed. Five species of Cardamine, all infrequent to common in rich woods, were recorded in BEH, but the elsewhere ubiquitous exotic Cardamine hirsuta L. was absent. Two native species of Lespedeza were found in BEH, but the exotic Lespedeza cuneata (Dum. Cours.) G. Don. was absent. Merely one native member (Polygonum virginianum) of a cohort of weedy smartweeds was recorded while Eurasian smartweeds, such as Polygonum persicaria L., were not found.

Invasive species that have frequently established themselves in other eastern Kentucky forests were also absent from BEH (Appendix 1). Examples of noteworthy absences include the woody plants Ailanthus altissima (Mill.) Swingle, Elaeagnus umbellata Thunb., Ligustrum sinense Lour, Paulownia tomentosa (Thunb.) Steud., and Rosa multiflora Thunb. Many nonnative and invasive grasses commonly found in secondary Cumberland Plateau woods, such as Miscanthus sinensis Anderss., Bromus secalinus L., Microstegium vimineum (Trin.) A. Camus., and Sorghum halapense (L.) Pers. were also absent. Weedy elements were present in BEH, but they consisted of native plants known to exploit edges and openings. Examples include Amphicarpaea bracteata, Galiutn aparine, Phytolacca americana, and Ranunculus abortivus. In fact, of the 263 species recorded here (Appendix 1), only Kentucky bluegrass (Poa pratensis) is generally considered non-native (Gleason and Cronquist 1991, Jones 2005) although some have challenged this classification (Wharton and Barbour 1991). As exotic species become increasingly widespread, the absence of exotics in oldgrowth systems such as BEH suggests an increasingly important role of these forests as reference ecosystems (McCarthy et al. 1987, McCarthy et al. 2001).

The plant species richness of BEH was less than that reported from natural areas where nonnatives add considerably to the species list, yet BEH exhibited impressive diversity within some groups and contained unusual native species. For instance, the native diversity of Aster, Solidago, and Viola in BEH was considerable. Big Everidge Hollow housed Solidago flaccidifolia and Astilbe biternata which are listed respectively as "uncommon" and "rare" in Kentucky, and Quercus marilandica which is uncommon in eastern Kentucky (Jones 2005). Several species that are typically restricted to the Appalachian Mountains such as Dryopteris carthusiana, Acer pensylvanicum, Solidago curtsii, and Cimicifuga americana were reported in our survey (Jones 2005). The relatively low total species richness of Big Everidge Hollow might lead to the untenable conclusion that management for biodiversity would include disturbing oldgrowth forest ecosystems to promote exotic species thereby increasing species diversity. However, the details of our flora suggest that BEH is a storehouse for native species diversity, and perhaps due to the absence of some species, it exhibits high level of "ecological integrity" (Angermeier 1994). Thus, the question for biodiversity preservation seems not to be "how much diversity?" but rather "which diversity?"

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Ryan W. McEwan1,2

Department of Environmental and Plant Biology, Ohio University, Athens, OH 45701

Robert D. Paratley

Department of Forestry, University of Kentucky, Lexington, KY 40546

Robert N. Muller

Santa Barbara Botanical Garden, Santa Barbara, CA 93105

Cynthia L. Riccardi

U.S. Forest Service, Pacific Northwest Research Station, Seattle, WA 98103

1 The authors thank Rob Watt, the grounds supervisor of the Lilley Cornett Woods Appalachian Research Station, for his hospitality and gracious assistance during the field surveys. Thanks also to Bill Martin and Eastern Kentucky University for access to the study site. We also thank Amy L. Goff-Yates, John Graham, Kevin Lewis, Jeff Lombardo, and two anonymous reviewers for helpful comments on earlier drafts of the manuscript. Finally, many thanks are owed to C. John Burk who carefully read and edited the original and revised manuscript, greatly improving its readability.

2 Author for correspondence, E-mail: ryan.w.mcewan.l @ohio.edu

Received for publication November 11, 2004, and in revised form July 27, 2005.

