Southeastern Naturalist R.L. Bartgis, E.A. Byers, R.H. Fortney, W. Grafton, and M.A. Berdine 2015 Vol. 14, Special Issue 7 158 the Valley & Environs 2015 Southeastern Naturalist 14(Special Issue 7):158–186 Rare Plants of Canaan Valley, West Virginia Rodney L. Bartgis1,5,*, Elizabeth A. Byers2, Ronald H. Fortney3, William Grafton4, and M. Ashton Berdine1, 6 Abstract - Canaan Valley (hereafter, the Valley), in northeastern West Virginia, supports large areas of wetland, upland forest, and upland non-forest habitats at relatively high elevations, providing potential habitat for a variety of rare plant species. The presence of 54 species of plants considered to be rare and of conservation concern in West Virginia plus 22 watchlist species has been confirmed in the Valley. No federally listed threatened or endangered plants have been found. One of the rare species is the globally critically imperiled Platanthera shriveri (Shriver’s Frilly Orchid) and 4 are globally vulnerable—Gymnocarpium appalachianum (Appalachian Oak Fern), Hypericum mitchellianum (Blue Ridge St. Johnswort), Euphorbia purpurea (Glade Spurge), and Polemonium vanbruntiae (Bog Jacob’s-ladder). Rare plants are found throughout the Valley; 80% occur in wetlands, and a significant assemblage is associated with wetlands on Greenbrier Limestone. Globally rare species are Appalachian endemics, but 41 of the Valley’s rare and watchlist plants are primarily northern in distribution. Extant native populations are known in West Virginia only from the Valley for 3 species—Carex atherodes (Awned Sedge), Gentianopsis crinita (Greater Fringed Gentian), and Viburnum trilobum (American Cranberry-bush)—and a significant portion of all known West Virginia occurrences for at least another 16 species are in the Valley. Several rare plant species grow in multiple places in the Valley, but others are known from only 1 or 2 sites and are quite vulnerable. Non-native insect pests threaten Abies balsamea (Balsam Fir) and Fraxinus nigra (Black Ash). There are a number of threats to the Valley’s rare plants: invasive plants, especially Typha latifolia (Broadleaf Cattail), Iris pseudoacorus (Yellow Flag), Phalaris arundinacea (Reed Canarygrass), and Microstegium vimineum (Japanese Stiltgrass); browsing by Odocoileus virginianus (White-tailed Deer); residential development; hydrologic changes to wetlands; and climate change. Introduction The flora of Canaan Valley (hereafter, the Valley) has long been recognized for its interesting mix of northern, eastern, and Appalachian plant species (Allard and Leonard 1952; Brooks 1957; Core 1966; Fortney 1975, 1993). Many of the Valley’s plants have been classified as rare in West Virginia and even rangewide (Fortney 1975). Efforts to conserve this unique flora, plus its associated unusual fauna, led to the establishment of the Canaan Valley National Wildlife Refuge. 1The Nature Conservancy, 194 Airport Road, Elkins, WV 26241. 2West Virginia Division of Natural Resources, PO Box 67, Elkins, WV 26241. 3Department of Civil and Environmental Engineering, West Virginia University, PO Box 6103, Morgantown, WV 26506 (deceased). 4West Virginia University Extension Service, PO Box 6125, Morgantown, WV 26506 (deceased). 5Current address - PO Box 894, Elkins, WV 26241. 6Current address - West Virginia Land Trust, PO Box 11823, Charleston, WV 25339-1823. *Corresponding author - rodbartgis@gmail.com. Southeastern Naturalist R.L. Bartgis, E.A. Byers, R.H. Fortney, W. Grafton, and M.A. Berdine 2015 Vol. 14, Special Issue 7 159 The Refuge’s mission, in part, is to preserve natural floral diversity and abundance. Its management objectives emphasize protecting wetlands and rare plant communities (USFWS 2011). In addition, Canaan Valley State Park lies in the southern part of the Valley. The purposes of West Virginia state parks include “preserving and protecting natural areas of unique or exceptional scenic, scientific ... significance” (West Virginia Code §20-5-3). To effectively conserve the Valley’s rare plants requires an understanding of their status and threats to their survival. Allard and Leonard (1952) and Fortney (1975) published the most thorough analyses of the Valley’s flora and plant communities. Previously published summaries of the Valley’s rare plants are now out of date (Fortney 1975, USFWS 1979). In this paper, we report the findings of a series of rare plant surveys conducted from 1990 through 2002, provide additional records from the West Virginia Division of Natural Resources (WVDNR) Natural Heritage Program, and review current information on the Valley’s rare plants. Study Area The Valley is a high-elevation depression within the Allegheny Mountains of Tucker County in northeastern West Virginia. This steep-sided, oval-shaped basin is sited within the Allegheny Mountains section of the Central Appalachian Broadleaf Forest-Coniferous Forest-Meadow Ecoregion Province (McNab and Avers 1994). The Valley’s elevations range from 3140 to 4307 ft (957–1313 m) above sea level; the Valley’s floor averages 3200 ft (975 m). Because of its high elevation, the Valley’s climate is classified as cool-temperate. The Valley is subject to the frost-pocket effect, in which cold air from the surrounding mountains flows downslope and pools on the Valley’s floor, often producing patches of unseasonably low temperatures. Frost is frequently reported as late as June and as early as August (Fortney 1975). The Valley underwent the same anthropogenic changes as the surrounding Allegheny Mountains, namely the destruction of its original forest by logging and fire (Allard and Leonard 1952). As it was cleared, an extensive area in the central part of the Valley was used for a large-scale cattle operation (Robinson 1953). Before the period of intensive logging and fires, the Valley supported an extensive conifer forest characterized by Picea rubens Sarg. (Red Spruce) (Allard and Leonard 1952, Brooks 1957). Today, the Valley’s vegetation is chiefly forb- and grass-dominated meadows, successional shrublands, and northern hardwood forests (Fortney and Rentch 2003). Red Spruce stands are restricted to small areas along the crests of surrounding mountains, on lower mountain slopes, in protected coves, and scattered within mixed conifer–hardwood forests in or near wetlands on the Valley’s floor. The upland hardwood forests are dominated by Acer saccharum Marshall (Sugar Maple), Fagus grandifolia Ehrh. (American Beech), and Prunus serotina Ehrh. (Black Cherry) (Fortney 1993). The Valley’s American Beeches are declining rapidly due to a disease complex called beech-bark disease. Southeastern Naturalist R.L. Bartgis, E.A. Byers, R.H. Fortney, W. Grafton, and M.A. Berdine 2015 Vol. 14, Special Issue 7 160 Although the primeval Red Spruce-dominated forests were likely majestic, the most distinguishing ecological feature of the Valley is the extent of its wetlands. The Valley contains about 9000 ac (3600 ha) of wetlands, which collectively comprise the largest wetland complex in the Central Appalachians (Fortney 1993). The Valley’s wetlands overlay three kinds of material: acidic substrates of peat and sandstone, circumneutral substrates associated with Greenbrier Limestone, and alluvium. Because of strong hydrologic influences, a high proportion of the Valley’s flora is composed of hydrophytic species, as is evident by the presence of numerous species of Scirpus (bulrush), Juncus (rush), Carex (sedge), and Glyceria (mannagrass). Many of the wetland species have strong affinities with more northern latitudes. In quantitative terms, Fortney (1975) reported that 80% of the Valley’s 523 plant species have generally northern distributions. Methods Natural Heritage staff of the WVDNR reviewed museum collections, herbarium records, and scientific articles regarding the rare plant species that had been reported in the Valley, including known locations and habitat information through 1989. Natural Heritage staff reviewed geologic and topographic maps and aerial photographs to identify additional sites that could potentially support rare species. We visited most of these sites in 1990, 1996, 1997, and 2002. When we encountered a rare plant, we mapped its location and described the population using the standard methods of NatureServe (2002). In 2014, we conducted a second review of herbarium and rare plant records available through the WVDNR. This review included: (1) the results of field studies conducted during several graduate theses (Bonner 2005, Ceperley 2002, Francl 2003); (2) records contributed by outside observers to the Natural Heritage database; and (3) field work by WVDNR staff as part of National Vegetation Classification surveys and conservation assessment of high-elevation wetlands and spruce forests in West Virginia. Conservation-status rankings follow those assigned by NatureServe for global (i.e., rangewide) ranks and by WVDNR for state-specific ranks (WVDNR 2014c). Each species was assigned a rank following a standardized system described in Faber-Langendoen et al. (2012). In a simplified sense, species ranked G1 are globally critically imperiled, G2 are globally imperiled, G3 are globally vulnerable, G4 are globally apparently secure, and G5 are globally secure. At the state level, species ranked S1 are considered to be critically imperiled within West Virginia, S2 are imperiled, S3 are vulnerable, S4 are apparently secure, and S5 are secure in the state. Consequently, a G5 S2 species, then, is secure when its whole range is considered, but is imperiled in West Virginia. We classified species ranked G1, G2, G3, S1, or S2 as being of conservation concern and S3 species with a global rank of G4 or G5 as watchlist species and worthy of monitoring because with further decline, they will become rare in the state. We viewed species with a status uncertain enough that it was assigned a spanning rank (such as G3G4 or S2S3) as if it had the less Southeastern Naturalist R.L. Bartgis, E.A. Byers, R.H. Fortney, W. Grafton, and M.A. Berdine 2015 Vol. 14, Special Issue 7 161 secure rank. We use the term “species” to include infraspecific taxa recognized and tracked by WVDNR. Our nomenclature generally follows Gleason and Cronquist (1991) and Harmon et al. (2006). Results and Discussion There are 54 species of conservation concern in the Valley (Appendix 1) and 22 watchlist species (Appendix 2). Five species are of global conservation concern. Platanthera shriveri P.M. Brown (Shriver’s Frilly Orchid) is globally critically imperiled, and 4 species are globally vulnerable: Euphorbia purpurea (Raf.) Fern. (Glade Spurge), Gymnocarpium appalachianum Pryer (Appalachian Oak Fern), Hypericum mitchellianum Rydb. (Blue Ridge St. Johnswort), and Polemonium vanbruntiae Britt. (Bog Jacob’s-ladder). Bog Jacob’s-ladder is a plant primarily of the Central and Northern Appalachians. The other 4 species of global conservation concern are endemic or nearly so to the Central and Southern Appalachians. Two other Central and Southern Appalachian endemics, Cuscuta rostrata Shuttlw. ex Englem. & Gray (Beaked Dodder) and Listera smallii Wieg. (Small’s Twayblade), are of conservation concern based on their state ranks. Another such endemic, Viola appalachiensis Henry (Appalachian Blue Violet), is a watchlist species. Additionally, the watchlist species Carex aestivalis M.A. Curtis ex Gray (Summer Sedge) is broadly an Appalachian endemic, found from New England to Alabama. The Central and Southern