Deforestation and Reforestation of Latin America and the Caribbean (2001–2010)

Forest cover change directly affects biodiversity, the global carbon budget, and ecosystem function. Within Latin American and the Caribbean region (LAC), many studies have documented extensive deforestation, but there are also many local studies reporting forest recovery. These contrasting dynamics have been largely attributed to demographic and socio‐economic change. For example, local population change due to migration can stimulate forest recovery, while the increasing global demand for food can drive agriculture expansion. However, as no analysis has simultaneously evaluated deforestation and reforestation from the municipal to continental scale, we lack a comprehensive assessment of the spatial distribution of these processes. We overcame this limitation by producing wall‐to‐wall, annual maps of change in woody vegetation and other land‐cover classes between 2001 and 2010 for each of the 16,050 municipalities in LAC, and we used nonparametric Random Forest regression analyses to determine which environmental or population variables best explained the variation in woody vegetation change. Woody vegetation change was dominated by deforestation (?541,835 km²), particularly in the moist forest, dry forest, and savannas/shrublands biomes in South America. Extensive areas also recovered woody vegetation (+362,430 km²), particularly in regions too dry or too steep for modern agriculture. Deforestation in moist forests tended to occur in lowland areas with low population density, but woody cover change was not related to municipality‐scale population change. These results emphasize the importance of quantitating deforestation and reforestation at multiple spatial scales and linking these changes with global drivers such as the global demand for food.     

Land‐use change outweighs projected effects of changing rainfall on tree cover in sub‐Saharan Africa

Global change will likely affect savanna and forest structure and distributions, with implications for diversity within both biomes. Few studies have examined the impacts of both expected precipitation and land use changes on vegetation structure in the future, despite their likely severity. Here, we modeled tree cover in sub‐Saharan Africa, as a proxy for vegetation structure and land cover change, using climatic, edaphic, and anthropic data (R² = 0.97). Projected tree cover for the year 2070, simulated using scenarios that include climate and land use projections, generally decreased, both in forest and savanna, although the directionality of changes varied locally. The main driver of tree cover changes was land use change; the effects of precipitation change were minor by comparison. Interestingly, carbon emissions mitigation via increasing biofuels production resulted in decreases in tree cover, more severe than scenarios with more intense precipitation change, especially within savannas. Evaluation of tree cover change against protected area extent at the WWF Ecoregion scale suggested areas of high biodiversity and ecosystem services concern. Those forests most vulnerable to large decreases in tree cover were also highly protected, potentially buffering the effects of global change. Meanwhile, savannas, especially where they immediately bordered forests (e.g. West and Central Africa), were characterized by a dearth of protected areas, making them highly vulnerable. Savanna must become an explicit policy priority in the face of climate and land use change if conservation and livelihoods are to remain viable into the next century.     

The savannization of moist forests in the Sierra Nevada de Santa Marta, Colombia

In the Rio Ranchería watershed of the Sierra Nevada de Santa Marta, between 500 and 1500 m, savanna vegetation is interspersed with moist forests. The savannas are composed of native savanna grasses like Aristida adscensionis L., Arundinella sp., Panicum olyroides Kunth, and Schyzachyrium microstachyum (Desv.) Roseng., Arrill & Izag and the African Melinis minutiflora P. Beauv. There is also Curatella americana L. and Byrsonima crassifolia (L.) H.B.K., two typical tree species of the neotropical savannas. Although moist forest patches occur more often on lower slopes and narrow valley bottoms, they can also be found on mid- and upper-slopes and less often on ridges. Thus, these forest patches are not gallery forests as are found throughout the neotropics, but the result of deforestation and fractionation of a continuous forest. A comparison of soil profiles between the savannas and remnant forest patches on the same slope, showed the disappearance of the A and B horizons (approx. 50 cm) under savanna vegetation. The sharp difference between the savanna and forest soils at the Rio Ranchería does not appear to be due to a change in soil water status along a toposequence or differences in the underlying bedrock. We hypothesize that the savannas of the Rio Ranchería watershed, are the result of deforestation and land practices on infertile soils derived from granite. The savannization process was likely initiated by Amerindians by means of the frequent use of fire or clearing lands for the cultivation of maize. The introduction of cattle by Spaniards (c. 1530) and the frequent use of fire to maintain grazing fields, contributed to further degradation of the habitat. While some tropical landscapes recovered their forest cover when human pressure was removed approximately 500 years ago, areas such as the Rio Ranchería watershed have suffered permanent damage. The savannas of this region are likely to remain unless fire is suppressed and soil restoration practices implemented.     