Appendix

The vascular flora of Big Everidge Hollow, an old-growth mixed mesophytic forest on the Cumberland Plateau of eastern Kentucky. Nomenclature follows Jones (2005). Abundance of the taxa in the watershed is noted by classification into one of five categories which are a modification of the system used by Jones (2005). Species which are widespread and abundant in the watershed are listed as common (Com.). Species which are easily found in the watershed, but absent from some areas were classified as frequent (Freq.). Species that were scattered throughout the watershed, or occasional in certain habitats, were listed as infrequent (Inf.). Species reliably found within a restricted habitat, in which they may have been plentiful, were listed as isolated (Iso.). Taxa occurring in small numbers whenever found and that were often absent in expected habitats were classified as "Rare". Other abbreviations are threatened (T), endangered (E), special concern (S) as listed for Kentucky by Jones (2005), and NR for new record.

FERNS AND FERN ALLIES

Aspleniaceae

Asplenium platyneuron (L.) BSP. Inf.

Dryopteridaceae

Athyrium filix-femina (L.) Roth. Inf.

Diplazium pycnocarpon (Sprengel) M. Broun. Inf., NR

Dryopteris carthusiana (ViIl.) H.P. Fuchs. Rare, S, NR

Dryopteris marginalis (L.) A. Gray. Freq.

Dryopteris goldiana (Hook.) A. Gray. Rare, NR

Polystichum acrostichoides (Michx.) Schott. Com.

Ophioglossaceae

Botrychium virginianum (L.) Sw. Inf.

Pteridaceae

Adiantum pedatum L. Inf.

Thelypteridaceae

Phegopteris hexagonoptera (Michx.) Fee. Inf.

Thelypteris noveboracensis (L.) Nieuwl. Inf.

CONIFERS

Cupressaceae

Juniperus virginiana L. Rare

Pinaceae

Pinus echinala Mill. Iso.

Pinus rigida Mill. Iso.

Pinus virginiana Mill. Rare.

Tsuga canadensis (L.) Carriere. Freq.

FLOWERING PLANTS DICOTS

Aceraceae

Acer nigrum Michx. F. Rare, NR

Acer rubrum L. Com.

Acer saccharum Marshall. Com.

Acer pensylvanicum L. Rare

Anacardiaceae

Toxicodendron radicans (L.) Kuntze. Com.

Annonaceae

Asimina trlloba (L.) Dunal. Freq.

Apiaceae

Cryptotaenia canadensis (L.) DC. Iso.

Osmorhiza claytonii (Michx.) C. B. Clarke. Freq.

Osmorhiza longistylis (Torr.) DC. Rare, NR

Sanicula canadensis L. Inf.

Sanicula trifoliata E. P. Bicknell. Rare, NR

Taenidia integerrima (L.) Drude Rare

Thaspium barbinode (Michx.) Nutt. Inf.

Thaspium trifoliatum (L.) A. Gray. Freq.

Aqufoliaceae

Ilex opaca Alton. Inf.

Araliaceae

Panax quinquefolius L. Rare

Aristolochiaceae

Aristolochia macrophylla Lam. Rare

Asarum canadense L. Iso.

Asclepiadaceae

Asclepias quadrifolia Jacq. Inf.

Asteraceae

Ageratina altissima (L.) R.M. King & H. Rob. Com.

Antennaria plantaginifolia (L.) Richardson. Inf.

Antennaria solitaria Rydb. Inf.

Coreopsis major Walter. Inf.

Doellingeria infirma (Michx.) Nees. Inf.

Elephantopus carolinianus Raeusch. Rare

Erigeron pulchellus Michx. Rare

Eupatorium purpureum L. Inf.

Eurybia divaricata (L.) G. L. Nelsom. Com.

Helianthus microcephalus Torr. & A. Gray Rare

Hieracium paniculatum L. Iso.

Hieracium venosum L. Iso.

Krigia biflora (Walter) S. F. Blake. Iso.

