Seed Bank Viability in Disturbed Longleaf Pine Sites

Some of the most species‐rich areas and highest concentrations of threatened and endangered species in the southeastern United States are found in wet savanna and flatwood longleaf pine (Pinus palustris Mill.) communities. Where intensive forestry practices have eliminated much of the natural understory of the longleaf ecosystem, the potential for reestablishment through a seed bank may present a valuable restoration opportunity. Longleaf pine sites converted to loblolly pine plantations and non‐disturbed longleaf sites on the Coastal Plain of North Carolina were examined for seed bank presence and diversity. Conducting vegetation surveys and examining the seed bank using the seedling emergence technique allowed for verification of the seed bank presence, as well as evaluation of the quality of the seed bank on disturbed longleaf pine sites. Forty‐three species and over 1,000 individuals germinated, and the seed banks of both the disturbed and non‐disturbed stand types contained species not noted in the vegetation survey. Although many of these species were considered weedy and typical of disturbance, numerous taxa were indicative of stable longleaf pine communities. This study confirms both the presence and quality of seed banks in highly disturbed former longleaf pine sites, suggesting that the seed bank may be an important tool in restoration efforts.     

Declines in plant species richness and endemic plant species in longleaf pine savannas invaded by <Emphasis Type="Italic">Imperata cylindrica</Emphasis>

Despite the widespread perception that non-native species threaten biodiversity, there are few documented cases of non-native species displacing rare or specialized native species. Here, I examined changes in plant species composition over 5 years during patch expansion of a non-native grass, Imperata cylindrica, in longleaf pine flatwoods in Mississippi, USA. I used a multivariate approach to quantify the degree of habitat specialization and geographic range of all species encountered. I examined losses of species collectively as a function of plant height (controlling for initial frequency) and then the relationship between height and the degree of association with longleaf pine flatwoods, disturbed habitats, and the outer Gulf Coastal Plain of the southeastern USA. Patch expansion resulted in dramatic declines in species richness and increases in ground-level shade at both sites in just 3 years. Most tall saplings, shrubs, and vines were not endemic to longleaf pine communities and were less likely to be displaced than short herbs, most of which were indicative of longleaf pine communities. These results suggest that invasion of longleaf pine communities by I. cylindrica will likely cause significant losses of short, habitat-specialists and reduce the distinctiveness of the native flora of these threatened ecosystems.     

Disturbance,seeds, restoration,and the importance of experiments and long‐term observations: the Editors' Award for 2005

Question: Can the geographic patterning of endemic plant species inform reserve selection in a region of high endemism? Location: The Southeastern Coastal Plain of North America, focusing primarily on the imperiled longleaf pine (Pinus palustris P. Miller) ecosystem. Methods: We documented the high level of plant endemism in the region, and characterized the endemic taxa into distributional subregions. Results: A total of 1630 plant taxa are endemic to the Coastal Plain, a large proportion of which are endemic to phytogeographical subregions within the Coastal Plain, with particularly large numbers of narrow endemics occurring in the East Gulf Coastal Plain and Florida Peninsula. Conclusions: This pattern of local endemism presents challenges in conserving the full biota of the region: a reserve system focusing on few and large conservation areas has theoretical benefits for long‐term management and viability, but will fail to capture many local endemics. We propose that the dispersed distribution of endemic species will require a mixture of large core reserves and smaller satellite reserves.     

Climate–tree growth relationships of longleaf pine (Pinus palustris Mill.) in the Southeastern Coastal Plain, USA

Knowledge of tree growth/climate response relationships is important to dendroecological studies and dendroclimatic reconstructions, particularly in the Southeastern Coastal Plain where few such studies have been attempted. To this end, we developed tree-ring chronologies of total ring width, earlywood width, and latewood width from longleaf pine (Pinus palustris Mill.) at three sites in the Southeastern Coastal Plain to examine the climate–growth relationships for this tree species. The length of these chronologies is unprecedented for southern pine chronologies in the Southeast. We compared the tree-ring chronologies to monthly temperature, precipitation, Palmer drought severity index (PDSI), and Palmer hydrological drought index (PHDI) data from the pertinent climate divisions. We found that PDSI and PHDI have the highest correlation with longleaf pine growth, and the strongest relationships between longleaf pine growth and these variables occur between July and November. Precipitation in the spring and summer was also positively related to growth at all sites. The relationship between temperature and growth was the weakest among all climate variables, but warm summer temperatures had a consistent, negative relationship with longleaf pine growth. The climate signal in the latewood was generally more robust than for total ring width and earlywood width.     

