Current and presettlement tree species composition of some upland forests in northern Mississippi
Brewer, J Stephen
Current and presettlement tree species composition of some upland forests in northern Mississippi1
BREWER, J. S. (Department of Biology, P. O. Box 1848, University, MS, 38677-1848). Current and Presettlement Tree Species Composition of Some Upland Forests in Northern Mississippi. J. Torrey Bot. Soc. 128:332349. 2001.–Combining the study of changes in mature forests with the study of secondary succession provides a unique opportunity to understand forests in a way not made possible by studying each alone. In this study, I examined tree species composition of two mature (>150 yr-old) upland hardwood forests with a long history of fire suppression and two younger second-growth upland forests (40 yr-old), also fire-suppressed. I then compared present-day tree species composition of the two old-growth forests and the two younger forests with the composition of bearing trees (i.e., trees identified by land surveyors in the 1830s) in upland and lowland areas within townships encompassing the four sites. Bearing tree samples were similar in composition to an explorer’s account of old-growth woodlands that dominated the region during sparse subsistence settlement by Chickasaw Indians. There were striking differences between present-day and bearing-tree samples. One of the old-growth forests (Bailey Woods) was more similar in species composition to the mid-successional secondgrowth forests than to bearing-tree samples. The shade-intolerant, but fire-tolerant, Quercus marilandica Muenchh. was nearly absent from present-day upland forests (= 10 cm dbh; range 0-3%) but was the most common species among presettlement upland trees (42% of all bearing trees in the township containing the field sites). Liquidambar styraciflua L., often regarded as both a pioneer species and a common constituent of alluvial floodplain and mesophytic terrace forests, was the single most abundant tree species in the understory and midcanopy of one of the old-growth forests (Bailey Woods) and in all size classes of trees of both younger forests. It was completely absent from presettlement records of upland trees, but was an abundant bearing tree in nearby sections containing lowland alluvial forests. I hypothesize that open oak woodlands dominated the upland landscape of Lafayette County, Mississippi before extensive settlement. These woodlands were neither mid-successional nor late-successional forests but were a unique fire-dependent community type that is absent from north Mississippi today.
Key words: fire, oak regeneration, succession, bearing tree, sweetgum, black jack oak, post oak, black oak, interior coastal plain.
The study of changes in species composition in forests has long been of interest to ecologists (e.g., see Clements 1916; Oosting 1942; Braun 1950; Beilmann and Brenner 1951; Keever 1953; Quarterman and Keever 1962; Christensen 1977; Poulson and Platt 1996; McCarthy et al. 2001). Numerous studies have focused on patterns of dominance and replacement of tree species in mature oak-hickory and pine-oak– hickory forests (e.g., Christensen 1977; Cain and Shelton 1994; references in Abrams 1992). There is widespread recognition that fires played a major role in maintaining oak- and pine-dominated communities in the eastern United States before EuroAmerican settlement (Quarterman and Keever 1962; Pyne 1982; Grimm 1984; Johnson and Schnell 1985; Lorimer 1985; Myers 1985; Dorney and Dorney 1989; Foti and Glenn 1990; Cutter and Guyette 1991; Abrams 1992; Delcourt and Delcourt 1998; Harrod and White 1998; Kirwan and Shugart 2000). Furthermore, some have argued that fires sharpened the ecological contrast between fire-dependent and fire– sensitive plant communities and thus increased biodiversity across the landscape in the past (Delcourt and Delcourt 1998). Hence, the ecological and conservation significance of longterm changes in mature forests must be interpreted in light of recent changes in fire regimes.
In addition to changes in mature forests, much is known about succession from oldfields and clearings to oak-hickory and pine-oak-hickory forests (e.g., Oosting 1942; Peet and Christensen 1987; Gill and Marks 1991; DeSteven 1991 a, b). Rapid colonization of oldfields and clearings by widely-dispersed, fast-growing species (e.g., Pinus taeda L., Liquidambar styraciflua L., hereafter, loblolly pine, sweetgum) is a characteristic feature of early-successional communities in the southeastern United States (Borman 1953; DeSteven 1991a, b). In contrast to the increasing interest in the role of fire in mature oak forests, however, most studies of oldfield succession have ignored the role of fire.
Although much is known about oldfield succession in areas previously dominated by oaks and pines, its relevance to the development of presettlement oak/pine communities has not been clearly established. Effective restoration and management of oak/pine-dominated communities will benefit from a reasonably accurate description of succession in presettlement landscapes. One hypothesis, hereafter, the “filter” hypothesis, emphasizes the importance of recurring fires in the development of oak- and pine– dominated communities (Chapman 1932; Schwartz 1994; Platt 1999). According to this hypothesis, advanced regeneration of slowgrowing, fire-tolerant upland oaks and pines follows major natural disturbances and is maintained as long as fires “filter out” fire-sensitive species (Platt 1999). With fire suppression, however, these fire-tolerant oaks and pines [e.g., Quercus stellata Wang. (post oak), Q. marilandica Muenchh. (black jack oak), Pinus palustris Mill. (longleaf pine)] cannot compete in large gaps with faster-growing, fire-sensitive pioneer species such as loblolly pine and sweetgum (Platt and Schwartz 1990; Frost 1993; Platt 1999). Thus, 2011-century patterns of oldfield succession in upland landscapes, for the most part, are a recent artifact of effective fire suppression and not necessarily indicative of how communities recovered from major disturbances prior to EuroAmerican settlement.
