Tropical and north Pacific teleconnections influence fire regimes in pine-dominated forests of north-eastern California, USA

Aim To assess the importance of drought and teleconnections from the tropical and north Pacific Ocean on historical fire regimes and vegetation dynamics in north‐eastern California. Location The 700 km² study area was on the leeward slope of the southern Cascade Mountains in north‐eastern California. Open forests of ponderosa pine (Pinus ponderosa var. ponderosa Laws.) and Jeffrey pine (P. jeffreyi Grev. & Balf) surround a network of grass and shrub‐dominated meadows that range in elevation from 1650 to 1750 m. Methods Fire regime characteristics (return interval, season and extent) were determined from crossdated fire scars and were compared with tree‐ring based reconstructions of precipitation and temperature and teleconnections for the period 1700–1849. The effect of drought on fire regimes was determined using a tree‐ring based proxy of climate from five published chronologies. The number of forest‐meadow units that burned was compared with published reconstructions of the El Niño/Southern Oscillation (ENSO) and the Pacific Decadal Oscillation (PDO). Results Landscape scale fires burned every 7–49 years in meadow‐edge forests and were influenced by variation in drought, the PDO and ENSO. These widespread fires burned during years that were dryer and warmer than normal that followed wetter and cooler years. Less widespread fires were not associated with this wet, then dry climate pattern. Widespread fires occurred during El Niño years, but fire extent was mediated by the phase of the PDO. Fires were most widespread when the PDO was in a warm or normal phase. Fire return intervals, season and extent varied at decadal to multi‐decadal time scales. In particular, an anomalously cool, wet period during the early 1800s resulted in widespread fires that occurred earlier in the year than fires before or after. Main conclusions Fire regimes in north‐eastern California were strongly influenced by regional and hemispheric‐scale climate variation. Fire regimes responded to variation that occurred in both the north and tropical Pacific. Near normal modes of the PDO may influence fire regimes more than extreme conditions. The prevalence of widespread teleconnection‐driven fires in the historic record suggests that variation in the Pacific Ocean was a key regulator of fire regimes through its influence on local fuel production and successional dynamics in north‐eastern California.     

A review of the relationships between drought and forest fire in the United States

The historical and presettlement relationships between drought and wildfire are well documented in North America, with forest fire occurrence and area clearly increasing in response to drought. There is also evidence that drought interacts with other controls (forest productivity, topography, fire weather, management activities) to affect fire intensity, severity, extent, and frequency. Fire regime characteristics arise across many individual fires at a variety of spatial and temporal scales, so both weather and climate – including short‐ and long‐term droughts – are important and influence several, but not all, aspects of fire regimes. We review relationships between drought and fire regimes in United States forests, fire‐related drought metrics and expected changes in fire risk, and implications for fire management under climate change. Collectively, this points to a conceptual model of fire on real landscapes: fire regimes, and how they change through time, are products of fuels and how other factors affect their availability (abundance, arrangement, continuity) and flammability (moisture, chemical composition). Climate, management, and land use all affect availability, flammability, and probability of ignition differently in different parts of North America. From a fire ecology perspective, the concept of drought varies with scale, application, scientific or management objective, and ecosystem.     

Three centuries of fire in montane pine‐oak stands on a temperate forest landscape

Question: What was the role of fire in montane pine‐oak (Pinus‐Quercus) stands under changing human land uses on a temperate forest landscape in eastern North America? Location: Mill Mountain in the central Appalachian Mountains, Virginia, US. Methods: A dendroecological reconstruction of fire history was generated for four stands dominated by xerophytic pine and oak species. The fire chronology began under presettlement conditions following aboriginal depopulation. Subsequent land uses included European settlement, iron mining, logging, and US Forest Service acquisition and fire protection. Results: Fires occurred approximately every 5 years until 1930 without any evidence of a temporal trend in fire frequency. Burning ceased after 1930. Area‐wide fires affecting multiple pine stands were common, occurring at intervals of approximately 16 years. Most living pines became established during the late 1800s and early 1900s. Dead pines indicated that an older cohort established ca. 1730. Most hardwoods were established between the 1920s and 1940s. Conclusions: Except for fire protection, changes in land use had no discernible influence on fire frequency. Lightning ignitions and/or large fire extent may have been important for maintaining frequent burning in the 1700s, while fuel recovery may have constrained fire frequency during later periods. The disturbance regime appears to be characterized by frequent surface fires and occasional severe fires, insect outbreaks or other disturbances followed by pine recruitment episodes. Industrial disturbances appear to have had little influence on the pine stands. The greatest impact of industrial society is fire exclusion, which permitted hardwood establishment.     

