Convergence of bark investment according to fire and climate structures ecosystem vulnerability to future change

Fire regimes in savannas and forests are changing over much of the world. Anticipating the impact of these changes requires understanding how plants are adapted to fire. In this study, we test whether fire imposes a broad selective force on a key fire‐tolerance trait, bark thickness, across 572 tree species distributed worldwide. We show that investment in thick bark is a pervasive adaptation in frequently burned areas across savannas and forests in both temperate and tropical regions where surface fires occur. Geographic variability in bark thickness is largely explained by annual burned area and precipitation seasonality. Combining environmental and species distribution data allowed us to assess vulnerability to future climate and fire conditions: tropical rainforests are especially vulnerable, whereas seasonal forests and savannas are more robust. The strong link between fire and bark thickness provides an avenue for assessing the vulnerability of tree communities to fire and demands inclusion in global models.     

Thinner bark increases sensitivity of wetter Amazonian tropical forests to fire

Understory fires represent an accelerating threat to Amazonian tropical forests and can, during drought, affect larger areas than deforestation itself. These fires kill trees at rates varying from < 10 to c. 90% depending on fire intensity, forest disturbance history and tree functional traits. Here, we examine variation in bark thickness across the Amazon. Bark can protect trees from fires, but it is often assumed to be consistently thin across tropical forests. Here, we show that investment in bark varies, with thicker bark in dry forests and thinner in wetter forests. We also show that thinner bark translated into higher fire‐driven tree mortality in wetter forests, with between 0.67 and 5.86 gigatonnes CO₂ lost in Amazon understory fires between 2001 and 2010. Trait‐enabled global vegetation models that explicitly include variation in bark thickness are likely to improve the predictions of fire effects on carbon cycling in tropical forests.     

When is a ‘forest’ a savanna,and why does it matter?

Savannas are defined based on vegetation structure, the central concept being a discontinuous tree cover in a continuous grass understorey. However, at the high‐rainfall end of the tropical savanna biome, where heavily wooded mesic savannas begin to structurally resemble forests, or where tropical forests are degraded such that they open out to structurally resemble savannas, vegetation structure alone may be inadequate to distinguish mesic savanna from forest. Additional knowledge of the functional differences between these ecosystems which contrast sharply in their evolutionary and ecological history is required. Specifically, we suggest that tropical mesic savannas are predominantly mixed tree–C₄ grass systems defined by fire tolerance and shade intolerance of their species, while forests, from which C₄ grasses are largely absent, have species that are mostly fire intolerant and shade tolerant. Using this framework, we identify a suite of morphological, physiological and life‐history traits that are likely to differ between tropical mesic savanna and forest species. We suggest that these traits can be used to distinguish between these ecosystems and thereby aid their appropriate management and conservation. We also suggest that many areas in South Asia classified as tropical dry forests, but characterized by fire‐resistant tree species in a C₄ grass‐dominated understorey, would be better classified as mesic savannas requiring fire and light to maintain the unique mix of species that characterize them.     

Trade‐offs between savanna woody plant diversity and carbon storage in the Brazilian Cerrado

Incentivizing carbon storage can be a win‐win pathway to conserving biodiversity and mitigating climate change. In savannas, however, the situation is more complex. Promoting carbon storage through woody encroachment may reduce plant diversity of savanna endemics, even as the diversity of encroaching forest species increases. This trade‐off has important implications for the management of biodiversity and carbon in savanna habitats, but has rarely been evaluated empirically. We quantified the nature of carbon‐diversity relationships in the Brazilian Cerrado by analyzing how woody plant species richness changed with carbon storage in 206 sites across the 2.2 million km² region at two spatial scales. We show that total woody plant species diversity increases with carbon storage, as expected, but that the richness of endemic savanna woody plant species declines with carbon storage both at the local scale, as woody biomass accumulates within plots, and at the landscape scale, as forest replaces savanna. The sharpest trade‐offs between carbon storage and savanna diversity occurred at the early stages of carbon accumulation at the local scale but the final stages of forest encroachment at the landscape scale. Furthermore, the loss of savanna species quickens in the final stages of forest encroachment, and beyond a point, savanna species losses outpace forest species gains with increasing carbon accumulation. Our results suggest that although woody encroachment in savanna ecosystems may provide substantial carbon benefits, it comes at the rapidly accruing cost of woody plant species adapted to the open savanna environment. Moreover, the dependence of carbon‐diversity trade‐offs on the amount of savanna area remaining requires land managers to carefully consider local conditions. Widespread woody encroachment in both Australian and African savannas and grasslands may present similar threats to biodiversity.     

