United States

Department of

Agriculture

Forest Service

Northeastern

Research Station

Research Paper NE-725

Characterization of High Elevation

Central Appalachian Wetlands

Karen E. Francl

W. Mark Ford

Steven B. Castleberry
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Abstract

We characterized 20 high elevation wetlands in the central Appalachian Mountains

in West Virginia and Maryland, in terms of vegetation, soils, hydrology, and geology.

Plant species were distributed along soil chemical (pH, conductivity) and physical

(organic matter depth) gradients across sites. Topography and geology appear to

explain differences among these wetlands, as reflected by soil and chemistry

measures and vegetation distribution. Our work provides substantial quantitative

baseline data for these uncommon high elevation wetlands, emphasizing the

importance and diversity of these isolated systems.

The Authors

KAREN FRANCL is the assistant director of the University of Notre Dame

Environmental Research Center and a research assistant professor, Department of

Biological Sciences, Notre Dame, IN 46556.

W. MARK FORD is a research wildlife biologist, USDA Forest Service Timber and

Watershed Laboratory, Parsons, WV 26287

STEVEN B. CASTLEBERRY is an assistant professor, Warnell School of Forest

Resources, University of Georgia, Athens, GA 30602

Manuscript received for publication 18 March 2004

Cover Photos, from left: Alder Bog, Yellow Creek, and Main Bog; photos by

Karen Francl.
1

Summary

We characterized 20 high elevation wetlands in the central Appalachian Mountains in West

Virginia and Maryland. We performed woody and herbaceous vegetation surveys by estimating

cover percentage in plots (10 x 10 m) and subplots (1 x 1 m), documented soil profiles and

texture within each vegetation plot, measured water chemistry parameters, monitored

hydrology with piezometers, and investigated geologic influences at each site. Plant species

were distributed along soil chemical (pH, conductivity) and physical (organic matter depth)

gradients across sites. Topography and geology explain the presence and condition of these

wetlands, as all sites were situated within natural basins, and 17 of 20 sites were underlain

primarily by sandstone that impedes drainage and creates a perched water table. The remaining

sites contained a greater proportion of limestone, which was reflected by differences in

vegetation and soil chemistry. Our research emphasizes the importance of small (< 5 ha)

wetlands, often overlooked and unprotected, that support as many wetland plants as their

larger counterparts on a per-unit-area basis. Our work provides substantial quantitative

baseline data for these high elevation wetlands, providing a foundation for understanding

the structure and function of these systems.
2

Introduction

Acidic, anoxic, nutrient-poor wetlands are intermittently

distributed across cool environments of the northern

United States and Canada. Plant and animal matter

production exceeds the annual decomposition rate in

these wetlands. Wetlands with these climate conditions

and plant communities are locally abundant in portions

of the central Appalachian Mountains (restricted in this

discussion to east-central and northern West Virginia

and western Maryland west of the Allegheny Front).

Despite similar plant communities in northern boreal

bogs and fens and central Appalachian wetlands, the

ontogeny of these regions’ wetlands is quite different.

Unlike the wetlands of the north, central Appalachian

wetlands were not influenced directly by continental

glacial recession (Ingham 1996). Continental glaciers

from the Wisconsin Ice Age did not extend any farther

south than Pennsylvania in the Appalachian region.

However, at high elevations (≥800 m), the climatological

conditions of cooler air and soil temperatures and greater

precipitation mimic the environment of wetlands at

more northern latitudes (Rigg and Strausbaugh 1949).

The locally cool climate acts as a refuge for many

northern plant species that were trapped as the last

continental glaciers receded. Several species reach their

southernmost extent in these central Appalachian

wetlands (Fortney 1975, Ingham 1996).

Fens and bogs are common throughout large portions of

the northern United States. However, these categories

fail to adequately describe or “match” conditions found

farther south in the central and southern Appalachians

(Moorhead and Rossell 1998). Typically, bogs of the

northern United States and Canada are defined by acidic

conditions (pH ≤4.1) and an ombrotrophic

(precipitation-fed) nature. Bog vegetation primarily

consists of Sphagnum mats, cranberries (Vaccinium spp.),

and other types of low-lying herbaceous plants (Core

1966). Classic northern fens are slightly less acidic and

are minerotrophic (groundwater-fed). Dominant

vegetation includes sedges (Carex spp.), rushes (Juncus

spp.) and several shrub species (Core 1966, Walbridge

1994). Central and southern Appalachian wetlands are

mostly minerotrophic, and therefore classified as fens,

but they support vegetation more similar to bog flora

(Moorhead and Rossell 1998). Additionally, these bogs

and fens lack the distinct concentric zonation of plant

communities found in many northern kettle bogs.

Indeed, southern Appalachian wetlands have been

described as an “irregular mosaic,” more patchy than

patterned (Wieder et al. 1981, Moorhead and Rossell

1998). Vegetational studies of central Appalachian

wetlands, such as Big Run Bog (Tucker County, WV)

support this description (Muzika et al. 1996), and

attribute plant distributional patterns to water chemistry

differences across the site (Wieder et al.1981, Walbridge

1994).

An important difference between northern boreal bogs

and central Appalachian wetlands is their original state

prior to anthropogenic activity. Rather than existing as

open, floating Sphagnum mats, many central

Appalachian wetlands (e.g., Canaan Valley, Tucker

County, WV) were forested, red spruce (Picea rubens)-

dominated wetlands prior to timber harvests of the late

1800s through the 1920s (Clarkson 1993). Additionally,

balsam fir (Abies balsamea) and rosebay rhododendron

(Rhododendron maximum) thickets were common in the

wetland canopy and understory, respectively, whereas the

forest floor consisted of wetland mosses atop a deep, rich

organic layer (Fortney 1993). Logging, mostly

clearcutting, exposed the organic soils, and subsequent

fires burned and removed the organic layers. As a result,

rock outcrops dominated these formerly forested

wetland landscapes (Fortney 1975). Tree removal also

created isolated pockets of raised water tables, so

formerly dry sites were too wet to sustain tree

regeneration, instead supporting shrub-scrub and open

wetlands. Currently, open wetlands, often beaver (Castor

canadensis) influenced, have reappeared on the

landscape, mainly in high elevation alluvial areas

(Fortney 1993, Walbridge 1994). Therefore, it is likely

that many central Appalachian wetlands are geologically

young or drastically altered from their natural state more

than a century ago. In very few exceptions, like

Cranberry Glades (Pocahontas County, WV), the lack of

local logging minimized disturbance to the organic layer,

which extends more than 2 m below the surface,

markedly deeper than logged, “young” wetlands

(Darlington 1943). Additionally, portions of The Glades

(Garrett County, MD) have been reported to contain

peat deposits greater than 3 m1

.

The vegetation, soils, and hydrology of a few large bogs

and fens in West Virginia (< 10 sites) have been studied

in the past (Darlington 1943, Gibson 1970, Walbridge

1994, Wieder 1985). Dominant plant associations and

short-term plant succession have been thoroughly

mapped in a two wetlands (Edens 1973, Walbridge

1994). Wieder (1985) and Yavitt (1994), working at Big

Run Bog, also examined the local wetland

biogeochemistry concluding that differences in

vegetative structure result from surface soil chemistry

1

Feller, D. 2003. Personal communication. Wildlife Biologist,

Maryland Department of Natural Resources, 10775 Savage

River Road, Swanton, MD 21561.
3

(i.e., calcium, nitrogen, and magnesium concentrations

and pH) variation at Big Run Bog. Walbridge’s (1994)

research at Big Run and three other wetlands in the area

support this relationship between plant species

composition, pH, and cation concentrations.

Other wetland soil chemistry studies in the region have

been cursorial, typically documenting nothing more

than pH and soil water temperature. Soil classification

maps are available for all counties in West Virginia and

Maryland (USDA Soil Conserv. Serv. 1959, USDA Soil

Conserv. Serv. 1982, USDA Natural Resources Conserv.

Serv. 2002a). However, due to scaling and poor-quality

aerial photographs used to make maps, soil surveys often

classify wetlands less than 2 ha according to dominant

terrestrial, xeric soils surrounding the area (Wakeley

1994). Those large enough to be recognized (e.g.,

Canaan Valley and Cranberry Glades) are generally

classified as muck and peat soils that are acidic and

poorly drained (Ingham1996, Moorhead and Rossell

1998). Darlington’s (1943) work at Cranberry Glades

showed great variation in peat depth, from 0.3 to 2.3 m

across the site, and peat type, including fine peat,

Sphagnum peat, and sedge peat.

Ingham (1996) performed a regional wetland survey that

examined general vegetation, soil, climatic, and geologic

conditions among 20 central Appalachian wetlands, but

limited research to qualitative comparisons. Additionally,

Diehl (1981) qualitatively examined the influence of

geologic factors on the presence and formation of nearly

50 wetlands throughout West Virginia. Until now, few

large-scale studies have been undertaken that

quantitatively assess the relationship of vegetation, soils,

hydrology, and geology of central Appalachian bogs and

fens.

Our goal was to survey high-elevation central

Appalachian wetlands and to provide baseline data for

each component that defines them as wetlands:

hydrophytic vegetation, hydric soils, and periodic

flooded conditions during the growing season

(Cowardin et al.1979). We wanted to determine how

variable these systems were, and if and how they differed

chemically and hydrologically. To accomplish this task,

we quantified vegetation structure, monitored water

levels with piezometers, and measured several soil

components (type, texture, chemistry, and vertical

profile) at 20 wetlands in northern West Virginia and

western Maryland. Additionally, we examined general

geologic and topographic patterns to determine how the

formation of these wetlands, underlying rock layers, and

placement on the land influence basic wetland

parameters.