Longleaf pine (Pinus palustris Mill.) fire scars reveal new details of a frequent fire regime

Question: How frequent and variable were fire disturbances in longleaf pine ecosystems? Has the frequency and seasonality of fire events changed during the past few centuries? Location: Kisatchie National Forest, Western Gulf Coastal Plain, longleaf pine–bluestem ecosystem, in relatively rough topography adjacent to the Red River, Louisiana, USA. Methods: Cross‐sections of 19 remnant pines exhibiting 190 fire scars were collected from a 1.2‐km² area. Tree‐rings and fire scars were precisely dated and analysed for the purpose of characterizing past changes in fire and tree growth. Temporal variability in fire occurrence and seasonality was described for the pre‐ and post‐European settlement periods. Seasonality of historic fires was determined by the scar position within the rings. The relationship between fire and drought was investigated using correlation and superposed epoch analysis. Results: The mean fire return interval for the period 1650‐1905 was 2.2 years (range 0.5 to 12 yr). Significant new findings include: evidence for years of biannual burning, temporal variability in fire seasonality, an increase in fire frequency and percentage of trees scarred circa 1790, and synchronous growth suppression and subsequent release of trees coinciding with land‐use changes near the turn of the 20th century. Drought conditions appeared unrelated to the occurrence of fire events or fire seasonality. Conclusions: Multi‐century fire history records from longleaf pine ecosystems are difficult to obtain due to historic land‐use practices and the species high resistance to scarring; however, our results indicate potential for reconstructing detailed fire histories in this ecosystem. Fire scars quantitatively documented one of the most frequent fire regimes known. Fire regime information, such as the temporal variability in fire intervals, prevalence of late‐growing season fire events and biannual burning, provide a new perspective on the dynamics of longleaf pine fire regimes.     

Reptile and Amphibian Responses to Restoration of Fire‐Maintained Pine Woodlands

Fire‐maintained woodlands and savannas are important ecosystems for vertebrates in many regions of the world. These ecosystems are being restored by forest managers, but little information exists on herpetofaunal responses to this restoration in areas dominated by shortleaf pine (Pinus echinata). We compared habitat characteristics and herpetofaunal communities in restored pine woodlands to relatively unmanaged, second‐growth forests in the Ouachita Mountains of western Arkansas, USA. We found woodland restoration with periodic burning affected species differently; some species benefited, some species appeared negatively affected, but most species did not respond clearly either way. Overall reptile captures were significantly (p = 0.041) greater in pine‐woodlands than in unrestored forest; one species of snake and three species of lizards were captured more often in woodlands than unrestored forests. Among anurans, we found no significant difference in captures between woodlands and unrestored forests for any species. Among salamanders, we captured western slimy salamanders (Plethodon albagula) almost exclusively in unrestored forest, but captures of other species did not differ between the two treatments. Historically, the Ouachita region likely consisted of a mosaic that included both fire‐maintained habitats (woodlands, savannas, and prairies) and areas of denser forest on mesic sites that were less likely to burn. Consequently, landscapes that retain both open woodlands and denser, less‐intensely burned forest (in the form of unharvested greenbelts or separate stands) would likely promote and maintain a greater diversity of herpetofauna.     