Lactuca floridana (L.) Gaertn. Rare

Packera obovata (Muhl. ex Willd.) WA. Weber and A. Love Inf.

Prenanthes altissima L. Com.

Sericocarpus asteroides (L.) BSP. Inf.

Solidago bicolor L. Inf.

Solidago caesia L. Com.

Solidago curtsii Torr. & A. Gray Rare, T, NR

Solidago erecta Pursh. Inf.

Solidago flaccidifolia Small. Rare, NR

Solidago flexicaulis L. Freq.

Solidago sphacelata Raf. Inf.

Symphyotrichum cordifolium (L.) G. L. Nelsom. Com.

Symphyotrichum lanceolatum (Willd.) G. L. Nelsom. Rare, NR

Symphyotrichum lateriflorum (L.) A. Love & D. Love Inf.

Symphyotrichum patens (Alton.) G. L. Nelsom. Iso.

Symphyotrichum undulatum (L.) G. L. Nelsom. Inf.

Balsaminaceae

Impatiens capensis Meerb. Iso.

Impatiens pallida Nutt. Iso.

Berberidaceae

Caulophyllum thalictroides (L.) Michx. Freq.

Podophyllum peltatum L. Inf.

Betulaceae

Betula lenta L. Inf.

Carpinus caroliniana Walter. Freq.

Ostrya virginiana (Miller) K. Koch. Inf.

Bignoniaceae

Bignonia capreolata L. Com.

Boraginaceae

Cynoglossum virginianum L. Rare

Brassicaceae

Arabis laevigata (Muhl.) Poiret. Inf.

Cardamine parviflora L. Rare

Cardamine pensylvanica Muhl. Ex Willd. Inf.

Dentaria laciniata Muhl. Ex Willd. Freq.

Dentaria heterophylla Nutt. Rare

Dentaria diphylla Michx. Inf.

Campanulaceae

Campanula divaricata Michx. Freq.

Campanulastrum americana (L.) Small. Rare

Lobelia inflata L. Rare

Lobelia spicata Lam. Rare, NR

Caprifoliaceae

Sambucus canadensis L. Rare

Viburnum acerifolium L. Freq.

Caryophyllaceae

Silene rotundifolia Nutt. Rare

Silene virginica L. Inf.

Stellaria pubera Michx. Freq.

Celestraceae

Euonymus americanus L. Com.

Clusiaceae

Hypericum hypericoides (L.) Crantz Inf.

Cornaceae

Cornus alternifolia L.f. Freq.

Cornus florida L. Com.

Crassulaceae

Sedum ternatum Michx. Freq.

Ebenaceae

Diospyros virginiana L. Rare, NR

Ericaceae

Epigaea repens L. Rare

Gaultheria procumb\ens L. Rare

Gaylussacia baccata (Wangenh.) K. Koch. Iso.

Kalmia latifolia L. Freq.

Oxydendrum arboreum (L.) DC. Freq.

Rhododendron calendulaceum (Michx.) Torr. Inf.

Rhododendron cumberlandense E.L. Braun Rare

Rhododendron maximum L. Iso.

Vaccinium corymbosum L. Inf.

Vaccinium pallidum Alton. Inf.

Vaccinium stamineum L. Inf.

Euphorbiaceae

Acalypha virginica L. Inf.

Fabaceae

Amphicarpaea bracteata (L.) Fern. Com.

Cercis canadensis L. Inf.

Desmodium glutinosum (Muhl.) A. Wood. Inf.

Desmodium nudiflorum (L.) DC. Freq.

Desmodium pauciflorum (Nutt.) DC. Rare

Lespedeza hirta (L.) Hornem. Rare

Lespedeza intermedia (S. Watson) Britton. Rare

Robinia pseudoacacia L. Rare

Vicia caroliniana Walter. Inf.

Fagaceae

Castanea dentata (Marshall) Borkh. Rare, E

Fagus grandifolia Ehrh. Freq.

Quercus alba L. Freq.

Quercus coccinea Muenchh. Inf.

