Fragment shape and tree species composition in tropical forests: a landscape level investigation

Fragmentation of tropical forest alters community composition as a result of changes in forest shape. This paper uses 22 hypotheses to test the effect of fragment shape on tree species composition in Ghana, West Africa, within biological categories of regeneration guild, rarity, phenology and dispersal. For both regenerating and mature trees, relationships between species composition and the shape of forest fragments were complex; almost half were significant but many failed to support the established hypotheses. Irregular shaped fragments had high proportions of regenerating, light‐demanding pioneers and mature, animal‐dispersed species. Species common to Ghana formed the foundation of communities in fragments of all shapes. Investigation at the landscape level indicated broad patterns of species change. Rigorous hypothesis testing is needed, following extensive demographic work on the ground, before population dynamics within tropical forest fragments can be comprehended fully and applied to conservation management.     

The fate of nutrients during fires in a tropical savanna

Abstract The nutrient loads contained in the grassy fuel before fires, and of ash subsequently, were compared to determine the fluxes of macronutrients, copper and zinc during fires at Kapalga in Kakadu National Park. The fluxes were estimated in three vegetation types: forest, woodland and open woodland. The magnitudes of the fluxes were greatest in the forest community where grassy fuel loads were highest at about 6.3 t ha^?1. In these sites, 54–94% of all measured nutrients in the fuel were transferred to the atmosphere during the fires. For each nutrient, the proportion transferred to the atmosphere as entrained ash was calculated by assuming that calcium was not volatilized during the fires. If the transfer of entrained ash represents local redistribution only, then rainfall accession and the deposition of these particu-lates should replace most of the losses of all nutrients except nitrogen (N). Estimated rates of biological fixation of N appear to be insufficient to replace the annual losses of N. It is therefore concluded that a regime of annual fires that completely burn the available grassy fuel would deplete N reserves in these savannas, unless there are other sources of biologically fixed N, which are unknown at present.     

片断化森林的边缘效应与自然保护区的设计管理

森林片断化引起了森林边缘物理环境与生物的变化。从森林边缘向林内,光辐射,温度、湿度、风等因素发生改变,这些反过 边缘上和边缘附近的生物产生极其重要的影响。有些研究表明,生物与非生物存在明显的边缘－核心的变化梯度,而有些则没有,除了影响边缘效应的客观因素（如边缘的取向、地势、年龄等）外,主观因素对这缘效应也有影响。边缘对生物与非生物的影响因片断化森林的面积、形状以及与其他森林片连接程度的不同而不同,     

Long-term potential for fires in estimates of the occurrence of savannas in the tropics

Aim   This study aims to improve the formulation and results of the Brazilian Center for Weather Forecasting and Climate Studies Potential Vegetation Model (CPTEC-PVM) by developing a new parameterization for the long-term occurrence of fire in regions of potential savannas in the tropics. Compared with the relatively slow processes of carbon uptake and growth in vegetation, fast mortality and biomass consumption by fires may favour grasses and reduce tree coverage.
Location   The tropics.
Methods   For finding large-scale relationships between fires and other environmental factors, we made two main simplifying assumptions. First, lightning is the most important source of ignition for natural fires. Second, over continental areas in the tropics, lightning is mainly related to the zonal flux of moisture transport.
Results   The parameterization of fire occurrence was built based on a simple empirical relationship, combining information on mean and intra-annual variance of the zonal wind.
Main conclusions   The implementation of this new relationship improved the formulation and the results of the CPTEC-PVM. As a result of this new parameter, the accuracy of the model in allocating the correct vegetation (seasonal forests) instead of savannas for large regions in India and Southeast Asia is now substantially higher than in previous studies.     