Wildfire refugia in forests: Severe fire weather and drought mute the influence of topography and fuel age

Wildfire refugia (unburnt patches within large wildfires) are important for the persistence of fire‐sensitive species across forested landscapes globally. A key challenge is to identify the factors that determine the distribution of fire refugia across space and time. In particular, determining the relative influence of climatic and landscape factors is important in order to understand likely changes in the distribution of wildfire refugia under future climates. Here, we examine the relative effect of weather (i.e. fire weather, drought severity) and landscape features (i.e. topography, fuel age, vegetation type) on the occurrence of fire refugia across 26 large wildfires in south‐eastern Australia. Fire weather and drought severity were the primary drivers of the occurrence of fire refugia, moderating the effect of landscape attributes. Unburnt patches rarely occurred under ‘severe’ fire weather, irrespective of drought severity, topography, fuels or vegetation community. The influence of drought severity and landscape factors played out most strongly under ‘moderate’ fire weather. In mesic forests, fire refugia were linked to variables that affect fuel moisture, whereby the occurrence of unburnt patches decreased with increasing drought conditions and were associated with more mesic topographic locations (i.e. gullies, pole‐facing aspects) and vegetation communities (i.e. closed‐forest). In dry forest, the occurrence of refugia was responsive to fuel age, being associated with recently burnt areas (<5 years since fire). Overall, these results show that increased severity of fire weather and increased drought conditions, both predicted under future climate scenarios, are likely to lead to a reduction of wildfire refugia across forests of southern Australia. Protection of topographic areas able to provide long‐term fire refugia will be an important step towards maintaining the ecological integrity of forests under future climate change.     

Long‐term variability and rainfall control of savanna fire regimes in equatorial East Africa

Fires burning the vast grasslands and savannas of Africa significantly influence the global carbon cycle. Projecting the impacts of future climate change on fire‐mediated biogeochemical processes in these dry tropical ecosystems requires understanding of how various climate factors influence regional fire regimes. To examine climate–vegetation–fire linkages in dry savanna, we conducted macroscopic and microscopic charcoal analysis on the sediments of the past 25 000 years from Lake Challa, a deep crater lake in equatorial East Africa. The charcoal‐inferred shifts in local and regional fire regimes were compared with previously published reconstructions of temperature, rainfall, seasonal drought severity, and vegetation dynamics to evaluate millennial‐scale drivers of fire occurrence. Our charcoal data indicate that fire in the dry lowland savanna of southeastern Kenya was not fuel‐limited during the Last Glacial Maximum (LGM) and Late Glacial, in contrast to many other regions throughout the world. Fire activity remained high at Lake Challa probably because the relatively high mean‐annual temperature (~22 °C) allowed productive C₄ grasses with high water‐use efficiency to dominate the landscape. From the LGM through the middle Holocene, the relative importance of savanna burning in the region varied primarily in response to changes in rainfall and dry‐season length, which were controlled by orbital insolation forcing of tropical monsoon dynamics. The fuel limitation that characterizes the region's fire regime today appears to have begun around 5000–6000 years ago, when warmer interglacial conditions coincided with prolonged seasonal drought. Thus, insolation‐driven variation in the amount and seasonality of rainfall during the past 25 000 years altered the immediate controls on fire occurrence in the grass‐dominated savannas of eastern equatorial Africa. These results show that climatic impacts on dry‐savanna burning are heterogeneous through time, with important implications for efforts to anticipate future shifts in fire‐mediated ecosystem processes.     

