CASTANEA 69(2): 116–124. JUNE 2004

Vascular Flora of a Southern Appalachian Fen and Floodplain Complex

1 1 2 ROBERT J. WARREN II, *J.DAN PITTILLO, and IRENE M. ROSSELL

1Department of Biology, Western Carolina University, Cullowhee, 28723-9646; 2Environmental Studies Department, University of North Carolina at Asheville, Asheville, North Carolina 28804-3299

ABSTRACT A survey of vascular flora was completed at wetland sites in the Tulula Creek floodplain in Graham County, North Carolina as part of a comprehensive ecological study. The vegetation survey was conducted in forested and unforested fen and floodplain wetlands in 1994 and 2001, and in a forested floodplain wetland in 2001 in order to document species occurring in these rare mountain habitats. A total of 107 taxa representing 52 families were identified. More than 66% of the taxa also have been reported in other non- alluvial wetlands in the region; about 31% of the taxa identified in the Tulula Creek wetland complex have been reported in non-alluvial wetlands in West Virginia and about 12% have been reported in the non- alluvial wetlands of northeastern and adjacent Canada. This paper documented the communities within this rare wetland complex before intensive stream restoration began in 2001.

INTRODUCTION Wetland plant communities in the Southern Blue Ridge Province are uncommon ecological islands within an expansive temperate deciduous forest; they often contain plant species that do not occur in the surrounding terrestrial and fluvial habitats (Weakley and Schafale 1994). Non-alluvial southern Appalachian wetlands are found in every mountain county in North Carolina (Weakley and Schafale 1994), yet they are infrequent ecosystems that cover less than one percent of the land area (Pittillo 1994). While it is unlikely that wetlands were ever extensive in the mountain areas, only about 200–300 ha of an estimated 2,000 ha of southern mountain wetlands remain in North Carolina (Weakley and Schafale 1994, Moorhead and Rossell 1998). There has been considerable debate over whether southern Appalachian wetlands should be classified as fens or bogs. The criteria used to distinguish fens and bogs in northern regions (Heinselmann 1970, Moore and Bellamy 1974) do not fit the southern ecosystems well. Generally, northern bogs are described as ombrotrophic, acidic (pH , 4.1) wetlands with a surface layer of (Sphagnum spp.) and other mosses, and with hydrology and ion input influenced primarily by precipitation. True ombrotrophic bogs develop peat layers higher than their surroundings and receive nutrients and other minerals exclusively from precipitation (Mitsch and Gosselink 2000). In contrast, northern fens are peatlands with hydrology derived from groundwater that has passed through mineral soil (minerotrophic), and often they are dominated by sedges (Moorhead and Rossell 1998). Many southern mountain wetlands have characteristics of both fens and bogs. Most, if not all, receive groundwater input, in addition to precipitation and surface inflow (Moorhead and Rossell 1998, Moorhead et al. 2001). Unlike the mineral soils of northern Europe and northern North America, which contain high levels of cations such as calcium and magnesium, the moun- tains of the southern Blue Ridge contain highly weathered, leached soils over nutrient-poor,

* email address: @earthlink.net

116 CASTANEA VOLUME 69 highly resistant bedrock. The long-term interactions of climate, topography and vegetation on these materials has produced regional soil complexes that are generally acidic and nutrient poor (Richardson and Gibbons 1993, Weakley and Schafale 1994, Pittillo et al. 1998). Wetlands associated with the southern end of the Appalachian Mountains (North Carolina and ) usually contain mineral soils with a lower organic matter concentration (7–40% of dry mass) and higher pH than their more northern counterparts in West Virginia (Wieder 1985, Walbridge 1994, Mowbray and Schlesinger 1988, Stewart and Nilson 1993, Moorhead and Rossell 1998, Moorhead et al. 2000). While the soil chemistry of most southern Appalachian wetlands more closely resembles northern bogs than fens, they lack the deep peat deposits, probably due to the warmer southern climate and longer dry periods during the growing season (Clymo 1984; Winston 1994; Wieder et al. 1989, 1994; Moorhead and Rossell 1998). Since they receive groundwater inputs from surrounding mineral soils, with the degree of minerotrophy varying from rich to poor, the authors propose ‘‘southern Appalachian fen’’ as the most parsimonious classification. Despite the scarcity of non-alluvial wetlands in the southern Appalachian Mountain region, these small systems contain diverse floral communities with many rare and uncommon species (Murdock 1994, Weakley and Schafale 1994, Moorhead and Rossell 1998, Rossell et al. 1999). Southern Appalachian wetlands contain almost one-fifth of 724 rare plant species monitored by the North Carolina Natural Heritage Program (Murdock 1994); however, pub- lished quantitative vegetation data are sparse. The plant communities of southern Appala- chian wetlands include endemics that exist in the unique microhabitats of the region, as well as species more commonly found in northern and Coastal Plain regions (Richardson and Gibbons 1993, Murdock 1994, Pittillo 1994, Weakley and Schafale 1994). The objective of this study was to inventory and document vascular floral species within Tulula Creek wetland complex. In addition, the wetland indicator status, rare or uncommon status, and the geographic affinity of each species were investigated.

