Longleaf Pine Vegetation of the Southern Atlantic and Eastern Gulf Coast Regions: A Preliminary Classification*

Robert K. Peet Department of Biology, CB"3280, University of North Carolina, Chapel Hill, NC 27599-3280

Dorothy J. Allard The Nature Conservancy, Southeast Regional Office, Box 2267, Chapel Hill, NC 27515

ABSTRACT

Quantitative data on the composition of natural longleaf pine-dominated vegetation collected across the range of the species east of the Mississippi River are used to develop a pre­ liminary, floristically-based, region-wide classification for use in conservation and preser­ vation planning.

The strongest compositional gradients appear related to soil moisture. We recognize four major series of longleaf-dominated vegetation, primarily differentiated with respect to this gradient (xeric, subxeric, mesic, and seasonally wet). These series are divided into twenty­ three communities, which correspond primarily to geographic position and physiographic province (the coastal plain and maritime fringe regions of the Atlantic and Gulf coasts re­ spectively, the piedmont I uplands, and the fall-line sandhills).

The five communities that belong to the Xeric Longleaf Woodland series occur on coarse, well-drained sands. The six Subxeric Longleaf Woodland communities made up the ma­ jority of the longleaf-dominated landscape of presettlement times . The four Mesic Longleaf Woodland communities are remarkably rich in species, but are uncommon in the modern landscape because they are largely confined to soils well-suited for agriculture . The eight Seasonally-Wet Longleaf Woodland communities contain both shrubby flatwoods and grassy, floristically-rich savannas.

Despite a visual dominance by longleaf pine, wiregrass, and scrub oaks, the greater longleaf pine ecosystem of the southeastern United States contains some of the most diverse plant communities known from the temperate zone. Longleaf Savannas were regularly observed with over 40 species of plants per square meter, and Mesic Longleaf Woodlands were found 2 with up to 140 species per 1000 m . Many of these species are largely confined to longleaf pine-dominated communities. These natural longleaf woodlands are being lost rapidly to a combination of land development and fire suppression.

'Botanical nomenclature follows Kartesz (1994), except we follow Peet (1993) in recognizing that the plants traditionally treated as Aristida stricta should be divided into a northern (A. stricta) and a southern species (A. beyrichiana) .

Proceedings of Ihe Tall Timbers Fire Ec%g!' Conference. No. 18, The Longleaf Pine Ecosystem: ecology, restoration and management, edited by Sharon M. Hermann, Tall Timbers Research Station, Tallahassee, FL, 1993

45 INTRODUCTION fire for stand maintenance and reproduction. Be­ fore fire suppression, regular, low-intensity surface Three centuries ago longleaf pine (Pinu s fires kept the pine woodlands open and relatively paiustris) dominated the coastal plain landscape of free of undergrowth. The presence of abundant the southeastern United States. However, settle­ grass, especially wiregrass (Aristida stricta in the ment of the region by Europeans dramatically al­ north, A. beyrichiana in the south; see Fig. 1) and tered the longleaf ecosystem (see Croker 1987, Frost bluestem grasses (Andropogon spp ., Schizachyrium 1993, Ware et al. 1993). As a consequence, much spp.) provided a ready source of spatially continu­ of the area once dominated by longleaf retains few, ous fuel which helped fire spread throughout the if any, longleaf trees. pine woodlands (Christensen 1988, Noss 1989, Stout and Marion 1993). Without fire, longleaf Initially, longleaf pine was heavily exploited stands develop a thick undergrowth of for tar, turpentine and rosin production. Most of broadleaved species under which pine regenera­ the mature trees that survived were eventually cut tion is impossible. In addition, fuel levels can build for timber. Pine reproduction failed, primarily be­ to the point that fire is catastrophic when it even­ cause of suppression of the fires that historically tually does occur. In the absence of fire, longleaf had controlled potential woody competitors, and vegetation declines in species diversity owing to because of the ubiquitous grazing of livestock, es­ decreased light and increased litter depth. Preser­ pecially hogs which voraciously consumed young vation of longleaf-dominated woodlands is not suf­ pines for their starchy taproots (Schwarz 1907, Hine ficient for preservation of the longleaf ecosystem 1925, Croker 1987, Lipscomb 1989, Frost 1993). Fi­ and its attendant biodiversity. Because the longleaf nally, because of the prevailing gentle topography, ecosystem is fire-maintained, only those few sites those areas with tillable soils were readily con­ that have continued to experience chronic fire re­ verted to agricultural production. In short, the tain a strong resemblance to the natural longleaf combined impact of the naval stores industry, lum­ systems of the Southeast. ber extraction, grazing and agriculture has served to remove longleaf from much of its former range. Examples of natural longleaf vegetation con­ This is p articularly true in the northern portion of taining both old-growth trees and an understory the longleaf range where the pines were exploited unaltered by fire suppression are almost nonexist­ first. Today, longleaf is nearly absent from the ent. Fortunately, fire has continued to be a tool for Neuse River in central North Carolina northward, land management in many longleaf areas with the despite the fact that this species once dominated result that examples of second-growth stands with much of the coastal plain of northeastern North the understory vegetation still intact can be found, Carolina and southeastern Virginia (Fig. 1; Pinchot particularly on public lands such as national and and Ashe 1897, Frost and Musselman 1987, Frost state forests, gamelands, and military bases. While 1993). over 70% of the remaining longleaf vegetation is in private ownership, fire suppression is more perva­ Longleaf pine is not the only distinctive spe­ sive in these generally smaller holdings. Over the cies of the once vast southeastern pinelands. The total original natural range of longleaf, less than 3% longleaf-dominated ecosystem also supports a of the natural upland vegetation remains in a semi­ great diversity of distinctive plant and animal spe­ natural, fire-maintained condition (Frost 1993). Fur­ cies which today persist only in the small frag­ ther, this residual fraction is not really ments of the original landscape that have representative of the original vegetation in that the managed to escape the bulk of the changes soils most conducive to agriculture were largely wrought by the growth of modern society. Not cleared of their natural vegetation well over a cen­ only has the exploitation of the longleaf resource tury ago (Pinchot and Ashe 1897, Mohr 1901, per se been devastating to this diversity, but other, Harper 1906, Frost 1993) with the consequence that more subtle changes have had equally significant the remaining fragments persist primarily on atypi­ impacts. The most important of these has been cally wet or dry sites. the elimination of chronic fire. More recently, me­ chanical damage to the understory of longleaf Longleaf vegetation, while widespread, has stands by pinestraw raking and mechanized tim­ been remarkably little studied (Noss 1988, Schafale ber removal has begun to significantly reduce and Weakley 1990, Stout and Marion 1993). Docu­ populations of many of the native species of the mentation of compositional variation can be found longleaf ecosystem. in the scientific literature for small portions of this system over limited ranges of soil conditions (e.g., Longleaf pine absolutely depends on frequent Bozeman 1971, Kologiski 1977, Taggart 1990, 1994).

46 o 100 200 300 400 MILES I I I I I I I I I I I I I 1'1'1 I I ' I I I ' I I I I 1'1 I I I I I I o 100 200 300 400 500 600 KILOMETERS

Figure 1. The 216 sample sites used in our final analysis were located in 44 counties scattered throughout much of the range of longleaf pine (Pinus pa/uslriSj east of the Mississippi River. The range of longleaf pine is indicated by stipples (after littlE) 1971 and Frost 1993) and the range of wiregrass (Arisl/da slricla and A. beyrichiana) by diagonal shading (after Peet 1993).

However, most of the longleaf region has not been longleaf pine ecosystem so that we will know what subjected to rigorous ecological study. For some needs to be preserved. Toward this end, the Natu­ regions it is now too late; for example, we can ral Heritage Programs in many of the southeastern hardly begin to describe the original longleaf veg­ states, in collaboration with The Nature Conser­ etation of northeastern North Carolina and south­ vancy, have developed classifications of natural eastern Virginia as virtually nothing is left to study communities, including those dominated by (but see Frost and Musselman 1987, Frost 1993). longleaf pine. Our goal in this paper is to combine Further, while descriptive treatments have been information from these qualitative state classifica­ published for various portions of the longleaf re­ tions with quantitative data collected by several gion (e.g., Harper 1906, Pessin 1933), there have independent researchers, to create a preliminary been no attempts to quantitatively document the classification of the natural longleaf-dominated floristic and structural variation in this ecosystem vegetation east of the Mississippi River. (Longleaf­ at a scale larger than a few counties. dominated communities west of the Mississippi are described by Bridges and Orzell 1989 and If a significant fraction of the biotic diversity Harcombe et al. 1993). of the longleaf pine ecosystem is to be preserved, we need to act rapidly. A critical early step in this process is documentation of the variation in the

47 METHODS nomy, plant species composition, geographic dis­ tribution, and important environmental factors Approach such as moisture and soil texture. Quantitative data collection in longleaf communities was initi­ Vegetation classification typically is a process ated prior to creation of the initial classification, of successive approximation. As our knowledge which allowed some field experience gained dur­ base increases, we can produce better descriptions ing that activity to influence the form of the classi­ and classifications, which in turn motivate new ob­ fication. Published literature on longleaf servations, which allow still better descriptions and communities was also used, but to a lesser degree. classifications. Several cycles of this process gen­ The initial classification included 15 community erally are required before the major patterns of types that spanned nearly all the major natural variation in a widely distributed vegetation type, communities in which longleaf pine dominates the such as the longleaf-dominated vegetation of the canopy or shares dominance with other species. southeastern United States, can be understood. Our somewhat informal method of classification Both the initial classification and our subse­ recognizes the importance of this successive ap­ quent preliminary classification were designed proximation approach. We further recognize that with the intent that protection of several high-qual­ our classification is only a preliminary effort, and ity examples of each of the communities, selected will certainly require revision as additional infor­ to represent the range of variation within each type, mation becomes available. should be sufficient to protect and preserve much of the biota of the greater longleaf-dominated eco­ Our approach to classification of longleaf veg­ system. This approach, when combined with both etation involved several steps. The first step was additional efforts to protect rare plant species that creation of an initial classification of longleaf pine occur in longleaf communities, and management of communities based on existing vegetation classifi­ large, longleaf - dominated landscapes to sustain cations and other descriptive information. This ecological processes such as fire, should provide an classification was developed by DJA as part of the effective strategy for protection of the longleaf pine creation of a "Southeastern United States Ecologi­ ecosystem and its biodiversity. cal Community Classification" for use by The Na­ ture Conservancy in protection of biodiversity at Vegetation data the community level (Allard 1990). The second step was to collect quantitative data on community We sought quantitative data on the species composition from across the range of longleaf to composition of longleaf pine vegetation from help refine and validate the initial classification. Si­ throughout the range of the species east of the Mis­ multaneously, quantitative data were collected by sissippi River. All stand data selected for inclusion the North Carolina Vegetation Survey (see Ac­ in the study included a complete list of the vascu­ knowledgments) from longleaf vegetation in the lar plant species in each sample plot, plus a mea­ North Carolina fall-line sandhills as part of an in­ sure of species importance that could be dependent project to validate and refine the transformed to approximate a ten-point cover/ North Carolina community classification of abundance scale. (Cover refers to the percentage of Schafale and Weakley (1990). These two datasets ground surface that would be covered by the leaf were supplemented with quantitative data from area projection of a particular species.) We in­ five other studies of longleaf-dominated vegetation cluded only stands that had not been subjected to to produce a dataset which included nearly 250 an extended period of fire suppression. After samples (Appendix I). In each case the botanical stands known to be degraded by fire suppression nomenclature was revised to conform to Kartesz or pinestraw raking were excluded, along with (1994). In the third step, these data were subjected some examples of types that were over-repre­ to various forms of multivariate analysis to refine sented, the final dataset included data from seven the initial classification and to allow better charac­ sources and contained 216 longleaf pine stands rep­ terization of the component communities. resenting 44 counties spread across all states within the range of longleaf pine east of the Mississippi The Southeastern Ecological Community Clas­ River, except Virginia (Fig. 1). Virginia was ex­ sification, from which our initial classification was cluded because the only known extant example of developed, was constructed primarily from the longleaf vegetation in Virginia has been strongly Natural Heritage Program classifications of the modified by fire suppression and logging (see Frost twelve southeastern states. Community attributes and Musselman 1987). Details of the datasets em­ used to create the classification included physiog­ ployed are summarized in Appendix 1.

48 Our standard cover/abundance scale is that cation (Allard 1990). As groups of stands were rec­ developed by the North Carolina Vegetation Sur­ ognized, those stands near the edges of groups or vey to provide maximum ease of interconversion that did not fit well were reexamined to see if they with other widely-used scales: 1 = trace, 2 = <1 % might better fit into another community type. cover,3 =1-2%,4 =2-5%, 5 =5-10%,6 =10-25%, 7 = 25-50%, 8 = 50-75%, 9 = 75-95%, 10 = > 95% cover. Our analysis and results are presented as a se­ For each of the seven datasets used, cover values ries of four two-dimensional ordination diagrams were transformed to approximate this scale. (Figs. 3-6). Symbols are used in these figures to in­ dicate the final community type assignments of the vegetation samples. These diagrams show stands Multivariate analysis arranged in ordination space, so the axes are di­ rectly interpretable only in terms of species com­ Ordination methods frequently are used to ar­ position. Nonetheless, correlations with range vegetation samples in an abstract, multidi­ environmental variables exist and are described in mensional space in such a fashion that samples the text. Further, the diagrams can be used to ex­ with similar species composition (and, therefore, amine the relationships among the recognized similar underlying environmental control) are lo­ community types and the degree to which the cated near each other, while dissimilar samples are types differ from each other. located far apart. This allows identification and visualization of the dominant trends in composi­ tion. In an ideal, perfectly orderly world, the vari­ PHYSIOGRAPHY OF THE LONGLEAF ous axes of the multidimensional space would be PINE REGION interpretable in terms of environmental variables responsible for the vegetation pattern observed. In Although longleaf pine dominated the prime­ practice, only the first one or two axes are usually val vegetation of much of the Southeast, the area interpretable, while the meaning of the remaining where it occurred was far from homogeneous. The variation is obscured by interactions and changing natural range of longleaf covers nearly all the importances of the critical factors with respect to southeastern coastal plain and spills over onto the the first few axes extracted. adjacent piedmont and interior uplands. Within the coastal plain, the species ranges from southeast To simplify interpretation of complex, multidi­ Virginia south to central Florida and west to Texas, mensional datasets, a strategy of progressive frag­ a large region that exhibits considerable variation mentation (Peet 1980) can be employed. Here, the in both geology and topography (Fig. 2). first one or two axes are examined and interpreted. Interpretation is based on knowledge of the sites, The coastal plain is a region of marine sedi­ and environmental data where they are available. ments, in many cases extensively reworked by Then, a portion the dataset that is seen in the first wave action. Because the coastal plain varies in to­ ordination to be readily interpretable in terms of pographic relief, it is convenient to recognize both some sort of environmental extreme is removed a region of coastal flatlands where local relief is less from the dataset so as to reduce its influence in the than 35 m and over 80% of the land surface is at subsequent ordinations. In this fashion, the dataset most gently sloping, and a region of rolling hills can be progressively simplified, and more subtle (see Fig. 2). This physiogtaphic division follows and deeply buried patterns can be exposed and in­ Hammond (1964), but also approximates the divi­ terpreted. sions recognized by Hodgkins (1965; Flatlands Coastal Plain and Undulating Coastal Plain) and by We employed a strategy of progressive frag­ Hodgkins et al. (1979; Middle Coastal Plain and mentation using Detrended Correspondence Hilly Coastal Plain). Analysis as an ordination technique (CANOCO 3.1; Hill and Gauch 1980, ter Braak 1987, see Peet et al. Coastal flatlands are best developed along the 1988). We also used a numerical classification pro­ Atlantic coastal plain, but a narrow band of low re­ duced using two-way indicator species analysis lief continues along the Gulf coast. Marine trans­ (TWINSPAN; Hill 1979) to help refine the divisions gressions across this flat landscape have left their in the dataset and to characterize the resulting clus­ marks in ways that strongly influence vegetation ters. At each step, tentative community types were composition. Much of this flat outer coastal plain recognized in the ordinations, with the first ap­ can be visualized as consisting of a series of old proximation based on the initial classification barrier dunes composed of coarse, siliceous sands, developed from the Nature Conservancy classifi­ behind which are old embayment areas with soils 49 Interior Uplands

