Firescape ecology: how topography determines the contrasting distribution of fire and rain forest in the south‐west of the Tasmanian Wilderness World Heritage Area

Aim To test the hypothesis that ‘islands’ of fire‐sensitive rain forest are restricted to topographic fire refugia and investigate the role of topography–fire interactions in fire‐mediated alternative stable state models. Location A vegetation mosaic of moorland, sclerophyll scrub, wet sclerophyll eucalypt forest and rain forest in the rugged, fire‐prone landscapes of south‐west Tasmania, Australia. Methods We used geospatial statistics to: (1) identify the topographic determinants of rain forest distribution on nutrient‐poor substrates, and (2) identify the vegetation and topographic variables that are important in controlling the spatial pattern of a series of very large fires (> 40,000 ha) that were mapped using Landsat Thematic Mapper (TM) satellite imagery. Results Rain forest was more likely to be found in valleys and on steep south‐facing slopes. Fires typically burned within highly flammable treeless moorland and stopped on boundaries with less flammable surrounding vegetation types such as wet sclerophyll forest and rain forest. Controlling for the effect of vegetation, fires were most likely to burn on flats, ridges and steep north‐facing slopes and least likely to burn in valleys and on steep south‐facing slopes. These results suggest an antagonism between fire and rain forest, in which rain forest preferentially occupies parts of the landscape where fire is least likely to burn. Main conclusions The distribution of rain forest on nutrient‐poor substrates was clearly related to parts of the landscape that are protected from fire (i.e. topographic fire refugia). The relative flammability of vegetation types at the landscape scale offers support to the proposed hierarchy of fire frequencies (moorland > scrub > wet sclerophyll > rain forest) that underpins the ecological models proposed for the region. The interaction between fire occurrence and a range of topographic variables suggests that topography plays an important role in mediating the fire–vegetation feedbacks thought to maintain vegetation mosaics in south‐west Tasmania. We suggest that these fire–topography interactions should be included in models of fire‐mediated alternative stable vegetation states in other fire‐prone landscapes.     

Abrupt fire regime change may cause landscape‐wide loss of mature obligate seeder forests

Obligate seeder trees requiring high‐severity fires to regenerate may be vulnerable to population collapse if fire frequency increases abruptly. We tested this proposition using a long‐lived obligate seeding forest tree, alpine ash (Eucalyptus delegatensis), in the Australian Alps. Since 2002, 85% of the Alps bioregion has been burnt by several very large fires, tracking the regional trend of more frequent extreme fire weather. High‐severity fires removed 25% of aboveground tree biomass, and switched fuel arrays from low loads of herbaceous and litter fuels to high loads of flammable shrubs and juvenile trees, priming regenerating stands for subsequent fires. Single high‐severity fires caused adult mortality and triggered mass regeneration, but a second fire in quick succession killed 97% of the regenerating alpine ash. Our results indicate that without interventions to reduce fire severity, interactions between flammability of regenerating stands and increased extreme fire weather will eliminate much of the remaining mature alpine ash forest.     

Examining forest resilience to changing fire frequency in a fire‐prone region of boreal forest

Future changes in climate are widely anticipated to increase fire frequency, particularly in boreal forests where extreme warming is expected to occur. Feedbacks between vegetation and fire may modify the direct effects of warming on fire activity and shape ecological responses to changing fire frequency. We investigate these interactions using extensive field data from the Boreal Shield of Saskatchewan, Canada, a region where >40% of the forest has burned in the past 30 years. We use geospatial and field data to assess the resistance and resilience of eight common vegetation states to frequent fire by quantifying the occurrence of short‐interval fires and their effect on recovery to a similar vegetation state. These empirical relationships are combined with data from published literature to parameterize a spatially explicit, state‐and‐transition simulation model of fire and forest succession. We use this model to ask if and how: (a) feedbacks between vegetation and wildfire may modify fire activity on the landscape, and (b) more frequent fire may affect landscape forest composition and age structure. Both field and GIS data suggest the probability of fire is low in the initial decades after fire, supporting the hypothesis that fuel accumulation may exert a negative feedback on fire frequency. Field observations of pre‐ and postfire composition indicate that switches in forest state are more likely in conifer stands that burn at a young age, supporting the hypothesis that resilience is lower in immature stands. Stands dominated by deciduous trees or jack pine were generally resilient to fire, while mixed conifer and well‐drained spruce forests were less resilient. However, simulation modeling suggests increased fire activity may result in large changes in forest age structure and composition, despite the feedbacks between vegetation–fire likely to occur with increased fire activity.     

