A trillion trees: carbon capture or fuelling fires?

Afforesting grassy systems for carbon gain using flammable plantation trees could shift the fire regime from lower intensity grass-fuelled fires to high-intensity crown fires. Future changes in climate will worsen this. We highlight the fire risk of trees planted for carbon and costs of fire protection using African examples.

Trees, carbon, and the psychology of landscapes.

Mitigating climate change while safeguarding biodiversity and livelihoods is a major challenge. However, rampant afforestation threatens biodiversity and livelihoods, with questionable benefits to carbon storage. The narrative of landscape degradation is often applied without considering the history of the landscape. While some landscapes are undoubtedly deforested, others existed in open or mosaic states before human intervention, or have been deliberately maintained as such. In psychology, a 'fundamental attribution error' is made when characteristics are attributed without consideration of context or circumstances. We apply this concept to landscapes, and then propose a process that avoids attribution errors by testing a null hypothesis regarding past forest extent, using palaeoecology and other long-term data, alongside ecological and stakeholder knowledge.

Quantifying the environmental limits to fire spread in grassy ecosystems.

Modeling fire spread as an infection process is intuitive: An ignition lights a patch of fuel, which infects its neighbor, and so on. Infection models produce nonlinear thresholds, whereby fire spreads only when fuel connectivity and infection probability are sufficiently high. These thresholds are fundamental both to managing fire and to theoretical models of fire spread, whereas applied fire models more often apply quasi-empirical approaches. Here, we resolve this tension by quantifying thresholds in fire spread locally, using field data from individual fires (n = 1,131) in grassy ecosystems across a precipitation gradient (496 to 1,442 mm mean annual precipitation) and evaluating how these scaled regionally (across 533 sites) and across time (1989 to 2012 and 2016 to 2018) using data from Kruger National Park in South Africa. An infection model captured observed patterns in individual fire spread better than competing models. The proportion of the landscape that burned was well described by measurements of grass biomass, fuel moisture, and vapor pressure deficit. Regionally, averaging across variability resulted in quasi-linear patterns. Altogether, results suggest that models aiming to capture fire responses to global change should incorporate nonlinear fire spread thresholds but that linear approximations may sufficiently capture medium-term trends under a stationary climate.

Feedbacks in ecology and evolution.

Ecology and evolutionary biology have focused on how organisms fit the environment. Less attention has been given to the idea that organisms can also modify their environment, and that these modifications can feed back to the organism, thus providing a key factor for their persistence and evolution. There are at least three independent lines of evidence emphasizing these biological feedback processes at different scales: niche construction (population scale); alternative biome states (community scale); and the Gaia hypothesis (planetary scale). These feedback processes make us rethink traditional concepts like niche and adaptation. We argue that organism-environment feedbacks must become a regular part of ecological thinking, especially now that the Earth is quickly changing.

Biome boundary maintained by intense belowground resource competition in world's thinnest-rooted plant community.

Recent findings point to plant root traits as potentially important for shaping the boundaries of biomes and for maintaining the plant communities within. We examined two hypotheses: 1) Thin-rooted plant strategies might be favored in biomes with low soil resources; and 2) these strategies may act, along with fire, to maintain the sharp boundary between the Fynbos and Afrotemperate Forest biomes in South Africa. These biomes differ in biodiversity, plant traits, and physiognomy, yet exist as alternative stable states on the same geological substrate and in the same climate conditions. We conducted a 4-y field experiment to examine the ability of Forest species to invade the Fynbos as a function of growth-limiting nutrients and belowground plant-plant competition. Our results support both hypotheses: First, we found marked biome differences in root traits, with Fynbos species exhibiting the thinnest roots reported from any biome worldwide. Second, our field manipulation demonstrated that intense belowground competition inhibits the ability of Forest species to invade Fynbos. Nitrogen was unexpectedly the resource that determined competitive outcome, despite the long-standing expectation that Fynbos is severely phosphorus constrained. These findings identify a trait-by-resource feedback mechanism, in which most species possess adaptive traits that modify soil resources in favor of their own survival while deterring invading species. Our findings challenge the long-held notion that biome boundaries depend primarily on external abiotic constraints and, instead, identify an internal biotic mechanism-a selective feedback among traits, plant-plant competition, and ecosystem conditions-that, along with contrasting fire regime, can act to maintain biome boundaries.