Post-Glacial Climate–Fire Interactions Control Tree Composition of Mesic Temperate Forests in Eastern North America

Stand-scale gap-phase dynamics is generally viewed as the main driver of development in mesic deciduous forests of the temperate biome. Soil charcoal of temperate forests in eastern North America are unnoticed in most surveys, thus explaining why fire is undervalued as a driver of forest succession. The extent to which gap-phase, fire, or other processes are responsible for the regeneration and maintenance of mesic deciduous forests is unknown because paleoecological evidence is lacking. We tested the fire-driven succession hypothesis on the development of this major forest type. Based on charcoal ¹⁴C dates of two sites, 44 and 55 fires occurred since early Holocene, with a mean interval of 170 to 215 years. The vegetation of both sites followed comparable post-glacial trajectories consisting of three distinct periods. Conifers dominated the two first periods during 5200–6000 years and were replaced by hardwoods–conifers over the last 3500 years. The first period was represented by boreal conifers, whereas the second period, dominated by white pine (Pinus strobus) forests, persisted during 3000–4300 years. The third period marked the development of hardwood (sugar maple, Acer saccharum) forests. Fires occurred continuously on the sites since early Holocene likely under dry conditions during the conifer periods and cooler and moister conditions during the hardwood–conifer period. Recurrent fires appear with climate as key drivers of the long-term dynamics of several temperate forests in eastern North America. Similar studies on other temperate forests should be pursued to test the hypothesis of climate–fire interactions influencing tree composition change.

     

Spatial patterns of Mexican pine-oak forests under different recent fire regimes

Patterns of spatial arrangement, tree density, and species composition were compared in three unharvested pine-oak forests under different recent fire regimes: (1) an uninterrupted frequent fire regime, (2) fire exclusion, and (3) fire exclusion followed by the return of fire. Regeneration was dense and highly aggregated at all sites but the frequent-fire overstory was random to uniform in spatial distribution and relatively open while the fire-excluded sites had clumped overstory trees with a high density of smaller trees. Dominance by sprouting species was greatest at the fire-excluded sites. Mortality was spatially aggregated at all sites, consistent both with thinning by fire and density-dependent mortality, but competitive self-thinning appeared insufficient to counteract the increased tree density without fire. The return of fire after 29 years of exclusion reduced tree density but left overstory trees aggregated and led to vigorous oak and alder sprouting. Frequent fire disturbance is considered essential to maintain open pine forests; fire exclusion with or without subsequent fire appears to lead to denser forests dominated by smaller trees of sprouting species.     

Fire disturbance and forest structure in old‐growth mixed conifer forests in the northern Sierra Nevada,California

Question: This study evaluates how fire regimes influence stand structure and dynamics in old‐growth mixed conifer forests across a range of environmental settings. Location: A 2000‐ha area of mixed conifer forest on the west shore of Lake Tahoe in the northern Sierra Nevada, California. Methods: We quantified the age, size, and spatial structure of trees in 12 mixed conifer stands distributed across major topographic gradients. Fire history was reconstructed in each stand using fire scar dendrochronology. The influence of fire on stand structure was assessed by comparing the fire history with the age, size, and spatial structure of trees in a stand. Results: There was significant variation in species composition among stands, but not in the size, age and spatial patterning of trees. Stands had multiple size and age classes with clusters of similar aged trees occurring at scales of 113 ‐ 254 m². The frequency and severity of fires was also similar, and stands burned with low to moderate severity in the dormant season on average every 9–17 years. Most fires were not synchronized among stands except in very dry years. No fires have burned since ca. 1880. Conclusions: Fire and forest structure interact to perpetuate similar stand characteristics across a range of environmental settings. Fire occurrence is controlled primarily by spatial variation in fuel mosaics (e.g. patterns of abundance, fuel moisture, forest structure), but regional drought synchronizes fire in some years. Fire exclusion over the last 120 years has caused compositional and structural shifts in these mixed conifer forests.     

Differences in Spider Community Composition among Adjacent Sites during Initial Stages of Oak Woodland Restoration

Fire suppression has altered the uplands of northern Mississippi (U.S.A.). Once blanketed by open oak woodlands, this region is now experiencing mesophytic tree invasion, canopy closure, reduced oak regeneration, and herbaceous understory loss. In an attempt to reestablish historical conditions, experimental restoration was initiated through thinning and burning treatments. Our study, part of a comprehensive monitoring effort, is the first to examine the impact of oak woodland restoration on the spider community and associated habitat structure. Samples measuring a variety of environmental variables and utilizing an array of spider collecting techniques were taken within four habitats located at the restoration site: fire‐suppressed forest, moderately treated forest, intensely treated forest, and old field. Two main conclusions resulted from this study. (1) Open‐habitat specialists responded positively to increased canopy openness regardless of the availability of herbaceous vegetation. (2) Woodland restoration increased spider diversity, perhaps through the formation of diverse habitat structure and/or by altering species dominance patterns. A rise in open‐habitat specialist diversity was observed as treatment intensity increased, with no compensatory reduction in the diversity of forest specialists. What remains to be seen is whether the continued transition to open woodland habitat will result in losses of forest specialist species. More aggressive overstory tree thinning is currently being administered to encourage the growth of herbaceous grasses and forbs, which will permit future tests of a hypothesized decline in forest specialists.     