The scarcity of information in nearly all aspects of

ecological concern provides countless research

opportunities in central Appalachian wetlands

(Moorhead and Rossell 1998). Once these habitats have

been thoroughly examined for floral composition and

physical and chemical parameters, informed

management decisions can be made regarding legal

protection of these sites.

Materials and Methods

Site Selection

Twenty sites were selected for this study (Fig. 1, Table 1).

Sites were identified from literature reviews and an

examination of topographic quadrangle maps. Site

selection was based on accessibility and permission to

work on private and public lands. Seventeen sites in

Randolph and Tucker Counties, WV, plus three in

Garrett County, MD and/or Preston County, WV, were

qualitatively categorized according to relative geographic

placement on the landscape (Big Run Bog, Canaan Loop

Road, Canaan Valley, Dolly Sods, Maryland/TNC, and

Otter Creek), and referred to as such in the text. Several

of the sites, including Big Run Bog, Alder Run, and

Cranesville Swamp, have been studied in the past

(Robinette 1964, Gibson 1970, Walbridge 1994).

However, many of the selected areas are small, isolated

habitats that have received little attention. The West

Virginia sites are located on federal- or state-owned land,

including the Monongahela National Forest (14 sites),

Canaan Valley National Wildlife Refuge (two), and

Canaan Valley Resort State Park (one). The three

wetlands in Garrett County, MD are owned by The

Nature Conservancy. Initially, all sites appeared to be

isolated depressional wetlands, but several sites also

contained flowing water (creeks and seeps less than 2 m

wide), and therefore likely were influenced by alluvial

inputs.

Vegetative Surveys

To determine the presence and relative importance of

hydrophytic vegetation, we followed Tiner (1999) and

Reed’s (1988) characterization techniques, incorporating

weighted values for each species, based on its degree of

wetland obligation. We sampled vegetation at each

wetland using plots and subplots, modified from

Walbridge’s (1994) nested-plot design. At each site, we

selected two to twenty-four 10 x 10 m plots as most

representative of community structure. The number of

plots per site was based on a combination of wetland size

and visible habitat diversity. The small sites were

sampled at approximately 1-2 plots/ha. Because the

larger sites could not be sampled with this same

intensity, plots representing each physiognomic type
4 Table 1.—Locality information for 20 wetland sites in West Virginia and Maryland, surveyed in 2001-2002

Site name Code Countya Geographic Latitude Longitude Elevation Area O

cluster (North) (West) (m) (ha)

Canaan Valley State ABER Tucker Canaan Valley 39° 00.930' 79° 27.800' 960 3.0 StPark, Abe Run TrailAlder Run ALDR Tucker Dolly Sods 39° 01.710' 79° 19.200' 1164 5.0 UCanaan Valley NWR, BEAL Tucker Canaan Valley 39° 04.273 79° 24.776' 954 1.2 UBeall TractBig Run Bog BIGR Tucker Big Run Bog 39° 07.017' 79° 34.554' 982 15.0 UCondon Run COND Randolph Otter Creek 38° 56.540' 79° 40.000' 918 3.0 UCranesville Swamp CRSW Preston Maryland/TNC 39° 31.981' 79° 29.199' 794 225.0 TFisher Springs Run FISH Tucker Dolly Sods 39° 00.890' 79° 19.510' 1158 25.0 UThe Glades GLAD Garrett, MD Maryland/TNC 39° 33.426' 79° 16.969' 775 240.0 THammel Glade HAMM Garrett, MD Maryland/TNC 39° 29.419' 79° 21.797' 737 20.0 TCanaan Valley NWR, HERZ Tucker Canaan Valley 39° 02.268' 79° 25.358' 983 10.0 UHerz TractHigh Ridge HIGH Tucker/ Dolly Sods 38° 57.647' 79° 21.235' 1216 28.0 U

Randolph

Canaan Loop, MAIN Tucker Canaan Loop Road 39° 04.371’ 79° 28.363’ 1111 30.0 UMain Rd.Moore Run MOOR Randolph, Otter Creek 39° 00.072' 79° 39.623' 991 2.0 UCanaan Loop, NORT Tucker Canaan Loop Road 39° 04.140' 79° 29.059' 1076 15.0 UNorth Rd.Canaan Loop, POWR Tucker Canaan Loop Road 39° 04.959' 79° 27.784' 1099 0.08 UPowerlineCanaan Loop, REDR Tucker Canaan Loop Road 39° 04.125' 79° 29.455' 1085 0.4 URed RunCanaan Loop, Trail 1 TRL1 Tucker Canaan Loop Road 39° 05.197' 79° 28.896' 1038 0.4 U(101)Canaan Loop, Trail 2 TRL2 Tucker Canaan Loop Road 39° 04.823' 79° 28.847’ 1067 0.2 U

(109/108)Canaan Loop, Trail 3 TRL3 Tucker Canaan Loop Road 39° 04.466' 79° 29.487 1085 1.3 U(109/701)Yellow Creek YELL Randolph Otter Creek 38° 57.742' 79° 40.600' 952 2.5 U

aWest Virginia unless otherwise noted
5

(i.e., shrub-scrub, open mossy, open grassy, and forested

patches) were surveyed. At least two plots of each

recognizable community type were sampled. Within

these plots, we visually estimated cover percentage for

each woody species (Walbridge 1994). Nine 1 m x 1 m

subplots were randomly selected within each plot, and

cover percentage for herbs, vines, and mosses was

estimated.

When possible, all plants were identified to species, and

taxonomy followed the PLANTS database (Strausbaugh

and Core 1978, USDA Natural Resources Conserv. Serv.

2002b. Sphagnum moss was identified to genus only. If a

plant was rare or easily identifiable by sight, photographs

served as voucher specimens. If a plant species was

common, a specimen was collected and prepared as an

herbarium specimen. Voucher specimens of pressed

plants were deposited in the herbarium at the University

of Georgia. Several moss specimens also were placed in

the herbarium at West Virginia University.

Vegetation was analyzed using weighted averages for each

species, where each species is assigned an appropriate

ecological indicator value, ranging from one (obligate

wetland species occurring more than 99 percent of the

time in wetlands) to five (obligate upland species found

in nonwetlands more than 99 percent of the time) (Reed

1988). If Reed (1998) did not assign a value to a species,

as in the case of nonvascular plants, we assigned an

appropriate value, based on literature review and values

assigned to other closely related species. For example, all

Sphagnum mosses were considered obligate wetland

species. A weighted average was calculated for each site:

Ws = Σ(Cis)(Ei

)/ Σ Cis

Where:

Ws = weighted average, or vegetative index value, for site s

Cis = percent cover for each species i at site s,

Ei

 = ecological index value for species i.

Site names and groupings 

 Maryland

 1 Cranesville Swamp 

 2 Hammel Glade 

 3 The Glades 

 Big Run 

 4 Big Run Bog 

 Canaan 

 5 North Bog (Canaan Loop) 

 6 Main Bog (C. Loop) 

 7 Red Run (C. Loop) 

 8 Trail 1 (C. Loop) 

 9 Trail 2 (C. Loop) 

 10 Trail 3 (C. Loop) 

 11 Powerline (C. Loop) 

 12 Abe Run (C. Valley) 

 13 Herz Tract (C. Valley) 

 14 Beall Tract (C. Valley) 

 Dolly Sods 

 15 Alder Run 

 16 Fisher Springs Run 

 17 High Ridge 

 Otter Creek 

 18 Moore Run 

 19 Yellow Creek 

 20 Condon Run 

Figure 1.—Twenty high elevation wetland study sites in the Allegheny Mountains of Preston, Tucker,

and Randolph counties, West Virginia and Garrett County, Maryland. Symbols refer to groups of

sites geographically clustered, and referred to as such in text.

×
6

The result was a weighted average of the site, ranging

from one to five. Weighted averages typically are less

than 2.0 in wetlands. For borderline averages (2.0–2.5),

it is necessary to use soil and hydrology parameters to

confirm that a site is a wetland (Tiner 1999).

To further analyze vegetation data, we calculated the

importance value (I) for each species, which equally

weighs its relative dominance and relative frequency

(Cox 1996). Dominance refers to the basal area covered

by a species (as estimated by the cover percentage within

each plot), and frequency to the fraction of plots

containing the particular species. Relative values are the

proportions for each species of the total for all species.

Soil Surveys

In our characterization surveys, we followed the hydric

soil morphology characters (soil color) explained by

Wakeley (1994), and supported by the USDA Soil

Conservation (1991) such as reduced, saturated

conditions result in gray-colored, low-chroma soils and

the accumulation of organic matter (surface organic layer

greater than 20 cm). Within the established 10 x 10 m

vegetation plots (2-22/site), we obtained soil profile

information for the surface and subsurface layers (2 plots

at the Glades were not sampled for soils because recent

acid mine drainage restoration construction filled in

these areas). We documented the depth, color (using a

Munsell Soil Color Chart), and particle size distribution

of each horizon to 1 m below the surface (Kollmorgen

Corporation1992). We measured the depth to gray

mottles and gray matrix to determine the extent of

reduced conditions in the soil. In measuring depth of

organic matter, we noted the percent of samples with

organic layers deeper than 20 cm (a histic epipedon)

(Hurt et al. 2003). We also calculated the percentage of

samples with organic layers greater than 40 cm, which

would classify as histosols under the U.S. soil

classification system (Soil Survey Staff 1998a, Hurt et al.