Contrasting demographics of tropical savanna and temperate forest eucalypts provide insight into how savannas and forests function. A case study using Corymbia clarksoniana from north‐eastern Australia

Eucalypts (Eucalyptus spp. and Corymbia spp.) dominate many communities across Australia, including frequently burnt tropical savannas and temperate forests, which receive less frequent but more intense fires. Understanding the demographic characteristics that allow related trees to persist in tropical savannas and temperate forest ecosystems can provide insight into how savannas and forests function, including grass–tree coexistence. This study reviews differences in critical stages in the life cycle of savanna and temperate forest eucalypts, especially in relation to fire. It adds to the limited data on tropical eucalypts, by evaluating the effect of fire regimes on the population biology of Corymbia clarksoniana, a tree that dominates some tropical savannas of north‐eastern Australia. Corymbia clarksoniana displays similar demographic characteristics to other tropical savanna species, except that seedling emergence is enhanced when seed falls onto recently burnt ground during a high rainfall period. In contrast to many temperate forest eucalypts, tropical savanna eucalypts lack canopy‐stored seed banks; time annual seed fall to coincide with the onset of predictable wet season rain; have very rare seedling emergence events, including a lack of mass germination after each fire; possess an abundant sapling bank; and every tropical eucalypt species has the ability to maintain canopy structure by epicormically resprouting after all but the most intense fires. The combination of poor seedling recruitment strategies, coupled with characteristics allowing long‐term persistence of established plants, indicate tropical savanna eucalypts function through the persistence niche rather than the regeneration niche. The high rainfall‐promoted seedling emergence of C. clarksoniana and the reduction of seedling survival and sapling growth by fire, support the predictions that grass–tree coexistence in savannas is governed by rainfall limiting tree seedling recruitment and regular fires limiting the growth of juvenile trees to the canopy.     

Climate regime shift and forest loss amplify fire in Amazonian forests

Frequent Amazonian fires over the last decade have raised the alarm about the fate of the Earth's most biodiverse forest. The increased fire frequency has been attributed to altered hydrological cycles. However, observations over the past few decades have demonstrated hydrological changes that may have opposing impacts on fire, including higher basin‐wide precipitation and increased drought frequency and severity. Here, we use multiple satellite observations and climate reanalysis datasets to demonstrate compelling evidence of increased fire susceptibility in response to climate regime shifts across Amazonia. We show that accumulated forest loss since 2000 warmed and dried the lower atmosphere, which reduced moisture recycling and resulted in increased drought extent and severity, and subsequent fire. Extremely dry and wet events accompanied with hot days have been more frequent in Amazonia due to climate shift and forest loss. Simultaneously, intensified water vapor transport from the tropical Pacific and Atlantic increased high‐altitude atmospheric humidity and heavy rainfall events, but those events did not alleviate severe and long‐lasting droughts. Amazonia fire risk is most significant in the southeastern region where tropical savannas undergo long seasonally dry periods. We also find that fires have been expanding through the wet–dry transition season and northward to savanna–forest transition and tropical seasonal forest regions in response to increased forest loss at the “Arc of Deforestation.” Tropical forests, which have adapted to historically moist conditions, are less resilient and easily tip into an alternative state. Our results imply forest conservation and fire protection options to reduce the stress from positive feedback between forest loss, climate change, and fire.     

Fire research for conservation management in tropical savannas: Introducing the Kapalga fire experiment