Savanna tree evolutionary ages inform the reconstruction of the paleoenvironment of our hominin ancestors.

Ideas on hominin evolution have long invoked the emergence from forests into open habitats as generating selection for traits such as bipedalism and dietary shifts. Though controversial, the savanna hypothesis continues to motivate research into the palaeo-environments of Africa. Reconstruction of these ancient environments has depended heavily on carbon isotopic analysis of fossil bones and palaeosols. The sparsity of the fossil record, however, imposes a limit to the strength of inference that can be drawn from such data. Time-calibrated phylogenies offer an additional tool for dating the spread of savanna habitat. Here, using the evolutionary ages of African savanna trees, we suggest an initial tropical or subtropical expansion of savanna between 10 and 15 Ma, which then extended to higher latitudes, reaching southern Africa ca. 3 Ma. Our phylogenetic estimates of the origin and latitudinal spread of savannas broadly correspond with isotopic age estimates and encompass the entire hominin fossil record. Our results are consistent with the savanna hypothesis of early hominin evolution and reignite the debate on the drivers of savanna expansion. Our analysis demonstrates the utility of phylogenetic proxies for dating major ecological transitions in geological time, especially in regions where fossils are rare or absent or occur in discontinuous sediments.

On the Three Major Recycling Pathways in Terrestrial Ecosystems.

Plants are the largest biomass component of most terrestrial ecosystems, and litter decomposition is considered the dominant process by which nutrients return to plants. We show that in terrestrial ecosystems, there are three major pathways by which plant biomass is degraded into forms that release nutrients again available to plants: microbial decomposition; vertebrate herbivory; and wildfires. These processes act at different spatial and temporal scales, have different niches, and generates different ecological and evolutionary feedbacks. This holistic view in which microbes, herbivores, and wildfires play a joint role in the functioning of ecosystems contributes to a better understanding of the diversity of mechanisms regulating the biosphere.

Mismatches between demographic niches and geographic distributions are strongest in poorly dispersed and highly persistent plant species.

The ecological niche of a species describes the variation in population growth rates along environmental gradients that drives geographic range dynamics. Niches are thus central for understanding and forecasting species' geographic distributions. However, theory predicts that migration limitation, source-sink dynamics, and time-lagged local extinction can cause mismatches between niches and geographic distributions. It is still unclear how relevant these niche-distribution mismatches are for biodiversity dynamics and how they depend on species life-history traits. This is mainly due to a lack of the comprehensive, range-wide demographic data needed to directly infer ecological niches for multiple species. Here we quantify niches from extensive demographic measurements along environmental gradients across the geographic ranges of 26 plant species (Proteaceae; South Africa). We then test whether life history explains variation in species' niches and niche-distribution mismatches. Niches are generally wider for species with high seed dispersal or persistence abilities. Life-history traits also explain the considerable interspecific variation in niche-distribution mismatches: poorer dispersers are absent from larger parts of their potential geographic ranges, whereas species with higher persistence ability more frequently occupy environments outside their ecological niche. Our study thus identifies major demographic and functional determinants of species' niches and geographic distributions. It highlights that the inference of ecological niches from geographical distributions is most problematic for poorly dispersed and highly persistent species. We conclude that the direct quantification of ecological niches from demographic responses to environmental variation is a crucial step toward a better predictive understanding of biodiversity dynamics under environmental change.

Alternative Biome States in Terrestrial Ecosystems.

