Southeastern Naturalist R.H. Fortney, S.L. Stephenson, and J.S. Rentch 2015 Vol. 14, Special Issue 7 121 Canaan Valley & Environs 2015 Southeastern Naturalist 14(Special Issue 7):121–135 Rare Plant Communities in Canaan Valley, West Virginia Ronald H. Fortney1, Steven L. Stephenson2,3, and James S. Rentch4,* Abstract - Canaan Valley (hereafter, the Valley) is a 34,600-ac (14,000-ha), high-elevation valley in the Central Appalachian Mountains of West Virginia. Its diverse wetland and upland habitats support a wide variety of plant communities, many of which are extremely rare. The prominence of rare communities is associated with the diversity of topographic settings, soils, geology, and hydrology, as well as the effects of human settlement and resource exploitation. Most of the rare plant communities are found in the wetlands of the Valley’s floor. Virtually all of the communities associated with the Valley’s extensive cold peatlands are rare, including (1) mixed conifer swamp-forests of Picea rubens (Red Spruce), Abies balsamea (Balsam Fir), and Tsuga canadensis (Eastern Hemlock), (2) mixed conifer-Fraxinus nigra (Black Ash) bog-forests in limestone-influenced wetlands in the central and southern parts of the Valley, and (3) extensive Sphagnum and Polytrichum bogs in the central and northern parts of Canaan Valley. Shrub communities such as Alnus incana ssp. rugosa (Speckled Alder), Viburnum recognitum (Smooth Arrowwood), and Salix discolor (Glaucous Willow) growing on mineral soils along waterways are also rare. Populus tremuloides (Trembling Aspen) groves, although abundant in the Valley, are extremely limited in the Appalachian region. Lastly, the grassand forb-dominated grass-bald communities on the surrounding mountain rims show an extremely limited distribution throughout the Central Appalachians. Introduction Canaan Valley (hereafter, the Valley), sited in southeastern Tucker County of north-central West Virginia, is an elongated, eroded, anticlinal valley in the Allegheny Mountain Section of the Appalachian Plateau Physiographic Province (Fenneman 1938). The Valley is 11.4 mi (19 km) long, ranges from 1.2 to 2.4 mi (2 to 4 km) wide, and covers approximately 35,000 ac (14,000 ha). Elevation ranges from 4528 ft (1372 m) on Cabin Mountain on the east to 3300 ft (1006 m) at the egress of the Blackwater River between Canaan and Brown mountains to the west. From an ecological point of view, the Valley has four principal distinguishing features. First, much of its flora is characterized as northern, interpreted as a remnant of the influence of a Pleistocene periglacial climate (Allard and Leonard 1951, Fortney 1975, Fortney and Rentch 2003, Goudy et al. 1969, Vogelmann 1978). Second, the Valley holds the largest inland freshwater wetland in the eastern United States, excluding glaciated regions to the north and the Atlantic 1Department of Civil and Environmental Engineering, PO Box 6103, West Virginia University, Morgantown, WV 26506 (deceased). 2Department of Biological Sciences, Fairmont State College, Fairmont WV 26554. 3Current address - Department of Biological Sciences, University of Arkansas, Fayetteville AR 72701. 4Division of Forestry and Natural Resources, PO Box 6125, West Virginia University, Morgantown, WV 26506. *Corresponding author - jrentch2@wvu.edu. Southeastern Naturalist R.H. Fortney, S.L. Stephenson, and J.S. Rentch 2015 Vol. 14, Special Issue 7 122 Coastal Plain to the east and south (Vogelmann 1978). Third, its once prominent Picea rubens Sarg. (Red Spruce) forest was one of the best developed of this type in the United States, producing up to 45,000 board ft/ac (262 m3/ha) (Brooks 1957). Even though the logging of the original Red Spruce forest was not unique to the Valley (e.g., Clarkson 1964), the events here may represent one of the more extreme examples of logging impacts and post-logging fires. And fourth, although the vegetation and habitats of the Valley are generally typical of highelevation ecosystems in the Central Appalachians, vegetation maps (Fortney and Rentch 2003) show high levels of plant community diversity and of spatial complexity, including a complex juxtaposition of upland and wetland habitats. Given the ecological distinctiveness of its vegetation, the identification of rare and otherwise unusual plant communities is important for managing the Valley’s public and private lands. A general goal of this paper is to identify and describe the Valley’s rare plant communities. We also have three more specific objectives: 1) identify rare or otherwise unusual plant communities, describing their distributions in the Valley; 2) list rare plant species associated with specific rare plant communities; and 3) compare the incidence of rare or otherwise significant plant communities in the Valley with those of other high-elevation wetland complexes in the Allegheny Mountain Section of the Appalachian Plateau Physiographic Province in West Virginia. Study Area Most descriptions of the presettlement, pre-logging vegetation suggest that while some graminoid-dominated openings existed on the Valley floor, the primary cover was mixed Red Spruce-Tsuga