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General
We here explore the occurrence of Hydrastis canadensis (Goldenseal) on two very different geologic substrates in the Appalachian Mountains. This herb, sought ever more intensively for its medicinal and herbal properties, is under grave threat of extirpation wherever it occurs. For this reason the location of one of these occurrences, first discovered in our Project inventories, and by far the largest, is not revealed.
Central to our study are the broad settings of the occurrences of the species in terms of geology, topography, soil and microclimate. Special attention is given to proximity to various rock types, other plants and to soil character, particularly with respect to pH. Also included is the variation in type of the enveloping forest. This section is also unashamedly opportunistic in its exposition on peripheral but interesting details of the occurrence of other species.
The first occurrence discussed is in calcareous terrain, specifically on dolomitic limestone with substantial inclusions of the siliceous rock chert. This formation, the Ordovician Beekmantown, exhibits a low grade of metamorphism dating to the Appalachian orogeny. Of the second occurrence we need only say that it has developed on soils derived from rock of granitic composition in a more mountainous terrain than the first occurrence. There is a difference between our treatment here of the two occurrences. While inventories of the calcareous area extended over years, those of the granitic area occurred during several days, were of greater intensity and were executed on specific traverses.
As in other sections of this work, both the scientific and common names of species are given on first mention assuming a species has a well-known common name but only the common name thereafter. Exceptions are made for certain genra represented by only one American species with a simple scientific name. Thus the genus name "Hydrastis" rather than "Goldenseal" is frequently used here, as is "Medeola" rather than "Indian Cucumberroot".
Click photos to see larger versions
April, Hydrastis canadensis on calcareous substrate
Late August, Hydrastis canadensis and Goodyera pubescens on granitic substrate
Early September, Hydrastis canadensis and Lycopodium flabelliforme on granitic substrate
Early September, Hydrastis canadensis on granitic substrate
Early September, Hydrastis canadensis on granitic substrate, photo by Mike Jones
Calcareous Substrate
This occurrence ofHydrastis is on soils derived from dolomitic limestone which also has conspicuous chert components (Rader, 1967) . Both limestone and chert are highly fractured, in part as a consequence of proximity to the Pulasky Staunton Fault, a major structural feature of the Great Valley of Virginia. Elevation of the occurrence is a little above 1600 feet (488 m) asl.
Hydrastis occurs in several small patches on the southeast facing slope of a small ridge approximately Ό mile (0.4 km) in length and which rises 120 feet (37 m) above the flood plain of Folly Mills Creek. This stream, a branch of the South Fork of the Shenandoah River, is near the middle of the Shenandoah Valley. An embayment of the flood Plain indents the ridge and is the site of a large artesian spring that feeds a calcareous wetland which is the site of a number of rare disjunct plants (Hunsucker and Mueller, 1998) .Both ridge and wetland fall in a State natural area preserve. Several distinct forest types as well as pasture land reverting to forest occupy the ridge. Forming the ridge's backbone, as exposed on its crest, are massive beds of fractured chert.which also extends down both northwest and southeast slopes as talus and scattered outcrops. This chert dominates the northwest slope, which has only minor outcrops of limestone. In contrast, the southeast slope is dominated by calcareous colluvium with numerous limestone outcrops. However, there are minor outcrops and substantial chert talus on this slope as well. Reverting pastureland on the ridge's southwest end bears few rock outcrops and appears to consist largely of calcareous but leached residual soils. This disposition of rock types has resulted in an ericaceous oak forest on the ridge crest and the northwest slope and a dry mesic calcareous forest on the southeast slope. Where more abundant limestone outcrops occur on the latter, there is a Chinquapin Oak (Quercus muehlenbergii) White Ash (Fraxinus americana) variant of the calcareous forest. The acid environment of most of the chert areas is enhanced by vertical leaching. However, this leaching also occurs in calcareous areas, and is even effective on outcrops of massive limestone. Where colluvial processes dominate, as on steep southeast slopes, deeper soils establish contact with and incorporate calcareous material, which is then brought into the shallow feeder root zone by down slope earth movement. Enhancing this effect is the lateral movement and bathing of the root zone by subsurface solutions that also originate in the calcareous depths. The result is a down slope enrichment of soils on these colluvial slopes. Soils here are of the mull type, brown and rich in clay and with organic matter that is well incorporated into the mineral matter to depths of a meter or more. Where organic matter is most abundant, they are a darker shade of brown.
The closed canopy forest of the ridge is a mature remnant of recent logging (~ 30 years before the present) which took an older but probably still secondary component. Consequently there is an un-equilibrated age distribution of trees, although the mix of trees and the floras in general clearly reflect substrate and aspect with fidelity. Few trees however exceed a foot (0.3 m) dbh or 70 years in age.
The canopy of the ericaceous oak forest of the ridge top and northwest slope is dominated by White and Black Oaks (Quercus alba and Q. velutina), with less common Pignut Hickory (Carya glabra), Red Hickory (C. ovalis), Mockernut Hickory (C. tomentosa), Red Maple (Acer rubrum) and Black Cherry (Prunus serotina) . Minor canopy species are White Pine (Pinus strobus), Virginia Pine (Pinus virginiana), Pitch Pine (Pinus rigida), Scarlet Oak (Quercus coccinea ), Chestnut Oak (Q. prinus) (one), Sassafras (Sassafras albidum), Tuliptree (Liriodendron tulipifera) (one), Canada Hemlock ( Tsuga canadensis) , Black Gum (Nyssa sylvatica) (one) and one American Chestnut (Castanea dentata) sapling. Small and understory trees are Downy Serviceberry (Amelanchier arborea) and Flowering Dogwood (Cornus florida) . Shrubs in the most acidic, rocky areas include Upland Low Blueberry (Vaccinium pallidum), Deer Berry (V. stamineum), Pinxter Azalea (Rhododendron nudiflorum ), Black Huckleberry (Gaylussacia baccata), Corymbed Spiraea (Spiraea corymbosa) and St Andrew's Cross (Hypericum hypericoides) . Some Mountain Laurel (Kalmia latifolia) may also have been present at one time but was likely browsed away by goats. Shrubs that occur throughout are Black Haw Viburnum (Viburnum prunifolium), Maple- leaf Viburnum (V. acerifolium) and Carolina Rose (Rosa Carolina) . Virginia Creeper (Parthenocissus quinquefolia), Poison Ivy (Rhus radicans), Summer Grape (Vitis aestivalis) and Northern Dewberry (Rubus flagellaris) are common vines except in the most acidic areas. A heavy leaf mat is everywhere present in this forest, and soils tend to be of the mor type (1). Here, as in most areas of the mountains, many Flowering Dogwood trees have been killed by Anthracnose (Discula destructiva).
The status of Coral Berry (Symphoricarpos orbiculatus), a somewhat calciphile shrub, in the acidic forest is unclear. While it apparently is excluded from the interior of the most acidic stands here, it occurs on their periphery in soils with pH less than 5.0.
However it appears to be excluded generally in the mountains where acid soils are of wide extent. It is possible that this plant satisfies its stability requirements through extended root contact with distant calcareous elements in soils. Perry (1982) documented and illustrated the enormous reach of tree roots and it has been postulated (Mueller, 2000) that roots of a given plant might well seek out soil volumes compatible with its nutrient requirements.
Herbs most characteristic of the most rocky and acidic areas are Spotted Wintergreen (Chimaphila maculata), Pipsisewa (C. umbellata), Rattlesnake Weed (Hieracium venosum), Downy Rattlesnake Plantain (Goodyera pubescens), Roundlobe Hepatica (Hepatica americana), and Creeping Five-leaf (Potentilla canadensis andP. simplex) . Other species of wider distribution are Plantain-leaf Pussytoes (Antennaria plantaginifolia), the alien Mouse-ear Chickweed (Cerastrum viscosum), Alumroot (Heuchera americana), American Pennyroyal (Hedeoma pulegioides), Common Speedwell (Veronica officinalis), Hairy Bedstraw ( Galium pilosum), Great Chickweed ( Stellaria pubera), Hairy Hawkweed (Hieracium gronovii), White Wood Aster (Aster divaricatus), Wavy-leaf Aster (A. undulatus), Curtis Goldenrod (Solidago curtisii ), Cut-leaf Goldenrod (S. arguta), Silverrod (S. bicolor), Venus Looking Glass (Specularia perfoliata), the tick trefoil Desmodium nudiflorum, Four-leaf Yam (Dioscorea quaternata), Rue Anemone (Anemonella thalictroides), Hairy-jointed Meadow Parsnip (Thaspium barbinode), Yellow Star Grass (Hypoxis hirsuta), Spiked Lobelia (Lobelia spicata var scaposa), Cutleaf Grapefern (Botrychium dissectum), Marginal Shield Fern (Dryopteris marginalis), Slender Gerardia (Gerardia tenuifolia), the sedges Carex pensylvanica, C. wildenowii, C. digitalis, C. rosea and the grasses Poverty Oat Grass (Danthonia spicata), Autumn Bent Grass (Agrostis perennans), Nimblewill (Muhlenbergia schreberi), Nodding Fescue (Festuca obtusa) and the alien Canada Bluegrass (Poa compressa) .Additionally Early Saxifrage (Saxifraga virginiensis), Common Many Knees (Polgonatum biflorum) and unidentified violets (Viola sp) are usually confined to tree bases. It is also interesting that neither Pipsisewa, a northern species, nor Roudlobe Hepatica ever occur on any south or southeast slope, but are confined to the northwest slope, although Spotted Wintergreen is common on acidic areas everywhere. Also, unlike Spotted Wintergreen, Pipsisewa has never been observed to flower here.
As might be expected, wherever limestone outcrops occur in the dominantly ericaceous oak forest, the area in proximity to them bears a richer, more mesic flora characteristic of calcareous, more alkaline soils. In one place, in the midst of an acidic area with numerous Red Maple seedlings, a small limestone outcrop is surrounded by a meter wide zone from which all Red Maple is excluded in a visual display of the stability relations of this species (see Fig. 1, Mueller, 2000) .