There is growing interest in the application of alternative stable state (ASS) theory to explain major vegetation patterns of the world. Here, we introduce the theory as applied to the puzzle of nonforested (open) biomes growing in climates that are warm and wet enough to support forests (alternative biome states, ABSs). Long thought to be the product of deforestation, diverse lines of evidence indicate that many open ecosystems are ancient. They have also been characterized as 'early successional' even where they persist for millennia. ABS is an alternative framework to that of climate determinism and succession for exploring forest/nonforest mosaics. This framework explains not only tropical forest-savanna landscapes, but also other landscape mosaics across the globe.

Lineage-based functional types: characterising functional diversity to enhance the representation of ecological behaviour in Land Surface Models.

Process-based vegetation models attempt to represent the wide range of trait variation in biomes by grouping ecologically similar species into plant functional types (PFTs). This approach has been successful in representing many aspects of plant physiology and biophysics but struggles to capture biogeographic history and ecological dynamics that determine biome boundaries and plant distributions. Grass-dominated ecosystems are broadly distributed across all vegetated continents and harbour large functional diversity, yet most Land Surface Models (LSMs) summarise grasses into two generic PFTs based primarily on differences between temperate C3 grasses and (sub)tropical C4 grasses. Incorporation of species-level trait variation is an active area of research to enhance the ecological realism of PFTs, which form the basis for vegetation processes and dynamics in LSMs. Using reported measurements, we developed grass functional trait values (physiological, structural, biochemical, anatomical, phenological, and disturbance-related) of dominant lineages to improve LSM representations. Our method is fundamentally different from previous efforts, as it uses phylogenetic relatedness to create lineage-based functional types (LFTs), situated between species-level trait data and PFT-level abstractions, thus providing a realistic representation of functional diversity and opening the door to the development of new vegetation models.

Comment on "The global tree restoration potential".

Bastin et al's estimate (Reports, 5 July 2019, p. 76) that tree planting for climate change mitigation could sequester 205 gigatonnes of carbon is approximately five times too large. Their analysis inflated soil organic carbon gains, failed to safeguard against warming from trees at high latitudes and elevations, and considered afforestation of savannas, grasslands, and shrublands to be restoration.

The Trouble with Trees: Afforestation Plans for Africa.

Extensive tree planting is widely promoted for reducing atmospheric CO2. In Africa, 1 million km2, mostly of grassy biomes, have been targeted for 'restoration' by 2030. The target is based on the erroneous assumption that these biomes are deforested and degraded. We discuss the pros and cons of exporting fossil fuel emission problems to Africa.

Does a tradeoff between trait plasticity and resource conservatism contribute to the maintenance of alternative stable states?

Phenotypic plasticity facilitates species persistence across resource gradients but may be limited in low-resource environments requiring resource conservation. We investigated the tradeoff between trait plasticity and resource conservatism across a biome boundary characterized by high turnover in nutrient and light availability, and whether this contributes to the maintenance of alternative stable states. Differences in plasticity were determined by comparing species' leaf and foliar nutritional trait responses to light, represented by leaf area index (LAI), and soil nutrient availability across forest-shrubland boundaries in South Africa. Although forest had higher LAI and soil nutrient availability than shrubland, forest species experienced greater resource variation. With increasing LAI and nutrient availability, forest species increased their leaf size, specific leaf area and leaf area/stem length, and decreased their foliar [N] and [K]. Although these responses are indicative of plasticity, shrubland species appeared to lack plasticity as evidenced by limited trait variation with environmental heterogeneity. Inhabiting diverse light environments imposed by forests probably selects for plasticity, whereas light-saturated, fire-prone, nutrient-poor environments that select for conservative leaf traits and below-ground investments compromise plasticity in shrubland species. This pattern suggests a tradeoff between trait plasticity and resource conservatism, which may support the stability of alternative vegetation states.

Resilience and restoration of tropical and subtropical grasslands, savannas, and grassy woodlands.