Multiple interacting ecosystem drivers: toward an encompassing hypothesis of oak forest dynamics across eastern North America

Many forests of eastern North American are undergoing a species composition shift in which maples (Acer spp.) are increasingly important while oak (Quercus spp.) regeneration and recruitment has become increasingly scarce. This dynamic in species composition occurs across a large and geographically complex region. The elimination of fire has been postulated as the driver of this dynamic; however, some assumptions underlying this postulate have not been completely examined, and alternative hypotheses remain underexplored. Through literature review, and a series of new analyses, we examined underlying assumptions of the “oak and fire” hypothesis and explored a series of alternative hypotheses based on well‐known ecosystem drivers: climate change, land‐use change, the loss of foundation and keystone species, and dynamics in herbivore populations. We found that the oak–maple dynamic began during a shift in climate regime‐from a time of frequent, severe, multi‐year droughts to a period of increased moisture availability. Anthropogenic disturbance on the landscape changed markedly during this same time, from an era of Native American utilization, to a time characterized by low population densities, to Euro‐American settlement and subsequent land transmogrification. During the initiation of the oak‐maple dynamic, a foundation species, the American chestnut, was lost as a canopy tree across a broad range. Several important browsers and acorn predators had substantial population dynamics during this period, e.g. white‐tailed deer populations grew substantially concurrent with increasing oak recruitment failure. In conclusion, our analyses suggest that oak forests are reacting to marked changes in a suite of interlocking factors. We propose a “multiple interacting ecosystem drivers hypothesis”, which provides a more encompassing framework for understanding oak forest dynamics.     

Human influences superseded climate to disrupt the 20th century fire regime in Jasper National Park,Canada

To enhance understanding of how climate and humans influenced historical fire occurrence in the montane forests of Jasper National Park, we crossdated fire-scar and tree age samples from 172 plots. We tested effects of drought and climatic variation driven by the El Niño-Southern Oscillation (ENSO) and Pacific North American (PNA) pattern on fire occurrence. We also tested whether local droughts were associated with ENSO, PNA, Pacific Decadal Oscillation and Atlantic Multidecadal Oscillation. We used a combination of instrumental and proxy-climate records to test whether climatic variation explained the absence of fire scars in our study area during the 20th century. From 1646 to 1915, 18 fires burned mainly during drier than average years. Drought years, but not fire years, were associated with positive ENSO and PNA indices, corresponding to warmer conditions with reduced snowpacks. Fire frequency varied through time, although no fire scars have formed since 1915. Potential recording trees present at all plots and climate conducive to fire over multiple years provide evidence that human influences superseded climatic variation to explain the lack of fire scars during the 20th century. Fire suppression significantly altered the fire regime after the formation of Jasper National Park, justifying the ongoing mechanical fuel treatments, prescribed and managed wildfires to improve forest resilience to climate change.     

Growth traits of juvenile American chestnut and red oak as adaptations to disturbance

American chestnut (Castanea dentata) was a dominant species in eastern North America prior to the importation of chestnut blight. In light of recent efforts to restore viable populations of chestnut in eastern forests, an increased understanding of its association with other co‐occurring, disturbance‐adapted oak species is necessary. We evaluated crown architecture and leaf morphology in juvenile chestnut and red oak (Quercus rubra) to assess potential differences in establishment strategies of both species. We also investigated differences in nonstructural carbohydrate reserves and whole tree biomass partitioning between species. Seedlings of both species were planted in forest stands treated either with midstory removal or small patch cuts, simulating potential restoration plantings. After 5–7 years, chestnut's allocation to its root system was lower than red oak's, with chestnut saplings instead diverting resources to branches and foliage. Chestnut had lower leaf area index, greater crown projection area, and higher specific leaf area than red oak, indicating the species may have an advantage in shaded understories. There were only minor differences in nonstructural root carbohydrate reserves, between red oak and American chestnut, indicating that chestnut may respond similarly to oak by resprouting after disturbances topkill young saplings. We suggest that American chestnut has morphological and physiological attributes that allow it to function as an opportunistic and plastic species that can utilize gaps to facilitate its canopy recruitment, yet still persist after occasional surface fire. This knowledge can guide restoration strategies for this iconic species of the eastern temperate forest region.     