Fire and drought: Shifts in bark investment across a broad geographical scale for Neotropical savanna trees

Savanna tree communities occurring in confluence zones with other biomes likely experience different environmental pressures, resulting in shifts in the selection of individual traits, the combinations of such traits, and species composition. In seasonally dry fire-prone environments, plant survival is presumably associated with adaptive changes in bark properties related to fire protection and water storage. Here, we integrated the multiple functions of the bark to investigate whether different selective pressures could influence patterns of variation in bark structure and allocation across species in a broad geographical range. We measured thickness, density, and water content of the inner and outer bark in branches and the main stem of the 51 most abundant species in three savanna communities differing in climatic aridity, one located at the core region of Cerrado in Central Brazil and the other two at its periphery, in the transition zones with Amazonia and Atlantic forest biomes. We found no difference in outer bark thickness but markedly difference in inner bark thickness between the three plant communities. In the central region, where dry season is long and fire is frequent, branches and main stem showed thicker inner bark. Contrastingly, in the south periphery region, where dry season is short, species showed thinner inner bark in both branches and main stem. Species from the north periphery region, where mean annual precipitation is higher, but fire is frequent and the dry season is also long, showed similar main stem inner bark thickness, but thinner branch inner bark compared to core region species. Our findings support the idea that investing in inner bark thickness and bark moisture may be the most advantageous strategy in plant communities that suffer from high evaporative demand during a long period and are at a high risk of fire.     

Expansion of gallery forests into central Brazilian savannas

Upland tropical forests have expanded and contracted in response to past climates, but it is not clear whether similar dynamics were exhibited by gallery (riparian) forests within savanna biomes. Because such forests generally have access to ample water, their extent may be buffered against changing climates. We tested the long‐term stability of gallery forest boundaries by characterizing the border between gallery forests and savannas and tracing the presence of gallery forest through isotopic analysis of organic carbon in the soil profile. We measured leaf area index, grass vs. shrub or tree coverage, the organic carbon, phosphorus, nitrogen and calcium concentrations in soils and the carbon isotope ratios of soil organic matter in two transitions spanning gallery forests and savanna in a Cerrado ecosystem. Gallery forests without grasses typically show a greater leaf area index in contrast to savannas, which show dense grass coverage. Soils of gallery forests have significantly greater concentrations of organic carbon, phosphorus, nitrogen and calcium than those of savannas. Soil organic carbon of savannas is significantly more enriched in ¹³C compared with that of gallery forests. This difference in enrichment is in part caused by the presence of C₄ grasses in savanna ecosystem and its absence in gallery forests. Using the ¹³C abundance as a signature for savanna and gallery forest ecosystems in 1 m soil cores, we show that the borders of gallery forests have expanded into the savanna and that this process initiated at least 3000–4000 bp based on ¹⁴C analysis. Gallery forests, however, may be still expanding as we found more recent transitions according to ¹⁴C activity measurements. We discuss the possible mechanisms of gallery forest expansion and the means by which nutrients required for the expansion of gallery forest might accumulate.     

Fire‐induced tree mortality in a neotropical forest: the roles of bark traits,tree size,wood density and fire behavior

Large‐scale wildfires are expected to accelerate forest dieback in Amazônia, but the fire vulnerability of tree species remains uncertain, in part due to the lack of studies relating fire‐induced mortality to both fire behavior and plant traits. To address this gap, we established two sets of experiments in southern Amazonia. First, we tested which bark traits best predict heat transfer rates (R) through bark during experimental bole heating. Second, using data from a large‐scale fire experiment, we tested the effects of tree wood density (WD), size, and estimated R (inverse of cambium insulation) on tree mortality after one to five fires. In the first experiment, bark thickness explained 82% of the variance in R, while the presence of water in the bark reduced the difference in temperature between the heat source and the vascular cambium, perhaps because of high latent heat of vaporization. This novel finding provides an important insight for improving mechanistic models of fire‐induced cambium damage from tropical to temperate regions. In the second experiment, tree mortality increased with increasing fire intensity (i.e. as indicated by bark char height on tree boles), which was higher along the forest edge, during the 2007 drought, and when the fire return interval was 3 years instead of one. Contrary to other tropical studies, the relationship between mortality and fire intensity was strongest in the year following the fires, but continued for 3 years afterwards. Tree mortality was low (≤20%) for thick‐barked individuals (≥18 mm) subjected to medium‐intensity fires, and significantly decreased as a function of increasing tree diameter, height and wood density. Hence, fire‐induced tree mortality was influenced not only by cambium insulation but also by other traits that reduce the indirect effects of fire. These results can be used to improve assessments of fire vulnerability of tropical forests.     