Not all forests are expanding over central Brazilian savannas

Recently we reported on the expansion of riparian forests into savannas in central Brazil. To enlarge the scope of the earlier study we investigated whether upland deciduous and xeromorphic forests behaved similarly. We investigated past vegetation changes that occurred in forest/savanna transitions using carbon isotope ratios (δ¹³C) measured in the soil organic matter as a tracer. We analyzed the ¹⁴C activity where δ¹³C showed major shifts in vegetation. The role of soil chemical and physical attributes in defining vegetation distribution is discussed. Structural changes in vegetation were found to be associated with shifts in the isotope composition (δ¹³C) of soil organic matter. This was attributed to intrinsic differences in the biomass of trees and grasses and allowed for the determination of past shifts in vegetation by evaluating δ¹³C at different depths. The deciduous forest decreased in area approximately 980 years ago. Tree cover increased in the xeromorphic forest, but the border stayed stable through time. The deciduous forest and adjacent savanna have eutrophic soils while the xeromorphic forest and adjacent savanna have dystrophic soils. However, greater organic carbon, nitrogen and phosphorus concentrations are observed in the forests. We provide concrete evidence of deciduous forest retreat unlike the stability observed in the xeromorphic forest/savanna boundary. These results contrast with the expansion of riparian forests recently reported in the same region.     

Asymmetric response of forest and grassy biomes to climate variability across the African Humid Period: influenced by anthropogenic disturbance?

A comprehensive understanding of the relationship between land cover, climate change and disturbance dynamics is needed to inform scenarios of vegetation change on the African continent. Although significant advances have been made, large uncertainties exist in projections of future biodiversity and ecosystem change for the world's largest tropical landmass. To better illustrate the effects of climate–disturbance–ecosystem interactions on continental-scale vegetation change, we apply a novel statistical multivariate envelope approach to subfossil pollen data and climate model outputs (TraCE-21ka). We target paleoenvironmental records across continental Africa, from the African Humid Period (AHP: ca 14 700–5500 yr BP) – an interval of spatially and temporally variable hydroclimatic conditions – until recent times, to improve our understanding of overarching vegetation trends and to compare changes between forest and grassy biomes (savanna and grassland). Our results suggest that although climate variability was the dominant driver of change, forest and grassy biomes responded asymmetrically: 1) the climatic envelope of grassy biomes expanded, or persisted in increasingly diverse climatic conditions, during the second half of the AHP whilst that of forest did not; 2) forest retreat occurred much more slowly during the mid to late Holocene compared to the early AHP forest expansion; and 3) as forest and grassy biomes diverged during the second half of the AHP, their ecological relationship (envelope overlap) fundamentally changed. Based on these asymmetries and associated changes in human land use, we propose and discuss three hypotheses about the influence of anthropogenic disturbance on continental-scale vegetation change.     

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.     

Pantropical geography of lightning‐caused disturbance and its implications for tropical forests