Appalachians are rich in endemic species. These species have restricted ranges and, although most Appalachian endemics are abundant enough to be considered secure, it is not surprising that nearly all of the Valley’s plants of global conservation concern are regional endemics. A large number of the species of interest are northern species that extend southward through the Central and Southern Appalachians, including 32 (59%) species of conservation concern and 9 (41%) watchlist species. All of these are otherwise secure across their entire range. Indeed, most have wide distributions and many can be found abundantly in cool regions across the northern US, Canada, Europe, and Asia. The southernmost known occurrence of Carex atherodes Spreng. (Awned Sedge) is in the Valley, as is the likely southernmost native occurrence of Viburnum opulus L. var. americanum Ait. (American Cranberry-bush). However, the native distribution of the latter in West Virginia has become uncertain due to the widespread planting of American Cranberry-bush and frequent confusion with the widely planted and similar-looking Viburnum opulus L. var. opulus Ait. (Eurasian Guelder-rose). Both varieties have escaped from cultivation to wetlands. Forty-four (81%) of the species of conservation concern and 16 (73%) of the watchlist species in the Valley are reported from wetland habitats, reflecting both the unique quality of the Valley’s wetland habitats and the relative scarcity of wetlands throughout West Virginia. Some rare species are known from very small populations in the Valley, including the orchids Small’s Twayblade, Cypripedium reginae Walt. Southeastern Naturalist R.L. Bartgis, E.A. Byers, R.H. Fortney, W. Grafton, and M.A. Berdine 2015 Vol. 14, Special Issue 7 162 (Showy Lady’s-slipper), and Shriver’s Frilly Orchid. Other rare species have one or more large occurrences in the Valley, each with thousands of stems. Of the species of conservation concern, Glyceria grandis S. Wats. (American Mannagrass), Bog Jacob’s-ladder, Rhamnus alnifolia L’Hér. (Alderleaf Buckthorn), Torreychloa pallida (Torr.) Church var. fernaldii (A.S. Hitchc.) Dore ex Koyama & Kawano (Fernald’s Mannagrass), and Zigadenus leimanthoides Gray (Bog Death-camas) have very large populations in the Valley. Among the watchlist species, Carex canescens L. (Silvery Sedge), Carex utriculata Boott (Beaked Sedge), Drosera rotundifolia L. (Roundleaf Sundew), Hasteola suaveolens (L.) Pojark. (False Indian-plantain), Scirpus atrocinctus Fern. (Blackgirdle Bulrush), Scirpus microcarpus J. & K. Presl (Red-tinge Bulrush), Vaccinium macrocarpon Ait. (Large Cranberry), and Vaccinium oxycoccos L. (Small Cranberry) have large populations. Five rare species previously recorded from the Valley have not been documented there in at least 40 years: Botrychium matricariifolium (A. Braun ex Dowell) A. Braun ex Koch (Daisy-leaved Grape-fern), Beaked Dodder, Geum aleppicum Jacq. (Yellow Avens), Lonicera canadensis Bartr. ex Marsh. (Fly Honeysuckle), and Taxus canadensis Marsh. (Canada Yew). Potential habitat still exists in the Valley, and these species may still occur here. All 5 species have extant populations elsewhere in West Virginia. A few species, listed alphabetically by genus below, are worthy of special comment. Abies balsamea (L.) P. Mill. (Balsam Fir) is ranked as G5 and S1. Balsam Fir grows in several kinds of habitats in the Valley, including circumneutral wetlands and alluvial wetlands, and less frequently in acidic wetlands and upland areas adjacent to wetlands (Ceperley 2002, Stephenson and Adams 1986). Prior to the large-scale logging a century ago, there was probably a large, robust Balsam Fir population mixed with Red Spruce and Tsuga canadensis (L.) Carriere (Eastern Hemlock) throughout the Valley. This population was almost certainly the largest one in West Virginia, but logging and subsequent fires severely reduced and fragmented its coverage. After nearly a century of limited recovery, this diminished Balsam Fir population has been attacked by an invasive insect, Adelges piceae Ratz. (Balsam Woolly Adelgid). Since the adelgid was first first recorded in Tucker County in 1992, many trees in the Valley have been infested and died (Ceperley 2002). Indeed, 43% of all Balsam Fir trees measured by Cherefko et al. (2015 [this issue]) had been killed by the Balsam Woolly Adelgid. Odocoileus virginianus Zimm. (White-tailed Deer) have also significantly reduced stem growth and density of the remaining Balsam Firs (Cherefko et al. 2015 [this issue]). In 1990, before the Balsam Woolly Adelgid infestation was significant, we observed that browsing by White-tailed Deer had almost completely eliminated the recruitment of Balsam Fir at 12 of 13 sites in the Valley. Because the adelgid kills older Balsam Fir, it appears very likely that browsing will significantly restrain the successful replacement of younger trees. Balsam Fir’s rapid decline during the last decade has caused its state conservation-status Southeastern Naturalist R.L. Bartgis, E.A. Byers, R.H. Fortney, W. Grafton, and M.A. Berdine 2015 Vol. 14, Special Issue 7 163 rank to shift from vulnerable to critically imperiled. Mortality has diminished over the last few years and there has been some regeneration, but the long-term outlook for this species is bleak in West Virginia. Although the Southern Appalachian endemic Beaked Dodder has been reported historically from damp habitats in the Valley (Allard and Leonard 1952), there are no recent records of its presence. In West Virginia, Beaked Dodder is known from several high-elevation sites where it typically grows in sunny openings parasitizing Solidago rugosa Mill. (Wrinkleleaf Goldenrod). Extant populations are known