Although the relatively dry character of this forest does not favor mosses, certain species are concentrated at tree bases on rocks and small elevations. Most conspicuous is a haircap moss, likely Polytrichum ohioense from its macroscopic features. Other common species on soil are Delicate Fern Moss (Thuidium delicatulum) and false haircap, likely Atrichum angustatum in part, while Hedwigia ciliata is, as is usual, found on siliceous rock, in this case chert. One or more species of Mnium also occur.
In the most acidic parts of the ericaceous oak forest pH values range from about 4.4 to 5.5, and these values are regularly obtained in close proximity to Hepatica, Downy Rattlesnake Plantain, Spotted Wintergreen, Pipsisewa, Rattlesnake Weed, the Blueberries, Black Huckleberry and Azalea. Among the lowest are those obtained from soil immediately beneathPolytrichum mats.
A large open area (pasture) on the ridge crest also has prevailingly acid soils. Common herbs there are Long-leaved Bluets (Houstonia longifolia), Pinweed (Lechea tenuifolia), Creeping Bush Clover (Lespedeza repens), Whorled Milkwort (Polygala verticillata), Bird-foot Violet ( Viola pedata), Ovate-leaved Violet (V. fimbriatula), Blue Curls ( Trichostema dichotomum), Six-week Fescue (Festuca octoflora), Tarweed (Cuphea petiolata), Indian Tobacco (Lobelia inflata) and Poverty Oat Grass. Where small elevations with a substantial chert component in the soil occur, there is a dense intergrowth of mosses and lichens as well as some of the aforementioned herbs. By far the most common moss here is Dicranum scoparium and the most conspicuous lichen a fruticose Cladina, likelyCladina subtenuis. Values of pH in these patches tend to be low, and in soil immediately beneath Dicranum is near 4.5.
At the southwestern edge of the ridge crest forest there is a stand of Virginia Pine that is encroaching on pasture. Although the interior of this stand has virtually no ground flora, the better lit edge has a cover of Poverty Oat Grass and a somewhat uncommon fruticose lichen Cetraria arenaria.
On the ridge, as in many areas inventoried by us, there appears to be a correlation between soil pH and the abundance and diversity of fungi, with acid soils being favored. Consequently mushrooms appear to be most abundant and in greater variety in the ericaceous oak forest as compared to the forest on the calcareous slope.
Although the canopy species of the ericaceous oak forest are also to be found on the southeast slope, additional species occur on the latter, particularly on calcareous colluvium. Especially abundant on the colluvium are White Ash (Fraxinus americana), Bitternut Hickory (Carya cordiformis), Black Walnut (Juglans nigra), Hackberry (Celtis occidentalis) of stunted form, Slippery Elm (Ulmus rubra), Northern Red Oak (Quercus rubrum) and Chinquapin Oak (Q. muehlenbergii). Of these Chinquapin Oak is closely associated with limestone outcrops and where these are particularly abundant, this oak and White Ash dominate the canopy. Other scattered canopy species are Red Mulberry (Morus rubra), Post Oak (Quercus stellata), Persimmon ( Diospyros virginiana), Black Locust (Robinia pseudoacacia), Sycamore (Platanus occidentalis) (one), Sugar Maple (Acer saccharum) (one) and Tuliptree (one) . Red Maple and Scarlet Oak occur in a few locations of high acidity such as chert outcrops and talus slopes. Virginia Juniper (Juniperus virginiana) is scattered throughout; there are a few White Pine and the alien Tree- of Heaven (Ailanthus altissima) occurs in a few openings. As in the ericaceous oak forest, Flowering Dogwood is a common understory tree, but Downy Serviceberry is much less common, and Redbud (Cercis canadensis) is an additional species here. Also, as previously indicated, Hackberry is of stunted form and really forms the most abundant understory in places. Other small trees and shrubs, mostly at the edges of this forest, are Black Haw Viburnum, Wild Plum (Prunus americana), American Hazelnut (Corylus americana), Coral Berry, Black Raspberry (Rubus occidentalis), Allegheny Blackberry (R. allegheniensis) and Lance-leaved Buckthorn (Rhmnus lanceolata) (one).
Japanese Honeysucle (Lonicera japonica) is the most common vine. Among natives Hispid Greenbrier (Smilax hispida), Virginia Creeper, Poison Ivy and Winter Grape (Vitis vulpina) dominate, but Summer Grape, Canada Moonseed (Menispermum canadense), Common Greenbrier (Smilax rotundifolia), Bittersweet (Celastrus scandens) and Northern Dewberry also occur..
Common herbs on the calcareous slopes are Lopseed (Phryma leptostachya), Honewort (Cryptotaenia canadensis), Starry Campion (Silene stellata), Plume Lily ( Smilacina racemosa), Nodding Fescue, Rattlesnake Fern (Botrychium virginianum), the tick-trefoils Desmodium glutinosum and D. paniculatum, Bloodroot (Sanguinaria canadensis), Hispid Buttercup (Ranunculus hispidus), Allegheny Crowfoot (R. allegheniensis), Kidney-leaf Crowfoot ( R. Abortivus), Hooked Crowfoot (R. recurvatus), the alien Bulbous Buttercup (R. bulbosa ), Hog Peanut (Amphicarpa bracteata ), Thimbleweed (Anemone virginiana), Mealy Bellwort (Uvularia perfoliata), Three-lobed Violet (Viola triloba), Tall Agrimony ( Agrimonia gryposepala), Black Cohosh (Cimicifuga racemosa), Smooth Sweet Cicely (Osmorhiza longistylis), Wild Geranium (Geranium maculatum), Wild Comfrey (Cynoglossum virginianum), Pale Indian Plantain (Cacalia atriplicifolia), Pilewort (Erechtites hieracifolia), Wild Liquorice (Galium circaezans), Cleavers (Galium aperine), Sweet-scented Bedstraw (Galium triflorum), Virginia Knotweed (Polygonum virginianum), Clearweed (Pilea pumila), Hairy Skullcap (Scutellaria elliptica), Wingstem (Verbesina alternifolia), May Apple (Podophyllum peltatum), Enchanter's Nightshade (Circaea quadrisulcata), Poverty Oat Grass, Carex rosea, C. sparganioides, C. laxiflora, Ebony Spleenwort (Asplenium platyneuron), Dotted St Johns Wort ( Hypericum dentatum), Upright Yellow Wood Sorrel (Oxalis stricta), Adder's Tongue (Ophioglossum vulgatum), Christmas Fern (Polystichum acrostichoides), Common Many Knees, Stickweed (Hackelia virginiana), Cranefly Orchid (Tipularia discolor) and the panic grass Panicum boscii. Under moist conditions Spotted Jewelweed (Impatiens capensis) seeds on the lower slope just above the wetland where it always occurs. Additionally, Wild Pink (Silene pensylvanica) is widely distributed in sunny exposures also on limestone outcrops - on the southeast slope as well as in ericaceous oak forest of the ridge crest, but appears to be absent on the northwest slope. Also Yellow Stargrass and the tick-trefoil Desmodium rotundifolium, species usually found in acidic terrain, were found in the Chinquapin Oak White Ash forest. Soil in close proximity to the former had pH=5.8., hinting that acid leaching may enable these species to grow here at the margins of their stability fields. A single clump of Lady Fern (Athyrium filix-femina) was observed in a chert talus area of the southeast slope.
The northeast end of the ridge is occupied by a dry mesic stand with a canopy little larger than sapling size, a consequence of recent reversion from pasture. Aspect here is almost due south and the slope gentle. Dominance is by the four species of hickories previously listed, with less common Black and White Oaks, Black Walnut, Slippery Elm and minor Chinquapin Oak and Red Mulberry. Black Haw Viburnum is common and a single small Spice Bush (Lindera benzoin) was seen. Many of the mesic herbs already listed for the calcareous slopes occur here, but some additions include the somewhat calciphile sedges Carex granularis, and C. frankii, Cutleaf Toothwort (Dentaria laciniata), Downy Wood Violet (Viola sororia ), Tinker's Weed (Triosteum perfoliatum), Squaw Weed (Senecio obovatus), Virginia Avens (Geum virginianum), Four-leaved Milkweed (Asclepias quadrifolia), Riparian Wild Rye (Elymus riparius), White Grass ( Leersia virginica) and the Blue-eyed Grass Sisyrinchium angustifolium. The two sedges, which are common in the nearby wetland, may be metastable ( Mueller, 2000) here, since they could have been established in an unusually wet year. Where this stand borders ericaceous oak forest and at certain leached spots in the interior of the stand, one may also find Spotted Wintergreen, Indian Pipe (Monotropa uniflora) and Pinesap (M. hypopithys) . Also locally, within the stand, are small, usually elevated areas, of apparently more intense leaching, and in these areas one may find small patches of Polytrichum moss.
Down slope from this stand, where a pasture opening with scattered Black Walnut is still reverting to forest, the following were noted: Oldfield Goldenrod (Solidago nemoralis), Canada Goldenrod ( S. canadensis), Wild Bergamot ( Monarda fistulosa), abundant New York Ironweed, Spanish Needles (Bidens bipinnata), Cudweed (Gnaphalium obtusifolium), Columbine (Aquilegia canadensis) and Purple Cliffbrake (Pellaea ochropurpurea) on limestone outcrops, Purpletop (Triodia flava), Kentucky Bluegrass (Poa pratensis), Hairy Panic Grass (Panicum lanuginosum), Few-flowered Panic Grass (Panicum oligosanthes), Canada Bluegrass, Smooth Ground Cherry (Physalis subglaberata) and Daisy Fleabane (Erigeron annuus).