Lightning is a major agent of disturbance, but its ecological effects in the tropics are unquantified. Here we used ground and satellite sensors to quantify the geography of lightning strikes in terrestrial tropical ecosystems, and to evaluate whether spatial variation in lightning frequency is associated with variation in tropical forest structure and dynamics. Between 2013 and 2018, tropical terrestrial ecosystems received an average of 100.4 million lightning strikes per year, and the frequency of strikes was spatially autocorrelated at local‐to‐continental scales. Lightning strikes were more frequent in forests, savannas, and urban areas than in grasslands, shrublands, and croplands. Higher lightning frequency was positively associated with woody biomass turnover and negatively associated with aboveground biomass and the density of large trees (trees/ha) in forests across Africa, Asia, and the Americas. Extrapolating from the only tropical forest study that comprehensively assessed tree damage and mortality from lightning strikes, we estimate that lightning directly damages c. 832 million trees in tropical forests annually, of which c. 194 million die. The similarly high lightning frequency in tropical savannas suggests that lightning also influences savanna tree mortality rates and ecosystem processes. These patterns indicate that lightning‐caused disturbance plays a major and largely unappreciated role in pantropical ecosystem dynamics and global carbon cycling.     

Herbaceous vegetation responses to experimental fire in savannas and forests depend on biome and climate

Fire–vegetation feedbacks potentially maintain global savanna and forest distributions. Accordingly, vegetation in savanna and forest ecosystems should have differential responses to fire, but fire response data for herbaceous vegetation have yet to be synthesized across biomes. Here, we examined herbaceous vegetation responses to experimental fire at 30 sites spanning four continents. Across a variety of metrics, herbaceous vegetation increased in abundance where fire was applied, with larger responses to fire in wetter and in cooler and/or less seasonal systems. Compared to forests, savannas were associated with a 4.8 (±0.4) times larger difference in herbaceous vegetation abundance for burned versus unburned plots. In particular, grass cover decreased with fire exclusion in savannas, largely via decreases in C₄ grass cover, whereas changes in fire frequency had a relatively weak effect on grass cover in forests. These differential responses underscore the importance of fire for maintaining the vegetation structure of savannas and forests.     

Holocene global warming and the origin of the Neotropical Gran Sabana in the Venezuelan Guayana

Aim The assumedly anomalous occurrence of savannas and forest–savanna mosaics in the Gran Sabana – a neotropical region under a climate more suitable for tropical rain forests – has been attributed to a variety of historical, climatic, and anthropogenic factors. This paper describes a previously undocumented shift in vegetation and climate that occurred during the early Holocene, and evaluates its significance for the understanding of the origin of the Gran Sabana vegetation. Location A treeless savanna locality of the Gran Sabana (4°30′–6°45′ N and 60°34′–62°50′ W), in the Venezuelan Guayana of northern South America, at the headwaters of the Caroní river, one of the major tributaries of the Orinoco river. Methods Pollen and charcoal analysis of a previously dated peat section spanning from about the Pleistocene/Holocene boundary until the present. Results Mesothermic cloud forests dominated by Catostemma (Bombacaceae) occupied the site around the Pleistocene/Holocene boundary. During the early Holocene, a progressive but relatively rapid trend towards savanna vegetation occurred, and eventually the former cloud forests were replaced by a treeless savanna. Some time after the establishment of savannas, a marked increase in charcoal particles indicates the occurrence of the first local fires. Main conclusions The occurrence of cloud forests at the Pleistocene/Holocene boundary contradicts the historical hypothesis according to which the Gran Sabana is a relict of the hypothetical widespread savannas that have been assumed to have dominated the region during the last glaciation. The first local fires recorded in the Holocene were on savanna vegetation, which is against the hypothesis of fire as the triggering factor for the establishment of these savannas. Climate change, in the form of global warming and a persistently drier climate, emerges as the most probable cause for the forest–savanna turnover.     

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.     

Savanna–forest hysteresis in the tropics

A simple dynamic model relating forest area in a region, its contribution to dry season precipitation and the effect on its own establishment was developed. The model equation shows hysteresis between forest and savannas as a function of imported dry season precipitation. Regions are either dominated by forests or savannas, with each ecosystem showing stability despite changes in imported dry season precipitation. Deforestation beyond a certain threshold value, however, could cause a collapse of forest ecosystems and replacement by savannas in marginal areas. The predictions of this model corroborate pollen core analysis in the Amazon basin, where historical stability of tropical forest cover has been shown despite global climate change.     

