THE PRACTICAL ASPECTS OF IMPLEMENTING A CONTROLLED BURNING SCHEME IN THE KIDEPO VALLEY NATIONAL PARK (SECOND YEAR OF OPERATION)

A controlled burning scheme, in conjunction with experimental plots, was necessary in order to investigate the effects of fire on tree regeneration in an area which otherwise suffers frequent uncontrolled fires and where an important tree species, Acacia gerrardii Benth.var. gerrardii, was failing to regenerate. The second year's results are described. The scheme was not successful in excluding uncontrolled fires. However, further experience, indicating yet more problems, was gained.     

Ecological effects of changes in fire regimes in Pinus ponderosa ecosystems in the Colorado Front Range

Question: What is the relative importance of low‐ and high‐severity fires in shaping forest structure across the range of Pinus ponderosa in northern Colorado? Location: Colorado Front Range, USA. Methods: To assess severities of historic fires, 24 sites were sampled across an elevation range of 1800 to 2800 m for fire scars, tree establishment dates, tree mortality, and changes in tree‐ring growth. Results: Below 1950 m, the high number of fire scars, scarcity of large post‐fire cohorts, and lack of synchronous tree mortality or growth releases, indicate that historic fires were of low severity. In contrast, above 2200 m, fire severity was greater but frequency of widespread fires was substantially less. At 18 sites above 1950 m, 34 to 80% of the live trees date from establishment associated with the last moderate‐ to high‐severity fire. In these 18 sites, only 2 to 52% of the living trees pre‐date these fires suggesting that fire severities prior to any effects of fire suppression were sufficient to kill many trees. Conclusions: These findings for the P. ponderosa zone above ca. 2200 m (i.e. most of the zone) contradict the widespread perception that fire exclusion, at least at the stand scale of tens to hundreds of hectares, has resulted in unnaturally high stand densities or in an atypical abundance of shade‐tolerant species. At relatively mesic sites (e.g. higher elevation, north‐facing), the historic fire regime consisted of a variable‐severity regime, but forest structure was shaped primarily by severe fires rather than by surface fires.     

Effects of 38 years of wildfires on tree density in the Blue Mountains,Australia

Forests are vital for biodiversity, carbon storage and ecosystem services, but can be potentially threatened by fires. Given the significance of forests and fire in a changing climate, research into the long-term effects of fire on forests plays an important role in understanding the global carbon cycle by the forests functioning as a large terrestrial carbon sink or source. In this study, we used aerial photography from 1975 and 2013 to count the change in the number of trees in 560 dry sclerophyll plots (40 × 40m) in the Blue Mountains of Australia. We analysed the relationship between the number of fires and severe fires in that period on the change in numbers of trees. We found that the average response was an increase of 1 tree per plot over 38 years. The number of fires had a small positive effect on tree numbers; plots with 2 or 3 severe fires had 1 and 2 extra trees, respectively, than those without fire. One exception was a severe fire in 2001 that did not show this positive effect, probably because it corresponded with extensive drought. Our findings suggest that number of forest canopy trees is resilient to the number of fires and number of severe fires.     