Abstract Fire is a dominant feature of tropical savannas throughout the world, and provides a unique opportunity for habitat management at the landscape scale. We provide the background and methodology for a landscape-scale savanna fire experiment at Kapalga, located in Kakadu National Park in the seasonal tropics of northern Australia. The experiment addresses the limitations of previous savanna fire experiments, including inappropriately small sizes of experimental units, lack of replication, consideration of a narrow range of ecological responses and an absence of detailed measurement of fire behaviour. In contrast to those elsewhere in the world, Australia's savannas are sparsely populated and largely uncleared, with fires lit primarily in a conservation, rather than pastoral, context. Fire management has played an integral role in the traditional lifestyles of Aboriginal people, who have occupied the land for perhaps 50 000 years or more. Currently the dominant fire management paradigm is one of extensive prescribed burning early in the dry season (May-June), in order to limit the extent and severity of fires occurring later in the year. The ecological effects of different fire regimes are hotly debated, but we identify geo-chemical cycling, tree demography, faunal diversity and composition, phenology, and the relative importance of fire intensity, timing and frequency, as critical issues. Experimental units (‘compartments’) at Kapalga are 15–20km² catchments, centred on seasonal creeks that drain into major rivers. Each compartment has been burnt according to one of four treatments, each replicated at least three times: ‘Early’- fires lit early in the dry season, which is the predominant management regime in the region; ‘Late’- fires lit late in the dry season, as occurs extensively in the region as unmanaged ‘wildfires’; ‘Progressive’- fires lit progressively throughout the dry season, such that different parts of the landscape are burnt as they progressively dry out (believed to approximate traditional Aboriginal burning practices); and ‘Unburnt’- no fires lit, and wildfires excluded. All burning treatments have been applied annually for 5 years, from 1990 to 1994. Six core projects have been conducted within the experimental framework, focusing on nutrients and atmospheric chemistry, temporary streams, vegetation, insects, small mammals, and vertebrate predators. Detailed measurements of fire intensity have been taken to help interpret ecological responses. The Kapalga fire experiment is multidisciplinary, treatments have been applied at a landscape scale with replication, and ecological responses can be related directly to measurements of fire intensity. We are confident that this experiment will yield important insights into the fire ecology of tropical savannas, and will make a valuable contribution to their conservation management.     

不同植被类型森林火灾及雷击火自组织临界性

利用黑龙江省大兴安岭林区呼中区 196 5～ 2 0 0 2年的雷击火数据、黑龙江省 1981～ 2 0 0 0年森林火灾数据及森林资源数据 ,对雷击造成的森林火灾的自组织临界性及不同植被类型条件下的自组织临界性作了研究 ,比较了在不同尺度和植被类型条件下火干扰的自组织临界性、自相似性 ,并与传统的森林火灾元胞自动机模型模拟的结果进行比较。结果表明 :中国黑龙江省不同森林类型的火干扰具有自组织临界行为 ,森林可燃物已经达到临界状态 ,其临界值在 1.8～ 2 .86之间 ,具有自相似性 ;当森林的面积过小时 ,森林火灾的“面积 -频率”分布曲线上会出现频率峰 ,表现出“有限面积效应”现象。     

Effects of fire and drought in a tropical eucalypt savanna colonized by rain forest

Aim This study documents the effects of multiple fires and drought on the woody structure of a north Australian savanna never grazed by domestic stock. Location The study was conducted in a 500 ha pocket of Eucalyptus‐dominated savanna surrounded by a late Quaternary lava flow. The flow is known as the Great Basalt Wall, located c. 50 km northeast of Charters Towers in semi‐arid north‐eastern Australia. This region was exposed to the largest 5‐year rainfall deficit on record between 1992 and 1996. Methods All individual woody plants were tagged within a 1.56 ha plot. Species were segregated into their habitat affinities (rain forest, ecotone, savanna) and regeneration strategy (resprouter, seeder). The survivorship of plants within these categories was analysed in relation to fire intensity from the first fire, and to each of four fires lit between 1996 and 2001. Results Before the first fire, the plot contained thirty‐one tree species including twenty‐one typical of the surrounding dry rain forest. These rain forest species were represented by small individuals and constituted <1% of the total basal area of woody plants. The basal area of savanna trees was 7.5 m² ha^?1 at the commencement of monitoring, although 31% had recently died and others had major crown damage. Further death of the drought debilitated savanna trees was substantial during the first year of monitoring and the basal area of live savanna trees declined to 1.1 m² ha^?1 after 5 years. Most species from both rain forest and savanna were classified as resprouters and are capable of regenerating from underground organs after fire. Species without this ability (rain forest seeders and ecotone seeders) were mostly eliminated after the first two consecutive fires. Among resprouters, survivorship declined as fire intensity increased and this was more pronounced for rain forest than for savanna species. Repeated burning produced a cumulative effect of decreasing survivorship for rain forest resprouters relative to savanna resprouters. Main conclusions The study provides evidence that savanna and rain forest trees differ in fire susceptibility and that recurrent fire can explain the restricted distribution of rain forest in the seasonally arid Australian tropics. The time of death of the savanna trees is consistent with the regional pattern after severe drought, and highlights the importance of medium term climate cycles for the population dynamics of savanna tree species and structure of Australian savannas.     