Coastal Flatlands

Figure 2. Longleaf pine (vertical lines) is distributed across several physiographic provinces, each with relatively distinct, longleaf-dominated communities (longleaf distribution after Little 1971 and Frost 1993; physiographic provinces modified from Hammond 1964 and Hodgkins 1965). that are much finer and often dominated by fine coastal plain allows better drainage with the con­ clayey sands. Soils derived from the barrier dune sequence that seasonally wet sites are less common, systems tend to be extremely dry due to the rapid mostly of local occurrence, and associated with percolation of water, whereas the soils of the near-surface impermeable and often indurated soil embayment regions tend to be seasonally saturated horizons. because the clay content of the soil and low relief make for poor drainage (DuBar et al. 1974, Daniels Along the inner-most portion of the coastal et al. 1984, Soller and Mills 1991). plain from central North Carolina around to the eastern edge of Alabama are found the fall-line Inland from the flatlands of the more recent sandhills. This mass of primarily Cretaceous-age marine terraces, the coastal plain is typically a re­ sandy sediments, in some places capped with Mi­ gion of low, rolling hills, often with loamy soils ocene dunes, is apparently the product of erosion (Fig. 2). Farther inland the landforms are older and of high mountains that once stood where today the topography is more hilly. A distinctive region there remain only the low hills of the piedmont. of clay hills occurs in Alabama and Mississippi Erosion of these piedmont hills has been so com­ (Hodgkins 1965, Hodgkins et al. 1979), which ex­ plete that the elevations of the sandhills now some­ tends a little into Georgia (Harper 1930). A simi­ times exceed those of the adjacent piedmont, lar but smaller area has also been recognized in erosion having been less intense because in the South Carolina (Myers et al. 1986). The more pro­ sandhills water drains readily into the sandy soil nounced topographic relief of the rolling inner rather than running off the surface. These coarse

50 sands also cause the prevailing sandhill soils to be rates almost perfectly samples from the Gulf highly permeable and consequently very droughty Coast states (upper right) from the Carolina for plant growth. However, embedded in these old coastal plain and fall-line sandhill samples (lower marine deltaic sands are frequent clay lenses that lo­ left). Our few sites outside the coastal plain seg­ cally inhibit drainage such that seeps occur where the regated at the far upper left with the mesic lenses outcrop (see Sohl and Owens 1991). coastal plain sites. In our nomenclature, we des­ ignate the coastal plain of the Gulf states (AL, FL, Inland from the sandhills, north and west of the GA, LA, MS), including the Atlantic coastal plain fall-line, is the piedmont region where marine sedi­ of Georgia, as "Southern", the coastal plain of the ments are replaced primarily by clay soils derived Carolinas (NC, SC) as "Atlantic", and the fall-line from weathering of ancient igneous and metamor­ sandhills (AL, GA, NC, SC) as ''Fall-line''. The At­ phic rocks. Most of these areas have been above sea lantic region primarily falls within the coastal level since well before the start of the Tertiary, with flatlands, but the southern segregate includes the consequent that the soils are highly weathered both the coastal flatlands of Georgia and Florida and infertile, and the drainage systems are well de­ and the rolling hills of the Gulf coastal plain. veloped. Farther west are the interior uplands of the Blue Ridge and the Ridge and Valley Provinces, again The xeric sandhill samples from western with ancient soils. Florida segregated perfectly in the second ordi­ nation (Fig. 4) and these are designated as South­ The longleaf vegetation of the coastal plain is ern Xeric Longleaf Woodland. The three mesic well known to vary with soil drainage from xeric san­ sites from the Gulf coastal plain also segregate dhill sites with coarse sandy soils to floristically rich well (Southern Mesic Longleaf Woodland), as do savannas and flatwoods of poorly drained flatlands two samples from an unusual Gulf Coast type (Mohr 1901, Harper 1914b, Wells 1932, Braun 1950, dominated by saw palmetto (Subxeric Longleaf Wharton 1978, Christensen 1988). This well-docu­ - Saw Palmetto Woodland; see Pessin 1933, Allen mented pattern led us to expect soil moisture to be 1956). All the other likely groups still exhibit a critical factor controlling composition of longleaf some overlap in their membership. The three vegetation. We also anticipated that composition distinct groups (within bold lines in Fig. 4) were would vary in an interpretable manner between removed from the ordination, along with the physiographic regions owing to differences in cli­ single but distinctive sample from serpentine mate, soil texture and soil fertility. soils of the Georgia piedmont (Serpentine Subxeric Longleaf Woodland).

VEGETATION PATTERNS The final dry-site ordination (Fig. 5) shows very little overlap of the final recognized clusters. Regional gradients The predominant gradients are again related to moisture and geography, but soil texture and nu­ Ordination of the complete dataset (Fig. 3) re­ trients also appear important. Xeric, well­ drained, infertile quartz sands are in the lower vealed a strong primary axis corresponding to soil moisture. Less pronounced sorting by latitude oc­ right while the clayey piedmont and upland sites curs along the second axes. This result led us to par­ (Piedmont/Upland Longleaf Woodland) segre­ tition the dataset almost exactly in the middle of the gate in the upper left, with the silty, mesic sites ordination (bold line in Fig. 3), the break separating (Fall-line Mesic Longleaf Woodland) on the far those sites that appeared to have seasonally-satu­ left. Subxeric sites between the two extremes of rated soils from those with better-drained soils. the soil texture gradient, sorted along an orthogo­ nal gradient corresponding to geographic loca­ Stands from the dry half of the dataset were tion (Fall-line, Atlantic and Southern Subxeric reordinated (Fig. 4). Again, a strong moisture gradi­ Longleaf Woodland). The final gradient is a geo­ ent is evident with the extremely xeric sites of coarse, graphic one of proximity to the coast. The fall­ well-drained sands concentrated in the lower right line sandhill samples occur at one extreme and and the mesic, more fertile sites with finer-textured the maritime fringe at the other, with the regu­ soils clustered in the upper left. A middle range of lar coastal plain samples in between. This is par­ moisture conditions occurs between the xeric and ticularly apparent among the more xeric samples mesic sites, corresponding well to the samples we where we recognize three types (Fall-line, Atlan­ initially characterized as subxeric. Orthogonal to the tic and Atlantic Maritime Xeric Longleaf Wood­ land). moisture axis is a latitudinal gradient, which sepa­

51 The first two axes of the wet-site ordination The four vegetation series and twenty-three (Fig. 6) together separate the flatwood sites (upper vegetation types extracted based on the above left) characterized by somewhat shrubby under­ analysis are listed in Table 1. In the following sec­ story vegetation and soils somewhat less sterile and tions, generalized descriptions and discussions are clayey than those of the herb-dominated savanna provided for each series. At the end of the discus­ sites. The "Southern," "Atlantic" and "Fall-line" sion of each series we provide summary sections Longleaf Savanna Woodlands segregate into three that list the dominant (high cover and high fre­ groups, reinforcing the significant differentiation quency) and most abundant (numerous individu­ with geographic position. als and high constancy, i.e. high between-stand frequency) species for each community type.

o

• + •

Figure 3. Ordination 1 contains all samples used in the final analysis. The first axis corresponds primarily to a moisture gradient with Xeric Longleaf Woodlands on the left, and moist savannas and flatwoods on the right. Among the moist sites, the Southern Longleaf Savannas (<) ) are most distinctive and extreme, but Atlantic Longleaf Savannas (.) separate from the Fall-line Longleaf Seepage Savannas (0) and Atlantic Longleaf Flatwoods (*). While Southern Xeric Longleaf Woodlands (.) are well separated, the Atlantic (+) and Fall-line Xeric Longleaf Wood­ lands (+) are inter-mixed with the Subxeric Longleaf Woodlands ( !). Piedmont/Upland Subxeric Longleaf Woodlands ( 0) are mixed with the Mesic Longleaf Woodlands ( .).

o

S2 Southern Subxeric Piedmont o 0 " Uplands " "

Fall-line Xeric

Figure 5. Ordination 3 resulted after reordinating the dry site dataset after removal of the Southern Xeric Longleaf, Subxeric Longleaf-Palmetto, and Southern Mesic Longleaf sites (defined by the bold lines in Figure 4) and the single Serpentine Subxeric Longleaf sample. A moisture gradient runs from the Xeric Longleaf Woodland sites in the lower right (0 = Fall-line; _ = Atlantic; <,1 = Atlantic Maritime) to the Fall-line Mesic (.) and Piedmont/Upland (0 ) sites in the upper left. The perpendicular gradient is principally geographic with Fall-line Slope (I) and Subxeric sites (*) in the lower left and Southern (....) and Atlantic Subxeric sites (,' ) in the upper right. At this pOint the various Xeric and Subxeric types are distinct and separate from the Mesic sites.

* ** * *"" * * • * * Atlantic Savannas * ** "*. * ** !It * Atlantic Flatwoods* * • * •• * Southern Savannas * .11<" *' • • ~ 0 0 00 * 00 * * .:. 0 * 0 • • • 0 .. Fail-line • ... Seepage Bogs o " " " o Fall-line " o Seepage Savannas "" o o

Figure 6. Ordination 5 illustrates the patterns of similarity among the Seasonally-wet Longleaf Woodlands. The more poorly-drained, nutrient-deficient Longleaf Savanna sites occur in the lower right (0 = Southern; • =Atlantic), while the somewhat more nutrient-rich, better-drained Longleaf Flatwood sites (*) occupy the upper left. The diagonal axis is largely one of distance from the coast with the fall-line sites at the bottom of the diagram (A= Seepage Bogs; 0= Seepage Savannas).

53 Xeric Longleaf Pine Woodlands tered shrubs (typically Myrica cerifera, Gaylussacia dumosa and Vaccinium spp.), and a sparse to mod­ The five communities ~hat comprise the Xeric erate cover of herbs and grasses can be expected Longleaf Woodlands all occur on deep, coarse, ex­ throughout. The grass layer of Xeric Longleaf cessively drained sands. These sites typically oc­ Woodlands usually is dominated by wiregrass cur on summits and shoulders of rises. The more (Aristida stricta north of the Congaree-Cooper River extreme xeric sites are associated with dune sys­ system of Sc, A. beyrichiana to the south), though tems such as occur on the east sides of Carolina these species are largely absent from central South bays (i.e., northeast of the primary axis of the de­ Carolina and from much of the Gulf coastal region pression) and along northeastern sides of large riv­ (see Fig. 1; Peet 1993). Bare sand typically is ers that flow into the Atlantic (e.g., Altamaha, Cape present at the soil surface, and species richness Fear, Pee Dee, Savannah; see Bozeman 1971, tends to be low. Christensen 1979, 1988). In addition, remnant old barrier island systems scattered across the outer Fall-line Xeric Longleaf Woodland can be coastal plain (Dubar et al. 1974) typically support found anywhere in the uplands of the fall-line Xeric Longleaf Woodlands. sandhills where soils originate from coarse, well­ drained sands (Christensen 1988, Stout and Marion Longleaf pine is widely scattered in the xeric 1993). The Atlantic and Southern Xeric Longleaf communities, and, owing to the extreme edaphic Woodlands can occur throughout the Atlantic and conditions, may not regenerate readily after cutting Gulf coastal plains respectively, though coarse or extended fire suppression. Only on the outer­ sands are more frequent close to the coast and most coastal plain of Georgia does longleaf cease along northeast sides of major rivers (see Bozeman to be the dominant species of the dry sand ridges 1971). Along the Gulf Coast flats there is a general (Bozeman 1971). Typically, there is a broad-leaved, soil-texture gradient such that the more western deciduous subcanopy with turkey oak (Quercus sites have siltier, less sandy soils. As a conse­ laevis) virtually ubiquitous and persimmon quence, Southern Xeric Longleaf Woodlands are (Diospyros virginiana) as a common associate. On more common and better developed in Florida somewhat finer-textured soils, bluejack oak (Q . than in coastal Mississippi or Louisiana. incana) also can be important. In addition, scat­

Tab!e 1. Longleaf pine Comm~ni!y Series and Types recognize~ in this study. Ad.ditional, undescribed communiti.es for which preliminary infor­ mation suggests recogmllon Will likely be necessary when suffiCient data are available are listed In parentheses Immediately after the commu­ nity within which they are currently included. . . .

Xeric Longleaf Pine Woodland Series Mesic longleaf Pine Wo,(ldland Series

Fall-line Xeric Longleaf Woodland Fall-line M~~i.c Longleaf Woodland Atlantic Xeric Longleaf Woodland Fall-line Slope Mesic Longleaf Woodland Southern Xeric Longleaf Woodland Atlantic Mesic longleaf Woodland Atlantic Maritime Longleaf Woodland Southern Mesic Longleaf Woodland Gulf Maritime Longleaf Woodland (Coosa Mesic Longleaf Woodland)

Subxeric longleaf Pine Woodland Series Seasonally-Wet longleaf Pine Woodland Series

Fall-line Subxeric Longleaf Woodland Fall-line Longleaf Seepage S~vanna Atlantic Subxeric Longleaf Woodland Fall-line Longleaf Seepage Bo,g Southern Subxeric Longleaf Woodland Atlantic Longleaf Savanna ' (Southern Clayhill Subxeric longleaf Woodland) Southern Longleaf Savanna (longleaf-Sand Pine Woodland) Southern Longleaf Seepage Savanna (Florida Subxeric Longleaf Woodland) Atlantic Longleaf Flatwood Subxeric Longleaf Saw Palmetto Woodland Southern Longleaf Flatwood Piedmont/Upland Subxeric Longleaf Woodland Piedmont Longleaf Flatwood (Upland Subxeric Longleaf Woodland) (Piedmont Subxeric Longleaf Woodland) (Fall-line Clayhill Subxeric Longleaf Woodland) Serpentine Subxeric Longleaf Woodland

