The relative importance of intrinsic and extrinsic factors in the decline of obligate seeder forests

Forests that regenerate exclusively from seed following high‐severity fire are particularly vulnerable to local extinction if fire frequency leaves insufficient time for regenerating plants to reach sexual maturity. We evaluate the relative importance of extrinsic (such as fire weather and climate cycles) and intrinsic (such as proneness to fire due to stand age and structural development) factors in driving the decline of obligate seeder forests. We illustrate this using obligate seeding alpine ash (Eucalyptus delegatensis) forests in the montane regions of Victoria, Australia, that were burnt by megafires in 2003 (142,256 ha) or 2007 (79,902 ha), including some twice‐burnt areas (11,599 ha). Geospatial analyses showed only a small effect of stand age on the remote sensing estimates of crown defoliation, but a substantial effect of forest fire weather, as measured by forest fire danger index (FFDI). Analysis of meteorological data over the last century showed that 5‐year increases in FFDI precede cycle major fires in the E. delegatensis forests. Such strong extrinsic climate/weather driving of high‐severity fires is consistent with the ‘interval squeeze model’ that postulates the vulnerability of obligate seeder forests to landscape‐scale demographic collapse in response to worsening fire weather under climate change.     

Aboveground carbon sequestration in dry temperate forests varies with climate not fire regime

The storage of carbon in plant tissues and debris has been proposed as a method to offset anthropogenic increases in atmospheric [CO₂]. Temperate forests represent significant above‐ground carbon (AGC) “sinks” because their relatively fast growth and slow decay rates optimise carbon assimilation. Fire is a common disturbance event in temperate forests globally that should strongly influence AGC because: discrete fires consume above‐ground biomass releasing carbon to the atmosphere, and the long‐term application of different fire‐regimes select for specific plant communities that sequester carbon at different rates. We investigated the latter process by quantifying AGC storage at 104 sites in the Sydney Basin Bioregion, Australia, relative to differences in components of the fire regime: frequency, severity and interfire interval. To predict the potential impacts of future climate change on fire/AGC interactions, we stratified our field sites across gradients of mean annual temperature and precipitation and quantified within‐ and between‐factor interactions between the fire and climate variables. In agreement with previous studies, large trees were the primary AGC sink, accounting for ~70% of carbon at sites. Generalised additive models showed that mean annual temperature was the strongest predictor of AGC storage, with a 54% near‐linear decrease predicted across the 6.1°C temperature range experienced at sites. Mean annual precipitation, fire frequency, fire severity and interfire interval were consistently poor predictors of total above‐ground storage, although there were some significant relationships with component stocks. Our results show resilience of AGC to frequent and severe wildfire and suggest temperature mediated decreases in forest carbon storage under future climate change predictions.     

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.     

Time‐since‐fire and climate interact to affect the structural recovery of an Australian semi‐arid plant community

How does time‐since‐fire influence the structural recovery of semi‐arid, eucalypt‐dominated Murray‐Mallee shrublands after fire, and is recovery affected by spatial variation in climate? We assessed the structure and dynamics of a hummock grass, Triodia scariosa N.T. Burb, and mallee eucalypts – two key structural components of mallee shrublands – using a >100 year time‐since‐fire chronosequence. The relative influence of climatic variables, both individually and combined with time‐since‐fire, was modelled to account for spatial variation in the recovery of vegetation structural components. Time‐since‐fire was the primary determinant of the structural recovery of T. scariosa and eucalypts. However, climate, notably mean annual rainfall and rainfall variability, also influenced the recovery of the eucalypt overstorey, T. scariosa cover and mean hummock height. We observed that (i) the mean number of live eucalypt stems per individual decreased while mean individual basal area increased, (ii) cover of T. scariosa peaked at ~30 years post‐fire and gradually decreased thereafter, and (iii) the ‘hummock’ form of T. scariosa occurred throughout the chronosequence, whereas the ‘ring’ form tended not to occur until ~30 years post‐fire. Time‐since‐fire was the key determinant of the structural recovery of eucalypt‐dominated mallee shrublands, but there is geographical variation in recovery related to rainfall and its variability. Fire regimes are likely to have different effects across the geographic range of mallee shrublands.     

