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.     

Long‐term post‐fire changes in the northeastern boreal forest of Quebec

Abstract. Natural dynamics in the boreal forest is influenced by disturbances. Fire recurrence affects community development and landscape diversity. Forest development was studied in the northeastern boreal forest of Quebec. The objective was to describe succession following fire and to assess the factors related to the changes in forest composition and structure. The study area is located in northeastern Quebec, 50 km north of Baie‐Comeau. We used the forest inventory data gathered by the Ministère des Ressources naturelles du Québec (MRNQ). In circular plots of 400 m², the diameter at breast height (DBH) of all stems of tree species greater than 10 cm was recorded and in 40 m² subplots, stems smaller than 10 cm were measured. A total of 380 plots were sampled in an area of 6000 km². The fire history reconstruction was done based on historical maps, old aerial photographs and field sampling. A time‐since‐fire class, a deposit type, slope, slope aspect and altitude were attributed to each plot. Each plot was also described according to species richness and size structure characteristics. Traces of recent disturbance were also recorded in each plot. Changes in forest composition were described using ordination analyses (NMDS and CCA) and correlated with the explanatory variables. Two successional pathways were observed in the area and characterized by the early dominance of intolerant hardwood species or Picea mariana. With time elapsed since the last fire, composition converged towards either Picea mariana, Abies balsamea or a mixture of both species and the size structure of the coniferous dominated stands got more irregular. The environmental conditions varied between stands and explained part of the variability in composition. Their effect tended to decrease with increasing time elapsed since fire, as canopy composition was getting more similar. Gaps may be important to control forest dynamics in old successional communities.     

Effects of long‐term fire exclusion and frequent fire on plant community composition: A case study from semi‐arid shrublands

Time since last fire and fire frequency are strong determinants of plant community composition in fire‐prone landscapes. Our study aimed to establish the influence of time since last fire and fire frequency on plant community composition and diversity of a south‐west Australian semi‐arid shrubland. We employed a space‐for‐time approach using four fire age classes: ‘young’, 8–15 years since last fire; ‘medium’, 16–34; ‘old’, 35–50; and ‘very old’, 51–100; and three fire frequency classes: burnt once, twice and three times within the last 50 years. Species diversity was compared using one‐way ANOVA and species composition using PERMANOVA. Soil and climatic variables were included as covariables to partition underlying environmental drivers. We found that time since last fire influenced species richness, diversity and composition. Specifically, we recorded a late successional transition from woody seeders to long‐lived, arid‐zone, resprouting shrub species. Fire frequency did not influence species richness and diversity but did influence species composition via a reduction in cover of longer‐lived resprouter species – presumably because of a reduced ability to replenish epicormic buds and/or sufficient starch stores. The distinct floristic composition of old and very old habitat, and the vulnerability of these areas to wildfires, indicate that these areas are ecologically important and management should seek to preserve them.     

Human disturbance and environmental factors as drivers of long‐term post‐fire regeneration patterns in Mediterranean forests

Question: What are the main forces driving natural regeneration in burned mature Mediterranean forests in the medium‐long term and what are the likely successional trajectories of unmanaged vegetation? Location: Valencia Region, eastern Spain. Methods: A wildfire burned 33 000 ha of Pinus halepensis and P. pinaster forest in 1979, and subsequent smaller wildfires took place between 1984 and 1996. The study was designed to sample the range of environmental and disturbance (fire recurrence and land use) conditions. The territory was classified into 17 different geomorphological and fire‐recurrence units. Vegetation cover and floristic composition were measured on a total of 113 plots (1000 m² each) randomly selected within these units. Results: The results show that 23 years after the fire the regenerated vegetation consists of successional shrublands, and that forest ecosystem resilience can be very low. The vegetation presents a strong correlation with most of the environmental variables, but fire (one or two fires), soil type and land use (in that order) are the main drivers of vegetation composition. Quercus coccifera shrublands persist on limestone soils while diverse types of other shrublands (dominated by seeder species) are found on marl soils. Conclusions: The results of this study indicate that disturbance factors strongly coupled to human activities, such as land use and fire, play a critical role in the current state of vegetation. Fire creates vegetation patches in different successional states while land use and soil type define the different types of shrubland in terms of their specific composition.     

