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

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

Evaluation of the heat pulse velocity technique for measurement of sap flow in rainforest and eucalypt forest species of south-eastern Australia

Sap flow in the stems of two cut saplings each of Eucalyptus maculata (a canopy eucalypt forest tree), Doryphora sassafras and Ceratopetalum apetalum (both canopy rainforest trees of south-eastern coastal Australia) was measured by the heat pulse velocity technique and compared with water uptake from a potometer. Scanning electron micrographs of wounding caused by implantation of temperature sensor and heater probes into the sapwood showed that wounding was similar in rainforest and eucalypt species and was elliptical in shape. A circular wound has been implicitly assumed in previous studies. Accurate measurements of sapling water use were obtained using the smaller transverse wound dimension rather than the larger longitudinal dimension because maximum disruption of sap flow through the xylem vessels occurred in the transverse plane. Accurate measurements of sap flux were obtained above a minimum threshold sap velocity. These velocities were 15·7,10·9 and 9·4 cm h^?1 for E. maculata, C. apetalum and D. sassafras, respectively. Below the threshold sap velocity, however, sap flow could not be accurately calculated from measurements of heat pulse velocity. The minimum threshold sap velocity appeared to be determined by probe construction and xylem anatomy. Despite the elliptical wounding and inaccurate measurement of sap flow below the threshold sap velocity, total sap flow over the experimental period for two saplings of each species was within 7% of water use measured by the potometer.     

长白山北坡森林群落结构组成及其海拔变化

对长白山北坡海拔700～1900m的13个森林群落进行了群落组成、结构随海拔上升变化情况的研究。结果表明,随着海拔的升高,群落的建群种组成、层次结构等,都表现出由复杂多样逐渐向简单单一过渡的变化趋势,从700m至1700m,针叶树所占的比例随海拔的升高呈明显的增加趋势,阔叶树比例随之减少,到林线时针阔叶树种均急剧减少,沿海拔梯度存在明显的树种更替现象。从群落的平均胸径及胸高断面积和可知。最大值出现于900～1100m及1600m,这2个海拔段正是不同森林群落类型间的过渡区。除海拔1800m的岳桦林之外,其它群落径级结构都呈倒J形分布,即各径级树木中,更新幼苗、幼树在数量上占绝对优势,随着胸径的增大,立木株数逐渐减少,这种径级结构表明,长白山北坡各海拔群落更新良好,群落处于稳定的发展状态。     

Climatic change and fire potential in South-Central British Columbia, Canada

The incidence and severity of forest fires are linked to the interaction between climate, fuel and topography. Increased warming and drying in the future is expected to have a significant impact on the risk of forest fire occurrence. An increase in fire risk is linked to the synchronous relationship between climate and fuel moisture conditions. A warmer, drier climate will lead to drier forest fuels that will in turn increase the chance of successful fire ignition and propagation. This interaction will increase the severity of fire weather, which, in turn, will increase the risk of extreme fire behaviour. A warmer climate will also extend fire season length, which will increase the likelihood of fires occurring over a greater proportion of the year. In this study of the North Okanagan area of British Columbia, Canada, the impacts of climate change of fire potential were evaluated using the Canadian Forest Fire Danger Rating System and multiple climate scenario analysis. Utilizing this approach, a 30% increase in fire season length was modelled to occur by 2070. In addition, statistically significant increases in fire severity and fire behaviour were also modelled. Fire weather severity was predicted to increase by 95% during the summer months by 2070 while fire behaviour was predicted to shift from surface fire‐intermittent crown fire regimes to a predominantly intermittent‐full crown fire regime by 2070 onwards. An increase in fire season length, fire weather severity and fire behaviour will increase the costs of fire suppression and the risk of property and resource loss while limiting human‐use within vulnerable forest landscapes. An increase in fire weather severity and fire behaviour over a greater proportion of the season will increase the risks faced by ecosystems and biodiversity to climatic change and increase the costs and difficulty of achieving sustainable forest management.     