nearby in Blackwater Falls State Park and on Dolly Sods. There is substantial potential habitat available in the Valley, so its apparent rarity here is remarkable. Showy Lady’s-slipper is known to live at only 2 sites in West Virginia. In the past, White-tailed Deer have substantially damaged this small population, reducing it from 2 stands in the 1980s to only 1 stand today. Fortunately, the remaining stand has been partly fenced and is relatively stable (E.A. Byers, pers. observ.). The population outside of the Valley is larger and has been successfully fenced to protect it from browsing by White-tailed Deer (Gregg 2004). At least two other West Virginia Showy Lady’s-slipper populations were destroyed by human activity in the Cheat Mountains during first half of the 20th century (M. Brooks, West Virginia University, Morganstown, WV, pers. comm.). Glade Spurge, which is often thought of as a wetland species (Strausbaugh and Core 1977), also occurs in uplands, including in the Valley. Here, the upland Glade Spurge sites are former pastures on limestone, typically with scattered Crataegus spp. (hawthorns) and a dense grass understory. At such sites, Danthonia compressa Austin (Mountain Oatgrass) and Anthoxanthum odoratum L. (Sweet Vernal Grass) are dominant grasses, except at one site where Poa compressa L. (Canada Bluegrass) is dominant. Glade Spurge also grows on limestone bedrock outcroppings. The Valley supports the second largest population of Glade Spurge known in West Virginia and one of the largest populations anywhere. Most other West Virginia populations are in montane wetlands, but the species’ largest reported population consists of thousands of plants growing in well-drained limestone hillside pastures near Osceola, Randolph County, WV. Fraxinus nigra Marsh. (Black Ash) occurs locally in the Valley wetlands, where it is often associated with Balsam Fir. Some stands are relatively large and may consist of a hundred or more trees. Until recently, Black Ash was considered vulnerable in West Virginia but has become imminently threatened by a beetle native to Eurasia, Agrilus planipennis Fair. (Emerald Ash Borer). The borer is a pest of North American ashes, including Black Ash, which have no natural defenses against it. First detected in North America in 2002, the borer has quickly expanded its range across North America and is expected to kill all ash trees within 10 years of infecting an area (Herms and McCullough 2014). As of October 2014, the Emerald Ash Borer was in 3 of the 9 West Virginia counties known to have Black Ash and was reported to be within 50 mi (80.5 km) of the Valley (USDA 2014). Southeastern Naturalist R.L. Bartgis, E.A. Byers, R.H. Fortney, W. Grafton, and M.A. Berdine 2015 Vol. 14, Special Issue 7 164 Emerald Ash Borer is likely to be documented in the Valley in the near future, and we expect that Black Ash will become functionally extinct here unless an effective means to control the beetle is found. Blue Ridge St. Johnswort was long-overlooked in West Virginia, probably due to Strausbaugh and Core (1977) incorrectly stating its petal color. In the last 10 years, we have observed it at multiple sites in the Cheat Range in Pocahontas and Randolph counties, Mount Porte Crayon in Randolph County, Spruce Mountain in Pendleton County, and the Yew and Allegheny mountains in Pocahontas County, in addition to the Valley. In West Virginia, it is typically found in small numbers in moist seeps and damp, erosional streamsides in spruce and northern hardwood forests. It also occurs in other partially shaded to sunny damp habitats. We think it will likely be found at other high-elevation sites. Shriver’s Frilly Orchid is a recently described species believed to be a stable taxon of ancient hybrid origin (Brown et al. 2008). Leah Ceperley (US Fish and Wildlife Service, Davis, WV) documented this species in the Valley on a grassy roadside and under nearby hardwoods, both typical habitats for the species. Although the population is in Canaan Valley National Wildlife Refuge, the site is in an area with intensive subdivision development and therefore it is extremely vulnerable to extirpation. Thelypteris simulata (Davenport) Nieuwl (Bog Fern) occurs in small, scattered patches in the Valley. Despite extensive searching by botanists, Bog Fern is otherwise known in West Virginia from only a few mountain-county wetlands. There is substantial potential habitat in the Valley, so its apparent rarity is curious. Large Cranberry and Small Cranberry, while locally abundant in the Valley, are absent from some additional, seemingly suitable peatlands, possibly reflecting the limited presence of appropriate non-forested habitat prior to logging. Roundleaf Sundew distribution follows a similar pattern. Bog Death-camas, primarily found on the coastal plain, is known from 3 occurrences in Sphagnum (peat moss) and Polytrichum (haircap moss) peatlands in the Valley. The largest occurrence comprises several thousand plants. Elsewhere in West Virginia, Bog Death-camas is found in peatlands on Dolly Sods, on the Allegheny Front, and along the shores of Shavers Fork of the Cheat River. There are some unresolved taxonomic issues. Questions remain about the identity of the Gymnocarpium (oak fern) populations in the Valley. Fortney et al. (2000) treated the occurrences here as the diploid Appalachian Oak Fern, but thought the populations may represent the circumboreal tetraploid Gymnocarpium dryopteris (L.) Newman (Northern Oak Fern) or triploid hybrids probably derived from the two. Natural Heritage botanists have identified some of the Valley’s populations as Northern Oak Fern. Colonies of the triploid hybrid oak fern have been reported elsewhere in West