火干扰因素对藏南地区高山松径向生长及气候响应特征的影响

森林火灾在二十一世纪内的发生频率逐步升高。大量研究发现森林火灾与树木生长之间存在有紧密的相关性。因此探究森林火灾对于树木生长的影响,分析火灾的不同强度对于树木径向生长是否有着显著的差异,将对于评估森林保护指标有着重要的实际意义。实验基于树木年轮学的研究方法,探究西藏林芝市本日山及九五六两场森林火灾对高山松径向生长的影响,分析在火灾前后高山松径向生长与气温及降水之间的响应关系。基于林芝市比日山及九五六两个火烧区域,建立了受轻度火影响和受中度火影响的高山松样地。利用树木年代学的方法,对年轮宽度指数与1961-2020年气温及降水分别进行相关分析,同时结合火灾的发生时间,将时间序列划分为:1961-2006年火灾前和2007-2021年火灾后。结果显示, 轻度火影响的高山松径向生长对气温和降水敏感,特别是在3-8月最高气温上呈现出显著性的增加。中度火影响的高山松径向生长则显著降低。火灾干扰明显抑制了高山松的径向生长,特别是火因子在平均气温和最高气温的干扰上对其径向生长具有明显抑制作用。受到轻度火影响的高山松径向生长在短期内有较为明显的增加趋势,但长期并不显著;而中度火影响的树木径向生长则在短期内不明显下降,但长期显著。受到轻度火影响的高山松中,其径向生长与最低气温之间呈显著的负相关,且使得高山松对外界环境的响应更为敏感,而中度火影响的高山松则并未表现出这种显著的响应状态。因此对于在高海拔地区而言,森林火灾对于树木径向生长有着显著的干扰的同时也提升树木与外界环境的响应程度。     

Modelling epiphyte metapopulation dynamics in a dynamic forest landscape

We combine simulations with spatial statistics to estimate the parameters of a metapopulation model for the epiphytic lichen Lobaria pulmonaria specializing on aspen ( Populus tremula ) and goat willow ( Salix caprea ) in Fennoscandian boreal old-growth forests. We estimated the parameters of a forest landscape model (FIN-LANDIS) by repeatedly running simulations and selecting the set of parameters for tree ecology and fire regime that reproduced empirical host tree density and spatial patterns. Second, we tested which variables were important in epiphyte colonization and estimated the dispersal kernel. Third, we run a metapopulation model for the lichen across the estimated landscape scenarios and selected values for the remaining parameters that reproduced the empirical patterns of epiphyte occurrence. There was little variation in predicted dynamics, occupancy and spatial patterns between replicate metapopulation simulations. However, more data would be required for accurately estimating the parameters of FIN-LANDIS primarily because of the inherent stochasticity in large scale forest fires. Following the beginning of fire suppression in the study area 150 years ago, the model predicts that lichen occupancy first increases but subsequently declines. The lower occupancy in the past than at present is explained by high rate of tree destruction by fires, which increases local extinction rate in patch-tracking metapopulations. In the absence of fires, the occupancy increases because of lower extinction rate, but without forest fires or alternative means of host tree regeneration, the lichen is predicted to go ultimately extinct because of severely reduced density of aspen and goat willow.     

Fire‐induced tree mortality in a neotropical forest: the roles of bark traits,tree size,wood density and fire behavior

Large‐scale wildfires are expected to accelerate forest dieback in Amazônia, but the fire vulnerability of tree species remains uncertain, in part due to the lack of studies relating fire‐induced mortality to both fire behavior and plant traits. To address this gap, we established two sets of experiments in southern Amazonia. First, we tested which bark traits best predict heat transfer rates (R) through bark during experimental bole heating. Second, using data from a large‐scale fire experiment, we tested the effects of tree wood density (WD), size, and estimated R (inverse of cambium insulation) on tree mortality after one to five fires. In the first experiment, bark thickness explained 82% of the variance in R, while the presence of water in the bark reduced the difference in temperature between the heat source and the vascular cambium, perhaps because of high latent heat of vaporization. This novel finding provides an important insight for improving mechanistic models of fire‐induced cambium damage from tropical to temperate regions. In the second experiment, tree mortality increased with increasing fire intensity (i.e. as indicated by bark char height on tree boles), which was higher along the forest edge, during the 2007 drought, and when the fire return interval was 3 years instead of one. Contrary to other tropical studies, the relationship between mortality and fire intensity was strongest in the year following the fires, but continued for 3 years afterwards. Tree mortality was low (≤20%) for thick‐barked individuals (≥18 mm) subjected to medium‐intensity fires, and significantly decreased as a function of increasing tree diameter, height and wood density. Hence, fire‐induced tree mortality was influenced not only by cambium insulation but also by other traits that reduce the indirect effects of fire. These results can be used to improve assessments of fire vulnerability of tropical forests.     