Structure and Composition of Historical Longleaf Pine Ecosystems in Mississippi,USA

Longleaf pine (Pinus palustris) historically was a widespread ecosystem composed of a simple tree canopy and grasslands ground layer. After widespread loss of this ecosystem due to logging and fire exclusion, little quantitative information exists about historical structure for restoration goals. We identified composition in De Soto National Forest and Pearl River County, Mississippi, USA, and density, basal area, and percent stocking in Pearl River County using General Land Office surveys and US Forest Service Forest Inventory and Analysis surveys. Historical longleaf ecosystems were about 85% pine, with lesser amounts of broadleaf evergreen and oak species. Densities were about 175 to 180 trees/ha, mean tree diameters were 45 cm, and stocking was around 60% to 65%, which suggested longleaf pines were closed woodlands. Current forests are 38% to 57% pine, primarily loblolly, while longleaf pine is 2% to 8% of composition. Indeed, current longleaf pine composition across the Coastal Plain averages 3% and does not reach 10% at smaller landscape scales. Fire-sensitive broadleaf species of water oak, sweetgum, yellow-poplar, and red maple increased from about 0.5% composition to 2% to 10% of composition. Forests became twice as dense, at about 280 trees/ha to 330 trees/ha, with mean tree diameters of 22 cm. These results characterize conversion from open old growth longleaf forests, resulting in part from human maintenance, to successional forests due to human disruption of the historical ecosystem. It is important to remember structure and composition of historical forests for restoration and recognize wholesale changes so that successional forests do not become the new social and cultural baseline.     

Patterns of abundance of fire ants and native ants in a native ecosystem

Abstract 1. This correlational study examines the relationship between the red imported fire ant (Solenopsis invicta) and native ants in a longleaf pine savanna. Fire ants are frequently associated with a decline in native ants throughout the invaded range, but fire ant invasion is often coupled with habitat disturbance. Invasion of fire ants into the longleaf pine savanna provides an opportunity to examine the structure of the ant community in the absence of habitat disturbance. 2. Pitfall trapping was conducted within the longleaf pine savanna as well as across a naturally occurring soil moisture gradient, in plots that had been artificially watered. 3. Species richness did not vary as a function of fire ant density. There was an inverse relationship between native ant density and fire ant density, but this abundance pattern does not necessarily imply a causal link between fire ant invasion and native ant decline. For individual species, fire ant densities were negatively correlated with the densities of only two native ant species, including Solenopsis carolinensis, a native species that potentially limits the invasion of fire ants. Additionally, fire ants and native ants respond differently to soil moisture, with native ants favouring drier conditions than fire ants. 4. The possible exclusion of fire ants by some native ants, as well as differences in habitat preferences, provide alternative explanations for the frequently observed negative correlation between fire ants and native ants.     

Conservation of genetic diversity in old‐growth forest communities of the southeastern United States

Question: How do studies of the distribution of genetic diversity of species with different life forms contribute to the development of conservation strategies? Location: Old‐growth forests of the southeastern United States. Methods: Reviews of the plant allozyme literature are used to identify differences in genetic diversity and structure among species with different life forms, distributions and breeding systems. The general results are illustrated by case studies of four plant species characteristic of two widespread old‐growth forest communities of the southeastern United States: the Pinus palustris – Aristida stricta (Longleaf pine – wiregrass) savanna of the Coastal Plain and the Quercus – Carya – Pinus (Oak‐hickory‐pine) forest of the Piedmont. Genetic variation patterns of single‐gene and quantitative traits are also reviewed. Results: Dominant forest trees, represented by Pinus palustris(longleaf pine) and Quercus rubra (Northern red oak), maintain most of their genetic diversity within their populations whereas a higher proportion of the genetic diversity of herbaceous understorey species such as Sarracenia leucophylla and Trillium reliquum is distributed among their populations. The herbaceous species also tend to have more population‐to‐population variation in genetic diversity. Higher genetic differentiation among populations is seen for quantitative traits than for allozyme traits, indicating that interpopulation variation in quantitative traits is influenced by natural selection. Conclusion: Developing effective conservation strategies for one or a few species may not prove adequate for species with other combinations of traits. Given suitable empirical studies, it should be possible to design efficient conservation programs that maintain natural levels of genetic diversity within species of conservation interest.     

