Assessing protected area vulnerability to climate change in a case study of South African national parks.

Climate change is challenging the ability of protected areas (PAs) to meet their objectives. To improve PA planning, we developed a framework for assessing PA vulnerability to climate change based on consideration of potential climate change impacts on species and their habitats and resource use. Furthermore, the capacity of PAs to adapt to these climate threats was determined through assessment of PA management effectiveness, adjacent land use, and financial resilience. Users reach a PA-specific vulnerability score and rank based on scoring of these categories. We applied the framework to South Africa's 19 national parks. Because the 19 parks are managed as a national network, we explored how resources might be best allocated to address climate change. Each park's importance to the network's biodiversity conservation and revenue generation was estimated and used to weight overall vulnerability scores and ranks. Park vulnerability profiles showed distinct combinations of potential impacts of climate change and adaptive capacities; the former had a greater influence on vulnerability. Mapungubwe National Park emerged as the most vulnerable to climate change, despite its relatively high adaptive capacity, largely owing to large projected changes in species and resource use. Table Mountain National Park scored the lowest in overall vulnerability. Climate change vulnerability rankings differed markedly once importance weightings were applied; Kruger National Park was the most vulnerable under both importance scenarios. Climate change vulnerability assessment is fundamental to effective adaptation planning. Our PA assessment tool is the only tool that quantifies PA vulnerability to climate change in a comparative index. It may be used in data-rich and data-poor contexts to prioritize resource allocation across PA networks and can be applied from local to global scales.

Resumen El cambio climático es un gran obstáculo para que las áreas protegidas (AP) logren sus objetivos. Para mejorar la planeación de las AP, desarrollamos un marco de trabajo para evaluar la vulnerabilidad de estas ante el cambio climático con base en la consideración de los impactos potenciales del cambio climático sobre las especies, sus hábitats y los recursos que usan. Además, determinamos la capacidad de las AP para adaptarse a estas amenazas climáticas mediante la valoración de las categorías efectividad de la gestión de las AP, las tierras adyacentes y la resiliencia económica. Los usuarios logran un puntaje y clasificación de vulnerabilidad específicas de la AP con base en las calificaciones de estas categorías. Aplicamos el marco de trabajo a los 19 parques nacionales de Sudáfrica. Ya que todos los parques se manejan como una red nacional, exploramos cómo pueden asignarse de mejor manera los recursos para lidiar con el cambio climático. Se estimaron la importancia de cada parque para la conservación de la biodiversidad de la red y la generación de ganancias. Después usamos las estimaciones para sopesar los puntajes y las clasificaciones generales de vulnerabilidad. Los perfiles de vulnerabilidad de los parques mostraron combinaciones distintivas de impactos potenciales del cambio climático y capacidades de adaptación; los impactos tuvieron una mayor influencia sobre la vulnerabilidad. El Parque Nacional Mapungubwe se ubicó como el más vulnerable ante el cambio climático, a pesar de tener una capacidad de adaptación relativamente alta, principalmente debida a grandes cambios proyectados para el uso de recursos y especies. El Parque Nacional Table Mountain tuvo el puntaje más bajo de vulnerabilidad generalizada. Las clasificaciones de vulnerabilidad al cambio climático difirieron notablemente una vez que se aplicaron los factores de importancia; el Parque Nacional Kruger fue el más vulnerable bajo ambos escenarios de importancia. La evaluación de vulnerabilidad al cambio climático es fundamental para la planeación efectiva de la adaptación. Nuestra herramienta de valoración de las AP es la única que cuantifica la vulnerabilidad de las AP al cambio climático en un índice comparativo. Puede usarse en contextos con muchos o pocos datos para priorizar la asignación de recursos en las redes de AP y puede aplicarse desde la escala local hasta la mundial.

Combined impacts of future climate-driven vegetation changes and socioeconomic pressures on protected areas in Africa.