2003).

From each point at which a soil profile was recorded, we

collected a soil sample at ca. 10 cm below the surface. So

that a minimum of three soil samples were taken from

each site, additional randomly selected points were

sampled at four of the smaller sites. Samples were dried

at 70° C for 48 hours, or until thoroughly dry. Particlesize analysis was completed following the hydrometer

method (Gee and Bauder 1982). Carbon and nitrogen

percentages were analyzed on a NC 2100 soil analyzer

(C.E. Elantech, Inc., Lakewood, NJ), which uses a gaschromatographic approach to quantitatively determine

concentrations. We compared percentage of clay to

percentage of organic carbon in each sample to further

classify the soils as organic, mineral, or mucky mineral

(Hurt et al. 2003). County soil maps also were analyzed

to determine the proportion of sites that were classified

as containing wetland soils (USDA Soil Conserv. Serv.

1959, USDA Soil Conserv. Serv. 1982).

To estimate organic matter from each sample, we

examined our carbon percentage results. Literature

suggests that soil type and depth of sample influence

percentage of carbon, which can be 40–59 percent of the

organic carbon (Nelson and Sommers 1982, Orlov

1985, Jenkinson 1988). In surface humic soils, the range

is typically 50–55 percent, and slightly less in subsoils2

(Orlov 1985). Because our samples were obtained at 10

cm below the surface, we used a conservative estimate of

48 percent in our calculations.

In early August 2001, we used a YSI 600XL with a

610DM data logger to obtain water chemistry readings

(YSI Inc., Yellow Springs, Ohio). Readings were taken at

two or more representative spots within each wetland,

and averaged for each site. At each site, we measured pH,

conductivity (mS/cm), dissolved oxygen (saturation

percentage), temperature (°C), and redox potential

(ORP, mV). To avoid bias from rainfall (Allan 1995),

data were recorded at all sites within a 36-hour period,

which occurred at least 48 hours after any measurable

precipitation.

Because dissolved oxygen and temperature fluctuate

throughout the day (Allan, 1995), we modified our

readings from all sites to estimate dissolved oxygen and

temperature at 1200 h, so that all sites could be

compared without temporal bias. To create a standard

curve for oxygen and temperature throughout the day,

we recorded temperature and dissolved oxygen at a

small, stagnant creek on the Herz Tract (Canaan Valley

NWR, Tucker County, WV). We recorded dissolved

oxygen and temperature at 2-hour intervals (0600 to

1800 h) on the day immediately following the 36-hour

recording period. No rainfall occurred during this

period.

Hydrological Monitoring

Criteria for wetland hydrology dictate that surface

saturation or inundation must exist for 5 percent or

more of the year (Wakeley 1994). To be considered

flooded or saturated, the water table must be within ca.

15–46 cm of the surface, with the specified depth

depending on soil drainage class and permeability

(Wakeley 1994). To account for yearly variability, we

2

Morris, L. 2002. Personal communication. Professor,

University of Georgia Warnell School of Forest Resources,

Athens, GA 30602.
7

monitored changes in the water table depth at each site

for two growing seasons. At each site, we installed a

piezometer, which was constructed of 5.2-cm-diameter

PVC with one end covered with water-permeable

double-folded landscaping fabric. A PVC cap was added

to the other end to prevent rainfall from entering the

pipe. Inside this pipe, we placed a 2.6-cm-diameter PVC

pipe as a dip-stick to measure the maximum water level

height since last measurement. The piezometer was

placed no more than 1 m deep, slightly projecting from

the surface (Morrison 1983). One instrument was

positioned in a representative area of each wetland (most

common microhabitat within the site), at least 10 m

from edge habitat and avoiding hummocks and direct

stream channels. If an impenetrable layer, such as a hard

rock or clay layer, was reached prior to 1 m deep, the

piezometer was established atop this layer. Using waterfinding paste to gauge current depth, and ground cork

particles to gauge maximum water levels since the last

measurement, piezometer readings were taken a

minimum of every 14 days from June to August 2001,

and checked once every 28 days thereafter through

September 2002, unless snowfall prevented access.

Previous wetland work indicated that sampling every 28

days results in an average error in daily stage estimates of

less than 0.05 m and 5 percent of the actual daily values

(Shaffer et al. 2000). For the purposes of comparing

these wetlands, monthly sampling provided a

representative description of water level fluctuations,

allowing us to examine intra- and inter-seasonal trends

and to infer if areas are continually flooded for 5 percent

of the year.

Piezometer trends were qualitatively compared to rainfall

data from the 2001 and 2002 growing seasons, as

recorded at the USDA Forest Service weather station on

Bearden Knob (Canaan Heights, Tucker County, WV;

elevation ca. 1150 m) , and to long-term (1980-1990)

precipitation trends obtained from the National Oceanic

and Atmospheric Administration station for Canaan

Valley (elevation ca. 955 m) (Stephenson 1993).

Geologic Analyses

The underlying geology of each site was determined

through an analysis of state geologic maps of West

Virginia and Maryland (Cardwell 1968, Maryland

Geological Survey 2000). Site-specific geologic

information also was obtained through published data

(Robinette 1964, Diehl 1981, Ingham 1996).

Dominant, secondary, and tertiary rock groups and

formations were recorded for each site. Quantitative

analyses were not performed, due to the scale, age, and

questionable accuracy of the maps. U.S. Geologic Survey

topographical quadrangle maps (1:24,000) were

examined to study trends in elevation within and

surrounding each site.

Statistical Analyses

Principal components analyses were performed using

NTSYS-pc version 2.02i (Applied Biostatistics, Inc.

1998). Plot and subplot plant data, measured in

proportions (I-values), were standardized using the

default methods in NTSYS. Initially, separate principal

components analyses were performed using the I-values

for each species for woody (plot) and herbaceous

(subplot) data (Ludwig and Reynolds 1988). However,

the first principal components of woody and herbaceous

analyses were strongly correlated (r = 0.922, P < 0.001).

Therefore, we performed a single principal components

analysis for all vegetation data, giving equal weight to the

woody and herbaceous I-values. For these analyses, we

calculated correlations among variables, then extracted

the first three principal components from the correlation

matrix. We chose to examine the first three axes in all

cases because the eigenvalues typically began to level off

at relatively low values at the fourth principal

component. Scatter diagrams were created by plotting

each site’s projection value for a particular component.

Although we violated the assumption of normality in the

data, we chose not to attempt data transformation

because this often makes the interpretation of principal

components difficult (Isebrand and Crow 1975).

Pearson product-moment correlations (α = 0.05),

performed using SYSTAT 8.0 (SPSS, Inc. 1998),

examined relationships among vegetation projection

values for each site, wetland indices, and all soil

chemistry measures (e.g., pH, dissolved oxygen, redox

potential). We also compared these values to the average

depth of the organic layer and average piezometer

recordings.

Results

Vegetation Surveys

We documented 50 species of trees, shrubs, and woody

vines in 133 plots and 130 species of herbs, mosses, and

liverworts in 1197 subplots (Appendix 1). Of these

plants, 41 were classified as wetland-obligate species and

23 were wetland facultative (Appendix 1) (Reed 1988).

Wetland index values were averaged for plot and subplot

data to equally weigh the influence of woody and

herbaceous plants. Values ranged from 1.69-3.29, with

the smaller, isolated sites rating higher (i.e., fewer

wetland-obligate species) than the larger, more open sites

(Table 2). However, wetland size did not influence the

average number of woody (r = -0.201; P = 0.396) and

herbaceous (r = -0.165; P = 0.486) plants per plot and

subplot, respectively (Table 2).
8

In a principal components analysis of the combined

vegetation plot and subplot data, the first component

explained 20.4 percent of total character variance, and

separated out Abe Run (Canaan Valley State Park,

Tucker County, WV) from the remaining 19 (Fig. 2).

Abe Run contained several disturbance-tolerant species

not restricted to wetland habitats, as well as additional

species unique to this site. Twenty-eight species exhibited

loadings above 0.98 (representative species listed in Fig.

2). No plant species exhibited similar high loadings on

the negative end of the axis, but herbs and mosses with

the most negative loadings included Sphagnum moss

(eigenvalue of -0.3853), cottongrass (Eriophorum

virginicum, -0.2377), and ground pine (Lycopodium

obscurum, -0.1933). Trees and shrubs with the most

negative loadings included groundberry (Rubus sp., -

0.4337), possumhaw (Viburnum nudum, -0.2828), black

chokeberry (Photinia melanocarpa, -0.2537), and

velvetleaf huckleberry (Vaccinium myrtilloides, -0.2528).

The second component explained 10.5 percent of the

variance, and separated High Ridge (Dolly Sods, Tucker/

Randolph County, WV) from the remaining sites.

Eighteen plant species exhibited loadings above 0.90;

however, no patterns were apparent among the

remaining 19 sites loaded on this axis. The third

component comprised 8.3 percent of the character

variance, and separated sites on a geographical basis.