Savannas after afforestation: Assessment of herbaceous community responses to wildfire versus native tree planting

Afforestation and fire exclusion are pervasive threats to tropical savannas. In Brazil, laws limiting prescribed burning hinder the study of fire in the restoration of Cerrado plant communities. We took advantage of a 2017 wildfire to evaluate the potential for tree cutting and fire to promote the passive restoration of savanna herbaceous plant communities after destruction by exotic tree plantations. We sampled a burned pine plantation (Burned Plantation); a former plantation that was harvested and burned (Harvested & Burned); an unburned former plantation that was harvested, planted with native trees, and treated with herbicide to control invasive grasses (Native Tree Planting); and two old-growth savannas which served as reference communities. Our results confirm that herbaceous plant communities on post-afforestation sites are very different from old-growth savannas. Among post-afforestation sites, Harvested & Burned herbaceous communities were modestly more similar in composition to old-growth savannas, had slightly higher richness of savanna plants (3.8 species per 50-m²), and supported the greatest cover of native herbaceous plants (56%). These positive trends in herbaceous community recovery would be missed in assessments of tree cover: whereas canopy cover in the Harvested & Burned site was 6% (less than typical of savannas of the Cerrado), the Burned Plantation and Native Tree Planting supported 34% and 19% cover, respectively. By focusing on savanna herbaceous plants, these results highlight that tree cutting and fire, not simply tree planting and fire exclusion, should receive greater attention in efforts to restore savannas of the Cerrado.     

Nitrogen deposition in tropical forests from savanna and deforestation fires

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

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

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

Bark thickness determines fire resistance of selected tree species from fire-prone tropical savanna in north Australia

We investigated the fire resistance conferred by bark of seven common tree species in north Australian tropical savannas. We estimated bark thermal conductance and examined the relative importance of bark thickness, density and moisture content for protecting the cambium from lethal fire temperatures. Eucalypt and non-eucalypt species were contrasted, including the fire-sensitive conifer Callitris intratropica. Cambial temperature responses to bark surface heating were measured using a modified wick-fire technique, which simulated a heat pulse comparable to surface fires of moderate intensity. Bark thickness was a better predictor of resistance to cambial injury from fires than either bark moisture or density, accounting for 68% of the deviance in maximum temperature of the cambium. The duration of heating required to kill the cambium of a tree (τ_c) was directly proportional to bark thickness squared. Although species did not differ significantly in their bark thermal conductance (k), the thinner barked eucalypts nevertheless achieved similar or only slightly lower levels of fire resistance than much thicker barked non-eucalypts. Bark thickness alone cannot account for the latter and we suggest that lower bark moisture content among the eucalypts also contributes to their apparent fire resistance. Unique eucalypt meristem anatomy and epicormic structures, combined with their bark traits, probably facilitate resprouting after fire and ensure the dominance of eucalypts in fire-prone savannas. This study emphasises the need to take into account both the thermal properties of bark and the mechanism of bud protection in characterising the resprouting ability of savanna trees.     

Massive mortality of aspen following severe drought along the southern edge of the Canadian boreal forest

Drought‐induced, regional‐scale dieback of forests has emerged as a global concern that is expected to escalate under model projections of climate change. Since 2000, drought of unusual severity, extent, and duration has affected large areas of western North America, leading to regional‐scale dieback of forests in the southwestern US. We report on drought impacts on forests in a region farther north, encompassing the transition between boreal forest and prairie in western Canada. A central question is the significance of drought as an agent of large‐scale tree mortality and its potential future impact on carbon cycling in this cold region. We used a combination of plot‐based, meteorological, and remote sensing measures to map and quantify aboveground, dead biomass of trembling aspen (Populus tremuloides Michx.) across an 11.5 Mha survey area where drought was exceptionally severe during 2001–2002. Within this area, a satellite‐based land cover map showed that aspen‐dominated broadleaf forests occupied 2.3 Mha. Aerial surveys revealed extensive patches of severe mortality (>55%) resembling the impacts of fire. Dead aboveground biomass was estimated at 45 Mt, representing 20% of the total aboveground biomass, based on a spatial interpolation of plot‐based measurements. Spatial variation in percentage dead biomass showed a moderately strong correlation with drought severity. In the prairie‐like, southern half of the study area where the drought was most severe, 35% of aspen biomass was dead, compared with an estimated 7% dead biomass in the absence of drought. Drought led to an estimated 29 Mt increase in dead biomass across the survey area, corresponding to 14 Mt of potential future carbon emissions following decomposition. Many recent, comparable episodes of drought‐induced forest dieback have been reported from around the world, which points to an emerging need for multiscale monitoring approaches to quantify drought effects on woody biomass and carbon cycling across large areas.     

