Cathedral State Park is a 133 acre (54 ha) remnant of ancient primary forest in West Virginia's Allegheny Mountain sub-province. Elevations in the Park range from 2410 to 2620 feet (735 to 799 m) asl, but Shaffer Mountain rises to 3236 feet (987 m) within two miles. Drainage is by branches of Rhine Creek, a branch of the Youghiogheny River, a part of the Ohio system. Underlying rock is probably entirely Mississippian Pocono Formation, which appears to be dominantly siliceous conglomeratic sandstone where exposed. Mississippian Greenbrier limestone overlies the Pocono immediately to the north, where it is characterized by open farmland. Devonian Hampshire Formation, which is also composed largely of siliceous rocks, underlies part of the drainage to the south (Cardwell et al, 1968) . Rocks appear to be gently dipping for the most part. However they are not well exposed in most of the Park.
The climate is humid continental and the region falls within the area of the cloudy day maximum of the eastern US (Reifsnyder and Lull, 1960) . Thus, cloud cover may contribute substantially to the microclimate. Normal daily average temperature in the general region for June is near 65 deg F (18.3 deg C) (National Climatic Data Center, 1966), which is similar to that of parts of inland New England. However the relatively high elevation of the Park, and its position in a valley in proximity to higher elevations, make it susceptible to additional cool air accumulation.
The Cathedral Forest was studied in some detail by Bieri and Anliot (1965), who gave most attention to canopy species. They identified several different communities: Hemlock Rhododendron, Hemlock Fern, Hemlock Duff and Maple Cherry Birch. Their determinations of dbh and basal area frequencies of Hemlock demonstrated the abundance of young, small diameter trees and the prominence of veterans in the dbh range of 40 inches (one m) or more. Also of considerable interest was their measurement of forest interior light intensities, which were found to be only 3.6% of those in an adjacent open field.
Because this remnant tract of primary forest is so small and is cut by a major highway (US 50), it has been subject to some damaging intrusions. While some of these are related to the highway and surrounding farming operations, others are also result from well intentioned but misguided management which makes use of limestone gravel on paths and drainways. Also, while the Hemlock Wooly Adelgid has apparently not made an incursion here as yet, it is a constant threat from the east.
It is also of interest to compare the Cathedral Forest with similar but larger tracts on the Allegheny Plateau of Pennsylvania. Perhaps most intensively studied of these was that on East Tionesta Creek (Hough, 1936; Hough and Forbes, 1943) . While these occurrences lie about 150 miles (240 km) north of Cathedral, regional temperatures in the two regions apparently do not differ greatly, since Appalachian isotherms tend to run north south rather than east west. Thus the average annual temperatures in both regions fall in the range of 48 deg F (8.8 deg C) . This similarity also appears to apply to average annual precipitation, which in both regions falls in the range of 45 inches (114 cm) (Hough and Forbes, 1943) . As a result of the elevation and the previously mentioned topographic conditions, Cathedral may actually be a bit cooler than the Pennsylvania occurrences. An additional parallel between Cathedral and the latter is the dominant presence of siliceous rocks underlying both occurrences (see references) . While cursory in nature, inventories reported here do present some material not previously investigated. This includes bryophytes in particular, but also a very limited survey of the fungi.
Inventories of Cathedral Park were conducted on 7-14-96, 9-22-98, 7-10-01 and 7-11-01. The first two of these were brief and in the nature of reconnaissance . During the last two inventories, particular attention was given to various substrates, the association of plants, particularly bryophytes, with these substrates, and the frequencies of bryophyte species in various habitats. Soil temperatures and pH values were also determined at a few stations.
Cathedral State Park 7-14-96
Note large snag in foreground.
Cathedral State Park 7-14-96
Cathedral State Park 7-14-96
Note Hemlock reproduction.
Our first impression of the Cathedral Forest on this day, during the height of summer's activity, was one of extraordinary odor, humidity and coolness, reflecting the tall Canada hemlock (Tsuga canadensis) canopy, the abundance of decaying down wood, fungi, mosses and liverworts. Songs of the Wood Thrush ( Hylocichla mustelina ) were unusually common, as were the sounds and sightings of Scarlet Tanager (Piranga olivacea), Red-eyed Vireo (Vireo olivaceus ), possible Solitary Vireo (V. solitarius ), Rufus-sided Towhee (Pipilo erythrophthalmus) and others, including unidentified warblers. Fish up to six inches (15 cm ) in length were seen in Rhine Creek.
While dominated by stately Hemlock, the canopy also included large White and Northern Red Oaks (Quercus alba and Q. rubra ) and Black Cherry ( Prunus serotina ), as well as smaller Yellow and Black Birches ( Betula allegheniensis and B. lenta ), American Beech (Fagus grandifolia ), Red and Sugar Maples ( Acer rubrum and A. saccharum ), Cucumbertree (Magnolia acuminata), Tuliptree (Liriodendron tulipifera ) and White Ash ( Fraxinus americana ) . A single Yellow Buckeye (Aesculus octandra) was seen. Understory trees noted were Striped Maple (Acer pensylvanicus ) and Mountain Ash ( Sorbus americana) . Other small trees or shrubs included Mountain Holly ( Ilex montana ), abundant Great Rhododendron (Rhododendron maximum ), some Mountain Laurel (Kalmia latifolia ), Witch Hazel (Hamamelis virginiana ), and, in moist areas, Winterberry Holly ( Ilex verticillata), Arrowwood Viburnum (Viburnum dentatum ) and Black Elderberry (Sambucus canadensis) . Quite abundant also, in moist, open areas near the stream, was a large-caned blackberry with a few straight spines (later identified as Rubus pensilvanicus) . Alternate-leaved Dogwood (Cornus alternifolia ) and a little hawthorn ( Crataegus sp) were observed at the edge of the tract, while the acidiphile Groundberry (Rubus hispidus) was common in moist areas. Although reported as present by Bieri and Anliot (1965), Red Elderbery (Sambucus pubens ) was not recorded by us in any of our inventories thus far.