canadensis (L.) Carr. (Eastern Hemlock) forest. Red Spruce tended to be more predominant on the surrounding mountain slopes and ridgetops where hemlock was less common. After the railroad-catalyzed logging of the late 1800s through early 1900s, much of the Valley and surrounding areas burned repeatedly, creating a distinctive pyric vegetation association of Populus tremuloides Michx. (Trembling Aspen), P. grandidentata Michx. (Big-tooth Aspen), and Prunus pensylvanica L. (Fire Cherry) that persisted in many areas for several decades (Allard and Leonard 1951). Severe wind and water erosion of topsoil added to the degradation, particularly on upland sites. In some upland areas, only boulder fields remain and they support extremely sparse plant communities. During the 1950s, parts of the Valley were temporarily converted to row crop and grazing lands. Since the 1970s, large areas in the southern end of the Valley have been converted to second-home developments and recreational areas. Concurrent with the decline of agricultural land uses and the increase in recreational development, but not necessarily in response to these changes, resident populations of Castor canadensis Kuhl (Beaver) and Odocoileus virginianus Zimmermann (White-tailed Deer) increased dramatically, further impacting the natural vegetation (Bonner et al. 2009, Michael 1992). Today, the Valley’s vegetation is a mosaic of open and forested habitats, consisting of both wetland and Southeastern Naturalist R.H. Fortney, S.L. Stephenson, and J.S. Rentch 2015 Vol. 14, Special Issue 7 123 upland cover types. Even with these past and present anthropogenic changes, the Valley’s vegetation has made a remarkable recovery, recapturing a high degree of naturalness in composition and structure. Its current vegetative cover is a combination of indigenous species and those introduced since the logging era ended. The vast wetland area on the valley floor distinguishes the Valley from other high-elevation wetlands in the Appalachian Plateau (Volgelmann 1978). The diverse assemblage of bogs, wet meadows, and swamps totals about 7400 ac (3000 ha) (Fortney and Rentch 2003). Forested wetland communities are largely restricted to small, scattered stands along streams running through the Valley. According to Fortney and Rentch (2003), the most extensive forested wetland communities are the mixed conifer swamp stands along streams and low areas, and the Trembling Aspen groves on poorly drained areas of the Valley. Extensive Spiraea alba DuRoi (Spiraea) and Hypericum densiflorum Pursh (Glade St. Johnswort) shrub thickets occur on the Valley’s poorly drained mineral soils. Alnus incana ssp. rugosa (DuRoi) Spreng. (Speckled Alder) shrub thickets are also extensive, mostly occurring on the floodplains of streams flowing through both mineral and organic soils. Vaccinium-Photinia (blueberry-chokeberry) shrub communities grow on soils with high organic content, but can occasionally develop on mineral soils. The shrub communities, particularly on organic soils, often have a nearly continuous bryophyte groundcover of Sphagnum spp. (sphagnum mosses). and Polytrichum spp. (haircap mosses). Communities dominated by herbaceous vegetation are highly variable. Floodplain communities range from nearly monotypic examples to those with diverse mixtures of sedges and grasses, and better-drained sites support mixed grass-forb communities. On organic soils, a continuous groundcover of bryophytes is typical, with other characteristic plants present, including sedges, rushes, bulrushes, and forbs like Solidago uliginosa Nutt. (Bog Goldenrod). The remaining better-drained upland areas of the Valley are old-field habitats vegetated chiefly by combinations of forbs, grasses, and ferns. Upland forest communities are predominantly northern hardwoods, with Acer saccharum Marsh. (Sugar Maple), A. rubrum L. (Red Maple), Fagus grandifolia Ehrh. (American Beech), and Prunus serotina Ehrh. (Black Cherry) as the more prominent species. Stands of Red Spruce are now found only in small areas on the upper slopes of the surrounding mountains and in sheltered coves; it is also intermixed with other conifers in isolated forested wetlands on the Valley floor. Today’s Valley vegatation consists of remnants of the original flora, species that have become established since the destruction of the original Red Spruce forest, and those introduced by humans. A large portion of the flora is composed of species with distinctly northern ranges (Fortney 1975). For example, Red Spruce and Abies balsamea (L.) Mill. (Balsam Fir), two boreal species typical of the original forest, are still common. Other boreal species include shrubs such as Viburnum opulus L. var. americanum Ait. (Highbush Cranberry) and Photinia melanocarpa (Michx.) K.R. Robertson & Phipps (Black Chokeberry), and herbs such as Chrysosplenium americanum Schwein ex Hook. (Golden Saxifrage) Southeastern Naturalist R.H. Fortney, S.L. Stephenson, and J.S. Rentch 2015 Vol. 14, Special Issue 7 124 and Eriophorum virginicum L. (Cottongrass). Plant diversity in the Valley is amplified by the presence of species with a Southern Appalachian