Africa's protected areas (PAs) are the last stronghold of the continent's unique biodiversity, but they appear increasingly threatened by climate change, substantial human population growth, and land-use change. Conservation planning is challenged by uncertainty about how strongly and where these drivers will interact over the next few decades. We investigated the combined future impacts of climate-driven vegetation changes inside African PAs and human population densities and land use in their surroundings for 2 scenarios until the end of the 21st century. We used the following 2 combinations of the shared socioeconomic pathways (SSPs) and representative greenhouse gas concentration pathways (RCPs): the "middle-of-the-road" scenario SSP2-RCP4.5 and the resource-intensive "fossil-fueled development" scenario SSP5-RCP8.5. Climate change impacts on tree cover and biome type (i.e., desert, grassland, savanna, and forest) were simulated with the adaptive dynamic global vegetation model (aDGVM). Under both scenarios, most PAs were adversely affected by at least 1 of the drivers, but the co-occurrence of drivers was largely region and scenario specific. The aDGVM projections suggest considerable climate-driven tree cover increases in PAs in today's grasslands and savannas. For PAs in West Africa, the analyses revealed climate-driven vegetation changes combined with hotspots of high future population and land-use pressure. Except for many PAs in North Africa, future decreases in population and land-use pressures were rare. At the continental scale, SSP5-RCP8.5 led to higher climate-driven changes in tree cover and higher land-use pressure, whereas SSP2-RCP4.5 was characterized by higher future population pressure. Both SSP-RCP scenarios implied increasing challenges for conserving Africa's biodiversity in PAs. Our findings underline the importance of developing and implementing region-specific conservation responses. Strong mitigation of future climate change and equitable development scenarios would reduce ecosystem impacts and sustain the effectiveness of conservation in Africa.

Las áreas protegidas (AP) de África son el último bastión de la biodiversidad distintiva del continente, pero cada vez están más amenazadas por el cambio climático, crecimiento sustancial de la población humana y cambio de uso de suelo. La planificación de la conservación enfrenta el reto de la incertidumbre de cuan fuerte y donde interactuarán estos factores a lo largo de las siguientes décadas. Investigamos los impactos futuros combinados de los cambios en la vegetación impulsados por el clima dentro de AP africanas y las densidades de población humana y el uso de suelo en sus alrededores en 2 escenarios hasta el final del siglo 21. Utilizamos las siguientes 2 combinaciones de las trayectorias socioeconómicas compartidas (SSP) y las trayectorias representativas de concentración de gases de invernadero (RCP): el escenario de "mitad del camino" SSP2-RCP4.5 y el escenario recurso intensivo "desarrollo impulsado por combustibles fósiles" SSP5-RCP8.5. Los impactos del cambio climático sobre la cobertura de árboles y el tipo de bioma (i. e., desierto, pastizal, sabana y bosque) fueron simulados con el modelo vegetación global dinámica adaptativo (aDGVM). En ambos escenarios, la mayoría de las AP fueron afectadas adversamente por lo menos por 1 de los factores, pero la coocurrencia de los factores fue mayoritariamente específica por región y escenario. Las proyecciones de MVGDa sugieren incrementos considerables en la cobertura de árboles impulsados por el clima en las AP en pastizales y sabanas actuales. Para AP en África Occidental, los análisis revelaron cambios en la vegetación impulsados por el clima combinados con sitios clave con numerosa población y gran presión de uso de suelo en el futuro. Excepto en muchos PA de África del Norte, los decrementos en la población y presiones de uso de suelo en el futuro fueron raros. A escala continental, SSP5-RCP8.5 condujo a mayores cambios impulsados por el clima en la cobertura arbórea y en la presión de cambio de uso de suelo, mientras que SSP5-RCP8.5 se caracterizó por una mayor presión demográfica en el futuro. Ambos escenarios SSP-RCP implicaron mayores retos para la conservación de la biodiversidad en AP africanas. Nuestros hallazgos subrayan la importancia de desarrollar e implementar respuestas de conservación específicas para cada región. Medidas sólidas para la mitigación del cambio climático así como escenarios de desarrollo equitativo podrían reducir los impactos en el ecosistema y sustentar la efectividad de la conservación en África.

Large uncertainties in future biome changes in Africa call for flexible climate adaptation strategies.