Patterns of Richness and Abundance in a Tropical African Leaf‐litter Herpetofauna1

I compared species richness and habitat correlates of leaf‐litter herpetofaunal abundance in undisturbed and selectively logged forests, and an abandoned pine plantation in Kibale National Park, Uganda. I sampled 50 randomly located 25 m² litter plots in each area during the wet and dry seasons in 1997. Ten anuran, five lizard, and three snake species were captured in plots over the study. Assemblage composition was most similar at logged and unlogged sites. The logged forest herpetofauna had higher species richness and abundance than the unlogged forest, but diversity was greater in the unlogged forest due to greater evenness. In contrast, the pine plantation site had the highest richness, abundance, and evenness of the three study sites, but species composition was distinct from the other areas. Herpetofaunal densities were significantly lower in all three areas during the dry season than in the wet season. During the dry season, soil moisture, litter mass, topography, shrub cover, and number of fallen logs were significant positive predictors of herpetofaunal presence in litter plots, but only soil moisture was significant in the wet season. The interaction of moisture and topography appears to be important in determining seasonal patterns of litter herpetofaunal distribution. Comparison of litter herpetofaunal studies across the tropics have shown that mid‐elevation faunas generally support fewer species than lowland faunas. Compared with other tropical mid‐elevation litter faunas, Kibale supports an intermediate number of species, but at lower densities than observed at any other mid‐elevation site reported in the literature.     

Quantifying Disturbance in Terrestrial Communities: Abundance–Biomass Comparisons of Herpetofauna Closely Track Forest Succession

Herpetofauna are frequently cited as ideal indicators for environmental and restoration monitoring due to their importance to ecological functioning, dominance of vertebrate biomass, and sensitivity to environmental change. Despite this relevance, herpetofauna have yet to be fully integrated into comprehensive, community‐scale restoration monitoring methodologies. We tested the applicability of one such method, abundance–biomass comparisons (ABCs), to monitoring restoration success in a longleaf pine ecosystem currently undergoing restoration via prescribed burning. W statistics (a metric of environmental perturbation) for herpetofaunal assemblages in longleaf pine stands managed under varying fire intervals closely tracked forest succession from recently burned longleaf pine stands to those characterized by longer times since last burn and high levels of hardwood encroachment. Our results suggest that ABCs may be an applicable method for terrestrial restoration monitoring that encompasses entire faunal assemblages and allows for generality in restoration or disturbance response. We emphasize, however, that applications of this method and others need to take into account the specific ecological characteristics of study organisms and the natural history of study sites to avoid incorrect interpretations of disturbance response and inappropriate management recommendations.     

Effects of longleaf pine planting density and other factors on stand structure and associated wildlife habitat

The primary objective of many longleaf pine (Pinus palustris) restoration programs is to enhance or restore habitat for wildlife dependent on herbaceous plant communities. Because herbaceous cover is inversely related to canopy cover, restoration programs often place restrictions on longleaf pine planting density. However, the influence of planting density on understory plant communities has been inadequately evaluated. Therefore, we initiated a study to examine the relative influences of planting density and other factors on overall understory composition and forage availability for white‐tailed deer (Odocoileus virginianus) and northern bobwhite (Colinus virginianus) in nine longleaf pine stands throughout the Coastal Plain of Alabama during 2017–2018. We found that coverage of herbaceous plants decreased 3.5%, coverage of woody plants decreased 2.4%, and coverage of northern bobwhite forage plants decreased 1.9% for each 1 m²/ha increase in longleaf pine basal area. However, planting density was not a significant predictor of current basal area, nor coverage of any functional group of plants we examined, likely because current longleaf pine density averaged only 46% (range = 30–64%) of seedling planting density. We did not detect an effect of prescribed fire on stand condition or understory plant communities, likely due to variability in fire timing and frequency. Our findings related to planting density were likely a function of low longleaf pine survival, which is not uncommon. Because of this and the inherent variability in growth rates for young longleaf pine stands, restoration programs should consider placing greater emphasis on post‐planting monitoring and management than planting density.     