distribution, including Vaccinium erythrocarpum Michx. (Southern Mountain Cranberry) and Euphorbia purpurea (Raf.) Fernald (Glade Spurge). The Valley’s bedrock geology has strongly influenced plant community development. The erosion-resistant Pottsville Sandstone caps much of the surrounding mountains, while softer, more erodible Mauch Chunk shales underlay the mountain slopes below the sandstone. Much of the Valley floor is underlain by Greenbrier Limestone. The erodibility of the Greenbrier Limestone produces a low-relief, poorly-drained watershed where saturation and inundation are common. Most of the wetlands in the Valley have developed over this bedrock. Furthermore, alkalinity from the limestone has influenced the distribution of plant species and plant communities. For example, patches of calciophiles, such as Fraxinus nigra Marsh. (Black Ash) and Polemonium van-bruntiae Britton (Jacob’s Ladder), overlay the Greenbrier Limestone. The types of soils have also affected the development of the Valley’s plant communities. Extensive areas of muck and peat soils are found in the Valley, with the largest and deepest occurring in central and northern sections (USDA 1960). Cameron (1970) reported peat depths of 14.1 ft (4.3 m) near the northern edge of Middle Ridge. Soils with histic-epipedons (Blago Series), and an organic surface layer (11.8 in [< 30 cm] thick) that overlays mineral strata, are extensive. These soils frequently host a vegetative cover of haircap mosses and an herb layer with mixtures of forbs and/or a shrub layer of blueberries and Viburnum spp. (viburnums). Upland soils, in contrast, appear to be less variable, and therefore, upland vegetative cover is more homogeneous. Methods This paper reviews vegetation studies conducted in the Valley and compares its plant communities with those of other high-elevation wetlands (>2800 ft [850 m]) in West Virginia. We accept Fortney (1975) as the authority for identifying and describing the distribution and composition of the Valley’s rare plant communities. We also use several more recent studies (Chambers 1996, Fortney 1997, Fortney and Rentch 2003, Rentch and Fortney 1997, and Stephenson and Adams 1986) and additional unpublished field data collected during 2001–2002. Species nomenclature follows Harmon et al. (2006). For a regional perspective, we review studies of other high-elevation areas in West Virginia that hold significant wetland habitats. These include Robinette (1964) for Cranesville Swamp; Darlington (1942), Edens (1973), and Kokesh (1988) for Cranberry Glades; Muzika et al. (1996) and Wieder et al. (1981) for Big Run Bog; and Walbridge and Lang (1982) for four relatively small wetlands in the Allegheny Mountain Section. To be scored as rare in our assessment, a plant community met at least one of the following three criteria: it (1) contained at least one dominant or co-dominant Southeastern Naturalist R.H. Fortney, S.L. Stephenson, and J.S. Rentch 2015 Vol. 14, Special Issue 7 125 species with a limited distribution in the Allegheny Mountain Section of West Virginia; (2) occurred in a habitat type with a limited or restricted distribution in the Allegheny Mountain Section (e.g., a wetland or grass bald); or (3) may have been common, but it typically supported one or more rare plant species. In addition, because wetland area throughout the Unglaciated Plateau is limited, communities that occur in wetlands are, by definition, more rare than those on uplands. Although our determination of rarity for plant communities was somewhat subjective, we based it on firsthand knowledge of the vegetation of the Valley and adjacent high-elevation areas, and on a comparison with plant communities in other wetlands of the region. In most instances, the name of a plant community is based on the dominant species in its uppermost stratum. Exceptions to this are communities with extensive bryophyte groundcover associated with organic soils. Operationally, to be scored as dominant, a species had to be: a) cited as a component of a community based on empirically derived data; b) included in a study where stratified sampling occurred using line transects, intercepts, or quadrats; or c) considered a dominant based on a calculation of some index of importance, dominance, or occurrence. Results and Discussion Fortney (1975) described 42 plant communities in the Valley, 32 of which were classified as wetlands and 10 as uplands. All of the wetland plant communities, except for those dominated by Spiraea, Glade St. Johnswort, and Typha latifolia L. (Broad-leaved Cattail), met at least one of the three criteria for rare plant communities, and therefore, we considered them to be rare. Further, based on recent fieldwork we added two additional communities, the Salix-tall shrub thicket and mixed conifer-Black Ash swamp-forest, to the list of rare communities. Four upland communities were designated as rare: (1) grass balds and (2) heath balds on exposed mountain summits, (3) Bigtooth Aspen groves, and (4) Black Cherry groves (See Table 1). The larger number of wetland vs. upland communities is likely related to the rich mosaic of soil types and