Relationships of subalpine forest fires in the Colorado Front Range with interannual and multidecadal‐scale climatic variation

Aim An understanding of past relationships between fire occurrence and climate variability will help to elucidate the implications of climate‐change scenarios for future patterns of wildfire. In the present study we investigate the relationships between subalpine‐zone fire occurrence and climate variability and broad‐scale climate patterns in the Pacific and Atlantic Oceans at both interannual and multidecadal time‐scales. Location The study area is the subalpine zone of Engelmann spruce (Picea engelmannii) and subalpine fir (Abies lasiocarpa), and lodgepole pine (Pinus contorta) in the southern sector of the Rocky Mountain National Park, which straddles the continental divide of the northern Colorado Front Range. Methods We compared years of widespread fire from AD 1650 to 1978 for the subalpine zone of southern Rocky Mountain National Park, with climate variables such as measures of drought, and indices such as the El Niño–Southern Oscillation (ENSO), the Pacific Decadal Oscillation (PDO), and the Atlantic Multidecadal Oscillation (AMO). Results Years of extensive subalpine‐zone fires are significantly related to climate variability, phases of ENSO, the PDO, and the AMO, as well as to phase combinations of ENSO, the PDO, and the AMO at both interannual and centennial time‐scales. Main conclusions Years of extensive fires are related to extreme drought conditions and are significantly related to the La Niña phase of ENSO, the negative (cool) phase of the PDO, and the positive (warm) phase of the AMO. The co‐occurrence of the phase combination of La Niña‐negative PDO‐positive AMO is more important to fire occurrence than the individual influences of the climate patterns. Low‐frequency trends in the occurrence of this combination of climate‐pattern phases, resulting from trends in the AMO, are the primary climate pattern associated with periods of high fire occurrence (1700–89 and 1851–1919) and a fire‐free period (1790–1850). The apparent controlling influence of the AMO on drought and years of large fires in the subalpine forests of the Colorado Front Range probably applies to an extensive area of western North America.     

Mesophication in temperate Europe: A dendrochronological reconstruction of tree succession and fires in a mixed deciduous stand in Białowieża Forest

1. The shift from shade‐intolerant species to shade‐tolerant mesophytic species in deciduous and mixed forests of the temperate zone is well described in studies from North America. This process has been termed mesophication and it has been linked to changes in fire regime. Fire suppression results in the cessation of establishment of heliophytic, fire‐dependent tree species such as oak (Quercus) and pine (Pinus). Due to the scarcity of old‐growth forests in Europe, data on long‐term compositional changes in mixed forests are very limited, as is the number of studies exploring whether fire played a role in shaping the dynamics.
2. The aim of this study was to reconstruct tree succession in a 43‐ha natural mixed deciduous forest stand in Bia?owie?a Forest (BF), Poland using dendrochronological methods. In addition, the presence of aboveground fire legacies (charred and fire‐scarred deadwood) enabled the fire history reconstruction.
3. Dendrochronological data revealed tree establishment (Quercus) back to the end of the 1500s and fires back to 1659. Under a regime of frequent fires until the end of the 18th century, only oak and pine regenerated, sporadically. A shift in the fire regime in the first half of the 19th century triggered oak and pine cohort regeneration, then gradually spruce (Picea) encroached. Under an increasingly dense canopy and less flammable conditions, regeneration of shade‐tolerant Carpinus, Tilia, and Acer began simultaneously with the cessation of oak and pine recruitment.
4. Synthesis. The study reports the first evidence of mesophication in temperate Europe and proves that fire was involved in shaping the long‐term dynamics of mixed deciduous forest ecosystems. Our data suggest that fire exclusion promoted a gradual recruitment of fire‐sensitive, shade‐tolerant species that inhibited the regeneration of oak and pine in BF.