An impression was also gained on this first visit that a considerable part of the upland under Hemlock had ground cover dominated by either Intermediate Shield Fern (Dryopteris intermedia ), White Wood Sorrel (Oxalis montana ) or Canada Mayflower (Maianthemum canadense ), with associated Hay-scented Fern (Dennstaedtia punctilobula), New York Fern (Thelypteris noveboracensis ), Ground Pine (Lycopodium flabelliforme), Indian Cucumberroot (Medeola virginiana ) and, although least conspicuous, very common Partridge Berry (Mitchella repens ) . Also seen was a little Wild Sarsaparilla (Aralia nudicaulis), scattered trillium, likely Trillium undulatum (Painted Trillium), Mountain Oat Grass (Danthonia compressa ) common along trails, Sweet-scented Bedstraw (Galium triflorum ), White Wood Aster (Aster divaricatus ), Indian Turnip (Arisaema triphyllum), an unidentified Sanicle ( Sanicula sp), Honewort (Cryptotaenia canadensis ), Shining Clubmoss (Lycopodium lucidulum) and the forest grass Brachyelytrum erectum. Common Greenbrier ( Smilax rotundifolia) was one of the few vines seen. Although Bieri and Anliot (1965) reported that Alpine Enchanter's Nightshade (Circaea alpina ) was an important constituent of the flora, we have, curiously, no record of this herb in any of our inventories!
Moist areas of the forest along the stream were characterized by abundant Crooked-stem Aster (Aster prenanthoides ), Wood Nettle ( Laportea canadensis ), an unidentified meadowrue ( Thalictrum sp), Interrupted Fern (Osmunda claytoniana ), Skunk Cabbage (Symplocarpus foetidus ), Tall Coneflower (Rudbeckia laciniata), Common Joe-pye Weed ( Eupatorium fistulosum ), Wide-leaved Joe-pye Weed (E. purpureum ), Clearweed (Pilea pumila ), American Water-pennywort (Hydrocotyle americana ), the alien Forget-me-not ( Myosotis scorpioides) in bloom, Northern Bugleweed ( Lycopus uniflorus ), Fowl Mannagrass (Glyceria striata ), the mannagrass G. melicaria, the bulrush Scirpus atrovirens and the sedge Carex folliculata. As we shall see, some of these herbs, such as Honewort and Wood Nettle in particular, may in some places be related to human-related soil enrichment.
An unusual species in a few moist areas was Star Violet ( Dalibarda repens ), also in bloom, and in association with other acidiphiles such as Partridge Berry, White Wood Sorrel and Ground Berry. According to Strausbaugh and Core ( 1977), the equally rare Lesser Rattlesnake Plantain ( Goodyera repens ) also occurs in the Park, although it was not seen by us. Also, Bieri and Anliot (1965) report Speckled Alder (Alnus rugosa ) as present. Additionally, Venable (undated) has Dwarf Cornel (Cornus canadensis ) as part of the flora. While this reporter is unaware of any other record of this plant, its presence would not be in conflict with this habitat.
Among bryophytes tentatively identified on this first visit were a haircap moss (Poltrichum sp) and the liverwort Bazzania trilobata. A fuller treatment is provided later.
Our brief inventory on this day began under threat of showers that are so much a part of this region.
We entered the trail system at the west end of the parking lot with initial observations of fungi that included the deadly Paxillus involutus, Xeromphalina campanella under decaying Hemlock, an unidentified Russula and Marasmiellus opacus. On moving west along the Cathedral Trail, we noted a very large White Oak and a species of the fungus Cordyceps, likely C. militaris, a common predator of larvae and pupae of butterflies and moths (Phillips, 1991) . We were then surprised by the unseasonable call of a Spring Peeper (Pseudacris crucifer ) from a nearby seep and noted unidentified species of the mosses Hypnum and Dicranum on soil.
We now made a right turn, still on the Cathedral Trail, and observed a few one foot (0.3 m) dbh dead, but still standing, Pitch Pine (Pinus rigida ) that probably once occupied an opening in the dominantly Hemlock forest, and in which fast growing hardwoods eventually shaded them out.
Continuing under threatening skies, we observed in succession the fungi Amanita citrina, with characteristic raw potato odor, and Trichaptum biforme;
then numerous seedlings of Hemlock and some of Red Maple as well; followed by the Honey Mushroom (Armillariella mellea ), and numerous small Hemlock saplings.
Now, arriving at a seep area, we noted Arrowwood Viburnum, the rush Juncus effusus, the bulrushScirpus atrovirens, the sedge Carex folliculata and Fowl Manna Grass, most of which had been observed in our inventory of 7-14-96. Here we again saw several standing but dead, Pitch Pine with bark still adhering, and nearby, the Hawthorn Crataegus flabellate. Delicate Fern Moss (Thuiddium delicatulum) appeared to be common on tree bases and unidentified species of the mossesFissidens and Brachythecium were noted as well. These were followed by a large patch of Tree Clubmoss (Lycopodium obscurum ) .