Wildfires affect tree growth and resilience in Northeastern China natural Dahurian larch forests: A dendrochronological perspective

Wildfires are natural and ubiquitous disturbances in boreal forests. Assessing their impacts on tree growth and resilience are particularly important to recognize the adaptation strategies of fire-tolerant species and forest succession in fire conditions. To date, the growth resilience of fire-tolerant species in boreal forests remains largely unquantified, and the drivers of resilience are poorly understood. Here, we measured the tree-ring widths of 99 fire-scarred trees from three sites in natural Dahurian larch (Larix gmelinii) forests. Three moderate-severity fire events in years 1987, 1990, and 2000 occurring at three sites were detected from the records of local forestry bureau. Based on tree-ring width data, we calculated resilience components (i.e., resistance, recovery, resilience and relative resilience) to quantify the responses of growth resilience in the larch trees to fires and analyzed their drivers at three sites. Results indicated that fires significantly reduced the tree growth. With the increasing tree age, these reductions were more pronounced. As for resilience components, our study showed a limited resistance but high recovery of tree growth against fires, and resistance tended to increase northwards but recovery showed the opposite, suggesting a growth-survival tradeoff was exhibited in Dahurian larch trees. With an increasing tree age, regional resistance and resilience showed a decreasing trend, whereas recovery and relative resilience showed an increasing trend. Resilience components were mainly affected by the climatic factors in spring. An increase in moisture availability enhanced resistance, a reduction in diurnal temperature range enhanced recovery, and an increase in mean temperature enhanced resilience and relative resilience. This study reveals that Dahurian larch could be even less favorable when faced with moderate or severe fire events, but a high capacity of recovery enables this species to adapt to the fire-prone condition. Moreover, this work highlights that the resilience of tree growth should be considered to understand tree behaviors and survival strategies of boreal forests following fires across fire-prone regions under future climate warming.     

Frequent fires reduce tree growth in northern Australian savannas: implications for tree demography and carbon sequestration

Tropical savannas are typically highly productive yet fire‐prone ecosystems, and it has been suggested that reducing fire frequency in savannas could substantially increase the size of the global carbon sink. However, the long‐term demographic consequences of modifying fire regimes in savannas are difficult to predict, with the effects of fire on many parameters, such as tree growth rates, poorly understood. Over 10 years, we examined the effects of fire frequency on the growth rates (annual increment of diameter at breast height) of 3075 tagged trees, at 137 locations throughout the mesic savannas of Kakadu, Nitmiluk and Litchfield National Parks, in northern Australia. Frequent fires substantially reduced tree growth rates, with the magnitude of the effect markedly increasing with fire severity. The highest observed frequencies of mild, moderate and severe fires (1.0, 0.8 and 0.4 fires yr^?1, respectively) reduced tree growth by 24%, 40% and 66% respectively, relative to unburnt areas. These reductions in tree growth imply reductions in the net primary productivity of trees by between 0.19 t C ha^?1 yr^?1, in the case of mild fires, and 0.51 t C ha^?1 yr^?1, in the case of severe fires. Such reductions are relatively large, given that net biome productivity (carbon sequestration potential) of these savannas is estimated to be just 1–2 t C ha^?1 yr^?1. Our results suggest that current models of savanna tree demography, that do not account for a relationship between severe fire frequency and tree growth rate, are likely to underestimate the long‐term negative effects of frequent severe fires on tree populations. Additionally, the negative impact of frequent severe fires on carbon sequestration rates may have been underestimated; reducing fire frequencies in savannas may increase carbon sequestration to a greater extent than previously thought.     