Canaan Loop Road sites clustered at the positive end of

the axis, whereas the Maryland sites were located toward

the negative end of the axis. Several representative species

most common at the Maryland sites, all with loadings of

-0.6965, included black cherry (Prunus serotina),

intermediate woodfern (Dryopteris intermedia),

blackgum (Nyssa sylvatica), smartweed (Polygonum

persicaria), and common oats (Avena sativa). Oats,

however, were planted at the site as part of an acid mine

drainage remediation effort. Species most often found at

the Canaan Loop Road sites include red spruce

(0.6531), lichens (0.6546), hygrohypnum moss

(Hygrohypnum eugyrium, 0.5902), bracken fern

(Pteridium aquilinum, 0.5830), possumhaw (Viburnum

nudum, 0.5621), velvetleaf huckleberry (0.5082), and

ground pine (Lycopodium obscurum, 0.4868).

Table 2.—Vegetative measurements for 20 wetland sites in West Virginia and Maryland, collected in 2001.

Wetland index value based on averaged plot and subplot values.

Site Plots sampled Herbaceous Woody species Mean herbaceous Mean woody Wetland

species species/plot species/plot index

ABER 5 44 14 7.8 5.4 2.82

ALDR 6 18 10 5.6 5.0 1.90

BEAL 4 39 7 6.0 4.0 2.07

BIGR 8 27 13 4.5 5.0 1.69

COND 4 26 10 5.6 6.0 2.12

CRSW 10 28 15 4.4 5.6 1.80

FISH 12 32 19 5.7 6.1 1.92

GLAD 24 47 18 3.7 3.9 2.04

HAMM 6 11 6 3.7 2.8 1.95

HERZ 8 25 7 4.4 3.4 2.20

HIGH 8 44 17 5.9 5.6 1.99

MAIN 10 21 16 3.7 8.0 1.93

MOOR 4 13 12 3.8 6.8 1.82

NORT 8 26 14 5.1 6.8 2.19

POWR 2 13 9 4.1 7.5 3.29

REDR 3 15 9 5.1 8.7 2.06

TRL1 2 16 10 5.1 7.0 2.91

TRL2 2 11 7 3.2 5.0 2.37

TRL3 3 13 9 3.0 7.0 2.08

YELL 4 15 6 2.9 3.5 2.02
9

Soil and Water Analyses

Several mean chemistry measures varied considerably

across sites, including soil pH (3.14–5.80), redox

potential (-24 – +370 mV), and dissolved oxygen (6.9–

110.7 percent; Table 3). Organic matter ranged from

20.3–75.6 percent, and trends were evident across sites

(Table 4). Sites with flowing water, the Beall Tract

(Canaan Valley NWR, Tucker County, WV) and

Condon Run (Otter Creek, Randolph County, West

Virginia), had lower organic matter. Sites with ponded

water, seeps, or no visible surface water had higher

organic matter.

Soil profiles were recorded for 2–22 points per site, one

in each vegetation plot (Table 5). Because depth to grey

matrix matched the depth just beyond the organic layer

for nearly every sample, we assumed in further

discussions that reduced conditions exist for all mineral

soils directly underlying the organic layer. The depth of

the organic layer was greater than 20 cm in plots at 19 of

20 sites and in 77.7 percent of all samples. The only

wetland lacking a 20-cm organic layer was the Powerline

site (Canaan Loop Road, Tucker County, WV), which

was the smallest (0.08 ha) of the sites. The organic-layer

depth was greater than 40 cm (a histosol) in plots at 12

of the 20 sites and in 30.0 percent of all samples.

Analyzing particle size, nine soil textural classes were

documented. Sandy clay loam and sandy loam were

found most often in the 130 samples (Table 5). In

comparing clay percentage to organic carbon percentage

within each sample, 9.3 percent were mineral soil, 25.6

percent were mucky mineral, and 65.1 percent were

organic (Fig. 3). Only Condon Run and the Beall Tract

(both containing creeks) produced samples that were

entirely mineral or mucky mineral soils. All samples

from four wetlands from the Canaan Loop Road

complex (Main, Red Run, Trail 3, and Powerline) were

organic. In comparison with other soil analyses, three of

these four sites (excluding Main) lacked histosols

(organic layer greater than 40 cm), and the Powerline site

had an even shallower (4-8 cm) organic layer. The

remaining 13 sites contained mixtures of mineral and

organic soils.

Component I 

(20.4%)

-0.15 -0.10 -0.05 0.00 0.05 2.00

Component III

(8.3%)

-0.75

-0.50

-0.25

0.00

0.25

0.50

ABER ALDR

BEAL

BIGR

COND

CRSW

FISH

GLAD

HAMM

HERZ

HIGH

MAIN

MOOR

NORT

POWR

REDRTRL1

TRL2

TRL3

YELL

Asclepius syriaca 

Betula nigra

Carex sp. (3) 

Impatiens capensis 

Leucanthemum vulgare

Scirpus atrovirens

Typha latifolia

Eriophorum virginicum

Kalmia latifolia

Lycopodium obscurum

Photinia melanocarpa

Rubus sp. (groundberry)

Sphagnum sp.

Vaccinium myrtilloides

Viburnum nudum

Hygrohypnum eugyrium

Lycopodium obscurum

Picea rubens

Pteridium aquilinum

unidentified lichen

Vaccinium myrtilloides

Viburnum nudum

Avena sativa

Dryopteris intermedia

Lolium perenne

Nyssa sylvatica

Polygonum persicaria

Prunus serotina

Rumex acetosella

Figure 2.—Principal components analysis of combined vegetation plot and subplot data

collected in 2001, evaluating I-values of each plant species at the 20 high elevation wetlands.

Encircled sites indicate those geographically close to one another. Plant species exhibiting

high positive or negative loadings are listed on appropriate axes.
10

Table 3.—Average water chemistry measures for 20 wetland sites in West Virginia and

Maryland in August 2001. Temperature and dissolved oxygen are standardized to

approximate readings at 1200.

 Site Temperature DO pH Conductivity ORP

(°C) (% saturation) (mS/cm) (mV)

ABER 14.8 110.7 5.80 0.141 149.1

ALDR 24.2 97.1 3.68 0.023 255.8

BEAL 11.4 79.1 4.23 0.010 249.7

BIGR 22.7 43.2 3.68 0.034 220.1

COND 17.5 18.4 3.95 0.022 234.9

CRSW 18.5 18.0 4.16 0.035 -3.7

FISH 20.4 31.7 4.82 0.018 134.8

GLAD 23.0 102.3 4.72 0.249 115.6

HAMM 17.3 18.6 5.89 0.133 -24.0

HERZ 21.9 35.7 4.49 0.091 128.7

HIGH 22.1 24.9 5.21 0.025 68.1

MAIN 21.3 53.6 4.43 0.051 251.7

MOOR 18.4 69.4 3.83 0.020 275.6

NORT 21.8 45.0 4.46 0.049 229.1

POWR 25.2 73.2 3.15 0.056 370.0

REDR 17.8 52.6 3.14 0.027 271.5

TRL1 21.6 47.2 3.41 0.012 306.2

TRL2 16.4 59.2 3.21 0.044 320.9

Table 4.—Soil and hydrology measures for 20 sites in West Virginia and Maryland, measured in 2001-2002.

Average water table depth computed from measures taken May 2001 – June 2002. Remaining averages based

on 3 - 24 soil samples analyzed per site. Categories “>20 cm organic” and “>40 cm organic” refer to percentage

of plots with organic depth greater than 20 and 40 cm, respectively.

Site % Carbon % Nitrogen C:N % Organic Organic >20 cm >40 cm Mean water table

ratio matter depth (cm) organic (%) organic (%) depth (cm)

ABER 18.4 0.96 19.2 38.4 31 100 20 -18.1

ALDR 24 1.37 17.5 50.0 33 100 17 -15.9

BEAL 9.7 0.63 15.4 20.3 20 50 0 -5.1

BIGR 18.7 0.98 19.1 38.9 38 88 38 -0.9

COND 11 0.55 20.1 23.0 52 100 75 -6. 0

CRSW 36.3 1.98 18.3 75.6 74 100 90 -26.8

FISH 19 1.11 17.1 39.5 30 100 0 -9.9

GLAD 27.9 1.59 17.6 58.2 52 91 50 -15.8

HAMM 15.7 1.2 13.1 32.7 55 100 33 -24.3

HERZ 17.4 0.86 20.2 36.2 15 13 13 -13. 0

HIGH 19.3 1.52 12.7 40.1 26 63 13 -15.9

MAIN 33.5 1.3 25.8 69.8 25 63 25 -10.9

MOOR 18.2 0.88 20.7 37.9 39 100 50 -11.4

NORT 20.2 0.88 22.9 42.0 20 50 0 -9

POWR 26.8 1.28 21.0 55.9 6 0 0 -10.8

REDR 30.1 1.33 22.6 62.6 21 100 0 -8.9

TRL1 13.3 0.72 18.5 27.7 17 50 0 -14.8

TRL2 18.8 1.05 17.9 39.2 25 50 0 -17.2

TRL3 24.4 1.37 17.8 50.8 19 33 0 -7.3

YELL 18.6 0.89 20.9 38.8 40 100 75 -11
11

Table 5.—Soil texture distribution for 20 wetland sites in West Virginia and Maryland, collected in 2001.

Numbers refer to proportion of plots in which textural class existed.

