The Vascular Flora of the Francis Beidler Forest
in Four Holes Swamp, Berkeley and Dorchester Counties, South Carolina
RICHARD D. PORCHER
Reprinted from Castanea 46:248-280. 1981.

• ABSTRACT
• INTRODUCTION
• DESCRIPTION OF PLANT COMMUNITIES
• ANNOTATED CHECKLIST OF THE VASCULAR FLORA
  OF BEIDLER FOREST AND ENVIRONS
• ACKNOWLEDGMENTS
• LITERATURE CITED

• Bio on Richard D. Porcher, Jr.

ABSTRACT

A survey of the vascular plants of The Francis Beidler Forest, A National Audubon Sanctuary in Four Holes Swamp in the Coastal Plain of South Carolina, is presented. Included are a description of major plant communities including an original growth bald cypress-tupelo gum swamp forest community, an annotated checklist of the vascular plants that includes ecological notes on rare and endangered species, and an historical account of Four Holes Swamp and Beidler Forest. Three hundred twelve species from 218 genera and 99 families are reported.

INTRODUCTION

The Francis Beidler Forest is a National Audubon Sanctuary in Four Holes Swamp, located in the Coastal Plain of South Carolina in Berkeley and Dorchester counties. Within the Sanctuary is the largest stand of original growth bald cypress (Taxodium distichum) and tupelo gum (Nyssa aquatica) in the United States (Brunswig and Winton, 1978). Four Holes Swamp is a low gradient, brown water, swamp-river floodplain system that runs for approximately sixty miles through the Coastal Plain of South Carolina. It averages approximately one to one and one-half miles in width along most of its length. From its origin near Cameron in Calhoun County, Four Holes Swamp runs in a southeast direction through Calhoun, Orangeburg, Dorchester, and Berkeley counties to a point along the Berkeley-Dorchester county line. Here the swamp makes a right turn and runs southwest for nine miles until it empties into the Edisto River near Ghivans in Dorchester County. Through the Edisto River the waters of Four Holes Swamp flow into the Atlantic Ocean. A curious feature of the swamp is the sudden right turn it makes. Instead of continuing its course southeasterly to the sea, it turns southwestward and runs parallel to the coast until it joins the Edisto River. One geological explanation (Carr, 1971) is that at a time when the seas were higher and the lower part of Four Holes was an estuarine arm of the pre-Pleistocene Atlantic, the right turn was probably formed by currents that built an off-shore bar. This bar then deflected the lower reach of the then Four Holes River into its present southwesterly course.

Lumbering since the turn of the century has removed the original growth timber along the entire run of Four Holes Swamp except for an 1800-acre section immediately above where the swamp makes its right turn. This section of untimbered original growth forest was owned by Francis Beidler I, a lumberman from Chicago, who died in 1924. With the prospect of the sale of the Beidler property due to liquidation of the estate, local conservationists' action to preserve the swamp led to action by the Nature Conservancy and National Audubon Society. In 1970 these two organizations arranged for the purchase of the 1800-acre section of original growth timber plus 1700 additional acres of adjacent timbered swamp to act as a buffer zone. Title to the properties was originally vested in the Nature Conservancy; however, under an agreement between the two organizations, a 60% interest was deeded to the National Audubon Society. Under the same agreement the Audubon Society became administrator of the properties, and maintains a Visitor's Center and a 6,500-foot boardwalk to allow public access to the major communities of the swamp. The public is not allowed access to the Sanctuary except via the boardwalk and managed canoe trips, in order to insure preservation of the swamp ecosystem. Field trips for educational purposes are allowed by appointment and scientific studies are permitted under the auspicies of the Sanctuary Manager. The 3500-acre tract has been named The Francis Beidler Forest in honor of Francis Beidler I.

The origin of the name "Four Holes Swamp'' is uncertain. It appears on pre-revolutionary maps where the only road through the swamp is near Highway 78. Local history has it that the name comes from four "holes" through the swamp forming seasonally dry passageways used by early pioneers. But the exact origin of the name remains a mystery and will remain so unless new documents are uncovered.

This survey of the vascular plants of Beidler Forest, supported by a grant from the National Audubon Society, is confined to the 1800-acre original growth tract, a cutover upland area where the Visitor's Center is located, portions of the bluff slope that border the eastern side of the swamp, and the bluff slope along the western side of the swamp above Mallard's Lake. Although the bluff slope is not part of the Four Holes Swamp floodplain, its ecological relationship with the swamp system and high aesthetic appeal warrants its inclusion in this study. Actually only a short distance of the bluff slope on the eastern side of the swamp is part of Beidler Forest. Future acquisitions to Beidler Forest, however, include the possibility of acquiring as much of the bluff slope as possible, both to act as a buffer zone to the fragile swamp floodplain and because of the rich and diverse mixed mesophytic hardwood forest on the bluff slope. For these reasons I included in the study the portions of the bluff slope that appear to be likely candidates for future acquisition.

This survey of Beidler Forest is primarily intended to be a base-line study on which future studies of swamp ecology can be conducted. Two factors of Beidler Forest make for an unique opportunity to the scientific community to study swamp ecology, especially studies of plant succession: (1) the presence of a large stand of original growth forest, which has been virtually untouched by man's activities, and (2) areas of cutover forest within the boundaries of the Forest which can be managed (managed in this concept being more to insure that man's activities in the cutover area are excluded or controlled) to let natural succession occur. In addition to the cutover areas within Beidler Forest there are numerous adjacent tracts of swamp forest, under various owners (Figure 2), that have been selectively timbered since the early 1930's. Records as to the dates and type of timber harvest are available for these adjacent tracts, thus considerable data already exists for studies on plant succession following disturbances. These tracts are possible future acquisitions to Beidler Forest, and if acquired, would expedite future studies. The untimbered section of Beidler Forest has been referred to in this study as "original growth" forest rather than a "virgin" forest. This latter designation is often interpreted as meaning that no vegetation, especially trees, has been removed by man's activities. Such is not the case in Beidler Forest. Certainly in colonial times and later, an occasional large cypress was removed to make shingles for roofs, or an occasional hardwood tree was removed for firewood or to make furniture. A few trees were probably felled to make trails or primitive roads into the swamp, and certainly an occasional still, deep in the swamp, necessitated the clearing of small trees. But these activities did not significantly alter the swamp ecosystem. Today the original growth section of Beidler Forest represents a mature stand of timber, virtually the same as it was when the first settlers moved inland from the coastal settlements and began the transformation of our native woodlands and swamps to fields and secondary growth forests.

In recognition of the national significance of the original growth forest described in this study, and the efforts that have been made to preserve it, The Francis Beidler Forest was designated a Registered National Natural Landmark in May of 1979 by the Department of Interior.

DESCRIPTION OF PLANT COMMUNITIES

The primary influence on the distribution and composition of communities in the Four Holes Swamp floodplain is the hydroperiod. Four Holes Swamp is a low gradient, creek or river swamp. From its headwaters near Cameron in Orangeburg County to a point below S. C. Highway 78, there is no discernible major channel. Instead the brown water flows slowly, slowly, through a series of small runs and streams that crisscross the floodplain and join and separate in a pattern that seems to have no purpose or identity. This pattern changes during heavy rains, however, as runoff from the adjacent uplands enters the floodplain, flooding it from upland to upland. Then the water moves swiftly in its relentless path to the sea. Only during severe droughts does the flow of water cease in Four Holes Swamp. During the summer of 1977 one could traverse the flood-plain without having to wade through any flowing water (although the lakes in the swamp retained water). Only where the swamp enters the Edisto River does a major channel develop.