Short‐term effects of herbicides and a prescribed fire on restoration of a shrub‐encroached pine savanna

Shrub encroachment occurring worldwide in savannas and grasslands has commonly been hypothesized to result from anthropogenically altered environments. Two disturbance‐based approaches to restoration have involved: (1) application of selective herbicides to reduce density/cover of shrubs; (2) reinstatement of natural fire regimes to generate environmental conditions favoring herbaceous species. We studied short‐term responses of native shrubs, vines, and grasses in a Louisiana pine savanna to herbicides followed by a prescribed fire and fire alone. Plots established in the summer, 2013, were hand‐sprayed in the fall with Imazapyr and Triclopyr, Triclopyr alone, or no herbicide, then prescribed burned the following spring. Numbers of species of shrubs and vines at scales of 1 and 100 m², numbers of stems and regrowth of stems produced by six common species of shrubs, and the number of flowering culms of perennial C₄ grasses were assessed postfire in 2014. Compared with fire alone, herbicides followed by fire resulted in (1) small reductions in species richness of shrubs and no effects on vines, (2) fewer stems comprising shrub genets, but similar postfire regrowth of resprouting shrub stems, and (3) fewer flowering culms of C₄ grasses. Underground storage organs of savanna shrubs and vines survived both aboveground disturbances. Thus, single applications of herbicides followed by fires reduced, but did not reverse shrub encroachment and negatively affected grasses. Because effects of herbicides overrode those of prescribed fires, these disturbances did not act synergistically, suggesting that reinstating natural fire regimes should be a priority in restoration of savannas and grasslands.     

Long‐term variability and rainfall control of savanna fire regimes in equatorial East Africa

Fires burning the vast grasslands and savannas of Africa significantly influence the global carbon cycle. Projecting the impacts of future climate change on fire‐mediated biogeochemical processes in these dry tropical ecosystems requires understanding of how various climate factors influence regional fire regimes. To examine climate–vegetation–fire linkages in dry savanna, we conducted macroscopic and microscopic charcoal analysis on the sediments of the past 25 000 years from Lake Challa, a deep crater lake in equatorial East Africa. The charcoal‐inferred shifts in local and regional fire regimes were compared with previously published reconstructions of temperature, rainfall, seasonal drought severity, and vegetation dynamics to evaluate millennial‐scale drivers of fire occurrence. Our charcoal data indicate that fire in the dry lowland savanna of southeastern Kenya was not fuel‐limited during the Last Glacial Maximum (LGM) and Late Glacial, in contrast to many other regions throughout the world. Fire activity remained high at Lake Challa probably because the relatively high mean‐annual temperature (~22 °C) allowed productive C₄ grasses with high water‐use efficiency to dominate the landscape. From the LGM through the middle Holocene, the relative importance of savanna burning in the region varied primarily in response to changes in rainfall and dry‐season length, which were controlled by orbital insolation forcing of tropical monsoon dynamics. The fuel limitation that characterizes the region's fire regime today appears to have begun around 5000–6000 years ago, when warmer interglacial conditions coincided with prolonged seasonal drought. Thus, insolation‐driven variation in the amount and seasonality of rainfall during the past 25 000 years altered the immediate controls on fire occurrence in the grass‐dominated savannas of eastern equatorial Africa. These results show that climatic impacts on dry‐savanna burning are heterogeneous through time, with important implications for efforts to anticipate future shifts in fire‐mediated ecosystem processes.     