Disturbance history and stand dynamics in tall open forest and riparian rainforest in the Central Highlands of Victoria

Abstract Spatial heterogeneity in the intensity of past disturbances has directly influenced the structure and composition of present‐day forests around the world. In south‐eastern Australia infrequent, high‐intensity wildfires are a major part of the historical disturbance regime. While these fires are often assumed to produce even‐aged stands, spatial heterogeneity in fire intensity due to highly variable topography may lead to more complex forest age structures. Our study describes the influence of disturbance on the age structure and dynamics of a mosaic of tall, open eucalypt forest, cool temperate rainforest and mixed species forest surrounding Bellel Creek in the Central Highlands of Victoria using dendrochronological techniques. We were particularly interested in the impacts of the 1939 Black Friday fire and its effects on forest age structure and subsequent stand development patterns. Within our study site tall open forest displayed two distinct age cohorts: (i) trees that established immediately after the 1939 fire and accounted for the majority of individuals in the forest, and (ii) scattered groups of older trees estimated to be approximately 200–250 years old. Cool temperate rainforest and mixed forest were also dominated by the post‐1939 fire age cohort. However, a greater proportion of trees in these forest types survived the 1939 fire relative to the tall open forest. The impact of the 1939 fire on the growth of surviving trees was highly variable but generally short‐lived. In most cases growth decreased after the 1939 fire, but generally returned to prefire levels within 1–3 years. Non‐fire disturbances were limited to small‐scale branch‐ and tree‐fall events, although the extreme snowstorm of 1977 appears to have caused extensive damage to rainforest communities. Our study demonstrates the opportunities for dendroecological studies to reconstruct historical dynamics and disturbance patterns in Australian forests and provides important insights into variation in landscape‐scale fire impacts and their effect on subsequent forest development patterns.     

Using a rainforest-flame forest mosaic to test the hypothesis that leaf and litter fuel flammability is under natural selection

We used a mosaic of infrequently burnt temperate rainforest and adjacent, frequently burnt eucalypt forests in temperate eastern Australia to test whether: (1) there were differences in flammability of fresh and dried foliage amongst congeners from contrasting habitats, (2) habitat flammability was related to regeneration strategy, (3) litter fuels were more flammable in frequently burnt forests, (4) the severity of a recent fire influenced the flammability of litter (as this would suggest fire feedbacks), and (5) microclimate contributed to differences in fire hazard amongst habitats. Leaf-level comparisons were made among 11 congeneric pairs from rainforest and eucalypt forests. Leaf-level ignitability, combustibility and sustainability were not consistently higher for taxa from frequently burnt eucalypt forests, nor were they higher for species with fire-driven recruitment. The bulk density of litter-bed fuels strongly influenced flammability, but eucalypt forest litter was not less dense than rainforest litter. Ignitability, combustibility and flame sustainability of community surface fuels (litter) were compared using fuel arrays with standardized fuel mass and moisture content. Forests previously burned at high fire severity did not have consistently higher litter flammability than those burned at lower severity or long unburned. Thus, contrary to the Mutch hypothesis, there was no evidence of higher flammability of litter fuels or leaves from frequently burnt eucalypt forests compared with infrequently burnt rainforests. We suggest the manifest pyrogenicity of eucalypt forests is not due to natural selection for more flammable foliage, but better explained by differences in crown openness and associated microclimatic differences.     

Fire as a Recurrent Event in Tropical Forests of the Eastern Amazon: Effects on Forest Structure,Biomass, and Species Composition1