Throughout the Four Holes Swamp floodplain the smaller runs and streams become deeper and wider where two or three merge forming numerous lakes of various sizes. It is along these runs, streams, and lakes that many of the cypress trees reach their largest size, and knees and buttresses obtain maximum development. Four such lakes occur in the original growth section of Beidler Forest: Goodson, Canoe, Singletary, and Mocossin. The largest of the lakes in Beidler Forest, Mallard's, occurs in the timbered section; however, a buffer zone of original growth was left along its northwest side. Another large lake, Mims Lake, occurs along the extreme southeast section of the cutover portion of Beidler Forest. Considerable marl outcrops occur along the edge of the bluff where it borders the lake on its southeast side.

Between the runs, streams, and lakes, eons of alluvial deposition on the floodplain and stream dissection have resulted in a gradual elevation of the soil resulting in three major hydroperiods: (1) flooded to extremely wet most of the year, (2) flooded for a few months from late winter to early spring, and (3) free from standing water most of the year, but flooded during heavy rains, and then draining very quickly. Under these conditions three climax bottomland forests, as described by Braun (1950), have developed in Four Holes Swamp: swamp forest, hardwood bottom, and ridge bottom. In light of recent studies by Dr. A. E. Radford, and his staff at UNC on natural diversity in our environment, the recognition of only three community types appears overly simplified. Even Braun (1950) recognized the simplification of the description of plant communities that prevailed during her time as she stated: "Most data on the composition of bottomland forests do not separate the communities of the several bottomland habitats. All hardwoods are considered together, thus developmental sequences and correlations with site factors are obscured." Dr. Radford and his students have stated the problem most eloquently (Radford et al., 1978): "We can no longer speak in terms of simply an American beech (Fagus grandifolia) forest, but must observe carefully, see relationships, and provide lists such as the following which truly demonstrate natural diversity: Fagus grandifolia/Kalmia latifolia, Fagus grandifolia /Aster Divaricatus, Fagus grandifolia / Mixed herbs, etc." It is obvious when reference is made to studies on swamp ecology that very little critical observation has been made of the distinctive communities that occur as the result of very slightly varying conditions such as elevation. Gaddy et al. (1976) did recognize eleven communities in their study on Congaree Swamp in South Carolina; however, they did not correlate the communities and the environmental parameters determining their distribution. This study of Beidler Forest followed a classical approach in describing the plant communities because time and financial considerations did not permit quantitative sampling. But as stated previously this study is intended as a base-line study on which to base future studies on swamp ecology in Beidler Forest. Plans are being formulated for a study of the original growth section of Beidler Forest in which correlations between plant communities and environmental parameters such as hydroperiod, soil, elevation, geology, etc., will be analyzed.

This approach should expand the list of distinct plant communities in Beidler Forest and determine the environmental parameters that determine their composition and distribution. Since this study will be made on an original growth bottomland forest system, an opportunity not available elsewhere, a significant contribution to the understanding of swamp ecology should result.

The following description of plant communities includes the three bottomland forest communities that occur on the flood-plain, a mixed mesophytic hardwood forest community on the bluff slope, and the seepage bogs at the base of the bluff slopes. As with the case with the hardwood bottom communities, the recognization of only one community on the bluff slope is simplified in light of the forementioned recent studies on natural diversity. This community will also be included in a future study of Beidler Forest.

1. Mixed Mesophytic Hardwood Forest. This community occurs on the bluff slope that flanks the eastern side of Four Holes Swamp and for a short distance along the southwest side above the northern end of Mallard's Lake. This community corresponds to Braun's (1950) mesophytic mixed hardwood community and Hunt's (1947) moist rich woods.

Along the eastern side of Four Holes Swamp, the bluff slope separates the floodplain from the adjacent uplands. In general the aspect of the slope is southwest; however, considerable local variation in aspect occurs along the entire slope due to numerous ravines dissecting the slope and acting as pathways for streams and branches that feed water into the swamp from the adjacent uplands. The bluff is approximately 25-30 feet above the flood-plain with a slope of 25-30 degrees along most of its length. A much steeper slope occurs along Mims Lake. The only comparable bluff slope on the southwest of the swamp is an area above the northern end of Mallard Lake's. Elsewhere along the southwest side of the swamp the uplands gently slope to the floodplain and no distinct mixed mesophytic hardwood forest occurs.

Only a small section of the bluff slope along the eastern side of the swamp is within the present boundary of Beidler Forest. Additional areas of the bluff slope were sampled, however, since future plans of the Audubon Society include acquisition of as much of this bluff slope as possible. During sampling of the bluff slope, it was noted that the majority of species found on the bluff slope outside of the Beidler Forest boundary were also found on the bluff slope within Beidler Forest.

It is at once evident that this mixed mesophytic hardwood community is more mesic than the oak-hickory, pine, or pine mixed-hardwood of the adjacent uplands, yet distinct from that of the alluvial bottomland of the floodplain. Canopy species include large specimens of Fagus grandifolia, Quercus michauxii, Q. falcata, &. alba, Q. nigra, &. schmardii, Liquidambar styraciflua, Liriodendron tulipifera, Juglans nigra, Acer rubrum, Nyssa syluatica, Ulmus americana Carya tomentosa, and C. glabra. An occasional loblolly pine (Pinus taeda) and spruce pine (P. glabra) are present. On the less mesic sites, oaks and hickories are more dominant, while on the more mesic sites Fagus grandifolia becomes more abundant. The bluff slope along Mims Lake actually fits the description of Braun's (1950) ravine slope forest because beech (Fagus grandifolia) comprises more than one-third of the canopy (based on field observation).

A distinct subcanopy is present, consisting of canopy species transgressives and the following common mesophytic subcanopy species: Cercis canadensis, Cornus florida, Carpinus caroliniana, Ilex opaca, and Asimina triloba.

The herbaceous species vary from the more shade tolerant species to the grasses, sedges, and legumes that became more abundant where the canopy becomes thinner. The herbaceous species growing under the dense canopy are particularly noteworthy since they include numerous of the more attractive wildflowers such as: Trillium cuneatum, Sanguinaria canadensis, Erythrina herbacea, Spigelia marilandica, Podophyllum peltatum, and six species of orchids, Malaxis spicata, M. uniflora, Tipularia discolor, Corallorhiza wisteriana, Hexalectris spicata, and Ponthieva racemosa.

The large size and old age of the canopy trees and the presence of canopy species transgressives indicate that the community is a mature, climax community, and will maintain itself barring any major disturbance. The absence of such introduced species as Lonicera japonica indicates that the community has been little disturbed in past times. The random mixture and composition of tree species throughout the community is probably due in part to the following factors: (1) its close proximity to a large wetland area and upland area whose hydrophytic (Ex.: Quercus nigra) and xerophytic (Ex.: Carya tomentosa) species respectively are present as waifs or carry-overs in a habitat much drier or wetter than typical for the species; (2) the localized variation in aspect and slope of the bluff, and (3) selective removal of species in past times. How much of the community represents original growth forest is not known, but judging from the size of many of the largest trees, it is likely that some of the trees are original growth. Many of the largest trees certainly date from colonial times. This community appears typical of the mixed mesophytic hardwood forest that occupies the slopes cut into the Pleistocene terraces of the Coastal Plain of S.C. (personal observation).

2. Seepage Bog. The upland area adjacent to Four Holes Swamp consists of sandy soil overlying a marl formation. Where the bluff slope intersects the level floodplain of the swamp, the marl is exposed as an outcrop. Groundwater from the upland sandy soil, unable to penetrate the substrata, spreads laterally and emerges as seepage where the substrata intersect the slope. This creates a bog-like condition which even in dry periods contains moisture since the flow of water is continuous. Even during the summer of 1977 when water ceased to flow in the swamp, the seepage bogs did not dry up. The water from the seepage undoubtedly tends to contribute to the water supply; however, this supply is minimal compared to the main water source of the swamp.