Combining the study of presettlement communities with the study of secondary and oldgrowth forests today provides a unique opportunity to infer the role that ecological processes such as fire played in structuring oak woodlands in the past. In this study, I examined two upland old-growth hardwood forests with a long (> 70 yr) history of active fire suppression (Bailey Woods and Jackson Strip) and two younger second-growth forests (40 yr-old; Oak Grove and Marijuana Fields Strip, hereafter MF Strip). All four stands were located within the city limits of Oxford, Mississippi (34 deg N, 89 deg W) in central Lafayette County, MS. In addition, I examined presettlement tree species composition in the vicinity of present-day Oxford, which was either uninhabited, or at most, sparsely settled by the Chickasaw Indians throughout most of the 300year period between first contact and settlement by U.S. citizens (1541 narratives of Ranjel and the Gentleman of Elvas in Malone 1922, Johnson 2000). The objectives of this study were: (1) to describe tree species composition of two oldgrowth forests and two mid-successional forests, 2) to compare present-day tree species composition of Bailey Woods, Jackson Strip, Oak Grove, and MF Strip with that of presettlement upland and lowland alluvial forests in central Lafayette County, Mississippi, and (3) to formulate a hypothesis about differences in recovery from major disturbances in pre- and postsettlement upland landscapes in northern Mississippi.
Methods. STUDY AREAS. The bulk of the current study was conducted in Bailey Woods, which is a tract of mature woods (approximately 20 ha) managed by the University of Mississippi. Part of Bailey Woods is a National Historic Landmark and the former estate of the late novelist, William Faulkner. The remaining part of Bailey Woods has been owned by the university since the date of its charter in 1844 (Sansing 1999). The university was built on forested land in 1848 (Sansing 1999). Aerial photographs of Bailey Woods and personal interviews with long-time residents of Oxford and relatives of Faulkner indicate that a mature deciduous forest existed as early as the late 1930s and has retained this appearance since. Faulkner, himself, commented that he had played in the woods as a child (early 1900s; Lawrence and Hise 1993). Local residents of Oxford referred to this area as “Bailey Woods” during the 1800s (Lawrence and Hise 1993). Some long-time residents of Oxford (including relatives of Faulkner) have suggested that the forest has never been cut. The oldest trees aged in this forest were white oaks (Quercus alba L.) and post oaks (Q. stellata Wang.) over 150 yr old, with diameters-at– breast-height (dbh) greater than 1.1 m. There is clear evidence of some human-caused disturbance within a small portion of the interior of the forest, however. Aerial photographs revealed that some selective logging was done to create a dirt road, which has been abandoned since the 1950s. Up until Faulkner’s death in 1962, the understory of the forest had experienced some grazing by dairy cows (Lawrence and Hise 1993). Fires have not occurred within the interior of the forest, however, for at least 70 years and according to long-time residents of Oxford, much longer. Since 1973, when the university bought the estate from Faulkner’s daughter, Jill Summers, there has been no significant human disturbance of the forest, although residential development adjacent to Bailey Woods continues today.
In addition to Bailey Woods, another mature forest stand was studied, Jackson Strip. This narrow strip of woods (from 50 to 200 m wide by -500 m long) occurs along a major road in Oxford (Jackson Avenue) and has been owned by the university since 1844. Like Bailey Woods, aerial photographs show these woods were mature in the 1930s (in contrast to the surrounding landscape not owned by the university). Some post oaks located at the edge of the forest, which were recently cut to widen Jackson Avenue, were over 120 years old. Bailey Woods and Jackson Strip are as old as any upland oak forest I have been able to locate in north central Mississippi.
In addition to Bailey Woods and Jackson Strip, tree species composition of two mid-successional sweetgum-oak-pine forests within Oxford were examined. One forest (Oak Grove; ~10 ha) was similar in shape to Bailey Woods. Another forest (MF Strip) was similar in size and shape to Jackson Strip. Both forests were the same age (~40 yr old; as deduced from stump rings of recently cut pines and aerial photographs). Both experienced considerable damage from an ice-storm in 1994, which appeared to have had the greatest negative impact on loblolly pine. Most of the dead and damaged pines were salvaged from MF Strip, however, creating a fairly open overstory canopy. As in similar communities in the southeastern United States, sweetgums and pines are among the first trees to become established within Andropogon/Schizachyrium scoparium-dominated oldfields in this area.
All four forests were located in a physiographic region recognized in Mississippi as the north central plateau of the interior coastal plain. The geological formation in central Lafayette County is primarily Eocene sands and clays (Hilgard 1860, Lowe 1921). All four field sites occur in an area that Hilgard (1860) described as being typical of oak-hickory forests in the western and northern portions of the north central plateau (the so-called called Yellow-Loam Hills and Table Lands). Pines were not a significant component of the early-19th century forest in the vicinity of the current field sites (Hilgard 1860). The soil of all stands was a readily erodible silty or sandy loam underlain by a reddish sand and clay (Hilgard 1860; Lowe 1921; Morris 1981).
All four stands were located on an east-west oriented ridge between the Little Tallahatchie River to the north and the Yocona River to the south (Fig. 1). Hence, none were located within a kilometer of a major river. Bailey Woods drains southward into the Yocona River watershed (Fig. 1). Jackson Strip, Oak Grove and MF Strip drain northward and westward into Davidson Creek and the East Goose Valley watershed, which drains into the Sardis Reservoir (formerly the Tallahatchie River).