云贵高原抚仙湖近13300年的花粉/炭屑记录*

抚仙湖是云贵高原著名的断陷深水湖,其沉积物蕴藏着流域地质历史时期丰富的环境信息。对钻取自该湖的900cm 湖泊沉积物岩芯进行花粉/炭屑分析及花粉数据的主成分分析表明,抚仙湖流域的植被、气候与火灾在过去的13 300年经历了5个阶段的变化：(1)13 300—10 400cal．a BP,植被以松林为主,伴有山地暗针叶林和常绿阔叶林,表明该时期气候较为冷湿,森林火灾多发,在后期随着温度和湿度的降低,森林火灾愈加频繁。(2)10 400—5 700cal．a BP,松林收缩,常绿阔叶林扩张,出现一定数量的落叶阔叶林,显示该时期气候偏暖偏干;此阶段早期随着气候变暖变干森林火灾的发生延续上阶段高发的状态,直到9 500cal．a BP后随着湿度的增加森林火灾明显减少。(3)5 700—1 800cal．a BP,松林变化较小,常绿/落叶阔叶林比重增大,首次出现了暖热性的枫香林,显示该时期暖湿的气候特征,火灾发生频率低。(4)1 800—500cal．a BP,松林扩张,阔叶林收缩,本阶段后期草本植被比重开始增加,显示该时期气候相对冷干,森林火灾发生频率较高。(5)500cal．a BP至今,松林收缩,落叶阔叶树种增多,草本植物花粉明显增多,显示该时期气候温凉偏干,森林火灾发生频率降低。     

Tree Mortality Following Boreal Forest Fires Reveals Scale-Dependant Interactions Between Community Structure and Fire Intensity

Fire disturbance patterns influence forest communities at a range of spatial scales. Forest community structure may also influence fire disturbance patterns, because tree species vary in their fuel value and in their tolerance to fire damage. However, the influence of community structure on fire disturbance likely depends on latent ecological differences between fires and on the spatial scale at which patterns are observed. Using data on fire intensity, community structure, and post-fire tree survival in four systematically sampled boreal forest fires, we tested the hypotheses that: (1) patterns in post-fire tree survival reflect interactions between fire intensity and community structure; (2) these relationships change with the spatial scale of observation. To test the first hypothesis, we used information theoretic methods to compare eight generalized linear mixed effects models describing the influence of community structure and fire intensity on tree survival in a 500 m² sample plot, accounting for latent fire-to-fire differences in response. To test the scaling hypothesis, we reaveraged the data at nine successively larger spatial resolutions up to approximately 2 km², at each resolution tracking the parameter values of the best model. When fit to the plot-level data, the dominant feature of the best model was a strong intensity–survival correlation which varied from fire to fire, and depended on plot-level community structure. In some fires, community structure and survival became more tightly coupled at larger scales, whereas fire intensity became less important. These results support the view that fire disturbance patterns are influenced by cross-scale interactions between community structure and fire intensity.     

Modelling Variable Fire Severity in Boreal Forests: Effects of Fire Intensity and Stand Structure

It is becoming clear that fires in boreal forests are not uniformly stand-replacing. On the contrary, marked variation in fire severity, measured as tree mortality, has been found both within and among individual fires. It is important to understand the conditions under which this variation can arise. We integrated forest sample plot data, tree allometries and historical forest fire records within a diameter class-structured model of 1.0 ha patches of mono-specific black spruce and jack pine stands in northern Québec, Canada. The model accounts for crown fire initiation and vertical spread into the canopy. It uses empirical relations between fire intensity, scorch height, the percent of crown scorched and tree mortality to simulate fire severity, specifically the percent reduction in patch basal area due to fire-caused mortality. A random forest and a regression tree analysis of a large random sample of simulated fires were used to test for an effect of fireline intensity, stand structure, species composition and pyrogeographic regions on resultant severity. Severity increased with intensity and was lower for jack pine stands. The proportion of simulated fires that burned at high severity (e.g. >75% reduction in patch basal area) was 0.80 for black spruce and 0.11 for jack pine. We identified thresholds in intensity below which there was a marked sensitivity of simulated fire severity to stand structure, and to interactions between intensity and structure. We found no evidence for a residual effect of pyrogeographic region on simulated severity, after the effects of stand structure and species composition were accounted for. The model presented here was able to produce variation in fire severity under a range of fire intensity conditions. This suggests that variation in stand structure is one of the factors causing the observed variation in boreal fire severity.     