hydrologic regimes in wetland areas (Fortney and Rentch 2003). The vegetation (Fortney 1997) and soils (Losche and Beverage 1967) maps of the Valley support this suggestion, showing that upland areas tend to have fewer soil types and vegetation units, each with a more extensive distribution than those of wetland areas. Further, the juxtaposition of wetland units, soil series, and vegetation types tends to be more complex in terms of variability per unit area and boundary conditions. The occurrence of rare and otherwise unusual plant communities is consistent with this pattern. Upland plant communities In this section, we outline the general features, resident plant species, and species associations for each of the Valley’s rare upland plant communities. Mixed grass-forb balds and low shrub bald. The region’s early explorers described mountain summits as open and grassy. According to Fortney (1975), Southeastern Naturalist R.H. Fortney, S.L. Stephenson, and J.S. Rentch 2015 Vol. 14, Special Issue 7 126 the area covered by these habitats likely expanded after the logging of the late 1800s through early 1900s, and subsequent fires (Allard and Leonard 1951). Core (1966) and Fortney (1975) described balds as open, grassy communities often dominated by Danthonia compressa Austin (Mountain Oat Grass). However, Rentch and Fortney (1997) concluded that the balds’ vegetation had changed since Fortney’s study (1975), becoming more varied, with an increase in the Table 1. Plant communities in Canaan Valley considered rare on a regional scale, based on studies by Fortney (1975), Rentch and Fortney (1997) and Rentch et al. 2013 [this issue]. * = occurs mostly on mineral soils, ** = occurs on either mineral or organic soils, *** = occurs mostly on organic soils, **** = may occur in mountain bald habitats. Upland communities Mixed grass-forb bald**** Low shrub thicket**** Populus grandidenta grove Prunus serotina grove Wetland swamp forest communities Picea-Tsuga-Abies swamp** Picea-Tsuga-Abies-Betula-Acer (Fraxinus ) swamp** Picea rubens swamp** Abies balsamea swamp** Populus tremuloides grove * Wetland shrub communities Salix-tall shrub-thicket* Alnus incana ssp. rugosa tall shrub thicket ** Low shrub thicket** Vaccinium-Photinia low shrub thicket ** Vaccinium-Photinia hummock bog*** Vaccinium-Polytrichum hummock bog*** Wetland herbaceous communities Sphagnum-Carex bog*** Sphagnum-Glyceria-Carex bog*** Sphagnum-Eriophorum bog*** Polytrichum hummock bog*** Polytrichum-Rubus hummock bog*** Polytrichum-Solidago hummock bog*** Polytrichum-Solidago-Rubus hummock bog*** Polytrichum-Eriophorum hummock bog*** Polytrichum-Glyceria hummock bog*** Polytrichum-Juncus hummock bog*** Polytrichum-Juncus-Carex-hummock-bog*** Polytrichum-Carex hummock bog*** Calamagrostis Canadensis wet meadow* Carex-Leersia-Calamagrostis-Glyceria wet meadow* Carex-Scirpus-Solidago-aster wet meadow* Carex-Solidago-Rubus wet meadow* Scirpus wet meadow/marsh* Juncus wet meadow/marsh* Carex wet meadow/marsh* Southeastern Naturalist R.H. Fortney, S.L. Stephenson, and J.S. Rentch 2015 Vol. 14, Special Issue 7 127 importance of shrubs, forbs, and Deschampsia flexuosa (L.) Trin. (Wrinkled Hairgrass). They identified Solidago rugosa Mill. (Wrinkled-leaf Goldenrod), Thelypteris noveboracensis (L.) Nieuwl. (New York Fern), Rubus canadensis L. (Blackberry), Vaccinium pallidum Ait. (Upland Low Blueberry), and V. angustifolium Ait. (Lowbush Blueberry) as additional common codominants. Rentch and Fortney (1997) further concluded that the balds are maintained in an open state by climatic factors, including extreme temperatures and physical damage to plants by wind, ice, and snow—factors that reduce the likelihood of tree establishment. Because of these factors, bald composition will likely remain stable for the foreseeable future, with combinations of grasses, forbs, and shrubs. Balds and their associated vegetation types are rare in the Central Appalachian Mountain region because they are restricted to only a few areas. Populus grandidentata grove-forest. Groves of Big-tooth Aspen are mainly restricted to well-drained Calvin and Dekalb soils at the bases of the Valley’s surrounding mountain slopes. These temporal and early successional communities have largely been replaced by an encroaching northern hardwood forest. The herbaceous stratum is often composed of old-field graminoids and forbs, such as Wrinkled-leaf Goldenrod, Dennstaedtia punctilobula (Michx.) Moore (Hayscented Fern), Pteridium aquilinum (L.) Kuhn (Bracken Fern), and Mountain Oat Grass. The shrub layer is generally poorly developed or absent. Prunus serotina grove-forest. Black Cherry was the Valley’s first tree species to be commercially exploited. Several groves of Black Cherry remain near Camp 72 and Big Cove. Found on well-drained areas that are adjacent to wetlands and along the bases of the surrounding mountains, this community type often occurs as pure stands or with a small component of Red Maple. Shrubs are usually lacking, and the herbaceous stratum is frequently comprised by large colonies of Hay-scented and New York ferns. Wetland plant communities Four conifer-dominated