Anthropogenic climate change is expected to impact ecosystem structure, biodiversity and ecosystem services in Africa profoundly. We used the adaptive Dynamic Global Vegetation Model (aDGVM), which was originally developed and tested for Africa, to quantify sources of uncertainties in simulated African potential natural vegetation towards the end of the 21st century. We forced the aDGVM with regionally downscaled high-resolution climate scenarios based on an ensemble of six general circulation models (GCMs) under two representative concentration pathways (RCPs 4.5 and 8.5). Our study assessed the direct effects of climate change and elevated CO2 on vegetation change and its plant-physiological drivers. Total increase in carbon in aboveground biomass in Africa until the end of the century was between 18% to 43% (RCP4.5) and 37% to 61% (RCP8.5) and was associated with woody encroachment into grasslands and increased woody cover in savannas. When direct effects of CO2 on plants were omitted, woody encroachment was muted and carbon in aboveground vegetation changed between -8 to 11% (RCP 4.5) and -22 to -6% (RCP8.5). Simulated biome changes lacked consistent large-scale geographical patterns of change across scenarios. In Ethiopia and the Sahara/Sahel transition zone, the biome changes forecast by the aDGVM were consistent across GCMs and RCPs. Direct effects from elevated CO2 were associated with substantial increases in water use efficiency, primarily driven by photosynthesis enhancement, which may relieve soil moisture limitations to plant productivity. At the ecosystem level, interactions between fire and woody plant demography further promoted woody encroachment. We conclude that substantial future biome changes due to climate and CO2 changes are likely across Africa. Because of the large uncertainties in future projections, adaptation strategies must be highly flexible. Focused research on CO2 effects, and improved model representations of these effects will be necessary to reduce these uncertainties.

Climate change promotes transitions to tall evergreen vegetation in tropical Asia.

Vegetation in tropical Asia is highly diverse due to large environmental gradients and heterogeneity of landscapes. This biodiversity is threatened by intense land use and climate change. However, despite the rich biodiversity and the dense human population, tropical Asia is often underrepresented in global biodiversity assessments. Understanding how climate change influences the remaining areas of natural vegetation is therefore highly important for conservation planning. Here, we used the adaptive Dynamic Global Vegetation Model version 2 (aDGVM2) to simulate impacts of climate change and elevated CO2 on vegetation formations in tropical Asia for an ensemble of climate change scenarios. We used climate forcing from five different climate models for representative concentration pathways RCP4.5 and RCP8.5. We found that vegetation in tropical Asia will remain a carbon sink until 2099, and that vegetation biomass increases of up to 28% by 2099 are associated with transitions from small to tall woody vegetation and from deciduous to evergreen vegetation. Patterns of phenology were less responsive to climate change and elevated CO2 than biomes and biomass, indicating that the selection of variables and methods used to detect vegetation changes is crucial. Model simulations revealed substantial variation within the ensemble, both in biomass increases and in distributions of different biome types. Our results have important implications for management policy, because they suggest that large ensembles of climate models and scenarios are required to assess a wide range of potential future trajectories of vegetation change and to develop robust management plans. Furthermore, our results highlight open ecosystems with low tree cover as most threatened by climate change, indicating potential conflicts of interest between biodiversity conservation in open ecosystems and active afforestation to enhance carbon sequestration.

Global ecosystems and fire: Multi-model assessment of fire-induced tree-cover and carbon storage reduction.

In this study, we use simulations from seven global vegetation models to provide the first multi-model estimate of fire impacts on global tree cover and the carbon cycle under current climate and anthropogenic land use conditions, averaged for the years 2001-2012. Fire globally reduces the tree covered area and vegetation carbon storage by 10%. Regionally, the effects are much stronger, up to 20% for certain latitudinal bands, and 17% in savanna regions. Global fire effects on total carbon storage and carbon turnover times are lower with the effect on gross primary productivity (GPP) close to 0. We find the strongest impacts of fire in savanna regions. Climatic conditions in regions with the highest burned area differ from regions with highest absolute fire impact, which are characterized by higher precipitation. Our estimates of fire-induced vegetation change are lower than previous studies. We attribute these differences to different definitions of vegetation change and effects of anthropogenic land use, which were not considered in previous studies and decreases the impact of fire on tree cover. Accounting for fires significantly improves the spatial patterns of simulated tree cover, which demonstrates the need to represent fire in dynamic vegetation models. Based upon comparisons between models and observations, process understanding and representation in models, we assess a higher confidence in the fire impact on tree cover and vegetation carbon compared to GPP, total carbon storage and turnover times. We have higher confidence in the spatial patterns compared to the global totals of the simulated fire impact. As we used an ensemble of state-of-the-art fire models, including effects of land use and the ensemble median or mean compares better to observational datasets than any individual model, we consider the here presented results to be the current best estimate of global fire effects on ecosystems.