Here we reached a trail junction, with the Cathedral Trail leading to the left and the Partridge Berry Trail to the right. The trail junction was marked by an uprooted tree with a rootwad that was comprised in part of a tan-colored sandy loam with little or no incorporated organic matter. As we continued along the Cathedral Trail, which here paralleled a branch of Rhine Creek, we observed the edible Deer Mushroom ( Pluteus cervinus ) and heard the calls of the American Crow (Corvus brachyrhynchos ) . Then, approaching another stream crossing, we became aware of the heavy undergrowth that bordered the stream here in the more available light, and which consisted largely of Great Rhododendron, but also of abundant large canes of Pennsylvania Blackberry. Also noted was the fungusTrogia crispa. This was soon followed by another stream crossing; then the edible fungus "Angelwings" (Pleurocybella porrigens ), a characteristic species on decaying conifer wood.
After yet another stream crossing, we came to the junction with the Giant Hemlock Trail, which we began to follow. Noted initially along this trail were fungi that included an unidentified Collybia and Mycena galericulata; then a large patch of 10 to 20 foot (3 to 6 m) -tall Hemlock saplings. These were followed by a fungus of the genus Pholiota and tiny seedlings of Hemlock and birch on a down bole that appeared to be covered on top by an unidentified species of the moss Hypnum and, on its side, by the leafy liverwort Bazzania trilobata. Next seen was the fungus Fomintopsis pinicola on dead conifer wood; then a large dead Hemlock with a heavy growth of the edible but virulent parasitic Honey Mushroom - also noted earlier elsewhere- on its roots. Our traverse was terminated prematurely here because of a downpour.
An impression was gained on this visit, after observation of many moss patches, that species of Hypnum were disposed to occupy horizontal surfaces on down boles, rock and soil, while Thuiddium delicatulum occupied tree bases. However, subsequent microscopic studies by R. Hunsucker on species collected during 7-01, showed that a species similar in appearance to Hypnum, namely Brotherella recurvans, appears to be an important species at Cathedral and may have been misidentified by this reporter.
Our traverse of 7-10-01 began early in the morning at the parking lot and proceeded west along the Cathedral Trail, as on our earlier visit. Weather was sunny, warm and humid. First experienced was the song of a Scarlet Tanager. Then we once again tallied canopy trees, the large Hemlock, Northern Red and White Oaks and Black Cherry, with smaller Yellow and Black Birches, Red Maple, Beech and a few Tuliptree, Cucumbertree and Sugar Maple. Unlike an early visit, White Ash-doubtless fortuitously - was not seen and Sugar Maple appeared very uncommon. However, the low soil pH values obtained by us in later inventories, indicate that habitat for White Ash may be very limited in areas inventoried by us and the pH values of the more acid soil enclaves may even lie outside of the stability field of Sugar maple (Mueller,2000) . A thin leaf mat consisted mostly of last season's Red and White Oak leaves and covered what appeared to be a mor type soil.
Numbered samples of bryophytes, soil temperatures and soils were taken at intervals along this and a subsequent traverse, An attempt was made to collect the bryophytes, which were identified by R. Hunsucker under the microscope, from a variety of habitats and substrates as described in what follows. Soil temperatures were taken from a depth of five inches (13 cm) and soil samples from a depth of 2-3 inches (5-8 cm) at selected locations. These depths were chosen as they were deemed both most accessible and significant, as they fall in the high density feeder root zone of plants and in the case of temperature, deep enough to be screened from superficial variations.
In the text liverworts (Hicks, 1992) are distinguished from mosses by the following sign (Li).
The first bryophyte sample (#1) was taken from the base of a large White Oak up to a height of 5 feet (1.5 m) and yielded the following species:
Anomodon attenuatus, Campylium chrysophyllum, Cololejeunea biddlecomiae (Li)
Ctenidium molluscum, Fissidens adianthoides, Lejeunea cavifolia (Li), Mnium cuspidatum, Neckera pennata.
As a consequence of the deep shade, little difference was noted in the bryophyte cover on different sides of the trunk.
A soil temperature reading (T-131) gave T=17.0 deg C. A soil sample (!) was taken from a grayish, organic clay with some organic matter beneath a two-inch (5 cm) mor layer. An initial pH value for this sample was 4.0, but 24 hours later this had decreased to 3.7, indicating very acid material indeed.
The ground flora was sparse beneath the large White Oak, but included Canada Mayflower, Indian Cucumber-root, Sweet-scented Bedstraw, a little jewelweed (Impatiens sp) and, of great interest, several plants of the Long-bracted Green Orchis (Habenaria viridis), a northern species which was not yet in bloom, but bore flower buds.
The second bryophyte sample (#2) was taken from down wood on the forest floor. Found were:
Brachythecium sp, Chiloscyphus profundus (Li), Hypnum curvifolium, Mnium cuspidatum, Mnium punctatum, Nowellia curvifolia (Li), Odontoschisma denudatum (Li), Pellia epiphylla (Li), Pellia sp ( sterile) (Li), Rhynchostegium serrulatum, Tetraphis pellucida, Thuidium delicatulum.
Our third bryophyte sample (#3) was taken from a mature Beech up to a height of 5 feet. It bore the following:
Brotherella recurvans, Brotherella tenuirostris, Campylium hispidulum, Chiloscyphus profundus (Li), Entodon brevisetus, Frullania asagrayana (Li), Plagiothecium laetum, Trichocolea tomentella (Li).