Quercus falcata Michx. Rare

Quercus marilandica Muenchh. Rare

Quercus montana Willd. Freq.

Quercus muhlenbergii Engelm. Rare

Quercus rubra L. Freq.

Quercus stellata Wangenh. Rare

Quercus velutina Lam. Inf.

Geraniaceae

Geranium maculatum L. Inf.

Hamamelidaceae

Hamamelis virginiana L. Inf.

Hippocastanaceae

Aesculus flava Ait. Inf.

Hydrangeaceae

Hydrangea arborescens L. Iso.

Hydrophyllaceae

Hydrophyllum macrophyllum Nutt. Inf.

Phacelia bipinnatifida Michx. Iso.

Juglandaceae

Carya cordiformis (Wangenh.) K. Koch. Inf.

Carya glabra (Miller) Sweet. Freq.

Carya ovata (Miller) K. Koch. Inf.

Carya tomentosa (Poiret) Nutt. Freq.

Jugions cinerea L. Rare, S

Juglans nigra L. Inf.

Lamiaceae

Collinsonia canadensis L. Inf.

Cunila origanoides (L.) Britton. Inf.

Meehania cordata (Nutt.) Britton. Rare

Monarda clinopodia L. Rare

Scutellaria elliptica Muhl. Inf.

Lauraceae

Lindera benzoin (L.) Blume. Iso.

Sassafras albidum (Nutt.) Nees. Inf.

Magnoliaceae

Liriodendron tulipifera L. Freq.

Magnolia acuminata (L.) L. Inf.

Magnolia macrophylla Michx. Inf.

Magnolia tripetala L. Inf.

Menispermaceae

Menispermum canadense L. Rare

Monotropaceae

Monotropa uniflora L. Rare

Moraceae

Morus rubra L. Rare

Nyssaceae

Nyssa sylvatica Marshall. Freq.

Oleaceae

Fraxinus americana L. Inf.

Fraxinus pennsylanica Marshall. Inf.

Onagraceae

Circaea lutetiana (L.) Asch. Inf.

Orobanchaceae

Conopholis americana (L.) Wallr. Rare

Epifagus virginiana (L.) Barton. Inf.

Oxalidaceae

Oxalis grandis Small. Inf.

Oxalis violacea L. Inf.

Papaveraceae

Sanguinaria canadensis L. Inf.

Phrymaceae

Phryma leptostachya L. Rare

Phytolaccaceae

Phytolacca americana L. Rare

Polemoniaceae

Phlox divaricata L. Inf.

Polygonaceae

Polygonum virginianum L. Inf.

Portulacaceae

claytonia virginica L. Freq.

Primulaceae

Lysimachia quadrifolia Sims. Inf.

Lysimachia tonsa (Alph. Wood) R. Knuth. Inf.

Pyrolaceae

Chimaphila maculata (L.) Pursh. Freq.

Ranunculaceae

Actaea pachypoda Elliot. Rare

Anemone quinquefolia L. Inf.

Anemonella thalictroides (L.) Spach. Inf.

Cimicifuga americana Michx. Rare, NR

Cimicifuga racemosa (L.) Nutt. Freq.

Clematis virginiana L. Rare

Hepatica acutiloba DC. Inf.

Ranunculus abortivus L. Rare

Ranunculus hispidus Michx. Inf.

Ranunculus recurvatus Poiret. Inf.

Thalictrum dioicum L. Rare, NR

Thalictrum pubescens Pursh. Rare

Rosaceae

Agrimonia pubescens Wallr. Inf.

Agrimonia rostellata Wallr. Inf.

Amelanchier arborea (F. Michx.) Fernald Inf.

Arnucus dioicus (Walter) Fernald Rare

Crataegus sp. L. Inf.

Geum canadense Jacq. Inf.

Potentilla canadensis L. Inf.

Potentilla simplex Michx. Inf.

Prunus serotina Ehrh. Inf.

Rosa Carolina L. Inf.

Rubus allegheniensis Porter Inf.

Rubus flagellaris Willd. Inf.