Post‐fire carbon dynamics in the tropical peat swamp forests of Brunei reveal long‐term elevated CH4 flux

Tropical peatlands hold about 15%–19% of the global peat carbon (C) pool of which 77% is stored in the peat swamp forests (PSFs) of Southeast Asia. Nonetheless, these PSFs have been drained, exploited for timber and land for agriculture, leading to frequent fires in the region. The physico‐chemical characteristics of peat, as well as the hydrology of PSFs are affected after a fire, during which the ecosystem can act as a C source for decades, as C emissions to the atmosphere exceed photosynthesis. In this work, we studied the longer‐term impact of fires on C cycling in tropical PSFs, hence we quantified the magnitude and patterns of C loss (CO₂, CH₄ and dissolved organic carbon) and soil‐water quality characteristics in an intact and a degraded burnt PSF in Brunei Darussalam affected by seven fires over the last 40 years. We used natural tracers such as ¹⁴C to investigate the age and sources of C contributing to ecosystem respiration (R_eco) and CH₄, while we continuously monitored soil temperature and water table (WT) level from June 2017 to January 2019. Our results showed a major difference in the physico‐chemical parameters, which in turn affected C dynamics, especially CH₄. Methane effluxes were higher in fire‐affected areas (7.8 ± 2.2 mg CH₄ m^?2 hr^?1) compared to the intact PSF (4.0 ± 2.0 mg CH₄ m^?2 hr^?1) due to prolonged higher WT and more optimal methanogenesis conditions. On the other hand, we did not find significant differences in R_eco between burnt (432 ± 83 mg CO₂ m^?2 hr^?1) and intact PSF (359 ± 76 mg CO₂ m^?2 hr^?1). Radiocarbon analysis showed overall no significant difference between intact and burnt PSF with a modern signature for both CO₂ and CH₄ fluxes implying a microbial preference for the more labile C fraction in the peat matrix.     

Short‐term insurance versus long‐term bet‐hedging strategies as adaptations to variable environments

Understanding how organisms adapt to environmental variation is a key challenge of biology. Central to this are bet‐hedging strategies that maximize geometric mean fitness across generations, either by being conservative or diversifying phenotypes. Theoretical models have identified environmental variation across generations with multiplicative fitness effects as driving the evolution of bet‐hedging. However, behavioral ecology has revealed adaptive responses to additive fitness effects of environmental variation within lifetimes, either through insurance or risk‐sensitive strategies. Here, we explore whether the effects of adaptive insurance interact with the evolution of bet‐hedging by varying the position and skew of both arithmetic and geometric mean fitness functions. We find that insurance causes the optimal phenotype to shift from the peak to down the less steeply decreasing side of the fitness function, and that conservative bet‐hedging produces an additional shift on top of this, which decreases as adaptive phenotypic variation from diversifying bet‐hedging increases. When diversifying bet‐hedging is not an option, environmental canalization to reduce phenotypic variation is almost always favored, except where the tails of the fitness function are steeply convex and produce a novel risk‐sensitive increase in phenotypic variance akin to diversifying bet‐hedging. Importantly, using skewed fitness functions, we provide the first model that explicitly addresses how conservative and diversifying bet‐hedging strategies might coexist.     

The effect of long‐term repeated burning and fire exclusion on above‐ and below‐ground Blackbutt (Eucalyptus pilularis) forest vegetation assemblages

We used a long‐term fire experiment in south‐east Queensland, Australia, to determine the effects of frequent prescribed burning and fire exclusion on understorey vegetation (<7.5 m) richness and density in Eucalyptus pilularis forest. Our study provided a point in time assessment of the standing vegetation and soil‐stored vegetation at two experimental sites with treatments of biennial burning, quadrennial burning since 1971–1972 and no burning since 1969. Vegetation composition, density and richness of certain plant groups in the standing and soil‐stored vegetation were influenced by fire treatments. The density of resprouting plants <3 m in height was higher in the biennially burnt treatment than in the unburnt treatment, but resprouters 3–7.5 m in height were absent from the biennial burning treatment. Obligate seeder richness and density in the standing vegetation was not significantly influenced by the fire treatments, but richness of this plant group in the seed bank was higher in the quadrennial treatment at one site and in the long unburnt treatment at the other site. Long unburnt treatments had an understorey of rainforest species, while biennial burning at one site and quadrennial burning at the other site were associated with greater standing grass density relative to the unburnt treatment. This difference in vegetation composition due to fire regime potentially influences the flammability of the standing understorey vegetation. Significant interactions between fire regime and site, apparent in the standing and soil‐stored vegetation, demonstrate the high degree of natural variability in vegetation community responses to fire regimes.     