Thinner bark increases sensitivity of wetter Amazonian tropical forests to fire

Understory fires represent an accelerating threat to Amazonian tropical forests and can, during drought, affect larger areas than deforestation itself. These fires kill trees at rates varying from < 10 to c. 90% depending on fire intensity, forest disturbance history and tree functional traits. Here, we examine variation in bark thickness across the Amazon. Bark can protect trees from fires, but it is often assumed to be consistently thin across tropical forests. Here, we show that investment in bark varies, with thicker bark in dry forests and thinner in wetter forests. We also show that thinner bark translated into higher fire‐driven tree mortality in wetter forests, with between 0.67 and 5.86 gigatonnes CO₂ lost in Amazon understory fires between 2001 and 2010. Trait‐enabled global vegetation models that explicitly include variation in bark thickness are likely to improve the predictions of fire effects on carbon cycling in tropical forests.     

Pyric tree spatial patterning interactions in historical and contemporary mixed conifer forests,California, USA

Tree spatial patterns in dry coniferous forests of the western United States, and analogous ecosystems globally, were historically aggregated, comprising a mixture of single trees and groups of trees. Modern forests, in contrast, are generally more homogeneous and overstocked than their historical counterparts. As these modern forests lack regular fire, pattern formation and maintenance is generally attributed to fire. Accordingly, fires in modern forests may not yield historically analogous patterns. However, direct observations on how selective tree mortality among pre‐existing forest structure shapes tree spatial patterns is limited. In this study, we (a) simulated fires in historical and contemporary counterpart plots in a Sierra Nevadan mixed‐conifer forest, (b) estimated tree mortality, and (c) examined tree spatial patterns of live trees before and after fire, and of fire‐killed trees. Tree mortality in the historical period was clustered and density‐dependent, because trees were aggregated and segregated by tree size before fire. Thus, fires maintained an aggregated distribution of tree groups. Tree mortality in the contemporary period was widespread, except for dispersed large trees, because most trees were a part of large, interconnected tree groups. Thus, postfire tree patterns were more uniform and devoid of moderately sized tree groups. Postfire tree patterns in the historical period, unlike the contemporary period, were within the historical range of variability identified for the western United States. This divergence suggests that decades of forest dynamics without significant disturbances have altered the historical means of pyric pattern formation. Our results suggest that ecological silvicultural treatments, such as forest restoration thinnings, which emulate qualities of historical forests may facilitate the reintroduction of fire as a means to reinforce forest structural heterogeneity.     

An empirical approach to the analysis of forest stratification

Natural forest communities consist of different overlapping elementary subpopulations. Using the results of forest stratification in the preceding study, the properties of mean tree height for subpopulation in a stratified forest stand were examined. Mean tree height decreased as tree density per subpopulation increased. This relation was described by a simple mathematical model consisting of a power equation of tree density and two coefficients. The first coefficient or exponent of tree density was close to −1/2 in its expectation, while the other coefficient depended on life forms, especially in tropical forests. For tropical deciduous forests which suffered from seasonal forest fires, the latter coefficient was smaller than those for tropical evergreen and Japanese forests. This difference of the coefficient was not clear between tropical evergreen and Japanese forests and between deciduous and evergreen forests in Japan. In conclusion, the proposed model is similar to the 1/2 power law of tree height in man-made forests with simple architecture, and is designated the quasi-1/2 power law of tree height.     