54 Vegetation immediately adjacent to both the Atlantic Xeric Longleaf Woodland (Figs. 7, 8). Atlantic and Gulf coasts experiences less extreme Dominant species include Pinus palustris, Quercus climatic conditions, with the consequence that most incana, Q. laevis, and Aristida stricta. Other common sites support closed forest. However, a distinctive species are Gaylussacia dumosa, Vaccinium tenellum, Maritime Longleaf Woodland can develop on bar­ Cnidoscolus stimulosus, Schizachyrium scoparium, Eu­ rier islands and other near-coastal dunes where phorbia ipecacuanhae, Asclepias humistrata, Ionactis deep, coarse sands occur. Unfortunately, we have linariifolius, Aster. tortifolius, Aureolaria pectinata, and quantitative data only from southern North Caro­ Pityopsis graminifolia. The driest sites often contain lina. Scattered barrier island longleaf populations Selaginella arenicola, Minuartia caroliniana and occur from northern North Carolina near Nags Stipulicida setacea. Head south to at least Cumberland Island, Geor­ gia (Hillestad et al. 1975, Wentworth et al. 1992). Southern Xeric Longleaf Woodland. Domi­ The Longleaf Woodland communities sampled nant species include Pinus palustris, Quercus laevis, along the North Carolina coastal fringe contain sig­ Q. incana, Sporobolus junceus, and Licania michauxii. nificant amounts of sand live oak and sand laurel Other common species are Diospyros virginiana, oak (Quercus geminata, Q. hemisphaerica), whereas Serenoa repens, Aristida beyrichiana, Cnidoscolus Clewell (1971) reports sand live oak to co-occur stimulosus, Eriogonum tomentosum, Pityopsis with myrtle oak (Q. myrtifolia) near the graminifolia, Yucca filamentosa, and Croton Apalachicola National Forest, Florida. Personal argyranthemus. Many species in this community, observations of this community type near Santa such as Ceanothus microphyllus, Asimina angustifolia Rosa on the Florida Gulf coast suggest a quite dif­ and A. obovata, Baptisia lecontei, Berlandiera ferent community from the Atlantic type; saw pal­ subacaulis, Aeschynomene viscidula, Rhynchosia metto (Serenoa repens), false rosemary (Conradina cytisoides, Palafoxia integrifolia, Chapmannia floridana, canescens), and gallberry (Ilex glabra) share domi­ Matalea pubiflora, Phoebanthus grandiflorus, Liatris nance in the shrub layer, together with such spe­ chapmanii, and Andropogon floridanus do not occur cies as American olive (Osman thus americanus), in the mid-Atlantic states. While these species are gopher apple (Licania michauxii), shiny blueberry never abundant in the Southern Xeric Longleaf (Vaccinium myrsinites) and numerous herbs. Fur­ Woodland, their presence makes it floristically ther information can be found in Harper (1914b) quite different from the Atlantic type. and Wolfe et al. (1988). Although we have no quantitative data from this type as it is represented Atlantic Maritime Longleaf Woodland. in Florida and the adjacent Gulf Coast states, our Dominant species include Pinus palustris, Quercus preliminary information suggests that both an At­ geminata, Q. hemisphaerica, Myrica cerifera, Persea lantic and a Gulf Maritime Longleaf Woodland borbonia, Sassafras albidum, Ilex opaca, and Aristida should be recognized. stricta. Other common species are Quercus laevis, Q. incana, Osmanthus americanus, Gaylussacia As with virtually all longleaf communities, fire dumosa, Vaccinium arboreum, V. tenellum, Smilax is required in Xeric Longleaf Woodlands for regen­ auriculata, Andropogon virginicus, Stipulicida setacea, eration of many of the component species, and for Cnidoscolus stimulosus, Euphorbia ipecacuanhae, suppression of broadleaved understory tree spe­ Asclepias humistrata, Bulbostylis capillaris, and cies, particularly turkey oak (Quercus laevis). Al­ Dichanthelium consanguineum. though frequent, low-intensity surface fires once were common in this community, the low fuel load Gulf Maritime Longleaf Woodland. Quanti­ would have restricted the frequency and intensity tative data are not available (see Harper 1914b, of fire relative to other longleaf types (see Clewell 1971, Wolfe et al. 1988). Christensen 1988, Frost 1993, Stout and Marion 1993). Subxeric Longleaf Pine Woodlands

Fall-line Xeric Longleaf Woodland. Dominant Subxeric Longleaf Woodlands, particularly the species include Pinus palustris, Quercus laevis, and Atlantic and Southern Subxeric Longleaf Wood­ Aristida stricta. Other common species are lands, dominated the presettlement landscape of Gaylussacia dumosa, Stipulicida setacea, Cnidoscolus most of the southeastern coastal plain (see Ware et stimulosus, Minuartia caroliniana, Euphorbia al. 1993). They occurred on most well-drained up­ ipecacuanhae, Asclepias humistrata, Aureolaria land sites, except for the extreme coarse sands oc­ pectinata, Bulbostylis capillaris, Carphephorus cupied by the Xeric Longleaf Woodlands. The soils bellidifolius, Chrysopsis gossypina, and Pityopsis underlying these sites, while generally infertile and graminifolia. containing a significant amount of sand, typically

55 have a greater content of silt and clay than do those though in the westernmost portion of its range of the Xeric Longleaf Woodlands, a pattern recog­ wire grass is largely restricted to the coastal tier of nized early on by Wells and Shunk (1931). counties. Those subxeric sites remaining for study Throughout the coastal plain and the fall-line are predominantly on coarse-textured soils, the sandhills, the general aspect of Subxeric Longleaf siltier soils of the inner coastal plain having long Woodlands is one of widely spaced pines with a ago been converted to agriculture (except for lim­ sparse, broad-leaved deciduous understory and a ited areas of Alabama, Mississippi and Louisiana). continuous, well-developed ground layer contain­ The original vegetation of such fine-textured soils ing a lush and diverse assemblage of grasses and perhaps will remain forever unknown. However, forbs. The few remnants we have been able to the greater abundance of bluestems (Andropogon sample have contained a large number of legume spp., Schizachyrium scoparium, and s. tenerum in the species. South) on the small patches on loamy soils that re­ main suggests that these grasses rather than The differences between the Atlantic and wiregrass may well have been the original ground­ Southern Subxeric Longleaf Woodlands can be seen layer dominants (Frost, Walker and Peet 1986). most readily in the ranges of the dominant species. West of the range of wiregrass in Mississippi and In the more northern type, the dominant grass is Louisiana, and most of central Alabama, bluestems the Carolina wiregrass (Aristida stricta), whereas in remain the most abundant grasses of the subxeric the Southern examples the dominant grass is pre­ pinelands (see Grelen and Duvall 1966), as is the dominantly the southern wiregrass (A. beyrichiann). case in central South Carolina, between the ranges Low blueberry species typically are abundant in of the two wiregrasses. both types, but Vaccinium crassifolium is essentially restricted to the northern variant, while V. Regional vegetation descriptions (e.g., Harper myrsinites barely enters South Carolina. 1906, 1943, Myers 1990, Schafale and Weakley 1990) often contrast two forms of what we call The broad-leaved understory generally is com­ Subxeric Longleaf Woodlands, corresponding to prised of scattered shrubby oaks, often including whether the underlying soil is predominantly sand bluejack, turkey and sand post oak (Quercus incana, or clay. Soil texture doubtless is important for ex­ Q. laevis, Q. margarettiae) on the sandier sites, post plaining compositional variation in these wood­ oak (Quercus stellata) on the clay hills of the inte­ lands, though the overriding importance of latitude rior Gulf coastal plain, and blackjack (Q. and moisture largely mask the importance of tex­ marilandica) throughout where the clay content is ture in the ordination analysis we present. A sepa­ particularly high. Persimmon (Diospyros virginiana) rate analysis of just the Fall-line Subxeric is also common. From Jackson County, Mississippi Woodlands revealed a strong gradient in soil tex­ eastward, wiregrass (Aristida beyrichiana, then A. ture with the siltier soils having greater herb diver­ stricta) dominates much of the grassy herb layer, sity, and the clayhills having a less well developed understory than the sandhills.

Figure 7. Atlantic Xeric Longleaf Woodland. The xeric extreme of longleaf vegetation on the Atlantic coastal plain is found on the eolian dune sands along the northeast sides of Carolina bays and major rivers. Salters Lake, Bladen County, North Carolina.

56 Figure 8. Atlantic Xeric Longleaf Woodland. This old­ growth stand of longleaf is typi­ cal of xeric sites in the spareness of its wiregrass (Aris/lda s/ric/dj and the pres­ ence of turkey oak (Quercus /aeviSj. Croatan National For­ est , Carteret County, North Carolina.

Although longleaf and wire grass dominate the mont and southern-most portions of the interior visual aspect of the Subxeric Longleaf Woodlands, uplands (see Fig. 2). Except for central Alabama, the herb layer can be impressively rich in species. longleaf habitats probably were always relatively Any inexperienced botanist attempting to catalog uncommon on these upland sites, and little remains the important forb species seems destined to be­ of this Piedmont/ Upland Longleaf Woodland be­ come lost in a confusing, though fascinating, col­ cause of the longer history of fire suppression in lection of trifoliate legumes and Asteraceous basal the piedmont and mountain regions. As a conse­ rosettes. Like other longleaf communities, the quence, much of the original diversity of this type Subxeric Longleaf Woodlands are fire-adapted, has been lost, and much of what remains is de­ with frequent, low-intensity, growing-season fires graded. We have lumped these various upland required to control understory hardwoods. In the types together, fully aware that further differentia­ absence of fire, oaks and other hardwoods quickly tion probably will be required if additional data assume dominance with the consequence that most ever become available. Some indication of their of the understory herbs of the open pinelands are original diversity can be found in Mohr's (1901) lost, as is the bulk of the wiregrass and virtually and Harper's (1943) summaries of the forests of all longleaf regeneration. Alabama, in which they discuss the longleaf forests of both the piedmont and the interior uplands. A particularly distinct form of longleaf vegeta­ tion found on well-drained sandy flatlands along Central Alabama has always contained the the Gulf Coast is the Sub xeric Longleaf - Saw Pal­ most extensive examples of the Piedmont/Upland metto Woodland. Identified by Pessin in 1933 as Longleaf Woodland (see Mohr 1901, Harper 1943, Xerophytic Coniferous Forest, and also recognized Golden 1979). Indeed, in Alabama longleaf origi­ by Allen (1956), this little-known community of the nally extended to an elevation of greater than 600 coastal flatlands from southeast Mississippi east to meters. southeast Georgia and south into central Florida is distinctive in appearance because of an almost con­ A few piedmont populations also remain in tinuous cover of saw palmetto (Serenoa repens), the Uwharrie Mountains of the North Carolina (see punctuated with scarlet balm (Calamintha coccinea) Schafale and Weakley 1990, Frost 1993). We recog­ and a scattering of other herb and shrub species. nize as closely related the vegetation of sheltered The best examples known to the authors are lo­ rocky slopes of the coastal plain where the flora cated in the DeSoto National Forest in Mississippi. shows close affinities with the clayier soils of the piedmont. Typically, such slopes occur where iron­ The original range of longleaf-dominated veg­ cemented sandstones formed over impermeable etation extended beyond the coastal plain onto the clay layers and now are evident at the surface due generally drier and clayier soils of the lower pied- to erosion of the overlying sands. This community

57 has moderately spaced pines in the canopy, with a Rhus copallinum, Aristida stricta, Andropogon spp. , scattered understory of oaks, a variable shrub layer, Schizachyrium spp., Pityopsis graminifolia, Solidago and a sparse to moderate herb layer. Typical un­ odora, and Toxicodendron pubescens. Other common derstory associates include black gum (Nyssa species are Quercus incana, Q. margarettiae, sylvatica), sparkleberry (Vaccinium arboreum), black­ Gaylussacia dumosa, Vaccinium tenellum, Liatris spp., jack oak (Quercus marilandica) and mountain laurel Ionactis linariifolius, Baptisia cinerea, Carphephorus (Kalmia latifolia) . Some examples are patchy, con­ bellidifolius, Cirsium repandum, Cnidoscolus taining small, grassy openings dominated by stimulosus, Coreopsis major, Dichanthelium ovaIe, bluestems (Andropogon spp.) in the middle of open Silphium compositum, Smilax glauca, and Tephrosia forest. The region of upper Clay Hills of Alabama virginiana. (sensu Hodgkins 1965), and sometimes the clay hills in the Carolina Fall-line Hills (Fenneman Atlantic Subxeric Longleaf Woodland. Domi­ 1938), appears quite distinct from the rest of the nant species include Pinus palustris, Quercus laevis, Subxeric Longleaf Woodlood (see Mohr 1901, Q. margarettiae, Q. incana, Q. marilandica, Vaccinium Beckett and Golden 1982) and is perhaps worthy arboreum, Vaccinium fuscatum, Gaylussacia dumosa, of recognition as a separate type. However, for the Rhus copallinum, Diospyros virginiana, Aristida stricta, lack of quantitative data, we tentatively include Schizachyrium scoparium, and Andropogon ternarius. these sites with the Piedmont/Upland Longleaf Other common species are Ionactis linariifolius, Woodlands. Hedyotis procumbens, Pityopsis graminifolia, Rhynchosia reniformis, Rhynchospora grayi, Solidago At Burke Mountain in Columbia County, Geor­ odora, Lechea spp., Stillingia sylvatica, Stylisma patens, gia, a particularly unusual vegetation type has de­ Cnidoscolus stimulosus, Desmodium spp., Lespedeza veloped over serpentine rock, which we designate spp., Mimosa quadrivalvis, Tephrosia spp., and as Serpentine Sub xeric Longleaf Woodland. The Pteridium aquilinum, although not all of these spe­ naturally droughty conditions of the soils associ­ cies are found throughout the range of the commu­ ated with serpentine (Whittaker 1954) probably ac­ nity. count for the occurrence of longleaf there (and pine in general on eastern North American serpentine). Southern Subxeric Longleaf Woodland Given that most serpentine soils support unusual, Dominant species include Pinus palustris, Quercus often disjunct plant species, it is not surprising that laevis, Q. margarettiae, Q. incana, Q. marilandica, Q. the Burke Mountain sample stands out as different falcata, Q. pumila, Vaccinium arboreum, V. elliottii, from all our other longleaf samples. We expect that Diospyros virginiana, Ilex vomitoria, Hypericum similar vegetation occurred on those few other sites hypericoides, Aristida beyrichiana, Aster tortifolius, with serpentine-like substrate, but we know of no Baptisia lanceolata, Dichanthelium ovaIe, Galactia other extant examples within the range of longleaf. regularis, Rhynchosia reniformis, Lespedeza repens, Pteridium aquilinium, Smilax bona-nox, Stylisma pat­ Our vegetation samples from peninsular ens, and Gelsemium sempervirens. Other common Florida are extremely restricted and insufficient for species are Vaccinium fuscatum, Ionactis linariifolius, construction of even a preliminary classification. Hedyotis procumbens, Pityopsis graminifolia, Nonetheless, published compositional data make Gymnopogon ambiguus, Rhynchospora grayi, Solidago clear that what is commonly known as "high pine" odora, Lechea paniculatum, Desmodium ciliare, Mimosa in north-central Florida is similar to our Southern quadrivalvis, and Tephrosia virginiana. Subxeric Longleaf Woodland (e.g., Sellards et al. 1915, Laessle 1942, Myers 1990, Stout and Marion Subxeric Longleaf - Saw Palmetto Woodland. 1993). What is less clear is whether there is suffi­ The dominant species in the two Mississippi sites cient longleaf - sand pine transition to justify rec­ are Pinus palustris, Quercus laevis, Q. incana, Q. ognition of this as a separate community. Similarly, marilandica, Cornus florida, Serenoa repens, Ilex the infrequently described scrubby flatwoods (see vomitoria, Vaccinium fuscatum, V. elliottii, V. Laessle 1942, Abrahamson and Hartnett 1990, Stout stamineum, Calamintha coccinea, Smilax pumila, and Marion 1993) also may occasionally be domi­ Schizachyrium scoparium, Galactia regularis, Pityopsis nated by longleaf, but the information available to graminifolia, Rhynchosia cytisoides, and Cyperus us currently is insufficient to justify recognition of retrofractus. Other common species are Aristida a longleaf-dominated form of this community. purpurascens, Ionactis linariifolius, Chamaecrista nictitans, Cnidoscolus stimulosus, Dalea pinnata, Fall-line Subxeric Longleaf Woodland (Figs. Desmodium strictum, Elephantopus elatus, Gaura 9, 10). Dominant species include Pinus palustris, filipes, Hedyotis procumbens, Hypericum hypericoides, Quercus laevis, Q. marilandica, Diospyros virginiana, Lechea spp., Opuntia humifusa, Dichanthelium

58 Figure 9. Fall-line Subxeric Longleaf Woodland. Frequently burned Subxeric Longleaf Wood­ lands typically have a well-devel­ oped sward of wiregrass (Aris/ida s/ric/ei) , punctuated with scattered small oaks (here turkey and bluejack oak; Quercus laevis, Q incana) , huckleberries (Gaylussacia spp) and bracken fern (P/eridium aquilinurrf). Nu­ merous herbaceous species can be found growing between the wiregrass clumps. Sandhills Gamelands , Scotland County, North Carolina. aciculare, Quercus faIcata, Sassafras albidum, Scleria sempervirens. In the Uwharrie National Forest in spp., Stylisma patens, Tephrosia chrysophylla, and the North Carolina piedmont, dominant species Toxicodendron pubescens. include Pinus palustris, Quercus marilandica, Nyssa sylvatica, Oxydendrum arboreum, Pinus virginiana, Piedmont/Upland Subxeric Longleaf Wood­ Pinus echinata, Quercus prinus, Vaccinium tenellum, land. Dominant species and other common spe­ Andropogon spp. and Schizachyrium spp. Other com­ cies vary significantly across the geographic range mon species are Diospyros virginiana, Pteridium of the community. An occurrence on the aquilinum, Dichanthelium spp., Pityopsis graminifolia, Oakmulgee District of the Talladega National For­ Tephrosia virginiana, and Solidago odora. At est in Alabama is dominated by Pinus palustris, Sugarloaf Mountain Recreational Area, Sandhills Nyssa sylvatica, Vaccinium arboreum, Kalmia latifolia, National Wildlife Refuge, South Carolina, domi­ Pteridium aquilinum, and Tephrosia virginiana. Other nant species include Pinus palustris, Pinus common species include Aster tortifolius, virginiana, Kalmia latifolia, Vaccinium arboreum, and Andropogon spp., Smilax glauca, and Gelsemium Vaccinium crassifolium. Other common species are

Figure 10. Fail-line Subxeric Longleaf Woodland Subxeric sites with significant quantities of silt or clay in the soil often support a well-developed de­ ciduous subcanopy, here in­ cluding turkey oak (Quercus /ae vis) , sand postoak (Q. margarehei), and pale hickory (Carya pallidei) . Fort Bragg, Hoke County, North Carolina.