Despite growing recognition of the conservation values of grassy biomes, our understanding of how to maintain and restore biodiverse tropical grasslands (including savannas and open-canopy grassy woodlands) remains limited. To incorporate grasslands into large-scale restoration efforts, we synthesised existing ecological knowledge of tropical grassland resilience and approaches to plant community restoration. Tropical grassland plant communities are resilient to, and often dependent on, the endogenous disturbances with which they evolved - frequent fires and native megafaunal herbivory. In stark contrast, tropical grasslands are extremely vulnerable to human-caused exogenous disturbances, particularly those that alter soils and destroy belowground biomass (e.g. tillage agriculture, surface mining); tropical grassland restoration after severe soil disturbances is expensive and rarely achieves management targets. Where grasslands have been degraded by altered disturbance regimes (e.g. fire exclusion), exotic plant invasions, or afforestation, restoration efforts can recreate vegetation structure (i.e. historical tree density and herbaceous ground cover), but species-diverse plant communities, including endemic species, are slow to recover. Complicating plant-community restoration efforts, many tropical grassland species, particularly those that invest in underground storage organs, are difficult to propagate and re-establish. To guide restoration decisions, we draw on the old-growth grassland concept, the novel ecosystem concept, and theory regarding tree cover along resource gradients in savannas to propose a conceptual framework that classifies tropical grasslands into three broad ecosystem states. These states are: (1) old-growth grasslands (i.e. ancient, biodiverse grassy ecosystems), where management should focus on the maintenance of disturbance regimes; (2) hybrid grasslands, where restoration should emphasise a return towards the old-growth state; and (3) novel ecosystems, where the magnitude of environmental change (i.e. a shift to an alternative ecosystem state) or the socioecological context preclude a return to historical conditions.

Human impacts in African savannas are mediated by plant functional traits.

Tropical savannas have a ground cover dominated by C4 grasses, with fire and herbivory constraining woody cover below a rainfall-based potential. The savanna biome covers 50% of the African continent, encompassing diverse ecosystems that include densely wooded Miombo woodlands and Serengeti grasslands with scattered trees. African savannas provide water, grazing and browsing, food and fuel for tens of millions of people, and have a unique biodiversity that supports wildlife tourism. However, human impacts are causing widespread and accelerating degradation of savannas. The primary threats are land cover-change and transformation, landscape fragmentation that disrupts herbivore communities and fire regimes, climate change and rising atmospheric CO2 . The interactions among these threats are poorly understood, with unknown consequences for ecosystem health and human livelihoods. We argue that the unique combinations of plant functional traits characterizing the major floristic assemblages of African savannas make them differentially susceptible and resilient to anthropogenic drivers of ecosystem change. Research must address how this functional diversity among African savannas differentially influences their vulnerability to global change and elucidate the mechanisms responsible. This knowledge will permit appropriate management strategies to be developed to maintain ecosystem integrity, biodiversity and livelihoods.

Steal the light: shade vs fire adapted vegetation in forest-savanna mosaics.

Shade cast by trees, which suppresses grass growth, and fire fuelled by grass biomass, which prevents tree sapling establishment, are mutually exclusive and self-reinforcing drivers of biome distribution in savanna-forest mosaics. We investigated how shade depth, represented by canopy leaf area index (LAI), is generated by adult trees across savanna-forest boundaries and how a shade gradient filters tree functioning, and grass composition and biomass. Forest trees exerted greater shading through increased stem density and greater light interception per unit biomass. A critical transition at LAI c. 1.5 was linked to tree shifts from savanna to forest species, functional shifts from fire-tolerant to light-competitive species, and grass composition shifts from C4 to C3 pathways. A second transition to grass fuel loads too low to support fires, occurred at a lower canopy density (LAI > 0.5), accompanied by shifts in C4 subtype dominance. This pattern suggests that shade suppression of grass biomass is an essential first step for the maintenance of alternative stable states.