Factors Controlling Vegetation Fires in Protected and Non-Protected Areas of Myanmar

Fire is an important disturbance agent in Myanmar impacting several ecosystems. In this study, we quantify the factors impacting vegetation fires in protected and non-protected areas of Myanmar. Satellite datasets in conjunction with biophysical and anthropogenic factors were used in a spatial framework to map the causative factors of fires. Specifically, we used the frequency ratio method to assess the contribution of each causative factor to overall fire susceptibility at a 1km scale. Results suggested the mean fire density in non-protected areas was two times higher than the protected areas. Fire-land cover partition analysis suggested dominant fire occurrences in the savannas (protected areas) and woody savannas (non-protected areas). The five major fire causative factors in protected areas in descending order include population density, land cover, tree cover percent, travel time from nearest city and temperature. In contrast, the causative factors in non-protected areas were population density, tree cover percent, travel time from nearest city, temperature and elevation. The fire susceptibility analysis showed distinct spatial patterns with central Myanmar as a hot spot of vegetation fires. Results from propensity score matching suggested that forests within protected areas have 11% less fires than non-protected areas. Overall, our results identify important causative factors of fire useful to address broad scale fire risk concerns at a landscape scale in Myanmar.     

Structural dynamics of a natural mixed deciduous forest in western Thailand

Abstract. Structural dynamics of a natural tropical seasonal – mixed deciduous – forest were studied over a 4-yr period at Mae Klong Watershed Research Station, Kanchanaburi Province, western Thailand, with particular reference to the role of forest fires and undergrowth bamboos. All trees > 5 cm DBH in a permanent plot of 200 m × 200 m were censused every two years from 1992 to 1996. The forest was characterized by a low stem density and basal area and relatively high species diversity. Both the bamboo undergrowth and frequent forest fires could be dominant factors that prevent continuous regeneration. Recruitment, mortality, gain (growth of survival tree plus ingrowth) and loss in basal area (by tree death) during the four years were 6.70%/yr, 2.91%/yr, 1.22%/yr and 1.34%/yr, respectively. Mortality was size dependent; middle size trees (30–50 cm) had the lowest mortality, while the smallest (5–10 cm) had the highest mortality. Tree recruitment was observed particularly in the first two years, mostly in the area where die-back of undergrowth bamboo occurred. The bias of the spatial distribution of recruitment to the area of bamboo die-back was significant and stronger than that to the forest canopy gaps. Successful regeneration of trees which survive competition with other herbs and trees after dieback of bamboo could occur when repeated forest fires did not occur in subsequent years. It is suggested that both the fire disturbance regime and bamboo life-cycle greatly influence the structure and dynamics of this seasonal tropical forest.     

火烧伤害对兴安落叶松树干径向生长的影响

在大兴安岭塔河林业局瓦拉干林场22支线截取3个兴安落叶松(Larix gmelinii)火疤圆盘,预处理和交叉定年后,测定垂直年轮(径向)和沿年轮(横向)方向2次火烧间的年轮宽度变化.通过生长趋势拟合、火烧后生长面积与正常(理论)生长面积比较,分析火烧伤害对兴安落叶松树干生长的影响.结果表明:火烧后横向年轮生长呈现出窄—宽—正常的趋势,这种趋势可以用3次曲线来拟合,即y=0.01x3-0.14x2+ 0.85x+ 0.56(DF=3,F=227.7,P＜0.0001,y为平均年轮宽度,x为偏离火烧点距离).在本文分析的3个圆盘中,火烧后的横向生长释放都出现在距火烧部位1/3范围内.火烧后径向上年轮宽度变化规律也较为明显,即在火灾后的几年时间内,年轮宽度值较火灾前都呈增加趋势.火烧后3个圆盘的径向生长释放持续时间分别为14、5a和11a.通过面积计算整体来看,火烧导致圆盘1较正常情况下损失了67.2％,圆盘2比正常情况下增加了6.6％,圆盘3增加了13.7％.上述分析表明,火烧导致的径向生长损失,可以较好地由横向和径向生长释放予以补偿,但是具体补偿多少因烧伤程度不同而有差异.火烧可直接改变树干径向生长变化,同时,这种变化也受其他因素影响,例如像树龄、树种特性、火灾间隔期等因素,这些因素的差异就可能导致补偿程度的不同.     