TREE SPECIES COMPOSITION. To determine tree species composition and potential stand replacement dynamics in Bailey Woods, four transects (140 to 260 m long) were oriented from the edge of the woods into the interior in mid-April, 1996. Later (in 1999), two additional transects were established atop ridges in the forest. Transects were placed so as to provide a representative sample of the forest as a whole and to examine differences associated with slope position and aspect. One transect was located parallel to a southfacing slope. A second transect was located parallel to the northfacing slope opposite the first. A portion of the second transect was located in a small floodplain of a small creek. A third transect was located perpendicular to a gentle southfacing slope and descending into a hollow. A fourth transect was located perpendicular to a southfacing slope, descending into a hollow, and then ascending a ridge within the interior of the woods. A fifth transect was located within the interior of the forest atop a ridge, and two additional points were located haphazardly atop another ridge. Altogether, 49 sampling points were placed within Bailey Woods.
In Jackson Strip, a single temporary (160 m) transect was oriented lengthwise >20 m from any edge. Approximately one-half of the length of the transect (containing 5 points) was located atop a ridge and the other half (containing 4 points) was located along an east-facing slope and associated hollow.
To provide a present-day contrast with the two mature forests, tree species composition of two mid-successional forests was quantified. Within each forest, a single temporary (140 m) transect was located along a west-facing slope at MF strip and an east-facing slope at Oak Grove. Each of these two transects contained 8 points.
Within each transect, I sampled tree species composition using a plotless (point-centered quarter) method (Cottam and Curtis 1956). Points were placed at 20-m intervals within each transect. Within each quarter, I located the two closest woody stems (taller than 1.5 m and >= 10 dbh) and measured the distance from the point to each stem, dbh, and identified the stem to species. Not all points contained 8 trees due to errors in sampling. These errors included sampling the same tree twice in two nearby points and accidental tagging of dead trees by student workers during spring censuses before leaf flush. At MF Strip, if two trees within a quarter could not be located before exiting the forest, then only one was located. Thus, at some points, fewer than 8 trees were sampled. I permanently tagged all stems at Bailey Woods for subsequent verification of their identity and to permit repeated censusing in the future. Altogether, I encountered 377 stems at Bailey Woods, 72 stems each at Jackson Strip, 64 stems at Oak Grove, and 59 stems at MF Strip. In addition to stems 10 cm dbh or greater, I counted all woody stems (except vines) greater than 1.5 m tall but less than 10 cm dbh within a 100 m2 circular pivot plot at each point (radius = 5.64 m). These stems included saplings of trees and stems of shrubs. For convenience, I call this size class, “saplings.”
PRESENT-DAY SIZE-CLASS DISTRIBUTION OF TREES. The density of trees in two size classes was determined for each species: “small” (i.e., 10-19.99 cm dbh, diameter at breast (1.5 m) height) and “large” (i.e., medium to very large trees from 20-113 cm dbh). Further subdivision of the large diameter class did not appear to provide much additional information on speciesspecific differences. I generated diameter class distributions separately for Bailey Woods and Jackson Strip and pooled trees from MF Strip and Oak Grove. I drew inferences about patterns of canopy replacement by comparing diameter– class distributions of common tree species with at least one stem >= 30 cm dbh.
RELATIONSHIP OF TREE SPECIES COMPOSITION To ELEVATION AND SOIL TEXTURE. To examine the relationship between species composition and the physical environment, I measured elevation above sea level and percent sand and each sampling point in Bailey Woods and Jackson Strip. I used a clinometer and measuring tape to measure relative elevation and related all measurements to benchmarks nearby. I did not obtain measurements at one point on a north-facing slope in Bailey Woods or one point on a slope at Jackson Strip. Percent sand was determined using a suspension method and a texture-dispersing reagent.
HISTORICAL TREE SPECIES COMPOSITION. To obtain quantitative estimates of tree species composition in the vicinity of the four forests in the 1830s (before extensive settlement by U.S. citizens), I tallied the frequency of bearing trees by species from data obtained from original surveyors’ notes in two adjacent townships containing the city of Oxford and an area south of Oxford (TBS, R3W and T9S, R3W; Fig. 1). In addition, I calculated the distance between survey posts and bearing trees to obtain a crude estimate of bearing tree density. Most of the trees were surveyed in 1834. None were surveyed after 1840. These dates coincided with a period immediately following the cession of this part of Mississippi by the Chickasaw Nation to the United States (Sansing 1999). Thus, tallying bearing-tree records provided the best available quantitative sample of trees representative of upland communities in this region before settlement by U. S. citizens.
All field sites were located within T8S, R3W, the township containing most of the city of Oxford, Mississippi. The township to the south (T9S, R3W) contained upland forest similar to that found in T8S, R3W and lowland vegetation associated with the Yocona River alluvial valley and associated tributaries in the watershed, including Burney Branch (Fig. 1). In addition, I tabulated data on a portion of T6S, R3W located within the alluvial valley of the Tallahatchie River (located ~ 40 km north of Oxford). Thus, bearing-tree species composition of the alluvial valleys of the Tallahatchie and Yocona rivers represented replicate samples of mesophytic forests of the early 19th century. I chose sections that, when combined, contained a gradient of forest types from lowland mesophytic forests (in the Tallahatchie and Yocona alluvial valleys) to upland forests on the ridge between the two watersheds, where Oxford is located. I excluded those sections in upland areas of the eastern and southeastern portion of T9S, R3W that were located on the Tallahatta and Neshobe Sand Formation and thus contained pine. I assumed that bearing tree records for sections containing upland areas in and around Oxford were representative of presettlement tree species composition in this region. I evaluated this assumption by comparing bearing-tree composition of this area to non-quantitative descriptions provided by Nutt ( in Jennings 1947) and Hilgard (1860).