59 Aronia arbutifolia, Asplenium platyneuron, Pityopsis ing between 100 and 140 vascular plant species per 2 graminifolia, Aristida stricta, Gelsemium sempervirens, 1000 m , these communities appear richer in spe­ and Pyxidanthera barbulata. cies at this scale than any other communities known from temperate North America (Peet et al. Serpentine Subxeric Longleaf Woodland. 1990). Dominant species include Pinus palustris, P. echinata, Quercus marilandica, Schizachyrium We examined two samples of Mesic Longleaf scoparium, and Calamintha georgiana. Other com­ Woodland from the Carolina fall-line sandhills that mon species are Baptisia alba, Chrysops is mariana, appear strikingly different from the other mesic Centrosema virginianum, and Gelsemium samples. These occurred on cool, steep, somewhat sempervirens . north-facing slopes in the buffer zone surrounding the Fort Bragg, NC artillery range where hot sum­ mer fires have been a regular occurrence for many Mesic Longleaf Pine Woodlands decades. Particularly unusual is the occurrence of such mountain zone species as mountain laurel Mesic Longleaf Woodlands generally differ (Kalmia latifolia) and galax (Galax urceolata) beneath from other longleaf communities in that they oc­ a relatively open canopy of longleaf with a few cur on moderately well-drained, often rolling up­ scattered blackjack oak (Quercus marilandica). We lands, but have relatively fertile, fine-textured, know of no other place where steep, cool, north­ usually loamy soils. Many of the examples sampled facing slopes retain an open, fire-maintained veg­ in the fall-line sandhills occurred on alluvial ter­ etation (though Kalmia latifolia occurs with some races. While appropriate upland soils exist in the regularity on the sandhill variant of the Piedmont/ fall-line region, virtually all of these sites have been Uplands Longleaf Woodland described earlier). cleared for agriculture, or have been fire sup­ Certainly this type was never common, and would pressed sufficiently long that the natural under­ have been among the first to be lost with a decline story long ago disappeared. We did not succeed in fire frequency. We only tentatively recognize the in finding data from any Atlantic coastal plain ex­ Slope Mesic Longleaf Woodland as a natural veg­ amples (Atlantic Mesic Longleaf Woodland), etation type since it may be largely an artifact re­ though appropriate soils are relatively common sulting from exceptionally high fire frequency. and we have recently seen two extant sites in Robeson County, NC. Most of these areas already Longleaf-dominated communities once oc­ had been converted to agriculture two centuries curred in the Coosa Valley at the southern end of ago. The few remaining areas are mostly fire sup­ the Ridge and Valley Province in Cherokee and pressed because they are isolated pockets in an ag­ Etowah counties, Alabama, and Floyd County, ricultural mosaic. The best and most numerous Georgia (Mohr 1897, Harper 1943, Wharton 1978). remaining examples of Mesic Longleaf Woodland While these communities are known only from his­ occur in the rolling hills of the Gulf coastal plain toric accounts, mesic, valley-bottom stands of and are classified here as Southern Mesic Longleaf longleaf probably were at one time abundant. Woodland. Finer-textured, loamy soils are more However, most of the lands likely to have con­ abundant in this region, and the conversion to ag­ tained this community were inundated by con­ riculture did not start as early or proceed as quickly struction of the Weiss Reservoir, while all other as on the Atlantic coastal plain (Frost 1993). occurrences of the community apparently have been destroyed by agriculture or development Mesic Longleaf Woodlands that have contin­ (Wharton 1978). If an example of the mesic ued to experience frequent fires are generally domi­ longleaf forests of the Coosa Valley were to be nated by sufficiently dense canopy pines that the found, a new community type might well need to individual trees are nearly in contact with each be recognized. other. Favorable growing conditions certainly would cause this vegetation, in the absence of fire, Fall-line Mesic Longleaf Woodland (Fig. 11). to quickly succeed to deciduous forest (see Veno Dominant species include Pinus palustris, Quercus 1976). The understory typically is lush, sometimes marilandica, Q. la evis, Q. margarettiae, Diospyros bordering on rank, with abundant herb species virginiana, Rhus copal/inum, Gaylussacia dumosa, mixed among the bluestem grasses (Schizachyrium Vaccinium tenellum, Schizachyrium scoparium, spp. and Andropogon spp.) and wiregrass (Aristida Aristida stricta, Ionactis linariifolius, Aster walteriana, stricta, A. beyrichiana). Particularly striking is the Eupatorium rotundifolium, Iris verna, Lespedeza repens, species-richness, and especially the legume-rich­ Pityopsis graminifolia, Solidago odora, Tephrosia ness, of the herb layer. With species counts rang­ virginiana, Toxicodendron pubescens, and Pteridium

60 Figure 11. Fall-line Mesic Longleaf Woodland. Mesic Longleaf Woodlands are rela­ tively rare today because sup­ pression of fire results in quick succession to dominance by shrubs and broad-leaved trees. Some of the best re­ maining examples are found on military bases in and adja­ cent to artillery ranges where hot summer fires are assured, and unexploded ordinance provides protection from devel­ opment. McPherson Danger Area, Fort Bragg, Hoke County, North Carolina. aquilinum. Other common species include Aster fuscatum, Gaylussacia dumosa, Ilex glabra, concolor, A. tortifolius, Desmodium linea tum, Schizachyrium scoparium, S. tenerum, Andropogon Eupatorium album, Euphorbia curtisii, Lespedeza gerardii, Andropogon ternarius, Aristida purpurascens capitata, Smilax glauca, Stylosanthes biflora, and many var. virgata, and Pteridium aquilinum. Aristida more. beyrichiana may dominate within its range. Other conunon species include Aletris aurea, Polygala nana, Fall-line Slope Mesic Longleaf Woodland. Eupatorium rotundifolium, E. semiserratum, Dominant species include Pinus palustris, Quercus Onosmodium virginianum, Gymnopogon ambiguus, G. marilandica, Diospyros virginiana, Nyssa sylvatica, brevifolius, Cn idoscolus s tim ulosus, Paspalum Kalmia latifolia, Gaylussacia dumosa, G. frondosa, setaceum, Dichanthelium spp., Stylosanthes biflora, Lyonia mariana, Vaccinium tenellum, Epigaea repens, Desmodium lineatum, Aster tortifolius, Pityopsis Aristida stricta, Schizachyrium scoparium, and Smilax graminifolia, Euphorbia corollata, Tragia urens, rotundifolia. Other common species include Stillingia sylvatica, Rhynchosia reniformis, Croton Oxydendrum arboreum, Carphephorus bellidifolius, argyranthemus, Carphephorus odoratissimus, Gentiana autumnalis, Hypericum hypericoides, Helianthus angustifolius, Hieracium gronovii, Pityopsis graminifolia, and Myrica cerifera. Hypericum hypericoides, and H. stans, among many others. Atlantic Mesic Longleaf Woodland. No quan­ titative data are available yet. However, the domi­ nant species probably include Pinus palustris, Seasonally-Wet Longleaf Pine Quercus stellata, Q.falcata, Quercus nigra, Liquidam­ Woodlands bar styraciflua, various shrubs, A ristida stricta, Schizachyrium scoparium, and Pteridium aquilinum. Poorly to moderately drained pinelands are Other conunon species are probably Quercus incana, common on the coastal flatlands of the southeast Quercus margarettiae, Q. marilandica, Q. pumila, Carya and are typically dominated by longleaf pine, pallida, C. alba, Ilex glabra, Gaylussacia frondosa, G. though slash pine (Pinus elliottii) will often share dumosa, Lyonia mariana, Persea palustn's, Gymnopogon or assume dominance on wetter sites from south­ brevifolius, Anthaenantia villosa, Dalea pinnata, Eu­ ern South Carolina south and across the Gulf states, phorbia corollata, Eupatorium rotundifolium, and Sol­ and pond pine (P. serotina) will assume dominance idago odora. in the wettest sites, usually those with organic soils, from Virginia to western Florida. In much of Southern Mesic Longleaf Woodland (Fig. 12). Florida and southeast Georgia, slash pine replaces Dominant species include Pinus palustris, Quercus longleaf completely on the wettest sites, thus lim­ marilandica, Quercus falcata, Q. incana, Q. iting the range of communities that we might re­ margarettiae, Diospyros virginiana, Vaccinium

61 Figure 12. Southern Mesic Longleaf Woodland. Mesic Longleaf Woodlands occur over relatively fine-textured soils and can support an ex­ traordinarily species-rich herb layer. Wade Tract, Thomas County, Georgia. fer to as longleaf types in that region (Clewell 1971, vannas contain some of the most species-rich com­ Gano 1917, Monk 1968). Westward along the Gulf munities known from temperate North America. coast in Alabama, Mississippi and Louisiana, slash pine was originally more narrowly distributed, oc­ Longleaf Savanna vegetation is most exten­ curring primarily on the edges of drainages with Sively developed on the flat terraces of the outer the flatwood and savanna lands almost exclusively coastal plain, but originally occurred throughout dominated by longleaf (Penfound and Watkins the coastal plain portion of the range of longleaf 1937). pine where drainage was restricted and fire was frequent. Nonetheless, extensive areas of savanna The vegetation of seasonally wet flatlands is appear to have been most frequent in Southeast called variously savanna or flatwoods. Within the North Carolina, and then from the Apalachicola ecological literature, the term "savanna" is used to region west along the Gulf Coast to Louisiana. describe a multiplicity of vegetation types, either Both regions have a number of endemic savanna lacking trees or containing widely spaced trees over species. For example, in the Carolina center one a well-developed grassland. In the Southeast, the finds such endemics as Dionaea muscipula, Gentiana term normally is used in the narrower sense of autumnalis, Lysimachia asperulaefolia, L. loomisii, Sol­ open, graminoid-dominated and largely shrub-free idago pulchra, S. verna, Tofieldia glabra, and the two pine woodland on seasonally-wet, oligotrophic dominant grasses Aristida stricta and Sporobolus sp. soils. Accordingly, in this treatment we use sa­ nov. (aft. teretifolius; personal communication, A. vanna to refer to seasonally-wet pinelands with Weakley). Endemics to the Gulf center include sev­ widely spaced trees on mineral soil with eral species each of Aster, Pinguicula, Sarracenia, and graminoid-dominated groundlayers, few shrubs Xyris and numerous others. A significant num­ and often an exceptionally species-rich herbaceous ber of species have disjunct ranges with occur­ layer. Flatwoods contrast with savannas in that rences in the Carolina center and again in the shrubs typically share dominance with the Florida panhandle and westward (e.g., Helianthus graminoids, or even surpass them, although shrub heterophyllus, Lilium iridollae, Parnassia caroliniana, density and size will vary with fire history. Pleea tenuifolia, Polygala hookeri, Rhynchospora breviseta, R. chapman ii, R. oligantha, Thalictrum Species counts of 40 or more per square meter cooletji). have been recorded for a number of savannas in the fall-line sandhills, the coastal flatlands of North This break in the distribution of savanna spe­ Carolina, and the lower coastal plain of MissiSSippi. cies is largely responsible for the compositional dif­ A few 100 m2 samples from the North Carolina fall­ ferences observed in our analysis which led us to line sandhills have in excess of 90 species. Thus, at distinguish separate Atlantic (Carolina and north both 1 m 2 and 100 m2 scales, the southeastern sa­ Georgia) and Southern Longleaf Savanna commu­