or mixed conifer-deciduous communities that occur in the Valley are regionally rare. They can develop on mineral or organic soils, and on soils with histic-epipedons (Blago Series). A bryophyte layer of Sphagnum spp. mosses is typically present on sites with organic soils. Picea-Tsuga-Abies swamp. Compared to presettlement estimates of the total area of this community type, today’s mixed conifer swamp-forests of Red Spruce, Eastern Hemlock, and Balsam Fir represent a small portion of the Valley’s vegetative cover. Over the past four decades, much of this forest type (as well as subtypes in which Red Maple and Betula allegheniensis Britt. [Yellow Birch] are components) has been innundated or degraded by the actions of Beavers. This is particularly apparent within Canaan Valley State Park and Big Cove. Associated canopy species include Black Cherry and Ilex montana Torr. & Gray (Mountain Holly). Picea-Tsuga-Abies-Betula-Acer (Fraxinus) swamp. We consider this plant community to be a subtype of the spruce-hemlock-fir swamp-forest described Southeastern Naturalist R.H. Fortney, S.L. Stephenson, and J.S. Rentch 2015 Vol. 14, Special Issue 7 128 above. The spruce-hemlock-fir-maple-birch (ash) swamp-forest is generally associated with other conifer-dominated communities, but it has a hardwood component typically represented by Red Maple and Yellow Birch. Black Cherry and Mountain Holly occasionally occur in the canopy, although their presence is unusual; Black Ash can be a principal component of the overstory. This community type, identified along Abe’s Run (Rentch et al. 2013 [this issue]) and the North Branch, appears to be associated with limestone. Several rare plants, including Glade Spurge, Rhamnus alnifolia L’Her. (Alder-leaved Buckthorn), Jacob’s Ladder, and Cypripedium reginae Walt. (Showy Lady’s Slipper), have been found in this community type. Picea rubens swamp. Although not frequent, Red Spruce can occur as the single, dominant species in a forest canopy. The Red Spruce swamp-forest is often associated with spruce-hemlock-fir swamp-forest. Abies balsamea swamp. We score this forest type as occasional throughout the Valley, where it is associated with the three wetland forest communities described above. Balsam Fir swamp-forest usually occurs in small stands (less than 2.5 ac [1 ha]). Infestations of Adelges piceae (Ratzeburg) (Balsam Woolly Adelgid) have reduced Balsam Fir in the overstory of most coniferous stands, particularly in the central and southern Valley. Between 1983 and 2002, a combination of adelgid mortality and inundation by beaver extirpated Balsam Fir from three stands—two in Canaan Valley State Park and one in the northern end of the Valley—where it had been a dominant or codominant component of the overstory (Fortney, unpubl. data). In addition, because deer overbrowse Balsam Fir seedlings, there has been virtually no recruitment. In combination, these stresses threaten the recovery and ultimate survival of this community type. Populus tremuloides grove. Trembling Aspen groves commonly develop in the Valley’s open areas where there is saturated mineral soil. Their most extensive growth occurs along the North Branch of the Blackwater River and areas adjacent to the Little Blackwater River and Glade Run. Some groves have become established in haircap moss bogs, but these stands tend to be short-lived due to nutrient shortages, competition, and cold injury (Fortney 1975). Available age data suggest that Trembling Aspen grove-forests are even-aged and clonal, i.e., all of the stems within a patch carry identical genetic material (Rentch and Anderson 2008). Understory species vary considerably along hydrologic gradients. Shrubs include Viburnum nudum L. (Wild Raisin), V. recognitum Fernald (Smooth Arrowwood), Vaccinium myrtilloides Michx. (Velvet-leaved Blueberry), and Spiraea. Common groundcover species include Rubus hispidus L. (Swamp Dewberry), Bog Goldenrod, Mountain Oat Grass, and Euthamia graminifolia (L.) Nutt. var graminifolia (Grass-leaved Goldenrod). Like the forest communities, each of the six shrub communities can occur on mineral or organic soils. On organic soils, shrub communities typically support a bryophyte layer of Sphagnum spp. and haircap mosses. Alnus incana ssp. rugosa-tall shrub thicket. Speckled Alder forms extensive shrub thickets along most of the watercourses of the Valley floor and along older Southeastern Naturalist R.H. Fortney, S.L. Stephenson, and J.S. Rentch 2015 Vol. 14, Special Issue 7 129 oxbows. Some of these communities are old, and are identifiable from aerial photographs taken in1945. The water table under alder thickets is usually at or near the ground’s surface for most of the growing season. Associated shrubs include Glade St. Johnswort, Spiraea, Ilex verticillata (L.) Gray (Winterberry), and Smooth Arrowwood. The speckled alder-tall shrub thicket is one of the most species-rich plant communities of the Valley (Fortney 1975). Its high biodiversity is probably related to the semi-open canopy of the shrub layer that allows filtered light, availability of abundant nutrients (these communities receive overbank flow, and alder