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Canaan Valley & Environs
2015 Southeastern Naturalist 14(Special Issue 7):187–202
Vegetation of Managed Grasslands in the Canaan Valley
National Wildlife Refuge
Kelly A. Chadbourne1,2 and James T. Anderson1,*
Abstract - Much of Canaan Valley was converted to agricultural uses following logging
in the early 1900s. More recently, some land has undergone succession from grassland
to scrub-shrub habitat. We evaluated vegetation and habitat structure in mowed and
unmowed hayfields and idle pastures during 1999 and 2000. We observed 71 plant species
on 3 hayfields and 3 pastures. Solidago ulginosa (Bog Goldenrod), Solidago rugosa
(Wrinkle-leaved Goldenrod), Achillea millefolium (Yarrow), Dactylis glomerata (Orchard
Grass), Phalaris arundinacea (Reed Canary Grass), and Hypericum densiflorum
(Glade St. John’s Wort) were the most common taxa. Species composition and abundance
varied by field type and mowing treatment. Vegetation was taller in pastures than in
hayfields; standing dead vegetation was greater in unmowed plots than in mowed plots.
Mowing is useful for maintaining vegetative structure for wildlife and may influence
plant species composition and abundance.
Introduction
The condition of West Virginia’s grasslands is driven by mixes of anthropogenic
changes and natural disturbances. Prior to European settlement, most of
West Virginia was forested, although open-land habitats occupied large patches
(Core 1949). Much of the treeless area was probably bog and other wetland
habitats, but grasslands and grass-bald communities comprised some of these
sites (Core 1949). Grass-bald communities are a unique high-elevation open-land
feature vegetated by grasses and forbs. Although the origin of these communities
is unknown, researchers have suggested that natural and anthropogenic disturbances
created them (Core 1949, Mark 1958, Rentch and Fortney 1997). Since
European settlement, the amount of grassland habitat in WV has increased. Surface
mining has resulted in the development of many grassland habitats over the
last 70 years (Whitmore 1980, Whitmore and Hall 1978). However, grasslands
created by farming—specifically pastures and hayfields—have also increased
dramatically since European settlement (Jones and Vickery 1997).
In the Appalachian region, idle farmlands provide important habitat for grassland-
dependent wildlife species (Bollinger et al. 1990). Idle hayfields and pastures
are increasing in the East due to changes in farming practices and the purchase of
farmland for other uses. These new and changing land practices provide habitats
that benefit a variety of wildlife (Farris and Cole 1981, McCoy et al. 2001). Shrubs
and trees become established on abandoned hayfields and pastures, and if open
1Wildlife and Fisheries Resources Program, Division of Forestry and Natural Resources,
West Virginia University, PO Box 6125, Morgantown, WV 26506. 2Current address - US
Fish and Wildlife Service, Great Bay National Wildlife Refuge, 100 Merrimac Drive,
Newington, NH 03801. *Corresponding author – jim.anderson@mail.wvu.edu.
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sites are not maintained through active management, habitat for grassland wildlife
species will be depleted and may eventually disappear in West Virginia.
Grazing, burning, and mowing or haying are 3 widely accepted techniques for
grassland management. Implementing these treatments is necessary to prevent
woody encroachment into prairie fragments (Burger et al. 1994). When haying is
used as a management technique, it often has the same effect as burning because
cut material is removed. Mowing fields can provide high-quality grassland habitat
by controlling woody vegetation, lowering vegetative height, and reducing
litter build-up, especially if the cuttings are collected and taken away (Sample
and Hoffman 1989). This activity halts succession and provides suitable wildlife
habitat, such as nesting grounds for grassland birds. When mowing is used as a
tool for grassland bird conservation, it is important to delay mowing until the
breeding season has concluded. Mowing has been implicated as a reason for
the declines of grassland nesting species (Bollinger et al. 1990, Frawley and Best
1991) because mowing early in the season disrupts nesting and will ultimately
lead to nest failure. Unfortunately, late season mowing often decreases the value
of hay to landowners and farmers who use the hay. Managers of natural areas
need quantitative estimates of vegetative structure and composition so that management
recommendations to enhance wildlife benefits can be achi eved.
Idle grasslands and hayfields have been documented on the Canaan Valley
National Wildlife Refuge (hereafter called “the Refuge”), but their structure and
importance to wildlife are relatively unknown. Fortney (1975) conducted a study
of the vegetative communities in Canaan Valley (hereafter, the Valley); however,
data on structural components important to wildlife and management effects were
neither sampled nor evaluated. The objective of this study was to estimate the
composition and structure of vegetation on the Refuge’s mowed and unmowed
hayfields and pastures.