     

Regeneration Patterns of European Oak Species (Quercus petraea (Matt.) Liebl., Quercus robur L.) in Dependence of Environment and Neighborhood

Quercus robur L. (pedunculate oak) and Quercus petraea (Matt.) Liebl. (sessile oak) are two European oak species of great economic and ecological importance. Even though both oaks have wide ecological amplitudes of suitable growing conditions, forests dominated by oaks often fail to regenerate naturally. The regeneration performance of both oak species is assumed to be subject to a variety of variables that interact with one another in complex ways. The novel approach of this research was to study the effect of many ecological variables on the regeneration performance of both oak species together and identify key variables and interactions for different development stages of the oak regeneration on a large scale in the field. For this purpose, overstory and regeneration inventories were conducted in oak dominated forests throughout southern Germany and paired with data on browsing, soil, and light availability. The study was able to verify the assumption that the occurrence of oak regeneration depends on a set of variables and their interactions. Specifically, combinations of site and stand specific variables such as light availability, soil pH and iron content on the one hand, and basal area and species composition of the overstory on the other hand. Also browsing pressure was related to oak abundance. The results also show that the importance of variables and their combinations differs among the development stages of the regeneration. Light availability becomes more important during later development stages, whereas the number of oaks in the overstory is important during early development stages. We conclude that successful natural oak regeneration is more likely to be achieved on sites with lower fertility and requires constantly controlling overstory density. Initially sufficient mature oaks in the overstory should be ensured. In later stages, overstory density should be reduced continuously to meet the increasing light demand of oak seedlings and saplings.     

Californian mixed-conifer forests under unmanaged fire regimes in the Sierra San Pedro Mártir, Baja California, Mexico

Aim This study appraises historical fire regimes for Californian mixed‐conifer forests of the Sierra San Pedro Mártir (SSPM). The SSPM represents the last remaining mixed‐conifer forest along the Pacific coast still subject to uncontrolled, periodic ground fire. Location The SSPM is a north–south trending fault bound range, centred on 31°N latitude, 100 km SE of Ensenada, Baja California. Methods We surveyed forests for composition, population structure, and historical dynamics both spatially and temporally over the past 65 years using repeat aerial photographs and ground sampling. Fire perimeter history was reconstructed based on time‐series aerial photographs dating from 1942 to 1991 and interpretable back to 1925. A total of 256 1‐ha sites randomly selected from aerial photographs were examined along a chronosequence for density and cover of canopy trees, density of snags and downed logs, and cover of non‐conifer trees and shrubs. Twenty‐four stands were sampled on‐the‐ground by a point‐centred quarter method which yielded data on tree density, basal area, frequency, importance value, and shrub and herb cover. Results Forests experience moderately intense understory fires that range in size to 6400 ha, as well as numerous smaller, low intensity burns with low cumulative spatial extent. SSPM forests average 25–45% cover and 65–145 trees per ha. Sapling densities were two to three times that of overstory trees. Size‐age distributions of trees ≥ 4 cm dbh indicate multi‐age stands with steady‐state dynamics. Stands are similar to Californian mixed conifer forests prior to the imposition of fire suppression policy. Livestock grazing does not appear to be suppressing conifer regeneration. Main conclusions Our spatially‐based reconstruction shows the open forest structure in SSPM to be a product of infrequent, intense surface fires with fire rotation periods of 52 years, rather than frequent, low intensity fires at intervals of 4–20 years proposed from California fire‐scar dendrochronology (FSD) studies. Ground fires in SSPM were intense enough to kill pole‐size trees and a significant number of overstory trees. We attribute long fire intervals to the gradual build‐up of subcontinuous shrub cover, conifer recruitment and litter accumulation. Differences from photo interpretation and FSD estimates are due to assumptions made with respect to site‐based (point) sampling of fire, and nonfractal fire intensities along fire size frequency distributions. Fire return intervals determined by FSD give undue importance to local burns which collectively use up little fuel, cover little area, and have little demographic impact on forests.     