Three species of emergent herbs appear to be confined to the seepage bogs and have not been found elsewhere: Gratiola virginiana (hedge hyssop), Micranthemum unbrosum, and Cardamine pennsylvanica. These species tend to occur at the headwaters of the seepage where the water tends to puddle. Additional emergent herbs occurring in the bog include Saururus cernuus, Hydrocotyle verticillata, Boehmeria cylindrica, Pilea fontana, Impatiens capensis, Bacopa monnieri, and Senecio glabellus. These latter species are not confined to the seepage bog, but occur in the swamp forest wherever moisture conditions are sufficient. They do tend to be more concentrated in the bog, however, because of the permanent water supply.

In late summer and in the fall, many of the seepage bogs tend to be overgrown by Pilea fontana and Impatiens capensis. Numerous species of the adjacent swamp forest and mixed mesophytic hardwood forest tend to be localized around these seepage bogs.

Although I have not made a quantitative determination, I would estimate that there is in excess of a hundred seepage bogs scattered along the base of the bluff slope on the eastern side of Four Holes Swamp between the extremes of the Beidler Forest boundary (approximately six miles). The occurrence of seepage bogs along the remaining run of Four Holes Swamp has not been documented, but undoubtedly they are not confined to the bluff slope adjacent to Beidler Forest.

The literature on seepage bogs in the Coastal Plain is sparse. Hunt (1947) described seepage slopes in his description of the plant habitats in the Charleston area. Hunt's seepage slopes correspond to the seepage bogs described in this study. Hunt listed only the woody flora around the seepage slopes and stated that Alnus serrulata and Rhus vernix characterized the seepage slopes. In this study only A. serrulata was found, and that around only one seepage bog. A significant contribution could be made to the botany of the coastal region of S.C. by documentating the occurrence of seepage bogs and describing their vegetation.

3. Swamp Forest. The swamp forest community occupies the constantly flooded (except in extreme dry periods) alluvial soil on the lowest level of the Four Holes Swamp floodplain. In the Beidler Forest portion of Four Holes Swamp, the swamp forest community is original growth timber since no commercial lumbering has occurred in Beidler Forest. This community corresponds to Wells' (1942) "swamp forest," Hunt's (1947) "cypress swamps," Penfound's (1952) "deep freshwater southern cypress-tupelo gum swamp," and Braun's (1950) "bald cypress-tupelo gum swamp forest."

The dominant trees of the swamp forest are Taxodium distichum (bald cypress) and Nyssa aquatica (tupelo gum). Bald cypress forms the canopy, with the tallest trees around 120 feet, while tupelo gum comprises a sub-canopy approximately 80 feet tall. The largest cypress trees are around five to six feet in diameter and have been estimated to be 700 to 800 years old (Dennis, 1970). In the deeper water at the lowest elevation bald cypress and tupelo gum occur together in almost pure strands. Elsewhere in the swamp at slightly higher elevations, a variety of tree species forms a conspicuous growth. Most common is Fraxinus caroliniana, a small tree that in some areas forms a subcanopy below the Nyssa aquatica, but in most areas is not dense enough to be considered a separate subcanopy. F. caroliniana occurs in slightly raised sites and generally grows from the bases of cypress and gum trees. Nyssa sylvatica var. biflora occurs scattered throughout, occasionally reaching canopy size. Two additional hydric species, Gleditsia aquatica and Planera aquatica, occur as small trees scattered throughout the swamp, either growing from the bases of the cypress or tupelo gum or rooted in the soil. Four species characteristic of the hardwood bottom occur throughout the community rooted on fallen logs, large cypress knees, and buttresses: Quercus laurifolia, Acer rubrum, Liquidambar styraciflua,and Ulmus americana. These four species occur as small trees since they are unable to obtain large size due to the uncertain foundation on which they grow. Bald cypress exhibits typical cone buttresses and knees, and tupelo gum exhibits typical buttress formation.

A well-developed shrub layer occurs on the higher sites with species rooted in the soil or growing on buttresses, knees, fallen logs, or stumps. Species include Itea virginica, Styrax americana Leucothoe racemosa, Lyonia lucida, Cornus stricta, Cephalanthus occidentalis, Viburnum dentatum var. dentatum, and Ilex decidua. These shrubs, along with lianas and transgressives of canopy and subcanopy species, form a dense growth in most areas of this habitat. In the lower sites of the swamp forest where standing water occurs most of the year, a sparse shrub layer occurs consisting of I. virginica, S. americana, C. occidentalis, L. racemosa, and L. lucida growing from the buttresses of Taxodium and Nyssa.

The herbaceous flora of the swamp forest is exceptionally rich and varied and is worth noting. In the deeper areas of the swamp, particularly along the edges of the lakes and runs, the herbaceous flora is confined to floating logs, stumps, knees, and buttresses. This habitat is given community status by some authors (Dennis and Batson, 1974); however, I prefer to consider it a microhabitat, or dry phase, of the swamp forest. Three species appear to characterize this habitat as they do not occur elsewhere in the swamp (or in the hardwood bottom and ridge bottom); Scutellaria lateriflora, Lycopus rubellus, and Hypericum virginicum. Additional species that occur in this microhabitat, as well as in the higher areas of the swamp forest and adjacent communities, include Woodwardia aerolata, Boehmeria cylindrica, Senecio glabellus, Onoclea sensibilis, Lobelia cardinalis, Diodia virginiana, and Hypericum densiflorum. Two weedy species of the upland area, Phytolacca americana and Eupatorium compositifolium, occur as rare components of this microhabitat, an adaptation that allows these primarily terrestrial upland species to grow in an aquatic habitat by being situated above the high water level.

In the higher sites in the swamp forest where the surface water is shallow or the soil extremely wet, or where isolated, low raised areas occur, the following herbs are found: Dracocephalum purpureum, Sanolus parviflorus, Asclepias perennis, Orontium aquaticun, Peltandra virginica, Scirpus divaricatus, S. fontinalis, Proserpinaca palustris, and Lobelia cardinalis. Also found in this habitat are the species of the fallen log, stump, knee, and buttress microhabitat discussed above (with the exception of S. lateriflora, L. rubellus, and H. virginicum).

An interesting feature of the swamp forest is the presence of an occasional sapling of Pinus glabra, spruce pine, growing on the buttresses and knees of Taxodium. No large specimens were observed as they are undoubtedly unable to reach large size because of the uncertain foundation. Spruce pine is characteristic of the nearby ridge bottom community which probably acts as the seed source. Although spruce pine is frequent in the coastal counties of South Carolina north to Georgetown County (Radford et al., 1968), I know of no reference documenting its occurrence in swamp forest as described above. It is interesting to note that the more common loblolly pine, P. taeda, was not observed under similar circumstances.

That this is a mature, climax community, maintaining itself, is apparent by the following observations: (1) a considerable size variation in the dominant trees, bald cypress and tupelo gum, ranging from seedlings to large trees, and (2) the presence of numerous fire-scarred trunks of living trees and charred remains of stumps, indicating periods when the swamp was dry.

4. Hardwood Bottom. As the elevation of the floodplain increases, the swamp forest grades into a hardwood bottom community. This community corresponds to Braun's (1950) glade or hardwood bottom which she states is most common along the floodplains of the larger rivers of the southeast such as the Congaree, Wateree, Santee, Savannah, etc.

The hardwood bottoms generally occur as islands scattered throughout the swamp floodplain wherever the elevation is raised above the level of the swamp forest. The hardwood bottom is only occasionally flooded, generally for a few months during the late winter and early spring when the runoff into the swamp is too great to be carried away by the swamp drainage. However, the bottom can be flooded any time of the year when rains are unusually heavy. The soil is alluvial and the water table is probably never more than six inches below the surface. The surface is relatively flat except where earthen mounds have been produced by the root systems of large trees that have been blown over.