Virginia (Pryer and Haufler 1993). All are morphologically very similar, and it seems prudent to more carefully study the plants in the Valley to confirm their identity. It is quite possible that all 3 oak fern forms occur here. Southeastern Naturalist R.L. Bartgis, E.A. Byers, R.H. Fortney, W. Grafton, and M.A. Berdine 2015 Vol. 14, Special Issue 7 165 Oak ferns have been documented from 4 sites overlaying Greenbrier Limestone in the southern third of the Valley. Of these, 3 were originally identified as the Appalachian Oak Fern. Two of these patches, one with 300–500 fronds and another with 2500–3000, occur in conifer swamps, where the oak fern grows on elevated root mounds, rocks, and woody debris. A third site, near a beaver pond, supports few plants (Fortney et al. 2000). The Appalachian Oak Fern is known from widely scattered wetland and cool forest sites in the state’s mountain counties and Ridge and Valley Province. At most upland sites, the plants are shallowly rooted in moss that covers rocks and woody debris. West Virginia’s largest known Appalachian Oak Fern populations grow on North Fork and Shenandoah mountains, both of which are in Pendleton County. Several historic locations have not been recently relocated (Fortney et al. 2000). The Appalachian Oak Fern’s preferred microhabitat was surely more abundant prior to logging. Fortney noted an adder’s-tongue fern in the Valley’s northern half, but neither photographed it nor collected a specimen (J. Rentch, pers. comm.). Ophioglossum pusillum Rafinesque (Northern Adder’s-tongue), which appears to be rare in West Virginia, is expected in cold, high-elevation sites (Bartgis and Breiding 1987). There are extant populations known near Cranberry Glades and at Blister Swamp, both in Pocahontas County, which resemble the Valley’s habitat. Northern Adder’s-tongue may actually be widespread in the Valley because it is easily overlooked and there is substantial potential habitat. However, Fortney reported the Valley’s population as Ophioglossum engelmannii Prantl (Limestone Adder’s-tongue), a southern species of dry, hot habitats at mostly low elevations. Limestone Adder’s-tongue is rare in West Virginia, where it is otherwise known only from cedar glades of the South Branch Potomac watershed (Bartgis 1993). An occurrence of Limestone Adder’s-tongue in the Valley would be extremely unusual. Although we expect that Northern Adder’s-tongue exists in the Valley, we have excluded Ophioglossum from our tables because we cannot confirm its identification. Two potentially rare hedgenettles, Stachys aspera Michx. (Hyssopleaf Hedgenettle) and S. tenuifolia Willd. (Smooth Hedgenettle), have been reported from the Valley. Hedgenettles also belong to a morphologically confusing group that has undergone substantial recent taxonomic revisions. Correct identities need to be confirmed for the Valley’s populations. The West Virginia rarity status of these 2 hedgenettle species is also uncertain. There are currently no plant species in the Valley that are listed as endangered or threatened by the US Fish and Wildlife Service. The Valley has suitable habitat for one threatened species, Trifolium stoloniferum Muhl. ex A. Eaton (Running Buffalo Clover), which occurs in modestly disturbed hardwood forests on or near limestone in other parts of Tucker County. However, it is not usually found in areas formerly dominated by conifers. Extant native populations for 3 species have been documented in West Virginia only from the Valley: Awned Sedge, Gentianopsis crinita (Froel.) Ma (Great Southeastern Naturalist R.L. Bartgis, E.A. Byers, R.H. Fortney, W. Grafton, and M.A. Berdine 2015 Vol. 14, Special Issue 7 166 Fringed Gentian), and American Cranberry-bush. A significant portion of the known West Virginia occurrences for at least another 16 species are in the Valley: Balsam Fir, Carex haydenii Dewey (Cloud Sedge), Carex lacustris Willd. (Lake Sedge), Showy Lady’s-slipper, Glade Spurge, Geum rivale L. (Purple Avens), American Mannagrass, False Indian-plantain, Bog Jacob’s-ladder, Alderleaf Buckthorn, Rubus pubescens Raf. (Dwarf Red Bramble), Blackgirdle Bulrush, Red-tinge Bulrush, Fernald’s Mannagrass, America Cranberry-bush, and Bog Death-camas. The survival of these plants in West Virginia depends on their persistence in the Valley. Based on botanical surveys conducted since 1990, several species previously considered rare enough to be ranked S1 or S2 have been redesignated as S3, based at least in part on their abundance in the Valley. These re-ranked species include the Silvery Sedge, False Indian-plantain, Blackgirdle Bulrush, and Redtinge Bulrush. Limestone-associated high-elevation wetlands are extremely rare in the Southern and Central Appalachians, and are most extensive in the region in West Virginia. Even here, they are restricted to a small area of wetlands underlain by Greenbrier Limestone that extends from eastern Preston County southward through eastern Tucker and Randolph counties into northeastern Pocahontas County. These wetlands are frequently fed by shallow limestone-influenced circumneutral groundwater, and are more mineral-rich than areas underlain by peat or sandstone. In addition to rare species, they support rare and unique plant communities that differ from those of other wetlands in the region (Byers et al. 2007). Development and construction activities that intercept or otherwise alter the movement of shallow groundwater may affect the wetlands that support rare plant species and natural communities (Rentch et al. 2008). Much of the Valley is underlain by Greenbrier Limestone (Cardwell et al. 1968). Our records indicate that in the Valley, Balsam Fir, Equisetum sylvaticum L. (Woodland Horsetail), Showy