We now discuss the communities which are located in closest proximity to or includeHydrastis as a component. This herb is characterized by a fleshy rhizome and dense, almost monocultural colonies. However, it is not clear that this mode of growth is a result of propagation by rhizomes. The following species were recorded within ten meters of the largest patch ofHydrastis on the calcareous slope:Black Walnut, Slippery Elm, Hackberry, White Ash, Bitternut Hickory, Mockernut Hickory, Red Hickory, Black Oak, White Oak, Black Cherry, Virginia Juniper, Red Maple (one seedling), Flowering Dogwood. Black Haw Viburnum, Coral Berry, Carolina Rose, Virginia Creeper, Japanese Honeysuckle, Winter grape, Poison Ivy, Hispid Greenbrier, Bloodroot, Hispid Buttercup, Allegheny Crowfoot, Kidney-leaf Buttercup, Bulbous Buttercup, Desmodium glutinosum, D. paniculatum, Hog Peanut, Thimbleweed, Mealy Bellwort, Three-lobed Violet, Tall Agrimony, Black Cohosh, Smooth Sweet Cicely, Wild Geranium, Wild Comfrey, Cleavers, Wild Liquorice, Sweet-scented Bedstraw, Hairy Skullcap, Rattlesnake Fern, New York Ironweed, Wingstem, Wavy-leaf Aster, Calico Aster, May Apple, Stickseed, Enchanter's Nightshade, Autumn Bent Grass, Panicum boscii, Common Plantain (Plantago rugelii), English Plantain (P. lanceolata), Daisy Fleabane, Carex rosea, Carex laxiflora, Common Ragweed (Ambrosia artimisiifolia), Nodding fescue, European Selfheal (Prunella vulgaris), Ebony Spleenwort, Common Dandelion (Taraxacum officinalis), Spring Cress (Barbarea verna), Wild Garlic (Allium vineale), Avens (Geum sp), Canada Bluegrass, Wedge Grass (Sphenopholis sp), Dotted St Johns Wort, Kentucky Bluegrass, Hedge Buckwheat (Polgonum cristatum), Brome Grass (Bromus sp), and Blunt-lobed Woodsia (Woodsia obtusa) .
Associated with the smaller patch of Hydrastis along the contour were the following:Northern Red Oak, White Basswood, Slippery Elm, Bitternut Hickory, Hackberry, Black Cherry, White Ash, Red Mulberry seedling, Red Maple seedling, Virginia Juniper seedling, Flowering Dogwood, Black Haw Viburnum, Coral Berry, Winter Grape, Japanese Honeysuckle, Hispid Greenbrier, Virginia Creeper, Wild Rye (Elymus sp), Black Cohosh, Wild Geranium, Three-lobed Violet, Cleavers, Rue Anemone, Nodding Fescue, Kidney-leaf Crowfoot, May Apple, Carex cephalophora, Carex laxiflora, Carex sparganoides, Carex rosea, Hog Peanut, Sweet-scented Bedstraw, Hedge Buckwheat, Bloodroot, Great Chickweed, Cut-leaf Goldenrod, Canada Bluegrass, Enchanter's Nightshade, Black Snakeroot (Sanicula canadensis), Common Many Knees, Ebony Spleenwort, Smooth Sweet Cicely, Wild Liquorice, Mealy Bellwort and Tall Agrimony.
The southeast slope is prevailingly calcareous and many pH values, particularly in the colluvial soils of the steeper slopes, fall in the range of 6.5 to 8.0. However there are many places, even in close proximity to limestone outcrops, where the soils are relatively acidic, likely as a result of vertical acid leaching. Here pH may fall below 5.0. In one perhaps not unusual case, a 6 inch (15 cm) deep depression on a massive limestone outcrop bore one plant each of the acidiphiles Spotted Wintergreen and Red Maple, with the latter being a tiny, thick-stemmed and possibly old "seedling". Soil within the depression was rich in organic matter and, although in close contact with both limestone and plant roots, had a pH of 5.0. Clearly, what is significant here is the position of these acidiphiles on top of massive, slow- to-dissolve limestone.
During September pH was determined for soil at a 2-3 inch (5-8 cm) depth within and in the vicinity of the Hydrastis patches. A value of 5.8 was obtained from within the largest patch. The same value was also determined in close proximity to Blunt-lobed Woodsia a little upslope from this patch. However, soil at the base of a Chinquapin Oak, also a little upslope, yielded an initial value of pH= 6.2 and 24 hours later one of 7.0. Soil from within the second and smaller patch had pH= 6.2 and again, after 24 hours this increased to 7.0.
Values of soil pH, again from depths of 2-3 inches in close proximity to plants of Ginseng (Panax quinquefolius) in the vicinity of the Hydrastis patches, were also determined. A soil sample from the calcareous slope had an initial value of 7.2, but 24 hours later this decreased to 6.3. A sample obtained from chert talus and part of a mesic herb flora, yielded a pH of 6.6, but 18 hours later one of 8.0. It seems likely that although the first sample was from a calcareous soil it contained enough siliceous material to lower the pH. Similarly, the second sample, although from an apparently siliceous talus, may have incorporated some fine calcareous material that reacted in this way. Of related interest was the presence, also within the chert talus, and less than a meter from the second sample, of Spotted Wintergreen.
The southwest end of the ridge consists of a mostly gently southeast sloping pasture that is reverting to forest. The primary woody pioneer species here is Black Cherry, but Virginia Pine, Pitch Pine, a single Table Mountain Pine (Pinus pungens), Black Walnut, hawthorn, including Cockspur Thorn (Crataegus crus-galli), Black Haw Viburnum, Coral Berry and the aliens Autumn Olive (Elaeagnus umbellata), Morrow's Honeysuckle (Lonicera morrowi), abundant Japanese Honeysuckle and Multiflora Rose ( Rosa multiflora ) also occur. Among other species observed were Virgin's Bower (Clematis virginiana), Black Raspberry, Allegheny Blackberry, Bittersweet, Broomsedge (Andropogon virginicus), Kentucky Bluegrass, abundant Bulbous Buttercup, Bracted Plantain (Plantago aristata), Wild Bergamot, Clearweed under Black Cherry, White Vervain (Verbena urticifolia), Bugleweed (Lycopus virginicus), Great Blue Lobelia (Lobelia siphilitica) and Poke (Phytolacca americana) . A number of these are characteristic of rich, moist soils, while others are typical of dry calcareous soils.
The acidity of soils in this reverting pasture ranges widely, and since slopes are gentle for the most part, these soils are largely residual and thus have been subject to considerable acid leaching. Measured pH values range from less than 5.0 under pine to greater than 7.0 under such species as Cockspur Thorn.
While their faunas are integral parts of the several forest types represented here, it has not been possible to make detailed correlations between them. Also, in the absence of systematic observations it is not possible to prioritize the relative contributions of the various animal species to the ecosystem. Most conspicuous and apparently most closely associated with the canopy are the tree squirrels. Of these the Eastern Gray (Sciurus carolinensis) appears to be the most abundant. However the Eastern Fox Squirrel (S. niger) is also common, as is the Eastern Chipmunk (Tamias striatus). The Red Squirrel (Tamiasciurus hudsonicus ) has been observed on a northern aspect of the neighboring ridge, but not here. Although not seen on this ridge, the Southern Flying Squirrel (Glaucomys volans) is likely to be present, as it is common in adjacent areas. It is significant that the frequency of squirrel nests on the ridge appears to be the greatest of any area viewed by this reporter and is likely more than one per acre. In a winter traverse of the ridge 25 nests were counted. The nest density appears to be highest in the calcareous southeast facing slopes, perhaps reflecting the richer soils as well as sun exposure there.
The Woodchuck (Marmota monax) is also common, as indicated by sightings and burrow frequency. Perhaps the dominant native herbivores are the Eastern Cottontail Rabbit (Sylvilagus floridanus) and the White-tailed Deer ( Odocoileus virginianus), with the latter in superabundance due to the lack of sufficient large predators. It is of particular interest to our inventory that while many plants are affected by deer browsing and grazing,Hydrastis may benefit from them. This is a result of the apparent shunning of this plant by all herbivores, including insects and domestic goats and horses as well!
Of the smaller mammalian herbivores mice are likely to be important, and of these, the White-footed (Peromyscus leucopus) and Deer (P. maniculatus) Mice are most frequently observed in the forest. Additionally, forest soils are everywhere kept well aerated by the Eastern Mole (Scalopus aquaticus). It is further recognized that below these in size there exists a vast community of arthropods and other invertebrates, vegetation and detritus consumers, that may outclass all vertebrates. Of these, most conspicuous, if not of greatest import in recent times, was the alien Gypsy Moth (Lymantria dispar), which five years before the present, had defoliated the ridge several years in succession, then vanished abruptly from natural causes. However the vigor of the ridge forest was such as to enable it to recover from the defoliation with minimal observed effect. A more constant, although not as severe a defoliator, especially of Black Cherry, is the Eastern Tent Caterpillar (Malacosoma americanum), which may be responsible for the stunted and crooked form of this tree in the Valley and Ridge physiographic province.
The small mammal herbivores are evidently well able to sustain their populations in the face of numerous small predators. Among mammals the most frequently observed are Raccoon (Procyon lotor), Gray Fox (Urocyon cinereoargenteus), Red Fox (Vulpes fulva), Opossum (Didelphis marsupialis), Striped Skunk (Mephitis mephitis), Longtail Weasel (Mustela frenata), various shrews and, by incursions from the nearby lowlands, Mink (Mustela vison) . These are joined by Great Horned Owl (Bubo virginianus), Screech Owl (Otus osio), Red Shouldered Hawk (Buteo lineatus) and Red-tailed Hawk (B. jamaicensis), as well as a number of less common raptors. Also contributing are such snakes as the Racer or "Black Snake" ( Coluber constrictor), which is the most commonly observed snake. Even more frequently observed is the largely vegetarian Eastern Box Turtle (Terrapene carolina), which reaches high densities in the area.
Because of the small size of the ridge forest, bird species exclusive of the wide ranging raptors are those adapted to edge habitats and well able to cope with predators. Among these are Yellow-billed Cuckoo (Coccyzus americanus), Rufous-sided Towhee ( Pipilo erythrophthalmus), Tufted Titmouse (Parus bicolor), White-breasted Nuthatch (Sitta caroliniensis), Great Crested Flycatcher (Myiarchus crinitus ), Carolina Wren (Thryothorus ludovicianus), Brown Thrasher (Toxostema rufum), Blue Jay (Cyanocitta cristata), Cardinal (Cardinalis cardinalis), American Robin (Turdus migratoris), American Crow (Corvus brachyrhnchos), and Mourning Dove (Zenoida macroura) . For the size of the bird and tree sizes on the ridge, the Pileated Woodpecker (Dryocopus pileatus) is extraordinarily common and has nested there, perhaps again reflecting the richness of habitat. However, the Red-bellied (Melanerpes carolinus) and the Downy (Picoides pubescens) are the most common woodpeckers.