The canopy consists of a mixture of deciduous hardwoods with oaks predominating and an occasional cypress carry-over from the adjacent swamp forest. Knee development of cypress is either lacking or very reduced, and both the cypress and tupelo gums lack the cone buttresses characteristic of the swamp forest. Canopy tree species include: Carya aquatica, Quercus lyrata, Q. nigra, Q. phellos, Q. laurifolia, Acer rubrum, Liquidambar styraciflua, Ulmus americana, Nyssa aquatica, Nyssa sylvatica var. biflora, Fraxinus tomentosa, and Celtis laevigata. Transgressives of the canopy species are present indicating the community is maintaining itself. The stands of hardwood bottom in Beidler Forest, like the swamp forest, represent original growth timber, although undoubtedly in past times a few trees were probably removed for fuel, furniture, etc.

A poorly developed subcanopy consists of Carpinus caroliniana, Persea borbonia, Ilex opaca, and Crataegus marshallii. A shrub layer is poorly developed except for dense stands of Sabal minor at higher elevations. Herbaceous plants and lianas are sparse.

5. Ridge Bottom. At a slightly higher elevation, the hardwood bottom grades into a third bottomland forest, the ridge bottom. The ridge bottoms are free of standing water most of the year, being flooded only during the heaviest rains, and then drain very quickly when the water subsides in the floodplain. This community corresponds to Braun's (1950) ridge bottom. The distribution and abundance of this community in the South Carolina Coastal Plain is not documented in the literature, as little work on swamp ecology has been done in this area. Hunt (1947) does not mention ridge bottoms in his descriptions of plant communities in the Charleston area. Based on available literature and the author's observations of the river systems of the Coastal Plain, ridge bottoms as described by Braun (1950) and as occur in the Beidler Forest section of Four Holes Swamp, may be very limited.

Throughout the swamp floodplain, the ridge bottoms occur as islands surrounded by hardwood bottom and swamp forest. Occasionally where the gradient is somewhat steeper, a zone of hardwood bottom is absent, and the ridge bottoms border the swamp forest. Six tree species delimit the ridge from the adjacent hardwood bottom: Quercus michauxii, Q. alba, Pinus glabra, P. taeda, Carya glabra, Cornus florida, and occasionally Fagus grancliflora on the higher ridge bottoms. Most of the species from the wetter hardwood bottom complete the canopy of very large trees: Liquidambar styraciflua, Acer rubrum, Quercus laurifolia, Q. nigra and Ulmus americana. Pinus taeda, loblolly pine, occurs scattered throughout the ridge bottom, becoming locally abundant in the ridge bottom adjacent to the boardwalk. It undoubtedly became established within the hardwood forest when the canopy was opened due to the death and falling of a large canopy tree. Many of the loblollys are extremely large. The dominant trees of the swamp forest, bald cypress and tupelo gum, are absent from the ridge bottom.

The subcanopy consists of Ilex opaca, Cornus florida, Carpinus caroliniana, Diospyros virginiana, Crataegus marshallii, Morus rubra, and canopy transgressives. Unlike the adjacent hardwood bottom, the shrub layer is highly developed and represented by the following species: Ilex decidua, Aesculus pavia, Vaccinium elliottii, Sabal minor, Myrica cerifera, Rhododendron canescens, and Hamamelis virginiana. A well-developed herbaceous layer consists of a variety of carices, panic grasses, and beak rushes plus the ephemeral wildflowers Podophyllum peltatum, Zephyranthes atamasco, Trillium cuneatum, Arisaema triphyllum, Listera australis, Sanguinaria canadensis, Clematis crispa, and Trillium pusillum var. pusillum. This latter species is botanically significant since it is listed as a candidate for endangered status by the Department of Interior (1975). One population was found in the ridge bottom that lies adjacent to the boardwalk and consisted of approximately a hundred stems in a half-acre area. The rarity of this species in the Coastal Plain is reported by Radford et al. (1968) and by Percher (1979). No other populations were found in Beidler Forest although the brief flowering period makes it likely that it could easily have been overlooked.

ANNOTATED CHECKLIST OF THE VASCULAR FLORA
OF BEIDLER FOREST AND ENVIRONS

Three hundred and twelve species of vascular plants, representing two hundred and eighteen genera and ninety-nine families, were collected in Beidler Forest and environs during this study. A summary of the flora is given at the end of this checklist. This checklist includes species found in the disturbed woods around the Visitor's Center. Plant collections were made primarily from March of 1976 through August of 1978, with a few made during the summer of 1979. One set of voucher specimens has been deposited in The Citadel Herbarium, Charleston, S.C. A second set is being maintained at The Citadel Herbarium, but will be ultimately maintained at the Visitor's Information Center at Beidler Forest when facilities are available. Nomenclature and sequence of families follows that of Radford et al. (1968). The habitat and frequency were determined by subjective field observations. Frequency of species is indicated by the relative terms: rare, occasional, locally abundant, and common. Frequency determinations are omitted from some species, such as Carer alata and Panicum dichotomiflorum, because of the difficulty in recognizing the species in the field.

For brevity the following numbers are used to represent the five communities: 1=mixed mesophytic hardwood forest, 2=seepage bog, 3=swamp forest, 4=hardwood bottom, and 5=ridge bottom.

PTERIDOPHYTA

Selaginellaceae

Selaginella apoda (L.) Spring. Meadow Spikemoss. Essentially throughout in moist places, but often overlooked.

Ophioglossaceae

Botrychium biternatum (Sav.) Underwood. Southern Grapefern. Occasional in 1 and 5.

Botrychium virginianum (L.) Swartz. Rattlesnake Fern. Occasional in 1 and 5.

Osmundaceae

Osmunda cinnamomea L. Cinnamon Fern. Common and scattered throughout 1, 3 and 5.

Osmunda regalis var. spectabilis (Willd.) Gray. Royal Fern. Occasional in 3 and 5.

Pteridaceae

Pteridium aquilinum (L.) Kuhn. Bracken Fern. Common in 1 and 5.

Aspidiaceae

Athyrium asplenioides (Michaux) A. A. Eaten. Southern Lady Fern. Common in 1 at lower slope; often locally abundant around 2.

Dryopteris ludouiciana (Kunze) Small. Southern Shield Fern. Locally abundant around 2 and in 1; found only on east side of swamp north of power line.

Onoclea sensibilis L. Sensitive Fern. Common throughout forested communities as terrestrial herb, often around edges of 2; common in 3 on stumps, fallen logs, cypress knees and buttresses.

Polystichum acrostichoides (Michaux) Schoot. Christmas Fern. Common in 1 and 5.

Thelypteris kunthii (Desv.) Morton. 1.

T. hexagonoptera (Michaux) Weatherby. Broad Beechfern. Common along base of slope in 1, primarily along eastern side of swamp.

Blechnaceae

Woodwardia areolata (L.) Moore. Netted Chain-fern. Common throughout all communities; often growing on fallen logs, stumps, knees, and buttresses in 3.

Woodwardia virginica (L.) Smith. Virginia Chain-fern. Occasional throughout.

Aspleniaceae

Asplenium platyneuron (L.) Oaks. Ebony Spleenwort. Common throughout all communities and often on fallen logs, stumps, knees, and buttresses in 3.

Polypodiaceae

Polypodium polypodioides (L.) Watt. Resurrection Fern. Common throughout as epiphyte on large trees and often in association with the epiphytic orchid Epidendrum conopseum.

SPERMATOPHYTA-GYMNOSPERMAE

Pinaceae

Pinus glabra Waiter. Spruce Pine. Scattered, but locally abundant in 5; rare in 1; rare in 3 growing as small trees on cypress knees and buttresses.

Pinus taeda L. Loblolly Pine. Common in 5; occasional in 1.

Taxodiaceae

Taxodiun distichum (L.) Richard. Bald Cypress. Common in 3; occasional in 4; largest trees estimated to be 800 years old.

Cupressaceae

Juniperus virginiana L. Red Cedar. Common in 1 and 5.

ANGIOSPERMAE-MONOCOTYLEDONEAE

Sparganiaceae

Sparganium americanum Nuttall. Bur-weed. Rare; edge of 3.