The effects of fire on forest structure and composition were studied in a severely fire-impacted landscape in the eastern Amazon. Extensive sampling of area forests was used to compare structure and compositional differences between burned and unburned forest stands. Burned forests were extremely heterogeneous, with substantial variation in forest structure and fire damage recorded over distances of <50 m. Unburned forest patches occurred within burned areas, but accounted for only six percent of the sample area. Canopy cover, living biomass, and living adult stem densities decreased with increasing fire inrensiry / frequency, and were as low as 10–30 percent of unburned forest values. Even light burns removed >70 percent of the sapling and vine populations. Pioneer abundance increased dramatically with burn intensity, with pioneers dominating the understory in severely damaged areas. Species richness was inversely related to burn severity, but no clear pattern of species selection was observed. Fire appears to be a cyclical event in the study region: <30 percent of the burned forest sample had been subjected to only one burn. Based on estimated solar radiation intensities, burning substantially increases fire susceptibility of forests. At least 50 percent of the total area of all burned forests is predicted to become flammable within 16 rainless days, as opposed to only 4 percent of the unburned forest. In heavily burned forest subjected to recurrent fires, 95 percent of the area is predicted to become flammable in <9 rain-free days. As a recurrent disturbance phenomenon, fire shows unparalleled potential to impoverish and alter the forests of the eastern Amazon.     

Coarse woody debris in Australian forest ecosystems: A review

Abstract Coarse woody debris (CWD) is the standing and fallen dead wood in a forest and serves an important role in ecosystem functioning. There have been several studies that include estimates of CWD in Australian forests but little synthesis of these results. This paper presents findings from a literature review of CWD and fine litter quantities. Estimates of forest‐floor CWD, snags and litter from the literature are presented for woodland, rainforest, open forest and tall open forest, pine plantation and native hardwood plantation. Mean mass of forest floor CWD in Australian native forests ranged from 19 t ha^?1 in woodland to 134 t ha^?1 in tall open forest. These values were generally within the range of those observed for similar ecosystems in other parts of the world. Quantities in tall open forests were found to be considerably higher than those observed for hardwood forests in North America, and more similar to the amounts reported for coniferous forests with large sized trees on the west coast of the USA and Canada. Mean proportion of total above‐ground biomass as forest floor CWD was approximately 18% in open forests, 16% in tall open forests, 13% in rainforests, and 4% in eucalypt plantations. CWD can be high in exotic pine plantations when there are considerable quantities of residue from previous native forest stands. Mean snag biomass in Australian forests was generally lower than the US mean for snags in conifer forests and higher than hardwood forest. These results are of value for studies of carbon and nutrient stocks and dynamics, habitat values and fire hazards.     

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.     

How do slow‐growing,fire‐sensitive conifers survive in flammable eucalypt woodlands?

Question: To what extent do low flammability fuel traits enhance the survival and persistence of fire‐sensitive (slowing‐growing, non‐serotinous, non‐resprouting) dominant trees in highly flammable landscapes, under varying fire‐weather conditions? Location: Mixed forests co‐dominated by flammable Eucalyptus species and fire‐sensitive Callitris glaucophylla in Pilliga State Forest, southeast Australia. Methods: The influence of vegetation composition (relative abundance of Callitris and flammable Eucalyptus) on fire intensity and survival of fire‐sensitive Callitris was assessed across gradients of Callitris abundance in mixed Eucalyptus–Callitris forests that burned under low‐moderate and extreme fire‐weather conditions. Results: In areas that burned under low‐moderate fire‐weather conditions, as Callitris abundance increased, fire intensity declined and Callitris survival increased (46%). By comparison, in extreme fire‐weather conditions, lower fire intensity at higher levels of Callitris abundance, was not sufficient to increase Callitris survival (4%). Callitris survival was also positively related to trunk diameter. Ground fuel type, but not biomass, varied with vegetation composition. Conclusions: These results demonstrate that flammable feedbacks, mediated by low flammability fuel traits of dominant trees, can provide an important mechanism for enhancing the survival and persistence of slow‐growing, non‐serotinous, non‐resprouting, fire‐killed trees in highly flammable landscapes. By modifying vegetation and fuel structure, patches of fire‐sensitive Callitris reduce fire intensity, and thereby reduce Callitris mortality, enhancing population persistence. However, this feedback loop is insufficient to ensure Callitris survival under extreme fire‐weather conditions, when fire intensity is greater. After burning, stands remain vulnerable to future fires, until trees grow large enough to modify fuel levels and reduce stand flammability.     