Experimental fire increases soil carbon dioxide efflux in a grassland long‐term multifactor global change experiment

Numerous studies have demonstrated that soil respiration rates increase under experimental warming, although the long‐term, multiyear dynamics of this feedback are not well constrained. Less is known about the effects of single, punctuated events in combination with other longer‐duration anthropogenic influences on the dynamics of soil carbon (C) loss. In 2012 and 2013, we assessed the effects of decadal‐scale anthropogenic global change – warming, increased nitrogen (N) deposition, elevated carbon dioxide (CO₂), and increased precipitation – on soil respiration rates in an annual‐dominated Mediterranean grassland. We also investigated how controlled fire and an artificial wet‐up event, in combination with exposure to the longer‐duration anthropogenic global change factors, influenced the dynamics of C cycling in this system. Decade‐duration surface soil warming (1–2 °C) had no effect on soil respiration rates, while +N addition and elevated CO₂ concentrations increased growing‐season soil CO₂ efflux rates by increasing annual aboveground net primary production (NPP) and belowground fine root production, respectively. Low‐intensity experimental fire significantly elevated soil CO₂ efflux rates in the next growing season. Based on mixed‐effects modeling and structural equation modeling, low‐intensity fire increased growing‐season soil respiration rates through a combination of three mechanisms: large increases in soil temperature (3–5 °C), significant increases in fine root production, and elevated aboveground NPP. Our study shows that in ecosystems where soil respiration has acclimated to moderate warming, further increases in soil temperature can stimulate greater soil CO₂ efflux. We also demonstrate that punctuated short‐duration events such as fire can influence soil C dynamics with implications for both the parameterization of earth system models (ESMs) and the implementation of climate change mitigation policies that involve land‐sector C accounting.     

Land‐use and fire history effects on post‐fire vegetation dynamics in eastern Spain

Question: Is post‐fire, medium‐term vegetation dynamics determined by land‐use or fire history prior to fire? Location: South‐facing slope in the Gallinera valley, Alicante province, eastern Spain. Methods: After mapping the land‐use and fire history of the study site using photo‐interpretation, we sampled vegetation structure on a set of plots representing the most frequent land‐use and fire history combinations on an area burned six years before sampling. We studied the effects of land‐use history, comparing the one‐fire land‐use trajectories. We analysed the effects of fire history; comparing one‐ and two‐fire plots for both previously cropped and uncropped areas. Results: Most variables were not significantly different between the earliest abandoned plots (abandoned at least 38 years before the fire) and the uncropped plots. On the most recently abandoned plots (abandoned between one and four years before the fire), the therophyte richness and the ratio of seeder: resprouter richness were significantly greatest. Different fire recurrences did not determine different post‐fire vegetation on either the uncropped or the early abandoned plots (all dominated by fire‐recruited seeder shrubs). The most recently abandoned plots had a lower resilience to fire. Conclusions: Land‐use history and recent pre‐fire land use, in particular, determined the post‐fire vegetation in the medium term. The vegetation composition converged during secondary succession among land‐use histories. Increasing fire recurrence had a small effect on mature plant communities, due to the combination of life‐history traits determining the response to fire of the dominant species.     