I examined the bearing tree composition of sections contained within the Tallahatchie and Yocona River valleys to provide clues as to the historical patterns of distribution of certain mesophytic pioneer tree species such as sweetgum, sassafras (Sassafras albidum (Nutt.) Nees.), water oak (Quercus nigra L.), and red maple (Acer rubrum L.).
Land surveyors used common names only, and many of these names lacked specificity or were used only by Mississippians at the time (Hilgard 1860). Because of the lack of specificity, it was impossible to gauge the relative importance of, say, the various species in the genus, Carya (hickory). Hilgard (1860) was extremely helpful in translating common names used by Mississippians to the more generally used common names. This in turn aided translation into modern scientific names. I only made assumptions about the specific name of a tree when I could verify the identity of this species in Hilgard (1860) or Waffles (1854).
The surveyors’ general descriptions of the landscape, along with their estimates of the sizes of bearing trees, revealed that the vast majority of surveyed areas in central Lafayette County had not yet been cleared for agriculture by U. S. citizens. Subsistence maize agriculture by Chickasaws had not been common in this area either (Jennings 1947, Johnson 2000). Thus, the survey examined here provided the best quantitative analysis available of the tree species composition of “virgin” forests in this region. This does not mean, however, that early-19th century forests escaped human disturbance, however. The Chickasaws used fires in their territory to aid in hunting and gathering and perhaps to range cattle and other livestock (Cushman 1899, Jennings 1947, Naime 1988, Johnson 2000).
COMPARISONS OF PRESENT-DAY AND HiSTORICAL FORESTS. I compared tree species composition of two mature forests (Bailey Woods and Jackson Strip) and two mid-successional forests (Oak Grove and MF Strip) with that of presettlement forests in Lafayette County. I subdivided the samples of old-growth trees at Bailey Woods according to slope position and aspect. Altogether, I constructed 13 unique samples, which included all bearing trees and those present-day trees >= 10 dbh. In addition, to address the impact of surveyors’ bias toward selecting large bearing trees (e.g., see Grimm 1984), I constructed 12 unique samples, which included all bearing trees and only those present-day trees 20 cm dbh. In the latter case, I pooled the two Jackson Strip samples into one sample.
Reconstructing historical tree species composition using survey data has been shown to be reasonably accurate and consistent with anecdotal accounts of early explorers in other studies (e.g., Quarterman and Keever 1962, Grimm 1984, Clewell 1986, Schwartz 1994). Nevertheless, there are several biases inherent in survey data (including a bias towards large trees), which complicate comparisons with present-day data (e.g., Bourdo 1956, Grimm 1984, Schwartz 1994). Suffice it to say that survey data, alone, do not provide an entirely accurate picture of presettlement tree composition.
MULTIVARIATE DATA ANALYSIS. I analyzed differences in tree species composition among samples using detrended correspondence analysis (DCA, Hill and Gauch 1980). Rare species were downweighted, and axes were rescaled. This analysis enabled me to sort samples graphically on the basis of similarity in species composition. I summarized the results of DCA graphically by plotting axis 1 and 2 scores of both samples and species. I examined the relationship between species composition of present-day old-growth forests and elevation and percent sand by regressing axis 1 and axis 2 scores against measurements of these variables taken at point samples. The amount of species turnover for an axis was estimated by the length of the gradient (in standard deviation units) after rescaling.
Results. TREE SPECIES COMPOSITION. Sweetgum (Liquidambar styraciflua) was the single most abundant species encountered at three of the four sites, including both second-growth sites and one old-growth site (Bailey Woods; Table 1). Southern red oak (Quercus falcata) dominated Bailey Woods and MF Strip in terms of basal area, whereas sweetgum and hickory occupied the greatest basal area at Oak Grove and Jackson Strip, respectively. Jackson Strip was unusual among the four sites, in that neither sweetgum nor southern red oak were common. Post oak (Quercus stellata) was important within ridge-top samples in both Bailey Woods and Jackson Strip (Table 1).
Among sapling-sized stems, hickory was the most abundant species/group at Bailey Woods (22% of 1004 stems) and the second most abundant at Jackson Strip (31.2% of 179 stems; Table 2). Also abundant at Bailey Woods were winged elm (Ulmus alata Michs. 12.8%) and black gum (Nyssa sylvatica Marsh. 12.1 %). Although privet (Ligustrum sinense Lour.), an invasive exotic species, was also abundant at Bailey Woods (10.5%), most stems of this species were restricted to 2 or 3 sampling points at the edge of the forest. It was otherwise absent. Black gum was the most abundant sapling-sized stem at Jackson Strip (32.4%). Winged elm was the most abundant sapling at Oak Grove (23.3% of 249 stems sampled). Also abundant were red maple (Acer rubrum, 18.9%), flowering dogwood (Cornus florida L., 16.1%) and sweetgum (10.4%). Oaks were not abundant as saplings in any of the above sites. None of the above sites contained relative densities of oak saplings greater than 4%.