DESCRIPTION OF THE STUDY AREA The 83-ha research site is located in the Tulula Creek floodplain (358169000N; 838429000W; elev. 800 m) in Graham County, North Carolina. The large, relatively flat floodplain is a mosaic of forested and unforested wetlands, including small depressions where Sphagnum spp. accumulate. The floodplain gradually intergrades into upland forests. The site is bordered by U.S. 129 and an abandoned railroad grade; it is crossed by two power lines and has clear vestiges of timbering and subsequent white pine (Pinus strobus) planting. However, it remained a wetland of regional significance (Gaddy 1981) until the mid-1980s when the United States Forest Service traded it to developers who planned to build a golf course and housing complex along the floodplain (Rossell et al. 1999, Rossell and Wells 1999, Moorhead et al. 2001). The developers dredged and channelized Tulula Creek, cut drainage ditches into the floodplain and cleared large areas. The project failed, and in 1994 the North Carolina Department of Transportation purchased the wetland for restoration as a wetland mitigation bank (Moorhead et al. 2001). The site is known locally as Tulula Bog, but it is a wetland complex that includes areas of southern Appalachian fen (ombrotrophic), red maple (Acer rubrum) swamp and open meadow (Moorhead and Rossell 1998, Warren 2002). The North Carolina Natural Heritage Program classified the site as a Swamp Forest-Bog Complex (Schafale and Weakley 1990), defined as poorly drained bottomlands with soils of alluvial origin now removed from regular flooding.

METHODS Vegetation sampling was conducted in an open canopy fen (open fen), closed canopy fen (closed fen), and an open canopy floodplain (open floodplain) in 1994 and 2001. A fourth site, a closed canopy floodplain (closed floodplain), was sampled in 2001. The two fen sites and the open floodplain adjoin each other, while the closed floodplain is approximately 1,000 m downstream from the other three. The fen receives a steady input of groundwater flow and is

JUNE 2004 117 removed from the stream while the floodplain sites are drier and immediately border the stream (Rossell et al. 1999, Rossell and Wells 1999, Moorhead et al. 2001). The 1994 fen sites were selected to investigate the vegetative communities in the forested and unforested portions of the fen, while the floodplain site was chosen for a separate investigation of red maple dynamics. The closed floodplain site was added in 2001 in order to document plant species in a late seral wetland community. A 0.8-ha grid of 100 m2 plots was established in the fen in 1994, with approximately half the plots located within the open canopy area and the other half within the closed canopy area. Twenty plots were randomly selected in each of the open and closed canopy sides of the fen. Trees with a diameter at breast height (DBH) .10 cm were inventoried in 10 3 10 m quadrats. Shrubs and saplings with a 2–10 cm DBH were inventoried in a 4 3 4 m quadrat nested within each 100 m2 plot. Herbaceous vegetation and woody seedlings ,2 cm DBH were inventoried in a 1 3 1m quadrat nested within each 100 m2 plot. The 1 3 1 m and 4 3 4 m quadrats did not overlap. Six 20 3 30 m plots were established in the open floodplain in 1994. Herbaceous vegetation was sampled using the same protocol as in the fen, with the exception that four 1 3 1 m quadrats were randomly placed in each of the six plots rather than in a grid. Shrub-layer and tree-layer vegetation was sampled using a 7 3 7 m (shrub-layer) and a 18 3 18 m (tree-layer) quadrat nested within each of the six plots. The fen and floodplain were re-sampled in 2001. A fourth site, the closed floodplain, was inventoried in 2001 using a 20 3 100 m linear grid of 40, 100 m2 plots. The linear grid was chosen to parallel hydrologic gradients in the floodplain. Vegetation was sampled using the same protocol as in the fen. were identified using Radford et al. (1968) and Weakley (2000). Voucher specimens were deposited at the Western Carolina University Herbarium (WCUH). Uncommon species were categorized based on species status reported in the North Carolina Natural Heritage Program Watch List (Amoroso and Finnegan 2002) and Weakley (2000). A species was defined as having a northern affinity if its geographic range primarily covers northern latitudes but is extended southward via the southern Appalachian Mountains. Coastal disjuncts were defined as those that occur mainly in the Coastal Plain and mountains but are scarce or absent in the Piedmont. These determinations were made, somewhat con- servatively, using Gleason and Cronquist (1963), Radford et al. (1968), Godfrey and Wooten (1979), Weakley and Schafale (1994) and Weakley (2000). The wetland indicator status of the vegetation at the Tulula wetland complex was determined using United States Fish and Wildlife Service (1996). Other sources consulted were United States Fish and Wildlife Service (1988) and United States Department of Agriculture (2001). Region 2 (Southeast) was used to determine indicator status. The wetland categories are: obligate (OBL; .99% in wetlands); facultative wetland (FACW; 67–99% in wetlands), facultative (FAC; 34–66% in wetlands), facultative upland (FACU; 1–33% in wetlands) and upland (UPL; ,1% in wetlands) (United States Fish and Wildlife Service 1996).