Short‐term carbon cycling responses of a mature eucalypt woodland to gradual stepwise enrichment of atmospheric CO2 concentration

Projections of future climate are highly sensitive to uncertainties regarding carbon (C) uptake and storage by terrestrial ecosystems. The Eucalyptus Free‐Air CO₂ Enrichment (EucFACE) experiment was established to study the effects of elevated atmospheric CO₂ concentrations (eCO₂) on a native mature eucalypt woodland with low fertility soils in southeast Australia. In contrast to other FACE experiments, the concentration of CO₂ at EucFACE was increased gradually in steps above ambient (+0, 30, 60, 90, 120, and 150 ppm CO₂ above ambient of ~400 ppm), with each step lasting approximately 5 weeks. This provided a unique opportunity to study the short‐term (weeks to months) response of C cycle flux components to eCO₂ across a range of CO₂ concentrations in an intact ecosystem. Soil CO₂ efflux (i.e., soil respiration or R_soil) increased in response to initial enrichment (e.g., +30 and +60 ppm CO₂) but did not continue to increase as the CO₂ enrichment was stepped up to higher concentrations. Light‐saturated photosynthesis of canopy leaves (A_sat) also showed similar stimulation by elevated CO₂ at +60 ppm as at +150 ppm CO₂. The lack of significant effects of eCO₂ on soil moisture, microbial biomass, or activity suggests that the increase in R_soil likely reflected increased root and rhizosphere respiration rather than increased microbial decomposition of soil organic matter. This rapid increase in R_soil suggests that under eCO_2, additional photosynthate was produced, transported belowground, and respired. The consequences of this increased belowground activity and whether it is sustained through time in mature ecosystems under eCO₂ are a priority for future research.     

The implication of irrigation in climate change impact assessment: a European‐wide study

This study evaluates the impacts of projected climate change on irrigation requirements and yields of six crops (winter wheat, winter barley, rapeseed, grain maize, potato, and sugar beet) in Europe. Furthermore, the uncertainty deriving from consideration of irrigation, CO₂ effects on crop growth and transpiration, and different climate change scenarios in climate change impact assessments is quantified. Net irrigation requirement (NIR) and yields of the six crops were simulated for a baseline (1982–2006) and three SRES scenarios (B1, B2 and A1B, 2040–2064) under rainfed and irrigated conditions, using a process‐based crop model, SIMPLACE . We found that projected climate change decreased NIR of the three winter crops in northern Europe (up to 81 mm), but increased NIR of all the six crops in the Mediterranean regions (up to 182 mm yr^?1). Climate change increased yields of the three winter crops and sugar beet in middle and northern regions (up to 36%), but decreased their yields in Mediterranean countries (up to 81%). Consideration of CO₂ effects can alter the direction of change in NIR for irrigated crops in the south and of yields for C3 crops in central and northern Europe. Constraining the model to rainfed conditions for spring crops led to a negative bias in simulating climate change impacts on yields (up to 44%), which was proportional to the irrigation ratio of the simulation unit. Impacts on NIR and yields were generally consistent across the three SRES scenarios for the majority of regions in Europe. We conclude that due to the magnitude of irrigation and CO₂ effects, they should both be considered in the simulation of climate change impacts on crop production and water availability, particularly for crops and regions with a high proportion of irrigated crop area.     

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.     

Impacts of climate change on fire activity and fire management in the circumboreal forest

Forest fires are a significant and natural element of the circumboreal forest. Fire activity is strongly linked to weather, and increased fire activity due to climate change is anticipated or arguably has already occurred. Recent studies suggest a doubling of area burned along with a 50% increase in fire occurrence in parts of the circumboreal by the end of this century. Fire management agencies' ability to cope with these increases in fire activity is limited, as these organizations operate with a narrow margin between success and failure; a disproportionate number of fires may escape initial attack under a warmer climate, resulting in an increase in area burned that will be much greater than the corresponding increase in fire weather severity. There may be only a decade or two before increased fire activity means fire management agencies cannot maintain their current levels of effectiveness.     

Prioritizing forest fuels treatments based on the probability of high‐severity fire restores adaptive capacity in Sierran forests

In frequent fire forests of the western United States, a legacy of fire suppression coupled with increases in fire weather severity have altered fire regimes and vegetation dynamics. When coupled with projected climate change, these conditions have the potential to lead to vegetation type change and altered carbon (C) dynamics. In the Sierra Nevada, fuels reduction approaches that include mechanical thinning followed by regular prescribed fire are one approach to restore the ability of the ecosystem to tolerate episodic fire and still sequester C. Yet, the spatial extent of the area requiring treatment makes widespread treatment implementation unlikely. We sought to determine if a priori knowledge of where uncharacteristic wildfire is most probable could be used to optimize the placement of fuels treatments in a Sierra Nevada watershed. We developed two treatment placement strategies: the naive strategy, based on treating all operationally available area and the optimized strategy, which only treated areas where crown‐killing fires were most probable. We ran forecast simulations using projected climate data through 2,100 to determine how the treatments differed in terms of C sequestration, fire severity, and C emissions relative to a no‐management scenario. We found that in both the short (20 years) and long (100 years) term, both management scenarios increased C stability, reduced burn severity, and consequently emitted less C as a result of wildfires than no‐management. Across all metrics, both scenarios performed the same, but the optimized treatment required significantly less C removal (naive=0.42 Tg C, optimized=0.25 Tg C) to achieve the same treatment efficacy. Given the extent of western forests in need of fire restoration, efficiently allocating treatments is a critical task if we are going to restore adaptive capacity in frequent‐fire 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 genome‐wide search for local adaptation in a terrestrial‐breeding frog reveals vulnerability to climate change