62 nities. Both centers appear to have distinctive in­ The fall-line sandhills and the coastal plain roll­ fertile flatland soils composed of fine clayey sands ing hills generally do not have the extensive flat that are largely absent in between. Our limited lands with impeded drainage necessary to support number of samples from the Georgia and southern true savanna. However, impermeable clay layers South Carolina coastal plain makes it difficult to are frequent in these regions and, where these lay­ know whether the phytogeographical break is ers approach the surface, seeps develop and the re­ strongest in central South Carolina as described for sulting wet mineral soils support Longleaf Seepage the Subxeric types and corresponding to the break Savannas, provided fire has been sufficiently fre­ between the ranges of the two wiregrass species, quent to keep out shrubs. These usually are simi­ or in Georgia corresponding to several of the dis­ lar to true coastal plain savannas in their species junctions listed in the previous paragraph. Our composition. Fall-line Longleaf Seepage Savanna choice of a central Georgia break must remain pro­ is best known from the Carolinas (e.g., Wells and visional until further data are available. Shunk 1931), but a couple of examples have been reported from as far west as the fall-line sandhilIs Savanna soils always are oligotrophic and sea­ of Alabama (Harper 1922). The similar Southern sonally saturated. Where a hardpan or other im­ Longleaf Seepage Savanna can be found in the permeable soil layer is present, soil conditions may coastal plain rolling hills of the Gulf states, but we be particularly xeric during drought periods. Al­ lack quantitative data for these sites. Bridges and though the texture of savanna soils can vary from Orzell (1989) have described such communities for relatively sandy to predominantly clay, the best the longleaf region west of the Mississippi River, developed and most floristically rich savannas are but descriptions are lacking for this community as invariably on finer-textured, poorly drained, soils it occurs farther east, though some mention can be (Walker and Peet 1983, Frost, Walker and Peet 1986, found in a number of more general works (e.g., Christensen 1988, Taggart 1990). Although several Eleuterius 1968, Folkerts 1982, Harper 1906, 1914a, authors recognize different forms of savannas as­ Plummer 1963). Where fall-line seepages develop sociated with clay and sand soils (e.g., Woodwell on sandier soils, often with more shrubs, we rec­ 1956, Taggart 1990, 1994), the sandier sites with sea­ ognize a separate community, the Fall-line Longleaf sonally wet soils generally clustered with Seepage Bog. This community might be viewed as flatwoods in our analysis. the fall-line analog of the flatwoods of the outer coastal plain. The wealth of showy herbaceous species of the Longleaf Savannas has attracted considerable flo­ Like "savanna", the term "£latwood" has a ristic attention, with the result that these now rela­ multiplicity of meanings and is often applied to tively rare communities are among the best known rather dry sites with abundant shrubs. We use the of the original longleaf community types (e.g., term more narrowly to refer to moist sites between Kologiski 1977, Folkerts 1982, Walker and Peet Mesic Longleaf Woodlands and Longleaf Savannas 1983, Norquist 1984, Taggart 1994). Nestled among where shrubs are moderately abundant (Le., wet­ the dominant grasses (Andropogon spp., Aristida mesic longleaf woodlands). These seasonally wet stricta and beyrichiana, Ctenium aromaticum, sites of low topographic relief differ from savannas Muhlenbergia capi/laris tricopodes, Sporobolus spp.) are in that the canopy is denser, shrubs and understory numerous basal-rosette composites (e.g., Balduina, trees are frequent, and the soil is somewhat more Bigelowia, Carphephorus, Coreopsis, Helianthus, Sol­ fertile and often sandier. Soils are often saturated idago), small sedges (e.g., Fimbristylis, Rhynchospora, during the winter and droughty during the grow­ Scleria), insectivorous plants (e.g., Drosera, Dionaea, ing season. Longleaf Flatwoods occur throughout Pinguicula, Sarra cenia, Utricularia), orchids (e.g., the range of longleaf pine in the Atlantic and Gulf Calopogon, Cleistes, Pia tan thera, Pogonia, Spiranthes ) coastal plains from North Carolina to Texas. The and lilies (e.g., Aletris, Lilium, Tofte/dia, Zigadenus) . relative abundance of shrubs on flatwood sites is Legumes are conspicuously absent from most sa­ little understood, though a somewhat higher fer­ vannas, a phenomenon noted by Gano (1917) and tility and better drainage than found in savannas Wells and Shunk (1931) and Taggart (1990, 1994). is probably important (see Christensen 1988, Stout The absence is made all the more notable by the and Marion 1993). wealth of legumes found in the mesic and subxeric community types, which is consistent with In addition to pines, hardwoods such as black Walker's (1985) and Taggart's (1990) reports of in­ gum (Nyssa bitlora), sweetgum (Liquidambar creased legume abundance on savannas that are sylvatica) and water oak (Quercus nigra) occur in better drained. flatwoods and can form a subcanopy. The shrub layer usually is well developed and dominated by

63 the same species that typically dominate bay for­ capillaris tricopodes and Sporobolus sp. nov. (aff. ests, such as sweet bay (Magnolia virginiana), red teretifolius). Other common species include bay (Persea palustris) gallberry (Ilex glabra), and titi Platanthera spp., Cleistes divaricata, Calopogon pallida, (Cyrilla racemiflora). Southward, running oaks C. tuberosus, Dionaea muscipula, Drosera capillaris, (Quercus minima, Q. pumila) often are dominant spe­ Pinguicula spp., Utricularia spp., Rhynchospora spp., cies in the shrub layer of drier flatwoods. How­ Fimbristylis spadacea, Lachnanthes caroliana, ever, from central South Carolina southward, the Lachnocaulon anceps, Xyris ambigua, X. caroliniana, characteristic species is saw palmetto (Serenoa Dichromena latifolia, Rhexia alifanus, R. petiolata, R. repens) which can at times form a solid understory lutea, Eriocaulon compressum, Liatris spp., canopy. Understory herbs are much less abundant Carphephorus paniculatus, C. tomentosus, Coreopsis than in the savannas because of the denser tree linifolia, Hypoxis spp., Dichanthelium spp., Agalinis canopy and increased competition from the shrub spp., Andropogon mohrii, Eryngium integrifolium, layer. Nonetheless, wiregrass and other plants of Eupatorium leucolepis, E. rotundifolium, Lycopodiella both Longleaf Savanna and Mesic Longleaf Wood­ caroliniana, Osmunda cinnamomea, O. regalis, Polygala land are frequent. spp., Sabatia spp., and Zigadenus glaberrimus. Bridges and Orzell (1989) and Taggart (1990) dis­ A type of longleaf vegetation occurs (or once cuss geographic differences in species composition occurred) in the eastern portions of the piedmont, of longleaf savannas. from North Carolina to Alabama (See Pinchot and Ashe 1897) with a species composition that places Southern Longleaf Savanna (Fig. 14). The it in the flatwood type. This Piedmont Longleaf most abundant species include Pinus palustris, P. Flatwood currently is known only from highly de­ elliottii, Bigelowia nudata, Carphephorus pseudoliatris, graded remnants in North Carolina that have been Chaptalia tomentosa, Coreopsis linifolia, Ctenium subjected to logging and fire suppression. The aromaticum, Helianthus heterophyllus, Ilex glabra, Lo­ community occurs on poorly drained upland flats belia brevifolia, Rhexia alifanus, Rhynchospora plumosa, that are themselves unusual in the piedmont. Little R. oligantha, Scleria reticularis, and Xyris ambigua. information is available on the original composi­ tion of this community. Remnant stands do sup­ Southern Longleaf Seepage Savanna. Quan­ port wiregrass (Aristida stricta) and creeping titative data are lacking for this community. How­ blueberry (Vaccinium crassifolium), but most of the ever, limited personal observation suggests that other original ground layer species are now gone common species of seepage savannas in southwest­ (see Schafale and Weakley 1990). ern Mississippi include Andropogon spp., Anthaenantia rufa, Aristida purpurascens virgata, Atlantic Longleaf Savanna (Fig. 13). Domi­ Cacalia ovata, Calopogon pallidus, C. tuberosus, Core­ nant species include Pinus pa/ustris, P. serotina, opsis linifolia, Chaptalia tomentosa, Ctenium Aristida stricta, Andropogon spp., Ctenium aromaticum, Eragrostis refracta, Eriocaulon aromaticum, Rhynchospora plumosa, Muhlenbergia compressum, E. dectangulare, Helianthus heterophyllus,

Figure 13. Atlantic Longleaf Savanna. Southeastern coastal plain flatlands with fine-textured, seasonally-satu­ rated soils contain among the highest small-scale species densities known from the Western Hemisphere. Where fire is frequent, the average species number can exceed 40 per square meter. Green Swamp, Brunswick County, North Carolina.

64 Figure 14. Southern Longleaf Savanna . A wealth of herba­ ceous species including numer­ ous orchids and insectivorous plants can be found in coastal plain pine savannas. Pitcher plants (Sarracenia a/a/a) and sundews (Orosera /racy~ domi­ nate in the foreground on this Gulf Coast savanna. Sandhill Crane National Wildlife Refuge, Jackson County, Mississippi.

Lachnanthes caroliana, Linum media, Lophiola aurea, South Carolina, and Quercus pumila is largely ab­ Lycopodiella alopecuroides, L. appressa, Dichanthelium sent from North Carolina. In addition, Aristida dichotomum ensifolium, Polygala lutea, Rhexia alifanus, stricta does not occur south of northern South Caro­ R. petiolata, Rhynchospora ciliaris, R. chapmanii, Sar­ lina. Other common species of Wet Longleaf Pine racenia alata, S. psittacina, Xyris ambigua, X. Flatwoods include Vaccinium crassifolium, baldwiniana, X. caroliniana, X. difformis, and Gaylussacia frondosa, Carphephorus odoratissimus, Zigadenus glaberrimus. Kalmia angustifolia, Lyonia mariana, Myrica cerifera, Cyrilla racemiflora, Pteridium aquilinum, Smilax spp., Fall-line Longleaf Seepage Savanna. The and Rhynchospora spp. most abundant species include Pinus palustris, P. serotina, Ilex glabra, Aristida stricta, Aster dumosus, Southern Longleaf Flatwood. Dominant spe­ Ctenium aromaticum, Drosera capillaris, Erigeron cies include Pinus palustris, Pinus elliottii, Myrica vermiS, Eupatorium rotundifolium, Lachnocaulon cerifera, llex glabra, Serenoa repens, and Aristida anceps, Osmunda cinnamomea, Pycnanthemum beyrichiana . Other common species are Pinus flexuosum, and Rhexia alifanus. Other common spe­ serotina, Kalmia hirsuta, Vaccinium myrsinites, Lyonia cies are Chaptalia tomentosa, Coreopsis linifolia, lucida, and Sabal palmetto (Wharton 1978, Eupatorium leucolepis, E. pilosum, Hypericum crux­ Abrahamson and Hartnett, 1990). Much variation andreae, Viburnum nudum, and Viola primulifolia. in species composition exists within this type.

Fall-line Longleaf Seepage Bog. Dominant Piedmont Longleaf Flatwood. Dominant spe­ species include Pinus palustris, P. serotina, Clethra cies in remnant occurrences include Pinus palustris, alnifolia, Lyonia lucida, Cyrilla racemiflora, Aronia Pinus taeda, Acer rubrum, Liquidambar styraciflua, arbutifolia, Ilex glabra, Arundinaria gigantea, Pteridium Gaylussacia frondosa, Lyonia mariana, Vaccinium aquilinum, Vaccinium crassifolium, and Aristida fuscatum, Ilex glabra, Vaccinium crassifolium, Panicum stricta. Other common species are Gaylussacia virga tum, Chasmanthium laxum, and Aristida stricta. frondosa, Symplocos tinctoria, Ilex opaca, Vaccinium Many species have coastal plain affinities. Other stamineum, Acer rubrum, Toxicodendron vernix, Mag­ common species include Quercus marilandica, Q. nolia virginiana, Persea palustris, Osmunda stellata, Nyssa sylvatica, Andropogon glomeratus, cinnamomea, and Woodwardia virginica. Eupatorium spp., Osmunda cinnamomea, Solidago odora, Rhynchospora spp., and Pityopsis graminifolia Atlantic Longleaf Flatwood. Dominant spe­ (Schafale and Weakley 1990). cies include Pinus palustris, P. elliottii, P. serotina, Ilex glabra, Serenoa repens, Quercus pumila, Ilex coriacea, Myrica cerifera, and Aristida stricta, although not all DISCUSSION AND CONCLUSIONS of these species occur throughout the range. For instance, Serenoa repens occurs only as far north as Although the once extensive Southeastern 65 longleaf pine woodlands may appear to the casual Glitzenstein, Cary Norquist, Rebecca Reed, John observer as a rather homogeneous expanse of Taggart, and Thomas Wentworth. longleaf pine, wire grass and scrub oak, this is de­ cidedly not the case. We have documented consid­ erable compositional variation which we have APPENDIX 1. DATA SOURCES summarized using 23 communities; we also antici­ pate a future need to recognize additional vegeta­ North Carolina Vegetation Su rvey data. We used tion types. The longleaf communities we recognize data from 69 plots sampled during June, 1989 and are largely separated along gradients correspond­ 1990 in the fall-line sandhills of North Carolina. ing to soil moisture, soil texture, and geographic The fundamental sampling unit was a 10 x 10 m region. module wherein the percent cover for each vascu­ lar plant species was recorded using the ten-point An equally important and little recognized as­ scale described in the methods section. Typically, pect of the remarkable diversity of longleaf ecosys­ a sample plot consisted of a block of 4 contiguous tems is found in the numbers of species present in modules, plus cover values of all additional spe­ individual samples. We report Mesic Longleaf cies encountered in a full 2x5 block of 10 modules, Woodlands with numbers of vascular plant species or 0.1 ha plot. Occasional plots were smaller, the per 1000 m2 ranging up to 140, the largest values smallest containing only a single 10 x 10 m mod­ yet reported for the temperate Western Hemi­ ule. In addition, we used data from four maritime sphere. Samples of 100 m2 with species counts over fringe longleaf pine communities collected as part 90 collected from Fall-line Longleaf Seepage Savan­ of a comprehensive study of barrier island mari­ nas also represent a new record for temperate time forests in May 1988 (see Wentworth et a1. North America. Finally, counts of more than 40 1992). The methods employed were identical to species per m 2 from Atlantic Longleaf Savannas those used in the fall-line sandhills study. (NC), Southern Longleaf Savannas (MS), and Fall­ line Longleaf Seepage Savannas (NC) exceed all Nature Conservancy data. Data were collected other values yet reported for the Western Hemi­ from 47 plots between April 1989 and November sphere. Many of these species are restricted to the 1990 in a study explicitly designed to provide in­ longleaf pine ecosystem. formation for refining the initial Nature Conser­ vancy classification. This study was coordinated The remarkable diversity of the greater through the Southeast Regional Office of The Na­ longleaf ecosystem is being lost rapidly, both ture Conservancy by DJA and involved ecologists through active habitat destruction and through ne­ from the regional and state offices of the Nature glect. Much habitat is being destroyed through Conservancy and the state Natural Heritage Pro­ development or conversion for greater economic grams. In this study longleaf-dominated commu­ yield. Simultaneously, much of what remains is nities were sampled in all states within the species' being lost through fire suppression, which quickly range except for Virginia, although time constraints leads to loss of many of the numerous species that did not allow all longleaf-dominated community inhabit the longleaf communities. If even a sub­ types in the initial classification to be included. stantial fraction of the diversity of the greater longleaf ecosystem is to be preserved, action must Permanent 20 x 50 m (0.1 ha) plots were estab­ be taken quickly to both preserve and manage the lished in relatively undisturbed longleaf pine com­ best remaining examples of each of the longleaf munities. Emphasis was placed on sampling sites communities. over the mesic to xeric portion of the moisture gra­ dient because fewer published data were available for these sites. As in the North Carolina Vegeta­ ACKNOWLEDGMENTS tion Survey study, quantitative data were collected from four contiguous 100 m 2 modules in each plot. For permission to use unpublished data we Cover class was recorded for each plant species in thank William Boyer, Cecil Frost, Cary Norquist, each module using the same 10-point scale, and James Snyder, John Taggart, and the members of species presence was noted for the remainder of a the North Carolina Vegetation Survey (Cecil Frost, full 0.1 ha plot. Michael Schafale, Alan Weakley, Thomas Wentworth, Peter White, and RKP). Earlier ver­ Frost data. Four plots from an unpublished sions of the manuscript benefited from the helpful data set collected by Cecil Frost as part of his doc­ comments of William Boyer, Cecil Frost, Jeff toral dissertation research were used in this analy­ sis. These data were from relatively undisturbed

66 areas of the Croatan National Forest in the outer tative set of 20 of the 40 stands that contained coastal plain of central North Carolina. Plots were longleaf pine. In each stand ten quadrats were 20 x 50 m CO.1 ha), with percent cover recorded for sampled for herbaceous data, while woody species all shrub and herb species in each of 25 0.5 x 2 m were recorded from 30 quadrats. Data originally subplots. Tree diameters were recorded and sub­ were recorded by 5 abundance classes which we sequently converted to cover using regression converted to match our ten-point scale. models developed from the North Carolina Vegeta­ tion Survey data. Norquist Data. Cary Norquist collected data from seven relatively undisturbed coastal savannas Taggart data. As part of his doctoral research, in southern Mississippi as part of her masters re­ John Taggart (1990, 1994) collected data from sea­ search (1984). Although Norquist did not record sonally-wet coastal plain savannas located between information on the sparse tree stratum, she did re­ the Congaree-Cooper river system in South Caro­ port that longleaf and slash pine (Pinus palustris, P. lina and the Neuse River in North Carolina. We elliottii) were the only important trees on any of include the 40 of his plots that contained longleaf her plots and were likely the original dominant pine. Sites were minimally disturbed; while past species (presently, the sites are dominated prima­ ditching and lumbering were allowed, soil distur­ rily by sparsely planted slash pine). Twenty 0.25 bance and prolonged fire suppression were not. m 2 quadrats were sampled at each savanna site, Tree diameters were measured in a 0.1 ha circular with presence recorded for each quadrat. plot, shrub cover values were recorded using a 6­ level scale in a 0.01 ha circular plot at the center of Snyder data . James Snyder collected extensive the tree plot, and frequency and cover of herbs data on the vegetation of the Croatan National For­ were recorded in 191m2 plots inside the shrub est on the outer coastal plain of central North Caro­ plot. lina as part of his masters research (1978, 1980). We used those 26 plots in his dataset that contained 2 Forest Service data. In a study of Florida pan­ longleaf pine. Plots were 10 x 20 m (200 m ) in size. handle sandhills vegetation, H.E. Grelen and oth­ Snyder recorded cover of each plant species using ers from the U. S. Forest Service collected data from a seven-point scale which we transformed to con­ 50 stands. Of these, we used data from a represen­ form to our ten-point scale.