Field Sites
This study was conducted on the Refuge in the Valley, Tucker County, WV.
The Valley is about 15 mi (24 km) long and 2–4 mi (3–6 km) wide and is oriented
on a northeast–southwest axis. The Valley’s floor lies at an elevation of 3150–
3248 ft (960–990 m) and is surrounded by mountains up to 1000 ft (305 m) above
the valley floor (USFWS 1979). A large wetland complex occurs on the Valley’s
floor (~6025 acres [2438 ha]), consisting of palustrine emergent, scrub-shrub,
forested, unconsolidated bottom, and unconsolidated shore wetlands (USFWS
1979).
The Valley’s climate is similar to that of northern New York, Vermont, New
Hampshire, and the northern half of Maine, with cold winters and cool summers
(Thornthwaite 1948). Many plants reach the southern limit of their ranges in
the Valley (Fortney 1993). The growing season is relatively short, with an average
of 92 frost-free days from 31 May–1 September (Fortney 1975). Summer
temperatures are moderate, with an average of 75–79 °F (24–26 °C) during the
day and 50–55 °F (10–13 °C) at night. Annual total rainfall is 45 in (114 cm);
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rainfall was 7 in (18 cm) below average in 1999 and at normal level in 2000
(NOAA 1999, 2000).
The Valley was first visited by European explorers in 1746, and people settled
as early as 1800 (Fortney 1993). Prior to logging, a forest of large Picea rubens
(Sarg.) (Red Spruce) with a dense understory of Rhododendron maximum L.
(Rhododendron) covered the Valley (Fortney 1993). Other species present here
and associated with a boreal climate included Abies balsamea L. (Balsam Fir),
Tsuga canadensis L. (Eastern Hemlock), Betula alleghaniensis (F. Michx.) (Yellow
Birch), Acer saccharum (Marsh.) (Sugar Maple), Fagus grandifolia (Ehrh.)
(American Beech), Vaccinium oxycoccos L. (Small Cranberry), Gaultheria
hispidula L. (Creeping Snowberry), Geum strictum (Aiton) (Yellow Avens),
Carex leptonervia (Fernald) (Nerveless Woodland Sedge), Scirpus microcarpus
(J. Presl. & C. Presl) (Panicled Bulrush), Scirpus atrocinctus (Fernald) (Blackgirdle
Bulrush), Caltha palustris L. (Marsh Marigold), and Arisaema triphyllum
(L.) Schott (Jack-in-the-Pulpit) (Strausbaugh and Core 1977). Within the forest,
natural openings or glades occurred, which consisted of grass balds or bogs too
wet for forest development. Deep layers of needles and other plant and animal
matter accumulated and, together with Sphagnum spp. (sphagnum mosses), built
humus-rich acidic soil.
Beginning in the 1880s, the railroad rendered the Valley more accessible and
thereby more susceptible to logging (Vogelmann 1978). The Red Spruce was
cut, leaving the Valley bare. By 1920, few trees remained and lumbering activity
throughout West Virginia declined dramatically (Vogelmann 1978). The microclimate
changed and the soil dried, creating a substrate where tree seedlings were
unable to become established and where fire proliferated (Fortney 1993, Vogelmann
1978). Once-forested bottomlands became sphagnum bogs on wetter sites
and Polytrichum spp. (haircap moss) hummocks on slightly drier sites (Fortney
1993). Drier uplands converted to forb and grass meadows and some heavily
burned areas were planted with mixed grasses (Vogelmann 1978).
Agriculture in the Valley has generally been unsuccessful, although some
farms exist today. Most crops produced low yields in the short growing season,
but cattle production was successful and has been the predominant form of livestock
since the area was logged (Vogelmann 1978). Pastures and hayfields are
found in the slightly drier southern portion of the Valley. Currently, the Refuge
grasslands consist of dry upland areas, Crataegus spp. (hawthorn) savannahs,
saturated wet meadows, and saturated scrub-shrub wetlands, all of which are
included in the broad definition of grassland systems (V ickery et al. 1999).