Restoring Perennial Warm‐Season Grasses as a Means of Reversing Mesophication of Oak Woodlands in Northern Mississippi

Fire suppression has removed an important ecological force previously responsible for shaping many plant communities throughout the world. Upland areas of north‐central Mississippi that have been protected from fire are now closed‐canopy forests including species known to be uncommon as bearing/witness trees in upland portions of the landscape (historically off‐site species) and sparse ground cover vegetation. Anecdotal evidence suggests that warm‐season grasses were prevalent in the understory of these communities, which could have provided more consistent fuel. We corroborate the historic presence of these grasses by looking at their natural co‐occurrence with oak regeneration (a requisite of self‐replacing stands of oaks found historically). Restoration of these communities has typically focused on burning and off‐site tree thinning. Utilizing a restoration experiment implementing these treatments, we found significantly reduced understory leaf litter in treatment areas. To test which variables associated with restoration treatments were most important for the survival of these grasses, we measured the effect of leaf litter removal and its interaction with environmental conditions on the survival of transplanted shoots. Survival of little bluestem increased with decreasing canopy density and decreasing leaf litter. Leaf‐litter removal did not increase survival, nor did it interact with either pre‐treatment leaf litter depth or canopy density. These results show that little bluestem benefits from conditions expected historically: increased light and possibly fire.     

Ecophysiological responses of two herbaceous species to prescribed burning, alone or in combination with overstory thinning

The oak-rich deciduous forests of the central Appalachian Mountains of eastern North America have changed significantly since the onset of effective fire suppression early in the 20th century. Those changes have resulted in progressively decreasing light and nutrient supplies to herbaceous perennial understory species. Application of ecological restoration treatments such as reintroduction of frequent dormant-season fire and overstory thinning to pre-suppression density often increase light, soil temperature and moisture, and short-term nutrient availability to pre-suppression levels. To persist in this environment, perennial understory herbs must be able to acclimate phenotypically to the very different resource supply combinations present with and without fire suppression. As part of a larger study of the response of the long-lived herbaceous perennials Desmodium nudiflorum and Panicum boscii to ecosystem restoration treatments in Ohio mixed-oak forests, this study examined the ecophysiological effects of prescribed burning (B) and the combination of burning and thinning (T + B) in mixed-oak forests in southern Ohio. Control (C) plants had significantly lower maximum photosynthetic rate (A(max)) than those in the treated plots. The enhancement of A(max) averaged 26.7% and 52.7% in the B and T + B treatments, respectively. Plants from the T + B plots had higher quantum yield, stomatal conductance, and photosynthetic nutrient use efficiency than B and C plants. B plants had greater intrinsic water use efficiency (WUE) than plants in the C or T + B treatments. Light saturation point (LSP), light compensation point (LCP), and "dark" respiration (DR) did not differ among treatments. Photosynthetic parameters did vary significantly between the species, but no significant treatment × species interactions were detected. Our results support the hypothesis that prescribed burning, especially when combined with overstory thinning, in these perennial herbs can result in phenotypic acclimation characterized by enhanced photosynthetic performance.     

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.     

Forest Pattern, Fire, and Climatic Change in the Sierra Nevada

In the Sierra Nevada, distributions of forest tree species are largely controlled by the soil-moisture balance. Changes in temperature or precipitation as a result of increased greenhouse gas concentrations could lead to changes in species distributions. In addition, climatic change could increase the frequency and severity of wildfires. We used a forest gap model developed for Sierra Nevada forests to investigate the potential sensitivity of these forests to climatic change, including a changing fire regime. Fuel moisture influences the fire regime and couples fire to climate. Fires are also affected by fuel loads, which accumulate according to forest structure and composition. These model features were used to investigate the complex interactions between climate, fire, and forest dynamics. Eight hypothetical climate-change scenarios were simulated, including two general circulation model (GCM) predictions of a 2 × CO₂ world. The response of forest structure,species composition, and the fire regime to these changes in the climate were examined at four sites across an elevation gradient. Impacts on woody biomass and species composition as a result of climatic change were site specific and depended on the environmental constraints of a site and the environmental tolerances of the tree species simulated. Climatic change altered the fire regime both directly and indirectly. Fire frequency responded directly to climate's influence on fuel moisture, whereas fire extent was affected by changes that occurred in either woody biomass or species composition. The influence of species composition on fuel-bed bulk density was particularly important. Future fires in the Sierra Nevada could be both more frequent and of greater spatial extent if GCM predictions prove true. Received 5 May 1998; accepted 4 November 1998.