In contrast to Bailey Woods, Jackson Strip, and Oak Grove, relative densities of saplings of oaks were high at MF Strip, where total overstory and subcanopy basal area was much lower than at the other three sites (9.65 m^sup 2^/ha vs. 20.85 m^sup 2^/ha at Bailey Woods, 18.93 m^sup 2^/ha at Jackson Strip, and 16.64 m2/ha at Oak Grove). “Red oak” saplings (i.e. Quercus falcata/velutina/coccinea) comprised 17.2% of the 548 stems encountered at MF Strip. Quercus alba comprised 10.8% of all stems. Nevertheless, despite the apparent positive effect of overstory thinning on oak saplings, sweetgum comprised the majority of sapling-sized stems at this site (22.3%). Also abundant at this site were red maple (12%), black cherry (Prunus serotina Ehrh., 5.5%), and black gum (5.3%).
PRESENT-DAY SIZE-CLASS DISTRIBUTION OF TREES. In general, oaks were better represented in the larger diameter classes than in smaller size classes at most sites. In Bailey Woods, most stems of southern red oak, post oak, and black oak (Quercus velutina) were >= 20 cm (Table 3). In contrast, sweetgum, hickory, American beech (Fagus grandifolia), and black gum were most abundant in the smallest diameter class of trees (10-19.99 cm; Table 3). The largest stem was white oak (Quercus alba), which was 113 cm dbh and contained greater than 150 growth rings. The largest sweetgum stem was 57 cm dbh and contained 48 growth rings. At Jackson Strip, hickory was most abundant in the smallest diameter class, whereas post oak was most abundant in the larger diameter classes (>= 20 cm dbh; Table 4). At the second-growth sites, sweetgum was abundant in all diameter classes (Table 3).
RELATIONSHIP OF TREE SPECIES COMPOSITION TO ELEVATION AND SOIL TEXTURE. Among point samples of old-growth trees, ridge samples differed substantially in species composition from samples on slopes or in hollows or small floodplains (Fig. 2). Samples sorted along axis 1 with respect to the relative abundance of four common species, southern red oak (Quefal), hickory (Carya), and flowering dogwood (Corflo) vs. post oak (Queste), and some species that were not common (e.g., Magnolia grandiflora (Maggra), a mesophytic species, vs. black jack oak (Quemar), a ridge species; Fig. 2). The eigenvalue for DCA axis 1 was 0.44 (45% of all variation in composition). The length of the gradient in species composition was 3.551. Low axis I scores corresponded to sampling points on ridges with silt soil, dominated by post oak (Fig. 2). Axis I was negatively correlated with elevation (r = -0.397, p = 0.002, n = 56) and positively correlated with percent sand (r = 0.4, p = 0.002). Percent sand ranged from 16.7 to 93.3 at Bailey Woods and from 23 to 79.3 at Jackson Strip. Elevation above sea level ranged from 131.66 to 149.15 m at Bailey Woods and from 138.47 to 155.82 at Jackson Strip. Samples sorted along axis 2 primarily with respect to relative abundance of American beech (Faggra) and white oak (Quealb) vs. hickory (Carya), flowering dogwood (Corflo), and sweetgum (Liqsty). The eigenvalue for axis 2 was 0.35 (36%). High axis 2 scores corresponded to lowlying, north-facing slopes dominated by American beech and white oak. Axis 2 scores were negatively correlated with elevation (r = -0.464, p = 0.0003) and not correlated with percent sand (r = 0.057, p = 0.676). The length of the gradient in species composition was 3.103.
COMPARISONS OF PRESENT-DAY AND HISTORIcAL FORESTS. Among samples that included bearing trees and present-day trees >= 10 cm dbh, upland bearing-tree samples differed most from lowland bearing-tree samples, and present-day forests were intermediate in species composition. Samples sorted along axis 1 with respect to the relative abundance of black jack oak (Blkjak), black oak (Blkoak), post oak (Pstoak), and scarlet oak (Scaoak) at one extreme vs. beech and holly at the other (Fig. 3; see Appendix for translations of bearing-tree names used by surveyors). The eigenvalue for DCA axis 1 was 0.55 (71%). The length of the gradient in species composition was 3.321. Samples of lowland bearing trees corresponded to low axis 1 scores and appeared to have occurred within climax beech-holly-oak forests. High axis 1 scores corresponded to samples of upland bearing trees. Present-day samples of trees corresponded to inter-mediate axis 1 scores. Most study sites contained a mixture of alluvial or mesophytic species (e.g., sweetgum, beech, white oak) and upland oak species (post oak, southern red oak; Tables 1 and 4). Some samples of present-day trees (e.g., Bailey Ridge and both Jackson Strip samples) were more similar in composition to upland bearing-tree samples than other presentday samples of trees (Fig. 3). This was due in large part to the abundance of post oak in these samples. Nevertheless, there was considerable overlap between ridge samples and others. Samples sorted along axis 2 primarily with respect to relative abundance of sweetgum (Swtgum), poplar and maple, vs. cedar, black gum (Blkgum), and elm (Fig. 3). The eigenvalue for axis 2 was 0.16 (21%). High axis 2 scores corresponded to the second-growth forests (Fig. 3). The patterns revealed in the ordination of bearing trees and present-day trees ? 20 cm dbh essentially were the same as those including all present-day trees >= 10 cm dbh, except that poplar and pine were associated with low axis 2 scores in the analysis that included only large trees (Fig. 4).