is a nitrogen-fixer), and the microtopographic variability of the alternating hummocks around the bases of alder clumps. The following rare species have been identified in this community: Thelypteris simulata (Davenport) Nieuwl. (Bog Fern), Glyceria grandis S. Wats. (American Mannagrass), Jacob’s Ladder, Hasteola suaveolens (L.) Pojark. (Sweet Indian Plantain), Scirpus atrocinctus Fernald (Black-girdled Wool-grass), and Equisetum fluviatile L. (Water Horsetail). Salix-tall shrub thicket. The willow thicket was not mentioned by Fortney (1975), but in the intervening years this community type has become more common. The willow-tall shrub thicket can include three species of willow. Salix sericea Marsh. (Silky Willow), the most common species, occurs in wet ditches and along the margins of beaver ponds. Associated with Speckled Alder, S. discolor Muhl. (Glaucous Willow) is a relatively rare species known from small communities along Abe’s and Glade Runs. Salix humilis Marsh. (Upland Willow) is ostensibly an upland species that grows in wetland areas of Big Cove. Vaccinium-Photinia-low shrub thicket and Vaccinium-Polytrichum bog. These two community types are often continuous with each other. They can occur on mineral soil or soil with a high organic content, and on hummocky surfaces with a nearly continuous groundcover of bryophytes. Blueberry- and chokeberrydominated bogs occur throughout the Valley, but they are best developed and most extensive in the Valley’s central and northern sections. The most common blueberries are Lowbush and Velvet-leaf; Black Chokeberry is often dominant or co-dominant. The principal viburnums are Wild Raisin and Smooth Viburnum. Also, Nemopanthus mucronata (L.) Loes. (Wild Holly) is a commonly associated shrub. Haircap mosses form the prominent groundcover layer in shrub-dominated bogs, and Dewberry typically forms a creeping layer. Less frequently, rare species such as Vaccinium oxycoccos L. and V. macrocarpon Ait. (Small and Large Cranberries, respectively) and Gaultheria hispidula (L.) Muhl. ex Bigelow (Snowberry) can occupy the surface layer. Herbaceous plant communities are mostly associated with muck and peat and with Blago soils. These herbaceous communities, along with the two aforementioned shrub communities, were previously classified as bogs or muskegs (e.g., Allard and Leonard 1951, Fortney 1975). However, based on the form of nutrient and water inputs, as well as the availability of soil nutrients (Chambers 1996, R. H. Fortney unpubl.), we conclude that these wetlands are more appropriately classified as fens. Like shrub bogs, these community types occur throughout the Valley, often forming extensive communities. Southeastern Naturalist R.H. Fortney, S.L. Stephenson, and J.S. Rentch 2015 Vol. 14, Special Issue 7 130 Sphagnum-Polytrichum bogs. In these communities, Sphagnum spp. or haircap mosses typically form a nearly continuous layer of groundcover. Sphagnum- haircap moss bog communities are rare, and among the least diverse in the Valley. The plants that emerge above the moss layer are highly variable and rarely occur in large numbers, although there are several species that occur either as associates or have a high fidelity to these moss bogs. The most important graminoid species include Carex utriculata Boott (Northwest Territory Sedge), C. stricta Lam. (Upright Sedge), C. scoparia Schkuhr ex Willd. (Broom Sedge), Cottongrass, Juncus effusus L. (Common Rush), J. filiformis L. (Thread Rush), and Scirpus microcarpus J. Presl & C. Presl (Bulrush). Important forb species are Drosera rotundifolia L. (Round-leaved Sundew), Eupatorium pilosum Walt. (Vervain Thoroughwort), Osmunda cinnamomea L. (Cinnamon Fern), Snowberry, Zigadenus leimanthoides Gray (Oceanorus), and Bog Goldenrod. Occasional shrubs and woody creepers are Wild Holly, Black Chokeberry, Velvetleaf Blueberry, Wild Raisin, Large and Small Cranberries, and Dewberry. As with shrub-bogs, hummocks often characterize these communities. The most common herb-dominated moss-bog communities include the Sphagnum-Carex bog, Sphagnum-Glyceria-Carex bog, Sphagnum-Eriophorum bog, Polytrichum hummock- bog, Polytrichum-Rubus hummock-bog, Polytrichum-Solidago hummockbog, Polytrichum-Solidago-Rubus hummock-bog, Polytrichum-Eriophorum hummock-bog, Polytrichum-Glyceria hummock-bog, Polytrichum-Juncus hummock-bog, Polytrichum-Juncus-Carex hummock-bog, and the Polytrichum- Carex hummock-bog. Graminoid- and forb-dominanted wetlands are common on mineral soils throughout the Valley. They are characteristically associated with the floodplains of streams that are or have been flooded by beaver ponds. They are also associated with old-field habitats that have seeps, and sites that experience seasonally high water tables. The species composition of these wetland communities is highly variable. Nearly monotypic communities are almost as common as those with two or more codominant species. In general, graminoid species are more important where the water table is persistently high or where overbank flow is frequent, whereas forbs are more common on better-drained sites. Carex marsh/wet meadow. This community