A little beyond this Beech we saw numerous saplings of Hemlock, White Baneberry (Actaea pachypoda) and White Snakeroot (Eupatorium rugosum) We were next attracted to a 30 inch (0.76 m) dbh Hemlock, which apparently bore but few bryophytes. In our sample (#4), taken from this tree, were:
Bazzania trilobata (Li), Hypnum curvifolium, Plagiothecium laetum, Pottiaceae (sterile), Tetraphis pellucida.
Seen in the vicinity was blooming Indian Pipe (Monotropa uniflora).
Our fifth bryophyte sample (#5) was from a 32 inch (0.81 m) dbh dead, but standing, White Oak, which yielded :
Anomodon attenuatus, Anomodon rostratus, Campylium chrysophyllum, Cololejeunea biddlecomiae (Li), Fissidens adianthoides, Fissidens osmundioides, Lejunea sp (cavifolia?) (Li), Mnium cuspidatum, Radula complanata (Li), Rhodobryum roseum.
A single plant of Wild Sarsaparilla was observed nearby.
Our next sample (#6) was from a down bole, which also bore abundant tiny Hemlock seedlings. Bryophytes recorded were:
Brotherella recurvans, Chiloscyphus profundus (Li), Dicranum fulvum, Dicranum viride, Mnium hornum, Nowellia curvifolia (Li), Polytrichum ohioense, Scapaia nemorosa (Li), Tetraphis pellucida, Trichocolea tomentella (Li).
An unidentified Atrichum moss was present on the soil here, but was not observed in the above sample. Also in the vicinity was an interesting white fungus related to slime molds (Ceratiomyxa sp) .
We now encountered a very large Northern Red Oak immediately adjacent to the trail, and bearing a large bracket fungus high up. The following bryophyte sample (#7) was taken from this oak:
Anomodon attenuatus, Brachythecium acuminatum, Brachythecium curtum, Brotherella tenuirostris, Eurhynchium hians, Rhynchostegium serrulatum, Thuidium delicatulum.
Now, at the trail's edge, we again came upon an occurrence of blooming Star Violet, and nearby, observed a good exposure of conglomeratic sandstone.
Continuing west along the trail, we saw the following fungi: the edible Amanita vaginata, the uncertainly edible Amanita flavoconia, the barely edible Lactarius lignyotis; also Xeromphalina campanella, the edible Tricholomopsis platyphylla ("Platterful"), the Deer Mushroom (Pluteus cervinus), also edible, and tiny Rickenella fibula, as usual, growing in moss. We were then greeted by the loud calls of a Pileated Woodpecker (Dryocopus pileatus) .
We next arrived at a very small, low terrace or flood plain segment. Plants observed on the segment were American Water-pennywort, Wide-leaved Joe-pye Weed, Intermediate Shield Fern, Skunk Cabbage, Wrinkle-leaf Goldenrod (Solidago rugosa), Hooked Crowfoot (Ranunculus recurvatus), Northern Swamp Buttercup ( Ranunculus septentrionalis), Tall Coneflower, a large unidentified Iris (Iris sp), Black Elderberry, Fowl Manna Grass, European Self-heal (Prunella vulgaris var vulgaris), a jewelweed, White Grass (Leersia virginica), Upright Yellow Wood Sorrel (Oxalis stricta), Deer-tongue Grass (Panicum clandestinum), the aliens Orchard Grass (Dactylis glomerata) and Garlic Mustard (Alliaria officinalis), a Northern Red Oak seedling, Virginia Knotweed (Polygonum virginianum), Rough Avens (Geum laciniatum) . Downy Many Knees (Polygonatum pubescens) and the alien forget-me-not. An unidentified thrush gave us a quick glimpse here as well.
A soil temperature determination (T-132) on the this segment of flood plain yielded T=16.0 deg C, while a soil sample (2) had an initial pH of 4.5 and the same value 24 hours later. Although sticky when wet, this soil contained some silt in addition to clay, as lumps formed on drying, could, with moderate effort, be broken with the fingers.
Moving through upland terrain again, we soon came to a patch of Stiff Clubmoss (Lycopodium annotinum), a species with a more northern distribution than other Central Appalachian clubmosses. It was followed in succession by the fungus Amanita muscaria, blooming Groundberry, Common Greenbrier, Shining Clubmoss, Pennsylvania Blackberry and a down bole with its barkless surface almost entirely covered by the liverwort Nowellia curvifolia.
We then arrived at the junction with the Giant Hemlock Trail, where our (#8) bryophyte sample was taken from a large Hemlock, resulting only in the followihg:
Cephalozia catenulate (Li), Nowellia curvifolia (Li), Tetraphis pellucida.
Our next sample (#9), from a large down bole of a mostly barkless Red Maple, was far richer:
Anomodon attenuatus, Bazzania trilobata (Li), Brotherella recurvans, Bryum sp, Dicranella heteromalla, Entodon macropodus, Isopterygiopsis muelleriana, Lejeunea cavifolia (Li), Mnium affine var ciliare, Nowellia curvifolia (Li), Platygyrium repens, Polytrichum ohioense, Thuidium delicatulum, Trichocolea tomentella (Li).
As we continued up the Giant Hemlock Trail, we saw our first Mountain Maple of the traverse and, nearby, a large bush of Wild Raisin (Viburnum cassinoides) . We were then attracted to the characteristic Hemlock bracket fungus Ganoderma tsugae at eye level on a large dying, but standing, Hemlock. Visible on this fungus were several 1.5 cm-long, boldly-patterned, red and black Pleasing Fungus Beetles (Megalodacne heros).
Still moving along this Trail, we came to an area in which the ground cover was dominated by Hay-scented and Intermediate Shield Ferns. Presumably this area was part of Bieri and Anliot's (1965) Hemlock-Fern Community.