One of the most striking differences in tree species composition between pre-settlement forests and present-day forests was the nearly complete absence of black jack oak from the presentday forests (including ridges; Tables 1 and 4, Fig. 5). Black jack oak was the most abundant bearing-tree species in the township containing Oxford, Mississippi (42.1%, Table 4). The secand most abundant upland bearing-tree species in this township was black oak at 22.4%; the third, post oak at 18.2% (Table 4). Post oak was abundant within upland bearing-tree samples of both townships and on ridges within Bailey Woods and Jackson Strip (Fig. 5).
The bearing trees closest to the field sites appeared to be representative of those sampled by surveyors within the township as a whole. Of the 18 bearing trees closest to (i.e. within 1.6 km of) Bailey Woods, there were 7 black oaks, 5 hickories, 4 black jack oaks, and 2 post oaks. Of the 8 bearing trees closest to the Jackson Strip, there were 5 black jack oaks, 2 hickories, and 1 black oak. Of the 10 bearing trees closest to the second-growth sites, there were 4 black jack oaks, 4 black oaks, and 2 hickories. A topographic map revealed that the elevations of the section midpoints and corners associated with these bearing trees were comparable to those of the field sites.
Southern red oak apparently was a common component of both upland and lowland presettlement forests (Table 4). However, after comparing upland samples of the township containing Oxford to those of the township south of Oxford, I saw that the abundance of southern red oak increased greatly going from the upland ridge areas in Oxford southward toward the Yocona River within Township 9S, 3W. In the vicinity of present-day sites in Oxford, bearing trees of southern red oak were not particularly common. Thus, unless southern red oak was confused with black oak by surveyors, it would appear that southern red oak has increased in frequency in upland forests in Oxford since the early 1800s.
Sweetgum was a frequently-sampled bearing tree in forests near major rivers. It apparently belonged to a mesophytic association dominated by American beech, white oak, American holly (Ilex opaca) and sweetgum. It would appear that sweetgum has increased in at least some upland forests in Oxford since the early 1800s (e.g., Bailey Woods, MF Strip, and Oak Grove; Fig. 5).
In addition to differences in species composition, upland bearing tree density was 4-6 times lower than the total density of trees >= 10 cm at either of the old-growth forests (52.06 bearing trees per ha vs. 295.22. stems per ha at Bailey Woods and 224.77 stems per ha at Jackson Strip). The bearing-tree density estimate was an underestimate of presettlement tree density because of selection bias. Nevertheless, these differences were substantial. Either the surveyors chose to ignore a substantial number of trees to get their bearings, or the density of canopy– sized trees was lower in presettlement times than it is in old-growth forests now.
Discussion. TREE SPECIES COMPOSITION OF UPLAND OAK COMMUNITIES IN NORTHERN MISSISSIPPI HAS CHANGED DRAMATICALLY SINCE THE EARLY 19TH CENTURY. Two striking observations reveal that upland old-growth forests bear little resemblance to presettlement oak communities in central Lafayette County in northern Mississippi. These are 1) the extraordinary of abundance of sweetgum in one of the oldgrowth forests (Bailey Woods), and the complete absence of sweetgum (Liquidambar styraciflua) among upland bearing trees, and 2) the numerical dominance of upland presettlement forests by black jack oak (Quercus marilandica), and the nearly complete absence of this species from both old growth forests today (Fig. 5). Another noteworthy difference is the greater abundance of hickory at one of the old growth sites (34.7% in Jackson Strip) compared with upland bearing trees (9%). The species composition of bearing trees in upland forests in and around Oxford, Mississippi is consistent with the anecdotal verbal descriptions of tree species composition and aspect of this area before European settlement in 1708 (Naime 1988), 1805 (Jennings 1947) and in the mid 1800s (Hilgard 1860), just following European settlement. Hilgard (1860) describes the species composition of the dominant trees in central Lafayette County, Mississippi as follows:
“The face of the country [is] either broken ridge lands with a very sandy soil, bearing an inferior growth, chiefly of Post Oak and Black Jack, commonly accompanied by the Scarlet … Oak and more or less, by the Spanish … Oaks; or broad ridges less high and abrupt, forming uplands more or less undulating, composed of Orange Sand covered by a stratum 2 to 4 feet in thickness, of yellow loam. The Spanish “Black” .. ., and Post Oak, and Hickory … form the prominent growth of these lands; but according to the scale of transition into the sandy ridge lands, more or less Black Jack and Scarlet Oak is added to the above [.] Page 296.