RESULTS A total of 107 taxa representing 52 families were documented at the fen and floodplain areas in 1994 and 2001. The plant type best represented at the Tulula Creek wetland sites was forbs (45 species). Other plant types included graminoids (22), shrubs (18), trees (10), ferns (7) and vines (5). The richest genera were Carex (7) and Dichanthelium (5), and the richest families were (12), (11), Rosaceae (9) and Poaceae (8). Almost half (49) of the species were wetland plants (either OBL or FACW), including 11 shrub species but no trees. The shrub and tree layers of most sites were dominated by Acer rubrum, which also was present as seedlings in the herbaceous layer in all sites except for the closed floodplain. Eight uncommon species were recorded at Tulula; they all were herbaceous, more than half were graminoids, and all were OBL except Carex festucacea (FACW). Four are regionally rare in the southern Appalachian Mountains but common elsewhere (Dichanthelium ensifolium var. ensifolium, decangulare, Peltandra virginica and Rhynchospora glomerata); three are on the North Carolina Natural Heritage Program Watch List (Amoroso and Finnegan

118 CASTANEA VOLUME 69 2002) (Carex festucacea, Dichanthelium ensifolium var. curtifolium and asprellum); and one is on the North Carolina Natural Heritage Program List of Rare Plants of North Carolina () (Amoroso and Finnegan 2002).

DISCUSSION Almost two-thirds (68) of the 107 taxa identified in the Tulula Creek wetland complex have been reported in other non-alluvial wetlands in the region, particularly in those classified as a Swamp Forest-Bog Complex, as determined by published species lists and inventories (Gaddy 1981, McLeod 1983, Schafale and Weakley 1990, Smith 1993, Richardson and Gibbons 1993, Pittillo 1994, Weakley and Schafale 1994). Most of the shared species were forbs (25) such as Arisaema triphyllum, patula and and woody species (23) such as Acer rubrum, ligustrina and Nyssa sylvatica. The cooler climate of the southern mountains often results in vegetative communities that are similar to northeastern flora (Rigg and Strausbaugh 1949, Gibson 1970, Pittillo 1994), and this is particularly true in the mountain wetlands (Weakley and Schafale 1994). Gibson (1982) found that about one-third of the taxa in a West Virginia montane wetland (Alder Run Bog) also occurred in northern boreal bogs, and she considered this to be evidence of a distinct northern affinity for the plant communities in the West Virginia wetland. The Tulula Creek wetland complex exhibits a similar northern affinity as about 29% (31) of the taxa identified in the wetland complex also were reported in non-alluvial wetlands in West Virginia, as determined by published species lists and inventories (Rigg and Strausbaugh 1949; Gibson 1970, 1982; Wieder et al. 1981, 1984; Ogle 1982; Walbridge and Lang 1982; Stewart and Nilsen 1993). Most of the species overlap between Tulula and West Virginia wetlands consisted of woody species (13) such as Acer rubrum, Viburnum cassinoides and Kalmia latifolia and forbs (12) such as Impatiens capensis, Sabatia campanulata and sagittatum. Interestingly, Gibson (1982) found little overlap between plant species in Alder Run Bog (WV) and southern Appalachian wetlands, and she suggested that the lack of overlap was due to considerably fewer climatic effects south of West Virginia from the most recent Ice Age (18,000 YBP). While the southern Appalachian Mountain region remained south of the ice sheets during the last glacial epoch, periglacial and tundra conditions may have extended down the higher elevations of the mountains, creating a southern corridor for boreal species (Delcourt and Delcourt 1985; Shafer 1986, 1988; Delcourt et al. 1993; Pittillo et al. 1998). As the glaciers retreated (about 12,000 YBP), flora with northern affinities may have colonized the new wetlands, which acted as climactic refugia as warming continued (Gibson 1982, Delcourt 1985, Delcourt and Delcourt 1985, Delcourt et al. 1993, Richardson and Gibbons 1993, Pittillo et al. 1998). There was much less of an overlap between the taxa in the Tulula wetland complex and non-alluvial wetland plant communities located north of West Virginia (northeastern United States and adjacent Canada), as determined by published species lists and inventories (Heinselmann 1970, Jeglum 1971, Hemond 1980, Nicholson 1995, Le Page and Keddy 1998, Mitsch and Gosselink 2000). Only 11% (12 species) of the taxa identified in the Tulula wetland complex also were reported in non-alluvial wetlands north of West Virginia, including Aster puniceus, Ilex verticillata, Osmunda regalis and Pinus strobus. The Tulula wetland communities, particularly the forested sites, also appeared to have floral similarities with red maple and headwater swamps throughout the Eastern United States (Gaddy 1981, Grafton and Eye 1982, Spooner 1982, Niering 1985, Weakley and Schafale 1994, Mitsch and Gosselink 2000). The reported vegetation structure of red maple swamps usually consists of an Acer rubrum-dominated canopy with a well-developed fern layer beneath (Mitsch and Gosselink 2000), which was similar to the structure in the closed fen and floodplain areas of the Tulula wetland complex (Warren 2002), as well as an overlap in ground-layer species (Gaddy 1981, Grafton and Eye 1982, Spooner 1982, Niering 1985, Weakley and Schafale 1994, Mitsch and Gosselink 2000). The absence of relictual and endemic species at the Tulula wetland complex may be a result of the intense anthropogenic disturbances, or it may indicate that the wetlands are of