The future distribution of the savannah biome: model-based and biogeographic contingency.

The extent of the savannah biome is expected to be profoundly altered by climatic change and increasing atmospheric CO2 concentrations. Contrasting projections are given when using different modelling approaches to estimate future distributions. Furthermore, biogeographic variation within savannahs in plant function and structure is expected to lead to divergent responses to global change. Hence the use of a single model with a single savannah tree type will likely lead to biased projections. Here we compare and contrast projections of South American, African and Australian savannah distributions from the physiologically based Thornley transport resistance statistical distribution model (TTR-SDM)-and three versions of a dynamic vegetation model (DVM) designed and parametrized separately for specific continents. We show that attempting to extrapolate any continent-specific model globally biases projections. By 2070, all DVMs generally project a decrease in the extent of savannahs at their boundary with forests, whereas the TTR-SDM projects a decrease in savannahs at their boundary with aridlands and grasslands. This difference is driven by forest and woodland expansion in response to rising atmospheric CO2 concentrations in DVMs, unaccounted for by the TTR-SDM. We suggest that the most suitable models of the savannah biome for future development are individual-based dynamic vegetation models designed for specific biogeographic regions.This article is part of the themed issue 'Tropical grassy biomes: linking ecology, human use and conservation'.

Climate change and long-term fire management impacts on Australian savannas.

Tropical savannas cover a large proportion of the Earth's land surface and many people are dependent on the ecosystem services that savannas supply. Their sustainable management is crucial. Owing to the complexity of savanna vegetation dynamics, climate change and land use impacts on savannas are highly uncertain. We used a dynamic vegetation model, the adaptive dynamic global vegetation model (aDGVM), to project how climate change and fire management might influence future vegetation in northern Australian savannas. Under future climate conditions, vegetation can store more carbon than under ambient conditions. Changes in rainfall seasonality influence future carbon storage but do not turn vegetation into a carbon source, suggesting that CO2 fertilization is the main driver of vegetation change. The application of prescribed fires with varying return intervals and burning season influences vegetation and fire impacts. Carbon sequestration is maximized with early dry season fires and long fire return intervals, while grass productivity is maximized with late dry season fires and intermediate fire return intervals. The study has implications for management policy across Australian savannas because it identifies how fire management strategies may influence grazing yield, carbon sequestration and greenhouse gas emissions. This knowledge is crucial to maintaining important ecosystem services of Australian savannas.

Fire in Australian savannas: from leaf to landscape.

Savanna ecosystems comprise 22% of the global terrestrial surface and 25% of Australia (almost 1.9 million km2) and provide significant ecosystem services through carbon and water cycles and the maintenance of biodiversity. The current structure, composition and distribution of Australian savannas have coevolved with fire, yet remain driven by the dynamic constraints of their bioclimatic niche. Fire in Australian savannas influences both the biophysical and biogeochemical processes at multiple scales from leaf to landscape. Here, we present the latest emission estimates from Australian savanna biomass burning and their contribution to global greenhouse gas budgets. We then review our understanding of the impacts of fire on ecosystem function and local surface water and heat balances, which in turn influence regional climate. We show how savanna fires are coupled to the global climate through the carbon cycle and fire regimes. We present new research that climate change is likely to alter the structure and function of savannas through shifts in moisture availability and increases in atmospheric carbon dioxide, in turn altering fire regimes with further feedbacks to climate. We explore opportunities to reduce net greenhouse gas emissions from savanna ecosystems through changes in savanna fire management.

Increasing atmospheric CO2 overrides the historical legacy of multiple stable biome states in Africa.