Alismataceae

Sagittaria sp. Arrowhead. Locally abundant in 3.

Poaceae

Arundinaria gigantea (Waiter) Muhl. Cane. Common in 1, 4 and 5.

Cinna arundinaceae L. Wood Reed. Common in 1 and 5.

Festuca obtusa Biehler. Fescue. Locally abundant in 1 on east side of swamp north of power line and on Neely Smith tract of Beidler Forest.

Melica mutica Waiter. Melic Grass. Common in 1 and 5.

Oplismenus setarius (Lam.) R. & S. Common in 1, 4 and 5.

Panicum boscii Poiret. Love Grass. Common in 1 and 5.

Panicum commutatum Schultes. Love Grass. Common in 1.

Panicum dichotomiflorum Michaux. Love Grass. Found along edge of Mims Lake.

Panicum laniflorum Lam. Love Grass. Common in 1 and 5.

Panicum virgatum L. Love Grass. 1.

Paspalum setaceum Michaux. Common in 1.

Trisetum pensylvanicum (L.) Beauvois ex R. & S. Edge of 2.

Uniola laxa (L.) BSP. Common in 1 and 5.

Cyperaceae

Carex alata Torrey. Edge of Canoe Lake in 3.

Carex bromoides Schkuhr. 3.

Carex debilis Michaux var. debilis. Common in 1.

Carex elliottii Schweintz & Torrey. 1.

Carex howei Mackenzie. Edge of 2.

Carex gigantea Rudge. 3.

Carex intumescens Rudge. 3.

Carex oxylepis Torrey & Hooker. 3.

Cyperus sesquiflorus (Torrey) Mattfeld & Kukenthal. Abandoned road in low area around Visitor's Center.

Dulichium arundinaceum (L.) Britton. Scattered and locally abundant in 3.

Eleocharis tortilis (Link) Schultes. Spike-rush. 2.

Rhynchospora caduca Ell. Beak Rush. 5.

Rhynchospora chalarocephala Fernald & Gale. Beak Rush. Edge of 2.

Rhynchospora corniculata (Lam.) Gray. Beak Rush. Occasional on edge of lakes in 3.

Rhynchospora miliacea (Lam.) Gray. Beak Rush. Common as ground cover in 5.

Rhynchospora mixta Britton ex Small. Beak Rush. 3.

Scirpus cyperinus (L.) Kunth. Bulrush. Rare on edge of 2.

Scirpus diuaricatus Ell. Bulrush. Common in 3.

Scirpus fontinalis Harper. Bulrush. Common in 3.

Scleria triglomerata Michaux. Nut Rush. Occasional in 1.

Arecaceae

Sabal minor (Jacquin) Persoon. Dwarf Palmetto. Common and often forming dense thickets in higher sites in 4 and lower sites in 5.

Araceae

Orontium aquaticum L. Golden Club. Occasional and widely scattered in 3.

Arisaema triphyllum (L.) Schoot. Jack-in-the-Pulpit. Common in 5 and lower slope of 1.

Peltandra virginica (L.) Kunth. Occasional in 3 around edge of lakes.

Xyridaceae

Xyris jupicai Richard. Yellow-eyed-grass, Collected once along abandoned road in low area around Visitor's Center.

Bromeliaceae

Tillandsia usneoides L. Spanish Moss. Common as epiphyte in 1, 3, 4 and 5.

Commelinaceae

Commelina comnunis L. Dayflower. Edge of Mims Lake in 3.

Tradescantia rosea Vent. var. rosea. Spiderwort. Common in 1 and 5.

Juncaceae

Juncus coriaceus Mackenzie. Common in 1 and along edge of 2.

Juncus megacephalus M.A. Curtis. 3.

Juncus repens Michaux. Occasional in wet depressions in 5.

Luzula echinata (Small) Hermann. Woodrush. Rare in 1 along Mims Lake.

Liliaceae

Medeola virginiana L. Indian Cucumber-root. Rare and appears to be restricted to flat, grassy ecotone between 1 and 3.

Smilacina racemosa (L.) Desf. False Solomoris Seal. Rare in 1.

Smilax bone-nox L. Common throughout all forested communities.

S. ecirrhata var. biltmoreana (Small) Ahles. Carrion-flower. Rare; collected in 1 along Mims Lake.

S. ecirrhata var. hugeri (Small) Ahles. Carrion-flower. Rare; collected in 1 on bluff east of Mims Lake.

S. glauca Waiter. Greenbrier. Common and scattered throughout all forested habitats.

S. hispida Muhl. Greenbrier. 3.

S. laurifolia L. Bamboo. Rare; scattered throughout 3.

S. pumila Waiter. Greenbrier. Common in 1, 5, and disturbed woods around Visitor's Center.

S. rotundifolia L. Greenbrier. Scattered throughout all forested habitats.

S. smallii Morong. Greenbrier. Occasional in 1.

S. walteri Pursh. Greenbrier. Common in 3.

Trilliun cuneatum Raf. var. cuneatum. Little Sweet Betsy. Common in 1 and 5.

Trilliun pusillun Michaux var. pusillum. Carolina Trillium. Rare in 5.

Uvularia floridana Chapman. Bellwort. Occasional in flat, grassy ecotone between 1 and 3.

Yucca filamentosa L. Bear-grass. Rare in flat grassy ecotone between 1 and 3.

Dioscoreaceae

Dioscorea villosa L. var. villosa. Wild Yam. Common in 1 and 5.

Amaryllidaceae

Hypoxis micrantha Pollard. Yellow Star-grass. Common in 4 and 5.

Zephyranthes atamasco (L.) Herbert. Atamasco Lily, Easter Lily. Common in 5 and flat, grassy ecotone between 1 and 3.

Iridaceae

Sisyrinchium mucronatum Michaux. Blue-eyed Grass. Common in 5.

Iris sp. Iris. Scattered throughout 3.

Orchidaceae

Corallorhiza wisteriana Conrad. Spring Coral-root. Rare; found in only one site in 1.

Epidendrum conopseun R. Brown. Green-fly Orchid. Common epiphytic in 3 on Taxodium distichum and a variety of hardwood trees.

Goodyera pubescens (Willd.) R. Brown. Downy Rattlesnake Plantain. Observed from two sites in ridge bottom along boardwalk. One population consisted of four specimens and the other of one specimen. G. pubescens was first reported from Dorchester County and the South Carolina Coastal Plain by Percher (1974) and, although common in the mountains, it is considered rare in the Coastal Plain.

Habenaria clavellata (Michaux) Sprengel. Small Green Wood-orchid. Common in drier areas of 3 and in grassy, flat ecotone between 1 and 3.

Hexalectris spicata (Welter) Barnart. Crested Coral-root. Rare; two populations found in 1 on eastern side of swamp, one north of power line and one on Neely Smith tract of Beidler Forest. Listera australis Lindley. Southern Twayblade. Found from one site in 5 along boardwalk; easily overlooked and probably more common.

Malaxis spicata Swartz. Florida Adder's Mouth. Appears to be restricted to the flat, grassy ecotone between 1 and 3. Only a few specimens found but may be more common since it is hard to detect growing in the dense ground cover of the ecotone.

M. unifolia Michaux. Green Adder's Mouth. Common in 1.

Ponthieva racemosa (Waiter) Mohr. Shadow-witch. Occasional in ecotone between I and 3.

Tipularia discolor (Pursh) Nuttall. Crane-fly Orchid. Common in 1 and 5.

ANGIOSPERMAE-DICOTYLEDONEAE

Saururaceae

Saururus cernuus L. Lizard's Tail. Common in drier areas of 2 and 3; occasional in 4.

Salicaceae

Populus heterophylla L. Swamp Cottonwood. Occasional in 3 and 5.

Salix caroliniana Michaux. Swamp Willow. Rare in 3.

Salix nigra Marshall. Black Willow. Rare on edge of 1.

Myricaceae

Myrica cerifera L. Wax Myrtle. Common in all communities.