A soil temperature determination (T-133) here gave a result of T-16.0 deg C, while a soil sample (3) had pH=3.7 both initially and 24 hours later.
Farther along the Trail we recorded in succession a Cucumbertree, a fallen Yellow Birch that bore the northern polypore Piptoporus betulinus (Phillips. 1991), then the alien Hemp Nettle (Galeopsis tetrahit), a Tuliptree sapling and, at the base of a living Black Cherry, the interesting and edible Berkeley's Polypore (Bondarzewia berkeleii), likely a parasite on this tree.
Not much farther along the Trail we came to a large patch of ground cover comprised in large part of Tree Clubmoss and Ground Pine, and beyond this, a small seep with Arrowwood Viburnum and Winterberry Holly and associated Cinnamon Fern (Osmunda cinnamomea), the large sedge Carex folliculata, the common manna grass Glyceria melicaria, the northern manna grass Glyceria laxa, ground Berry and Skunk Cabbage. On the adjacent upland we saw Painted Trillium, the grass Brachyelytrum erectum, a down bole apparently covered by a Hypnum moss, a species of the fungus Ramaria on dead conifer wood, and a conspicuous occurrence of unidentified species of Atrichum and Polytrichum mosses on the soil.
During the morning of 7-11-01 another traverse was made along the Cathedral Trail. Post-cold front conditions prevailed after a rainy night. As our traverse began, successively noted species were the fungus Tylopolis felleus ( Bitter Bolete), Round-leaf Violet (Viola rotundifolia), a large patch of Medeola under Northern Red Oak and the possibly poisonous fungus Amanita pantherina. Many unidentified bird voices in the canopy as well as exposures of non-conglomeratic sandstone were also noted.
Our next bryophyte sample (#10) was now taken from a one foot (0.3 m) dbh Yellow Birch, with the following result:
Bazzania trilobata (Li), Brotherella recurvans, Dicranum montanum, Hypnum pallescens, Plagiothecium laetum, Tetraphis pellucida.
A bryophyte sample (#11) from a 3 foot (0.9 m) dbh White Oak exhibited the following species:
Anomodon rostratus, Campylium chrysophyllum, Fissidens poypodioides, Mnium cuspidatum, Thuidium delicatulum.
We now arrived again at the location of previously cited Long-bracted Green Orchis. Moving west from there along the trail, we noted in succession the fungus Xeromphalina kauffmanii on hardwood, an unidentified vereo-like bird call, the abundance of Red Maple seedlings on the ground and those of Hemlock both on the ground and among mosses on the larger down boles.
The next bryophyte sample (#12) was taken from a decayed stump, which, to the unaided eye, appeared covered by some species of Atrichum and Polytrichum. While the latter was apparently fortuitously excluded in our sample, the following were found under the microscope:
Atrichum undulatum, Brotherella recurvans, Dicranella heteromalla, mats of Dicranella heteromalla protonema
Following this a sample (#13) was taken from native sandstone, which had:
Brotherella recurvans, Dicranum viride, Mnium punctatum, Sematphyllum demissum, Tetraphis pellucida.
Approaching the stream area once more, we noted Great Rhododendron in bloom, the fungi Polyporus varius on dead hardwood and an unidentified Russula and a seedling of Mountain Ash.
A bryophyte sample (#14) from rocks in mid-stream bore the following:
Amblystegium riparium, Eurhynchium riparioides, Mnium punctatum, Pollia epiphylla (Li), Racomitrium aciculare.
Moving along the Trail again, we identified in succession the Trail Rush (Juncus tenuis) on the beaten path, the mosses Thuidium delicatulum and Mnium hornum, although tentatively, on soil, saw a two foot (0.6 m) dbh Yellow Birch and on dead Hemlock the fungi Boletus badius (edible), Dadaleopsis confragosa and the unusual but edible Pseudohydnum gelatinosum. An impression was also gained here of the widespread, if subordinate, occurrence of the liverwort Bazzania trilobata.
We now came to a large patch of Hay-scented Fern that locally dominated the ground flora. Our next bryophyte sample (#15) was taken, considerably off the Trail, from rock among the Ferns. Only the following species were found:
Bazzania trilobata (Li), Dicranum fulvum, Isopterygium elegans.
Additional species here were the grass Brachyelytrum erectum, Mountain Oat Grass, a seedling of Cucumbertree and another Berkeley's Polypore, this time at the base of a Hemlock, and bearing a large rove beetle of unidentified species.
A soil temperature determination (T-134) was also made in the in the general area and a soil sample (4) taken. A temperature T= 17.0 deg C was obtained. The soil sample had pH=3.7 initially and this increased to 4.0 twenty four hours later. This soil had a one inch (2.5 cm) -thick mor layer over pinkish-gray mineral soil. Hard lumps formed on drying, indicating high clay content. Some species in the immediate sample area were a two foot ( 0.6 m) dbh Northern Red Oak, seedlings of Red Maple, Pennsylvania Blackberry and Intermediate Shield Fern. A large fritillary butterfly was also seen here.
Still off the Trail, we came upon a patch of seedlings of the Asiatic water Pepper (Polygonum cespitosum) which may have had its origin in Black Bear (Ursus americanus) or other animal scat. Independent patches of White Wood Sorrel were present in the area as well.
Our last bryophyte sample (#16) was from a four foot (1.22 m) dbh Hemlock snag. Only the following were found:
Bazzania trilobata (Li), Brotherella recurvans, Nowellia curvifolia (Li), Tetraphis pellucida.