The dominant vegetation over much of the global land surface is not predetermined by contemporary climate, but also influenced by past environmental conditions. This confounds attempts to predict current and future biome distributions, because even a perfect model would project multiple possible biomes without knowledge of the historical vegetation state. Here we compare the distribution of tree- and grass-dominated biomes across Africa simulated using a dynamic global vegetation model (DGVM). We explicitly evaluate where and under what conditions multiple stable biome states are possible for current and projected future climates. Our simulation results show that multiple stable biomes states are possible for vast areas of tropical and subtropical Africa under current conditions. Widespread loss of the potential for multiple stable biomes states is projected in the 21st Century, driven by increasing atmospheric CO2 . Many sites where currently both tree-dominated and grass-dominated biomes are possible become deterministically tree-dominated. Regions with multiple stable biome states are widespread and require consideration when attempting to predict future vegetation changes. Testing for behaviour characteristic of systems with multiple stable equilibria, such as hysteresis and dependence on historical conditions, and the resulting uncertainty in simulated vegetation, will lead to improved projections of global change impacts.

Intermediate coupling between aboveground and belowground biomass maximises the persistence of grasslands.

Aboveground and belowground biomass compartments of vegetation fulfil different functions and they are coupled by complex interactions. These compartments exchange water, carbon and nutrients and the belowground biomass compartment has the capacity to buffer vegetation dynamics when aboveground biomass is removed by disturbances such as herbivory or fire. However, despite their importance, root-shoot interactions are often ignored in more heuristic vegetation models. Here, we present a simple two-compartment grassland model that couples aboveground and belowground biomass. In this model, the growth of belowground biomass is influenced by aboveground biomass and the growth of aboveground biomass is influenced by belowground biomass. We used the model to explore how the dynamics of a grassland ecosystem are influenced by fire and grazing. We show that the grassland system is most persistent at intermediate levels of aboveground-belowground coupling. In this situation, the system can sustain more extreme fire or grazing regimes than in the case of strong coupling. In contrast, the productivity of the system is maximised at high levels of coupling. Our analysis suggests that the yield of a grassland ecosystem is maximised when coupling is strong, however, the intensity of disturbance that can be sustained increases dramatically when coupling is intermediate. Hence, the model predicts that intermediate coupling should be selected for as it maximises the chances of persistence in disturbance driven ecosystems.

Next-generation dynamic global vegetation models: learning from community ecology.

Dynamic global vegetation models (DGVMs) are powerful tools to project past, current and future vegetation patterns and associated biogeochemical cycles. However, most models are limited by how they define vegetation and by their simplistic representation of competition. We discuss how concepts from community assembly theory and coexistence theory can help to improve vegetation models. We further present a trait- and individual-based vegetation model (aDGVM2) that allows individual plants to adopt a unique combination of trait values. These traits define how individual plants grow and compete. A genetic optimization algorithm is used to simulate trait inheritance and reproductive isolation between individuals. These model properties allow the assembly of plant communities that are adapted to a site's biotic and abiotic conditions. The aDGVM2 simulates how environmental conditions influence the trait spectra of plant communities; that fire selects for traits that enhance fire protection and reduces trait diversity; and the emergence of life-history strategies that are suggestive of colonization-competition trade-offs. The aDGVM2 deals with functional diversity and competition fundamentally differently from current DGVMs. This approach may yield novel insights as to how vegetation may respond to climate change and we believe it could foster collaborations between functional plant biologists and vegetation modellers.

Atmospheric CO2 forces abrupt vegetation shifts locally, but not globally.

It is possible that anthropogenic climate change will drive the Earth system into a qualitatively different state. Although different types of uncertainty limit our capacity to assess this risk, Earth system scientists are particularly concerned about tipping elements, large-scale components of the Earth system that can be switched into qualitatively different states by small perturbations. Despite growing evidence that tipping elements exist in the climate system, whether large-scale vegetation systems can tip into alternative states is poorly understood. Here we show that tropical grassland, savanna and forest ecosystems, areas large enough to have powerful impacts on the Earth system, are likely to shift to alternative states. Specifically, we show that increasing atmospheric CO2 concentration will force transitions to vegetation states characterized by higher biomass and/or woody-plant dominance. The timing of these critical transitions varies as a result of between-site variance in the rate of temperature increase, as well as a dependence on stochastic variation in fire severity and rainfall. We further show that the locations of bistable vegetation zones (zones where alternative vegetation states can exist) will shift as climate changes. We conclude that even though large-scale directional regime shifts in terrestrial ecosystems are likely, asynchrony in the timing of these shifts may serve to dampen, but not nullify, the shock that these changes may represent to the Earth system.

Fire and fire-adapted vegetation promoted C4 expansion in the late Miocene.