Juglandaceae

Carya aquatica (Michaux f.) Nuttall. Water Hickory. Common in 3, 4 and 5.

Carya cordiformis (Wang.) K. Koch. Bitternut Hickory. 5.

Carya glabra (Waiter) Sweet. Pignut Hickory. Common in I and 5.

Carya pallida (Ashe) Engler & Graebner. Pale Hickory. Rare; found only in 1 along Mims Lake.

Carya tomentosa (Poiret) Nuttall. Mockernut Hickory. Common in 1 and 5.

Juglans nigra L. Black Walnut. Occasional in less mesic sites in 1.

Betulaceae

Alnus serrulata (Aiton) Willd. Tag Alder. Rare on edge of 1.

Carpinus caroliniana Waiter. Ironwood. Common in all forested habitats except 1 where it is occasional on fallen logs and stumps.

Fagaceae

Castanea pumila (L.) Miller. Chinquapin, Occasional in less mesic sites

Fagus grandifolia Ehrhart. Beech. Locally abundant in more mesic sites in 1 and higher sites in 5.

Quercus alba L. White Oak. Occasional in 5; common in 1.

Q. austrina Small. Bluff Oak. Rare; less mesic sites in 1.

Q. falcata Michaux. var. falcata. Southern Red Oak. Occasional in 1 and 5.

Q. falcata var. pagodaefolia Ell. Cherrybark Oak. Rare in 1.

Q. laurifolia Michaux. Laurel Oak. Common in 4; occasional in 1 and 5.

Q. lyrata Waiter. Overcup Oak. Common in 4; occasional in lower sites in Q. michauxii. Nuttall. Swamp Chestnut Oak. Common in 1 and 5. This species is a good indicator of the demarkation between ridge bottoms and hardwood bottoms since it does not occur in the hardwood bottom.

Q. nigra L. Water Oak. Common in 1, 4 and 5.

Q. shumardii Buckley. Swamp Red Oak. Common in 1; occasional in 5.

Q. virginiana Miller. Live Oak. Only one specimen observed--edge of 1 along Mims Lake.

Ulmaceae

Celtis laeuigata Willd. Hackberry. Occasional in 1, 4 and 5.

Planera aquatica Waiter ex J. F. Gmelin. Water-elm. Common and scattered throughout 3.

Ulmus alata Michaux. Winged Elm. Occasional in 1; rare in 5.

U. americana L. American Elm. Common in 1, 4 and 5; occasional in 3 growing on fallen logs, stumps, knees, and buttresses.

Moraceae

Morus rubra L. Red Mulberry. Common in 1; rare in 5.

Urticaceae

Boehmeria cylindrica (L.) Swartz. False Nettle. Common in 3 on fallen logs, stumps, buttresses, and cypress knees; common in 2.

Pilea fontana (Lunell) Rydberg. Clearweed. Common in 3 on fallen logs, stumps, buttresses, and cypress knees; common in 2 and in drier sites in 3.

Loranthaceae

Phoradendron serotinum (Raf.) M. C. Johnston. Mistletoe. Common in all forested communities.

Aristolochiaceae

Aristolochia serpentaria L. Birthwort. Occasional in 1, 3, 4 and 5.

Hexastylis arifolia (Michaux) Small. Heart Leaf. Common in 1 and 5.

Polygonaceae

Polygonum hydropiperoides Michaux. Common in all communities.

P. setaceum Baldwin ex Ell. 2.

Tovara virginiana (L.) Raf. Jumpseed. Common in 4 and 5, and ecotone between 1 and 3.

Phytolaccaceae

Phytolacca americana L. Pokeweed. Occasional on fallen logs in 3 and common wherever openings are created in all forested communities.

Ranunculaceae

Anemone virginiana L. Thimbleweed. Rare in 1.

Clematis crispa L. Leather-flower. Occasional in 5.

Ranunculus carolinianus DC. Buttercup. 4.

R. pusillus Poiret. Buttercup. Common in grassy ecotone between 1 and 3.

Thalictrum sp. Meadow Rue. Rare; found only in 1 along Mims Lake.

Nymphaeaceae

Nyphar luteum ssp. macrophyllum (Small) E. O. Beal. Spatterdock. Rare; found only in Mallard's Lake.

Berberidaceae

Podophyllum peltatum L. May-apple. Common in more mesic areas of 1.

Menispermaceae

Cocculus carolinus (L.) DC. Coralbeads. Occasional where 1 borders fields.

Magnoliaceae

Liriodendron tulipifera L. Tulip Tree. Common in 1; rare in 5.

Magnolia grandiflora L. Bull Bay. Occasional in 5.

M. virginiana L. Sweet Bay. Occasional in 5.

Annonaceae

Asimina triloba (L.) Dunal. Pawpaw. Found in one dense stand in 1 on Weatherford Bluff.

Lauraceae

Lindera benzoin (L.) Blume. Spicebush. Common in drier areas of 3 and in ecotone between 1 and 3; occasional in 5.

Persea borbonia (L.) Sprengel. Red Bay. Common in all forested communities.

Sassafras albidium (Nuttall) Nees. Sassafras. Common in less mesic areas of 1, especially along borders of fields.

Papaveraceae

Sanguinaria canadensis L. Bloodroot. Common in 1; occasional in 5.

Brassicaceae

Cardamine pensylvanica Muhl. Bitter Cress. Common in 2 and apparently confined to this habitat.

Saxifragaceae

Decumaria barbara L. Climbing Hydrangea. Common in 3 and 5.

Itea virginica L. Virginia Willow. Common in 3 growing around bases of trees and on fallen logs, stumps, cypress knees, and buttresses; common around edge of 2.

Hamamelidaceae

Hamamelis virginiana L. Witch-hazel. Common in 1 and 5.

Liquidambar styraciflua L. Sweet Gum. Common in 1, 4 and 5; occasional in 3 as small tree on fallen logs, stumps, cypress knees, and buttresses.

Rosaceae

Agrimonia gryposepala Wallroth. Harvest-lice. Common in II

Amelanchier arborea (Michaux f.) Fernald. Juneberry. Rare; only two small trees found growing in cutover woods around Visitor's Center.

Crataegus narshallii Eggl. Hawthorn. Common in 4 and 5.

C. punctata Jacquin. Hawthorn. 1.

Geum canadense Jacquin. Avens. Occasional in 1.

G. laciniatum Murray. Avens. Rare in 1.

Prunus americana Marshall. Wild Plum. Occasional in less mesic sites

P. serotina Ehrhart var. serotina. Black Cherry. Common in 1 and 5.

P. umbellata Ell. Hog Plum. Rare; only two specimens found, both along edge of 1 where it borders field along Mims Lake.

Rubus betulifolius Small. Blackberry. Common in 3 on stumps, fallen logs, buttresses, and cypress knees.

R. trivialis Michaux. Dewberry. Occasional in 1 and 5.

Sorbus arbutifolia (L.) Heynhold var. arbutifolia. Red Chokeberry. Occasional in 1, 5, and disturbed woods around Visitor's Center.

Fabaceae

Amphicarpa bracteata (L.) Fernald. Hog Peanut. Rare; less mesic sites.

Cercis canadensis L. Redbud. Common in 1.

Clitoria mariana L. Butterfly Pea. Occasional in less mesic sites in 1.

Desomodium rotundifolium DC. Dollarleaf. Common in 1.

Desomodium sp. Beggar Lice. Common in 1.

Erythrina herbacea L. Cherokee Bean. Rare in 1.

Gleditsia aquatica Marshall. Water Locust. Common in 3; occasional in

Pueraria lobata (Willd.) Ohwi. Kudzu. A stand of Kudzu is rapidly invading the mixed mesophytic hardwood forest near the landing at Mims Lake; not found elsewhere.

Wisteria frutescens (L.) Poiret. Wisteria. Scattered throughout 3.