Colombian dry moist forest transitions in the Llanos Orientales—A comparison of model and pollen-based biome reconstructions

Colombian vegetation, at the ecological level of the biome, is reconstructed at six sites using pollen data assigned a priori to plant functional types and biomes. The chosen sites incorporate four savanna sites (Laguna Sardinas, Laguna Angel, El Piñal and Laguna Carimagua), a site on the transition between savanna and Amazon rainforest (Loma Linda) and a site within the Amazon rainforest (Pantano de Monica). The areal extent of tropical moist forest, tropical dry forest and steppe have been subject to significant change: differential responses of the vegetation to climatic shifts are related to changes in plant available moisture, duration of dry season and edaphic controls on the vegetation. The record from El Piñal shows that the present-day savanna vegetation, dominated by steppe (Poaceae) with little occurrence of woody savanna taxa (e.g. Curatella, Byrsonima), was present since the last glacial period of the northern hemisphere. Unfortunately, El Piñal is located on an edaphic savanna and is not particularly responsive to registering change. Most records cover the early Holocene; one site records the El Abra stadial (Younger Dryas equivalent), when forest expansion reflects more humid climatic conditions and higher plant available moisture. During the early and middle Holocene, the maximum expansion of steppe and tropical dry forest occurred, indicating that dry climatic conditions continued to around 4000 ¹⁴C BP. The following period, from shortly before 4000 ¹⁴C BP, is characterised by an increase in forest and gallery forests, reflecting a wetter period probably with a shorter annual dry season. Anthropogenic influence on the vegetation is recorded by all the records over the last millennial, particularly characterised by a reduction in forest cover and high amplitude changes in vegetation.Biome transitions from one type to another, and the environmental controls on this shift, are investigated by applying a vegetation model (BIOME-3). The model uses climatic data from six meteorological stations that, encompass a range of environments within lowland Colombia, which are similar to the pollen data. The signals of vegetation change can be translated to the main environmental controls of temperature and moisture to indicate the degree of change needed in these parameters to record the vegetation change depicted by the pollen data. Moisture balance is the dominant control on driving vegetation change whether under seasonal or annual control. The combined reconstruction from pollen data and model output of biome-scale vegetation dynamics for lowland Colombia allows an understanding of the environmental controls to be developed.     

Synergistic effects of drought and deforestation on the resilience of the south-eastern Amazon rainforest

The south-eastern Amazon rainforest is subject to ongoing deforestation and is expected to become drier due to climate change. Recent analyses of the distribution of tree cover in the tropics show three modes that have been interpreted as representing alternative stable states: forest, savanna and treeless states. This situation implies that a change in environmental conditions, such as in the climate, could cause critical transitions from a forest towards a savanna ecosystem. Shifts to savanna might also occur if perturbations such as deforestation exceed a critical threshold. Recovering the forest would be difficult as the savanna will be stabilized by a feedback between tree cover and fire. Here we explore how environmental changes and perturbations affect the forest by using a simple model with alternative tree-cover states. We focus on the synergistic effects of precipitation reduction and deforestation on the probability of regime shifts in the south-eastern Amazon rainforest. The analysis indicated that in a large part of the south-eastern Amazon basin rainforest and savanna could be two alternative states, although massive forest dieback caused by mean-precipitation reduction alone is unlikely. However, combinations of deforestation and climate change triggered up to 6.6 times as many local regime shifts than the two did separately, causing large permanent forest losses in the studied region. The results emphasize the importance of reducing deforestation rates in order to prevent a climate-induced dieback of the south-eastern Amazon rainforest.     