Role of mixed‐species flocks in the use of adjacent savannas by forest birds in the central Cerrado,Brazil

Abstract: We examined the role of mixed‐species flocks for forest birds during their breeding and non‐breeding seasons in the use of savannas adjacent to forests in central Cerrado, Brazil. Transect surveys (n = 64) were conducted in eight savanna patches. Distances of birds from forests were estimated. Recorded birds were classified as members or not of mixed‐species flocks. About half of the bird species recorded in savannas were found in at least one mixed‐species flock. As distance from the forest increased, the number of species in mixed‐species flocks tended not to vary, while the number of species foraging alone or in mono‐specific groups decreased. Thus, for some forest species, participation in mixed‐species flocks allowed a greater use of more distant savannas. This tendency of being in mixed‐species flocks at greater distances from forests also can be interpreted as a reluctance to forage alone or in mono‐specific groups due to higher predation risk in less protective vegetation distant from cover. There was strong seasonal variation in the participation of bird species in mixed‐species flocks. There were significantly more species in mixed‐species flocks than out of these associations in the non‐breeding season, while differences in the breeding season were not significant. These patterns occurred, in part because mixed‐species flocks tended to be more frequent, to have more species and to forage at greater distances from forests during the early non‐breeding season than in other periods. This study suggests that the formation of mixed‐species flocks plays an important role in promoting the use of adjacent savannas by forest birds at forest/savanna boundaries in Cerrado. It also pointed out a novel advantage gained by birds with participation in mixed‐species flocks – greater use of adjacent vegetation patches.     

Fire deficits have increased drought sensitivity in dry conifer forests: Fire frequency and tree‐ring carbon isotope evidence from Central Oregon

A century of fire suppression across the Western United States has led to more crowded forests and increased competition for resources. Studies of forest thinning or stand conditions after mortality events have provided indirect evidence for how competition can promote drought stress and predispose forests to severe fire and/or bark beetle outbreaks. Here, we demonstrate linkages between fire deficits and increasing drought stress through analyses of annually resolved tree‐ring growth, fire scars, and carbon isotope discrimination (Δ¹³C) across a dry mixed‐conifer forest landscape. Fire deficits across the study area have increased the sensitivity of leaf gas exchange to drought stress over the past >100 years. Since 1910, stand basal area in these forests has more than doubled and fire‐return intervals have increased from 25 to 140 years. Meanwhile, the portion of interannual variation in tree‐ring Δ¹³C explained by the Palmer Drought Severity Index has more than doubled in ca. 300–500‐year‐old Pinus ponderosa as well as in fire‐intolerant, ca. 90–190‐year‐old Abies grandis. Drought stress has increased in stands with a basal area of ≥25 m²/ha in 1910, as indicated by negative temporal Δ¹³C trends, whereas stands with basal area ≤25 m²/ha in 1910, due to frequent or intense wildfire activity in decades beforehand, were initially buffered from increased drought stress and have benefited more from rising ambient carbon dioxide concentrations, [CO₂], as demonstrated by positive temporal Δ¹³C trends. Furthermore, the average Δ¹³C response across all P. ponderosa since 1830 indicates that photosynthetic assimilation rates and stomatal conductance have been reduced by ~10% and ~20%, respectively, compared to expected trends due to increasing [CO₂]. Although disturbance legacies contribute to local‐scale intensity of drought stress, fire deficits have reduced drought resistance of mixed‐conifer forests and made them more susceptible to challenges by pests and pathogens and other disturbances.     