Clay Loam Sand 

Site clay sandy clay clay loam silty clay sandy clay loam sandy loamy sand

loam loam loam sand

ABER – – – – 0.400 – 0.400 0.200 –

ALDR – – – – 0.333 – 0.667 – –

BEAL – – – – 1.000 – – – –

BIGR – – – 0.125 0.625 – 0.250 – –

COND – – – – 0.250 – 0.500 0.250 –

CRSW – 0.200 – – 0.300 – 0.300 0.200 –

FISH 0.083 0.250 – – 0.500 – 0.167 – –

GLAD – – – – 0.364 – 0.364 0.091 0.182

HAMM – – – – – – 0.667 0.167 0.167

HERZ 0.250 0.250 0.125 – 0.125 – 0.250 – –

HIGH – – – – 0.500 0.125 0.250 – 0.125

MAIN – – – – 0.375 – 0.500 0.125 –

MOOR 0.250 – – – 0.250 – 0.250 0.250 –

NORT – – – – 0.500 – 0.500 – –

POWR – 0.333 – – 0.667 – – – –

REDR – – – – – – 0.333 0.667 –

TRL1 – – – – 1.000 – – – –

TRL2 – – – – 0.333 – 0.333 0.333 –

TRL3 – – – – – – 0.667 – 0.333

YELL – – – – 0.500 – – 0.500 –

Percentage clay

0 10 20 30 40 50 60

Percentage carbon

0

5

10

15

20

25

30

35

40

45

50 Organic

Mucky

mineral 

Mineral

Figure 3,—Percentage organic carbon versus percentage clay for 130 soil samples

collected at 10 cm depth in 2001 from the 20 high elevation wetland sites. Soil

samples are classified as organic, mineral, or mucky mineral.
12

Hydrological Measures

Considerable interseasonal variation (Fig. 4B, 4C) to

almost no variation (Fig. 4A, 4D) was observed. Eight of

the 20 sites showed a marked height decrease (-30 cm) in

the water table outside of the growing season. Three sites

(Herz Tract, High Ridge, and Trail 2) fell below -50 cm.

Using the most restrictive definition of 15 cm from the

surface for 5 percent of the year, then 19 of 20 sites met

the requirements for a wetland in 2001 and 2002. At the

remaining site, Cranesville Swamp, the water table

remained approximately 20 cm below the surface for

most of the 2001 growing season but met the

requirement in the 2002 growing season. Additionally,

Date

6/1/01

8/1/01

10/1/01 

12/1/01 

2/1/02

4/1/02

6/1/02

8/1/02

Water table level (cm)

-15

-12

-9

-6

-3

0

3

6

A

Date

6/1/01 

8/1/01 

10/1/01 

12/1/01 

2/1/02 

4/1/02 

6/1/02 

8/1/02 

Water table level (cm)

-80

-60

-40

-20

0

B

Date

6/1/01 

8/1/01

10/1/01 

12/1/01 

2/1/02 

4/1/02 

6/1/02 

8/1/02 

Water table level (cm)

-40

-30

-20

-10

0

C

Date

7/1/01 

9/1/01 

11/1/01 

1/1/02 

3/1/02 

5/1/02 

7/1/02 

Water Level (cm)

-25

-20

-15

-10

-5

0

D

D a te

5/1/01 

7/1/01 

9/1/01 

11/1/01 

1/1/02 

3/1/02 

5/1/02 

7/1/02 

9/1/02 

Rainfall (cm)

0

2

4

6

8

E

Figure 4.—Water table depth versus time for four wetland sites in the Allegheny Mountains of West

Virginia and Maryland, monitored May 2001 – September 2002: (A) BIGR, (B) HERZ, (C) TRL3,

and (D) ABER (see text). Solid circles, connected by solid line, indicate current readings taken on

specified dates. Open circles plotted above solid circles indicate highest level reached since last reading.

Dashed horizontal line indicates soil surface. Also plotted is rainfall hydrograph (E) for sampling period.
13

several areas within 10 m of the installed

piezometer remained flooded for a large

portion of both seasons.

Long-term precipitation trends revealed that

2001 and 2002 were drier than the long-term

average of 134 cm of precipitation per year

(Stephenson 1993). The hydrologic years of

2000 (May 2000 – April 2001), 2001 (May

2001 – April 2002) and 2002 (May 2002 –

April 2003) recorded 98, 118, and 123 cm of

precipitation, respectively. When examining

day-by-day records of precipitation for the

periods in which piezometers were installed and

monitored, it is unclear whether wetland water

levels responded to specific rain events (Fig.

4E).

Geologic Interpretation

Fifteen of the study wetlands were dominated

by underlying layers from the Pottsville Group,

that contain a sandstone and shale layer that is

highly erosion-resistant (Table 6, Appendix 2).

Two sites in Canaan Valley contained an older

sandstone layer from the Pocono group. The

three remaining sites, Abe Run, Cranesville

Swamp, and Hammel Glade (Garrett County,

MD), contained rock from the Greenbrier

Group, that includes a significant amount of

limestone mixed with sandstone and shale. In

an examination of topographical and soil maps,

several trends were noted. All sites were situated

within small basins, up to 30 m lower in elevation than

surrounding landscapes. Additionally, many of the sites

were located near the headwaters of streams. Only one

site of the 17 examined for mapped soils in West

Virginia contained muck and peat—Fisher Springs Run

(Dolly Sods, Tucker County, WV). The remaining sites

were designated as a variety of series that contain poorly

draining and/or acidic soils. These include the Lickdale

(deep, poorly-drained soils in upland depressions), Blago

(deep, poorly-drained soils formed in acidic conditions

and sandstone weathering), and Brinkerton (deep,

somewhat poorly drained soils developed in acidic

material) series.

Comparison Among Vegetation, Soils,

and Hydrology

Pearson’s correlations revealed significant positive

relationships among several vegetative variables

(principal components loadings) and water chemistry

variables (i.e., pH, dissolved oxygen, conductivity),

indicating chemical gradients along vegetative

community types (Table 7). Water chemistry factors

were closely correlated to one another. For example,

increases in pH corresponded to an increase in

conductivity and a decrease in redox potential (Table 7).

Additionally, pH readings were negatively correlated to

average water table depth and the C:N ratio (table 7).

Organic matter depth was negatively correlated to the

wetland vegetative index, indicating that sites with

deeper organic layers contained a greater proportion of

wetland-obligate or wetland-facultative species (Table 7).

Organic depth also was related positively to the third

principal component of the vegetation analysis,

indicating a relationship between soil composition and

vegetative community composition (Table 7).

In a principal components analysis of vegetation, soils,

and hydrology compared in Table 7, the first component

explained 33.0 percent of total character variance (figure

5). Variables with high negative loadings for this first

component include pH (-0.7324), and conductivity

(-0.8433). Water-table depth (0.6395), ORP (0.6996),

and the third component of the vegetation PCA

(0.7736) exhibited high positive loadings for this

component. This component separated out the three

Maryland sites, plus Abe Run, from the remaining

Table 6.—Principle underlying geology for 20 wetlands in West

Virginia and Maryland. Primary layer indicates that which

dominates by wetland area. Secondary and tertiary layers, if

present, are found in increasingly smaller portions of the site.

Appendix 2 lists code descriptions.

Site Primary Secondary Tertiary

layer layer layer

ABER Mg

ALDR Ppv

BEAL Mp

BIGR Pa Pk

COND Pnr Pk

CRSW Mg Mp Mmc

FISH Pc

GLAD Pc Ppv Pa

HAMM Mg Mmc Mp

HERZ Mp

HIGH Pk Pa Pc

MAIN Ppv

MOOR Pnr Pk

NORT Ppv

POWR Ppv

REDR Ppv

TRL1 Ppv

TRL2 Ppv

TRL3 Ppv

YELL Pnr Pk 
14 Table 7.—Correlation analysis of soil, hydrology, and vegetation measures from 20 wetland sites in West Virginia and 

Correlation (r) listed for each pair with P-value in parentheses (P ≤ 0.1 in bold).

 Vegetation Soils and Hydrology

Veg. Veg Veg. Wetland ORP Temp. Cond’y DO% pH C:N 

PC III PC II PC I Index Ratio 

Soils/Hydrology

Mean water 0.276 0.028 -0.184 -0.075 0.489 0.091 -0.377 -0.008 -0.388 0.354 

depth (0.238) (0.907) (0.438) (0.752) (0.029) (0.704) (0.101) (0.973) (0.091) (0.126) 

Organic depth -0.595 0.094 0.007 -0.530 -0.505 -0.194 0.296 -0.148 0.268 -0.246 

(0.006) (0.695) (0.976) (0.016) (0.023) (0.413) (0.204) (0.533) (0.254) (0.295) 

Mean %C 0.057 0.193 -0.156 -0.128 -0.158 0.378 0.159 0.028 -0.129 0.394 

(0.811) (0.415) (0.510) (0.590) (0.507) (0.100) (0.503) (0.906) (0.588) (0.085) 

Mean %N -0.210 -0.126 -0.153 -0.224 -0.505 0.338 0.241 -0.094 0.136 -0.159

(0.375) (0.598) (0.520) (0.342) (0.023) (0.145) (0.306) (0.693) (0.568) (0.503) 