Shifts in functional traits elevate risk of fire‐driven tree dieback in tropical savanna and forest biomes

Numerous predictions indicate rising CO₂ will accelerate the expansion of forests into savannas. Although encroaching forests can sequester carbon over the short term, increased fires and drought‐fire interactions could offset carbon gains, which may be amplified by the shift toward forest plant communities more susceptible to fire‐driven dieback. We quantify how bark thickness determines the ability of individual tree species to tolerate fire and subsequently determine the fire sensitivity of ecosystem carbon across 180 plots in savannas and forests throughout the 2.2‐million km² Cerrado region in Brazil. We find that not accounting for variation in bark thickness across tree species underestimated carbon losses in forests by ~50%, totaling 0.22 PgC across the Cerrado region. The lower bark thicknesses of plant species in forests decreased fire tolerance to such an extent that a third of carbon gains during forest encroachment may be at risk of dieback if burned. These results illustrate that consideration of trait‐based differences in fire tolerance is critical for determining the climate‐carbon‐fire feedback in tropical savanna and forest biomes.     

Increased tree densities in South African savannas: >50 years of data suggests CO2 as a driver

For the past century, woody plants have increased in grasslands and savannas worldwide. Woody encroachment may significantly alter ecosystem functioning including fire regimes, herbivore carrying capacity, biodiversity and carbon storage capacity. Traditionally, increases in woody cover and density have been ascribed to changes in the disturbance regime (fire and herbivores) or rainfall. Increased atmospheric CO₂ concentrations may also contribute, by increasing growth rates of trees relative to grasses. This hypothesis is still heavily debated because usually potential CO₂ effects are confounded by changes in land use (disturbance regime). Here we analyse changes in woody density in fire experiments at three sites in South African savannas where the disturbance regime (fire and herbivores) was kept constant for 30 and 50 years. If global drivers had significant effects on woody plants, we would expect significant increases in tree densities and biomass over time under the constant disturbance regime. Woody density remained constant in a semiarid savanna but tripled in a mesic savanna between the 1970s and 1990s. At the third site, a semiarid savanna near the southern limits of the biome, tree density doubled from the mid 1990s to 2010. Interpretation of the causes is confounded by population recovery after clearing, but aerial photograph analysis on adjacent non‐cleared areas showed an accompanying 48% increase in woody cover. Increased CO₂ concentrations are consistent with increased woody density while other global drivers (rainfall) remained constant over the duration of the experiments. The absence of a response in one semiarid savanna could be explained by a smaller carbon sink capacity of the dominant species, which would therefore benefit less from increased CO₂. Understanding how savannas and grasslands respond to increased CO₂ and identifying the causes of woody encroachment are essential for the successful management of these systems.     

Classifying the Neotropical savannas of Belize using remote sensing and ground survey

Aim This paper evaluates a method of combining data from GPS ground survey with classifications of medium spatial resolution LANDSAT imagery to distinguish variations within Neotropical savannas and to characterize the boundaries between savanna areas and the associated gallery forests, seasonally dry forests and wetland communities. Location Rio Bravo Conservation Area, Orange Walk District, Belize, Central America. Methods Dry season LANDSAT data for 10 April 1993 and 9 March 2001 covering a conservation area of 240,000 acres (97,459 ha), were rectified to sub‐pixel accuracy using ground control points positioned by GPS ground survey. The 1993 image was used to assess the accuracy with which the boundaries between the savanna matrix and gallery forests, high forests, wetlands and water bodies could be discriminated. The image was classified by a maximum likelihood (ML) classifier and the shapes and areas of forest and wetland classes were compared with an interpretation of these land cover types from 1 : 24,000 aerial photography, mapped at 1 : 50,000 scale in 1993. The 2001 image was used to assess whether different subtypes of savanna could be distinguished from LANDSAT data. This required the creation of a reference (‘ground truth’) data set for testing classifications of the image. One hundred and sixty sample patches (650 ha, distributed over an area of 7000 ha) of ten sub‐types of savanna vegetation and associates identified using a physiognomic classification scheme, were delineated on the ground by GPS and divided into two subsets for training and testing. Continuous classifications of LANDSAT data covering the savannas were developed that estimated potential contributions from up to five sub‐types of land cover (grassland, wetland, pine woodland, gallery forest and palmetto). The accuracy of each classification was assessed by comparison against ground data. An ML classification was also produced for the 2001 image using the same areas for training. This allowed a comparison of the relative accuracy of both continuous and Boolean ML methods for classifying savanna areas. Results The boundary between savannas and evergreen forests, gallery forests and open water in the study region could be delineated by the ML classifier to within 2 pixels (60 m) using LANDSAT imagery. However, the constituent sub‐types within the savanna were poorly discriminated. Whilst the shape and extent of closed canopy forest, gallery forest, wetlands and water bodies agreed closely with the distributions interpreted from aerial photography, classes such as ‘open pine savanna’ or ‘grassland’ were only 45–65% accurate when tested against ground data. A continuous classification, estimating the proportions of three savanna vegetation subtypes (grassland, marshland and woodland) present in each pixel, correctly classified more of the ground data for these cover types than the comparable ML result. Proportional mixtures of the land cover estimated by the continuous classifier also compared realistically with the vegetation formations observed along ground transects. Main conclusions By using GPS, a ground survey of vegetation cover was accurately matched to remotely sensed imagery and the accuracy of delineating boundaries and classifying areas of savanna was assessed directly. This showed that ML classification techniques can reliably delineate the boundaries of savannas, but continuous classifiers more accurately and realistically represent the distribution of the subtypes comprising savanna land cover. By combining these ground survey and image classification methods, medium spatial resolution satellite sensor data can provide an affordable means for land managers to assess the nature, extent and distribution of savanna formations. Over time, using the archives of LANDSAT (and SPOT) data together with marker sites surveyed in the field, quantitative changes in the extents and boundaries of savannas in response to both natural (e.g. fire, hurricane and drought) and anthropogenic (e.g. cutting and disturbance) factors can be assessed.     