Rubiaceae

Galium aparine L. Inf.

Galium circaezans Michx. Inf.

Galium latifolium Michx. Inf.

Galium triflorum Michx. Freq.

Houstonia longifolia Gaertn. Inf.

Mitchella repens L. Inf.

Santalaceae

Pyrularia pubera Michx. Iso.

Saxifragaceae

Astilbe biternata (Vent.) Britton. Rare

Heuchera americana L. Inf.

Heuchera villosa Michx. Iso.

Tiarella cordifolia L. Iso.

Scrophulariaceae

Aureolaria laevigata (Raf.) Raf. Freq.

Pedicularis canadensis L. Inf.

Tiliaceae

Tilia americana L. Freq.

Ulmaceae

Ulmus americana L. Inf.

Ulmus rubra Muhl. Inf.

Urticaceae

Laportea canadensis (L.) Wedd. Iso.

Pilea pumila (L.) A. Gray Rare

Violaceae

Viola labridorica Schrank. Rare

Viola cucullata Aiton. Rare, NR

Viola palmata L. Inf.

Viola pubescens Aiton. Rare

Viola rostrata Pursh. Freq.

Viola sororia Willd. Inf.

Viola striata Aiton. Inf.

Vitaceae

Parthenocissus quinquefolia (L.) Planch. Com.

Vitis aestivalis Michx. Inf.

MONOCOTS

Araceae

Arisaema triphyllum (L.) Schott. Freq.

Convallariaceae

Polygonatum biflorum (Walter) Elliott Inf.

Maianthemum racemosum (L.) Link Freq.

Cyperaceae

Carex communis L. H. Bailey Freq.

Carex digitalis Willd. Freq.

Carex laxiflora Lam. Inf.

Carex normalis Mackenzie Rare, NR

Carex platyphylla Carey. Rare, NR

Carex prasina Wahlenb. Inf.

Carex virescens Muhl. Ex Willd. Inf.

Dioscoreaceae

Dioscorea villosa L. Freq.

Iridaceae

Iris cristata Soland. ex Aiton. Iso.

Sisyrinchium angustifolium Mill. Rare

Juncaceae

Luzula echinata (Small) FJ. Herm. Inf., NR

Liliaceae

Erythronium americanum Ker Gawl. Iso.

Medeola virginiana L. Rare

Orchidaceae

Aplectrum hyemale (Muhl. Ex Willd.) Torr. Rare

Cypripedium acaule Aiton. Rare

Goodyera pubescens (Willd.) R. Br. Freq.

Tipularia discolor (Pursh) Nutt. Rare

Poaceae

Agrostis perennans (Walter) Tuck. Freq.

Brachyelytrum erectum (Schreber) P. Beauv. Inf.

Bromus pubescens Muhl. Rare, NR

Danthonia spicata (L.) E Beauv. Inf.

Diarrhena americana P. Beauv. Inf.

Dichanthelium boscii (Poir.) Gould & C. A. Clark Inf.

Dichanthelium clandestinum (L.) Gould Inf.

Dichanthelium commutatum (Schult.) Gould Inf.

Dichanthelium dichotomum (L.) Gould Inf.

Elymus hystrix L. Inf, NR

Elymus riparius Wiegand Rare

Festuca subverticillata (Pers.) E. Alexeev. Inf.

Poa cuspidata Nutt. Inf.

Poa pratensis L. Inf.

Poa sylvestris A. Gray. Inf.

Smilacaceae

Smilax glauca Walter. Inf.

Smilax rotundifolia L. Freq.

Smilax tamnoides L. Inf.

Trilliaceae

Trillium erectum L. Iso.

Trillium grandiflorum (Michx.) Salisb. Iso.

Uvulariaceae

Prosartes lanuginosa (Michx.) D. Don. Inf.

Prosartes maculata (Buckley) A. Gray. Rare, S

Uvularia perfoliata L. Freq.

Copyright Torrey Botanical Society Oct-Dec 2005

Source: Journal of the Torrey Botanical Society

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