67 0\ 00 APPENDIX 2: COMMUNITY COMPOSITION

This table contains the frequencies of species that occurred in the samples included in our analysis.The number of samples included is shown at the top of each column. Only species that had afrequency of at least .50 in one community, or that occurred in at least 4communities, are inciuded.The full table, induding all rare species, is available from the authors upon request. Nomenclature follows Kartesz (1994). ..., ..., ..., ..., c c c ..., ..., ..., lii .." ..., c c c c ..., ~ '8 1i '8 .." C ..., c '8 0 0 c c '5 'g ;0: .. '8 ~ ..c ~ '8 ~ c = 0 --' m 1i 0 1i ~ == 0. 1i OX> "" 1i ~ ~ ::l ~ --' 0 > "8 --' --' u --' u ~ c ;0: ~ ..... ::l ..... ' c en ~ c '8 en ~ --' ..... ~ --' 0 go u u .'" .." --' ..... --' ~ --' ..... --' -:ii ~ c --' ..... > 10 = --' ..... u ~ .'"m u u en .g .", m ~ --' ,; (J) ell'" I --' ~ >< c en c u 'E :::E ::l ::l en --' u ~ u ~ u m ~ c .= ~ ." '" c c ~ c ~ 1l i ~ ~ ~ ~ ~ ~ ~ :2 ~ ~ ~ ~ 0 ~ 1'i 1'i ~ ::i' (J) ::i' ~ ::i' J! en ~ j ~ ~ (J) en ::i' ::i' ~ ~

Sa]pe lize 11 10 28 20 14 35 33 16

AC.AlYPHA GRACILENS 0200 1.000 0.333 PCER RlIBRUM 0.050 02:Jl 0.286 0.333 0.111 0.429 0200 0.429 0.6118 AG.AJJNS RUCAUUS 0.556 AG.AJJNIS PURPURfA 0.050 0.071 0.057 0.265 AGERAnNA AROMATICA 0.074 02:Jl 0.357 0.333 .AlETR~ AUREA 0.667 0.\\1 0.088 0.143 METRlS FARINOSA 0.050 0.:00 0.333 0.200 0.647 0.571 0.1 25 METRlS LUTEA 0200 0.556 ANDRC!'CGON GERARDiI 0071 0.667 0.111 0.125 ANDROPOGON GLOMEflAlUS GlAUCOPSIS 0556 ANDROPOGON ~mRlI 0.778 ANOOOPOGON SP. 0.727 0.400 0.001 I.OOl 0.200 0.500 0.250 0.857 0556 0.429 I.GOO 0.571 0.563 /lNDROPOGON TERNAA US 0.185 0.400 0.:00 0.667 0222 MIIlROPOGON VlRGIN!:US 0. \\1 0.750 0.200 0.600 02:Jl 0.143 0.778 0.229 0.375 AN1l1AEN.ANTlI. VI LLOSA 0.037 0.050 0.400 0.667 0.029 0.059 ARlsnDA BEYRICHl>\NA 0.444 0.001 0.250 0.333 ARlsnDA PURPUflASCENS PURPURASCENS 0.037 0.050 1.000 1.000 0.125 ARlsnDA PURPURASCENS VAR. VlRGATA 0037 0.400 0.143 0.667 0.778 0.11 4 0.294 ARlSnDA STRICTA 1.000 l.eoo U()J 1.000 0500 1.000 U()) 0.823 0.824 0.571 nS75 ARISTctOCHlI. SERPENTARl>\ 0.037 D.n50 0.600 1.000 0333 ARONIA ARBU11FOO\ 0.250 0.050 0.143 0.333 0.333 0657 0.412 0.571 0.7:Jl ARU~lNARlI. GK,ANTEA 0.001 0.250 0071 0.314 0.353 0.429 0.438 ASCLE Pl>\S AMPlfXICAUUS O.WI 0.037 0.30) 0.143 0.333 ASCLEPl>\S HUMlSTflATA 0.364 0200 0.185 0250 0.050 0200 0.:00 ASCLEPl>\S TUBEROSA 0.050 0.071 0.063 ASTERACW\TLIS 0.037 0.050 0200 0600 1.000 0.111 ASTER COlmOR 0296 0.J50 0.786 0.333 0.C66 0.029 0.143 ASTER DUMOSUS 0.037 0050 02:Jl 0.643 1.000 0.a37 02&i 0.471 0.657 0.1 88 ASTER lATERIFlORUS 0.500 0200 0.500 0.333 ASTER PALLIlOSUS 0.333 0057 0.353 0.143 0.003 ASTER PATENS 0200 0.250 1.00) "C "C "C "C C C C ~ "C "C m ~ "C :a 1;1 C c c "C ~ '8 '8 ~ "C C 1;1 "C -'" ~ m m C ~ -g '8 3: c 0> m 0 ~ ~ J J!i 'g c 0 '8 3: -' 0 -' ~ -g m '8 ~ -' ! -' 0 ~ ! ~ c m ;:; -' -' ~ -' -g j ~ -' -' .~ E -' . ~ en ~ -' c c < ~ ~ ~ b'l'" c ::E j ~ -' :::I ~ ::It c en en u -' :::I Jl E ~ E ~ u u ~ u ~ u ~ '"' c .'" i c ~ c .~ .~ = ~ ~ = "E .~ :I -l5 :I ~ :I :I ~ ~ ~ -'" ~ 0 -'" 0 ~ ~ ien en ~ ~ ~ ~ ~ en ~ ~ .. ~ en ~ ~ ~ .. Sampl1!ize 11 10 28 20 14 35 33 16

ASTER PATERNUS OAoo 02:0 0.:00 0.643 0.029 0.029 0.0&'3 ASTER SE~CEUS 0.667 ASTER SOUDAG1NEUS 0.182 0.1:1.) 0.500 0.571 0.143 0.029 02M 0.313 ASTER SJRCULOSUS 1.000 0029 ASTER TORTIFOUUS 0.364 O.OCO 0.148 0.:00 1.000 1.000 1.000 0.:00 O.7M Of~7 0.171 0235 0.143 02[1) ASTER WA1.TERI 0.:00 O!iOO 1.000 0.371 0.412 0.429 02[1) AUREOLARVI PECTINATA 0.636 0.100 0.037 0250 0.250 BftlilU1NALtlRORA 0.778 O.OM 0.059 IW'llSIA ~1BA 1.000 IW'llSVI CALYCOSA V1110SA 0.444 IW'llSVI CINEREA o.m 0.700 0.643 0.029 0.059 0.125 IW'llSVI LOC£ClATA 0.074 0.800 IW'llSIA TItltTORVI 0.182 0.2()) 0.143 0.057 0.029 0.125 BARTONIA V1RGh~ 0.556 0.176 BlGELO'/IVI NLOATA 1.000 0143 0824 0.571 BOLTO~\~ DfFUSA 0.",7 o.rl2 BUlOOSlYUSCAPIUARIS 0.091 0.074 0.2:1.) 0.:00 0.063 BLOOSlYUS C1LVlTIf()IJA 0.182 0.148 0.050 0.:00 CA1.A~JNIHA COCCINEA 1.000 CALM'JNIHA GECflGlANA 1000 CAWCARPA /llIER K: ~ 0400 0.:00 0.667 CiWSlA GRAM1NEA 0273 0.:00 OJ] 0.214 CA1.0PCGON PAlillUS &BVlRBATUS 0.333 0.882 0285 CALCl'CGON TUlEROSUIS 0571 0125 CARPHEPHORUS BEW~ FOUUS 1.000 0.600 02:1.) 0.7:0 02:0 0.:00 O.seo 0.114 0.143 0.125 CAAPHEPHORUS ODOAATISSI MUS O.ll) 0.007 0.600 0:00 02:0 0.333 0.543 CARPHEPHORUS PANClAATUS 0.100 0.071 0.429 0.7(0 O.COl CARPHEPHORUS PSELOOLIATRIS IDoo CARPlfPHORUS TC+.t:NTOSUS 0314 0.:00 O.COl CARYAPUlA 0.100 0:00 0.307 1000 CARYA PAlillA 0.182 0.050 0.:00 0.500 O.COl CENIOTHUS MIERICAMJS 0.074 0.300 0200 0.214 0667 0029 0.053 CENTEUA ASIATICA 0.778 O.OM 0.147 0.286 0.063 CENTROSBM VIRG1NlAMJM 0.037 0.:00 1.000 0.667 0.029 0.029 CfWMECRISTA FASCI CULATA O.~ 1.000 0.200 0.029 CHM\l\ECR~TA NICTfTANS 0.100 1.000 0500 0.429 1.000 0.143 O.COl CHAPTAL1A TOMENTOSA 1.000 0.057 0.382 0.714 CHASlMNTHIUM lAXlIA 0.200 0.071 0.057 O.COl CHRYSOPSIS GOSSYPiNA 0.455 0.556 0.3:0 O!iOO

0\ \0 -.J 0 .., .., "C .., c c c ~ .., .., .., ~ "C c c c 'C 'C .., -'" .., c .., .., 'C ~ c ~ c c ~ ~ ;;: ~ 'C ~ ~ g ~ ~ c = 1'i ~ ;;: ~ 0 1;;: l!:. dl ;;, ~ ~ ::J 1'i ~ ~ u .2 c .., m ~ ~ 1 ~ ~ :::j .!.! ~ ;;: ~ ~ ~ g ~ = ;;, ~ i! ."m c ~ m u u .., '"~ ~ m ~ ~ ~ ~ '"> ·c ~ ::J u .. > = g. ~ ~ u ~ .g u u OK: u ~ .;;; ~ cl!l ." .. ... ~ ~ .~ '" ~ ;It i ::c ~ ~ ~ ~ c?I '" '" ~ ~ ~ >< ~ c u :!! ~ ~ c ~ Jl ~ m '" m m u m m u '" '" ." f c c m m c .""E c .""E ..,E ~ = c ~ ~ = ~ ~ ~ -'­ ~ -'" 0 -'" -'" i I ~ ;t ~

Samp~ ~e 11 10 23 ::;J 14 35 33 16

alRYSOPSIS fMRIANA 0.100 0.200 02!>! 1000 0.429 1.000 0.111 0.086 0.382 0.143 CIRSIUM HORRllutLlA 0.500 0.657 CIRSIUM REPANDUM 0818 01 00 0.750 0.571 0.063 ClflSTES DrYN'iICATA 0.0 71 0.029 0.118 0.143 0063 ClfTHRA All1lFOUA 0.250 oI!>! 02:11 0.:1)0 0286 0143 0429 0.875 CUTORIA MI,AlANA 02!>! 05".,j 0.500 0357 0.125 CNIDOSCOLUS STIMLlOSUS 1000 0.900 0.481 0.500 0.6:11 0.500 0400 1.000 0071 0007 0029 COREOPSIS UNIFOUA 0.071 1000 0.029 0794 0714 0.188 cmOPSIS MAKlR 0.031 0.500 I.O;() O.2"W 02!>! 0.357 0333 0.029 03 13 COREOFSISVERTICILLATA OO!>! 0.2!>! 0.50 0.423 0.023 0.143 CORNUS flORIDA 0.037 O.IW O.roJ 100l 0.250 0265 0.007 0063 CRATAEGUS UNlfLORA 0001 0.370 0.100 040l 0.2 14 CROTAlAAlA PURSHII 0. 148 0.050 10CI1 0:1)0 0.007 0.029 0.1 47 CROTAlARIA ROTUNDIFOlJA 0222 0. 500 0.:1)0 0.333 CROTftLARlA SAGIlTAUS 0.667 CROTON ARGYRANII\EMUS 0.852 0.4CI1 CTENlJM ARO~\I,ncll~ 0.2O:J 0.071 0333 1. 000 0.0~9 0.529 0.857 0.2!>! CYFEAIJS PlUKENETI 1000 0.:1)0 CYRIllA RACEM:FLORA 0071 0.111 0.200 0.147 0.250 DAlEA PINNATA 0.182 0444 0. 100 0.500 1000 DAtf1HONIA SERICEA 02!>! 0.643 0.029 0.265 0.063 DESMODIUMWARE 0150 0.:1)0 0200 0.250 05".,j 0333 DESMODIU\\ lAEVIGATUM 0.09 1 0.050 0.2O:J 0.214 0.333 DESI.lJDIUM UNEATUM 0.091 01:11 0500 0714 1000 0.111 0.057 0.029 0.063 DESMOIJIUM MA~WUllCUM O.I!>! 0.500 0.143 0333 0.029 DESMOOIUMOBTUSUM O.O!>! 0.500 0.071 0.333 0.029 DESMOOIJM PA}J K:LlATUM 0182 0.050 0.500 0.2O:J 02:11 0.071 0.667 DESI.lJDIUM STRICTUM 0.162 0.037 0200 1.000 0.071 0.333 0.143 0063 DESMO[JLlA TENUroJUM 0.100 0.214 0111 0257 0.176 0.286 0125 Di:HANTHEUIJMACICUlARE 0148 0.100 1000 O.a:\) 10CI1 0.357 1000 0.333 0.1 14 O.C63 DICHANTHEUUMcotllAUTATUM 0.350 02!>! 1000 0.571 0.125 IlICHANTHEUUM CCtlSANGUINEUM 0.100 0.750 0.050 0.111 0229 Ofl29 0143 DlCHANTHEUUM DICHOTOMl.I.I DICHOTOMl.I.I 0200 0.:1)0 0.250 I.Ceo 0.143 0143 0.250 DICHANTHEUuMDICHOTC!.IJMENSIFOUl.lA 02!Xl 0214 n667 0.778 0.114 0700 0.857 0313 OICHANTHEUUM IlICHOTOMl.IA TEttJE 0.100 0.357 0.333 0.143 0.250 DICHANTHEUUM lONGIUG ULATUM 0.778 OIalANrnEUUM OliGOSANTH ES O.I!>! 0200 0.143 0.063 DICHANTHEUUMOVALE 0273 0.100 0.148 0.6:11 1000 1000 02:11 0.929 lew 0147 0.143 0.375 Oi;HANllfUUI.I SAB UlORLM 0.200 0.037 0.071 0.114 O:CHANTHEUl.I.I SPHAEROCARPON 0.667 .., c .., .., ~ ~ .., ~ m -g c c c .., .., 'g m '8 .., .., c .., m '8 .., c co co m 'g ~ c m m m ~ ~ ~ ~ ~ 'g co ~ '" c =0 'g --' 0 ~ --' 'g a> =g ~ ~ ~ ~ --' --' g. .., ~ --' --' --' 0 ~ --' .g ~ '"c ~ --' --' --' E --' in ~ --' c 0 1§ ~ g. ~ --' 0 0 n; .., --' --' --' --' ." D­ c :::l --' 0 g. :::l 0 ~ ." m ~ --' l'4 ~ .§ 0 0 en ~ ~ 0 0 .'" 'g: en .'" ~ ~ .g .g en ,. --' u:: 1 ~ Jen ~ >< ~ --' --' --' >< ~ ~ en en 0 ~c co ~ ~ ::::l --' ! --' ~ 0 ~ 0 ~ 0 0 ~ ~ c 0 c .= 0 c c c c c c E c c ~ m :z m ~ m m :z ~ ~'" ~ ~ ~ ~ en~'" en~ ~ Ien ~ ~ ien ien ~ ~ ~ ~