Simulating Deforestation in Minas Gerais,Brazil, under Changing Government Policies and Socioeconomic Conditions

Agricultural expansion is causing deforestation in Minas Gerais, Brazil, converting savanna and tropical dry forest to farmland, and in 2012, Brazil’s Forest Code was revised with the government reducing deforestation restrictions. Understanding the effects of policy change on rates and locations of natural ecosystem loss is imperative. In this paper, deforestation in Minas Gerais was simulated annually until 2020 using Dinamica Environment for Geoprocessing Objects (Dinamica EGO). This system is a state-of-the-art land use and cover change (LUCC) model which incorporates government policy, landscape maps, and other biophysical and anthropogenic datasets. Three studied scenarios: (i) business as usual, (ii) increased deforestation, and (iii) decreased deforestation showed more transition to agriculture from shrubland compared to forests, and consistent locations for most deforestation. The probability of conversion to agriculture is strongly tied to areas with the smallest patches of original biome remaining. Increases in agricultural revenue are projected to continue with a loss of 25% of the remaining Cerrado land in the next decade if profit is maximized. The addition of biodiversity value as a tax on land sale prices, estimated at over $750,000,000 USD using the cost of extracting and maintaining current species ex-situ, can save more than 1 million hectares of shrubland with minimal effects on the economy of the State of Minas Gerais. With environmental policy determining rates of deforestation and economics driving the location of land clearing, site-specific protection or market accounting of externalities is needed to balance economic development and conservation.     

Cascading biodiversity and functional consequences of a global change–induced biome switch

Aim At a regional scale, across southern Africa, woody thickening of savannas is becoming increasingly widespread. Using coupled vegetation and faunal responses (ants), we explore whether major changes in woody cover in savannas represent an increase in the density of savanna trees (C₄ grass layer remains intact) or a ‘regime shift’ in system state from savanna to thicket (=dry forest) where broad‐leaved, forest‐associated trees shade out C₄ grasses. Location Hluhluwe Game Reserve, South Africa. Methods We sampled paired open (low woody cover) and closed (high cover that have undergone an increase in tree density) sites. Vegetation was sampled using belt transects, and a combination of pitfall trapping and Winkler sampling was used for ants. Results Closed habitats did not simply contain a higher density of woody savanna species, but differed significantly in structure, functional composition (high prevalence of broad‐leaved trees, discontinuous C₄ grasses) and system properties (e.g. low flammability). Ant assemblage composition reflected this difference in habitat. The trophic structure of ant assemblages in the two habitats revealed a functional shift with much higher abundances of predatory species in the closed habitat. Main conclusions The predominance of species with forest‐associated traits and concomitant reduction of C₄ grasses in closed sites indicate that vegetation has undergone a shift in fundamental system state (to thicket), rather than simply savanna thickening. This biome shift has cascading functional consequences and implications for biodiversity conservation. The potential loss of many specialist savanna plant species is especially concerning, given the spatial extent and speed of this vegetation switch. Although it is not clear how easily the habitat switch can be reversed and how stable the thicket habitats are, it is likely in the not‐too‐distant future that conservation managers will be forced to make decisions on whether to actively maintain savannas.     