Methods
Vegetation monitoring
We studied 6 grassland plots, labeled the Beall, Cortland, Freeland, Harper,
Herz, and Thompson tracts, which totaled 730 ac (295 ha) (mean = 121.6 ac/
field, SE = 31.4; [49.2 ha/field, SE = 12.7]). We classified each plot according
to its previous land use as idle hayfield (Beall [230 ac (93 ha)], Harper [180 ac
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(73 ha)], and Thompson [59 ac (24 ha)]) or as idle pasture (Cortland [40 ac (16
ha)], Freeland [69 ac (28 ha)], and Herz [151 ac (61 ha)]). At the conclusion of
the 1999 bird breeding-season, in late August, Refuge staff mowed 50% of each
tract except Herz to determine the effects of habitat manipulation on vegetative
structure and composition. The Herz tract was not mowed because it was too wet
for mowing equipment. We placed transects for vegetation surveys on mowed
and unmowed portions of each field. Transects ran from edge to edge and parallel
to the long axis of each field. Placement depended on plot size, and we placed the
transects 164 ft (50 m) away from the short axis of each field ( Warren 2001).
We sampled the vegetation once per month during June–August of 1999
and 2000 following the methods of Best et al. (1997). We measured the grassland
vegetation’s capacity to obstruct vision (i.e., vertical density [cm]) and
maximum height (cm), the litter depth (cm) using a 3.28-ft (1-m) tall Robel
pole (Robel et al. 1970), and the canopy coverage using a Daubenmire frame
(Daubenmire 1959). We recorded vertical density at 13 ft (4 m) from the Robel
pole. We measured ground cover, classified as canopy, litter, or bare ground,
to the nearest 5% based on Best et al. (1997). We classified canopy coverage
as either living or standing-dead vegetation and we classified vegetative
cover as forb, grass, or woody plant (Best et al. 1997). Thus, the sum of forbs,
grasses, and woody vegetation percentages equaled the percent canopy cover,
as did the sum of the percent living plus percent standing-dead vegetation. Litter
included all dead and decomposing plant material on the soil surface. We
included all dead plant material found above the litter layer in standing-dead
vegetation and considered it in maximum height and vertical density readings.
We recorded measurements every 16.4–65.6 ft (5–20 m) depending on field
size and shape (Warren 2001). In general, we recorded the above measurements
every 16.4 ft (5 m) on small fields (less than 69 ac [28 ha]) and every 65.6 ft (20 m) on
large fields (≥151 ac [61 ha]) (Warren 2001). Regardless of field size or shape,
we recorded species of vegetation touched by a 0.2-in- (0.5-cm-) thick rod
placed every 3.28 ft (1 m) along each transect.
Statistical analyses
We checked the data for normality by using the Shapiro-Wilk statistic and
for homogeneity of variances by plotting residuals (Cody and Smith 1991). To
improve normality and the distribution of variances, we used an arcsine transformation
on canopy, live vegetation, dead vegetation, wood, and vertical density,
and used a log (x+1) transformation on percent bare ground, vertical density, maximum
height, and frequency data (Zar 1999).
We analyzed habitat characteristics, the dependent variables, of plots using
three-way multivariate analysis of variance (MANOVA) or three-way analysis
of variance (ANOVA) between treatments (mowed and unmowed), habitat types
(pastures and hayfields), and months (June, July, and August) (independent variables).
Then, we analyzed the data using three-way ANOVA to compare habitat
characteristics between habitat types (pastures and hayfields), years (1999 and
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2000), and months (June, July, and August). For combined 1999 and 2000 data,
we used all data from 1999, but only data from the unmowed portion of each
field in 2000. We analyzed vertical density, maximum height, and litter depth
(dependent variables) using 3 separate ANOVAs. We used MANOVA to analyze
cover type (canopy, litter, or bare ground), growth state (standing live or standing
dead), and vegetative type (forb, grass, or woody plant); these categories were
transformed because the variables within the groups were correlated (Warren
2001). We compared the frequency of occurrence (number of points occupied/
number of points sampled) between hayfields and pastures for selected species
during July 1999 and 2000. We compared frequency between years and field
types using two-way ANOVA. We considered all tests to be significant at P less than
0.05. Following a significant ANOVA, we used Tukey’s multiple comparison test
to separate means.
Results
We identified 71 plant species, including 67 on hayfields and 58 on pastures
during 1999 and 2000 (Appendix 1). Species with the most coverage in hayfields
and pastures were similar for both years (Appendix 1). The dominant grassland
plants present on the Refuge consisted of Dactylis glomerata (Orchard Grass),
Danthonia compressa (Flattened Oatgrass), Anthoxanthum odoratum. (Sweet
Vernal Grass), Phleum pratense (Timothy), Solidago ulginosa (Bog Goldenrod),
Solidago rugosa (Wrinkle-leaved Goldenrod), Hypericum densiflorum (Bushy
St. Johnswort), and Spiraea alba (Narrow-leaved Meadowsweet). Frequency of
occurrence of plant species was similar between pastures and hayfields based
on July 1999 and 2000 data. The biggest difference was for Reed Canary Grass,
which occurred on 9.7% of the points on hayfields but only on less than 0.0001% of
points in pastures (Table 1).