type, a variant of the graminoid meadow, is common throughout the Valley. The most frequent sedges are Carex folliculata L. (Northern Long Sedge), Broom Sedge, C. vulpinoidea Michx. (Fox Sedge), Upright Sedge, and Northwest Territory Sedge. The latter two species often form nearly circular monotypic colonies that are clearly visible in aerial photographs. Several large colonies of the rare sedge C. cannescens L. (Silvery Sedge) occur in the Valley center, and a C. atherodes Spreng. (Wheat Sedge) colony occurs on Glade Run. Associated species are those listed for other wet meadow types. The sedge marsh/wet meadow community is typically associated with floodprone areas of streams and the edges of beaver ponds. Juncus marsh/wet meadow. This community type is usually dominated by Common Rush, a species that seems to be associated with recently disturbed habitats, Southeastern Naturalist R.H. Fortney, S.L. Stephenson, and J.S. Rentch 2015 Vol. 14, Special Issue 7 131 particularly streamsides impacted by ATV activity. Common Rush is also frequently a dominant emergent species in bogs with extensive bryophyte groundcover. Mixed graminoid-forb wet meadows. This highly variable mixture of common graminoid and forb species occurs on a variety of mineral soils and under different hydrologic regimes. It may occur in large expanses near streams and rivers, in abandoned fields with seasonally high water tables, or as meadows that occur as an intermediate wetland type between uplands and shrub- or forested wetlands. In general, forbs are more important on the drier sites. The most common graminoids include the grasses American Mannagrass, Leersia oryzoides (L.) Sweet (Rice Cut-grass) Agrostis gigantea Roth (Redtop) and Calamagrostis canadensis (Michx.) P. Beauv. (Bluejoint Grass); the sedges Northern Long Sedge, C. lurida Wahlenb. (Sallow Sedge), Broom Sedge, Upright Sedge, Fox Sedge, and C. gynandra Schwein. (Nodding Sedge); the bulrushes Scirpus atrovirens Willd. (Dark Green Bulrush), and Black-girdled Wool-grass; and the rushes Common Rush and Juncus subcaudatus (Engelm.) Coville and Blake (Woodland Rush). Prominent forbs and ferns include Galium obtusum Bigelow (Bluntleaf Bedstraw), Swamp Dewberry, Bog Goldenrod, Grass-leaved Goldenrod, Doellingeria umbellata (P. Mill.) Nees var. umbellata (Flat-topped White Aster), Thelypteris palustris Schott (Marsh Fern), and Cinnamon Fern. The following common assemblages of graminoids and forbs occur as three definable communities: Carex-Leersia-Calamagrostis-Glyceria wet meadow, Carex-Scirpus-Solidagoaster wet meadow, and Carex-Solidago-Rubus wet meadow. Graminoid wet meadow/marshes. This type is one of the few wetland graminoid communities for which we have evidence of a pre-logging-era occurrence in the Valley—some Bluejoint Grass communities were cut for hay by the Valley’s early settlers (Core 1966). The graminoid wet meadow/marsh type is similar to the graminoid-forb meadow, but differs in that it has a clear dominance of one or more graminoid species, and it generally develops under wetter hydrologic regimes than does the mixed graminoid-forb wet meadow. Also, graminoid wet meadow/marsh communities are often associated with disturbed streamsides with overbank flooding, and sites where Beaver activity has raised the water table. Wet meadows are often dominated by Bluejoint grass, Rice Cutgrass, or a combination of many of the same sedges and grasses listed for the mixed graminoid-forb wet meadow. Bluejoint Grass typically dominates meadows along streams with frequent overbank flow as well as old beaver ponds. Associated species are the same as those found in mixed graminoid-forb wet meadows. Several rare species (e.g., American Mannagrass and Veronica scutellata L. [Marsh Speedwell]) are typically found in these habitats. The following three common associations of graminoids and forbs often occur as definable communities—bluejoint grass wet meadow, rice cutgrass wet meadow/ marsh, and bulrush wet meadow/marsh. Southeastern Naturalist R.H. Fortney, S.L. Stephenson, and J.S. Rentch 2015 Vol. 14, Special Issue 7 132 Comparison of the region’s wetland plant communities The Valley’s rare wetland plant communities are not unlike those of other high-elevation wetlands in the Allegheny Mountain Section. Tables 2 and 3 list the major plant communities and dominant plant species of Cranesville Swamp (Robinette 1964), Cranberry Glades (Edens 1973, Kokesh 1988), Big Run Bog (Muzika et al. 1996, Weider et al. 1981), and Cupp Run and Laurel Run (Walbridge and Lang 1982). The most obvious differences among these six places are the presence of a species in only one area and the large number of communities in the Valley compared to the other sites. Balsam Fir is present only in the Valley; Larix laricina (Du Roi) K. Koch (Larch) grows only in Cranesville Swamp. Twenty-six of the 28 wetland community types and 31 of 32 of the region’s community dominants grow in the Valley. Table 2. Plant communities cited in studies of six high-elevation wetlands in the Allegheny Mountain region of West Virginia. See text for study citations. P = present, TST = tall shrub-thicket, LST = low shrub-thicket, WM = wet meadow). Big Canaan Cranesville Cranberry Run Cupp Laurel Valley Swamp Glades Bog Run Bog Forest communities Mixed-conifer hardwood P P P P P P Picea rubens P P P P Abies-Picea-Tsuga P Larix-Picea-Tsuga P Mixed conifer-Fraxinus-Betula P P Tsuga canadensis swamp P P P P Populus tremuloides grove P Shrub Communities Alnus incana TST P P P P P Spiraea shrub thicket P P P Vaccinium-Photinia LST P P P P Hypericum LST P P Salix thicket P P Herbaceous Communities Sphagnum-Carex bog P P P P P P Sphagnum-Glyceria-Carex bog P Sphagnum-Eriophorum bog P P Sphagnum-Dulichium bog P P Polytrichum-Solidago-Rubus bog P P P Polytrichum-Eriophorum bog P P P Polytrichum-Carex-Juncus bog P P P Polytrichum-Solidago-Eriophorum bog P P P P Mixed Graminoid Forb WM P Juncus-Carex WM P Calamagrostis canadensis WM P Typha-Osmunda WM P P P P Typha marsh P P Leersia marsh P P P P P P Totals 26 17 11 12 8 10 Southeastern Naturalist R.H. Fortney, S.L. Stephenson, and J.S. Rentch 2015 Vol. 14, Special Issue 7 133 The Valley’s many types of plant communities are related in part to its large area. The Valley is five to 20 times bigger than each of the other areas, so more communities and species would be expected there. The diversity and complexity of the Valley’s habitats are also notable. The Valley’s large number of habitats reflects (1) the juxtaposition of acid sandstones and alkaline limestone, (2) the Table 3. Dominant plant species cited in studies of six high-elevation wetlands in the Allegheny Mountain region of West Virginia. See text for study citations. P = present. Big Canaan Cranesville Cranberry Run Cupp Laurel Valley Swamp Glades Bog Run Bog Tree Species Tsuga canadensis P P P P P P Picea rubens P P P P Betula allegheniensis P P P P Prunus serotina P Acer rubrum P P P Abies balsamea P Fraxinus nigra P P Larix laricina P Shrub Species Alnus incana ssp. rugosa P P P P Spiraea alba P Hypericum densiflorum P P P P P P Ilex verticillata P P P Viburnum spp. P P P P P Vaccinum spp. P P P P Photinia melanocarpa P P P P P Salix spp. P P P P Herbaceous Species Carex utriculata P P P Carex stricta (Upright Sedge) P P Carex canescens (Silver Sedge) P P P P Carex folliculata P P P P P P Dulichium arundinaceum (L.) Britton P P (Threeway Sedge) Eriophorum virginicum P P P Rhynchospora alba L. (White Beak-sedge) P P P Juncus effusus P P P P Juncus brevicaudatus (Engelm.) Fernald P P (Narrow-panicle Rush) Juncus subcaudatus (Woodland Rush) P P P Glyceria spp. P P Rubus hispidus P P P P P Solidago uliginosa P P P P P Calamagrostis canadensis P Scirpus spp. P Osmunda cinnamomea P P P Totals 31 22 13 17 11 9 Southeastern Naturalist R.H. Fortney, S.L. Stephenson, and J.S. Rentch 2015 Vol. 14, Special Issue 7 134 rock structure of the eroded anticline, and (3) the pattern of human disturbances. Overall, there are many similarities among the plant communities of these six areas, but the Valley is unique in that it supports examples of almost all of the communities found in the 5 other areas, as well as the majority of the dominant species listed for any of the other wetlands. Compared to other places in the Allegheny Mountain Section and nearby montane areas with extensive wetland habitats, such as Cranberry Glades, the Valley supports an extraordinary array of plant communities. A large percentage of the Valley’s community types are considered rare in the Allegheny Mountain Section, largely because they are wetland communities and wetland habitats are limited in this region. Another habitat-related factor is the presence of Greenbrier Limestone as the surface rock layer, which adds to habitat diversity. Also contributing to rarity are the many northern plant species that are at or near the southernmost limits of their distributions. Lastly, the Valley’s numerous rare plant communities may reflect the level of detail at which the plant communities have been studied. Management Implications The rich diversity of the Valley’s plant communities, and the many regionally or globally rare plant species deserve special consideration by the Valley’s land managers. Management goals should be developed to sustain this diversity. For example, we recommend that management practices target keystone species, like White-tailed Deer and Beaver, which can directly and indirectly impact rare plant communities, positively as well as negatively. Literature Cited Allard, H.A., and E.C. Leonard. 1952. The Canaan and the Stony River valleys of West Virginia, their former magnificent spruce forests, their vegetation, and floristics today. Castanea 17:1–60. Bonner, J.L., J.T. Anderson, J.S. Rentch, and W.N. Grafton. 2009. Vegetative composition and community structure associated with beaver ponds in Canaan Valley, West Virginia, USA. Wetlands Ecology and Management 17: 543–554. Brooks, M.G. 1957. Canaan Valley. West Virginia Conservation 21:7–10. Cameron, C.C. 1970. Peat resources of the unglaciated uplands along the Allegheny structural front in West Virginia, Maryland, and Pennsylvania. US Geological Survey, Professional Paper 700-D:D153-D161. Chambers, D.B. 1996. 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