Arthropod community composition along snowmelt gradients in snowbeds in the Snowy Mountains of south‐eastern Australia

Environmental gradients drive variation in community composition across a range of spatial scales. In alpine regions, areas of long‐lasting snow (‘snow patches’) create snowmelt gradients that drive considerable change in vegetation structure and composition over small spatial scales. This study examined whether there is parallel variation in arthropod communities using snowmelt gradients in the Australian Alps. Mites (Acarina) were the most common arthropods in snow patches, followed by springtails while, among the insects, the orders Hymenoptera (primarily Formicidae), Diptera, Coleoptera (primarily Carabidae) and Hemiptera (primarily Cicadellidae) dominated. Along the snowmelt gradient, arthropod assemblages changed from having equal proportions of predators and herbivores in early‐melting zones to being predator‐dominated in late‐melting zones, particularly early in the growing season. This followed a transition in vegetation cover and composition and was driven by higher numbers of predacious carabid beetles in later‐melting zones. Overall, however, our results suggest that snowbed arthropod communities in the Australian alpine zone are more sensitive to short‐term effects, such as time since snowmelt, than to differences in vegetation structure and composition or long‐term patterns of snowmelt. Continued advancement of snowmelt timing due to warmer spring temperatures is therefore likely to have more impact on the seasonality of snowbed arthropod communities than on the overall community composition.     

Changes in soil moisture drive soil methane uptake along a fire regeneration chronosequence in a eucalypt forest landscape

Disturbance associated with severe wildfires (WF) and WF simulating harvest operations can potentially alter soil methane (CH₄) oxidation in well‐aerated forest soils due to the effect on soil properties linked to diffusivity, methanotrophic activity or changes in methanotrophic bacterial community structure. However, changes in soil CH₄ flux related to such disturbances are still rarely studied even though WF frequency is predicted to increase as a consequence of global climate change. We measured in‐situ soil–atmosphere CH₄ exchange along a wet sclerophyll eucalypt forest regeneration chronosequence in Tasmania, Australia, where the time since the last severe fire or harvesting disturbance ranged from 9 to >200 years. On all sampling occasions, mean CH₄ uptake increased from most recently disturbed sites (9 year) to sites at stand ‘maturity’ (44 and 76 years). In stands >76 years since disturbance, we observed a decrease in soil CH₄ uptake. A similar age dependency of potential CH₄ oxidation for three soil layers (0.0–0.05, 0.05–0.10, 0.10–0.15 m) could be observed on incubated soils under controlled laboratory conditions. The differences in soil CH₄ uptake between forest stands of different age were predominantly driven by differences in soil moisture status, which affected the diffusion of atmospheric CH₄ into the soil. The observed soil moisture pattern was likely driven by changes in interception or evapotranspiration with forest age, which have been well described for similar eucalypt forest systems in south‐eastern Australia. Our results imply that there is a large amount of variability in CH₄ uptake at a landscape scale that can be attributed to stand age and soil moisture differences. An increase in severe WF frequency in response to climate change could potentially increase overall forest soil CH₄ sinks.     

Root trait variation along water gradients in the Cape Floristic Region

Questions

Roots are responsible for essential plant functions including water uptake. However, the extent to which root traits (size and structure) determine plant presence in water-limited environments is still poorly understood. Here we investigated how root traits vary across water availability gradients within a dry South African biome.

Location

South Africa.

Methods

We measured six below-ground (root) and eight above-ground (leaf + stem) traits of 124 individuals of nine dominant woody shrub species from wetter and drier sites (600–700 vs 250–300 mm annual precipitation) in the Fynbos biome of the Cape Floristic Region. Within sites, we sampled from recently burnt and unburnt/more mature vegetation and at three locations along topographical gradients.

Results

Drier regions showed greater maximum rooting depth, length, root dry matter content and root to shoot ratio. These trait patterns were consistent at an intraspecific level, along locally drier topographical locations and in post-fire environments. Roots accounted for significant whole-plant trait variation. Additionally, in drier conditions, we found increased root allocation deviating from expected global allometric relationships. Our study also demonstrates that the combination of fire and drought in the driest locations results in poor above-ground vegetation recovery in terms of plant size, cover and individual counts with only resprouters persisting.

Conclusions

Our research suggests that root investment in Fynbos shrubs will likely be key for coping with a drier and warmer future and should be a focus of more research for dryland biomes.     