Soil temperature and depth of legume germination during early and late dry season fires in a tropical eucalypt savanna of north‐eastern Australia

Abstract Temperatures that significantly increase seed germination of some tropical legumes (i.e. 80–100°C) were documented in the topsoil during the passage of early (May) and late (October) dry season fires in a tropical eucalypt savanna of north‐eastern Australia. Elevated temperatures penetrated at least 30 mm into the soil during the higher‐intensity, late dry season fires, but were only detected at 10 mm during the early dry season fires. The depth from which germination of two native legume forbs Galactia tenuiflora and Indigofera hirsuta occurred was positively related to the temperature elevation in the topsoil and was greater after late compared with early dry season fires. A broader range in germination depth, resulting in higher seedling densities, was recorded for I hirsuta after late dry season fires. These results suggest that seedling emergence of native leguminous forbs is likely to occur at a greater density after late rather than early dry season fires in tropical eucalypt savannas of north‐eastern Australia. Therefore, the season of burning, as a result of its relationship to fire intensity, can influence species composition through its effect on seed germination.     

中度强度森林火灾对马尾松次生林土壤有机碳密度的影响

森林火灾作为森林非连续的干扰因子, 是生物地球化学循环的驱动因子, 显著改变生态系统的结构和功能及养分循环与能量传递, 引起森林碳库与碳分配格局的变化, 进而影响森林演替进程及固碳能力。该研究以广东省马尾松(Pinus massoniana)次生林为研究对象, 采用相邻样地比较法和空间代替时间法, 以野外调查采样与室内试验分析为主要手段, 定量研究突发性森林火灾对土壤有机碳密度的影响, 探讨森林火灾对土壤有机碳固持的影响机制。结果表明: 与对照相比, 森林火灾后的幼龄林、中龄林和成熟林的土壤有机碳密度分别为35.12、40.80和52.34 t·hm^-2, 依次降低了10.93%、8.52%和7.56%。相比对照, 幼龄林、中龄林和成熟林土壤剖面(0-60 cm)的土壤有机碳密度变化范围分别为5.04-7.76、5.26-10.27和6.33-13.58 t·hm^-2, 依次降低了2.51%-16.83%、1.31%-11.85%和1.09%-12.50%; 森林火灾显著降低了幼龄林和中龄林0-30 cm的土壤有机碳密度, 显著降低了成熟林0-20 cm的土壤有机碳密度。马尾松次生林土壤有机碳密度与土壤理化性质具有显著相关关系。通径分析表明, 对照样地和过火样地中, 土壤全氮含量均对土壤有机碳密度的直接作用最大, 土壤细根生物量对土壤有机碳密度的直接作用较小, 但其通过土壤全氮含量对土壤有机碳密度的影响均表现在间接作用上。嵌套方差分析表明, 土壤深度解释了土壤有机碳密度变异的70.60%, 林龄解释了其变异的25.35%, 森林火灾解释了其变异的2.34%。研究发现: 森林火灾减少了马尾松次生林各林龄的土壤有机碳密度。在水平方向上, 随着林龄增长, 土壤有机碳密度的减少幅度降低; 在垂直方向上, 土壤有机碳密度随着土壤土层深度加深而降低, 且随林龄增长减少幅度下降。研究森林火灾对森林生态系统土壤有机碳的影响, 有助于理解森林生态系统土壤碳固持和碳循环过程, 对制定旨在减缓全球变化的科学合理的林火管理策略具有重要意义。     

Fire alters diversity,composition, and structure of dry tropical forests in the Eastern Ghats

Fire is known to have dramatic consequences on forest ecosystems around the world and on the livelihoods of forest‐dependent people. While the Eastern Ghats of India have high abundances of fire‐prone dry tropical forests, little is known about how fire influences the diversity, composition, and structure of these communities. Our study aimed to fill this knowledge gap by examining the effects of the presence and the absence of recent fire on tropical dry forest communities within the Kadiri watershed, Eastern Ghats. We sampled plots with and without evidence of recent fire in the Eswaramala Reserve Forest in 2008 and 2018. Our results indicate that even though stem density increases in the recently burned areas, species richness is lower because communities become dominated by a few species with fire resistance and tolerance traits, such as thick bark and clonal sprouting. Further, in the presence of fire, the size structure of these fire‐tolerant species shifts toward smaller‐sized, resprouting individuals. Our results demonstrate that conservation actions are needed to prevent further degradation of forests in this region and the ecosystem services they provide.     