Fire-tolerance mechanisms of common woody plant species in a semiarid savanna in south-western Zimbabwe

The frequency of fire has increased in savannas yet few studies have assessed how plants persist when subjected to long‐term disturbance by fire. We investigated the contributions of bark thickness and resprouting to the persistence of woody plants in two fire trials that were started in 1948 and 1949. The number of resprouts per individual, bark thickness, basal diameter and height of woody plants were measured in unburnt plots and those burnt annually, triennially and quinquennially during the late dry season. Changes in tree density, number of resprouts and individuals in different height classes between 1963 and 2002 were assessed. Bark thickness varied among species and also increased with increases in basal diameter. Generally, plants with thick bark survived fire more than those with thin bark. Resprouting was the major fire survival strategy for most species. The number of resprouts produced per plant ranged from 4 ± 3 (Acacia rehmanniana) to 14 ± 9 (Pseudolachnostylis maprouneifolia). Fire reduced species richness in plots burnt annually and triennially by 47% and 6% respectively. Species richness increased in unburnt plots (5%) and those burnt quinquennially (16%). Most woody species survived fire through a combination of traits.     

Phylogenetic structure of Brazilian savannas under different fire regimes

Questions: Fire is a strong filter in fire‐prone communities and is expected to assemble closely related species when functional traits are conserved in plant lineages. Do frequent fires assemble savannas with closely related species (phylogenetic clustering)? If so, what are the clades pruned by fire in the phylogenetic trees? Are species of semi‐deciduous seasonal forests, where fires are not frequent, less related than expected by chance (phylogenetic over‐dispersion)? Are life forms conserved in the phylogeny of the species? Location: Central and SE Brazilian savannas (Emas National Park, 18°18′S, 52°54′W; Brasília, 15°56′–15°57′S, 47°53′–47°56′W and Corumbataí‐Itirapina, 22°13′–22°15′S, 47°37′–47°39′W); and close semi‐deciduous seasonal forests (in Pirenópolis, 15°45′S, 49°04′W; Brasília, 15°33′S, 47°51′W; and São Carlos, 21°55′S, 47°48′W). Methods: We recorded woody species in savannas under different fire regimes and in semi‐deciduous seasonal forests. We obtained data from the literature and from field sampling. We compared mean phylogenetic distance of species of savanna and of nearby semi‐deciduous seasonal forest sites. We obtained significance by randomizing the species among the tips of phylogenetic trees. We also assessed whether life forms were evolutionary conserved across phylogeny of the studied plants (phylogenetic signal) with tests based on the variance of phylogenetic independent contrasts. Results: Some sites of savanna under high fire frequency were characterized by phylogenetic over‐dispersion of woody species whereas, in contrast, some sites of semi‐deciduous seasonal forest were characterized by phylogenetic clustering. We found phylogenetic signals in the traits across the phylogeny of the 801 species investigated. Conclusion: Fire may have different roles in assembling plant species in Brazilian savannas than in other fire‐prone communities. We postulate that the absence of phylogenetic clustering in the cerrado is mainly due to the persistence of long‐lived resprouting species from different plant lineages.     

Effects of Fire on Seedling Diversity and Plant Reproduction (Sexual vs. Vegetative) in Neotropical Savannas Differing in Tree Density

Little is known about the effects of fire on the structure and species composition of Neotropical savanna seedling communities. Such effects are critical for predicting long‐term changes in plant distribution patterns in these ecosystems. We quantified richness and density of seedlings within 144 plots of 1 m² located along a topographic gradient in long‐unburned (fire protected since 1983) and recently burned (September 2005) savannas in Brazil. These savannas differ in tree density and canopy cover. Sites along the gradient, however, did not differ in species composition prior to the fire. In recently burned savannas we also evaluated the contribution of vegetative reproduction relative to sexual reproduction by quantifying richness and density of root suckers. Finally, we tested seed tolerance to pulses of high temperatures—similar to those occurring during fires on the soil surface and below—of five dominant savanna tree species. Seedlings were more abundant and diverse in unburned than in burned savannas. Seedling species composition differed among unburned and burned savannas probably reflecting early differences in root: shoot biomass allocation patterns. In recently burned savannas, root suckers were more abundant and diverse than seedlings. Relatively long exposures (>10 min) of temperatures of 90 °C reduced seed germination in all studied species suggesting a negative effect of fire on germination of seeds located at or aboveground level. Because vegetative reproduction contributes more than sexual reproduction in burned environments, frequent fires are likely to cause major shifts in species composition of Neotropical savanna plant communities, favoring clonally produced recruits along tree density/topographic gradients.     