Climate change,fuel and fire behaviour in a eucalypt forest

A suite of models was used to examine the links between climate, fuels and fire behaviour in dry eucalypt forests in south‐eastern Australia. Predictions from a downscaled climate model were used to drive models of fuel amount, the moisture content of fuels and two models of forest fire behaviour at a location in western Sydney in New South Wales, Australia. We found that a warming and drying climate produced lower fine fuel amounts, but greater availability of this fuel to burn due to lower moisture contents. Changing fuel load had only a small effect on fuel moisture. A warmer, drier climate increased rate of spread, an important measure of fire behaviour. Reduced fuel loads ameliorated climate‐induced changes in fire behaviour for one model. Sensitivity analysis of the other fire model showed that changes in fuel amount induced changes in fire behaviour of a similar magnitude to that caused directly by sensitivity to climate. Projection of changes in fire risk requires modelling of changes in vegetation as well as changes in climate. Better understanding of climate change effects on vegetation structure is required.     

Variation in spatial and temporal gradients in zooplankton spring development: the effect of climatic factors

1. We examined the temporal and spatial heterogeneity of zooplankton in lake surface waters during the spring of 3 years in Lake Washington, U.S.A., a large lake with a high production of sockeye salmon fry. 2. We show large within‐season and among‐year variation in the horizontal distribution of temperature, chlorophyll a concentration, and zooplankton in the lake. The main pattern, a delay in zooplankton population increase from the north‐ to the south‐end of the lake, recurred in each year and was persistent within each spring. 3. The delay is primarily caused by the development of a temperature gradient during spring warming, as cold mountain water enters the south end of the lake, while warm water enters the north end via a river draining a nearby lake. Climate factors, such as air temperature and precipitation during winter and spring, appear to influence the extent of the delay of zooplankton increase. 4. If the climate continues to warm, the temporal disconnection in zooplankton development between lake areas immediately influenced by cold river inflow and areas that are influenced by spring warming may increase in magnitude. Thus, the different areas of the lake may not contribute equally to fish production.     

The effect of climatic gradients, topographic variation and species traits on the beta diversity of rain forest trees

Aim  We assessed the rates of turnover of tree species with distance (beta diversity) in wet forests of the Western Ghats (WG) complex of India to see whether climate, topographic variation or species traits influence beta diversity.
Location  The Western Ghats is a chain of mountains about 1600 km in length, running parallel to the western coast of the Indian Peninsula from above 8° N to almost 21° N latitude.
Methods  We used data from 60 small plot inventories concentrated in three regions: the southernmost part of the Western Ghats (SWG) (8°24' to 9°37' N), the Nilgiri Hills (11°12' to 11°14' N), and the central Western Ghats (CWG) (12°32' to 14°51' N). We used Sorensen's index (SI) to estimate the similarity in species composition between two plots and regressed SI against the logarithm of the distance between plots to assess beta diversity. A bootstrapping procedure provided confidence intervals for regression coefficients. To test for the effects of climate, we regressed seasonality differences between plots against SI for low-elevation (< 800 m) plots along the north–south axis, and all plots in the SWG. We assessed the impact of the rainfall gradient in the Kogar region.
Results  Among all three regions, beta diversity was highest along the latitudinal axis, and along the rainfall gradient in the Kogar region. Differences in seasonality between sites were strongly related to beta diversity along the north–south seasonality gradient and within the SWG. Within the three regions, beta diversity was highest in the region with the strongest rainfall gradient and lowest for the topographically heterogeneous SWG. Beta diversity did not differ between forest strata and dispersal modes.
Main conclusions  We conclude that climate, particularly seasonality, is probably the primary driver of beta diversity among rain forest trees of the Western Ghats complex.     

Floristic variation in ecotonal areas: Patterns,determinants and biogeographic origins of subtropical forests in South America