Altered ignition catchments threaten a hyperdiverse fire‐dependent ecosystem

Human activities affect fire in many ways, often unintentionally or with considerable time‐lags before they manifest themselves. Anticipating these changes is critical, so that insidious impacts on ecosystems, their biodiversity and associated goods and services can be avoided, mitigated or managed. Here we explore the impact of anthropogenic land cover change on fire and biodiversity in adjacent ecosystems on the hyperdiverse Cape Peninsula, South Africa. We develop a conceptual framework based on the notion of an ignition catchment, or the spatial extent and temporal range where an ignition is likely to result in a site burning. We apply this concept using fire models to estimate spatial changes in burn probability between historical and current land cover. This change layer was used to predict the observed record of fires and forest encroachment into fire‐dependent Fynbos ecosystems in Table Mountain National Park. Urban expansion has created anthropogenic fire shadows that are modifying fire return intervals, facilitating a state shift to low‐diversity, non‐flammable forest at the expense of hyperdiverse, flammable Fynbos ecosystems. Despite occurring in a conservation area, these ecosystems are undergoing a hidden collapse and desperately require management intervention. Anthropogenic fire shadows can be caused by many human activities and are likely to be a universal phenomenon, not only contributing to the observed global decline in fire activity but also causing extreme fires in ecosystems where there is no shift to a less flammable state and flammable fuels accumulate. The ignition catchment framework is highly flexible and allows detection or prediction of changes in the fire regime, the threat this poses for ecosystems or fire risk and areas where management interventions and/or monitoring are required. Identifying anthropogenic impacts on ignition catchments is key for both understanding global impacts of humans on fire and guiding management of human‐altered landscapes for desirable outcomes.     

The role of succession in the evolution of flammability

Fire-prone ecosystems contain plants that are both fire-adapted and flammable. It has been hypothesized that these plants were under selection to become more flammable, but it is unclear whether this could be adaptive for an individual plant. We propose arrested succession as a robust mechanism that supports the evolution of flammability in surface fire ecosystems without the need to invoke group selection or additional fitness benefits. We used the natural history of lodgepole pine (Pinus ponderosa) forests, longleaf pine (Pinus palustris) forests, and tall grass prairies to create a general mathematical model of surface fire ecosystems and solved for the evolutionarily stable strategy (ESS) level of flammability. In our model, fires always kill understory plants and only sometimes kill overstory plants. Thus, more flammable plants suffer increased mortality due to fires, but also more frequently arrest succession by clearing their understory of late successional competitors. Increased flammability was selected for when the probability of an overstory plant dying from an individual fire was below a maximum threshold and the rate of succession relative to fires was above a minimum threshold. Future studies can test our model predictions and help resolve whether or not plants have been selected to be more flammable.     

Explaining fire‐driven landscape transformation during the Initial Burning Period of New Zealand's prehistory

At the time of Māori settlement, ca. 750 years ago, New Zealand's ecosystems experienced catastrophic change, including the introduction of fire to ignition‐limited ecosystems and the resulting widespread loss of forest. While high‐resolution sediment‐charcoal analyses suggest this forest loss was rapid, Māori populations were small and transient during the Initial Burning Period and there is evidence for widespread fire activity in places where there is little archaeological evidence of human presence. These observations beg the question ‘how did small populations manage to transform large areas so rapidly?’ Using a simulation model, we demonstrate how the relationship between time since fire and flammability in New Zealand's forests drives positive feedbacks that allow for rapid and extensive deforestation. Under ignition scenarios mirroring prehuman conditions, the model did not produce significant deforestation – thus, it is extremely unlikely that deforestation could have occurred without human‐initiated burning. Scenarios where ignition was spatio‐temporally random also failed to result in deforestation. Rapid and widespread forest loss occurred in scenarios incorporating spatio‐temporally savvy selection of ignition locations. Targeting ignitions in flammable vegetation was more important than targeting ignitions in years with favourable climatic conditions. However, targeting in space and time concurrently, such that flammable vegetation was ignited during favourable climatic years was the most efficient strategy of those simulated. Following the Initial Burning Period decadal ignitions would have been sufficient to maintain a deforested shrubland/grassland landscape. New Zealand's Initial Burning Period is one of many that occurred across eastern Polynesia following human settlement, and these events have left long‐term legacy effects that remain evident in contemporary landscapes. Improving understanding of how humans shaped environments in New Zealand in the past has implications for eastern Polynesia as a whole.     