The song of a Wood Thrush was now heard in the distance and that of a Red-eyed Vireo nearby.
Taking another trail, we began our return to our point of origin. Successively tallied along this trail were a fungus of the genus Collybia on hardwood twigs, seedlings of Mountain Holly, the poisonous puffball Scleroderma citrinum; then moving along the stream, the previously unreported shrub Ninebark (Physocarpus opulifolius), Turtlehead (Chelone glabra), White Wood Aster, Sweet White Violet (Viola blanda) and the edible fungus Amanita fulva. We terminated our traverse here.
A pH determination was also made of water from the branch of Rhine Creek that flows past the parking lot. This water was very acidic, with pH= 4.5, indicating that acidic substrate also prevails up-stream from the main tract of the Cathedral forest. It seems likely, however, that this acidity is moderated where the stream enters the watershed on Greenbrier Limestone to the north. Such moderation may also explain the fish observed on our 7-14-96 visit.
The following is a table of bryophyte frequencies in the sixteen numbered habitats (see text) from Cathedral State Park.
|7||Brotherella recurvans||7||Tetraphis pellucida||5||Bazzania trilobata (Li)||5||Nowellia curvifolia (Li)||4||Anomodon attenuatus||4||Mnium cuspidatum||4||Thuidium delicatulum||3||Campylium chrysophyllum||3||Chiloscyphus profundus (Li)||3||Mnium punctatum||3||Trichocolea tomentella (Li)||2||Anomodon rostratus||2||Brotherella tenuirostris||2||Cololejeunea biddlecomiae (Li)||2||Dicranella heteromalla||2||Dicranum fulvum||2||Dicranum viride||2||Fissidens adianthoides||2||Hypnum curvifolium||2||Lejeunea cavifolia (Li)||2||Pellia epiphylla (Li)||2||Plagiothecium laetum||2||Polytrichum ohioense||1||Amblystegium riparium||1||Atrichum undulatum||1||Brachythecium acuminatum||1||Brachythecium curtum||1||Brachythecium sp||1||Bryum sp||1||Campylium hispidulum||1||Cephalozia catenata (Li)||1||Ctenidium molluscum||1||Dicranum montanum||1||Entodon brevisetus||1||Entodon macropodus||1||Eurhynchium hians||1||Eurhynchium riparioides||1||Fissidens osmundioides||1||Fissidens polypodioides||1||Frullania asagrayana (Li)||1||Hypnum pallescens||1||Isopterygiopsis muelleriana||1||Isopterygium elegans||1||Lejeunea sp (Li)||1||Mnium affine var ciliare||1||Mnium hornum||1||Neckera pennata||1||Odontoschisma denudatum (Li)||1||Pellia sp(Li)||1||Platygyrium repens||1||Pottiaceae||1||Racomitrium aciculare||1||Radula complanata (Li)||1||Rhodobryum roseum||1||Scapania nemorosa (Li)||1||Sematophyllum demissum|
Central here, as in most of our studies, is the question of species and community stability. Cathedral State Park is old growth primary forest and such forests are sometimes regarded, or at least treated, as a forest type, as distinguished from secondary forest. However, as emphasized by Braun (1950) and others, forest types are products of climate, topography and soils, or ultimately of the interaction of climate and the geologic base.
The Cathedral forest is characterized by a large number of northern / montane and moisture-loving vascular plants. Examples, not all of which were seen by us, are Yellow Birch, Mountain Ash, Mountain Maple, Red Elderberry, Yew, Hobblebush, Speckled Alder, Dwarf Cornell, Canada Mayflower, Yellow Clintonia, White Wood Sorrel, Star Violet, Alpine enchanter's Nightshade, Painted Trillium, Large-bracted Green Orchis, Glyceria Laxa and Stiff Clubmoss. It is thus of great interest to compare the non-vascular flora, in particular bryophytes, to see if it too fits this pattern. In the following discussion an examination is made of the known environmental and substrate preferences of the observed bryophytes. Particular attention is given the mosses, drawing upon the classic work of Crum and Anderson (1981). Where cited in this section, this work is indicated by the symbol (C&A). Reference is also made to a high elevation oak forest near Reddish Knob, to oak forests of the North River Valley and to a mesic deciduous forest at Tea Creek on the Allegheny Plateau. Installation of our inventories in these forests is pending.
The four most frequently occurring bryophytes are all characteristic of acidic, cool northern or montane habitats.
Brotherella recurvans is found in "generally coniferous woods at higher elevations", :"A northern and montane species" (C&A). According to Cogbill (1996) this is one of the characteristic mosses of spruce-fir forests throughout the northeast.
Tetraphis pellucida is "An acidiphile", " in moist coniferous forests".(C&A) An unusual feature at Cathedral is this species common occurrence on tree trunks, especially Hemlock. It was not found at all by us in deciduous forest types at Reddish Knob, the North River Valley or at Tea Creek, although it is part of the Hemlock forest at Ramsey's Draft as well as some other Central Appalachian forests with boreal components.
Bazzania trilobata and Nowellia curvifolia are liverworts characteristic of cool, moist forests, with the former particularly common in spruce-fir forests (Cogbill, 1996) . It is frequently the most abundant bryophyte in Central Appalachian Red Spruce forests (Carvell, 1993) .
Anomodon attenuatus, Mnium cuspidatum and Thuidium delicatulum are widely distributed species, particularly in the northern and eastern US.
Chiloscyphus profundus and Lejeunea cavifolia were not found by Breil (1996) on the Virginia piedmont, a likely indication of northern / montane preference.