JUNE 2004 119 recent origin. In either case, the plants in the Tulula Creek wetland complex may be colonizers from the surrounding area rather than holdouts from centuries past. A second explanation for the low number of relictual and endemic species is that woody encroachment reduced herbaceous wetland communities to the point that stochastic extirpations eliminated the rare species. Weakley and Schafale (1994) reported a higher proportion of northern, coastal and endemic species in southern Appalachian non-alluvial mountain wetlands than were recorded at the Tulula Creek wetland site. Three uncommon species of interest were Carex trichocarpa, Eriocaulon decangulare and Galium asprellum. In North Carolina, C. trichocarpa had previously only been found in Haywood and Ashe Counties, both in the southern Appalachian mountains. Carex trichocarpa primarily grows in low ground and marshy areas north to Canada (Radford et al. 1968, Beal 1977, United States Department of Agriculture 2001), and its presence at the Tulula Creek wetland in Graham County represents its southernmost known population. Carex trichocarpa is significantly rare in North Carolina but more common in the northern portions of its range (Amoroso and Finnegan 2002). Eriocaulon decangulare is largely a Coastal Plain species with a disjunct mountain range; it chiefly occurs in acidic, peaty soils in bogs and pinelands at the coast and in non-alluvial wetland in the mountains (Radford et al. 1968, Beal 1977, Godfrey and Wooten 1979). The occurrence of E. decangulare in the open fen was infrequent and sparse. Conversely, Galium asprellum is largely a northern species that reaches southward along the Appalachian Mountains in bogs and moist places (Radford et al. 1968, Godfrey and Wooten 1979). Galium asprellum occurred throughout the Tulula Creek wetland complex and was most prevalent in the open fen. Because this inventory was part of a larger ecological study, the list of taxa is comprehensive but not exhaustive, and rare species may be underrepresented. For example, Utricularia subulata L. and Lilium canadense L. were known to occur at the site, but neither appeared in the study plots. Despite the extensive anthropogenic disturbance at the Tulula Creek wetland site, only two non-native species were recorded ( hydropiper and Trifolium repens), and both were very infrequent at the site. Trifolium repens is a non-wetland (FACU) plant seeded into the area by the North Carolina Department of Transportation for erosion control during restoration activities at the site, and the small amount of cover may be a holdover from recent construction projects. Persicaria hydropiper is a wetland plant species (OBL) from Eurasia that was only recorded in open floodplain in 2001. The lack of exotic plant species may be attributable to the isolated nature of the wetland, which was far from any major urban or agricultural centers, even in the presence of heavy disturbance and typical exotic species corridors such as the roadways and power lines. While anthropogenic activities have altered the structure of the Tulula Creek wetland complex, it remains a significant example of a habitat type that is increasingly rare in the southern Appalachian region. This paper documented the vascular flora within this rare wetland complex before intensive stream restoration activities were initiated in 2001.