C:N Ratio 0.481 0.477 -0.026 0.130 0.542 0.205 -0.103 0.166 -0.441

(0.032) (0.033) (0.912) (0.585) (0.013) (0.386) (0.664) (0.485) (0.052) x

pH -0.518 -0.279 0.479 -0.150 -0.755 -0.241 0.483 -0.086

(0.019) (0.233) (0.033) (0.529) (0.001) (0.306) (0.031) (0.718) x

DO% -0.161 0.216 0.446 0.296 0.468 -0.058 0.405

(0.499) (0.360) (0.049) (0.204) (0.038) (0.807) (0.077) x

Conductivity -0.662 0.235 0.307 0.102 -0.330 0.070

(0.001) (0.318) (0.188) (0.669) (0.155) (0.770) x

Temperature 0.177 -0.070 -0.409 0.073 0.093

(0.455) (0.771) (0.073) (0.758) (0.697) x

ORP 0.553 0.214 -0.106 0.399

(0.011) (0.366) (0.657) (0.081) x

VegetationWetland Index 0.443 0.150 0.356

(0.051) (0.529) (0.123) x

Veg. PC I 0.000 0.000

(1.000) (1.000) x

Veg. PC II 0.000

(1.000) x
15

wetlands. The second component explained 18.3 percent

of total character variance. Nitrogen percentage (0.5049)

and the organic layer depth (0.4676) exhibited high

positive loadings on this axis, whereas percent dissolved

oxygen (-0.7318), ORP (-0.5315), wetland index

(-0.7021), and the first component of the vegetation

PCA (-0.5929) had high negative loadings. This

component separated Abe Run and Cranesville Swamp

(at opposite extremes) from the remaining sites. The

third component explained 14.7 percent of total

character variance, but no major trends among sites or

among variables were evident.

Discussion

Assessment of Current Data

When examining vegetation, soils, and hydrology as a

whole, all sites fit the biological definition of a wetland

(Cowardin et al. 1979). Nearly all sites (18 of 20)

exhibited wetland vegetative index values less than 2.5,

and those with values more than 2.5 met all hydrology

and hydric soils requirements. Topography and geology

appear to be the driving factors for the presence and

condition of these wetlands. Topographically, all sites

were situated within locally concave pockets, which

provide a pooling site for precipitation and encourage

growth of wetland-obligate and wetland-facultative

vegetation. Geologically, 17 of the 20 sites are underlain

by a hard sandstone and shale layer that impedes

drainage and encourages water table levels to be near the

soil surface. The remaining three sites, Abe Run,

Hammel Glade, and Cranesville Swamp, also contain

limestone, which typically is associated with less acidic

soils. This was true for Abe Run and Hammel Glade,

whose soil pH readings were markedly higher (5.80 and

5.89, respectively) than other sites. However, at the

sample sites chosen for Cranesville Swamp, Pocono

Sandstone overlaid the Mauch Chunk (limestone) layer.

Therefore, the direct influence of limestone was not

represented in the soil pH.

Piezometer data revealed that most sites are

hydrologically similar. Nearly all wetlands maintained a

water table at or near the surface for substantial portions

of the 2001 and 2002 growing seasons. Hydrologic data

indicate that these study years, and the few preceding

them, were relatively dry when compared to long-term

trends. Though we acknowledge that the difference in

elevation between recording stations at Canaan Valley

Principal Component I 

(33.0%)

-1.5 -1.0 -0.5 0.0 0.5

Principal Component II 

(18.3%)

-1.0

-0.5

0.0

0.5

1.0

ABER

ALDR

BEAL

BIGR

COND

CRSW

FISH

GLAD

HAMM

HERZ

HIGH

MAIN

MOOR

NORT

POWR

REDR

TRL1

TRL2

TRL3

YELL

Vegetation PC III

WaterTable Depth

ORP

pH

Conductivity

Wetland Index

Vegetation PC I

DO%

ORP

Organic Depth

%Nitrogen

Figure 5.—Principal components analysis of vegetation, soils, and hydrology at the 20 wetland sites.

Variables exhibiting high positive or negative loadings listed on appropriate axes.
16

and Beardon Knob might account for some of the

differences in precipitation, the recent years were still

markedly drier than normal. Thus, despite the lower

precipitation recorded in each year, these sites still

maintained water table levels at appropriate levels for

typical wetlands.

It appears that even sites greatly influenced by limestone

contain an underlying clay or rock layer that maintained

a perched water table. Therefore, bedrock might not

directly influence the presence of a wetland in all cases,

but clearly affects localized soil chemistry (Webb et al.

1997).

Webb et al. (1997) acknowledged that bedrock type

influences water chemistry and soils. Because vegetation

communities partially are driven by soil properties, one

would observe an indirect relationship between the

bedrock layer and vegetation distribution. Indeed, we

found that many of these study sites are similar for a

number of vegetation, soils, and hydrology parameters.

These wetlands exhibited a great deal of intrasite

variability, in terms of soil texture, organic-layer depth,

and nutrient concentrations. Even sites smaller than 2 ha

contained up to three soil textural classes and a variable

range of organic depths (e.g., 5-30 cm at Beall Tract, 27-

100+ cm at Hammel Glade). Similarly, Darlington

(1943) found 2 m differences in peat depth across

Cranberry Glades. Intrasite variation could be attributed

to many physical and geological factors recorded at our

sites, including direct input from creeks, local

underlying rock types, or slight elevational changes,

which would influence depth to the water table. The

frequency of fire also would regulate peat accumulation

in these systems. It is possible that sizeable within-site

differences could be found with additional survey work

that examined the organic layer below 1 m.

Previous estimates of peatland area failed to acknowledge

the presence of histosols in West Virginia, indicating that

many of the smaller wetlands examined here were

overlooked (Soil Survey Staff, 1998b). However, the

presence of histosols at more than half the sites

(including six sites less than or equal to 5 ha) illustrates

the lack of small-scale scrutiny.

Therefore, wetland size appears to be an influential

factor in delineating wetlands, but fails to acknowledge

the equally important smaller sites that show similar

wetland soil characteristics. Additionally, vegetative

assessments showed that wetland size did not influence

the number of species per unit area or the diversity

index, emphasizing that small wetlands should not be

overlooked in preservation efforts.

Wetlands: Past and Present

A review of the geologic literature available for this

region supports our findings for the presence and

formation of these wetlands. Diehl’s (1981) examination

of 49 wetlands in the unglaciated Allegheny Plateau

(west-central West Virginia) and the Allegheny

Mountain (eastern West Virginia) provinces revealed that

hard sandstone layers directly underlie the majority of

sites within our study area. These wetlands often are at

headwaters of mature rivers that travel across originally

flat-lying strata. Typically, water flows across these

horizontal, unsloped strata until a basin is eroded away

and water ponds up. The basin reaches its maximum

depth when eroding strata, such as Mauch Chunk or

Greenbrier limestone, are removed and less penetrable

sandstone layers are exposed (Diehl 1981). Given the

low permeability of some sandstones, the water table

becomes perched and can reach the surface.

Given the stable geology of the systems, one must

examine other factors that have shaped the current state

of these wetlands. Logging of red spruce forests a century

ago markedly altered the landscape. Additionally,

wetlands in the Dolly Sods Scenic Area and Cranesville

Swamp were influenced by livestock grazing, fire, beaver

activity, and anthropogenic species introductions

(Fortney 1975). Furthermore, the impact of exotic

insects such as the balsam woolly adelgid, (Adelges piceae)

or hemlock woolly adelgid, (Adelges tsugae), heavy deer

browse on saplings, and recent human activities, such as

coal mining and subsequent acid-mine drainage, inhibits

some species’ ability to sustain healthy populations

(Michael 2000). One site in particular, the Glades, has

been notably impacted by acid mine drainage and is

currently undergoing long-term remediation (West

Virginia Department of Environmental Protection 1997).

Validation for Wetland Preservation

When Fortney (1975) studied the plant communities of

Canaan Valley, he found that some wetland habitats

contained the highest plant species diversity. Speckled

alder (Alnus incana) thickets, such as those found at

Alder Run (Dolly Sods, Tucker County, WV), High

Ridge, and the Glades, commonly contained at least 45

associated species of herbaceous vegetation. Other

associations dominated by meadowsweet (Spiraea alba)

or shrubby St. John’s wort (Hypericum densiflorum), such

as Herz Tract, may contain fewer plant species, but

provide sufficient cover and nesting sites for several

uncommon small mammals (Fortney 1993), including

meadow jumping mice (Zapus hudsonius) and southern

bog lemmings (Synaptomys cooperi). Wetlands containing

open water, including Moore Run (Otter Creek,

Randolph County, West Virginia) and Fisher Springs,
17

also are valuable sites for breeding herpetofauna

(Semlitsch and Bodie 1998).

In our study, we found that small, isolated wetlands

exhibit hydrological and soil characteristics similar to

larger, recognized sites and serve as patchily distributed

refugia for wetland-obligate species in the area. Semlitsch

and Bodie (1998) address the importance of small

wetlands in the Southeast, emphasizing that they are not

expendable if regional biodiversity is to be maintained.

Sites smaller than 1.2 ha comprised 46.4 percent of the

371 Carolina bays examined, and nearly 90 percent were

less than 4.0 ha. Similar high proportions were found in

Maine wetlands, in which 62 percent of the sites were <

4.05 ha (Gibbs 1993). Small wetlands such as these are

significant contributors to amphibian species richness

and diversity, and harbor large numbers of wetland

plants and aquatic insects with limited dispersal abilities

(Semlitsch and Bodie 1998). The destruction of such

habitats would increase dispersal distances between

suitable sites and likely isolate these plants and animals

from larger wetlands serving as source populations. As

local extinction rates increased, genetic diversity could

decline (Semlitsch and Bodie 1998). Lower genetic

diversity would mean that the ability of a species to

adapt to changing environmental conditions also would

decrease (Hansen et al. 2001).