Terrestrial‐breeding amphibians are likely to be vulnerable to warming and drying climates, as their embryos require consistent moisture for successful development. Adaptation to environmental change will depend on sufficient genetic variation existing within or between connected populations. Here, we use Single Nucleotide Polymorphism (SNP) data to investigate genome‐wide patterns in genetic diversity, gene flow and local adaptation in a terrestrial‐breeding frog (Pseudophryne guentheri) subject to a rapidly drying climate and recent habitat fragmentation. The species was sampled across 12 central and range‐edge populations (192 samples), and strong genetic structure was apparent, as were high inbreeding coefficients. Populations showed differences in genetic diversity, and one population lost significant genetic diversity in a decade. More than 500 SNP loci were putatively under directional selection, and 413 of these loci were correlated with environmental variables such as temperature, rainfall, evaporation and soil moisture. One locus showed homology to a gene involved in the activation of maturation in Xenopus oocytes, which may facilitate rapid development of embryos in drier climates. The low genetic diversity, strong population structuring and presence of local adaptation revealed in this study shows why management strategies such as targeted gene flow may be necessary to assist isolated populations to adapt to future climates.     

Genome‐wide scans detect adaptation to aridity in a widespread forest tree species

Patterns of adaptive variation within plant species are best studied through common garden experiments, but these are costly and time‐consuming, especially for trees that have long generation times. We explored whether genome‐wide scanning technology combined with outlier marker detection could be used to detect adaptation to climate and provide an alternative to common garden experiments. As a case study, we sampled nine provenances of the widespread forest tree species, Eucalyptus tricarpa, across an aridity gradient in southeastern Australia. Using a Bayesian analysis, we identified a suite of 94 putatively adaptive (outlying) sequence‐tagged markers across the genome. Population‐level allele frequencies of these outlier markers were strongly correlated with temperature and moisture availability at the site of origin, and with population differences in functional traits measured in two common gardens. Using the output from a canonical analysis of principal coordinates, we devised a metric that provides a holistic measure of genomic adaptation to aridity that could be used to guide assisted migration or genetic augmentation.     

Accelerating regrowth of temperate‐maritime forests due to environmental change

To understand how environmental changes have influenced forest productivity, stemwood biomass (B) dynamics were analyzed at 1267 permanent inventory plots, covering a combined 209 ha area of unmanaged temperate‐maritime forest in southwest British Columbia, Canada. Net stemwood production (ΔB) was derived from periodic remeasurements of B collected over a 40‐year measurement period (1959–1998) in stands ranging from 20 to 150 years old. Comparison between the integrated age response of net stemwood production, ΔB(A), and the age response of stemwood biomass, B(A), suggested a 58 ± 11% increase in ΔB between the first 40 years of the chronosequence period (1859–1898) and the measurement period. To estimate extrinsic forcing on ΔB, several different candidate models were developed to remove variation explained by intrinsic factors. All models exhibited temporal bias, with positive trends in (observed minus predicted) residual ΔB ranging between of 0.40 and 0.64% yr^?1. Applying the same methods to stemwood growth (G) indicated residual increases ranging from 0.43 and 0.67% yr^?1. Higher trend estimates corresponded with models that included site index (SI) as a predictor, which may reflect exaggeration of the age‐decline in SI tables. Choosing a model that excluded SI, suggested that ΔB increased by 0.40 ± 0.18% yr^?1, while G increased by 0.43 ± 0.12% yr^?1 over the measurement period. Residual G was significantly correlated with atmospheric carbon dioxide (CO₂), temperature (T), and climate moisture index (CMI). However, models driven with climate and CO₂, alone, could not simultaneously explain long‐term and measurement‐period trends without additional representation of indirect effects, perhaps reflecting compound interest on direct physiological responses to environmental change. Evidence of accelerating forest regrowth highlights the value of permanent inventories to detect and understand systematic changes in forest productivity caused by environmental change.     