There were no differences between mowed and unmowed treatments for
percent cover-type (Wilks’ mean = 0.786, P = 0.196), percent vegetative type
(Wilks’ mean = 0.796, P = 0.218), vertical density (F1, 21 = 2.37, P = 0.138), and
maximum height (F1, 21 = 0.20, P = 0.657) (Table 2). There were differences between
treatments for percent growth state (Wilks’ mean = 0.710, P = 0.033), with
standing-dead vegetation taller in unmowed than in mowed plots (F1, 21 = 6.05,
P = 0.023) (Table 2).
There were no differences between habitat types for ground cover (Wilks’
mean = 0.865, P = 0.352), growth state (Wilks’ mean = 0.926, P = 0.411), vegetative
cover (Wilks’ mean = 0.734, P = 0.074), or vertical density (F1, 24 = 0.62, P =
0.440) for combined 1999–2000 data (Table 3). Vegetation was taller in pastures
than in hayfields (F 1, 24 = 6.10, P = 0.021; Table 3). However, litter was deeper
in hayfields than in pastures (F1, 24 = 15.95, P < 0.001; Table 3). There were no
differences between years for growth state (Wilks’ mean = 0.927, P = 0.416) and
litter depth (F1, 24 = 0.27, P = 0.606). Differences were detected between years
for ground cover (Wilks’ mean = 0.609, P = 0.011). Of the percent groundcover
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Table 1. Frequency (%; number of points occupied/number of points observed) of selected abundant
species of vegetation on pastures (3) and hayfields (3) on the Canaan Valley National Wildlife
Refuge, July 1999 and 2000. tr = trace
Pastures Hayfields
Species Mean SE Mean SE F P
Achillea millefolium 2.52 1.65 3.42 0.80 1.92 0.203
Agrostis gigantea 1.11 0.31 0.12 0.12 9.65 0.015
Anthoxanthum odoratum 2.97 1.59 2.84 1.94 0.37 0.561
Asclepias syriaca 0.51 0.51 0.19 0.14 0.12 0.743
Bromus spp. 0.15 0.15 0.19 0.19 0.02 0.897
Carex spp. 2.11 1.03 2.62 0.96 0.18 0.679
Cirsium spp. 0.18 0.18 0.02 0.02 0.68 0.433
Crataegus spp. 0.04 0.04 0.03 0.03 0.04 0.849
Dactylis glomerata 6.76 4.27 4.20 2.69 0.34 0.574
Danthonia compressa 3.67 2.09 2.12 1.38 0.38 0.678
Daucus carota 0.26 0.26 0.17 0.11 0.02 0.903
Dichanthelium clandestinum 1.27 1.27 0.33 0.17 0.10 0.759
Euthamia graminifolia 2.74 2.00 1.88 1.11 0.01 0.924
Schedonorus arundinaceus 1.86 1.26 0.44 0.26 0.94 0.362
Fragaria spp. 2.12 1.78 0.48 0.13 0.58 0.469
Hieracium spp. 0.87 0.62 0.33 0.22 0.59 0.460
Holcus lanatus 2.74 1.55 2.95 0.76 0.43 0.530
Houstonia caerulea tr - 0.08 0.07 1.00 0.347
Hypericum densiflorum 6.34 4.28 3.28 2.08 0.04 0.838
Hypericum ellipticum 0.08 0.08 tr - 1.00 0.347
Juncus effusus 1.01 0.65 1.49 0.70 0.30 0.597
Leucanthemum vulgare 0.45 0.33 1.45 0.53 2.67 0.118
Lolium perenne 0.91 0.54 1.94 0.97 0.67 0.436
Lotus corniculatus 2.01 1.14 6.44 3.95 2.49 0.153
Oxalis stricta 0.32 0.25 0.51 0.26 0.46 0.517
Packera aurea 0.17 0.17 0.06 0.04 0.29 0.604
Phalaris arundinacea tr - 9.72 4.39 12.91 0.007
Phleum pratense 9.79 3.92 6.95 3.50 0.05 0.821
Plantago virginica 0.09 0.09 tr - 1.00 0.347
Poa palustris 0.03 0.03 0.21 0.21 0.61 0.457
Potentilla spp. 3.69 1.62 13.34 3.36 6.23 0.037
Prunella vulgaris 0.17 0.13 0.39 0.29 0.45 0.523
Pteridium aquilinum 0.28 0.22 0.08 0.08 0.57 0.473
Ranunculus spp. 0.34 0.19 0.57 0.25 0.58 0.456
Rumex acetosella 0.23 0.12 0.09 0.07 0.80 0.399
Salix sericea tr - 0.42 0.34 1.60 0.242