Contrasting fire-related resilience of ecologically dominant ants in tropical savannas of northern Australia

This paper examines the role of fire in mediating the relative abundance of two of the world's major ecologically dominant ant genera, Iridomyrmex and Oecophylla, where they coexist across the tropical savanna landscapes of northern Australia. These taxa have contrasting biogeographical histories, which are predicted to lead to contrasting responses to fire. Iridomyrmex is an autochthonous Australian genus that has radiated primarily in the arid zone; as such, its abundance is predicted to be promoted by frequent fire because this maintains an open habitat. In contrast, Oecophylla is a genus of leaf‐nesting ants occurring in the canopies of Old World tropical rainforest, and is a recent arrival to Australia in geological time; the abundance of these ants is predicted to decline under frequent fire. We test these predictions using results from a landscape‐scale fire experiment, where three experimental fire regimes (including no fire) were applied to replicated subcatchments over a 5‐year period. Using sweep nets, ants were sampled in the grass layer (the habitat layer of greatest overlap between Iridomyrmex and Oecophylla) in eucalypt woodland (canopy cover < 30%) and open eucalypt forest (canopy cover about 50%) habitats. A total of 27 species from 11 genera were collected during the study; eight were common enough for statistical analysis, and the abundances of four of these were significantly affected by fire treatment. As predicted, the abundance of Iridomyrmex was promoted by fire, whereas that of Oecophylla declined. These changes occurred only under late‐season (relatively high intensity) fires, and for Oecophylla occurred only in open forest (not woodland) habitat. This fire‐mediated relationship between Iridomyrmex and Oecophylla mirrors the much broader, ecosystem‐wide dynamic between eucalypt‐dominated savanna and rainforest in tropical Australia, with savannas dominated by fire‐resistant sclerophyll elements of Australian origin, and rainforest dominated by fire‐sensitive mesophyll elements of South‐East Asian origin.     

Edge Structure Determines the Magnitude of Changes in Microclimate and Vegetation Structure in Tropical Forest Fragments1

Edge structure is one of the principal determinants of the extent and magnitude of edge effects in forest fragments. In central Amazonia, natural succession at forest edges typically produces a dense wall of vegetation dominated by Cecropia spp. that buffers the forest interior. Fire encroachment into forest edges, however, eliminates the soil seed bank, enhances plant mortality, and promotes succession to an open, Vismia–dominated edge that does not buffer the forest interior. Contrasting open, fire–encroached forest edges and closed, non–fire–encroached edges were examined in central Amazonia to assess the effects of edge structure on microclimate and vegetation structure in tropical forest fragments. Edge penetration distances for most microclimate and vegetation structure variables were as much as two to five times greater at open edges than at closed edges. The magnitude of these differences suggests that edge structure is one of the main determinants of microclimate and vegetation structure within tropical forest fragments. Edge effects also varied systematically with fragment area. For a given edge type, 100–ha fragments had consistently lower canopy height, higher foliage density, higher temperature, a higher rate of evaporative drying, lower leaf litter moisture content, and lower litter depth than continuous forest, at all distances from the forest edge. These differences, however, were relatively minor compared to the striking differences in edge penetration between open and closed forest edges. For organisms in small fragments, the difference between open and closed edges may be the difference between total edge encroachment on one hand and an effective nature reserve on the other, relatively independent of absolute fragment area.     