Landscape dynamics in Mediterranean oak forests under global change: understanding the role of anthropogenic and environmental drivers across forest types

The Mediterranean region is projected to be extremely vulnerable to global change, which will affect the distribution of typical forest types such as native oak forests. However, our understanding of Mediterranean oak forest responses to future conditions is still very limited by the lack of knowledge on oak forest dynamics and species‐specific responses to multiple drivers. We compared the long‐term (1966–2006) forest persistence and land cover change among evergreen (cork oak and holm oak) and deciduous oak forests and evaluated the importance of anthropogenic and environmental drivers on observed changes for Portugal. We used National Forest Inventories to quantify the changes in oak forests and explored the drivers of change using multinomial logistic regression analysis and an information theoretical approach. We found distinct trends among oak forest types, reflecting the differences in oak economic value, protection status and management schemes: cork oak forests were the most persistent (62%), changing mostly to pines and eucalypt; holm oak forests were less persistent (53.2%), changing mostly to agriculture; and deciduous oak forests were the least persistent (45.7%), changing mostly to shrublands. Drivers of change had distinct importance across oak forest types, but drivers from anthropogenic origin (wildfires, population density, and land accessibility) were always among the most important. Climatic extremes were also important predictors of oak forest changes, namely extreme temperatures for evergreen oak forests and deficit of precipitation for deciduous oak forests. Our results indicate that under increasing human pressure and forecasted climate change, evergreen oak forests will continue declining and deciduous oak forests will be replaced by forests dominated by more xeric species. In the long run, multiple disturbances may change competitive dominance from oak forests to pyrophytic shrublands. A better understanding of forest dynamics and the inclusion of anthropogenic drivers on models of vegetation change will improve predicting the future of Mediterranean oak forests.     

Interactive direct and plant‐mediated effects of elevated atmospheric [CO2] and temperature on a eucalypt‐feeding insect herbivore

Understanding the direct and indirect effects of elevated [CO₂] and temperature on insect herbivores and how these factors interact are essential to predict ecosystem‐level responses to climate change scenarios. In three concurrent glasshouse experiments, we measured both the individual and interactive effects of elevated [CO₂] and temperature on foliar quality. We also assessed the interactions between their direct and plant‐mediated effects on the development of an insect herbivore of eucalypts. Eucalyptus tereticornis saplings were grown at ambient or elevated [CO₂] (400 and 650 μmol mol^?1 respectively) and ambient or elevated ( + 4 °C) temperature for 10 months. Doratifera quadriguttata (Lepidoptera: Limacodidae) larvae were feeding directly on these trees, on their excised leaves in a separate glasshouse, or on excised field‐grown leaves within the temperature and [CO₂] controlled glasshouse. To allow insect gender to be determined and to ensure that any sex‐specific developmental differences could be distinguished from treatment effects, insect development time and consumption were measured from egg hatch to pupation. No direct [CO₂] effects on insects were observed. Elevated temperature accelerated larval development, but did not affect leaf consumption. Elevated [CO₂] and temperature independently reduced foliar quality, slowing larval development and increasing consumption. Simultaneously increasing both [CO₂] and temperature reduced these shifts in foliar quality, and negative effects on larval performance were subsequently ameliorated. Negative nutritional effects of elevated [CO₂] and temperature were also independently outweighed by the direct positive effect of elevated temperature on larvae. Rising [CO₂] and temperature are thus predicted to have interactive effects on foliar quality that affect eucalypt‐feeding insects. However, the ecological consequences of these interactions will depend on the magnitude of concurrent temperature rise and its direct effects on insect physiology and feeding behaviour.     

Altered fire regimes cause long‐term lichen diversity losses

Many global ecosystems have undergone shifts in fire regimes in recent decades, such as changes in fire size, frequency, and/or severity. Recent research shows that increases in fire size, frequency, and severity can lead to long‐persisting deforestation, but the consequences of shifting fire regimes for biodiversity of other vegetative organisms (such as understory plants, fungi, and lichens) remain poorly understood. Understanding lichen responses to wildfire is particularly important because lichens play crucial roles in nutrient cycling and supporting wildlife in many ecosystems. Lichen responses to fire have been little studied, and most previous research has been limited to small geographic areas (e.g. studies of a single fire), making it difficult to establish generalizable patterns. To investigate long‐term effects of fire severity on lichen communities, we sampled epiphytic lichen communities in 104 study plots across California's greater Sierra Nevada region in areas that burned in five wildfires, ranging from 4 to 16 years prior to sampling. The conifer forest ecosystems we studied have undergone a notable increase in fire severity in recent decades, and we sample across the full gradient of fire severity to infer how shifting fire regimes may influence landscape‐level biodiversity. We find that low‐severity fire has little to no effect on lichen communities. Areas that burned at moderate and high severities, however, have significantly and progressively lower lichen richness and abundance. Importantly, we observe very little postfire lichen recolonization on burned substrates even more than 15 years after fire. Our multivariate model suggests that the hotter, drier microclimates that occur after fire removes forest canopies may prevent lichen reestablishment, meaning that lichens are not likely to recolonize until mature trees regenerate. These findings suggest that altered fire regimes may cause broad and long‐persisting landscape‐scale biodiversity losses that could ultimately impact multiple trophic levels.