CATALOG OF VASCULAR PLANTS Most nomenclature follows Weakley (2000) except when taxonomically unclear, in which case Kartesz (1994) was used. Families and genera are arranged alphabetically. Uncommon species according to the North Carolina Natural Heritage Program Watch List (Amoroso and Finnegan 2002) and Weakley (2000) are preceded by an asterisk (*). Significantly rare species (Amoroso and Finnegan 2002) are preceded by a double asterisk (**). Wetland indicator status (OBL-obligate, FACW-facultative wetland, FAC-facultative, FACU-facultative upland, UPL- upland, NA-not available) and geographic affinity (N-northern affinity; C-coastal disjunct) are printed after each species in the list when available and appropriate. Five plants identifiable to genus but not species due to lack of reproductive structures were omitted from catalog.

120 CASTANEA VOLUME 69 ACERACEAE CYPERACEAE Acer rubrum L., FAC Carex atlantica Bailey, FACW, C Carex communis Bailey, NA, N ADOXACEAE Carex debilis Michx., FACW Sambucus canadensis L., FACW *Carex festucacea Schkuhr ex Willd., FACW Viburnum cassinoides L., FACW, N Carex intumescens Rudge, FACW Carex stricta Lam., OBL, N ALISMATACEAE **Carex trichocarpa Muhl. ex Willd., OBL, N Eleocharis obtusa (Willd.) J.A. Shultes, OBL Sagittaria latifolia Willd. var. pubescens (Muhl. ex Nutt.) *Rhynchospora glomerata (L.) Vahl., OBL J.G. Sm., OBL Scirpus expansus Fern. Strong, OBL, N ANACARDIACEAE DRYOPTERIDACEAE Rhus copallinum L., FACU Athyrium asplenioides (Michx.) A.A. Eat., NA Toxicodendron radicans (L.) Kuntze, FAC Dryopteris intermedia (Muhl. Ex Willd.) A. Gray, FACU, N AQUIFOLIACEAE Ilex opaca Ait., FAC Kalmia latifolia L., FACU Ilex verticillata (L.) A. Gray, FACW Lyonia ligustrina (L.) DC, FACW ARACEAE Oxydendrum arboreum (L.) DC, UPL Vaccinium corymbosum L., FACW, C Arisaema triphyllum L., FACW *Peltandra virginica (L.) Schott, OBL

ARALIACEAE *Eriocaulon decangulare L., OBL, C

Aralia spinosa L., FAC FABACEAE

ASTERACEAE Apios americana Medik., FACW Trifolium repens L., FACU Ageratina altissima King & H.E. Robins., FACU Ambrosia artemisiifolia L., FACU FAGACEAE Aster pilosus Willd., FAC Quercus rubra L., FACU Aster puniceus L., OBL, N Quercus velutina Lam., UPL Eupatorium fistulosum Barratt, FAC Eupatorium purpureum L., FAC GENTIANACEAE Lactuca canadensis L., FACU Solidago patula Muhl. ex Willd., OBL Sabatia campanulata (L.) Torr., FACW, C Solidago rugosa P. Miller, FAC IRIDACEAE Vernonia novaborcensis (L.) Michx., FAC, N Sisyrinchium mucronatum Michx., FACW, N AZOLLACEAE JUNCACEAE Woodwardia areolata (L.) T. Moore OBL, C Juncus debilis A. Gray, OBL, C BALSAMINACEAE Juncus effusus L., FACW Impatiens capensis Meerb.. FACW Juncus tenuis Willd., FAC

BETULACEAE LAMIACEAE Alnus serrulata (Ait.) Willd., FACW Pycnanthemum muticum (Michx.) Pers., FAC

CAESALPINIANCEAE LAURACEAE Chamaecrista nictitans (L.) Moench, FACU Lindera benzoin (L.) Blume, FACW

CELASTRACEAE LYCOPODIACEAE Euonymus americana L., FAC Lycopodium obscurum L., FACU, N

CLUSIACEAE MAGNOLIACEAE Hypericum mutilum L., FACW Liriodendron tulipifera L., FAC

CONVALLARIACEAE MELASTOMATACEAE Polygonatum biflorum (Walt.) Ell., FAC Rhexia mariana L., FACW

CORNACEAE NYSSACEAE Cornus amomum P. Miller, FACW Nyssa sylvatica Marsh., FAC

JUNE 2004 121 ONAGRACEAE Aronia arbutifolia (L.) Pers., FACW Aronia melanocarpa (Michx.) Ell., FAC, N Ludwigia alternifolia L., OBL Malus angustifolia (Ait.) Michx., NA OSMUNDACEAE Potentilla simplex Michx., FACU Prunus serotina Ehrh., FACU Osmunda cinnamomea L., FACW Rosa palustris Marsh., OBL Osmunda regalis L. var. spectabilis (Willd). A. Gray, OBL Rubus argutus Link, FACU Rubus hispidus L., FACW, C OXALIDACEAE Oxalis stricta L., UPL