The most frequently heard forest interior bird is the Red -eyed Vireo (Vireo olivaceus), while others, such as Scarlet Tanager (Piranga olivacea), Wood Thrush (Hylocichla mustlina) and Ruffed Grouse (Bonasa umbellus) are only rarely heard or seen due to the limited area, and probably do not nest on the ridge. An important indicator species, the Whip-poor-will (Caprimulgus vociferous) was frequently heard several decades ago, but has not been seen or heard since. There remain however, two large birds that are frequently seen on the ridge. They are the Wild Turkey (Meleagris gallopavo) and the Turkey Vulture (Cathartes aura) . It is unlikely that either of these nest on the ridge. Of the two, Wild Turkey is apt to be most attracted by the many feeding opportunities provided by the ridge's productive nut trees, while large groups of the Vultures at times perch overnight.
Just as the species that are present on the ridge reflect the general environment and microhabitats there, so also those which are absent but occur on neighboring ridges contribute to our knowledge as well. Thus the steep northern aspect of a 300 foot (92 m) high ridge across the valley supports a rich mesic flora that includes the following: Beech ( Fagus grandifolia), basswoods ( Tilia heterophylla and T. americana), Butternut (Juglans cinerea), Hophornbeam (Ostrya virginiana), Muscletree (Carpinus caroliniana), Spice Bush in abundance, Wood Nettle (Laportea canadensis), Horse Balm (Collinsonia canadensis), Indian Turnip (Arisaema triphyllum), Pale Jewelweed (Impatiens pallida), Wreath Goldenrod ( Solidago caesia), Virginia Spring Beauty (Claytonia virginiana), Smooth Yellow Violet (Viola pensylvanica), Large Yellow Ladyslipper (Cypripedium pubescens), Intermediate Shield Fern (Dryopteris intermedia) and Hay-scented Fern (Dennstaedtia punctilobula) . Of these, only White Basswood and Spice Bush were observed on the Hydrastus-bearing ridge, and then only as a single plant each. Since the neighboring ridge is also underlain by limestone in this case by Cambrian Elbrook Formation we must look to such factors as moisture and temperature influences to explain the presence of these species on one ridge and not the other. It seems likely that the northern aspect and the expected higher moisture content and lower temperatures of the soil are involved, since most of the listed species for the neighboring ridge demand such soils. It must not be assumed, however, that because Hydrastis does not occur on the neighboring ridge, that the environment there excludes it. We shall see that the other occurrence reported on for this plant here is in a quite cool, moist environment as indicated by coexisting species and general conditions.
Granitic Substrate
Our second occurrence of Hydrastis, here on a substrate derived in large part from granitic rocks, lies in a small stream valley. This valley opens to a little east of south, resulting in generally south southeast aspects. Slopes are gentle on a terrace of granitic boulder and cobble alluvium, and elevations are similar to those of the occurrence on calcareous terrain. Although it lies in mountainous terrain, maximum elevations in the watershed are only moderate by Appalachian standards. All our inventory traverses but one were in proximity to the stream, here fictitiously called "Main Branch", but one also crossed the "Side Branch" that joins Main Branch from the northwest and passed through upland forest for a distance. The main body ofHydrastis occupies the alluvial terrace, but the plant is also abundant in places on the natural levee of Main Branch.
A mature but not old closed canopy forest occupies the Main Branch Valley and this forest is dominated by mesic species. In most areas of our inventories Tuliptree appears to be the most common species, but in some areas Beech is equally important. White Pine is more conspicuous than abundant as large isolated trees. Of less importance but common are Red Maple, Black Birch, White Ash, Black Gum, Black Locust, Black Cherry, American and White Basswoods, Canada Hemlock, Cucumbertree (Magnolia acuminata), Sassafras, Black Walnut, Slippery Elm and Sycamore along streams. Oaks are relatively unimportant, but Northern Red, Black and White Oaks occur, with the latter being most abundant on bordering uplands. Shagbark Hickory (Carya ovata) is almost the only hickory identified and appeared to occur mostly as seedlings and small saplings. A single Mockernut was however seen in upland forest. American Chestnut sprouts have only a minor presence. Of special interest is the scattered occurrence of Yellow Biirch (Betula alleghaniensis) and Butternut, both with decided concentrations near the stream. Of equal interest is the presence of a little Black Ash (Fraxinus nigra).
Understory species are Muscletree, Flowering Dogwood, Downy Seviceberry and Alternate-leaf Dogwood (Cornus alternifolia) . Maple-leaf Viburnum appears to be far and away the most common shrub, but there are also Coral Berry, Black Haw Viburnum, Spice Bush, Witch Hazel (Hamamelis virginiana), Winterberry Holly (Ilex verticillata), a little azalea, likely Pinxter, and Beaked Hazelnut (Corylus cornuta). Smooth Alder (Alnus serrulata) occurs along the stream and there are Upland Low Blueberry and Deerberry in a few places. Vines are not prominent in this forest, but Virginia Creeper and Poison Ivy are common as ground flora. Common Greenbrier is the only woody greenbrier noted, and there is a little Bittersweet, Summer Grape and Fox Grape (Vitis labrusca), with the latter apparently confined to the proximity of streams.
Before discussing the inventories in detail, we present a summary list of observed herbs. These are listed in the approximate order in which they were first encountered during the traverses and thus give some indication of association. Species marked with an * indicate those which we did not encounter in our inventory of the calcareous terrain. Note also, that in most of the text, in the interest of brevity, the scientific genus name "Medeola"is substituted for the common name "Indian Cucumberroot".
A late May traverse Crossing Main Branch and Terrace
*New York Fern (Thelypteris noveboracensis), Christmas Fern,* Showy Orchid (Orchis spectabilis), * Carrion Flower (Smilax herbacea), * Indian Cucumberroot (Medeola virginiana), *Dwarf Iris (Iris verna), * Carex scabrata, Hispid Buttercup, a blue-eyed grass (Sisyrinchium sp), Nodding Fescue, Black Cohosh, *Showy Skullcap (Scutellaria serrata), Hairy Bedstraw, Thimbleweed, Hog Peanut, *Miterwort (Mitella diphylla), Rattlesnake Fern, Pale Indian Plantain, *Sensitive Fern (Onoclea sensibilis) , *Ground Pine (Lycopodium flabelliforme), May Apple, *a wedge grass (Trisetum pensylvanicum), *Carex prasena, * Carex laevivaginata, *Silvery Glade Fern (Athyrium thelypterioides), * Pennsylvania Bitter Cress (Cardamine pensylvanica), Virginia Knotweed, Lady Fern, * Maidenhair Fern (Adiantum pedatum), Downy Rattlesnake Plantain, White Wood Aster, *Tree Clubmoss (Lycopodium obscurum), *Broad-leaf Goldenrod (Solidago flexicaulis), * a woodrush (Luzula echinata), * Sweet White Violet (Viola blanda), *Shining Clubmoss (Lycopodium lucidulum), * Painted Trillium ( Trillium undulatum), Ginseng, Great Chickweed, Virginia Strawberry, Mealy Bellwort, Goldenseal, the forest grass *Brachyelytrum erectum, Wild Comfrey and *Tasslerue (Trautvetteria caoliniensis)
An early September Traverse Along Main Branch
Wingstem, *Hairy Sweet Cicely (Osmorhiza claytoni), Tall Agrimony, White Grass, Lopseed, *Bottlebrush Grass ( Elymus hystrix), *Indian Turnip, the tick-trefoil Desmodium nudiflorum, Honewort, Sweet-scented Bedstraw, Field Basil, Downy Wood Violet, a panic grass Panicum boscii, Common Speedwell. Ebony Spleenwort, * Intermediate Shield Fern, Spotted Wintergreen, *a ladyslipper (Cypripedium sp), *Tall Coneflower (Rudbeckia laciniata), * Tall Meadowrue (Thalictrum pubescens), *Wood Nettle, * Northern White Violet (Viola blanda), * Lettuce Saxifrage (Saxifraga micranthidifolia), Calico Aster, Great Blue Lobelia, *Golden Ragwort (Senecio aureus), *False Nettle (Boehmeria cylindrica), Clearweed, Spotted Jewelweed, *Crested Shield Fern (Dryopteris cristata), Cutleaf Grapefern, Curtis Goldenrod, *Scirpus polyphyllus , *Interrupted Fern ( Osmunda claytoniana), *Wood Reed Grass (Cinna arundinacea), *Small Coralroot (Corallhorhiza odontorhiza), *Beech Drops (Epifagus virginiana), *Deertongue Grass (Panicum clandestinum), * Bushy Panic Grass (Panicum dichtomum) , *Partridge Berry (Mitchella repens), *Wild Sarsaparilla ( Aralia nudicaulis) and Pinesap.
An Early September Traverse to the South, Crossing Side Branch
*Broad-glumed Brome Grass(Bromus latiglumis), Wild Liquorice, *Philadelphia Fleabane (Erigeron philadelphicus), *Round-leaved Pyrola (Pyrola rotundifolia), *Carex virescens, *Carex gracillima, *deciduous Spinulose Shield Fern (Dryopteris carthusiana ), *Canada Brome Grass (Bromus pubescens), Carex laxiflora, *Horse Balm, Autumn Bent Grass, *Wrinkle-leaf Goldenrod ( Solidago rugosa), *Water Hemlock (Cicuta maculata), *Purple-stem Aster (Aster puniceus ), Common Plantain, *Golden Saxifrage (Chrysosplenium americanum) and *Scouring Rush (Equisetum hyemale) .