Scirpus atrocinctus 0.14 0.13 tr - 1.34 0.280
Sisyrinchium spp. 0.67 0.35 0.18 0.17 1.22 0.302
Solidago rugosa 9.78 3.19 6.32 1.45 0.35 0.573
Solidago ulginosa 6.94 3.68 4.45 2.83 0.19 0.673
Spiraea alba 3.76 3.07 1.47 0.84 0.04 0.840
Stellaria graminea 0.09 0.09 0.32 0.28 0.52 0.493
Taraxacum officinale 1.62 0.70 0.19 0.10 3.71 0.090
Trifolium aureum 0.22 0.14 0.46 0.21 0.85 0.383
Trifolium pratense 0.01 0.01 0.92 0.64 2.40 0.160
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variables, bare ground did not differ between years (F1, 24 = 1.26, P = 0.273), but
the percentages of canopy cover and litter were different. Percent canopy-cover
was higher in 2000 than in 1999 (F1, 24 = 8.39, P = 0.008; Fig. 1), but percent litter
was higher in 1999 than in 2000 (F1, 24 = 12.33, P = 0.002; Fig. 1). Vertical density
(F1, 24 = 17.71, P < 0.001) and maximum height (F1, 24 = 24.95, P < 0.001) were
higher in 2000 than in 1999 (Fig. 2).
Table 2. Vegetative characteristics for mowed and unmowed treatments on the Canaan Valley National
Wildlife Refuge, Tucker County, WV, June–August 2000.
Mowed Unmowed
Variable Mean SE Mean SE
Groundcover (%)
Canopy 79.20 3.31 73.21 3.27
Litter 17.81 3.14 20.19 3.30
Bare ground 3.10 1.39 6.68 2.37
Growth state (%)
Live 77.28 3.16 66.01 4.83
Dead 1.20 0.38 3.09 0.61
Vegetative cover (%)
Forbs 37.92 2.02 35.94 2.60
Grasses 39.84 2.20 33.07 3.48
Wood 0.73 0.40 4.20 1.52
Vertical density (cm) 18.43 2.65 22.65 2.76
Maximum height (cm) 50.57 4.98 51.80 4.56
Litter depth (cm) 1.73 0.16 2.67 0.20
Table 3. Vegetative characteristics for hayfields and pastures on the Canaan Valley National Wildlife
Refuge, Tucker County, WV, June–August, 1999 and 2000.
Hayfields Pastures
Variable Mean SE Mean SE
Groundcover (%)
Canopy 75.35 2.84 76.63 4.02
Litter 20.38 2.83 17.58 3.73
Bare ground 4.30 1.33 5.96 2.81
Growth state (%)
Live 71.97 2.49 70.13 6.31
Dead 2.83 0.56 1.52 0.56
Vegetative cover (%)
Forbs 34.21 1.94 39.40 4.30
Grasses 38.55 1.80 33.26 4.30
Woody plants 1.99 0.82 3.37 1.71
Vertical density (cm) 18.14 2.20 23.83 3.24
Maximum height (cm) 47.45 3.83 55.79 5.57
Litter depth (cm) 2.53 0.23 1.90 0.16
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Discussion
When the Refuge’s grasslands were subjected to mowing, we found that
vegetative structure was similar for most variables in mowed and unmowed
fields at one year after treatment. McCoy et al. (2001) found that most mowed
fields in northeastern Missouri showed no difference in vegetative structure
between the years before and after mowing. The higher percentage of standingdead
vegetation in the unmowed plots, specifically in pastures, provided cover
for grassland birds at the start of their breeding season when maximum height
and vertical density were lower. However, a decrease in the percentage of
standing-dead vegetation on mowed plots appeared to depress bird territory establishment
on the mowed sites during the summer of 2000 (Warren 2001). On
the Refuge, standing-dead vegetation found on unmowed treatments contributed
to increased vertical density early in the bird-breeding season. This taller
vegetation may have served as a cue that attracted some birds as they chose
their nest sites and territories.