Sa~jk~ II 10 28 20 14 35 33 16

CXCHAtllHELIUMS TRIGOSUMLElJCOBlEPHAR!S 0.51)) 0.071 0.667 0.257 0.382 0.429 0.063 DICHAtIlHELIUM STRIGOSUMS TRIGOSUM 0.071 0444 0.057 0.188 DIOtlAEA MUSCIPULA 0.588 0.063 DIOSPYROS VIRGINIANA 0.727 0.:00 0.815 0.500 O.S;) 1000 1.0IXl 0.500 1000 0.929 I.COO 0111 0.257 0.438 DROSERA BREVIFOlJII 0.071 0111 0.029 0.143 0.063 DROSERA CM'IlARkS 0778 0.OS7 0.500 0.714 0.063 DYSCHORISTE OBLCl'I3IFOUA 1.01Xl B.EPHANTOPUS ELATUS 0.074 0200 lOIXl 0.333 B.EPIWITOPUS NUDATUS 0.400 0286 0.086 0.118 0.250 B.EPHANTOPUS TOMENTOSUS 0100 0.2SO 0114 0.667 0.063 EPIG.AEA REPENS 0.354 0.500 1.000 0.143 0.063 ERIGERON STRIC-ClSUS 02"1Al 0.500 1.0IXl 0071 0.333 0063 ERKlERON I'ERNUS 0.556 0.057 0.853 0.714 0063 ERKlCAULON COMPRESSU ~j 0667 ERKlCAUlON IICANGUlARE 0778 0.2(£ 0.143 ERKlGO NLI.~ TOMENTOSLM 0.852 0.400 ERYNGILIA INTEGRlFOUUM 0.556 0.029 0.500 0571 ERYNGIUM YUCCIFOULI.~ 0200 0071 0333 0.111 0.059 EUPATOOUM /lIBUM 0.100 0.450 0.600 0.714 1000 0.029 0.029 0.125 EUPATORIUM COMPOSmFOLIUM 0.182 0.037 0.050 1000 0.200 0214 0.029 0.143 0.125 EUPATORIUM GLAUCESCENS 0.182 0150 0.071 0.003 EUPATORUM lEUCOLEPiS 0.091 0143 0.556 0.314 0.794 0.714 0.188 EUPATORIUM MOHRII 0.091 O.OSO 0.500 0057 0.143 0.375 EUPATOR!JM PWSUM 0.150 0571 0216 0.714 0.500 EUPATORIUMROMDIFOUUM 0091 O.ISO 02 00 01SO 0.857 1000 0. 111 0.400 061 8 0.857 0.875 EUPATORIUM SEMISERRATl!.~ 0.500 0.500 0.667 0. 111 EUPI1JR!M COROUlATA 0.100 0.037 0.250 0.300 0.500 0.2"C() 0.500 02SO 1.000 0.143 1.000 0.003 EUFHORBlIl CURTISIl 0.364 0222 0.400 0,250 0.786 0.2C£ 0.375 EUFHORBlIl EXSERTA 0.150 0.571 0.143 EUFHORBlIl FlOODAtIA 0.667 0.60) EUFHORBlIllPECAClJMIlIIE 1.00) 0.500 0.500 0.650 O.1V1 TEN~FOUA 1.0IXl 0,111 0.171 0.059 0.003 GALACTlIl ERECTA 0.091 0.259 0.2X1 0.400 0.571 0.667 0.003 GAlACTlIl REGUlARlS 06:l6 03(/) 0.593 0.450 1.(0) 0.600 0500 0.500 0.286 0.333 0.029 0.143 0.125 GAlACTlIl VOLUBI.IS 0.037 0.500 0200 0.071 0.667 GAlAX URCEOLATA 0.500 GALIUM HISPIDUlUM 0.050 GAlIUM PlOSUM 0.091 0.148 0.400 0.600 0.571 0.667 0063 GAMOCHAETA PURPUREA 0.091 0.050 0.143 0.029 -..J -.l N

c c =c =c =m =m m m c =c c '0 = =m m =m '8 m '8 '8 c =c c =m ~ =m =lii =lii -'" '8 ~ ~ ~ c = -g ~ ~ 0 ~ --' ;: ;: ;: --' ~ m .l'l '8 '8 ~ :::l ~ ;: ID --' 0 ~ -g m ~ ~ --' :::l :::l in ;: c = ~ ~ --' :::l ." ~ m C ID 0 0 :::l '" ~ --' --' ~ = :::l :::l 0 ~ ~ :::l --' 0 J§ ." '" m 0 0 .;;; ~ [ 0 0 J .;;; .;;; ll; "'­ --' il: Jl ~ => ~ '"--' --' 3l --' ~ ~ ~ '" --' '" --' ~ 0 ~ ~ --' --' ~ ~ '"E '" § -~ 0 ID ~ '" '"0 ." ~ c ID ID '"c ~ :§'" .~ E '" ~ ~ '" ~ ~ :;: 5: '" :!2 ~ ~ ~ ~ -'" ! .s ~ 0 ~ ::i ::i i ell 8: ~ c7l en ::i « ..u. &! &! « '" ~ '" ~ &!

Sam~~ze 11 10 28 20 14 35 33 16

GAYLLGSAClII DUMOSA 0.818 0.700 0.037 O.seD 0.750 10::\) 0.600 0.500 1.0:11 lOCO 0.667 0657 0. 3~ 0413 0625 GAYlUSSAClII FROIroSA 0.091 0.700 02SO 0.050 0.500 0.250 1.000 0.357 0.800 0.471 0.429 0.813 GELSH.IUM SEMPERI1RENS 01CO 0.111 0.500 0050 10:11 0.50) 0250 lOCO 0.286 0.333 0.343 0.125 GENTlANA AIJThI,~'!J\lIS 0200 0500 0.429 0.086 0.313 GRAnolA PILOSA 0.143 0.029 0.147 0.143 0125 G'fl,~OPOGON iWBIGJLG 0222 0500 0.20) lOCO 0063 G\1~ON BREVIFUJUS 0.091 0.150 0.429 1.000 0.556 0.147 0.286 0.125 HEl.lIINTIIEMUM CfiNlillENSE 0091 O.SCC 0.400 0.143 H~LG ANGUSTIFOUUS 0.037 0.333 0.2,"/:1 0200 0.429 HEl.lIINTIILG ATRORlmENS 0.091 0.050 0.571 0.029 HEUANllJUS HETEROPHYUUS lOCO ' 0.029 0.559 0.143 HIERACIUM GRO~'Il111 0.091 0.100 0.074 0200 1000 0.400 0.500 0.250 0.214 0.667 0.086 00&3 HIERACILIA XM4RlANUM 0.091 0.400 0.500 0.500 0.125 hDUSTONIA LONGIFOLIA 1.0:.1 0.071 HOUSTONlA PROCLI.~BENS 0.185 0600 1.000 0.333 HYPERK:UM CRUX·ANDREAE 0200 0.071 lOCO 0.222 0257 0.235 0.857 0375 HYPERlCLI.~ IlENTICUlAnJM 10:11 HYPERICUM GENTIANOIDES 0273 0.111 0.200 ICoo 0.071 0.063 HYPERICUM HYPERICOIDES 0.939 0.100 0.185 02SO 0500 10:11 0.800 lOCO 0.500 0.500 0.57 1 0.667 0.114 0.250 HYPOXIS HIRSUTA 0.250 100) 0.143 0.333 HYPOXiS MICIlAIIrnA 0.300 0.143 0.471 0511 IlEX CORIACEA 0143 0333 0286 0029 0.2;() IUEXGlABRA 0.200 0.037 0.2SO 0.100 0800 0500 0.357 0.333 0.889 0~1 4 0.794 0.857 0.625 IlEX OPACA 0.100 0.037 0.750 0.050 0.003 0071 0.333 0.2fr) 0.125 IlEX VOMrTORlll 0.074 0.500 ICOO 10:11 0667 IONACns UNARlfOUUS 0455 0,800 0.074 0.750 lOCO 0600 1000 o.m 1.00) 0.486 0,324 0.286 0.186 IRIS VERNA VEf\\IA 0273 0.500 0.250 0.857 0.457 0.235 0.286 0.5€3 JUNCUS BIFLORLG ODSO 0,071 0.029 0.0&3 KAU,RI\ lATIFOUA 0.250 10:11 lACHtWllHES CAROUANA 0.556 0057 0176 lACHt-IXAULON ANCEPS 0333 0.086 0.765 0.857 0.438 lfCHEA MIt()R 0.037 0200 1000 0.029 LECHEA SEssmORA 0.6)J 100) LESPEDEZA ANGUSTIFOUA 0.050 0.429 0.114 0029 0.125 LESPEDEZA CAPrTATA O.ml 0.250 0.714 0.667 0.111 0.314 0265 0.429 0.250 LESPEDEZA HIRTA 0.100 0.074 0.200 0200 0.333 0.029 OWl LESPEDEZA MERMEDIA 0.050 0.071 0.063 LESPEDEz.I PROCU~'IlENS O.WI 0.037 0.050 0.200 lESPEDEz.I REPENS 0.091 0.122 0.400 0.500 0.800 0.250 0~29 1000 0.029 0.059 0.125 ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ -'" ~ ~ :g ~ -g ~ ~ 'g ~ ~ ~ 'C'" "8 ~ ~ ~ :;:: ~ ~ :a ~ ~ ~ ~ J 'g ~ go 'is ~ ~ ~ --' 0 --' ~ '" --' .~ j -'" <.> .~ ~ ~ [ <.> ·c .-;: ;0 a. 1 (J) ~ .g '" 1l :> .~ --' u: JI ::J :> (J) &l --' --' ,;; (J) ~(J) ~ ~ ::l ::l JI --' ~ ::J E E .g '" i '"E E ~ ~ ~ '"~ '"~ ~ .2! '" .2! '" .2! ~ .""E :;: :;: ~ -i :::L ~ ~ ~ ~ ~ ~ .s ~ -'" -'" ~ ~ ~ ~ ~ ~ en en &! eni ~ ~ ien en « « ~ ~

~size 10 28 20 14 35 33 16

LESPECEA ~RGNK:A 02:<) O.2!O 0.500 0.571 0.667 0.029 0280 0.063 lJATRIS G~I~FCUII 0.727 ~.3CO 0.111 O.7!O 0.500 0.571 0.235 0.571 0.063 lJATRIS SPICATA 0091 0.280 Off) 0.1IIIl 0.176 0.143 llATRIS SOOAPROSA O.O!O 1.000 lJATRIS SQUARRl.lOSA 0.1137 LX;ANIA MICfI/IUXlI ·.0.852 0.400 0.500 Lm1lAMBAR ST\'1lACFLlJA 0200 0.2:<) 0.214 0.333 0.457 0.059 0.125 lOBElIA BREVlFCUII 0.889 L08® NUTTMUI 0.143 Ql14 0.765 0.429 0.375 LOB8JA PUBERUA 11:00 0.111 LOP'rlWAlJRfA 0.556 ll.O'IIlGlA HIRT8lA O.O!O 0.111 0.029 0286 0.063 LLUIIIGIA VIffiATA 0.1IIIl O.I IB 0.286 0.063 LYOO'OOElJ).ALOPECIJROIctS 0.071 0.111 0.029 0.471 0.429 LYccroDI8lA ~IANA 0.111 0.324 0.143 0.188 LYONJA lXlUSTRINA 0.214 0.114 0.429 0.438 LYONIA MIRIANA 0.100 02!l.l 0.400 0.500 02:<) 1.000 0.571 0.457 O.I1B 0.286 O~ f.\\OO.IA VlRGINiANA 0.2!o O.O!O 0.500 0286 0.486 0.353 0.286 0.2!I.l ~WfR£DA VIRGINCA 1.000 0.333 WJlSHAI.UA GRM~NIFOlIA 0.100 0.029 0.412 0.429 WJ1SHAllIA QBOVATA 0.091 0.100 0286 0.143 WJ1SHALlJA fW!IlSA 1.000 /Ij ~I:JSA QUADRrlALVIS 0.519 0200 Q400 0.357 0.667 0.063 ~JMJARTIA CAROUNIANA 0.727 0.185 O.O!O 0063 MUlLE~ERGIA CllPllARIS TRICHOPOOES 0~74 0100 0.071 o.m 0.676 0.429 0.125 MYRCA CERIFERA 0&0 0.037 1.000 0.0:<) 1.000 0.500 0.500 1.000 0.444 0.829 0.735 0.143 0.063 t.M'ICA HETEROP'rlYlJ). 0.143 0.343 0.441 U571 0.5&) NYSSA BIFlORA 0.100 0.111 0.114 0.176 0.2!I.l NYSSA SYl.VAnCA 0.100 0.2:<) O.1!O 0200 0.2:<) 1.000 0.500 UOO 0.200 0.375 OENOTHERA FRlIT'COISA O.1!O 1.000 0286 0.029 0.029 0.143 OPLII'1lA HUML'USA 0.182 0.148 O.O!O 1.000 ORllEXiJJM PEOOt-.aJLATUM PSOIWJ(J:DES 0.091 01~ 0.429 0.147 0286 0.3 13 OS~NTHlIS MlERlC/lNUS 0.7:<) 0200 0.50:1 0.333 OSMUMlA CNNN.+:lMEA O.O!O 0.143 0.257 0.029 0.714 O.~ OXYDENDRUM APBORELIA 0.100 02:<) 0.500 0.071 0.125 OXYPOUS RUFOR~tS 0.556 OXYPOUS TERNATA 0.059 0.429 0.125 PANICUM MtEPS 0.500 0400 1.000 0.222 0.029 PANW.I VlRGATUM 0.091 0.259 0200 0.071 0.556 0.029 0.088 0.286 O.HI8 PASPM.UMlAE'It O.O!O 0.667 0.111 0.088 -.J V-l -...J +>­

~ -g ~ =~ = = ~ -'" ~ -'" ~ =~ =~ '0 =~ "0 "0 = ~ ~ g ~ = 8 ~ ~ =~ =~ ffi ~ ~ ~ ~ ~ ~ "8 ~ ~ --'== == '8 =0 ] "8 ~ :::j== ~ --' 8­ ] < ~ ~