Mid-July Traverses on the Terrace and Along Main Branch
Smooth Sweet Cicely, White Vervain, *Tall Bellflower (Campanula americana), *Moccasin Flower (Cypripedium acaule), *a manna grass (Glyceria melicaria), *Carex laxiculmis, *Small Green Wood Orchis (Habenaria clavellata), *Cinnamon Fern (Osmunda cinnamomea), *Groundnut (Apios americana), * Cowbane (Oxypolis rigidior) *Carex gynandra, *American Selfheal (Prunella vulgaris var lanceolata), *White Clintonia ( Clintonia umbellulata), *Yellow Passion Flower (Passiflora lutea), *Basil Balm (Monarda clinopodia), *Tree Clubmoss (Lycopodium obscurum), *Broad Beech Fern (Phegopteris hexagonoptera), Plume Lily and *Royal Fern (Osmunda regalis) . The northern disjunct *Bog Bluegrass (Poa paludigena) has also been reported in the area (G. Fleming, personal communication), but was not seen by us.
The Late May Traverse Across Main Branch and the Terrace Details
Entering the forest above the Side Branch confluence, we crossed Main Branch and moved west across the terrace. At the stream crossing we noted Muscletree, Black Haw Viburnum, Witch Hazel, New York and Christmas Ferns, Showy Orchid, Carrion Flower, Medeola, Dwarf Iris and Carex Scabrata. On the terrace we saw Hispid Buttercup, a blue-eyed grass, Nodding Fescue and uncovered a Seal Salamander (Desmognathus monticola) under forest debris. Continuing, we noted the abundance of Tuliptree, an azalea (likely Pinxter), Red Maple, Black Cohosh, Showy Skullcap, Hairy Bedstraw, Thimbleweed, Hog Peanut, more Medeola and Miterwort.
Still moving west on the terrace, we came under tall, young Tuliptrees with a ground cover of Rattlesnake Fern, more Showy Orchid and Pale Indian Plantain. Delicate Fern Moss (Thuidium delicatulum) was common on down boles.
We now reached the edge of the terrace at the base of a steep slope with an easterly aspect. Here we saw the first Black Birch and a single Yellow Birch, perhaps bespeaking an increase in moisture from colluvial sources. Other species were Sensitive Fern, scattered small patches of Ground Pine, Nodding Fescue, May Apple, the wedge grass Trisetum pensylvanicum, an indicator of abundant moisture, Carex prasina, an inhabitant of "wet woods and thickets" (Strausbaugh and Core, 1977) ; as also is the coexisting Carex laevivaginata. With these were Silvery Glade Fern, Beech, the fungi Rickenella fibula and Hygrophorus margiatus, Pennsylvania Bittercress, Virginia Creeper, Hog Peanut, abundant Virginia Knotweed and common Medeola. We now heard an Ovenbird (Seiurus aurocapillus) and soon after saw our first patch of Tree Moss (Climacium americanum), which is usually an indicator of rich, moist soils. Next seen was Lady Fern, then a little Maidenhair Fern, a small clump of Downy Rattlesnake Plantain, White Wood Aster and, on trees, the lichen Lobaria pulmonaria. Following these were a few unhealthy appearing Canada Hemlock saplings and more Tree Moss. We now reached a small ravine in which grew a single Sweet White Violet, Shining Clubmoss, Painted Trillium and the fungus Leccinum albellum. A Scarlet Tanager was heard here.
Directing our traverse toward the stream again, we encountered our first Ginseng, here under Red Maple and with nearby Hog Peanut, Maple-leaf Viburnum, Medeola, Virginia Strawberry and Mealy Bellwort. Immediately following was our first Hydrastis, accompanied by a little of the forest grass Brachyelytrum erectum; then more Ginseng, a single mature Yellow Birch and Wild Comfrey. Also seen here was a small frog, possibly a tree frog, on the forest floor. We next came to an area in which the forest floor was dominated by Ground Pine, but with some Hydrastis as well. The latter soon increased and formed large patches with a sprinkling of Ginseng. As we again reached the stream, somewhat upstream from our initial crossing, we noted a luxuriant growth of herbs, including large, robust Hydrastis and very large Tasslerue.
Early September Traverse North Along Main Branch Details
This traverse began in the area of greatest concentration of Hydrastis on the terrace. It was then estimated that this plant occupied most of the area covered by ground flora for a distance of perhaps 600feet (180 m) parallel to the stream and 150 feet (46 m) perpendicular to it. Thus the area of major concentration may be of the order of two acres (0.8 ha) . Additionally, many small patches and individual plants of Hydrastis occur outside this major concentration, Some patches of the plant in the major area of concentration are of the order of 100 feet (30 m) across, and within them it forms 90% of the ground flora.
The canopy in the major area of concentration of Hydrastis is dominated by Tuliptree punctuated by large White Pine. Other canopy species are Beech, Black Birch, Red Maple, Black Locust, Northern Red Oak, White Ash, Black Cherry of poor form, Cucumbertree, Black Walnut, Sassafras, a few Yellow Birch, Shagbark Hickory and Black Oak, with the last two practically confined to seedlings. Muscletree dominates the understory, but Flowering Dogwood and Downy Serviceberry also occur.
Although Spice Bush and Black Haw Viburnum may be the most conspicuous shrubs, Maple-leaf Viburnum is almost omnipresent as seedlings and small plants, although these appear to be less frequent within the Hydrastis patches. While Coral Berry too is widespread, it is not as common, nor does it attain the large size that it does in calcareous terrain. Next to Virginia Creeper and Poison Ivy, Common Greenbrier appears to be the most common vine. The little Summer Grape observed appeared to be concentrated near the stream.
Hydrastis was observed to extend virtually to the base of a 34 inch (0.86 m) dbh White Pine (see later for a pH determination here) . Other herbs within the Hydrastis patches included very common Virginia Knotweed, blooming White Wood Aster, Hog Peanut, Scattered Wingstem, Hairy Sweet Cicely, Tall Agrimony, White Avens, Mealy Bellwort, Ginseng, Rattlesnake Fern, Lopseed in seed, Bottlebrush Grass, Carrion Flower, IndianTurnip, Desmodium nudiflorum, Honewort, Sweet-scented Bedstraw, Field Basil, Downy Wood Violet and Panicum boscii.
Herbs less clearly associated with Hydrastis are Christmas Fern, Ground Pine in patches that interdigitate with it, Cutleaf Grapefern, Medeola, Common Speedwell, Ebony Spleenwort, Downy Rattlesnake Plantain, a very little Intermediate Shield Fern, Spotted Wintergreen and Shining Clubmoss. Red Maple seedlings appear to be more common outside the patches. Mosses noted were Delicate Fern Moss, one or more species of Hypnum on down boles, Mnium affine and an unidentified Entodon on rocks.
Pileated Woodpeckers were heard, a Gray Treefrog (Hyla versicolor/chrysocelis) called once and a male Box Turtle was found in the area.
As we moved north from the main Hydrastis patches, we encountered a small Beech sapling with numerous large, white wooly aphids that covered a 0.5 inch (1.3 cm) thick branch. These aphids waved their posteriors vigorously in the air. An associated fungus may have resulted from honeydew. Another fungus on the ground in the vicinity was an unidentified species of Suillus.
We now reached the cobble levee that bordered Main Branch and noted again that Hydrastis in this environment was the most robust of any ever seen by us. As far as we could determine by a rather cursory examination, individual plants of the species were not connected by rhizomes, but at least five leaf scars were present on some of them. Other species on the levee were Tall Meadowrue, Wood Nettle, abundant large Christmas Fern, Tall Coneflower, Tree Moss, Spice Bush, Virginia Knotweed, White Ash, Calico Aster and Sweet White Violet. Although not recorded here, Tasslerue had been found in a similar location along this stream on our May visit.
Farther along the stream edge were Lettuce Saxifrage, abundant Carex scabrata, Great Blue Lobelia in bloom, Golden Ragwort, False Nettle, Clearweed, Spotted Jewelweed, also in bloom, and a Yellow Birch sapling.
On higher ground again, and with little or no Hydrastis in view, we saw the fungi Naematoloma fasciculare on buried White Pine, Scleroderma citrinum, Mycena haematopus and Amanita virosa; then an unidentified species of Hypnum moss on a down bole, a two foot (0.6 m) dbh Tuliptree, an unexpected single plant of Crested Shield Fern, Small Coralroot Orchid, Cutleaf Grapefern, a little Hydrastis, an unidentified Lepiota fungus, Curtis Goldenrod, an 11 inch (0.28 m) dbh Beech and Beech Drops in bloom.
As our traverse once again took us to the stream edge, we noted there Scirpus pollyphyllus, Winterberry Holly, Lady Fern, Silvery Glade Fern, Shining Clubmoss, Sensitive and Interruped Ferns, Smooth Alder, more Crested Shield Fern, Great Chickweed, a 16 inch (0.41 m) dbh Beech and Wood Reed Grass.
Then, once more ascending the terrace slope, we saw the fungus Collybia confluens, a saprophyte on fallen leaves, abundant Medeola, Downy Rattlesnke Plantain, Deertongue Grass, Bushy Panic Grass and unidentified Xerula and Leccinum fungi.
The Valley now narrowed greatly and the canopy consisted of Beech, Tuliptree, Red Maple, large White Pine, Black and Yellow Birches and Hemlock. Also seen were the mossBryoandersonia illecebra and a large (3.5 inch/ 9 cm ), faintly striped dark brown millipede with no detectable odor ( Narceus americanus ?) . These were followed by an unidentified Psathyrella fungus, a 19 inch (0.48 m) dbh Beech (very healthy appearing, as were all Beech in the area) ; also abundant Ground Pine and Medeola, common Yellow Birch and the fungi Pluteus cervinus and Green Stain (Chlorociboria aeruginascens) with fruiting bodies both on decaying wood. Another spot inventory of the canopy yielded, in order of importance, Beech, Tuliptree, Black and Yellow Birches and, in the distance, on the bordering west slope, a very large White Oak with spreading crown.