Impacts of climate variability on tree demography in second growth tropical forests: the importance of regional context for predicting successional trajectories

Naturally regenerating and restored second growth forests account for over 70% of tropical forest cover and provide key ecosystem services. Understanding climate change impacts on successional trajectories of these ecosystems is critical for developing effective large‐scale forest landscape restoration (FLR) programs. Differences in environmental conditions, species composition, dynamics, and landscape context from old growth forests may exacerbate climate impacts on second growth stands. We compile data from 112 studies on the effects of natural climate variability, including warming, droughts, fires, and cyclonic storms, on demography and dynamics of second growth forest trees and identify variation in forest responses across biomes, regions, and landscapes. Across studies, drought decreases tree growth, survival, and recruitment, particularly during early succession, but the effects of temperature remain unexplored. Shifts in the frequency and severity of disturbance alter successional trajectories and increase the extent of second growth forests. Vulnerability to climate extremes is generally inversely related to long‐term exposure, which varies with historical climate and biogeography. The majority of studies, however, have been conducted in the Neotropics hindering generalization. Effects of fire and cyclonic storms often lead to positive feedbacks, increasing vulnerability to climate extremes and subsequent disturbance. Fragmentation increases forests’ vulnerability to fires, wind, and drought, while land use and other human activities influence the frequency and intensity of fire, potentially retarding succession. Comparative studies of climate effects on tropical forest succession across biogeographic regions are required to forecast the response of tropical forest landscapes to future climates and to implement effective FLR policies and programs in these landscapes.     

Fire Severity and Long-term Ecosystem Biomass Dynamics in Coniferous Boreal Forests of Eastern Canada

The objective of this study was to characterize the effects of soil burn severity and initial tree composition on long-term forest floor dynamics and ecosystem biomass partitioning within the Picea mariana [Mill.] BSP-feathermoss bioclimatic domain of northwestern Quebec. Changes in forest floor organic matter and ecosystem biomass partitioning were evaluated along a 2,355-year chronosequence of extant stands. Dendroecological and paleoecological methods were used to determine the time since the last fire, the soil burn severity of the last fire (high vs. low severity), and the post-fire tree composition of each stand (P. mariana vs. Pinus banksiana Lamb). In this paper, soil burn severity refers to the thickness of the organic matter layer accumulated above the mineral soil that was not burned by the last fire. In stands originating from high severity fires, the post-fire dominance by Pinus banksiana or P. mariana had little effect on the change in forest floor thickness and tree biomass. In contrast, stands established after low severity fires accumulated during the first century after fire 73% thicker forest floors and produced 50% less tree biomass than stands established after high severity fires. Standing tree biomass increased until approximately 100 years after high severity fires, and then decreased at a logarithmic rate in the millennial absence of fire. Forest floor thickness also showed a rapid initial accumulation rate, and continued to increase in the millennial absence of fire at a much slower rate. However, because forest floor density increased through time, the overall rate of increase in forest floor biomass (58 g m⁻² y⁻¹) remained constant for numerous centuries after fire (700 years). Although young stands (< 200 years) have more than 60% of ecosystem biomass locked-up in living biomass, older stands (> 200 years) sequester the majority (> 80%) of it in their forest floor. The results from this study illustrate that, under similar edaphic conditions, a single gradient related to time since disturbance is insufficient to account for the full spectrum of ecosystem biomass dynamics occurring in eastern boreal forests and highlights the importance of considering soil burn severity. Although fire severity induces diverging ecosystem biomass dynamics in the short term, the extended absence of fire brings about a convergence in terms of ecosystem biomass accumulation and partitioning.     

Fire weather risk differs across rain forest—savanna boundaries in the humid tropics of north‐eastern Australia

Alternative stable state theory has been applied to understanding the control by landscape fire activity of pyrophobic tropical rain forest and pyrophytic eucalypt savanna boundaries, which are often separated by tall eucalypt forests. We evaluate the microclimate of three vegetation types across an elevational gradient and their relative fire risk as measured by McArthur's Forest Fire Danger Index (FFDI). Microclimatic data were collected from rain forest, tall eucalypt forest and savanna sites on eight vegetation boundaries throughout the humid tropics in north Queensland over a 3‐year period and were compared with data from a nearby meteorological station. There was a clear annual pattern in daily FFDI with highest values in the austral winter dry season and lowest values in the austral summer wet season. There was a strong association of the meteorological station FFDI values with those from the three vegetation types, albeit they were substantially lower. The rank order of FFDI values among the vegetation types decreased from savanna, tall eucalypt forest, then rain forest, a pattern that was consistent across each transect. Only very rarely would rain forest be flammable, despite being adjacent to highly flammable savannas. These results demonstrate the very strong effect of vegetation type on microclimate and fire risk, compared with the weak effect of elevation, consistent with a fire–vegetation feedback. This study is the first demonstration of how vegetation type influences microclimate and fire risk across a topographically complex tropical forest–savanna gradient.     