Legal Implications

Isolated wetlands are considered some of the most

important and threatened ecosystems in the country

(Osmond et al. 1995). Unfortunately, few state or federal

laws protect them. Since a 2001 U.S. Supreme Court’s

decision (Solid Waste Agency of Northern Cook County v.

U.S. Army Corp of Engineers), Section 404 of the Clean

Water Act no longer protects isolated wetlands such as

those examined in this study (Bedford and Godwin

2003). Because the U.S. Army Corps of Engineers lacks

jurisdiction over isolated sites, like those in this study,

any further regulation must be the responsibility of the

states (through Section 404(g)), which may implement

their own permit programs over non-navigable waters

(Osmond et al. 1995). However, no recognized statutes

or programs in West Virginia have been created explicitly

to protect isolated wetlands as has been done in many

states (Christie and Hausman 2003), unless affected by

acid mine drainage from coal-mining operations (West

Virginia Annotated Code § 22-11-7a, 2001). In

Maryland, state wetlands law is limited to navigable

waters, and reiterates similar restrictions of the federal

Clean Water Act.

Due to the lack of explicit legal protection of isolated

wetlands, it is important to determine how central

Appalachian wetlands should be protected. The selected

study sites are either government-owned (e.g., part of the

Monongahela National Forest, Canaan Valley National

Wildlife Refuge, Canaan Valley Resort State Park) or

privately owned by The Nature Conservancy. Both

owner-types protect the sites through government

regulations or self-regulated policies. However, many

privately owned wetlands exist within this region and do

not fall under any protection guidelines (Diehl 1981,

Ingham 1996). Currently, the conventional method to

preserving wetlands regionally on private property is for

private conservation groups, such as The Nature

Conservancy, or governmental agencies to purchase the

land. In 2002, the USDI Fish and Wildlife Service

purchased a large parcel in Canaan Valley from

Allegheny Power, increasing the refuge’s total wetlands to

2175 ha3

. However, not all wetlands can be or should be

purchased and preserved by public means. Potentially

beneficial programs include educating private

landowners in the utility of these wetlands (e.g.,

floodwater storage and filtration of heavy metals and

organic materials) or exploring potential future financial

incentives through mitigation banking or carbon

sequestration.

Conclusions

Although other researchers have studied several of these

wetlands previously, our study is the first to compare a

large number of sites on a quantitative basis. Many of

the quantitative differences could be explained

qualitatively in terms of geology and topography. These

two factors combined to create favorable conditions for

wetlands. Variability in geology led to differentiation

among vegetative communities and soil and water

properties.

These data support previous research, in that central

Appalachian fens and bogs are similar to boreal bogs and

fens of the northern United States and Canada. There

exists an overlap of many vegetative species and chemical

characteristics. However, all study sites from the central

Appalachians lacked the concentric zonation of many

northern bogs and exhibited a greater degree of withinsite variation in terms of soil texture, organic depth, and

distribution of vegetation. Moorhead and Rossell’s

(1998) description of southern Appalachian sites as an

“irregular mosaic” also appears to fit these 20 central

Appalachian wetlands (Wieder et al. 1981).

Based upon the survey of vegetation, soils, and

hydrology, we conclude that all sites qualify as wetlands.

3

Sturm, K. 2003. Personnal communication. Wildlife

Biologist, U.S. Fish and Wildlife Service, Canaan Valley

National Wildlife Refuge, HC 70 Box 200, Davis, WV 26260.
18

Future management decisions should incorporate the

impacts on these isolated systems when making

landscape-level changes. Furthermore, wetland size

should not be a determining factor in decisions to

preserve sites, as we have shown that small wetlands are

just as important ecologically as thelarger counterparts.

Localized management efforts, such as removing

invading upland or exotic species or minimizing foot

traffic in these wetlands, might be necessary. However, in

most cases simple preservation and protection may be

the best tool to sustain or reach desired conditions.

Acknowledgments

We thank the Georgia Museum of Natural History, the

Phillips Soil Lab, and the D. B. Warnell School of Forest

Resources at the University of Georgia for providing

funding and/or equipment necessary to conduct this

research. The Canaan Valley NWR also provided in-kind

contributions of housing, equipment, and field

assistance and we are grateful to K. Sturm and his staff

for their helpful efforts. The Nature Conservancy

allowed access to its properties for this study. We thank

S. Dumpert, R. Kotecki, H. Hadley, J. Wykle, N.

Castleberry, J. Rodrigue, R. Smith, K. Warren, and K.

Dodge for assistance in the field. Lastly, M. B. Adams, L.

Morris, M. Campbell, C. R. Jackson and S. Studlar

provided needed methodical and analytical assistance.

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Appendix 1

Plants collected in vegetation surveys in 2001 at 20 wetlands in West Virginia and Maryland. Names listed in

accordance with PLANTS database (USDA—NRCS, 2002). Values indicate ecological indicator values used in

weighted analyses.

Family Scientific name Author Value

Non-vascular plants

Amblystegiaceae Hygrohypnum eugyrium (Schimp. in B.S.G.) Loeske 

Dicranaceae Dicranum scoparium Hedw.

Fontinalaceae Fontinalis dalecarlica Schimp. in B.S.G. 

Hypnaceae Hypnum imponens Hedw.

Lepidoziaceae Bazzania trilobata (L.) Gray

Leucobryaceae Leucobrym albidum (Brid. ex P. Beauv.) Lindb.

Leucobryaceae Leucobrym glaucum (Hedw.) Angstr.

Polytrichaceae Polytrichum commune Hedw.

Scapaniaceae Scapania undulata (L.) Dumort. 

Sphagnaceae Sphagnum capillifolium (Ehrh.) Hedw. 1.00

Sphagnaceae Sphagnum fallax (Klinggr.) Klinggr. 1.00

Sphagnaceae Sphagnum palustre L. 1.00

Sphagnaceae Sphagnum sp. 1.00

Vascular plants

Aceraceae Acer pensylvanicum L. 4.00

Aceraceae Acer rubrum L. 3.00

Aquifoliaceae Ilex montana Torr. & Gray ex Gray

Aquifoliaceae Ilex verticillata (L.) Gray 1.67

Aquifoliaceae Nemopanthus mucronatus (L.) Loes. 1.00

Araceae Arisaema triphyllum (L.) Schott 2.33

Araceae Orontium aquaticum L. 1.00

Araceae Symplocarpus foetidus (L.) Salisb. ex Nutt. 1.00

Asclepiadaceae Asclepias syriaca L. 4.33

Aspleniaceae Asplenium montanum Willd. 5.00

Asteraceae Achillea millefolium L. 4.00

Asteraceae Symphyotrichum praealtum var. (Poir.) Nesom 2.00

praealtum

Asteraceae Symphyotrichum puniceum var. (L.) A.& D. Löve 1.00

puniceum

Asteraceae Eupatorium fistulosum Barratt 2.00

Asteraceae Euthamia graminifolia (L.) Nutt. 3.00

Asteraceae Leucanthemum vulgare Lam. 5.00

Asteraceae Solidago rugosa P. Mill 3.00

Asteraceae Solidago uliginosa Nutt. 1.00

Balsaminaceae Impatiens capensis Meerb. 2.00

Betulaceae Alnus incana ssp. rugosa (L.) Moench (Du Roi) Clausen 2.00

Betulaceae Betula lenta L. 4.00

Betulaceae Betula nigra L. 2.00

Caprifoliaceae Viburnum dentatum L. 3.00

Caprifoliaceae Viburnum nudum L. 1.00

Caryophyllaceae Galium obtusum Bigelow 1.67

Caryophyllaceae Stellaria graminea L. 4.33

Clusiaceae Hypericum canadense L. 2.00

Clusiaceae Hypericum densiflorum Pursh 2.67

Continued
23

Clusiaceae Hypericum ellipticum Hook 1.00

Clusiaceae Hypericum mutilum L. 2.00

Clusiaceae Hypericum punctatum Lam. 3.33

Clusiaceae Hypericum sp.

Clusiaceae Triadenum virginicum (L.) Raf. 1.00

Convolvulaceae Ipomoea sp.