Prior to acquisition by the Refuge in 1994, the pastures and hayfields were
actively used for agriculture, thus maintaining much of both field types in
similar successional stages during our study. In general, at our sites on the
Refuge, plant species frequency was similar between hayfields and pastures,
which reduced the distinction between the two habitat types. However, Reed
Figure 1. Comparison of percent canopy and percent litter between years on the Canaan
Valley National Wildlife Refuge, Tucker County WV, 1999–2000.
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Canary Grass, an exotic invasive species (Kecher and Zadler 2004), was found
at a greater rate on hayfields than on pastures. Both hayfields and pastures
had exotic and native grasses that may have simulated the now-rare North
American natural tallgrass prairie (Steinauer and Collins 1996). Madden et al.
(2000) found that Dolichonyx oryzivorus L. (Bobolink) mainly used hayfields
or pastures with exotic, tall grasses. These tall, rhizomatous, graminoids are
structurally similar to the native grass species they replaced. Associated bird
species have adopted this introduced vegetation as breeding habitat (Madden et
al. 2000). McCoy et al. (2001) stated that succession on idle grass fields often
results in monotypic stands of dense vegetation with limited habitat value for
wildlife species. Similarly, on the Refuge, grassland songbirds used Reed Canary
Grass when other plant species were present, but they did not use dense,
monotypic stands of Reed Canary Grass for territory establishment or nest location
(Warren 2001). Grassland birds treated idle hayfields and idle pastures
equally with regard to territory establishment and bird densities, although they
showed a preference for unmowed sites when selecting territories in 2000 (Warren
2001). The quality of vegetation in actively managed grasslands may be
more desirable than in unmanaged sites, but wildlife used idle grass habitats for
foraging, territory establishment, breeding, protection from weather and predators,
and bedding (Warren 2001).
Figure 2. Comparison of vertical density (cm) and maximum height (cm) between years
on the Canaan Valley National Wildlife Refuge, Tucker County, WV, 1999–2000.
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While the Cortland and Herz pastures offered abundant shrub cover not
typical of grasslands, in general, the hayfields and pastures that we studied had
similar species composition and structure. Furthermore, grassland birds appeared
to respond similarly to both habitat types. According to Knopf et al. (1988), birds
within a vegetative association may respond to differences in plant species and
structural variables when selecting habitats. Vegetation was taller in pastures
than hayfields due to the predominance of shrubby St. Johnswort and Narrowleaved
Meadowsweet in the pastures. Also, litter depth was greater in hayfields
than pastures, possibly because there was more grass and forb biomass in hayfields
that contributed to the litter layer (Warren 2001). However, grassland birds
on the Refuge selected hayfields and pastures regardless of any potential finescale
differences in plant species and structure. Some bird species, like Sturnella
magna L. (Eastern Meadowlark), avoided the shrub-dominated parts of pastures
and were found on grass-dominated areas of hayfields. Pastures had more shrubs,
which increased the amount of edge area and vertical diversity, and provided additional
habitat for wildlife species.
An increase in canopy cover, vertical density, and maximum height from 1999
to 2000 can largely be attributed to differences in weather conditions between
the two years. West Virginia experienced a drought in 1999 and normal rainfall
in 2000 (NOAA 2000). In the Valley, May–August total precipitation was 8.6 in
(21.9 cm) greater in 2000 than in 1999. This difference in precipitation may have
influenced grassland plant species growth and wildlife distributions in the Valley
and surrounding areas in both years (O’Connor et al. 1999). In 1999, grassland
plants produced seed and died back early in response to the drought. Additionally,
these plants did not achieve their maximum height or form a complete canopy
because of their physiological responses to the drought (Holtzer et al. 1988). In
2000, grassland plants showed a positive growth response to the increased rainfall
and thereby provided better cover for a longer time. However, even with a
drought effect, plant species composition appeared to be similar between 1999
and 2000 on unmowed fields.
Management implications
It is important to actively manage the grasslands to provide high-quality grassland
habitat for the Refuge’s wildlife. To benefit grassland species, a combination
of mowing, grazing, and prescribed burning should be implemented to reset succession
and maintain an open setting. These management techniques will prevent
woody encroachment into grassland habitat fragments (Burger et al. 1994). Herkert
et al. (1996) found that providing a mosaic of mowed and unmowed, grazed
and ungrazed, and burned and unburned areas provides a full range of habitats
for wildlife species. Such habitat diversity complements the different responses
of grassland plant and wildlife species to management techniques. Mowing may
be the most feasible option because it eliminates the need for coordinating with
livestock farmers. It also avoids having to deal with unpredictable weather for
prescribed burning. Sample and Hoffman (1989) found that mowing can be used
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to control woody vegetation, reduce vegetative height, and reduce litter build-up
if the cuttings are harvested. Mowing should be conducted on a rotational basis,
leaving fields or portions of fields idle for one to two growing seasons. On the
Refuge, mowing should be conducted in mid-to-late August to avoid most of the
grassland birds’ nesting seasons.