This description of tree species composition and soils is very similar to those of Rush Nutt in 1805 (Jennings 1947). Nutt adds, however, that the groundcover of the uplands was dominated by “sedge-grass” (most likely Andropogon spp. or Schizachyrium scoparium), suggesting that presettlement oak-dominated communities resembled thin woodlands or savannas and not the closed-canopy forests seen today. Nutt described the region around Oxford 30 years before cession to the United States. Furthermore, Nutt saw few settlements of Chickasaws near Oxford and no cultivation of sandy or sandy– clay ridges anywhere in north-central Mississippi (Jennings 1947; Johnson 2000). Nevertheless, Chickasaws might have set fires to maintain these areas as hunting grounds (Cushman 1899, Naime 1988). They were observed using fires to drive game (Nairne’s  observations in Naime 1988). Most Chickasaws settled primarily in three or four towns in the fertile BlackBelt Prairie (approximately 70 km east of Oxford) throughout most of the historic period before cession (Johnson 2000). In 1708, the area around their towns consisted of open oak woodlands, savannas, and prairies (Nairne 1988). Nairne claimed that the open character of the oak woodlands characterized most of the upland landscape of the Chickasaw Nation (including unsettled areas around present-day Oxford), but this claim was based on hearsay.
The circumstantial evidence presented here suggests that fire suppression in the 20,b century is largely responsible for the observed differences between old-growth and presettlement forests. Three of the four most common oak species in presettlement forests in central Lafayette County, Mississippi, black jack oak, post oak (Quercus stellata), and black oak (Q. velutina) are all generally associated with either frequently burned areas or extremely poor soils. Southern red oak (Q. falcata) appears to be common over a wide range of ecological conditions (Table 6, Hilgard 1860). Black jack and post oak frequently co-occur with longleaf pine in fire-prone savannas of the outer coastal plain of the southeastern United States (Harper 1914; Schwartz 1994). Historically, frequently-burned post oak savannas were common in the Ozark mountains and in Oklahoma (Rice and Penfound 1959; Cutter and Guyette 1991). In the northern midwest, many of the frequently-burned oak savannas are dominated by black oak (Curtis 1971; Dorney and Dorney 1989; Abrams 1992). All three species produce thick bark and are capable of resprouting after fires (Hodgkins 1958; Ferguson 1957, 1961; Harmon 1984; Lorimer 1985). Thus, three of the most common tree species in presettlement upland communities in this region of Mississippi also are the three most fire– tolerant in the species pool.
The most compelling indirect evidence in support of the hypothesis that fire suppression was responsible for the observed changes in tree species composition in northern Mississippi is the dramatic decline in black jack oak in presentday forests. Even allowing for the bias in bearing tree records, the difference between 42% and 1% relative frequency of black jack oak is substantial. Indeed, 42% may be an underestimate of the relative frequency of black jack oak in presettlement forests, because its short stature may have discouraged its use as a bearing tree (Schwartz 1994). Casual field observations throughout Lafayette County and adjacent Panola County to the west indicate that mature stands of black jack oak are now restricted to poor soils on exposed ridge tops, in open areas, at the edges of forests or near roads. The data presented here, however, indicate that black jack oak is uncommon even on ridge tops in Bailey Woods and Jackson Strip. Part of this may be due to the high silt content of the ridge soils sampled in this study. Black jack oak is apparently favored on sandier soils (Hilgard 1860).
In addition to the low abundance of black jack oak in present-day upland forests, most of the upland red oak species [(e.g., black oak, southern red oak, and scarlet oak (Q. coccinea)] are poorly represented in the smaller diameter classes in the old-growth forests. This is also true of post oak. These data and observations suggest that the light environment of presettlement upland forests in this part of Mississippi was very different than it is now. This is especially likely to be true if the low bearing tree densities are at all indicative of presettlement canopy-tree density. I hypothesize that many of the closed-canopy forests currently dominated by loblolly pine, sweetgum, southern red oak, and hickories historically had more open canopies and lower stem densities of mesophytic species in the understory. Such reduced competition likely permitted the regeneration of upland oaks in presettlement forests (Lorimer 1993). Replacement of oaks by fire-sensitive mesophytic species on all but the most xeric or infertile sites following extended fire-suppression appears to be occurring throughout the eastern United States (Lorimer 1985; Patterson and Sassaman 1988; Abrams 1992; Harrod and White 1998).
Another observation implicating the importance of fire suppression in upland forests today is the greater proportion of fire-sensitive species known to have been restricted primarily to forests on alluvial terraces, steep ravines, or floodplains before settlement (e.g., sweetgum, red maple, Lorimer 1984). Mesophytic forests in ravines are thought not to have experienced frequent fires before EuroAmerican settlement (McCarthy et al. 2001). Although apparently not common in upland forests in northern Mississippi in the early to mid 1800s (Nutt  in Jennings 1947; Hilgard 1860), sweetgum was quite common in this region by the 1900s, following a prolonged period of land clearing for cotton agriculture (Harper 1913; Dunston 1913; Lowe 1921). It was especially common in cut-over areas and oldfields (Lowe 1921). Thus, the increased abundance of sweetgum in some of the last remaining old-growth fragments of oak forest in northern Mississippi may reflect a greater abundance of this species in the region as a whole. In addition, however, active fire suppression may have contributed to the increased abundance of sweetgum in upland forests throughout the 20th century. Fires ignited by lightning or Native Americans likely played an important role in preventing the establishment of sweetgum in disturbed areas before extensive settlement by US citizens.