Cyperaceae Carex atlantica ssp. atlantica Bailey 1.67

Cyperaceae Carex baileyi Britt. 1.00

Cyperaceae Carex brunnescens (Pers.) Poir. 2.00

Cyperaceae Carex canescens L. 1.00

Cyperaceae Carex debilis Michx. 3.00

Cyperaceae Carex echinata Murr. 1.00

Cyperaceae Carex folliculata L. 1.00

Cyperaceae Carex gynandra Schwein. 1.00

Cyperaceae Carex hirtifolia Mackenzie 

Cyperaceae Carex interior Bailey 1.00

Cyperaceae Carex leptalea Wahlenb. 1.00

Cyperaceae Carex lurida Wahlenb. 1.00

Cyperaceae Carex normalis Mackenzie 4.00

Cyperaceae Carex pauciflora Lightf. 1.00

Cyperaceae Carex rostrata Stokes 1.00

Cyperaceae Carex scoparia Schkuhr ex Willd. 2.00

Cyperaceae Carex stipata Muhl. ex Willd. 1.00

Cyperaceae Carex trisperma Dewey 1.00

Cyperaceae Carex vesicaria L. 1.00

Cyperaceae Eleocharis tenuis Willd.) J.A. Schultes 1.67

Cyperaceae Eriophorum virginicum L. 1.00

Cyperaceae Rhynchospora alba (L.) Vahl 1.00

Cyperaceae Scirpus atrocinctus Fern. 1.67

Cyperaceae Scirpus cyperinus (L.) Kunth 1.67

Dennstaedtiaceae Pteridium aquilinum (L.) Kuhn 4.00

Droseraceae Drosera rotundifolia L. 1.00

Dryopteridaceae Dryopteris cristata (L.) Gray 1.67

Dryopteridaceae Dryopteris intermedia (Muhl. ex Willd.) Gray 4.00

Dryopteridaceae Dryopteris marginalis (L.) Gray 4.33

Dryopteridaceae Dryopteris x boottii (Tuckerman) Underwood (pro sp.) 2.00

Dryopteridaceae Onoclea sensibilis L. 2.00

Ericaceae Epigaea repens L.

Ericaceae Gaultheria hispidula (L.) Muhl. ex Bigelow 2.00

Ericaceae Gaultheria procumbens L. 4.00

Ericaceae Kalmia latifolia L. 4.00

Ericaceae Rhododendron maximum L. 3.00

Ericaceae Vaccinium angustifolium Ait. 4.00

Ericaceae Vaccinium corymbosum L. 2.33

Ericaceae Vaccinium macrocarpon Ait. 1.00

Ericaceae Vaccinium myrtilloides Michx. 3.00

Ericaceae Vaccinium oxycoccos L. 1.00

Ericaceae Vaccinium pallidum Ait. 2.33

Fabaceae Trifoluim sp.

Family Scientific name Author Value

Appendix 1—continued.

Continued
24

Fagaceae Quercus rubra L. 4.33

Gentianaceae Gentiana linearis Froel. 1.00

Hamamelidaceae Hamamelis virginiana L. 4.00

Iridaceae Sisyrinchium angustifolium P.Mill. 2.33

Juncaceae Juncus brevicaudatus (Engelm.) Fern. 1.00

Juncaceae Juncus canadensis J. Gay ex Laharpe 1.00

Juncaceae Juncus effusus L. 1.67

Juncaceae Juncus subcaudatus (Engelm.) Coville & Blake 1.00

Juncaceae Juncus sp.

Juncaceae Luzula acuminata Raf. 3.00

Lamiaceae Lycopus uniflorus Michx. 1.00

Liliaceae Veratrum viride Ait. 1.67

Lycopodiaceae Lycopodium clavatum L. 3.00

Lycopodiaceae Lycopodium obscurum L. 4.00

Nyssaceae Nyssa sylvatica Marsh 3.00

Oenagraceae Ludwigia palustris (L.) Ell. 1.00

Oleaceae Fraxinus pennsylvanica Marsh 2.00

Onagraceae Oenothera fruticosa L. 3.00

Ophioglossaceae Ophioglossum vulgatum L. 2.00

Orchidaceae Cypripedium acaule Ait. 4.33

Orchidaceae Habenaria clavellata (Michx.) Luer 3.67

Orchidaceae Spiranthes cernua (L.) L.C. Rich. 2.00

Osmundaceae Osmunda cinnamomea L. 2.00

Oxalidaceae Oxalis stricta L. 5.00

Pinaceae Abies balsamea (L.) P. Mill. 3.00

Pinaceae Picea abies (L.) Karst. 

Pinaceae Picea rubens Sarg. 4.00

Pinaceae Pinus resinosa Soland. 4.00

Pinaceae Pinus rigida P. Mill 4.00

Pinaceae Pinus strobus L. 4.00

Pinaceae Tsuga canadensis (L.) Carr. 4.00

Poaceae Agrostis gigantea Roth 2.33

Poaceae Anthoxanthum odoratum L. 4.00

Poaceae Avena sativa L. N/A

Poaceae Brachyelytrum erectum (Schreb. ex Spreng.) Beauv. N/A

Poaceae Calamagrostis canadensis (Michx.) Beauv. 1.67

Poaceae Danthonia spicata (L.) Beauv. ex Roemer & 4.00

J.A. Schultes

Poaceae Dichanthelium acuminatum (Sw.) Gould & C.A. Clark 2.00

Poaceae Dichanthelium clandestinum (L.) Gould 2.67

Poaceae Dulichium arundinaceum (L.) Britt. 1.00

Poaceae Glyceria melicaria (Michx.) F.T. Hubbard 1.00

Poaceae Glyceria striata (Lam.) A.S. Hitchc. 1.00

Poaceae Lolium perenne L. 4.33

Poaceae Phalaris arundinacea L. 2.00

Poaceae Phleum pratense L. 4.00

Polemoniaceae Polemonium vanbruntiae Britt. 2.00

Polygonaceae Polygonum persicaria L. 2.00

Polygonaceae Polygonum sagittatum L. 1.00

Family Scientific name Author Value

Appendix 1—continued.

Continued
25

Polygonaceae Rumex acetosella L. 5.00

Ranunculaceae Caltha palustris L. 1.00

Rosaceae Amelanchier laevis Wieg. 2.33

Rosaceae Crataegus sp.

Rosaceae Dalibarda repens L. 3.00

Rosaceae Photinia floribunda (Lindl.) Robertson & Phipps 2.00

Rosaceae Photinia melanocarpa (Michx.) Robertson & Phipps 3.00

Rosaceae Potentilla norvegica L. 4.00

Rosaceae Potentilla simplex Michx. 4.33

Rosaceae Prunus serotina Ehrh. 4.00

Rosaceae Prunus sp.

Rosaceae Rosa multiflora Thunb. ex Murr. 4.00

Rosaceae Rubus–blackberry group

Rosaceae Rubus–dewberry group 2.00

Rosaceae Spiraea alba Du Roi 1.67

Rosaceae Spiraea tomentosa L. 2.33

Salicaceae Populus tremuloides Michx.

Salicaceae Salix alba L. 2.00

Salicaceae Salix nigra Marsh 1.67

Salicaceae Salix sericea Marsh 1.00

Sarraceniaceae Sarracenia purpurea L. 1.00

Sparganiaceae Sparganium erectum ssp. stoloniferum (Graebn.) Hara 1.00

Thelypteridaceae Thelypteris palustris Schott 1.67

Thelypteridaceae Thelypteris simulata (Davenport) Nieuwl. 2.00

Typhaceae Typha latifolia L. 1.00

Betulaceae Betula alleghaniensis Britt. 3.00

Ulmaceae Ulmus sp.

Violaceae Viola sp.

Family Scientific name Author Value

Appendix 1—continued.
26 Appendix 2

Key to geologic codes and dominant rock types within each group or formation. Geologic periods and rock stratyoungest to oldest in geologic time. General lithology also listed for each category (as described by Webb et al. 1Geologic Period Group Formation General Lithology Upper Pennsylvanian Conemaugh 67% Shale/clay; 28% Sandstone; 4% Coal; 1% LimesMiddle Pennsylvanian Allegheny 63% Sandstone, 22% Shale/clay, 10% Coal, 5% LimeLower Pennsylvanian Pottsville Sandstone (some conglomeratic), Shale, Coal Lower Pennsylvanian Pottsville Kanawha 73% Sandstone, 21% Shale/clay, 4% Coal, 0-2% LimLower Pennsylvanian Pottsville New River 72% Sandstone, 23% Shale/clay, 5% Coal Upper Mississippian Mauch Chunk 63% Shale/clay, 33% Sandstone, 4% Limestone Middle Mississippian Greenbrier 94% Limestone, 4% Shale/clay, 2% Sandstone 

Limestone

Lower Mississippian Pocono Sandstone, shale 
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Francl, K.E.; Ford, W.M.; and Castleberry, S.B. 2004. Characterization of high

elevation central Appalachian wetlands. Res. Pap. NE-725. Newtown Square,

PA: U.S. Department of Agriculture, Forest Service, Northeastern Research

Station. 26 p.

We characterized 20 high elevation wetlands in the central Appalachian Mountains

in West Virginia and Maryland, in terms of vegetation, soils, hydrology, and geology.

Plant species were distributed along soil chemical (pH, conductivity) and physical

(organic matter depth) gradients across sites. Topography and geology appear to

explain differences among these wetlands, as reflected by soil and chemistry

measures and vegetation distribution. Our work provides substantial quantitative

baseline data for these uncommon high elevation wetlands, emphasizing the

importance and diversity of these isolated systems.

Key Words: bog, central Appalachian Mountains, fen, geology, hydric soils,

hydrophytic vegetation, wetland characterization
Headquarters of the Northeastern Research Station is in Newtown Square,

Pennsylvania. Field laboratories are maintained at:

Amherst, Massachusetts, in cooperation with the University of Massachusetts

Burlington, Vermont, in cooperation with the University of Vermont

Delaware, Ohio

Durham, New Hampshire, in cooperation with the University of New Hampshire

Hamden, Connecticut, in cooperation with Yale University

Morgantown, West Virginia, in cooperation with West Virginia University

Parsons, West Virginia

Princeton, West Virginia

Syracuse, New York, in cooperation with the State University of New York,

College of Environmental Sciences and Forestry at Syracuse University

Warren, Pennsylvania

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