The Initial Phase of a Longleaf Pine‐Wiregrass Savanna Restoration: Species Establishment and Community Responses

The significant loss of the longleaf pine‐wiregrass ecosystem in the southeastern United States has serious implications for biodiversity and ecosystem functioning. In response to this loss, we have initiated a long‐term and landscape‐scale restoration experiment at the 80,125 ha (310 mi²) Department of Energy Savannah River Site (SRS) located near Aiken, South Carolina. Aristida beyrichiana (wiregrass), an important and dominant grass (i.e., a “matrix” species) of the longleaf pine savanna understory, and 31 other herbaceous “non‐matrix” species were planted at six locations throughout SRS in 2002 and 2003. Of the 36,056 transplanted seedlings, 75% were still alive in June 2004, while mean 1–2 year survival across all planted species was 48%. Lespedeza hirta (hairy lespedeza) exhibited the greatest overall survival per 3 × 3 m cell at 95%, whereas Schizachyrium spp. (little bluestem) exhibited the greatest mean cover among individual species at 5.9%. Wiregrass survival and cover were significantly reduced when planted with non‐matrix species. Aggregate cover of all planted species in restored cells averaged 25.9% in 2006. High rates of survival and growth of the planted species resulted in greater species richness (SR), diversity, and vegetative cover in restored cells. Results suggest that the loss of the longleaf pine‐wiregrass ecosystem may be ameliorated through restoration efforts and illustrate the positive impact of restoration plantings on biodiversity and vegetative cover.     

Effects of Understory Vegetation Management on Brood Habitat for Northern Bobwhites

Abstract: Northern bobwhite (Colinus virginianus) populations have experienced severe declines for several decades, and declines have been particularly precipitous in the southern United States. These declines are partially attributable to large-scale conversions of potential habitat to short-rotation pine (Pinus spp.) forests managed for wood fiber production and fire exclusion in pine-dominated landscapes. We used standard arthropod sampling techniques, human-imprinted bobwhite chicks, and vegetation response to evaluate effects of different understory vegetation management practices on brood habitat quality within a commercially managed pine forest in Louisiana, USA, during 2002–2005. Specifically, we evaluated effects of mowing, prescribed fire during the growing season, prescribed fire in combination with imazapyr application, and no vegetation management on arthropod abundance and diversity, vegetation response, and the probability of bobwhite chicks successfully capturing an arthropod. Bobwhite chicks were more successful at capturing arthropods, and arthropod abundance and diversity were greatest, in plant communities managed using prescribed fire and imazapyr. Forest stands managed using a combination of fire and imazapyr were managed primarily to benefit the federally endangered red-cockaded woodpecker (Picoides borealis; RCW). Our findings suggest that management directed toward improving forest condition for RCWs improves habitat quality for brooding bobwhites. However, bobwhite chicks in our study area were less successful at capturing arthropods than were chicks in other studies in the southeastern United States. Brood-rearing habitat in pine forests similar to those we studied may be of generally poor quality, and could be related to precipitous declines of bobwhites in the West Gulf Coastal Plain. Managers should recognize that creating high-quality brood habitat in forests similar to those we studied will require more intensive management of understory vegetation than we studied.     

Herpetofaunal Responses to Restoration Treatments of Longleaf Pine Sandhills in Florida

The hypothesis that habitat restoration will provide for community reestablishment and the creation of habitat heterogeneity was examined with regards to the herpetofauna of longleaf pine sandhills in northwest Florida. The herpetofaunal response to restoration was examined in fire‐suppressed, hardwood‐dominated areas treated with (1) spring fire; (2) felling or girdling; or (3) a granular form of the herbicide hexazinone. No‐treatment controls were also included. Felling or girdling and herbicide plots were burned for fuel reduction two dormant seasons after initial treatment application. Additionally, data were collected in frequently burned reference sandhills to establish the target condition or restoration goal. Vegetation variables and herpetofaunal capture rates were compared among control and treatment areas. Two similarity indices were utilized to compare treatments and controls with reference sites, to examine restoration success. Restoration treatment effects were observed through reduced hardwood densities. Litter composition varied among control and treatment plots, with leaf litter being highest in areas lacking recent fire. Capture rates of some herpetofaunal species varied significantly among treatment plots. In 1997 similarity indices showed that spring‐burned and felling or girdling plots were more similar to the reference sandhills than the other plots. Treated plots were not significantly different from controls in 1998, a year of a severe drought.     