PHYTOLACCACEAE *Galium asprellum Michx., OBL

Phytolacca americana L., FACU SALICACEAE PINACEAE Salix sericea Marsh., OBL, N Pinus strobus L., FACU, N SCROPHULARIACEAE POACEAE Agalinis purpurea (L.) Pennell, FACW Calamagrostis coarctata (Torr.) Eat., OBL, C Danthonia spicata (L.) Beauv. Ex Roemer & J.A. Schultes, SMILACEAE NA Smilax glauca Walt., FAC Dichanthelium clandestinum (L.) Gould, FACW Dichanthelium commutatum (J.A. Shultes) Gould, FAC SOLANACEAE Dichanthelium dichotomum (L.) Gould, FAC *Dichanthelium ensifolium (Baldw.) Gould var. curtifo- Solanum carolinense L., FACU lium,NA *Dichanthelium ensifolium (Baldw.) Gould var. ensifolium, THELYPTERIDACEAE NA Muhlenbergia schreberi J.F. Gmel., FAC Thelypteris noveboracensis (L.) Nieuwl., FAC, N

POLYGONACEAE TRILLIACEAE Fallopia scandens (L.) Holub, FAC Trillium undulatum Willd., FACU, N Persicaria hydropiper (L.) Opiz, OBL Polygonum sagittatum L., OBL VIOLACEAE

PRIMULACEAE Viola cucullata Ait., OBL, N Viola primulifolia L., FACW Lysimachia lanceolata Walt., FAC

RANUNCULACEAE VITACEAE Clematis virginiana L., FAC Parthenocissus quinquefolia (L.) Planch., FAC Xanthorhiza simplicissima Marsh., FACW Vitis labrusca L., FAC

ROSACEAE XYRIDACEAE Amelanchier laevis Wieg., NA, N Xyris torta J.M. Smith, OBL, N

ACKNOWLEDGMENTS The authors gratefully acknowledge the contributions of Kevin Moorhead, Henry Mainwaring, Tom Martin and Dan Tinker for manuscript suggestions; David Losure, C. Reed Rossell Jr., Joe-Ann McCoy and Shannon Konrad for field assistance; Paul A. Schmalzer and two anonymous reviewers for helpful suggestions that clarified this manuscript; and Western Carolina University Graduate School and North Carolina Department of Transportation and Center for Transportation and Environment for funding.

LITERATURE CITED

AMOROSO, J.L. and J.T. FINNEGAN (eds.). 2002. Natural Heritage Program list of the rare plants of North Carolina. North Carolina Natural Heritage Program, Division of Parks and Recreation, Raleigh, North Carolina.