Fire in the tropical gallery forests of Belize

ABSTRACT. Historical records of burning, field observations, and a manipulation experiment were used to evaluate the extent and impact of fire in a system of gallery forests in the Mountain Pine Ridge savanna, Belize. The outer boundaries of gallery forests are fire-prone zones, but fires rarely intrude into these forests. This is attributed to the existence of fire-tolerant trees in the outer zone, which preserve a forest interior of low flammability. Occasional fire incursions are patchily distributed and partially inhibited by slope convexities. Intrusions consume litter and root mats and destroy seedlings and saplings, but create a wide variety of subsequent light regimes depending upon the degree of canopy destruction. At most sites, partial canopy cover persists and seedlings of a subset of forest tree species establish preferentially. Early survivorship of these seedlings is comparable to those established in undamaged forest. Where canopy opening is severe, a secondary succession is initiated, with large numbers of herbaceous plants deriving from the seed bank. Gallery forests contain core zones into which fire very rarely intrudes, and peripheral zones that experience fire incursions that are patchily distributed in space and time. In the latter zones fire incursions play a role comparable to that of canopy gaps in continuous forests, but also create a unique class of micro-habitats to which a subset of tree species is specialized. The fire regime over the recent past in this gallery forest system appears to have had an enriching, rather than a depauperizing, effect on the forest communities, and such systems represent plausible refugia for forest species in fire-prone landscapes.     

Fire effects and ecological recovery pathways of tropical montane cloud forests along a time chronosequence

Tropical montane cloud forests (TMCFs) harbour high levels of biodiversity and large carbon stocks. Their location at high elevations make them especially sensitive to climate change, because a warming climate is enhancing upslope species migration, but human disturbance (especially fire) may in many cases be pushing the treeline downslope. TMCFs are increasingly being affected by fire, and the long‐term effects of fire are still unknown. Here, we present a 28‐year chronosequence to assess the effects of fire and recovery pathways of burned TMCFs, with a detailed analysis of carbon stocks, forest structure and diversity. We assessed rates of change of carbon (C) stock pools, forest structure and tree‐size distribution pathways and tested several hypotheses regarding metabolic scaling theory (MST), C recovery and biodiversity. We found four different C stock recovery pathways depending on the selected C pool and time since last fire, with a recovery of total C stocks but not of aboveground C stocks. In terms of forest structure, there was an increase in the number of small stems in the burned forests up to 5–9 years after fire because of regeneration patterns, but no differences on larger trees between burned and unburned plots in the long term. In support of MST, after fire, forest structure appears to approximate steady‐state size distribution in less than 30 years. However, our results also provide new evidence that the species recovery of TMCF after fire is idiosyncratic and follows multiple pathways. While fire increased species richness, it also enhanced species dissimilarity with geographical distance. This is the first study to report a long‐term chronosequence of recovery pathways to fire suggesting faster recovery rates than previously reported, but at the expense of biodiversity and aboveground C stocks.     

边缘效应及其对森林生态系统影响的研究进展

边缘效应是生态学和生物保护的重要概念之一,它在研究生态系统尺度和景观生态系统尺度的能量流和物质流等生态过程中具有重要作用.本文对边缘效应的内涵、特征、定量评价(包括定量分析基础、强度、影响区、模型等)、应用研究等方面进行阐述,分析了边缘效应研究中存在的不足,总结了边缘效应对森林生态系统的影响及其研究方向,以期为森林经营、保护区管理等生产实践提供借鉴.     

Nitrogen deposition in tropical forests from savanna and deforestation fires

We used satellite‐derived estimates of global fire emissions and a chemical transport model to estimate atmospheric nitrogen (N) fluxes from savanna and deforestation fires in tropical ecosystems. N emissions and reactive N deposition led to a net transport of N equatorward, from savannas and areas undergoing deforestation to tropical forests. Deposition of fire‐emitted N in savannas was only 26% of emissions – indicating a net export from this biome. On average, net N loss from fires (the sum of emissions and deposition) was equivalent to approximately 22% of biological N fixation (BNF) in savannas (4.0 kg N ha^?1 yr^?1) and 38% of BNF in ecosystems at the deforestation frontier (9.3 kg N ha^?1 yr^?1). Net N gains from fires occurred in interior tropical forests at a rate equivalent to 3% of their BNF (0.8 kg N ha^?1 yr^?1). This percentage was highest for African tropical forests in the Congo Basin (15%; 3.4 kg N ha^?1 yr^?1) owing to equatorward transport from frequently burning savannas north and south of the basin. These results provide evidence for cross‐biome atmospheric fluxes of N that may help to sustain productivity in some tropical forest ecosystems on millennial timescales. Anthropogenic fires associated with slash and burn agriculture and deforestation in the southern part of the Amazon Basin and across Southeast Asia have substantially increased N deposition in these regions in recent decades and may contribute to increased rates of carbon accumulation in secondary forests and other N‐limited ecosystems.     