Continuing up-valley we saw the mosses Dicranum scoparium and an unidentified Hypnum on rock; then a single 9 inch (0.23 m) Butternut, followed by Shagbark Hickory, the inedible "Cheese Polypore" (Tyromyces chioneus) fungus, Partridge Berry in fruit, White Wood Aster in bloom and Silvery Glade Fern. At a seep were Fowl Manna Grass, an unidentified Spikerush (Eleocharus sp), an unidentified Atrichum moss, a 15 inch (0.38 m) dbh Yellow Birch, New York Fern and a peculiar "epiphytic" fungus that resembled a sponge on a twig.
Our traverse, which had extended up-valley for perhaps Ό mile (0.4 km), had been confined to the west side of the stream. On our return south we made a few additional observations on the stream's east side. These included a 25 inch ( 0.64 m) dbh Beech, a Black Gum, Wild Sarsaparilla, Pinesap and the first noted blueberries, namely Deerberry and Upland Low Blueberry.
Additional fungi collected on this traverse and identified by R. Hunsucker are as follows:
Amanita cokeri
Amanita farinose
Amanita virosa bis pongera group (frequent)
Austroboletus gracilis
Baeospera myosura on White Pine cones
Boletus ornatipes
Cantharellus cinnabarinus
Clavaria cristata
Clitocybe clavipes
Clitocybe nuda
Clitocybe spp (3)
Collybia dryophylla
Cortinarius corrugatus
Cortinarius spp (2 3)
Crepidotus applanatus
Daedaliopsis ambigua
Daedaliopsis confragosa
Favolus alveolaris
Hygrophoropsis aurantiaca
Lactarius argillaceifolius
Lactarius spp
Leccinum albellum
Lepiota clypeolaria
Lycoperdon perlatum
Marasmius spp
Melaoleuca alboflavida
Mycena leaiana
Mycena pura
Pholiota sp
Pleurotus ostreatus
Pluteus cervinus
Polyporus varius
Russula spp
Stereum complicatum
Stereum striatum
Stropharia hardii
Suillus granulatus
Trametes versicolor
Trichaptum biforme
Xylaria radicata
Xylaria polymorpha
Early September Traverse South Across Side Branch Details
Our traverse to the south began at about 10:00 AM at the origin of our previous day's traverse to the north. Our traverse began by observing a Northern Cricket Frog (Acris crepitans ) hopping across the forest floor. We then successively observed the following species as we moved south: Maple-leaf Viburnum, Ebony Spleenwort, Broad-glumed Brome Grass, possible Showy Orchid, Wild Liquorice, seedlings of White Oak all more or less associated with Hydrastis. Then entering a more elevated area, we saw an unidentified Crepidotus fungus under a 32 inch (0.81 m) dbh White Pine, with scattered Hydrastis in pine needle duff; then a 32 inch dbh Tuliptree followed by leaf rosettes of Philadelphia Fleabane, an Alternate-leaf Dogwood, the fungi Suillus granulatus and Suillus americanus, both species being commonly associated with White Pine ( Phillips, 1991) . These were followed by Round-leaved Pyrola near White Pine, Carex virescens and Beaked Hazelnut near a 37 inch (0.94 m) dbh White Pine. Here also a mouse or vole was observed running across the forest floor. Next seen was Carex gracillima, the deciduous Spinulose Shield Fern Dryopteris carthusiana, the fungus Hygrophoropsis aurantiaca (partial to decaying conifers), the mosses Rhynchostegium serrulatum on soil andRhodobryum roseum on rock and Downy Serviceberry in the understory. A spot inventory here also yielded Tuliptree, Black Birch, Red Maple, Beech, Black Locust and Shagbark Hickory in the canopy and Muscletree in the understory, while the ground flora included, in addition, Canada Brome Grass, Nodding Fescue, Carex laxiflora and a little Hydrastis.
We soon reached an old tote road along which we found Wingstem, Pale Indian Plantain, Horse Balm, Autumn Bent Grass, Bottlebrush Grass, Wrinkle-leaf Goldenrod and the fungi Crucibulum leave and Lycoperdon perlatum.
As we arrived at Side Branch we noted a 15 inch (0.38 m) dbh Yellow Birch on the stream bank with Sycamore, Red Maple and a 10 inch ( 0.25 m) Muscletree. Moving upstream for a short distance, we saw Carex scabrata, Water Hemlock, very abundant Tree Moss and a Dusky Salamander ( Desmognathus sp) in the stream bed, while the far bank was occupied by Northern Red Oak, abundant Christmas Fern and Delicate Fern Moss as the dominant moss. Also seen in the general area were Witch Hazel, Lady and Sensitive Ferns, abundant Tree Moss, Wood Reed Grass, Miterwort, Thimbleweed, Purple-stem Aster and a Black Walnut. A single Mockernut Hickory stood on a nearby elevation.
We now began to cross a more steeply sloping terrace or bench to the south of Side Branch. The canopy here was dominated by Tuliptree, with Black Birch, Beech, Cucumbertree and White ash, while Flowering Dogwood and saplings of White Pine and Red Maple formed an understory. A single Chestnut Oak sapling, the only representative of the species observed in the area, was seen here. Seedlings of Shagbark Hickory, Black Oak and Red Maple were common, as was Maple-leaf Viburnum. Hog Peanut and Medeola were conspicuous in the ground flora.
We next came to a shallow swale in which lay a foot (0.3 m) thick, very long down bole entirely covered by Delicate Fern Moss. Also seen were an unidentified hot tasting Lactarius fungus, a 27 inch (0.69 m) dbh Tuliptree, Wild Liquorice, a Red-bellied Woodpecker, Ground Pine, Medeola, generally abundant Tuliptree, seedlings of basswood and hawthorn, Virginia Knotweed, Cutleaf Grapefern, delicateFern Moss on rock, Black Haw Viburnum, Muscletree and a single plant of Hydrastis.
Moving onward, we continued to see abundant Tuliptree, with White Pine, Black Cherry of characteristic crooked form, Black Locust, the fallen remains of American Chestnut, Flowering Dogwood, Poison Ivy, more Small Coralroot orchid, a 36 inch (0.9 m) dbh White Pine and fungi that included an unidentified species of Pholiota and the edible Xerula fufuracea and Suillus pictus.
We now came to a springbrook at which we saw Great Blue Lobelia in bloom, Golden Saxifrage, Lettuce Saxifrage and, virtually on the springbrook bank, a dense, five meter square patch of Hydrastis. Nearby in a low area there was Slippery Elm, Scouring Rush and a large vine of Fox Grape with ripe, tart but quite flavorful grapes.
Additional fungi collected on this traverse and identified by R. Hunsucker are as follows:
Agaricus sp
Hygrophorus cantharellus
Hygrophorus flavescens
Hygrophorus marginatus
Inonotus radicatus
Leccinum sp
Mycena sp
Panellus stipticus
Phaeolus schweinitzii
Psathyrellus sp
Stereum ostrea
Tricholoma aurantium
Mid-July Traverses Details
At about 10:30 AM, under hazy skies and cool temperatures, a foray was made to the vicinity of the largest Hydrastis patches with the chief purpose of collecting soil samples for pH determinations. After noting the presence of two Box Turtles in the area, the first sample (no 1) was collected from a depth of 2-3 inches (5-8 cm) under a large White Pine and among Hydrastis. Other species within a few meters included Maple-leaf Viburnum, Poison Ivy, Spice Bush, a Beech sapling, Muscletree, Medeola, Tuliptree, and Ebony Spleenwort. Delicate Fern Moss was the most conspicuous bryophyte. On drying, this soil sample, which was apparently quite rich in organic matter, appeared, from its lack of coherence, to be silty. It had an initial pH of 5.5 and 5.4 twenty-four hours later. It should be kept in mind that soil here is confined to narrow interstices between cobbles and boulders so that profiles are difficult to observe. However it everywhere had a "rich " appearance, with organic matter well integrated with mineral matter.
Species noted in a nearby large patch ofHydrastis were abundant Virginia Knotweed, somewhat stunted Tall Agrimony with a few blossoms, Desmodium nudiflorum, scattered Wingstem, Smooth Sweet Cicely and seedlings of Shagbark Hickory. Associated mature trees were Red Maple, American Basswood and White Ash.
Sample no2 was taken from a depth of 2-3 inches next to Downy Rattlesnake Plantain in association with sweet cicely, Hog Peanut and Wild Liquorice. Hydrastis occurred at a distance of a foot from the sample area. This sample had an initial pH of 5.8 and 24 hours later 5.7.
Other species seen within a nearby Hydrastis patch were Mealy Bellwort, stunted Rattlesnake Fern and stunted and defoliated Coral Berry.
Sample no3 was taken from, as usual, a 2-3 inch depth in a "rich" appearing semi-opening with abundant Wingstem, blooming White Vervain, Hog Peanut, Spice Bush and other mesic herbs. Black Walnut and a patch of Hydrastis were nearby. This soil sample yielded an initial pH of 5.8 and, after 24 hours, 5.7.
It may be significant that, as distinguished from its occurrence on calcareous substrate, very few of the Hydrastis plants anywhere here bore fruit, perhaps a consequence of the generally deep shade, more acidic substrate or greater competition. Also, the only bird seen or heard on this foray was a Red-eyed Vireo.
In the late afternoon a return was made to the vicinity of the Hydrastis patches, but this time on their western fringe where we had seen the most Ginseng. A soil sample (no 4) was then taken from close to a plant of Ginseng from a depth of 2-3 inches and amongHydrastis, with nearby Medeola, Red Maple, Indian Turnip, Muscletree, Wild Comfrey and Entoloma murrai, a fungus said to inhabit "damp woods and swamps" (Phillips, 1991) . An initial pH value of 5.9 was obtained, but after 24 hours this had increased to 6.2.
A second sample (no5) was also taken from next to another Ginseng plant with Spice Bush in close proximity. Also noted in the general vicinity were Spotted Wintergreen, Tall Bellflower in bloom and the deadly fungus Amanita virosa. This sample proved to have an initial pH of 5.9, with no later determination made. It appears that, as in calcareous terrain, pH in the vicinity of Ginseng is slightly but consistently higher than is general in the vicinity of Hydrastis.