Long-term impact of a stand-replacing fire on ecosystem CO2 exchange of a ponderosa pine forest

Ponderosa pine (Pinus ponderosa) forests of the southwestern United States are a mosaic of stands where undisturbed forests are carbon sinks, and stands recovering from wildfires may be sources of carbon to the atmosphere for decades after the fire. However, the relative magnitude of these sinks and sources has never been directly measured in this region, limiting our understanding of the role of fire in regional and US carbon budgets. We used the eddy covariance technique to measure the CO₂ exchange of two forest sites, one burned by fire in 1996, and an unburned forest. The fire was a high‐intensity stand‐replacing burn that killed all trees. Ten years after the fire, the burned site was still a source of CO₂ to the atmosphere [109±6 (SEM) g C m^?2 yr^?1], whereas the unburned site was a sink (?164±23 g C m^?2 yr^?1). The fire reduced total carbon storage and shifted ecosystem carbon allocation from the forest floor and living biomass to necromass. Annual ecosystem respiration was lower at the burned site (480±5 g C m^?2 yr^?1) than at the unburned site (710±54 g C m^?2 yr^?1), but the difference in gross primary production was even larger (372±13 g C m^?2 yr^?1 at the burned site and 858±37 g C m^?2 yr^?1at the unburned site). Water availability controlled carbon flux in the warm season at both sites, and the burned site was a source of carbon in all months, even during the summer, when wet and warm conditions favored respiration more than photosynthesis. Our study shows that carbon losses following stand‐replacing fires in ponderosa pine forests can persist for decades due to slow recovery of the gross primary production. Because fire exclusion is becoming increasingly difficult in dry western forests, a large US forest carbon sink could shift to a decadal‐scale carbon source.     

Coupled effects of wind‐storms and drought on tree mortality across 115 forest stands from the Western Alps and the Jura mountains

Damage due to wind‐storms and droughts is increasing in many temperate forests, yet little is known about the long‐term roles of these key climatic factors in forest dynamics and in the carbon budget. The objective of this study was to estimate individual and coupled effects of droughts and wind‐storms on adult tree mortality across a 31‐year period in 115 managed, mixed coniferous forest stands from the Western Alps and the Jura mountains. For each stand, yearly mortality was inferred from management records, yearly drought from interpolated fields of monthly temperature, precipitation and soil water holding capacity, and wind‐storms from interpolated fields of daily maximum wind speed. We performed a thorough model selection based on a leave‐one‐out cross‐validation of the time series. We compared different critical wind speeds (CWSs) for damage, wind‐storm, and stand variables and statistical models. We found that a model including stand characteristics, drought, and storm strength using a CWS of 25 ms^?1 performed the best across most stands. Using this best model, we found that drought increased damage risk only in the most southerly forests, and its effect is generally maintained for up to 2 years. Storm strength increased damage risk in all forests in a relatively uniform way. In some stands, we found positive interaction between drought and storm strength most likely because drought weakens trees, and they became more prone to stem breakage under wind‐loading. In other stands, we found negative interaction between drought and storm strength, where excessive rain likely leads to soil water saturation making trees more susceptible to overturning in a wind‐storm. Our results stress that temporal data are essential to make valid inferences about ecological impacts of disturbance events, and that making inferences about disturbance agents separately can be of limited validity. Under projected future climatic conditions, the direction and strength of these ecological interactions could also change.     