We present the first quantification of forest community composition and its relationship with environmental factors in South American subtropical Atlantic Forests. In this region, rain, seasonally dry and mixed forests form an ecotonal zone near the parallel of latitude 30°S. To investigate how well current knowledge on climatic effects and biogeographic distribution apply to subtropical ecotones, we tested the following expectations: (i) there is a floristic longitudinal gradient correlated to altitudinal and climatic gradients; (ii) climatic variables are more important than soil factors in shaping floristic composition; and (iii) there are three floristic regions in the southernmost limit of the Atlantic Forest biome that are expected to be distinct in composition, structure and biogeographical origin. We examined floristic composition and its relationship with environmental factors across 52 1‐ha permanent study areas in subtropical Brazil, containing in total 269 tree species ≥ 9.5 dbh (diameter at breast height). Climatic data, related to rainfall seasonality and temperature, as well as soil properties, were compiled from published sources or global data banks. Expectations one and two were confirmed, but expectation three was only partially met. Hierarchical cluster analysis divided the southernmost Atlantic Forests into four major groups (Rain, Seasonally Dry, Western Mixed and Eastern Mixed Forests). Overall, the tested environmental variables differed significantly among the four regions. Using indicator species analysis, we distinguished 46 indicator species, which had significant environmental preferences for one floristic region. These species can be used as indicators of environmental conditions or to determine to which floristic region a certain forest belongs. Biogeographic distributions differed between floristic groups, supporting the interpretation that Eastern Mixed Forests are relict forests of a temperate forest of Andean origin that occurred during colder palaeoclimates. Western Mixed Forests represent the main floristic ecotone between Seasonally dry and Eastern Mixed Forests.     

The effect of fire and permafrost interactions on soil carbon accumulation in an upland black spruce ecosystem of interior Alaska: implications for post‐thaw carbon loss

High‐latitude regions store large amounts of organic carbon (OC) in active‐layer soils and permafrost, accounting for nearly half of the global belowground OC pool. In the boreal region, recent warming has promoted changes in the fire regime, which may exacerbate rates of permafrost thaw and alter soil OC dynamics in both organic and mineral soil. We examined how interactions between fire and permafrost govern rates of soil OC accumulation in organic horizons, mineral soil of the active layer, and near‐surface permafrost in a black spruce ecosystem of interior Alaska. To estimate OC accumulation rates, we used chronosequence, radiocarbon, and modeling approaches. We also developed a simple model to track long‐term changes in soil OC stocks over past fire cycles and to evaluate the response of OC stocks to future changes in the fire regime. Our chronosequence and radiocarbon data indicate that OC turnover varies with soil depth, with fastest turnover occurring in shallow organic horizons (～60 years) and slowest turnover in near‐surface permafrost (>3000 years). Modeling analysis indicates that OC accumulation in organic horizons was strongly governed by carbon losses via combustion and burial of charred remains in deep organic horizons. OC accumulation in mineral soil was influenced by active layer depth, which determined the proportion of mineral OC in a thawed or frozen state and thus, determined loss rates via decomposition. Our model results suggest that future changes in fire regime will result in substantial reductions in OC stocks, largely from the deep organic horizon. Additional OC losses will result from fire‐induced thawing of near‐surface permafrost. From these findings, we conclude that the vulnerability of deep OC stocks to future warming is closely linked to the sensitivity of permafrost to wildfire disturbance.     

A biogeographic model of fire regimes in Australia: current and future implications

Aim Patterns of fire regimes across Australia exhibit biogeographic variation in response to four processes. Variations in area burned and fire frequency result from differences in the rates of ‘switching’ of biomass growth, availability to burn, fire weather and ignition. Therefore differing processes limit fire (i.e. the lowest rate of switching) in differing ecosystems. Current and future trends in fire frequency were explored on this basis. Location Case studies of forests (cool temperate to tropical) and woodlands (temperate to arid) were examined. These represent a broad range of Australian biomes and current fire regimes. Methods Information on the four processes was applied to each case study and the potential minimum length of interfire interval was predicted and compared to current trends. The potential effects of global change on the processes were then assessed and future trends in fire regimes were predicted. Results Variations in fire regimes are primarily related to fluctuations in available moisture and dominance by either woody or herbaceous plant cover. Fire in woodland communities (dry climates) is limited by growth of herbaceous fuels (biomass), whereas in forests (wet climates) limitation is by fuel moisture (availability to burn) and fire weather. Increasing dryness in woodland communities will decrease potential fire frequency, while the opposite applies in forests. In the tropics, both forms of limitation are weak due to the annual wet/dry climate. Future change may therefore be constrained. Main conclusions Increasing dryness may diminish fire activity over much of Australia (dominance of dry woodlands), though increases may occur in temperate forests. Elevated CO₂ effects may confound or reinforce these trends. The prognosis for the future fire regime in Australia is therefore uncertain.     