On the following sunny but cool morning we initiated a traverse a little north of the major Hydrastis occurrence in the edge of the upland forest to the east of the stream. This forest was dominated by Beech, mostly small, but in some trees attaining 2 feet (0.6 m) dbh or more. Black Birch, usually not over a foot (0.3 m) dbh, was also common, and large White Pine were scattered throughout. Other species were Red Maple, Cucumbertree, Witch Hazel a few American Chestnut sprouts, Upland low Blueberry, Maple-leaf Viburnum, Partridge Berry and scattered Spotted Wintergreen and Downy Rattlesnake Plantain. Now descending again to the stream side, we noted a canopy dominated by Beech, Black Birch and Tuliptree. but with scattered Yellow Birch. Other species were Black Gum, Spice Bush, Tasslerue, abundant Lady and Christmas Ferns, Carex Scabrata, commonMedeola and a patch of Moccasin Flower. Moving downstream, we suddenly came upon several small saplings of Black Ash. One of these saplings, with a stem diameter of an inch (2.5 cm) stood in the shallow bed of a springbrook next to a much larger Black Birch on the spring bank. An unidentifiedHabenaria orchid grew nearby. Water from this springbrook was found to have a pH of 6.2.
Continuing downstream across a low terrace west of the stream, we saw round-leaf Pyrola in bloom, abundant Virginia Knotweed; then on the stream bank, patches of Tree Moss inter-grown with Partridge Berry. These were followed successively by the Old Man of the Woods (Strobilomyces floccopus ) fungus, patches of an unidentified Atrichum moss, a little Shining Clubmoss and the fungusCantharellus cinnabarrinus, which occurred as numerous bright red, tiny mushrooms distributed along the very edge of the stream bank for a considerable distance.
Next seen was an unidentified Clitocybe fungus, Tall Meadowrue, the moss Mnium cuspidatum and the thallus liverwort Conocephalum conicum with an unidentifiedPorella liverwort in the stream bed. Also present here was an unidentified species of the mossBrachythecium, as well as Hypnum curvifolium. In places in proximity to the stream the manna grass Glyceria melicaria formed dense patches and Carex laxiculmis was a common sedge. Birds heard and/or seen here included a Wood Pewee ( Contopus virens), a Rufus-sided Towhee and a Blue Jay.
Lady and Christmas Ferns continued particularly abundant and New York and Silvery Glade Fern were both common. Accompanying these were Mealy Bellwort, the mosses Anomodon attenuatus on the base of a large Tuliptree, Delicate Fern Moss and an unidentified Atrichum on soil. Also present was somewhat stunted Wood Nettle; then a few Hemlock, Yellow Birch, Summer Grape and a few plants of Bittersweet on the ground but no vines. Here also there was a single plant of what appeared to be the grape fern Botrychium biternatum. We now began to see Butternut, one of which was 15 inches (0.38 m) dbh and quite healthy in appearance. With these were Tasslerue, Muscletree, Small Green Wood Orchid in bloom, Cowbane and the fungus Amanita vaginata. These were followed by Cinnamon Fern, Groundnut, Winterberry Holly, Crested Shield Fern; then another healthy appearing Butternut at 18 inches (0.56 m) dbh. In a more than usually moist spot grew tall Carex gynandra, unidentified jewelweed, White Ash and a vigorous vine of Fox Grape with green fruit. A soil sample (no 6) was taken from a low mucky area near the Fox Grape. This soil yielded an initial pH of 5.8, but 24 hours later this had decreased to 5.4.
On again reaching the area of large Hydrastis patches, we noted the edible fungus Pluteus admirabilis, as well as a species of Melanoleuca., American Self-heal ; and near the Hydrastis, the white coral fungus Ramariopsis kunzei, a species partial to humus.
After observing an over-flight of what appeared to be two calling Sandhill Cranes (Grus canadensis) in the afternoon of the same day, we began another traverse up Main Branch. The first part of this traverse was through upland forest, again on the valley's east side, but farther upslope than before. The forest here was open and dominated by Beech, most of which was mature but not large. Accompanying it were larger White Pine, Northern Red and White Oaks and Tuliptree. One White Pine measured 34 inches (0.86 m) dbh, while a White Oak attained 36 inches (0.91 m) .Also present, and generally of smaller sizes, were Red Maple and Black Birch. There were few shrubs and little ground flora. Partridge Berry was concentrated under White Pine and Beech Drops were noted. The traverse through the upland forest was terminated perhaps Ό mile (0.4 km) above the major Hydrastis patch and a return traverse was begun downstream.
Noted first near the stream was a patch of vigorous Clintonia, likely the White species; then Yellow Birch, Cancerroot, a patch of Tree Moss / Partridge Berry intergrowth at the stream edge, abundant Christmas Fern and Poison Ivy. A soil sample (no 7) was taken on an adjacent terrace west of the stream. This sample had an initial pH of 5.7, but 24 hours later it was 5.5. Next seen on our progress downstream was the only vine of Passion Flower seen on any of our traverses. Also in the vicinity were Basil Balm, Broad Beech Fern with abundant Lady Fern and the fungus Galiella rufa. On the nearby stream bank we saw a small patch of Tree Clubmoss, the only occurrence of this species seen in the area. Also noted here was the abundance of Beech, mostly small. but including 18 inch ( 0.44 m) and 22 inch (0.55 m) veterans. Black Birch, Downy Rattlesnake Plantain and Common Greenbrier were also conspicuous. Here we reached a springbrook that discharged into the stream from the west. Recorded in the vicinity was a 32 inch (0. 81 m) Tuliptree, as well as Plume Lily and Witch Hazel, both of which we had earlier observed in scattered occurrences along the stream.
There appeared to be few tree seedlings in this forest and, as observed earlier, Water Hemlock and the Tree Moss Partridge Berry intergrowth were common near the stream, as was also Virginia Creeper. Other features of this spot included the only Royal Fern encountered by us, the abundance of seedlings and small plants of Indian Turnip, common but isolated patches of Ground Pine and the excellent edible mushroom Cantharellus fallax.
We ended our traverse by taking a water sample of Main Branch. This sample proved to have a pH a little above 6.0, perhaps 6.1 or 6.2, values in agreement with that obtained for the springbrook earlier and for other streams in the area. A mental note had also been made of a general feature of all our traverses on this terrain, namely the abundance of small lepidoptera, which, while quite substantial, did not seem as great as on calcareous terrain.
Additional fungi collected on these July traverses and identified by R. Hunsucker are as follows:
Morning Traverse
Amanita brunnescens on soil
Amanita flavoconia on soil
Amanita virosa complex on soil
Clavaria vermicularia - on soil
Clitocybe spp in humic soil
Collybia dryophylla on soil
Entoloma murrai on soil
Hygrocybe coccinea on soil
Lycoperdon pyriforme on rotting oak
Marasmius sp on dead maple twigs
Melanoleuca melaleuca on soil
Polyporus varius on decaying stems of hardwoods
Rickenella fibula in moss
Steccherinum adustum on stems of fallen hardwoods
Stereum striatum on dead branches of Muscletree
Suillus pictus on soil under White Pine
Trametes versicolor on rotting boles and branches of fallen trees
Tremella mesenterica on rotting oak
Trichaptum biforme on rotting wood
Xeromphalina kauffmanii on rotting oak
Xerula radicata on soil
Afternoon Traverse
Boletus ornatipes on soil
Cantharellus cinnabarinus on soil
Cantharellus cibarius on soil
Collybia confluens on rotting leaves
Coltricia cinnamomea on soil
Hydnum repandum ( Sweet Tooth) on soil, conifers, hardwoods
Hygrophorus ovinus on soil
Lactarius corrugis on soil
Lactarius piperatus on soil
Lactarius subplinthogalus on soil
Lactarius subvellereus on soil
Bryophytes (From the Granitic Substrate)
The following is a listing of bryophytes from 8 habitat groups that were collected and identified by Dr. Robert Hunsucker. Liverworts are distinguished by the symbol (L).
1. from the bases of trunks and exposed roots of large White Pine:
Hypnum fertile
Pottiaceae (unidentified)
2. from rotting White Pine down boles, both decorticated and with bark:
Brotherella tenuirostris
Chiloscyphus profundus (L)
Entodon compressus (on bark)
Hypnum imponens
Nowellia curvifolia (L) (on decorticated down bole)
Platygyrium repens (on bark)
Thuidium delicatulum (on bark)
3. from dead, standing deciduous snags:
Aneura pinguis
Frulannia eboracensis (L)
Halohymenium triste
Mnium cuspidatum
Platygyrium repens
Ulota crispa
4. from rotting deciduous trees, shrubs and stumps on the ground:
Amblystegium varium
Anomodon attenuatus
Anomodon rostratus
Dicranum flagwllare
Entodon compressus
Frullania eboracensis (L)
Halohymenium triste
Hypnum imponens
Hypnum pallescens
Mnium cuspidatum
Nowellia curvifolia (L)
Platygyrium repens
Pylaisiella selwynii
Rhynchostegium serrulatum
Thuidium delicatulum
Ulota crispa
5. from soil, mostly near, but not in small stream:
Atrichum angustatum
Climacim americatum
Conocephalum conicum (L) (on bank of stream)
Eurhynchium hians
Mnium cuspidatum
Plagiothecium cavifolium
Rhynchostegium serrulatum
Thuidium delicatulum
6. from rock and soil over rock:
Amblystegium serpens
Amblystegium tenax
Anomodon attenuatus
Anomodon rostratus
Brachythecium sp
Bryoandersonia illecebra
Conocephalum conicum (L) (near stream)
Entodon compressus
7. from bark of trunk and exposed roots of deciduous trees and shrubs such as Tuliptree, Black and Yellow Birches, White Oak, Red Hickory (Carya ovalis), Beech, White Ash,. Flowering Dogwood and Black Haw Viburnum.