Euphorbiaceae

Acalypha gracilens Gray. Three-seeded Mercury. Rare in 1.

Cnidoscolus stimulosis (Michaux) Engelm. & Gray. Bull Nettle. Occasional in less mesic sites in 1.

Euphorbia corollata L. Flowering Spurge. Common in less mesic sites in 1.

Anacardiaceae

Rhus copallina L. Winged Sumac. Common in 1 and 5.

R. radicans L. Poison Ivy. Common in all forested communities; common in 3 on stumps, fallen logs, cypress knees and buttresses.

Aquifoliaceae

Ilex decidua Waiter. Possum Haw. Common in 4; occasional in 3 and 5.

Ilex opaca Aiton. Holly. Common in 1, 4 and 5.

Celastraceae

Euonymus anericanus L. Strawberry Bush. Common in 1 and 5.

Aceraceae

Acer rubrum L. Red Maple. In 3 it occurs occasionally on fallen logs, cypress knees, stumps, and buttresses; common in 1, 4 and 5.

Hippocastanaceae

Aesculus pavia L. Red Buckeye. Common in 1 and 5.

Balsaminaceae

Impatiens capensis Meerb. Jewel-weed. Common in 2, generally forming dense thickets in the fall.

Rhamnaceae

Berchemia scandens (Hill) K. Koch. Supple-jack Vine. Common in all forested communities.

Vitaceae

Ampelopsis arborea (L.) Koehne. Pepper-vine. Common in all forested communities.

Parthenocissus quinquefolia (L.) Planchon. Virginia Creeper. Common in 1, 4, 5, and disturbed woods around Visitor's Center.

Vitis rotundifolia Michaux. Muscadine. Common in ail forested communities.

V. vulpina L. Frost Grape. Rare; found only once on edge of 1 where it borders field.

Hypericaceae

Hypericum densiflorum Pursh. St. John's-wort. Occasional in 3, usually growing on cypress knees, fallen logs, buttresses, and stumps.

H. hypericoides (L.) Grantz. St. Andrews Cross. Occasional in 1.

H. mutilum L. St. John's-wort. Rare in 2.

H. punctatum Lam. St. John's-wort. Rare in 2.

H. virginicun L. St. John's-wort. Common on fallen logs in 3.

Violaceae

Viola palmata L. Violet. Common in 1 and 5; occasional in 4.

V. primulifolia L. Violet. 1.

Passifloraceae

Passiflora lutea L. Passion-flower. Occasional in 1 and 5.

Lythraceae

Cuphea carthagensis (Jacquin) Macbride. Waxweed. Rare; found on old abandoned road in low area around Visitor's Center.

Melastomataceae

Rhexia mariana L. var. mariana. Meadow Beauty. Rare; found on old abandoned road in low area around Visitor's Center.

Onagraceae

Ludwigia decurrens Waiter. Rare; found on abandoned road in low area around Visitor's Center.

L. palustris (L.) Ell. Common throughout 3 along edge of lakes and creeks.

L. repens Forster. 3.

Haloragaceae

Proserpinaca palustris L. Common in higher sites of 3, often occurring in dense mats.

Araliaceae

Aralia spinosa L. Hercules Club. Occasional in 1.

Apiaceae

Cicuta maculata L. Water Hemlock. Rare; found along edge of Mims Lake.

Hyclrocotyle verticillata Thunberg. Common in 2 and in higher sites in 3.

Ptilimnium capillaceum (Michaux) Raf. Rare; found along abandoned road in low area around Visitor's Center.

Sanicula canadensis L. Snakeroot. Common in 1.

Siun suave Waiter. Water Parsnip. Occasional along lake edges in 3.

Thaspium barbinode (Michaux) Nuttall. Meadow Parsnip. Rare in 1.

Nyssaceae

Nyssa aquatica L. Tupelo Gum. Common in 3; rare in 4.

N. sylvatica var. biflora (Waiter) Sargent. Occasional in 3.

N. sylvatica Marshall var. sylvatica. Black Gum. Occasional in 1.

Cornaceae

Cornus asperifolia Michaux. Dogwood. Rare; disturbed woods around Visitor's Center.

C. florida. L. Flowering Dogwood. Common in 1 and 5.

C. stricta Lam. Swamp Dogwood. Common in 1, 4 and 5.

Clethraceae

Clethra alnifolia var. tomentosa (Lam.) Michaux. Sweet Pepperbush. Common in 1 and 5.

Ericaceae

Chimaphila maculata (L.) Pursh. Spotten Wintergreen. Common in 1 and 5.

Gaylussacia frondosa (L.) T. & G. Dangleberry. Rare; found in disturbed woods around Visitor's Center and in 1.

Leucothoe axillaris (Lam.) D. Don. var. axillaris. Leucothoe. Rare; found only along road leading to Visitor's Center.

L. racemosa (L.) Gray. Fetter-bush. Common throughout 3, often growing from base of cypress buttresses.

Lyonia lucida (Lam.) K. Koch. Fetter-bush. Common in 3 growing from buttresses of cypress trees and as terrestrial shrub in elevated sites.

Rhododendron alabamense Rehder. Wild Azalea. Disturbed woods around Visitor's Center; rare.

R. canescens (Michaux) Sweet. Wild Azalea. Occasional in 1 and 5.

Vaccinium amoenum Aiton. Rare; disturbed woods around Visitor's Center.

V. arboreum Marshall. Sparkleberry. Common in 1 and 5.

V. corymbosum L. Highbush Blueberry. Rare; disturbed woods around Visitor's Center.

V. elliottii Chapman. Common in 1, 4 and 5.

V. tenellum Aiton. Rare; found in less mesic sites in 1.

Primulaceae

Samolus parviflorus Raf. Water Pimpernel. Common in elevated sites in 3 and around 2.

Sapotaceae

Bumelia lycioides (L.) Persoon. Buckthorn, Ironwood. Rare; few scattered small trees found on less mesic sites in 1 south of powerline on east side of swamp.

Ebenaceae

Diospyros virginiana L. Persimmon. Common in I and 5; occasional in 4 where some specimens reach large size.

Symplocaceae

Symplocos tinctoria (L.) L'Her. Horse Sugar. Rare; 1 and 5.

Styracaceae

Halesia carolina L. Silverbell. Rare; one plant found in 1 bordering Mims Lake.

Styrax amer'cana Lam. Storax. Common in 3; occasional in 4 and 5.

Oleaceae

Chionanthus virginicus L. Fringe-tree. Rare; one specimen found along road leading into Visitor's Center.

Fraxinus americana L. var. americana. White Ash. Rare in 1.

F. caroliniana Miller. Water Ash. Common in 3 as small tree and forming dense subcanopy in more elevated sites.

F. tonentosa Michaux f. Red Ash. Rare in 3.

Loganiaceae

Cynoctonum mitreola (L.) Britton. Mitterwort. Common in 2, 3, 4 and 5.

Gelsemium semperuirens (L.) Aiton. Yellow Jessamine. Common in 1 and 5.

Spigelia marilandica L. Indian Pink. Common in I and 5.

Gentianaceae

Sabatia calycina (Lam.) Heller. Occasional in 3, 4 and 5.

S. dodecandra (L.) BSP. Sabatia. Rare in 3.

Apocynaceae

Apocynum cannabinun L. Indian Hemp. Rare in 1.

Asclepidaceae

Ascelpias perennis Waiter. Milkweed. Common 3 and 4.

A. uariegata L. Milkweed. Rare in 1.

Matelea suberosa (L.) Shinners. Rare in 1.

Convolvulaceae

Ipomoea hederacea (L.) Jacquin var. hederacea. Morning Glory. Rare in 1.

Polemoniaceae

Phlox glaberrima L. Phlox. Rare in 1.

Verbenaceae

Callicarpa americana L. French Mulberry. Common in 1 and 5.

Verbena urticifolia L. Occasional in 1 and 5.