Determinants of growth of the flammable grass,Triodia scariosa: Consequences for fuel dynamics under climate change in the Mediterranean region of South Eastern Australia

Environmental conditions may influence the presence and strength of competitive interactions between different life forms, thereby shaping community composition and structure, and corresponding fuel dynamics. Woodland and shrubland communities of the Mediterranean climate region of South Eastern Australia contain a varied mixture of herbaceous and woody plants. The ratio of herbaceous to woody plants changes along gradients of temperature, moisture and soil fertility. This study aimed to experimentally examine the relative importance of, and interactions between environmental controls (moisture and soil fertility) on the balance of dominant herbaceous (Triodia scariosa) and woody plants (e.g. Acacia ligulata and Leptospermum coriaceum) and their ultimate effects on fuel and fire regimes. The results suggest that environmental determinants of the growth of T. scariosa are likely to be more important than interactions with shrubs in controlling the distribution of T. scariosa. The growth of T. scariosa was consistently higher under hot temperatures and on the less fertile yellow sands, which dominate the south of the region. The results suggest that there is strong potential for the distribution and abundance of T. scariosa to be altered in the future with changes in temperature associated with climate change. The distribution of soil types across the Mediterranean climate region of South Eastern Australia may be predisposed to favour the southerly expansion of T. scariosa‐dominated communities in the future under a warmer climate.     

Evidence of neutral and adaptive genetic divergence between European trout populations sampled along altitudinal gradients

Species with a wide geographical distribution are often composed of distinct subgroups which may be adapted to their local environment. European trout (Salmo trutta species complex) provide an example of such a complex consisting of several genetically and ecologically distinct forms. However, trout populations are strongly influenced by human activities, and it is unclear to what extent neutral and adaptive genetic differences have persisted. We sampled 30 Swiss trout populations from heterogeneous environments along replicated altitudinal gradients in three major European drainages. More than 850 individuals were genotyped at 18 microsatellite loci which included loci diagnostic for evolutionary lineages and candidate markers associated with temperature tolerance, reproductive timing and immune defence. We find that the phylogeographic structure of Swiss trout populations has not been completely erased by stocking. Distinct genetic clusters corresponding to the different drainages could be identified, although nonindigenous alleles were clearly present, especially in the two Mediterranean drainages. We also still detected neutral genetic differentiation within rivers which was often associated with the geographical distance between populations. Five loci showed evidence of divergent selection between populations with several drainage-specific patterns. Lineage-diagnostic markers, a marker linked to a quantitative trait locus for upper temperature tolerance in other salmonids and a marker linked to the major histocompatibility class I gene were implicated in local adaptation and some patterns were associated with altitude. In contrast, tentative evidence suggests a signal of balancing selection at a second immune relevant gene (TAP2). Our results confirm the persistence of both neutral and potentially adaptive genetic differences between trout populations in the face of massive human-mediated dispersal.     

Regional scale gradients of climate and nitrogen deposition drive variation in ectomycorrhizal fungal communities associated with native Scots pine

Ectomycorrhizal fungi commonly associate with the roots of forest trees where they enhance nutrient and water uptake, promote seedling establishment and have an important role in forest nutrient cycling. Predicting the response of ectomycorrhizal fungi to environmental change is an important step to maintaining forest productivity in the future. These predictions are currently limited by an incomplete understanding of the relative significance of environmental drivers in determining the community composition of ectomycorrhizal (ECM) fungi at large spatial scales. To identify patterns of community composition in ECM fungi along regional scale gradients of climate and nitrogen deposition in Scotland, fungal communities were analysed from 15 seminatural Scots pine (Pinus sylvestris L.) forests. Fungal taxa were identified by sequencing of the ITS rDNA region using fungal‐specific primers. Nonmetric multidimensional scaling was used to assess the significance of 16 climatic, pollutant and edaphic variables on community composition. Vector fitting showed that there was a strong influence of rainfall and soil moisture on community composition at the species level, and a smaller impact of temperature on the abundance of ectomycorrhizal exploration types. Nitrogen deposition was also found to be important in determining community composition, but only when the forest experiencing the highest deposition (9.8 kg N ha^?1 yr^?1) was included in the analysis. This finding supports previously published critical load estimates for ectomycorrhizal fungi of 5–10 kg N ha^?1 yr^?1. This work demonstrates that both climate and nitrogen deposition can drive gradients of fungal community composition at a regional scale.