Using Multiscale Spatial Models to Assess Potential Surrogate Habitat for an Imperiled Reptile

In evaluating conservation and management options for species, practitioners might consider surrogate habitats at multiple scales when estimating available habitat or modeling species’ potential distributions based on suitable habitats, especially when native environments are rare. Species’ dependence on surrogates likely increases as optimal habitat is degraded and lost due to anthropogenic landscape change, and thus surrogate habitats may be vital for an imperiled species’ survival in highly modified landscapes. We used spatial habitat models to examine a potential surrogate habitat for an imperiled ambush predator (eastern diamondback rattlesnake, Crotalus adamanteus; EDB) at two scales. The EDB is an apex predator indigenous to imperiled longleaf pine ecosystems (Pinus palustris) of the southeastern United States. Loss of native open-canopy pine savannas and woodlands has been suggested as the principal cause of the species’ extensive decline. We examined EDB habitat selection in the Coastal Plain tidewater region to evaluate the role of marsh as a potential surrogate habitat and to further quantify the species’ habitat requirements at two scales: home range (HR) and within the home range (WHR). We studied EDBs using radiotelemetry and employed an information-theoretic approach and logistic regression to model habitat selection as use vs. availability. We failed to detect a positive association with marsh as a surrogate habitat at the HR scale; rather, EDBs exhibited significantly negative associations with all landscape patches except pine savanna. Within home range selection was characterized by a negative association with forest and a positive association with ground cover, which suggests that EDBs may use surrogate habitats of similar structure, including marsh, within their home ranges. While our HR analysis did not support tidal marsh as a surrogate habitat, marsh may still provide resources for EDBs at smaller scales.     

Reptile, amphibian, and lemur diversity of the Malahelo Forest, a biogeographical transition zone in southeastern Madagascar

Ambatorongorongo Mountain lies at the historical intersection betweenhumid, spiny, and littoral forests in southeastern Madagascar. We report theresults of surveys of the herpetofauna and lemurs occurring in Malahelo Forest,a small (<25 ha) forest fragment lying on the western slope ofAmbatorongorongo Mountain. There are at least 41 reptile, 11 amphibian, and 7lemur species in this forest, including several that are endemic to southeasternMadagascar and are at severe risk of extinction. The species richness of theMalahelo fauna is comparable to that of even the largest forest reserves in theregion. We also evaluate the similarity of the Malahelo herpetofauna to that ofnearby humid, spiny, and littoral forests to assess the biogeographic affinitiesof its amphibians and reptile assemblages. Both groups contain speciescharacteristic of each of the three surrounding forest types, but thebiogeographic patterns appear to differ for amphibians and reptiles. Overall,the herpetofauna and lemurs of the Malahelo Forest indicate that it is a remnantof a biogeographic transition zone between the major forest types ofsoutheastern Madagascar. The combination of high species richness, regionalendemics, and unique herpetofaunal and lemur assemblages should make MalaheloForest a high conservation priority, and we give recommendations for protectingwhat remains of this important transitional forest.     