Impacts of wind disturbance on fragmented tropical forests: A review and synthesis

Abstract Wind disturbance is an important ecological force in the tropics, especially in the cyclonic and hurricane zones from about 7–20° latitude. Damage from intense winds may be especially severe in fragmented forests because of their abrupt artificial margins and denuded surrounding landscapes. We review available information on the effects of windstorms on fragmented forests, synthesizing studies from Australasia, Amazonia and elsewhere in the tropics. Wind damage in fragmented landscapes can be influenced by a range of factors, such as forest‐edge orientation, edge structure, the size of nearby clearings and local topography. We argue that wind disturbances are likely to interact with, and exacerbate, a range of deleterious environmental changes in fragmented forests. Among the most important of these are altered forest structure, shifts in plant species composition, exotic‐plant invasions, reduced carbon storage and elevated vulnerability to fire. The damaging impacts of winds on fragmented forests could potentially increase in the future, particularly if global warming leads to increasingly severe or frequent windstorms.     

Coexistence and divergence of tropical dry forests and savannas in southern Mexico

Aim The purpose of the study was to assess the degree of floristic differentiation between tropical dry forest (TDF) and savanna occurring in a single landscape. This comparison provides information on the responses of vegetation to the prevailing environmental conditions, while it also allows us to make inferences about large‐scale events and processes, both biogeographical and evolutionary. Our approach included three levels of analysis: (1) taxonomic, (2) morphological and (3) vegetational. Location The seasonal dry tropical landscape in the Nizanda region, Oaxaca State, southern Mexico. The landscape comprises a complex vegetation mosaic in which tropical dry forest and savannas are the most conspicuous components. Methods Comparisons between TDF and savanna were based on inventories for these communities produced after 8 years of botanical survey. At the taxonomic level, the relative representation of taxa of different hierarchical levels in each community was examined. Morphological analyses required the classification of species on each of three criteria: (1) growth form, (2) life form and (3) growth habit. Vegetation level analysis was based on the frequencies of taxa in one hundred 100‐m² composition plots with which matrices of binary data were constructed for species, genera and families. These were subjected to classification analysis with Ward's method and using Euclidean distances as the dissimilarity algorithm. Results The combined flora for both communities comprised 600 species, 375 genera and 94 families; between them they shared 31, 40 and 34 taxa, respectively. The corresponding Sørensen similarity values were 10%, 21% and 72%, respectively. Ranking genera and families according to their species richness displayed large differences between savanna and TDF. Large differences between these communities were observed for Acanthaceae, Cactaceae, Euphorbiaceae and Mimosaceae, whereas Fabaceae and Asteraceae had similar high ranks according to the species richness in the two systems. The growth form spectrum diverged between the two communities, with TDF having more trees, shrubs and climbers. Savanna was characterised by forbs and graminoid herbs. Growth habit spectra revealed a clear dominance of herbaceous and suffruticose plants in savanna, and of woody elements and epiphytes in TDF. Regarding Raunkiaer's life forms, savanna had relatively more hemicryptophytes, and TDF more phanerophytes. Classification analyses showed that savanna and TDF forest samples kept their identities, regardless of taxonomic level (species, genera and families) at which the analyses were performed. Main conclusions The TDF and savanna of Nizanda represent two floristic systems with a large degree of differentiation at all taxonomic levels and patterns of morphological attributes. This suggest that the two floristic sets have evolved independently for extended periods of time, despite their close proximity. One important implication of this floristic differentiation is the large joint contribution made by these communities to the regional flora.