Curiously, despite the fact that wildfires set by settlers were common throughout Mississippi in the early 1900s (Dunston 1913), sweetgum was still common in cut-over areas in northern Mississippi at this time (Harper 1913; Lowe 1921). Most fires set by settlers occurred between growing seasons, however (Komarek 1964; Pyne 1982, Robbins and Myers 1992). Sweetgum resprouts more readily following fires occurring between growing seasons (winter, early spring) than following fires during the growing season (late spring, summer; Lotti 1956; Ferguson 1961). Hence, the clearing of forests combined with a lack of growing-season burning could have contributed to sweetgum’s dramatic increase in upland areas at this time. Most lightning-started fires appear to be started during the growing-season months in the eastern United States (March to September; Foti and Glenn 1990, Robbins and Myers 1992, Howe 1994, Olson and Platt 1995, Ruffner and Abrams 1998). There is considerable evidence that large areas burn during drought years or dry times of the year following lightning ignitions in the northeast (Ruffner and Abrams 1998), southeast (Robbins and Myers 1992), and midwest (Howe 1994) in the United States. Many fire-tolerant grassland and savanna species in the eastern United States show more dramatic increases in flowering following growing-season fires than following winter fires (Streng et al. 1993; Brewer and Platt 1994; Howe 1994). Some of these species [e.g. Pityopsis graminifolia (Michx.) Nutt.] occur in mature oak forests in northern Mississippi, but consistently flower at rates of less than 1% and are declining in these forests (J. S. Brewer, unpublished data). Most evidence suggests that lightning fires have been comparatively less common than human-set fires in oak forests (e.g., see Barden and Woods 1974; Martin 1990; Cutter and Guyette 1991, Delcourt and Delcourt 1998). Nevertheless, lightning fires accompanying occasional growing-season droughts may have been an important selection pressure in presettlement oak-dominated landscapes, and their importance should not be discounted.
FIRE-FILTERED RECOVERY OF OAK-DOMINATED COMMUNITIES. Here, I present a hypothesis of oak community development, offered as an alternative to models of forest succession based on shade tolerance, inhibition, or facilitation (Connell and Slayter 1977; Peet and Christensen 1987). Although the details of the mechanisms vary, in general, most succession-based models of oak forest development predict that rapidlygrowing, widely-dispersed, shade-intolerant species are eventually replaced by slower-growing shade-tolerant species during succession. In contrast, the filter hypothesis predicts that, repeated fires “filter out” seedlings of invasive, fire-sensitive pioneer species such as loblolly pine and sweetgum and promote advanced regeneration of fire-tolerant oaks (Harrington and Stephenson 1955; Lotti 1956; Ferguson 1961; Arthur et al. 1998). With fire suppression, however, these slow-growing oaks most likely could not compete in large gaps with faster-growing, fire-sensitive pioneer species (Peet and Christensen 1987; DeSteven 1991a, b). Thus, it is useful to view the fire regime in presettlement oak communities as a filter or sieve (sensu van der Valk 1981; Weiher and Keddy 1999; see also Gleason 1926) that directs changes in community structure, rather than just as a disturbance that “arrests” succession.
Shade tolerance is not the only variable affecting patterns of succession in upland forests of north-central Mississippi. The shared dominance of the canopy by large, slow-growing oaks (e.g., post oak, southern red oak, white oak) with smaller, younger (
Restoration of oak woodlands or savannas provides an excellent opportunity to test the fire– filtered recovery hypothesis. Prescribed burning is currently gaining acceptance as a management tool to restore presettlement oak woodlands (Laatsch and Anderson 2000). Much interest has focused on the effectiveness of restoring fire to closed-canopy oak forests to promote oak regeneration and groundcover plant diversity. The success of such restoration efforts so far is mixed (e.g., see Wendel and Smith 1986; Van Lear and Waldrop 1989; Watt et-al. 1993; Arthur et al. 1998; Laatsch and Anderson 2000). Alternatively, most studies of oldfields and early-successional precursors to oak forests of the eastern United States have focused on competitive interactions among oaks and mesophytic pioneers (e.g., DeSteven 1991a, b), microclimate of forest edges (Matlack 1993), herbivory (Gill and Marks 1991), or dispersal limitation (Lawson et al. 1999). Curiously, the role that fire plays in the recovery of oak woodlands from early-successional grasslands and savannas (e.g. “wildlife clearings”, oldfields, seed-tree regeneration cuts, naturally-disturbed gaps) has largely been ignored. Studies of ecophysiology, abiotic variables, competitive interactions, herbivory, and dispersal limitation in early-successional trees have yielded important insights on the mechanisms of succession (e.g., Bazzaz 1979; Facelli and Pickett 1991; DeSteven 1991a, b; Gill and Marks 1991; Matlack 1993; Lawson et al. 1999). Given the current interest in oak woodland restoration, fire is another important variable to be considered in studies of succession in oak communities.
1 This research was funded by the Department of Biology at the University of Mississippi in support of the laboratory portion of the General Ecology class. The USDA Sedimentation Laboratory in Oxford allowed me to look at aerial photographs, and the Lafayette County Circuit Clerk Office made the surveyor’s notes available to me.
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J. Stephen Brewer 2
Department of Biology, University of Mississippi, firstname.lastname@example.org
2 I thank the General Ecology classes of 1996-2000 at Ole Miss, and the following students in particular, Allison Hailman, Adam Sisk, Brian Talley, Ben Seale, Reagan Schiffer, Ashley Cain, and Jason Martin. I thank Steve Aquilani, Steve Ashley, and Edgar Leighton for field assistance, Cynthia Shearer for permission to work in Bailey Woods, Bonnie Krause for providing some historical documents, and M. C. Falkner and B. R. Brown for providing historical information.
Received for publication February 14, 2001, and in revised form July 2, 2001.
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