Lady’s-slipper, Glade Spurge, Black Ash, Purple Avens, Bog Jacob’s-ladder, Alderleaf Buckthorn, and American Cranberry- bush are especially associated with wetlands that are influenced by the limestone substrate. Wetlands overlying alluvium share some of the same rare and uncommon plants that live in circumneutral wetlands. These wetlands often occur as a mosaic of physiognomic types. Further, their soils appear to be nutrient-rich compared to wetlands on peat and other acidic substrates (Byers et al. 2007). Some species, such as False Indian-plantain and American Mannagrass, occur most frequently in the Valley in alluvial wetlands. Wetlands with peatmoss or sandstone at the surface are often acidic, nutrientpoor fens, dominated by peat and haircap mosses (Byers et al. 2007, Fortney 1975). These areas are often colloquially referred to as bogs, but there are no true, exclusively rain-fed bogs in the Valley. Prior to deforestation, most of these areas were likely dominated by Red Spruce, but some non-forested openings probably existed. In the Valley, Pogonia ophioglossoides (L.) Ker-Gawl. (Rose Pogonia), Southeastern Naturalist R.L. Bartgis, E.A. Byers, R.H. Fortney, W. Grafton, and M.A. Berdine 2015 Vol. 14, Special Issue 7 167 Large Cranberry, Small Cranberry, Roundleaf Sundew, and Bog Death-camas grow primarily in acidic, nutrient-poor fens. Fens dominated by peat and haircap mosses are widespread in the higher mountains of West Virginia and often support additional populations of the rare plant species that grow in the Valley’s fens. Allard and Leonard (1952) and Fortney (1975) presented limited data for assessing the historic trends of the Valley’s plant populations. A comparison of their comments with our observations suggests that over the last half century Silvery Sedge, False Indian-plantain, Red-tinge Bulrush, and Small Cranberry have become more abundant in the Valley, while Beaked Dodder has become rarer. In addition, when we examined aerial photographs taken in 1945 and 2000, we observed notable increases in the number and extent of clones of Carex utriculata Bott (Beaked Sedge), which are readily identifiable in the imagery (Fortney and Rentch 2003). We caution that for some of these species the apparent temporal trends may reflect biases caused by the observers’ familiarity with the species, the timing of field work, and/or the variable locations of floral surveys. Nine species of conservation concern and 1 watchlist species are associated with active or abandoned beaver ponds, including the only known West Virginia colony of Awned Sedge. Castor canadensis Kuhl (North American Beaver, hereafter Beaver), extirpated from West Virginia by 1835, were reintroduced into Tucker County in 1935 (Lesser and Cromer 2015 [this issue]). They flourished, and 127 colonies were counted in and around the Valley in 1974 (Lesser and Cromer 2015 [this issue]). It is highly likely that the abundance of species such as Fernald’s Mannagrass increased with the Beaver’s return. The Beaver-pond cycle creates a mosaic of early to mid-successional wetlands that provide important beta diversity to the natural communities of the Valley (Bonner et al. 2009). However, inundation by Beaver also has eliminated many conifer-dominated areas in the Valley (Fortney 1975, Rentch and Fortney 2003), including stands of Balsam Fir. Beaver also have the potential to eliminate highly localized rare species populations in the Valley. For example, Beaver dams associated with road construction have substantially altered a highly diverse circumneutral wetland complex in the Canaan Valley State Park. Inundation resulted in extensive Red Spruce and Balsam Fir mortality, deposition of a thick mineral surface over peat and other original surficial layers, increased downcutting that led to locally drier conditions, and a substantial expansion of Typha latifolia L. (Broadleaf Cattail) (Rentch et al. 2008). All or most of the Valley’s known populations of Carex bushii Mackenzie (Bush’s Sedge), Carex normalis Mackenzie (Greater Straw Sedge), Showy Lady’s-slipper, Woodland Horsetail, Purple Avens, Greater Fringed Gentian, Appalachian Oak Fern, Small’s Twayblade, Alder-leaved Buckthorn, Saxifraga pensylvanica L. (Eastern Swamp Saxifrage), Schizachne purpurascens (Torr.) Swallen (False Melicgrass), and American Cranberry-bush are restricted to small areas of the Canaan Valley State Park, including sites immediately adjacent to the impacted area described above. These sites, which also support large populations of Glade Spurge and Bog Jacob’s-ladder, are susceptible to hydrologic changes, Southeastern Naturalist R.L. Bartgis, E.A. Byers, R.H. Fortney, W. Grafton, and M.A. Berdine 2015 Vol. 14, Special Issue 7 168 herbivory by White-tailed Deer, Balsam Fir mortality, and colonization by invasive plants. The loss of these sites would substantially diminish the viability of a significant number of the Valley’s rare plants. The majority of other species of conservation concern and watchlist species have at least 1 population within the Canaan Valley National Wildlife Refuge. However, all or a significant proportion of the known occurrences of Blue Ridge St. Johnswort and Rose Pogonia in the Valley are on private property. We observed negative impacts on wetland habitats caused by all-terrainvehicles (ATVs) throughout most of the Valley during our 1990 field surveys. Our records suggest that ATV activity degraded the Valley’s wetlands by killing plants; churning the peat mat, which allows it to dry and decay; damming sites with churned soil or peat; and causing severe sedimentation in streams and standing water. In 1990, we encountered 15 populations of 6 species of rare and uncommon plants that had been