Multiple Scales of Control on the Structure and Spatial Distribution of Woody Vegetation in African Savanna Watersheds

Factors controlling savanna woody vegetation structure vary at multiple spatial and temporal scales, and as a consequence, unraveling their combined effects has proven to be a classic challenge in savanna ecology. We used airborne LiDAR (light detection and ranging) to map three-dimensional woody vegetation structure throughout four savanna watersheds, each contrasting in geologic substrate and climate, in Kruger National Park, South Africa. By comparison of the four watersheds, we found that geologic substrate had a stronger effect than climate in determining watershed-scale differences in vegetation structural properties, including cover, height and crown density. Generalized Linear Models were used to assess the spatial distribution of woody vegetation structural properties, including cover, height and crown density, in relation to mapped hydrologic, topographic and fire history traits. For each substrate and climate combination, models incorporating topography, hydrology and fire history explained up to 30% of the remaining variation in woody canopy structure, but inclusion of a spatial autocovariate term further improved model performance. Both crown density and the cover of shorter woody canopies were determined more by unknown factors likely to be changing on smaller spatial scales, such as soil texture, herbivore abundance or fire behavior, than by our mapped regional-scale changes in topography and hydrology. We also detected patterns in spatial covariance at distances up to 50–450 m, depending on watershed and structural metric. Our results suggest that large-scale environmental factors play a smaller role than is often attributed to them in determining woody vegetation structure in southern African savannas. This highlights the need for more spatially-explicit, wide-area analyses using high resolution remote sensing techniques.     

Seasonal leaf dynamics across a tree density gradient in a Brazilian savanna

Interactions between trees and grasses that influence leaf area index (LAI) have important consequences for savanna ecosystem processes through their controls on water, carbon, and energy fluxes as well as fire regimes. We measured LAI, of the groundlayer (herbaceous and woody plants <1-m tall) and shrub and tree layer (woody plants >1-m tall), in the Brazilian cerrado over a range of tree densities from open shrub savanna to closed woodland through the annual cycle. During the dry season, soil water potential was strongly and positively correlated with grass LAI, and less strongly with tree and shrub LAI. By the end of the dry season, LAI of grasses, groundlayer dicots and trees declined to 28, 60, and 68% of mean wet-season values, respectively. We compared the data to remotely sensed vegetation indices, finding that field measurements were more strongly correlated to the enhanced vegetation index (EVI, r ²=0.71) than to the normalized difference vegetation index (NDVI, r ²=0.49). Although the latter has been more widely used in quantifying leaf dynamics of tropical savannas, EVI appears better suited for this purpose. Our ground-based measurements demonstrate that groundlayer LAI declines with increasing tree density across sites, with savanna grasses being excluded at a tree LAI of approximately 3.3. LAI averaged 4.2 in nearby gallery (riparian) forest, so savanna grasses were absent, thereby greatly reducing fire risk and permitting survival of fire-sensitive forest tree species. Although edaphic conditions may partly explain the larger tree LAI of forests, relative to savanna, biological differences between savanna and forest tree species play an important role. Overall, forest tree species had 48% greater LAI than congeneric savanna trees under similar growing conditions. Savanna and forest species play distinct roles in the structure and dynamics of savanna–forest boundaries, contributing to the differences in fire regimes, microclimate, and nutrient cycling between savanna and forest ecosystems.