Stability in the patterns of long‐term development and growth of the Canadian spruce–moss forest

Aim The spruce–moss forest is the main forest ecosystem of the North American boreal forest. We used stand structure and fire data to examine the long‐term development and growth of the spruce–moss ecosystem. We evaluate the stability of the forest with time and the conditions needed for the continuing regeneration, growth and re‐establishment of black spruce (Picea mariana) trees. Location The study area occurs in Québec, Canada, and extends from 70°00′ to 72°00′ W and 47°30′ to 56°00′ N. Methods A spatial inventory of spruce–moss forest stands was performed along 34 transects. Nineteen spruce–moss forests were selected. A 500 m² quadrat at each site was used for radiocarbon and tree‐ring dating of time since last fire (TSLF). Size structure and tree regeneration in each stand were described based on diameter distribution of the dominant and co‐dominant tree species [black spruce and balsam fir (Abies balsamea)]. Results The TSLF of the studied forests ranges from 118 to 4870 cal. yr bp . Forests < 325 cal. yr bp are dominated by trees of the first post‐fire cohort and are not yet at equilibrium, whereas older forests show a reverse‐J diameter distribution typical of mature, old‐growth stands. The younger forests display faster height and radial growth‐rate patterns than the older forests, due to factors associated with long‐term forest development. Each of the stands examined established after severe fires that consumed all the soil organic material. Main conclusions Spruce–moss forests are able to self‐regenerate after fires that consume the organic layer, thus allowing seed regeneration at the soil surface. In the absence of fire the forests can remain in an equilibrium state. Once the forests mature, tree productivity eventually levels off and becomes stable. Further proof of the enduring stability of these forests, in between fire periods, lies in the ages of the stands. Stands with a TSLF of 325–4870 cal. yr bp all exhibited the same stand structure, tree growth rates and species characteristics. In the absence of fire, the spruce–moss forests are able to maintain themselves for thousands of years with no apparent degradation or change in forest type.     

Species mixture effects on flammability across plant phylogeny: the importance of litter particle size and the special role for non‐Pinus Pinaceae

Fire affects and is affected by plants. Vegetation varies in flammability, that is, its general ability to burn, at different levels of ecological organization. To scale from individual plant traits to community flammability states, understanding trait effects on species flammability variation and their interaction is important. Plant traits are the cumulative result of evolution and they show, to differing extents, phylogenetic conservatism. We asked whether phylogenetic distance between species predicts species mixture effects on litterbed flammability. We conducted controlled laboratory burns for 34 phylogenetically wide‐ranging species and 34 random two‐species mixtures from them. Generally, phylogenetic distance did not predict species mixture effects on flammability. Across the plant phylogeny, most species were flammable except those in the non‐Pinus Pinaceae, which shed small needles producing dense, poorly ventilated litterbeds above the packing threshold and therefore nonflammable. Consistently, either positive or negative dominance effects on flammability of certain flammable or those non‐flammable species were found in mixtures involving the non‐Pinus Pinaceae. We demonstrate litter particle size is key to explaining species nonadditivity in fuelbed flammability. The potential of certain species to influence fire disproportionately to their abundance might increase the positive feedback effects of plant flammability on community flammability state if flammable species are favored by fire.     

Humid tropical rain forest has expanded into eucalypt forest and savanna over the last 50 years

Tropical rain forest expansion and savanna woody vegetation thickening appear to be a global trend, but there remains uncertainty about whether there is a common set of global drivers. Using geographic information techniques, we analyzed aerial photography of five areas in the humid tropics of northeastern Queensland, Australia, taken in the 1950s and 2008, to determine if changes in rain forest extent match those reported for the Australian monsoon tropics using similar techniques. Mapping of the 1950s aerial photography showed that of the combined study area (64,430 ha), 63% was classified as eucalypt forests/woodland and 37% as rain forest. Our mapping revealed that although most boundaries remained stable, there was a net increase of 732 ha of the original rain forest area over the study period, and negligible conversion of rain forest to eucalypt forest/woodland. Statistical modeling, controlling for spatial autocorrelation, indicated distance from preexisting rain forest as the strongest determinant of rain forest expansion. Margin extension had a mean rate across the five sites of 0.6 m per decade. Expansion was greater in tall open forest types but also occurred in shorter, more flammable woodland vegetation types. No correlations were detected with other local variables (aspect, elevation, geology, topography, drainage). Using a geographically weighted mean rate of rain forest margin extension across the whole region, we predict that over 25% of tall open forest (a forest type of high conservation significance) would still remain after 2000 years of rain forest expansion. This slow replacement is due to the convoluted nature of the rain forest boundary and the irregular shape of the tall open forest patches. Our analyses point to the increased concentration of atmospheric CO(2) as the most likely global driver of indiscriminate rain forest expansion occurring in northeastern Australia, by increasing tree growth and thereby overriding the effects of fire disturbance.