killed by ATVs. Nondescript plants including sedges seem to have been most at risk. However, many populations of Bog Jacob’s-ladder were close to being damaged by ATVs in 1990. Subsequently, private and public landowners restricted ATV access to most of the Valley, which reduced the magnitude of the harmful impacts. In 2002, we again noted some local impacts to rare plant populations caused by ATV use. We also observed that some of the sites that had been damaged a decade earlier had not fully recovered. Even as of 2014, damage persisted in some locations. As previously noted, overbrowsing by White-tailed Deer also harm some of the Valley’s rare plants, including Showy Lady’s-slipper. We have seen local, severe damage to Bog Jacob’s-ladder in the Valley, but the intensity of herbivory on it appears to vary substantially across sites and among years. We have noted little American Cranberry-bush recruitment within the Valley, and suspect that it is due to White-tailed Deer browsing on young plants. Balsam Fir and Black Ash recruitment are also negatively impacted by herbivory. Canada Yew populations were apparently declining due to White-tailed Deer herbivory as early as 1952 (Rossbach 1952). White-tailed Deer have damaged populations of Glade Spurge elsewhere (Loeffler and Wegner 2000), but we have not observed such damage in the Valley. Invasive plants have also become a substantial threat to the Valley’s native plants, especially over the last 30 years (Grafton and Fortney 2015 [this issue]). Circumneutral wetlands have been most heavily impacted. Although it is native to eastern North America, Broadleaf Cattail is probably not native to the high-elevation wetlands of the Central Appalachians. In fact, it is absent or rare in relatively undisturbed high-elevation wetlands. We suspect its abundance in the Valley may have been promoted by the widespread grazing of cattle in the Valley’s wetlands during the first half of the 20th century, and it has substantially increased in abundance in the Valley since 1945 (Fortney and Rentch 2003). Broadleaf Cattail responds swiftly to the hydrologic changes caused by damming by Beavers, road construction, upslope development, and nutrient enrichment (Baldwin and Cannon 2007). We have observed Southeastern Naturalist R.L. Bartgis, E.A. Byers, R.H. Fortney, W. Grafton, and M.A. Berdine 2015 Vol. 14, Special Issue 7 169 many places in the Valley where rare plants are being encroached upon by extensive stands of Broadleaf Cattails, especially in circumneutral wetlands. These stands are often so extensive and dense that few other plants co-occur. Broadleaf Cattails also flourish in the Valley’s peatland areas. At 1 peatland site where an extensive Boradleaf Cattail infestation was carefully killed off with herbicide, Bog Death-camas became significantly more abundant within 3 years (R.L. Bartgis, pers. observ.). The non-native Iris pseudacorus L. (Yellow Flag), while more limited, is also expanding in the Valley. We are particularly concerned about 2 other invasive plant species. Since the 1970s, we have observed a notable increase in the abundance in the Valley of Phalaris arundinacea L. (Reed Canarygrass) (Grafton and Fortney 2015 [this issue]). Native to eastern North America, Reed Canarygrass has come to form large monotypic stands in the Valley, especially adjacent to disturbances and waterways. Microstegium vimineum (Trin.) A. Camus (Japanese Stiltgass), a native of Asia, was not noted in the Valley as recently as 2005. An annual, it now appears locally but sometimes abundantly along roads and trails, especially in the northeastern portion of the Valley. This is a highly aggressive species in West Virginia (WVDNR 2009) and, unlike most non-native species, thrives in shade as well as open habitats (Fryer 2011). It spreads along roads, trails, streams, and other corridors and can be dispersed by people, machines, and animals. Whitetailed Deer do not eat it and encourage its growth by preferentially feeding on other species (Fryer 2011). If not controlled, Japanese Stiltgrass has the potential to become a dominant species in moist, but not inundated, habitats, especially on limestone and alluvium. It can also suppress or displace native herbaceous plants and prevent the regeneration of woody plants (Fryer 2011). Invasive Broadleaf Cattail, Yellow Flag, and Reed Canarygrass threaten the most significant rare plant stands within Canaan Valley State Park. Other potentially harmful invasive species that are also becoming more common in the Valley are Dipsacus sylvestris Huds. (Common Teasel), Tussilago farfara L. (Coltsfoot), Elaeagnus umbellata Thunb. (Autumn Olive), and Rosa multiflora Thunb. (Multiflora Rose). Our most significant concern is climate change. Mean annual temperatures in the the Valley region may increase by 6 °F (14 °C) or more by the end of the 21st century (TNC et al. 2009). The species with primarily northern ranges are especially vulnerable. Balsam Fir, Awned Sedge, Coptis trifolia (L.) Salisb. (Threeleaf Goldthread), Cornus canadensis L. (Canadian Bunchberry), Purple Avens, American Mannagrass, Juncus filiformis L. (Thread Rush), Alderleaf Buckthorn, Dwarf Red Bramble, and American Cranberry-bush are species that we suspect are likely at or near the southern limit of their climatic tolerance in the Valley because they are rare or absent at lower elevations or farther southward (Appendix 1). The species associated with high-elevation limestone wetlands, including the globally uncommon Glade Spurge and Bog Jacob’s-ladder, may be the most vulnerable. There is little suitable habitat available in West Virginia outside of the Valley, and none at a significantly higher elevation in which they could Southeastern