Large proportions of the Earth's land surface are covered by biomes dominated by C(4) grasses. These C(4)-dominated biomes originated during the late Miocene, 3-8 million years ago (Ma), but there is evidence that C(4) grasses evolved some 20 Ma earlier during the early Miocene/Oligocene. Explanations for this lag between evolution and expansion invoke changes in atmospheric CO(2), seasonality of climate and fire. However, there is still no consensus about which of these factors triggered C(4) grassland expansion. We use a vegetation model, the adaptive dynamic global vegetation model (aDGVM), to test how CO(2), temperature, precipitation, fire and the tolerance of vegetation to fire influence C(4) grassland expansion. Simulations are forced with late Miocene climates generated with the Hadley Centre coupled ocean-atmosphere-vegetation general circulation model. We show that physiological differences between the C(3) and C(4) photosynthetic pathways cannot explain C(4) grass invasion into forests, but that fire is a crucial driver. Fire-promoting plant traits serve to expand the climate space in which C(4)-dominated biomes can persist. We propose that three mechanisms were involved in C(4) expansion: the physiological advantage of C(4) grasses under low atmospheric CO(2) allowed them to invade C(3) grasslands; fire allowed grasses to invade forests; and the evolution of fire-resistant savanna trees expanded the climate space that savannas can invade.

The stability of African savannas: insights from the indirect estimation of the parameters of a dynamic model.

Savannas are characterized by a competitive tension between grasses and trees, and theoretical models illustrate how this competitive tension is influenced by resource availability, competition for these resources, and disturbances. How this universe of theoretical possibilities translates into the real world is, however, poorly understood. In this paper we indirectly parameterize a theoretical model of savanna dynamics with the aim of gaining insights as to how the grass-tree balance changes across a broad biogeographical gradient. We use data on the abundance of trees in African savannas and Markov chain Monte Carlo methods to estimate the model parameters. The analysis shows that grasses and trees can coexist over a broad range of rainfall regimes. Further, our results indicate that savannas may be regulated by either asymptotically stable dynamics (in the absence of fire) or by stable limit cycles (in the presence of fire). Rainfall does not influence which of these two classes of dynamics occurs. We conclude that, even though fire might not be necessary for grass-tree coexistence, it nonetheless is an important modifier of grass: tree ratios.

Partitioning of root and shoot competition and the stability of savannas.

A classic problem in coexistence theory is how grasses and trees coexist in savannas. A popular deterministic model of savannas, the rooting niche separation model, is based on an assumption that is not empirically supported in many savannas. Alternative models that do not rely on the rooting niche assumption invoke intricate stochastic mechanisms that limit their attractiveness as general models of savannas. In this article we develop an alternative deterministic model of grass-tree interactions and use it to analyze the conditions under which grass-tree coexistence is possible. The novel feature of this model is that it partitions aboveground and belowground competition and simulates the fact that fire and herbivory remove only aboveground biomass. The model predicts that stable coexistence of grasses and trees is possible, even when grasses and trees do not have separate rooting niches. We show that when aboveground competition is intense, grasses can be excluded by trees; under such conditions, fire can prevent grasses from exclusion and induce a stable savanna state. The model provides a general framework for exploring the interactive effects of competition, herbivory, and fire on savanna systems.

Effects of four decades of fire manipulation on woody vegetation structure in Savanna.

The amount of carbon stored in savannas represents a significant uncertainty in global carbon budgets, primarily because fire causes actual biomass to differ from potential biomass. We analyzed the structural response of woody plants to long-term experimental burning in savannas. The experiment uses a randomized block design to examine fire exclusion and the season and frequency of burn in 192 7-ha experimental plots located in four different savanna ecosystems. Although previous studies would lead us to expect tree density to respond to the fire regime, our results, obtained from four different savanna ecosystems, suggest that the density of woody individuals was unresponsive to fire. The relative dominance of small trees was, however, highly responsive to fire regime. The observed shift in the structure of tree populations has potentially large impacts on the carbon balance. However, the response of tree biomass to fire of the different savannas studied were different, making it difficult to generalize about the extent to which fire can be used to manipulate carbon sequestration in savannas. This study provides evidence that savannas are demographically resilient to fire, but structurally responsive.