Mnium punctatum is usually found "in swampy, eutrophic coniferous woods" (C&A)
Campylium chrysophyllum, Fissidens polypodioides and Ctenidium molluscum are seemingly anomalous in that they appear to be markedly calciphile (C&A). However, in our sampling at Cathedral they were all found only on White Oak and not on clearly acidic substrate.
Many of the minor species are well-known acidiphiles. Examples are Brotherella tenuirostris, Dicranum fulvum, Isopterygium elegans, Racomitrium aciculare and Sematophyllum demissum (C&A).
A possible mechanism at work here is the transfer by capillary action or otherwise of hydrogen ions from the acid soils to overlying moist down wood and tree bases.
Brachythecium curtum is found "Newfoundland to Minnesota, south in the higher mountains to North Carolina"
It is also interesting to compare the two Hypnum species. The most common species, Hypnum curvifolium is characteristic of "moist shady places", while Hypnum pallescens is found " in dry or moist woods" (C&A). The latter was observed by us to be the most frequent species in high elevation oak forest presumably dryer than Cathedral - near Reddish Knob.
Another minor species, Platygyrium repens, had a high frequency in all oak forests studied by us, and in at least one case, was the most frequent bryophyte.
There is also a small element of southerners at Cathedral. The best example is Entoodon macropodus- an exception that proves the rule-" Widespread in the southeastern United States (from Virginia to Florida and west to Arkansas and Texas) " (C&A).
Cooper-Ellis (1998) undertook a comparative study of bryophytes in old growth and secondary forests of western Massachusetts. Her paper also includes an extensive bibliography of previous work. Six different forest types were included. However, these were combined in the two groups of old growth and secondary forest. Thus a comparison with our study program is difficult. She found that the two dominant species at Cathedral, Brotherella recurvans and Tetraphis pellucida, showed no particular pattern of distribution between trees, down boles, rocks and soil, while Hypnum pallescens, a minor species at Cathedral, was among the most common in both old growth and secondary forests. The lack of pattern of B. recurvans and T. pellucida may be a consequence of her combining Tsuga canadensis Betula allegheniensis forest types with types dominated entirely by hardwoods.
The evidence of good reproduction of Hemlock found at Cathedral State Park by Bieri and Anliot (1965) is perhaps not surprising for the time of their study. However its continuance to the present time, as observed by us, is unusual, given the current widespread destructive Whitetail Deer (Odocoileus virginianus) browsing of this tree throughout the Central Appalachians. The effects of this Deer browsing on the forests of Pennsylvania's Allegheny Plateau has been the subject of extensive investigations by Bjorkbom and Larson (1977) and by Whitney (1984), and they report large effects indeed. Another evergreen that has fared less well at Cathedral, and in the Central Appalachians in general, is the Yew (Taxus canadensis). Although Bieri and Anliot reported its presence, not a vestige was seen by us, and it seems likely that it has been virtually eliminated by the recently greatly increased populations of Deer.
While there is ample evidence of disturbance at Cathedral in the form of remains of openings invaded by Pitch Pine and hardwoods, this disturbance appears to have been on a small scale and confined to normal gap formation. Runkle (1991) places the area in a region dominated by small gap disturbance as distinguished from Pennsylvania's Allegheny Plateau, where he finds larger scale blow-downs and fires to be of equal or greater importance. Evidence of such large scale disturbances in the latter region were presented by Hough (1936) and Hough and Forbes (1943). A point made by these authors is that fire was less prevalent in Hemlock than in other less moist forests and in forests on exposed topography. However, if preceded by a blow-down, any Hemlock forest became vulnerable to fire. In any case, little or no evidence of fire was observed by us at Cathedral.
Our experience in the Cathedral forest supports observations of the clear relationship between individual species, forest type, topography and geologic substrate. First, the great majority of observed species of vascular plants are either of wide occurrence or are markedly acidiphile, in agreement with measured soil pH and the observed substrate of conglomeratic sandstone. Secondly, a large number of these species, perhaps 17 or more, are also northern or montane in distribution, in harmony with the regional climate as modified by the local topography. Finally, both the observed species of bryophytes and their frequencies in a variety of microhabitats reflect the same environmental factors as do the vascular plants. Unfortunately, it was not possible in the time available, to investigate possible small scale variations in the geologic substrate and related variations, if any, in the flora, such as were found by us at The Gaudineer Scenic Area and in The Fanny Bennett Hemlock Grove. We did not, for example, have an opportunity to explore the southern extension of the Park. Also, less clear than the readily observed relations described above, may be a general ambiance and feedback that integrates light intensity and quality, moisture, substrate chemistry, micro-topography and biologic elements to produce a community that would not exist but for the presence of all of these elements. More evidence of this interpretation will be provided in future presentations of our inventories in other forest types.
Our attention was drawn however to floral elements apparently out of harmony with the dominant oligotrophic, acidic community, particularly on slopes below trails and route US 50. This takes the form of dense growths of mesic, nutrient demanding native herbs such as Wood Nettle and Honewort as well as introduced species. It seems apparent from these growths that nutrients and alkaline solutions have originated in path and road gravels and washed down-slope. It is also possible that some other species, first recorded by Bieri and Anliot (1965), and which also seem in conflict with the dominant flora, originated in this way, Restoration of the natural community would require cessation of the use of limestone gravels on paths and barriers to prevent road-wash. Ideally, as much as possible of the offending material would also be removed from paths and drainways.