Naturalist R.L. Bartgis, E.A. Byers, R.H. Fortney, W. Grafton, and M.A. Berdine 2015 Vol. 14, Special Issue 7 170 find climatic refuge. In a systematic review of a limited number of species, Byers and Norris (2011) determined that Glade Spurge was extremely vulnerable and Shriver’s Frilly Orchid was highly vulnerable to the effects of climate change. The biological and ecological characteristics that place these species at such high risk are shared by many of the Valley’s rare plants. In the Valley, 19 (35%) of the plant species of concern are associated with conifer-dominated swamps. Some, such as Small’s Twayblade, have a strong affinity to these habitats. Unfortunately, these communities are highly threatened in the Valley. Only about 2% of the Valley now supports conifer swamps, and only 2% supports upland Red Spruce forests (Fortney and Rentch 2003). In addition to degradation from historic logging and fires, Red Spruce forests are in decline from the loss of two major components, Balsam Fir and Eastern Hemlock, to non-native insect pests. Adelges tsugae Annand. (Hemlock Woolly Adelgid), originally from Asia, appeared in the eastern US in the 1950s and in Tucker County in 2001 (USDA Forest Service 2001). Like the Balsam Woolly Adelgid situation, the mortality of Eastern Hemlocks from Hemlock Woolly Adelgid is substantial. Most mature trees are killed within 4–6 years of their infestation, but the mortality rate of the trees is lower in much colder climates (Paradis et al. 2008). Unfortunately, the cold temperatures required to limit the Hemlock Woolly Adelgid rarely occur in the Valley. To date, Eastern Hemlock mortality in the Valley has not been significant but is substantial in adjacent areas. A warming climate will likely benefit the Hemlock Woolly Adelgid (Paradis et al. 2008). Higher temperatures also place Red Spruce, the third and main conifer component of the Valley’s forested wetlands, at risk (Byers and Norris 2011). To increase the resilience of rare plants in the Valley to a warming climate, we strongly urge management actions that reduce other stressors such as invasive species, expand existing conifer cover (primarily Red Spruce) to assure continued conifer-swamp habitat and provide for locally cooler microclimates, and otherwise safeguard the larger and more viable rare plant populations. However, global action to reduce future warming is the only longterm strategy to ultimately keep temperatures within the thermal tolerances of the Valley’s rare plants. Our knowledge of the distribution, ecology, and status of the Valley’s rare plants remains incomplete. Because of the vast extent of the wetlands in the Valley and the remoteness of some sections, more rare plant species and populations will probably be found. For example, there is substantial potential habitat for Poa paludigena Fernald (Bog Bluegrass), a rare grass of more northern wetlands that is currently known in West Virginia only from Cranberry Glades (WVDNR 2014a). Even the most studied sites have seldom been visited throughout the growing season. There is a risk that some species or populations may be inadvertently lost from the Valley simply because their presence has not been documented when development or management decisions are made. Southeastern Naturalist R.L. Bartgis, E.A. Byers, R.H. Fortney, W. Grafton, and M.A. Berdine 2015 Vol. 14, Special Issue 7 171 Acknowledgments Brian McDonald, Sam Norris, and Ruth Thornton helped with initial field surveys, which were funded by The Nature Conservancy and the Canaan Valley Institute. James T. Anderson, Allison W. Cusick, Brian McDonald, and an anonymous reviewer provided useful comments on an early version of the manuscript. 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Byers, R.H. Fortney, W. Grafton, and M.A. Berdine 2015 Vol. 14, Special Issue 7 174 WVDN R. 2014a. Biotic natural heritage database records of rare species and natural communities. Wildlife Diversity Unit, West Virginia Division of Natural Resources, Elkins, WV. Oracle/ArcView geodatabase. WVDNR. 2014b. Plots 2-WV database of community ecology plots. Wildlife Diversity Unit, West Virginia Division of Natural Resources, Elkins, WV. MS-Access database. WVDNR. 2014c. Rare plants tracked by the West Virginia Natural Heritage Program. Wildlife Diversity Unit, West Virginia Division of Natural Resources, Elkins, WV. 10 pp. WVDNR. 2014d. Unpublished data maintained by the West Virginia Natural Heritage Program. Wildlife Diversity Unit, West Virginia Division of Natural Resources, Elkins, WV. Access and ArcGIS databases. WVDNR. 2014e. WV Curatorial Database System, maintained by West Virginia Natural Heritage Program. Label database of herbarium specimens from West Virginia University Herbarium, Youngstown State University Herbarium, Marshall University Herbarium, West Virginia Natural Heritage Program Herbarium, Davis and Elkins College Herbarium, Carnegie Museum of Natural History Herbarium, Virginia Polytechnic Institute and State University, University of South Carolina Herbarium, and the private collections of Paul J. Harmon, Scott Shriver, Al Shriver, and Clete Smith. Wildlife Diversity Unit, West Virginia Division of Natural Resources, Elkins, WV. FoxPro database. Southeastern Naturalist R.L. Bartgis, E.A. Byers, R.H. Fortney, W. Grafton, and M.A. Berdine 2015 Vol. 14, Special Issue 7 175 Southeastern Naturalist R.L. Bartgis, E.A. Byers, R.H. Fortney, W. Grafton, and M.A. Berdine 2015 Vol. 14, Special Issue 7 176 Southeastern Naturalist R.L. Bartgis, E.A. Byers, R.H. Fortney, W. Grafton, and M.A. Berdine 2015 Vol. 14, Special Issue 7 177 Southeastern Naturalist R.L. Bartgis, E.A. Byers, R.H. 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