122 CASTANEA VOLUME 69 BEAL, E.O. 1977. A manual of marsh and aquatic vascular plants of North Carolina. North Carolina Agricultural Experiment Station Technical Bulletin 247. North Carolina State University, Raleigh, North Carolina. CLYMO, R.S. 1984. The limits to peat bog growth. Phil. Trans. Royal Soc. London 303:605–654. DELCOURT, H.R. and P.A. DELCOURT. 1985. Quaternary palynology and vegetational history of the southeastern United States. p. 1–28. In: Bryant, V.M., Jr. and R.G. Holloway (eds.). Pollen records of late-Quaternary North American sediments. American Association of Stratigraphic Palynologists Foundation, Dallas, . DELCOURT, P.A. 1985. The influence of later-quaternary climatic and vegetational change on paleohydrology in unglaciated eastern North America. Ecologia Medit. Tome IX (Fascicule 1):17–26. DELCOURT, P.A., H.R. DELCOURT, D.F. MORSE, and P.A. MORSE. 1993. History, evolution and organization of vegetation and human culture. p. 47–49. In: Martin, W.H., S.G. Boyce, and A.C. Echternacht (eds.). Biodiversity of the southeastern United States: lowland terrestrial communities. John Wiley and Sons, , New York. GADDY, L.L. 1981. The bogs of the southwestern mountains of North Carolina. Report to North Carolina Natural Heritage Program, Raleigh, North Carolina. GIBSON, J.R. 1970. The flora of Alder Run Bog, Tucker County, West Virginia. Castanea 35:81–98. GIBSON, J.R. 1982. Alder Run Bog, Tucker County, West Virginia: its history and vegetation. p. 101–115. In: McDonald, B.R. (ed.). Proceedings of the symposium on wetlands of the unglaciated Appalachian region, Morgantown, West Virginia. GLEASON, H.A. and A.C. CRONQUIST. 1963. Manual of vascular plants of northeastern United States and adjacent Canada. Willard Grant Press, Boston, Massachusetts. GODFREY, R.K. and J.W. WOOTEN. 1979. Aquatic and wetland plants of southeastern United States. University of Press, Athens, Georgia. GRAFTON, W.N. and O.L. EYE. 1982. Vascular flora of eight selected West Virginia wetlands with special references to rare species. p. 107–121. In: McDonald, B.R. (ed.). Proceedings of the symposium on wetlands of the unglaciated Appalachian region, Morgantown, West Virginia. HEINSELMANN, M.L. 1970. Landscape evolution, peatland types, and the environment in the Lake Agassiz Peatlands Natural Area, . Ecol. Monogr. 40:235–260. HEMOND, H.F. 1980. Biochemistry of Thoreau’s Bog, Concord, Massachusetts. Ecol. Monogr. 50:507–526. JEGLUM, J.K. 1971. Plant indicators of pH and water level in peatlands at Candle Lake, Saskatchewan. Can. J. Bot. 49:1661–1676. KARTESZ, J.T. 1994. A synonymized checklist of the vascular flora of the United States, Canada, and Greenland. Timber Press, Portland, Oregon. LE PAGE, C. and P.A. KEDDY. 1998. Reserves of buried seeds in beaver ponds. Wetlands 18:242–248. MCLEOD, D.E. 1983. The vascular flora of a small southern Appalachian bog-fen. Assoc. Southeast. Biol. Bull. 30:70–71. MITSCH, W.J. and J.G. GOSSELINK. 2000. Wetlands, 3rd ed. Van Nostrand Reinhold, New York, New York. MOORE, P.D. and D.J. BELLAMY. 1974. Peatlands. Springer-Verlag, Inc., New York, New York. MOORHEAD, K.K. and I.M. ROSSELL. 1998. Southern mountain fens. p. 379–403. In: Messina, M.G. and W.H. Conner (eds.). Southern forested wetlands. Lewis Publishers, Boca Raton, . MOORHEAD, K.K., I.M. ROSSELL, J.W. PETRANKA, and C.R. ROSSELL,JR. 2001. Tulula wetlands mitigation bank. Ecol. Res. 19:74–80. MOORHEAD, K.K., R.E. MOYNIHAN, and S.L. SIMPSON. 2000. Soil characteristics of four southern Appalachian fens in North Carolina, USA. Wetlands 20:560–564. MOWBRAY, T. and W.H. SCHLESINGER. 1988. The buffer capacity of organic soils of the Bluff Mountain Fen, North Carolina. Soil Sci. 146:73–79. MURDOCK, N.A. 1994. Rare and endangered plants and animals of southern Appalachian wetlands. Water Air Soil Poll. 77:385–405. NICHOLSON, B.J. 1995. The wetlands of Elk Island National Park: vegetation classification, water chemistry and hydrotopographic relationships. Wetlands 15:119–133. NIERING, W.A. 1985. Wetlands. Alfred A. Knopf, Inc., New York, New York. OGLE, D.W. 1982. Glades of the Blue Ridge in southwestern Virginia. p. 143–147. In: McDonald, B.R. (ed.). Proceedings of the symposium on wetlands of the unglaciated Appalachian region. West Virginia University, Morgantown, West Virginia. PITTILLO, J.D. 1994. Vegetation of three high-elevation southern Appalachian bogs and implications of their vegetational history. Water Air Soil Poll. 77:333–348. PITTILLO, J.D., R.D. HATCHER,JR., and S.W. BUOL. 1998. Introduction to the environment and vegetation of the Southern Blue Ridge Province. Castanea 63:202–216.