It should also be pointed out that, while soils here are very acidic and appear to have a mor-type structure, they lack the thick, dark, highly organic upper horizon that is characteristic of purely coniferous old growth forests. They share this feature with the acidic soils of The Gaudineer Scenic Area, which also has a prominent deciduous component, in that case with Red Spruce.
The kind procurement by Christina Wulf of copies of a number of documents important to this study is greatly appreciated.
Bieri, Robert and Sture F. Anilot (1965) The Structure and Floristic Composition of a Virgin Hemlock Forest in West Virginia. Castanea 30, 205-226.
Bjorkbom, J. C. and R. G. Larson (1977) The Tionesta Scenic and Research Natural Areas, Forest Service General Technical Report NE-31, Upper Darby, Pa.
Braun, E. Lucy (1950) Deciduous Forests of Eastern North America. Macmillan Publishing Co., New York, N. Y.
Breil, David A. (1996) Liverworts and Hornworts of the Virginia Piedmont. Banisteria, no 8, 3-28.
Cardwell, Dudley H., Robert B. Erwin, Herbert P. Woodward and Charles W. Lotz, compilers (1968) Geologic Map of West Virginia, slightly revised 1986,. West Virginia Geologic and Economic Survey, Morgantown, West Va.
Carvell, Kenneth L. (1993) Bryophytes and Their Ecological Role in the Upland Forests. in Upland Forests of West Virginia, Steven L. Stephenson editor. pp 87 100.McClain Printing Co., Parsons, west Va.
Cogbill, Charles V. (1996) Black Growth and Fiddlebutts: The Nature of Old-Growth Red Spruce in Eastern Old-Growth Forests. Prospects for Rediscovery and Recovery. Mary Byrd Davis, editor. pp 113-125. Island Press, Washington, DC.
Cooper-Ellis, Sarah (1998) Bryophytes in old-growth forests of western Massachusets. Journal of the Torrey Botanical Society. 125 (2), 117-132.
Crum, Howard A. and Lewis E. Anderson (1981) Mosses of Eastern North America, In Two Volumes. Columbia University Press, New York, N. Y.
Hicks, Marie L. (1992) Guide to the Liverworts of North Carolina. Duke University Press, Durham, North Carolina.
Hough, A. F. (1936) A Climax Forest Community on East Tionesta Creek in Northwestern Pennsylvania. Ecology 17 (1), 9-28.
Hough, A. F. and R. D. Forbes (1943) The Ecology and Silvics of Forests in the High Plateaus of Pennsylvania. Ecological Monographs, 13 (3), 299-320.
Mueller, R. F. (2000) Stability relations in Forests. Forests of the Central Appalachians Project. Virginians for Wilderness web Site.
Phillips, Roger (1991) Mushrooms of North America. Little Brown and Co., Boston, Mass.
Reifsnyder, William E. and Howard W. Lull (1965) Radiant Energy in Relation to Forests. Technical Bulletin No. 1344, U.S. Dept. of Agriculture, Forest Service. Washington, DC.
Runkle, James R. (1996) Central Mesophytic Forests. in Eastern Old-Growth Forests Prospects for Rediscovery and Recovery, Mary Byrd Davis, editor. pp 161-177, Island Press, Washington, DC.
Strausbaugh, P. D. and Earl L. Core (1977) Flora of West Virginia, second edition, Seneca Books, Inc., Grantsville, West Va.
Venable, Norma Jean (undated) Cathedral State Park. West Virginia University Cooperative Extension Service, Morgantown, West Virginia.
July 31, 2001
John R. Pope, Jr, Chief
Div. of Natural Resources
Parks and Recreation Section
Capitol Complex, Bldg. 3, Rm. 714
1900 Kanawha Blvd. East
Charleston WV 25305-0662
Dear Mr. Pope,
I have a few things to report regarding our recent inventory of Cathedral State Park. Dr Hunsucker and I appreciate your permission to inventory this important natural area.
First, we do believe that all employees connected to the Park in your Division are doing their utmost to protect and enhance this great public resource. However we believe certain management practices as well as outside influences are seriously jeopardizing the ecological features of the Park that are important to keeping it in a pristine condition, especially with regard to its value for scientific studies. Among the most damaging practices is the use of limestone gravels on the walking paths. These gravels are also washed farther into the Park by the hard rains that are frequent in the area. The original rock that underlies the Park is conglomeratic sandstone, which creates a very acidic substrate for plant growth. Thus acidiphile plants dominate its flora. However limestone is alkaline, acts to neutralize the original acid and contributes additional nutrients, with the result that around, and particularly down-slope from these paths, the original flora is being replaced by lime tolerant and more nutrient demanding plants such as Wood Nettle (Laportea canadensis), Honewort (Cryptotaenia canadensis) and other natives which originally may have occurred in very limited parts of the Park, or by alien species such as Common Burdock (Arctium minus) and Coltsfoot (Tussilago farfara), which should not occur at all.
A similar process is occurring as a result of erosion of and wash from the embankments of Route 50, and this may be even more serious than the limestone paths as chemicals in addition to lime may be involved
The use of limestone for paths or as lining for drainways should immediately be discontinued and the removal of old limestone attempted as much as is practicable. The West Va Department of Transportation should be contacted to see about stabilization of the road embankments. Sandstone or quartzite gravels should be substituted in the Park and in all drainways that lead into the Park. Native stone similar to that that underlies the Park would be preferred. We hope you will consider taking these measures to restore and protect the future integrity of this natural and national treasure.
We hope in the near future to make available to you a more complete report on our inventories this year. We also hope to continue these studies in the future. Once again, thanks for the opportunity to work in this park.
F. Mueller PhD