When the same is not the same: phenotypic variation reveals different plant ecological strategies within species occurring in distinct Neotropical savanna habitats

Variations in abiotic characteristics such as soil water availability and fertility impose different selective pressures on plant populations. This may produce intraspecific variability in functional traits, even at a fine spatial scale. We investigated whether functional traits related to water-use efficiency, resource-retention strategy, soil nutrient acquisition, and fire tolerance differ in species that occur in two different habitats of Brazilian Cerrado: rocky savannas and savanna woodlands. Rocky savannas occur over sandstone, quartzite outcrops and have shallow nutrient-poor and low-moisture rocky soils, while savanna woodlands occur over well-drained and deep soils with frequent fire regimes. We measured nine functional traits of 40 tree species that occur in both habitats. Rocky savanna individuals exhibited a greater water-use efficiency strategy. The resource-retention strategy in rocky savanna individuals was corroborated by lower adult maximum height. However, despite the lower nutrient availability in rocky savanna soils, we only detected lower leaf phosphorus content in individuals from this habitat. Furthermore, individuals from both habitats had equally thick bark, suggesting that the fire-defense strategy is related to a stable, rather than plastic trait. Overall, our results highlight the central role of contrasting soil water availability patterns in driving phenotypic plasticity within species. We conclude that savanna species are responding to water and nutrient availabilities, via plasticity in traits related to the resource-retention strategy, and preparing for future fires, via uniformly thick bark. Wide plant distribution in contrasting habitats is possible for species that can shift ecological strategies to survive in nutrient- and water-limited habitats such as rocky savannas.     

Chronic water stress reduces tree growth and the carbon sink of deciduous hardwood forests

Predicted decreases in water availability across the temperate forest biome have the potential to offset gains in carbon (C) uptake from phenology trends, rising atmospheric CO₂, and nitrogen deposition. While it is well established that severe droughts reduce the C sink of forests by inducing tree mortality, the impacts of mild but chronic water stress on forest phenology and physiology are largely unknown. We quantified the C consequences of chronic water stress using a 13‐year record of tree growth (n = 200 trees), soil moisture, and ecosystem C balance at the Morgan–Monroe State Forest (MMSF) in Indiana, and a regional 11‐year record of tree growth (n > 300 000 trees) and water availability for the 20 most dominant deciduous broadleaf tree species across the eastern and midwestern USA. We show that despite ~26 more days of C assimilation by trees at the MMSF, increasing water stress decreased the number of days of wood production by ~42 days over the same period, reducing the annual accrual of C in woody biomass by 41%. Across the deciduous forest region, water stress induced similar declines in tree growth, particularly for water‐demanding ‘mesophytic’ tree species. Given the current replacement of water‐stress adapted ‘xerophytic’ tree species by mesophytic tree species, we estimate that chronic water stress has the potential to decrease the C sink of deciduous forests by up to 17% (0.04 Pg C yr^?1) in the coming decades. This reduction in the C sink due to mesophication and chronic water stress is equivalent to an additional 1–3 days of global C emissions from fossil fuel burning each year. Collectively, our results indicate that regional declines in water availability may offset the growth‐enhancing effects of other global changes and reduce the extent to which forests ameliorate climate warming.     

Precipitation gradients,plant biogeography,and the incidence of drip-tips in Cerrado plant species

Examining how both climate and species distribution patterns correlate with leaf morphology can give insights into the ecological and evolutionary patterns that drive adaptive selection of leaf form and function. Drip-tips are a common feature of leaves in rain forest tree species; they are thought to be an adaptation that aids leaf drying and maximizes photosynthesis in areas with high-rainfall climates. We tested whether this macroecological pattern holds true across the precipitation gradients in a non-rain forest region—the woodland savannas of Brazil known as the Cerrado—and compared our results with previous studies from Amazonia. Drip-tips were, as expected, less common overall in the drier Cerrado than in Amazonia. In addition, within the Cerrado, drip-tips were more prevalent in areas with higher rainfall as well as in Cerrado sites that were closer to Amazonia. Moreover, species that occurred across both the Cerrado and Amazonia had drip-tips more often than species that were found only in the Cerrado. These findings support the hypothesis that drip-tips are adaptive and that either the cost of retaining drip-tips is low or that in drier regions they have other benefits.