Further research should be conducted on the Valley’s grasslands to determine
the effect of surrounding active farmland management on overall vegetative
structure and composition, as well as to investigate wildlife species’ responses to
different management regimes. Additionally, the vegetative structure and composition
of these grasslands should be monitored to determine the long-term effects
of mowing.
Acknowledgments
We thank the US Fish and Wildlife Service (Canaan Valley National Wildlife Refuge);
West Virginia Division of Natural Resources; West Virginia University’s Division of Forestry
and Natural Resources; the Davis College of Agriculture, Natural Resources, and
Design at West Virginia University (Brown Faculty Development Fund); and the West
Virginia Agricultural and Forestry Experiment Station (McIntire-Stennis) for funding.
We are indebted to the late W.N. Grafton for field help in identifying plants. We also thank
C.A. Rhoads and S.K. Reilly for assisting with data collection. R.C. Whitmore and L.
Butler reviewed an earlier version of this paper. This is manuscript number 3201 of the
West Virginia University Agricultural and Forestry Experiment Station.
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Appendix 1. Percent cover of vegetative species found on Canaan Valley National Wildlife Refuge’s
idle hayfields and pastures, Tucker County, WV (1999–2000). H = hayfields and P = pastures.
1999 2000
Scientific name Species H P H P
Family Apiaceae
Daucus carota L. Queen Anne’s Lace 0.34 0.00 0.54 0.13
Zizia aptera (A. Gray) Golden Alexander 0.04 0.05 0.00 0.00
Family Asclepidaceae
Asclepias syriaca L. Common Milkweed 0.05 0.00 0.03 0.18
Family Asteraceae
Achillea millefolium L. Yarrow 6.46 3.18 4.27 0.48
Anaphalis margaritacea L. Pearly Everlasting 0.00 0.00 0.04 0.00
Aster spp. Aster 0.12 0.30 0.04 0.02
Cirsium spp. Thistle 0.19 0.29 0.21 0.22
Erigeron pulchellus (Michx.) Daisy Fleabane Trace 0.00 0.12 0.00
Euthamia graminifolia (L.) Nutt. Goldenrod, Grass-leaved 1.12 2.03 0.90 0.57
Hieracium spp. Hawkweed 0.15 0.00 1.00 1.08
Leucanthemum vulgare Lam. Ox-eye Daisy 1.46 0.22 1.69 0.24
Packera aurea L. Golden Ragwort 0.00 0.05 0.14 0.00
Rudbeckia hirta L. Black-eyed Susan 0.01 0.02 0.07 0.08
Solidago bicolor L. Silverrod 0.00 0.01 0.00 0.05
Solidago rugosa (Mill.) Wrinkle-leaved Goldenrod 5.32 6.25 7.04 3.65
Solidago uliginosa (Nutt.) Bog Goldenrod 6.42 13.60 8.37 13.49
Taraxacum officinale (F.H. Wigg.) Dandelion 0.62 1.14 0.68 0.68
Tragopogon pratensis L. Yellow Goat’s Beard 0.04 0.00 0.19 0.00
Family Brassicaceae
Brassica rapa L. Bird’s Rape 0.00 0.00 0.01 0.00
Family Caryophyllaceae
Dianthus armeria L. Deptford Pink Trace 0.00 0.00 0.00
Stellaria graminea L. Lesser Stitchwort 0.02 0.00 0.22 0.03
Stellaria longifolia (Muhl.) Longleaf Stitchwort 0.00 0.00 0.00 0.00
Family Cyperaceae
Carex spp. Sedge 1.07 2.39 0.71 2.26
Scirpus atrocinctus (Fernald) Blackgirdle Bulrush 0.00 0.00 0.01 0.10
Family Dennstaedtiaceae
Pteridium aquilinum L. Bracken Fern 0.02 0.12 0.10 0.10
Family Ericaceae