Phrymaceae

Phryma leptostachya L. Lop-seed. Common in more mesic sites in 1.

Lamiaceae

Collinsonia tuberosa Michaux. Scattered but locally abundant in 1.

Dracocephalum purpureum (Waiter) McClintock. Obedient Plant. Occasional and scattered throughout 3 in more elevated sites.

Lycopus rubellus Moench var. rubellus. Common and apparently restricted to fallen logs, stumps, cypress knees, and buttresses in 3.

L. virginicus L. Rare; collected along abandoned road in low area around Visitor's Center.

Perilla frutescens (L.) Britton. Rare; found only on disturbed site around camp ground at Mims Lake and along lake edge.

Prunella vulgaris L. Rare and widely scattered in less mesic sites in 1.

Scutellaria integrifolia L. var. integrifolia. Skullcap. Occasional in less mesic sites in 1.

S. lateriflora L. Skullcap. Common and apparently confined to fallen logs, stumps, cypress knees and buttresses in 3.

Solanaceae

Solanum americanum Miller. Nightshade. Rare and widely scattered in less mesic sites in 1.

Scrophulariaceae

Aureolaria virginica (L.) Pennell. Occasional in less mesic sites of 1.

Bacopa monnieri (L.) Pennell. Common in 2.

Gratiola virginiana L. Common and apparently confined to 2.

Micranthenum umbrosum (J. F. Gmelin) Blake. Common in 2 and wet areas in 4.

Mimulus alatus Aiton. Rare; found along abandoned road in low area near Visitor's Center.

Bignoniaceae

Anisostichus capreolata (L.) Bureau. Cross Vine. Common in all forested communities.

Campsis radicans (L.) Seemann. Trumpet Vine. Common in all forested communities.

Orobanchaceae

Conopholis anericana (L.) Wallroth. Cancer-root. Occasional in 1.

Epifagus virginiana (L.) Barton. Beech-drops. Parasite on roots of Fagus grandifolia; occurs in more mesic sites of 1 where beech occurs.

Acanthaceae

Justicia ouata (Waiter) Lindau. Common in elevated sites in 3.

Ruellia caroliniensis (Waiter) Steudel. Common in 1.

Rubiaceae

Cephalanthus occidentalis L. Button Bush. Common in 3, especially along edge of lakes.

Diodia teres Waiter. Found along abandoned road in low area near Visitor's Center.

D. virginiana L. Occasional on fallen logs in 3.

Galium circaezans Michaux. Bedstraw. Common in 1.

G. tinctoriun L. Bedstraw. Common in 1 and 5.

G. triflorum Michaux. Bedstraw. Common in 1 and 5.

Caprifoliaceae

Lonicera japonica Thunberg. Japanese Honeysuckle. Occasional in forested communities where they border fields or have openings created by disturbances.

L. sempervirens L. Coral Honeysuckle. Occasional in 1; disturbed woods around Visitor's Center.

Sambucus canadensis L. Elderberry. Occasional in 1 and 5; disturbed woods around Visitor's Center.

Viburnum dentatum L. var. dentatum. Common in 4 and 5; occasional in elevated sites in 3.

V. obovatum Waiter. Occasional in 4 and 5.

V. prunifolium L. Black Haw. Rare; found only along road leading to Visitor's Center.

Cucurbitaceae

Melothria pendula L. Creeping Cucumber. Occasional in 1.

Campanulaceae

Lobelia cardinalis L. Cardinal Flower. Common in 3 in higher sites, where it often grows in colonies, and on fallen logs and stumps.

L. elongata Small. Lobelia. Scattered throughout 3 in elevated sites.

Asteraceae

Baccharis halimifolia L. Sea-Myrtle. Occasional in 1, 5 and edge of lakes in 3.

Bidens bipinnata L. Spanish Needles. 1.

Carduus lecontei (L.) Waiter. Thistle. 1.

Chrysogonum virginianum L. Green-and-Gold. Occasional in less mesic sites in 1.

Elephantopus carolinianus Willd. Elephant's-foot. 1.

E. nudatus Gray. Elephant's-foot. 1.

E. tomentosus L. Elephant's-foot. Common in 1.

Erechtites hieracifolia (L.) Raf. Fireweed. Occasional in 1.

Eupatorium coelestinum L. Ageratum. 1.

E. compositifolium Waiter. Dog-fennel. Rare; fallen log in 3.

E. dubium Willd. Rare in 1.

E. perfoliatum L. Boneset. Abandoned road in low area around Visitor's Center; rare.

Heliopsis helianthoides (L.) BSP. Ox-eye. Locally abundant in 1 north east of power line on east side of swamp.

Lactuca floridana (L.) Gaertner. Wild Lettuce. Rare in 1.

Melanthera hastata Michaux. Rare; less mesic sites in 1.

Mikania scandens (L.) Willd. Climbing Hempweed. Occasional in 3 and in 1 along borders with fields.

Pluchea camphorata (L.) DC. Marsh-fleabane. Rare; collected along abandoned road in low area around Visitor's Center.

Senecio glabellus Poiret. Butterweed. Common in 2; scattered throughout 3.

Solidago arguta Aiton. Goldenrod. 1.

Verbesina occidentalis (L.) Waiter. Locally abundant in scattered sites in 1.

V. virginica L. var. virginica. Locally abundant in scattered sites in 1.

ACKNOWLEDGMENTS

LITERATURE CITED

BRUNSWIG. N. L., and S. G. WINTON. 1978 The Francis Beidler Forest in Four Holes Swamp. National Audubon Society, New York, N.Y.

CARR, ARCHIE. November 1971. Black water-green light. In: Audubon: The magazine of the National Audubon Society. New York, N.Y.

DENNIS, J. V. 1970. Four Holes Swamp, Berkeley and Dorchester counties, S.C.: study of the natural history and matters pertaining to acquisition of one of the last large virgin bottomland swamps in the South. Unpub. report to the Nature Conservancy, Arlington, Va

DENNIS, W. M., and W. T. BATSON. 1974. The floating log and stump communities in the Santee Swamp of South Carolina. Castanea 39(2):166-170

DEPARTMENT OF THE INTERIOR, FISH AND WILDI.LFE SERVICE 1975 Threatened or endangered fauna or flora. Reviewof status or vascular plants and determination of "critical habitat." Federal Register 40(127) Part V: 27824-27924.

GADDY, L. L., T. S. KOHLSAAT, E. A. LAURENT, and K. B. STANSELL. 1976 A vegetation analysis of preserve alternatives involving The Beidler Tract of Congaree Swamp. South Carolina Wildlife and Marine Resources Department, Columbia, S.C.

HUNT, K. W. 1947. The Charleston woody flora. The American Midland Naturalist 37(31:670-756

PENFOUND, W. T. 1952. The southern swamps and marshes. Bot. Rev. 18:413-446.

PORCHER, R. D. 1974. Some notes and distribution records for Orchidaceae from the South Carolina Coastal Plain. J. Elisha Mitchell Sci. Sec. 90(2):73-74.
------------ 1979. Identifying, locating, and inventorying populations of Proposed endangered and threatened vascular plants of the Francis Marion National Forest in Berkeley and Dorchester counties, S.C. Report submitted to U.S. Forest Service, Atlanta, Ga 17 pp.

RADFORD, A. E., H. E. AHLES, and C. R. BEI.L.1968. Manual of the vascular flora of the Carolinas. University of North Carolina Press, Chapel Hill. 1183 pp.

RADFORD, A. E., L. J. OTTE. and D. S. OTTE. 1978. Ecological diversity classification and inventory. In: Natural heritage classification and inventory. University of North Carolina Student Stores, Chapel Hill.

WELLS, B. W. 1942. Ecological problems of the southeastern U.S. Coastal Plain. Bet.Rev. 8:533-62.

THE CITADEL
BIOLOGY DEPARTMENT
CHARLESTON, S.C. 29409