Samp~ iize II 10 28 20 14 35 33 16

PASPALUM PRAECOX O.~ 0.059 PASPPWMSETACEUM 0.037 0.400 1.000 03..13 0222 0029 0.029 PENSTE II.ONAUSTRALIS 0.037 0.200 0.500 0.286 0.063 PERSEA BORBONIA 0.300 0.750 O.())j 0.071 0.333 0.571 PERSEA PALUSTRIS 005:1 0059 0.143 0.500 PltlGlJCUlA SPP. 0.444 o.m; 0.143 O.oru P~US Ern INATA 0.050 0.500 lool 0.333 O.oru PINUS PALUSTRS I.Coo I.O:X) 1000 I.Coo 1000 1000 lool IO"J) ICOO 1.000 1000 I.Irn 1.000 Hoo 1000 1.000 I.COO l.e:xJ PINUS SEROrnA O.IOC 0.214 0.257 0.647 0.429 0500 PIM.IS TAEOA 0. 182 0.200 0.037 1.000 0.1 so 0.4C() 0500 1.000 0.286 0333 0222 0.200 O.oru PIIYCI'SIS GR.AMU1IFOUA 1000 0.100 0.889 Q2SO 0.9:xJ 1.000 0.000 1000 0.250 1000 O.SOl O.gx.) 1.0:xJ 0333 0.114 0.882 0.286 0.250 POlYGALA CRtmTA o.m 0.029 0.382 POLYGALA LUTEA 0.2S0 0071 0.371 0~ 12 0.571 0.375 PCLYGALA MARIANA 0500 0.029 0.029 POLYGAi.A NANA O.roJ 0.667 POLYGALA POLYGM.~ 0.091 POLYGALA fW.+:lSA 0.444 0200 0.143 POLYGONEllA GRACIUS 0704 POlYGONEllA POlYGM.~ 0.091 0.037 0.500 0.500 0.071 POTEmuA CAN.ADENSIS 0.2S0 0.571 0.029 0.286 0.063 PRUNUS SEROnNA 0.037 O.fI'Jl 0.500 OroJ 0.2S0 0.500 0.429 1.000 0.188 PTERIO:UMAQU!.INUM 0.4 00 0.407 0.2S0 04C() 0.500 O.BOO 0.500 0.500 0857 0.333 0.657 0.647 0.571 0.813 PTEROCAULON VIRGAMI 0.1 48 0250 1.000 0. 171 0.118 PYCNAN1lfEI.lJM FlEXUOSUM 0. 286 0.086 0.088 0.857 0.375 PYXJ OANTH ERA BARBUlATA 0.091 0100 0.500 0.114 0.125 OO ERCUS FALCATA 0.300 0. 150 0.50) O.IIJJ 1000 0.2S0 0.071 1000 0.057 OOERCUS GEMINATA 0.148 10:!) n500 0.029 OOERCUS HE MiSPflAERICA 0.100 0037 1.c\''1l O.())j I.e] 0.400 nsoo 0.071 0.333 0.057 O.oru mucus IttPi'IA O~ 1.000 OB I5 0.750 0.1iiil ICOO 0.800 1.000 0.571 0667 0.143 0.188 QUERCUS lAEVIS I.Irn 0.500 1.0lC 0.750 o.m 1.000 O.BOO 1.000 0250 0.500 0.857 0.333 0. 188 OOERCUS MARGAREillAE 0.273 O.3llO O . ~ 0250 0.700 1000 OBOO 1000 0250 Ogx.) 0333 0.057 0250 OOERCUS MARI.ANIlICA 0.900 0.400 0500 1.000 1.000 I.COO 1000 0.111 0.114 0.059 0.143 O~ OOERCUS NKlRA 0.100 0.100 0.200 0.143 Hoo 0200 O.oru QUERCUS PU~flA 0.800 0.057 0.029 OOERCUS STEllATA 0.100 0.050 0.200 0.500 0286 0.001 0086 flij EXIAAUFAN US 0.200 0.571 Of,,7 10:0 0.543 0.941 0.857 0.938 RHEXIA MARIANA 0.071 0.333 O.rd 0.057 0118 0.313 RHEXl4PETlO lATA 0.071 0086 0.4 71 0.429 n313 RHOOODE~DRON ATIW/T1C UM 0.050 0.143 0.029 0.059 0250 .., .., .., ~ ~ ~ ~ .., .., .., ~ ~ .., ~ ~ ~ ~ ~ 'g .., ~ '5 ~ 'g ~ ..!!! 'g .., .., ~ '5 '5 ~ ~ ~ ~ ~ 'g ~ ~ ~ "8 ~ ~ ~ 0> ~ ~ 'g ~ 0 'g ~ 0 ~ 'g 'g 'g ~ ~ ~ ~ ~ ~ g. ~ .., ~ -= ~ '-' ~ ~ ~ '" ~ :::l ~ '-' ~ in ~ 0 ~ ~ :::l ~ ." ~ ~ ~ ~ ~ ":ii .., ~ ~ ~ ~ ~ ~ '-' '-' ~ :::l .~ ~ ~ ~ ~ ~ ~ ~ '-' 1 '-' ~ .g '-' '-' ;0, '-' '-' -~ ~ ~ ." ~ ~ ~ ~ i ". ~ ~ ~ j &'l ~ ~ '" ! '" ~ :!!l ~ ~ ~ ~ :.i ~ ~ :!l! '"E '-' ~ I ~ ~ '-' '" '-' ~ ~ '-' ~ i ~ J!! E = ~ ~ = = ~ ~ ~ ~ ~ ~ ~ "5 ~ e. ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ '" .!t'" <1l ~ ~ ~ ~ ~ ~ ~ ~

Samp~size II 10 28 20 14 35 33 16

RHUS CCI'AWNUM 0.182 0.100 0.074 0.800 1.000 0400 0500 0000 1.1:00 0.500 0.929 1.000 0.314 0429 0.438 RHYNCHOSIA CYllSOIDES 0.593 0.200 1.1:00 RHYNCHOSIA RENIFORMIS 0273 0.100 0.333 0.400 I.COO 0800 0.500 0.2141 (1) RHYNCHOSIA TOMENTOSA 0.500 0.286 0.333 RHYNCHOSPORA BAlDWiNIl 02:i1 0.667 0.412 RHYNCHOSPORA BREVlS8A 0.500 0.143 RHYNCHOSPORA CHAPMANII 0.667 0.529 0.571 O~ RHYNCHOSPORA CIUARIS 0.333 0.111 0.066 0.853 RHYNCHOSPORA FASCIClJ.ARIS 0.778 0.029 0.235 RHYNCHOSPORA GLOBUlARlS 0.667 0.111 0.088 RHYNCHOSPORA GRAY! 0.635 0.222 0.150 0.400 0.500 0.357 0.333 0.143 RHYNCHOSPORA LATIFOLIA 0.0:iI 0.556 0.059 0.143 RHYNCHOSPORA OLIGANTHA 0.889 0.029 RHYNCHOSPORA PLUMJSA 1.1:00 0.171 0.765 0286 0.063 RHYNCHOSPORA RARIFLORA 0.091 0.556 0.029 0285 RUBUS ARGUTIUS 0.100 1.1:00 0071 0.567 0.057 0.125 RUBUS CUNEIFOUUS 0.500 0.2:i1 0071 RUBUS FLAGELLARIS Ol:il 0.400 0.333 0.029 0.188 RUBUS lRlVlAUS 0.400 0.071 0.057 0.059 O.C63 RUDBECKIA HIRTA 0.657 SABAnA CAMPANlJ.ATA 0.778 SARRACENIA AlATA 0667 SARRACENIA FLAVA 0.071 0.086 0.2!11 0.429 O.C63 SARRACENIA PURPUREA 0.029 0.118 0.143 02:i1 SASSAFRAS AlBIDUM 0.182 0.300 0.037 100) 0.500 0.500 0200 1.000 0.2:i1 0.500 0500 1.1:00 0.029 0500 SCH~HYRUM SCOPARIUM 0273 0.600 0.074 02:i1 0.300 0200 IOOC 1.0;(1 1.000 0.214 1.1:00 0.557 0.314 0.143 0.188 SCH~HYRIUM lENtRUM 0.259 1.1:00 0.111 SCLERIA CIUATA 0273 0074 0.300 0.250 1.000 0.357 0.333 0.029 0.143 O.~ SCLERIA MINOR 0.071 0.086 0.412 0.286 0.063 SCLERIA PAUCIFLORA 0.091 0.214 0.333 0778 0.086 0.529 0.571 0.125 SCLERIA RETIClJ.ARIS O.O:il 0.889 0.059 0.143 SCLERIA TRIGLOMERATA 020) 0.400 0.500 0.714 0.114 0.375 SERENOA REPENS 0.530 1.000 SEYMERIA CASSKJIDES 0.200 0500 0071 0.086 0.529 SIUPHIUM COMPOSmJM 0.354 0.185 0.7:i1 0.500 0500 0.643 0.125 SISYR~CHIUM AlBIDUM 0.074 0.100 0071 0.333 0029 OC53 SISYR~CHIUM NASHII 0182 0.100 0.143 0.143 0125 SMIlAX AURiCULATA 0259 1.000 0.400 0.333 Sf~lAX BONA·NOX 0.100 0037 1.0:0 1.1:00 0.029 0053

-.l VI -.J 0\

"C "C -g C C ~ ~ "C ~ "C C -g c "C "C ~ '§ "C C C ~ '§ "C "C -'" c ~ ~ '" c ~ '8 'g ~ c ~ ~ ! ~ c 8' .... 0 :::l== == a:> ~ '§ ~ ~ I .... 0= ~ g...... 0 ~ ~ ~ ~ ...... , ~ :::l en J c

SMILAXGWU 0.400 0.250 0.600 0.200 0.500 1.000 0.500 0.714 1.000 0.111 0.451 0.625 SMILAX lAURmJA 0.500 0.333 0229 0.0<'> 0286 0.500 SMIAX PUMllA 0.074 1.000 0.333 S).ILAX ROTUNOlFOllA O.OSO 1.000 0.071 0.667 00<'> SMILAX SMAWI 0.400 0.667 SOUllAGQ OOORA 0.364 0.630 0.900 1.000 0.&1) 0.500 02SO 0.500 0.857 1.000 0.111 0.143 0.059 0.571 0.563 SOUOAGO RUGOSA 0.667 SOUDAGO SlRCTA &PULCHRA 0.086 0.1.82 0571 SORGHAS1IUM NlIT/INS 0.200 0200 1.000 0.429 0.333 o.m 0.0<'> 011.8 SPOROOCWS JJfO'US 0.364 0.815 0.200 0.500 1.000 0.071 0.667 STIW'l'GIA SYlVATlCA 0.182 0370 O.1SO 0.600 STPUUCIDA SETACEA 1.000 02SO 0.1 SO 0.071 0.063 STROPHOSl'IlES UI£EUlATA 0.667 ST'iUSIM PATENS 0.818 0.300 om 0.2SO 0.350 1.000 0.Il00 1.000 0.357 0.333 smmCAANEUS O.OSO 0.500 0200 0071 STYlOSANTHES BIFLORA 0.182 0.556 0.5SO 1.000 0.400 0.500 1.000 0.786 1.000 0.143 0294 0.286 0.125 S'rWlOCOS TIt-tTOPIA 0.100 0.500 0.500 0.2SO 0071 0.257 o.m 0.125 TEPHROSlo\ CHRYSOPHYllA 0.593 1.000 TEPHROSIA FLORIDA 0.455 0200 0.148 0.300 1.000 0.429 1.000 0.057 0.188 TEPHROSIA HISPIDUlA 0.100 0.500 0.200 0.294 TEPHROSlo\ OOOORYCH(]IlES 0.667 0.111 TEPHROSlI, SPICATA o.oso 0.429 0.667 0.114 0.143 TEPHROSIA VlRGINlI,NA 0.091 0.100 0.074 0.650 0.500 0.600 0500 0.500 0.929 0.667 0.114 om 0.143 02SO TCflElDlI, RACE/.KlSA 0.222 O.7t1l 0.429 TOXKXXlENcroN PUBESCENS 0.455 0.100 0.074 0.800 0.400 1.000 0.857 1.000 0.313 TOXiCOCtNDROO RADtANS 0200 0.250 0200 0.071 0.143 0.125 TOXlCOCtNDRON VERN~ 0.500 0286 QI1.8 TRAGIA URENS 0.636 0.400 0.222 0.350 1.000 0.800 0.500 0.643 0333 0.125 lITRICULARlI, SUlllATA 0.556 o.m 0.143 VACCINIUM ARBOREUM 0.091 0200 0.111 1.000 O.lfXl 1.000 1000 Om:) 0.250 0.214 1.000 VACCIf.ll.IA CRASSIFOULlj 0.500 02SO 0.200 1.000 0.286 0.743 0.618 0.429 0.375 VACCINIUM DARROWII 0200 0[£7 VACCINUM Ewom 0.111 1.000 1.000 0.250 1.000 0071 1.000 VACCINIUM FORMOSUM 0250 0.214 0029 0.4<'> 0.563 VACCIf.ll,Ij FUSCATUM 0.100 0.037 OOSO 0.500 0.600 1.000 0.500 0.143 0.486 0.0<'> 0.375 VACCl'lll.l.1 MYRSiNlTES 0.296 O.OSO 0200 VACCINUM STM111II:1.I.1 0.091 0.600 0.148 0.250 0.100 1.000 0.400 1.000 0.250 1.000 I.OCO o.m VACCINIJM TENEllL!.1 0.091 OBO) 0.750 0.&1) 0250 1.000 1.000 0.800 0.059 0.429 0.625 VERNONlI, ACAUUS 02SO 1.000 0.571 0.057 0.059 0188 LL

E'lE'l~g gg~ «< 'i3 ~~ >"'H'Hn :o eee i5 j' ;Z: ;Z: :::!J ~ ~$: c>~~ ;::R = !(l. :'ii:'ii~e 5!!3S ~"'~'" ~ ~m ~;;~g ~ ~~ ~ ~i>; 3~~ s;l >::! Cl 55 ::offiQ» 5> ~ ~ ! 'S" ~

~~ =: Fall-line Xeric LL Woodland

8 8 <5 Allanti cXeric LL Woodland

p p p ~ . ~. Ml Sl Southern Xeric LL Woodland

§ Atlanlic MarilimeLL Wood land

p p 8 8" ~§ ~ Fall-line Subxeric LL Woodland

§ § Allantic Subxeric LL Woodland

§ §~ Southern Subexeric LL Woodland

Subxeric Saw PalmettoWood land

§ PeidmontlUpland LL Woodland

§§ § Serpentine Subxeric LL Wood land

Fall-lineMesic LL Slope Woodland

:;;: Fall-line Mesic LL Woodland ~ ~. ~~ ~

~ ~ Southern Mesic LL Woodland

~~. ~ i ~ Southern LL Savanna

p= ~~. ~ - lHl i§ i§ ~ AllanticLL Flatwood

p= h"" . ~ §l ~ . ~~ ~. ~ Allantic LL Savan na

~~ ~ . ~~. ~e;"" ~. Fail-line Seepage Savanna

p = = :..-... N . c; w ~. g;:8 ~;~ Fall-line LL Seepage Bog LITERATURE CITED Daniels, RB., H.J. Kleiss, S.w. Buol, H.J. Byrd and J.A Phillips. 1984. Soil systems in North Carolina. Abrahamson, W.G. and D.C Hartnett. 1990. Pine North Carolina Agricultural Research Service, flatwoods and dry prairies. Pages 103-149 in Bulletin 467, Raleigh, NC Ecosystems of Florida. RL. Myers and J.J. Ewel, eds. The University of Central Florida Press, DuBar, J.R, H.S. Johnson Jr., B.G. Thorn and W.O. Orlando. Hatchell. 1974. Neogene stratigraphy and morphology, south flank of the Cape Fear arch, Allard, D.J. 1990. Southeastern United States North and South Carolina. Pages 139-173 in Post­ ecological community classification. Interim Miocene stratigraphy, Central and Southern report, Version 1.2. The Nature Conservancy, Atlantic Coastal Plain. RQ. Oaks Jr. and J.R Southeast Regional Office, Chapel Hill, NC DuBar, eds. Utah State University Press, Logan, UT.

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81