Customary Fire Regimes and Vegetation Structure in Gabon’s Bateke Plateaux

Most fires in Africa are anthropogenic yet remain understudied. Studies typically address managed fire, or the ??fire triad?? of early dry season-late dry season-suppression, and fire regimes which are annual or less, leaving unstudied the anthropogenic fire regimes that occur in the majority of African savannas. I take the case of the Bateke Plateaux area where burning today occurs both annually and semi-annually and measure the impacts of these regimes on savanna structure, measuring stem survival post fire and post fire regeneration of resprouts of the dominant savanna tree. While annual fires are hot and burn completely, semi-annual fires are cooler and patchy, favouring re-sprout survival and an escape route for small stems to mature into trees. This work extends the fire triad model to include an anthropogenic semi-annual regime which favours tree survival. The integration of local fire regimes into future studies will help increase our understanding of climate, vegetation dynamics as well as help orient policy and conservation.     

Ecological responses to el Niño-induced surface fires in central Brazilian Amazonia: management implications for flammable tropical forests

Over the past 20 years the combined effects of El Niño-induced droughts and land-use change have dramatically increased the frequency of fire in humid tropical forests. Despite the potential for rapid ecosystem alteration and the current prevalence of wildfire disturbance, the consequences of such fires for tropical forest biodiversity remain poorly understood. We provide a pan-tropical review of the current state of knowledge of these fires, and include data from a study in a seasonally dry terra firme forest of central Brazilian Amazonia. Overall, this study supports predictions that rates of tree mortality and changes in forest structure are strongly linked to burn severity. The potential consequences for biomass loss and carbon emissions are explored. Despite the paucity of data on faunal responses to tropical forest fires, some trends are becoming apparent; for example, large canopy frugivores and understorey insectivorous birds appear to be highly sensitive to changes in forest structure and composition during the first 3 years after fires. Finally, we appraise the management implications of fires and evaluate the viability of techniques and legislation that can be used to reduce forest flammability, prevent anthropogenic ignition sources from coming into contact with flammable forests and aid the post-fire recovery process.     

Fire frequency and tree canopy structure influence plant species diversity in a forest-grassland ecotone

Disturbances and environmental heterogeneity are two factors thought to influence plant species diversity, but their effects are still poorly understood in many ecosystems. We surveyed understory vegetation and measured tree canopy cover on permanent plots spanning an experimental fire frequency gradient to test fire frequency and tree canopy effects on plant species richness and community heterogeneity within a mosaic of grassland, oak savanna, oak woodland, and forest communities. Species richness was assessed for all vascular plant species and for three plant functional groups: grasses, forbs, and woody plants. Understory species richness and community heterogeneity were maximized at biennial fire frequencies, consistent with predictions of the intermediate disturbance hypothesis. However, overstory tree species richness was highest in unburned units and declined with increasing fire frequency. Maximum species richness was observed in unburned units for woody species, with biennial fires for forbs, and with near-annual fires for grasses. Savannas and woodlands with intermediate and spatially variable tree canopy cover had greater species richness and community heterogeneity than old-field grasslands or closed-canopy forests. Functional group species richness was positively correlated with functional group cover. Our results suggest that annual to biennial fire frequencies prevent shrubs and trees from competitively excluding grasses and prairie forbs, while spatially variable shading from overstory trees reduces grass dominance and provides a wider range of habitat conditions. Hence, high species richness in savannas is due to both high sample point species richness and high community heterogeneity among sample points, which are maintained by intermediate fire frequencies and variable tree canopy cover.