JUNE 2004 123 RADFORD, A.E., H.E. AHLES, and C.R. BELL. 1968. Manual of the vascular flora of the Carolinas. The University of North Carolina Press, Chapel Hill, North Carolina. RICHARDSON, C.J. and J.W. GIBBONS. 1993. Pocosins, Carolina bays and mountain bogs. p. 257–310. In: Martin, W.H., S.G. Boyce, and A.C. Echternacht (eds.). Biodiversity of the Southeastern United States: lowland terrestrial communities. John Wiley and Sons, New York, New York. RIGG, G.B. and P.D. STRAUSBAUGH. 1949. Some stages in the development of sphagnum bogs in West Virginia. Castanea 14:129–148. ROSSELL, C.R., I.M. ROSSELL, J.W. PETRANKA, and K.K. MOORHEAD. 1999. Characteristics of a partially disturbed southern Appalachian forest-gap bog complex. Virginia Mus. Nat. Hist. spec. publ. 7:81–89. ROSSELL, I.M. and C.L. WELLS. 1999. The seed banks of a southern Appalachian fen and an adjacent degraded wetland. Wetlands 19:365–71. SCHAFALE, M.P. and A.S. WEAKLEY. 1990. Classification of the natural communities of North Carolina, third approximation. North Carolina Natural Heritage Program, Division of Parks and Recreation, Raleigh, North Carolina. SHAFER, D.S. 1986. Flat Laurel Gap Bog, Pisgah Ridge, North Carolina: late Holocene development of a high- elevation heath bald. Castanea 51:1–9. SHAFER, D.S. 1988. Late quaternary landscape evolution at Flat Laurel Gap, Blue Ridge Mountains, North Carolina. Quat. Res. 30:7–11. SMITH, A.B. 1993. A survey of mountain wetland communities. Report to North Carolina Natural Heritage Program, Raleigh, North Carolina. SPOONER, D.M. 1982. Wetlands in Teays-stage valleys in extreme southeastern Ohio: formation and flora. p. 89–99. In: McDonald, B.R. (ed.). Proceedings of the symposium on wetlands of the unglaciated Appalachian region. West Virginia University, Morgantown, West Virginia. STEWART, C.N., JR. and E.T. NILSEN. 1993. Association of edaphic factors and vegetation in several isolated Appalachian peat bogs. Bull. Torrey Bot. Club 120:128–135. UNITED STATES DEPARTMENT OF AGRICULTURE,NATURAL RESOURCES CONSERVATION SERVICE. 2001. The PLANTS Database, Version 3.1. National Plant Data Center, Baton Rouge, . http://plants.usda.gov. UNITED STATES FISH AND WILDLIFE SERVICE,NATIONAL WETLANDS INVENTORY. 1988. The 1988 national list of vascular plant species that occur in wetlands. United States Fish and Wildlife Service biological report 88. United States Fish and Wildlife Service, Washington, D.C. UNITED STATES FISH AND WILDLIFE SERVICE,NATIONAL WETLANDS INVENTORY. 1996. The 1996 national list of vascular plant species that occur in wetlands. United States Fish and Wildlife Service, Washington, D.C. WALBRIDGE, M.R. 1994. Plant community composition and surface water chemistry of fen peatlands in West Virginia’s Appalachian Plateau. Water, Air, Soil Poll. 77:247–269. WALBRIDGE, M.R. and G.E. LANG. 1982. Major plant communities and patterns of community distribution in four wetlands of the unglaciated Appalachian region. p. 131–142. In: McDonald, B.R. (ed.). Proceedings of the symposium on wetlands of the unglaciated Appalachian region. West Virginia University, Morgantown, West Virginia. WARREN, R.J. II. 2002. Impact of woody vegetation succession upon herbaceous communities in a southern Appalachian wetland. M.S. thesis, Western Carolina University, Cullowhee, North Carolina. WEAKLEY, A.S. 2000. Flora of the Carolinas and Virginia (Working draft). The Nature Conservancy, Southern Resource Office, Durham, North Carolina. WEAKLEY, A.S. and M.P. SCHAFALE. 1994. Non-alluvial wetlands of the Southern Blue Ridge—diversity in a threatened ecosystem. Water, Air, Soil Poll. 77:359–93. WIEDER, R.K. 1985. Peat and water chemistry at Big Run Bog, a peatland in the Appalachian mountains of West Virginia, U.S.A. Biogeochemistry 1:277–302. WIEDER, R.K., A.M. MCCORMICH, and G.E. LANG. 1981. Vegetational analysis of Big Run Bog, a nonglaciated sphagnum bog in West Virginia. Castanea 46:16–29. WIEDER, R.K., C.A. BENNETT, and G.E. LANG. 1984. Flowering phrenology at Big Run Bog, West Virginia. Amer. J. Bot. 71:203–209. WIEDER, R.K., J.P. YAVITT, G.E. LANG, and C.A. BENNETT. 1989. Aboveground net primary productivity at Big Run Bog, West Virginia. Castanea 54:209–216. WIEDER, R.K., M. NOVAK, W.R. SCHELL, and T. RHODES. 1994. Rates of peat accumulation over the past 200 years in five Sphagnum-dominated peatlands in the United States. J. Paleolimnol. 12:35–47. WINSTON, R.B. 1994. Models of the geomorphology, hydrology and development of domed peat bodies. Geol. Soc. Amer. Bull. 106:1594–1604.

Received October 31, 2002; Accepted May 7, 2003.

124 CASTANEA VOLUME 69