Climate and the inter‐annual variability of fire in southern Africa: a meta‐analysis using long‐term field data and satellite‐derived burnt area data

Aim This study investigates inter‐annual variability in burnt area in southern Africa and the extent to which climate is responsible for this variation. We compare data from long‐term field sites across the region with remotely sensed burnt area data to test whether it is possible to develop a general model. Location Africa south of the equator. Methods Linear mixed effects models were used to determine the effect of rainfall, seasonality and fire weather in driving variation in fire extent between years, and to test whether the effect of these variables changes across the subcontinent and in areas more and less impacted by human activities. Results A simple model including rainfall and seasonality explained 40% of the variance in burnt area between years across 10 different protected areas on the subcontinent, but this model, when applied regionally, indicated that climate had less impact on year‐to‐year variation in burnt area than would be expected. It was possible to demonstrate that the relative importance of rainfall and seasonality changed as one moved from dry to wetter systems, but most noticeable was the reduction in climatically driven variability of fire outside protected areas. Inter‐annual variability is associated with the occurrence of large fires, and large fires are only found in areas with low human impact. Main conclusions This research gives the first data‐driven analysis of fire–climate interactions in southern Africa. The regional analysis shows that human impact on fire regimes is substantial and acts to limit the effect of climate in driving variation between years. This is in contrast to patterns in protected areas, where variation in accumulated rainfall and the length of the dry season influence the annual area burnt. Global models which assume strong links between fire and climate need to be re‐assessed in systems with high human impact.     

Restorative mowing on an abandoned semi‐natural meadow: short‐term and predicted long‐term effects

Abstract. When management, in the form of cattle grazing and mowing, ceases the abundance of competitively superior plant species tends to increase in abandoned semi‐natural meadows. Litter accumulation elevates the soil nutrient levels and hinders seedling recruitment. We surveyed changes in plant cover and species composition of a formerly grazed meadow in permanent plots for six years. Some plots were unmown, while others were mown and raked annually in August. The cover of grasses decreased and herb cover remained unchanged regardless of the treatment. Mowing and raking significantly reduced litter accumulation and increased the number of ground layer species. The expected long‐term effects of abandonment and restorative mowing were studied by calculating the transition probabilities for unmown and mown plots and simulating the course of succession as projected by the transition matrices. During a simulation period of 30 yr, abandonment led to (1) a decrease in the cover of small herbs, (2) a slight increase in the cover of tall herbs and (3) a slight decrease in the cover of grasses. In contrast, the cover of small herbs on the mown plots remained unchanged or slightly increased during the course of simulation. These results suggest that mowing late in the season is primarily a management tool for the maintenance of the existing species diversity and composition. However, it may not be an effective restorative tool to induce overall changes in the resident vegetation of abandoned grass‐dominated meadows. Grazing or mowing early in the season may be more effective in this respect. Consequently, mowing early or, alternatively, late in the season may provide management strategies for the maintenance and restoration of species diversity, respectively.     

Carbon dynamics in the future forest: the importance of long‐term successional legacy and climate–fire interactions

Understanding how climate change may influence forest carbon (C) budgets requires knowledge of forest growth relationships with regional climate, long‐term forest succession, and past and future disturbances, such as wildfires and timber harvesting events. We used a landscape‐scale model of forest succession, wildfire, and C dynamics (LANDIS‐II) to evaluate the effects of a changing climate (A2 and B1 IPCC emissions; Geophysical Fluid Dynamics Laboratory General Circulation Models) on total forest C, tree species composition, and wildfire dynamics in the Lake Tahoe Basin, California, and Nevada. The independent effects of temperature and precipitation were assessed within and among climate models. Results highlight the importance of modeling forest succession and stand development processes at the landscape scale for understanding the C cycle. Due primarily to landscape legacy effects of historic logging of the Comstock Era in the late 1880s, C sequestration may continue throughout the current century, and the forest will remain a C sink (Net Ecosystem Carbon Balance > 0), regardless of climate regime. Climate change caused increases in temperatures limited simulated C sequestration potential because of augmented fire activity and reduced establishment ability of subalpine and upper montane trees. Higher temperatures influenced forest response more than reduced precipitation. As the forest reached its potential steady state, the forest could become C neutral or a C source, and climate change could accelerate this transition. The future of forest ecosystem C cycling in many forested systems worldwide may depend more on major disturbances and landscape legacies related to land use than on projected climate change alone.