Vaccinium spp. Blueberry 1.09 1.14 1.68 8.87
Family Fabaceae
Lotus corniculatus L. Bird’s-foot Trefoil 0.00 0.00 0.00 1.24
Medicago sativa L. Alfalfa Trace 0.00 0.00 0.54
Trifolium aureum Pollich Yellow Hop Clover 0.38 0.00 0.36 0.08
Trifolium pratense L. Red Clover 0.68 0.07 1.00 0.19
Trifolium repens L. White Clover 0.00 0.00 0.01 0.00
Family Gentianaceae
Gentiana clausa Closed Gentian 0.00 0.00 0.03 0.00
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1999 2000
Scientific name Species H P H P
Family Hypericaceae
Hypericum densiflorum (Pursh) Bushy St. Johnswort 4.28 10.30 5.25 11.69
Hypericum ellipticum (Hook.) Elliptic-leaved St. Johnswort 0.00 0.07 0.00 0.05
Family Iridaceae
Sisyrinchium spp. L. Blue-eyed Grass 0.01 0.32 0.00 0.11
Family Juncaceae
Juncus effusus L. Common Rush 0.11 0.62 1.55 2.61
Family Labiatae
Mentha spp. L. Mint 0.13 1.06 0.11 0.02
Prunella vulgaris L. Heal-all 0.10 0.07 0.29 5.19
Satureja vulgaris L. Field-basil 0.28 0.04 0.22 5.34
Stachys palustris L. Marsh Woundwort 0.00 0.00 0.00 2.79
Family Lycopodiaceae
Lycopodium digitatum (Fernald) Fan Clubmoss 0.00 0.16 0.01 2.98
Lycopodium spp. Clubmoss 5.48 4.56 1.37 0.00
Family Onagraceae
Oenothera perennis L. Small Sundrops 0.07 0.00 0.04 0.00
Family Oxalidaceae
Oxalis stricta L. European Yellow Wood Sorrel 0.16 0.37 0.24 11.69
Family Plantaginaceae
Plantago virginica L. Buck Plantain, Buck 0.15 0.48 0.12 0.05
Family Poaceae
Agrostis giantea Roth Redtop 0.18 1.24 1.44 1.18
Anthoxanthum odoratum L. Sweet Vernal Grass 0.08 4.31 3.01 6.18
Dactylis glomerata L. Orchard Grass 6.35 1.62 6.81 5.19
Danthonia compressa Austin Flattened Oatgrass 0.44 1.69 6.94 5.34
Dichanthelium clandestinum Deertongue 0.00 0.02 0.25 0.19
(L.) Gould
Holcus lanatus L. Velvetgrass 0.28 1.88 1.50 1.24
Lolium perenne L. Eastern Ryegrass 0.32 0.41 2.09 0.54
Phalaris arundinacea L. Reed Canary Grass 11.93 0.00 13.12 0.00
Phleum pratense L. Timothy 1.99 5.84 3.21 2.98
Poa palustris L. Fowl Bluegrass 2.40 0.02 0.00 0.00
Schedonorus arundinaceus Tall Fescue 0.00 0.00 1.36 2.79
(Schreb.)
Bromus spp. Brome 0.20 0.20 0.00 0.00
Family Polygalaceae
Polygala sanguinea L. Rose Polygala 0.00 0.00 0.14 0.00
Family Polygonaceae
Rumex acetosella L. Field Sorrel, Sheep Sorrel 0.00 0.00 0.17 0.19
Family Ranunculaceae
Ranunculus spp. Buttercup 0.17 0.38 1.41 1.41
Family Rosaceae
Crataegus spp. Hawthorne 0.00 0.00 0.08 0.08
Fragaria spp. Wild strawberry 0.33 2.34 0.61 0.61
Potentilla spp. Cinquefoil 11.12 4.20 9.86 9.86
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Scientific name Species H P H P
Rubus spp. Dewberry 0.55 2.41 1.22 1.22
Rubus spp. Blackberry 0.00 0.00 0.00 0.00
Spiraea alba (Du Roi) Narrow-leaved Meadowsweet 1.90 7.56 2.59 2.59
Family Rubiaceae
Galium mollugo L. White Bedstrawe 3.61 2.02 0.96 0.96
Houstonia caerulea L. Bluet 0.08 0.26 0.06 0.06
Family Salicaceae
Salix sericea (Marshall) Silky Willow 0.43 0.04 1.01 1.01
Family Scrophulariaceae
Linaria vulgaris (Mill.) Yellow Toadflax 0.00 0.00 0.00 0.00
Family Violaceae
Viola sororia var. sororia Willd. Common Blue Violet 0.02 0.00 0.03 0.03