Amblystegium varium
Anomodon attenuatus
Brachythecium acuminatum
Brachythecium oxycladon
Bryhnia graminicolor
Cololejeunea biddlecomiae (L)
Entodon sp
Frullania riparia (L)
Halohymenium triste
Metzgeria furcata (L)
Mnium cuspidatum
Platygyrium repens
Thuidium delicatulum
8. from bases and exposed roots of trees such as Tuliptree, Black and Yellow Birches, Red Hickory and White Oak:
Amblystegium varium
Anomodon attenuatus
Anomodon rostratus
Brachythecium sp (sterile) Brotherella tenuirostris
Callicladium haldenianum
Cololejeunea biddlecomiae (L)
Dicranella heteromalla
Entodon compressus
Frullania brittoniae (L)
Frullania eboracensis (L)
Haplohymenium triste
Homalotheciella subcapillata
Hypnum fertile
Hypnum pallescens
Mnium cuspidatum
Orthotrichum sp (sterile)
Plagiothecium cavifolium
Platygyrium repens
Porella platyphylla (L)
Pylaisiella polyantha
Rhynchostegium serrulatum
Rhodobryum roseum
Thelia hirtella
Thuidium delicatulum
The following shows the frequencies of the bryophyte taxa in the 8 habitat groups:
5 Anomodon attenuatus
5 Mnium cuspidatum
5 Platygyrium repens
5 Thuidium delicatulum
4 Entodon compressus
4 Haplohymenium triste
3 Amblystegium varium
3 Anomodon rostratus
3 Frullania eboracensis (L)
3 Rhynchostegium serrulatum
2 Brotherella tenuirostris
2 Cololejeunea biddlecomiae (L)
2 Conocephalum conicum (L)
2 Hypnum fertile
2 Nowellia curvifolia (L)
2 Plagiothecium cavifolium
2 Ulota crispa
1 Amblystegium serpens
1 Amblystegium tenax
1 Aneura pinguis
1 Atrichum angustatum<br> 1 Brachythecium acuminatum
1 Brachythecium oxycladon
1 Bryhnia graminicolor
1 Bryoandersonia illecebra
1 Callicladium haldenianum
1 Chiloscyphus profundus (L)
1 Climacium americanum
1 Dicranella heteromala
1 Dicranum flagellare
1 Eurhynchium hians
1 Frullania brittoniae (L)
1 Frullania riparia (L)
1 Homalotheciella subcapillata
1 Metzgeria furcata
1 Orthotrichum sp
1 Porella platyphylla (L)
1 Pottiaceae
1 Pylaisiella polyantha
1 Pylaisiella selwynii
1 Rhodobryum roseum
1 Thelia hirtella
Summary and Conclusions
The habitats of the two occurrences of Hydrastis discussed here are quite different in terms of physical setting, chemical character of substrate and the enveloping floras. While the occurrence on calcareous terrain occupies a rather steep, southeast slope in dry mesic forest, that on granitic terrain occupies a gently sloping terrace in moist mesic forest. The canopy of the former is dominated by oaks and hickories, whereas that of the latter consists overwhelmingly of mesic species such as Tuliptree, Beech, birches and Red Maple. In general character the enveloping floras of the occurrence on calcareous terrain are southern Appalachian, with only a few species of markedly northern distribution. By contrast those on granitic terrain have a generous component of northern or cool climate species such as Yellow Birch, Black Ash, Crested Shield Fern, Painted Trillium and Wild Sarsaparilla, as well as numerous moisture dependents like Indian Turnip, Wood Nettle, Horse Balm, Indian Cucumberroot and a variety of ferns that were not part of the floras on calcareous terrain. Sedges and grasses in particular differ greatly in the two occurrences, Only Carex laxiflora among sedges was identified in both; while the 16 and 14 grasses recorded in the calcareous and granitic terrains respectively had only three clearly identified species in common. This may however in part reflect our limited surveys, particularly on the granitic terrain.
While the calcareous terrain includes some highly acidic enclaves, the granitic terrain appears less, if more uniformly, acidic. Measured soil pH on the calcareous terrain ranges from 4.4 to 8.0. Although our pH determinations on the granitic terrain are few in number, the values obtained, in conjunction with the observed vegetation, lead us to believe that they generally lie between 5.5 and a little more than 6.0. This conclusion is most strongly supported by the spring and stream waters, which yield the higher values. It seems reasonable that the lower values obtained for soils, result from vertical acid leaching. The apparent absence of pH values as low as those from the calcareous terrain may be a consequence of the absence of highly siliceous rocks such as chert and the relatively subdued microtopography. It may also be an artifact of our failure to sample near the few occurrences of blueberries that we saw in the granitic terrain. A likely contributing factor on the granitic terrain is the ready availability of bases. Granite undergoing weathering might be expected to be a constant source of potassium, sodium and calcium ions, since these elements are abundant in granitic minerals such as feldspar. Magnesium, on the other hand, is likely to be less abundant than in calcareous rock.
The distribution of a number of species in the two areas of Hydrastis occurrence provide, in terms of pH and available moisture, additional information on their stability fields. Reference is here made to diagrams of inferred stability relations (Mueller, 2000) based on a variety of field occurrences. From these diagrams it appears that in both occurrences Red Maple ranges to its upper pH limit, and the same is true of Beech in granitic terrain. The common associations of Beech, White Ash and Red Maple on the latter is consistent with moderately acidic, moist soils. The virtual absence of Chestnut Oak (only one sapling seen) may also be a consequence of high soil moisture. The absence of Sugar Maple on the granitic terrain is not as easily explained however. Although this maple's inferred stability field has been placed at slightly lower soil moisture levels than that of Beech, its apparent total absence here is more likely a result of other factors perhaps biological, such, as among others, unfavorable mycorhizal associations. Because of their rarity, knowledge of the stability fields of Goldenseal and Ginseng is of special interest. However this rarity militates against seeing these species throughout their ranges in habitat. Both plants appear to be quite demanding of nutrients and well developed mull type soils. While found in both moist and dry mesic habitats, as documented here, their moisture limits are not established. The same is true of pH, although in terms of the limited measurements of this parameter reported here, as well as all unreported observations of this reporter, the upper pH limit of Ginseng appears to lie above 7.0. The even more limited observations of Goldenseal, may indicate a slightly lower pH range for this species, but this is little more than conjecture.
The limited occurrence of heath family shrubs and certain other acidiphiles on the granitic terrain may be a consequence of a relatively high base content of the ground water and the deeper soils, placing them outside the stability fields of rhododendrons and Mountain Laurel. We do gain further insight into the stability relations of Spotted Wintergreen and Downy Rattlesnake Plantain. As previously surmised, the upper pH limit of the former appears to lie a little below 6.0 in most environments in which it has been observed, perhaps a little below that of Red Maple. However here this species also appears stable under moist mesic conditions, in which it has not been previously seen by this reporter. Additionally, our observations here apparently extend the upper pH stability limit of Downy Rattlesnake Plantain to only a little below 6.0 as well also in contradiction of this reporter's previous view of the matter.
Considering now the bryophytes of the Granitic Substrate, we note a contrast with those of the Calcareous Substrate of the Oak-Hickory Ridge Forest of the Folly Mills Watershed. If C represents the number of taxa in common with our standard of comparison, the Mesic Slope Forest of the Folly Mills Watershed, and T the total number of taxa in the Granitic Substrate bryophyte assemblage, the index of similarity CxC / T is 4.5, while that of the Calcareous Substrate is 4.1 (see our section on the Folly Mills Watershed).
A word also needs to be said about the herbal and medicinal properties of Goldenseal. As pointed out earlier, this plant is strongly shunned by all herbivores in the occurrences observed by us. To this reporter there is a message here regarding its suitability for internal human consumption. This caution is also supported by Russell et al (1997) who state that it "causes only low toxicity if eaten" but "all parts" are regarded as poisonous. A toxic principle of the plant includes Isoquinolin alkaloids, and these authors discourage its use for self medication. Of course, this in no way invalidates the potential of Goldenseal for medical use. However, it appears that this use has not as yet been adequately explored in a scientific way. Additional references are listed at the site of natureserve at http://www.natureserve.org/
Acknowledgements
We appreciate the many contributions of Dorothy Simkins to our observations of the flora and fauna, particularly with respect to birds and the fungi, which so readily escape our efforts. We also thank Gary Fleming for calling our attention to the Black Ash and Bog Bluegrass in one of the areas inventoried.
Note
(1) The terms "mor" and "mull", dating to Hundeshagen in 1830 ( Armson, 1979), have been found by this reporter to provide the clearest and most convenient field descriptions of forest soils.
References
Armson, K. A. (1979) Forest Soils: properties and processes. University of Toronto Press, Toronto.
Fleming, Gary P. (1999) Plant Communities of Limestone, Dolomite, and Other Calcareous Substrates in the George Washington and Jefferson National Forests, Virginia. Virginia Natural Heritage Technical Report 99-4. Virginia Dept. of Conservation and Recreation, Div. Of Nat. Heritage, Richmond, Va.
Heritage Technical Report 99-4. Virginia Dept. of Conservation and Recreation, Div. Of Nat. Heritage, Richmond, Va.
Hunsucker, Robert and R. F. Mueller (1998) Folly Mills Calcareous Wetland, Augusta County, Virginia. Forests of the Central Appalachians Project, Virginians for Wilderness Web Page.
Mueller, R. F. (2000) Stability Relations in Forests. Forests of the Central Appalachians Project, Virginians for Wilderness Web Page.
Perry, Thomas O. (1982) The Ecology of Tree Roots and the Practical Significance Thereof,Journal of Arboriculture, 8 (8), 197 211.
Phillips, Roger (1991) Mushrooms of North America. Little Brown and Co., Boston.
Rader, Eugene K. (1967) Geology of the Staunton, Churchville, Greenville and Stuarts Draft Quadrangles, Report of Investigations 12. Virginia Div. Of Mineral Resources, Charlottesville, Va.
Russell, Alice B., James W. Hardin, Larry Grand, and Angela Fraser (1997) Poisonous Plants of North Carolina. North Carolina State University Web Site.
Strausbaugh, P. D. and Earl L. Core (1977) Flora of West Virginia. second ed., Seneca Books Inc. Grantsville, West Va.