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

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Longleaf Pine Vegetation of the Southern Atlantic and Eastern Gulf Coast Regions: a Preliminary Classification* 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 naturcliiongleaf pin~~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 / 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~r,i.Fh 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 fegularly observed with over 40 species of plants per square meter, and Mesic Longleaf Woodlands were found with up to 140 species per 1000 m2• 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. *Botanicalnomenclature follows Kartesz (1994), except we follow Peet (1993) in, recognizing that the plants traditionally treated as Aristi~a,stricta should be divided into a northern (A. strieta) and a southern species (A. b~yrichiana). Proceedings of the Tall Timbers Fire Ecology 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 (Pinus fires kept the pine woodlands open and relatively palustris) 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 particularly 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 ofsecond- 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 B I! ! I I iii I iii I • I ' I iii Iii iii 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 palust(iSj 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 (Aristida stricta and A. beyr/chianti) by diagonal shading (after Peet 1993). However, most ofthe 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
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