Soil nitrogen dynamics as an indicator for longleaf pine restoration

Assessing the status of soil nutrients with their corresponding microbial communities provides important information about degraded soils during the restoration of coastal wet pine forests. Net nitrogen mineralization, nitrogen‐oxidizing bacteria (NOB), and soil microbial biomass were compared with patch‐derived volume along a 110‐year longleaf pine (Pinus palustris Mill.) chronosequence for identifying a trajectory and ecological benchmark during forest restoration. Net nitrogen mineralization rates decreased significantly in the maturing‐aged, pine patches, driven by a larger drop in net nitrification. Net nitrification and abundance of NOB were higher in young pine patches compared to soils from the maturing (86–110 years) pine patches. Gross nitrate fluxes followed the nonfungal portion of the soil microbial biomass along the chronosequence, declining in 64‐year‐old pine patches. Microbial biomass peaked in patches 17–34 years of age, but significantly declined in the older patches. Fungal biomass leveled off without decline. Ammonium was the major source of nitrogen within the maturing pine patches as well as the wetland patches, indicating that ammonium maintains longleaf pine during growth‐limiting conditions. Nitrate dominated during rapid tree growth, optimally in mesic conditions. The relative amounts of available ammonium to nitrate can be used to model nitrogen cycling in facultative‐wetland pine forests of the coastal United States as soils alternate between wet and mesic conditions. A key restoration benchmark occurred after 86 years of pine development when pine patch growth rates slowed, with lower numbers of NOB, when the nonfungal biomass leveled off, and net nitrification rates are at a minimum, during pine maturation.     

Scaling species dynamics in Pinus palustris communities: Effects of pine straw raking

Abstract. In the southeastern USA, harvest of pine straw sometimes involves mechanical raking of natural Pinus palustris (longleaf pine) communities. Little is known about the effects of raking nor how these effects may vary in time and space. In a two yr experiment, we examined the effects of mechanized raking on Pinus palustris dominated communities (scrub oak, dry savanna, and mesic savanna) by monitoring vegetation at seven spatial scales (0.01–100 m²). We measured floristic similarity and the proportion of species initially present that were gained (i.e. new species) or lost during four sampling periods. Relationships between spatial scale and these community attributes were analyzed using a repeated measures approach and functional response curves. Spatial scale clearly affected observed rates of species loss and floristic similarity; losses declined and floristic similarity increased as scale increased. We relate these patterns to expanding population sizes with scale and our inability to detect species reductions in large populations. Scale had little influence on species gains. The effects of raking did not differ across scales, but raking caused greater mean losses of species and greater mean changes in floristic similarity when mean values were calculated over all scales. Raking also increased the mean rate of species gains in the mesic savanna during one period. Otherwise, interaction effects of community and raking were largely absent from both mean values and response curves. Despite significant short‐term effects of raking, changes in species richness were minor.     

Effects of long-term fire exclusion on tree species composition and stand structure in an old-growth Pinus palustris (Longleaf pine) forest

Frequent fire is an integral component of longleaf pine ecosystems, creating environmental conditions favoring survival and growth of juvenile pines. This study examined stand structure, species composition, and longleaf pine regeneration in an old-growth tract of longleaf pine forest (Boyd Tract) experiencing long-term (>80 yr) fire exclusion in the Sandhills of North Carolina. Sampling of woody stems (i.e., 2.5 cm diameter at breast height) and tallies of longleaf pine seedlings were carried out in plots established randomly on upland, mesic areas and lowland, xeric areas within the Boyd Tract. Dominant woody species in mesic plots were black oak, hickories, and large, sparse longleaf pines. Xeric plots had high densities of turkey oak with the large longleaf pines, as well as higher frequencies of smaller longleaf stems. These differences between areas were associated with higher clay content of upland soils and higher sand content of lowland soils. Age-class frequency distributions for fire-suppressed longleaf pine following the last wildfire at the Boyd Tract approximately 80 yr ago contrasted sharply with data from an old-growth longleaf tract in southern Georgia (Wade Tract) that has been under a long-term frequent fire regime. Post-burn recruitment for the Boyd Tract wildfire appears to have been initially high on both site types. Longleaf pine recruitment diminished sharply on the mesic site, but remained high for 60 yr on the xeric site. Currently, longleaf pine regeneration is minimal on both site types; several plots contained no seedlings. Sharp contrasts in longleaf pine dominance and stand structure between the Boyd and Wade Tracts demonstrate the importance of large-scale disturbance, especially hurricanes and fire, in shaping the structure and function of longleaf pine ecosystems of the southeastern United States. In particular, long-term exclusion of fire on the Boyd Tract has altered stand structure dramatically by permitting hardwoods to occupy at high densities the characteristically large gaps between longleaf stems that are maintained by fire and other disturbances.