The impact of shrub encroachment on savanna bird diversity from local to regional scale

Aim  Evidence is accumulating of a general increase in woody cover of many savanna regions of the world. Little is known about the consequences of this widespread and fundamental ecosystem structural shift on biodiversity.
Location  South Africa.
Methods  We assessed the potential response of bird species to shrub encroachment in a South African savanna by censusing bird species in five habitats along a gradient of increasing shrub cover, from grassland/open woodland to shrubland dominated by various shrub species. We also explored historical bird species population trends across southern Africa during the second half of the 20th century to determine if any quantifiable shifts had occurred that support an ongoing impact of shrub encroachment at the regional scale.
Results  At the local scale, species richness peaked at intermediate levels of shrub cover. Bird species composition showed high turnover along the gradient, suggesting that widespread shrub encroachment is likely to lead to the loss of certain species with a concomitant decline in bird species richness at the landscape scale. Finally, savanna bird species responded to changes in vegetation structure rather than vegetation species composition: bird assemblages were very similar in shrublands dominated by Acacia mellifera and those dominated by Tarchonanthus camphoratus .
Main conclusions  Shrub encroachment might have a bigger impact on bird diversity in grassland than in open woodland, regardless of the shrub species. Species recorded in our study area were associated with historical population changes at the scale of southern Africa suggesting that shrub encroachment could be one of the main drivers of bird population dynamics in southern African savannas. If current trends continue, the persistence of several southern African bird species associated with open savanna might be jeopardized regionally.     

Stochastic losses of fire‐dependent endemic herbs revealed by a 65‐year chronosequence of dispersal‐limited woody plant encroachment

The factors responsible for maintaining diverse groundcover plant communities of high conservation value in frequently burned wet pine savannas are poorly understood. While most management involves manipulating extrinsic factors important in maintaining species diversity (e.g., fire regimes), most ecological theory (e.g., niche theory and neutral theory) examines how traits exhibited by the species promote species coexistence. Furthermore, although many ecologists focus on processes that maintain local species diversity, conservation biologists have argued that other indices (e.g., phylogenetic diversity) are better for evaluating assemblages in terms of their conservation value. I used a null model that employed beta‐diversity calculations based on Raup–Crick distances to test for deterministic herbaceous species losses associated with a 65‐year chronosequence of woody species encroachment within each of three localities. I quantified conservation value of assemblages by measuring taxonomic distinctness, endemism, and floristic quality of plots with and without woody encroachment. Reductions in herb species richness per plot attributable to woody encroachment were largely stochastic, as indicated by a lack of change in the mean or variance in beta‐diversity caused by woody encroachment in the savannas studied here. Taxonomic distinctness, endemism, and floristic quality (when summed across all species) were all greater in areas that had not experienced woody encroachment. However, when corrected for local species richness, only average endemism and floristic quality of assemblages inclusive of herbs and woody plants were greater in areas that had not experienced woody encroachment, due to the more restricted ranges and habitat requirements of herbs. Results suggest that frequent fires maintain diverse assemblages of fire‐dependent herb species endemic to the region. The stochastic loss of plant species, irrespective of their taxonomic distinctness, to woody encroachment suggests that the relevance of niche partitioning or phylogenetic diversity to the management of biodiversity in wet pine savannas is minimal.     

Woody Encroachment Removal from Midwestern Oak Savannas Alters Understory Diversity across Space and Time

Recovering biodiversity is a common goal during restoration; however, for many ecosystems, it is not well understood how restoration influences species diversity across space and time. I examined understory species diversity and composition after woody encroachment removal in a large-scale savanna restoration experiment in central Iowa, United States. Over a 4-year time series, restoration had profound effects across space and time, increasing richness at local and site-level scales. Restoration sites had increased α (within sample) Simpson's diversity and α and γ (site level) species richness relative to control sites, although γ and β (among sample) Simpson's diversity, β richness, and α species evenness were not affected. Changes in richness were driven by graminoids at the α and γ scales and woody species (and some evidence for forbs) at the α scale. Interestingly, indicator species analysis revealed that at least some species from all functional groups were promoted by restoration, although no species were significant indicators of pre-treatment or control sites. Both savanna and nonsavanna species were indicators of restored sites. Restoration promoted exotic species at both scales, although species with spring phenologies were unaffected. Woody encroachment removal may be a means to promote species establishment in savannas; however, in this study, it resulted in establishment and proliferation of native and exotic and savanna and nonsavanna species. Future work might consider reintroduction of key savanna species to supplement those that have established. Work like this demonstrates the utility of restoration experiments for conducting research on large- and multiscale processes, such as species diversity.     

Trade‐offs between carbon stocks and biodiversity in European temperate forests

Policies to mitigate climate change and biodiversity loss often assume that protecting carbon‐rich forests provides co‐benefits in terms of biodiversity, due to the spatial congruence of carbon stocks and biodiversity at biogeographic scales. However, it remains unclear whether this holds at the scales relevant for management, and particularly large knowledge gaps exist for temperate forests and for taxa other than trees. We built a comprehensive dataset of Central European temperate forest structure and multi‐taxonomic diversity (beetles, birds, bryophytes, fungi, lichens, and plants) across 352 plots. We used Boosted Regression Trees (BRTs) to assess the relationship between above‐ground live carbon stocks and (a) taxon‐specific richness, (b) a unified multidiversity index. We used Threshold Indicator Taxa ANalysis to explore individual species’ responses to changing above‐ground carbon stocks and to detect change‐points in species composition along the carbon‐stock gradient. Our results reveal an overall weak and highly variable relationship between richness and carbon stock at the stand scale, both for individual taxonomic groups and for multidiversity. Similarly, the proportion of win‐win and trade‐off species (i.e., species favored or disadvantaged by increasing carbon stock, respectively) varied substantially across taxa. Win‐win species gradually replaced trade‐off species with increasing carbon, without clear thresholds along the above‐ground carbon gradient, suggesting that community‐level surrogates (e.g., richness) might fail to detect critical changes in biodiversity. Collectively, our analyses highlight that leveraging co‐benefits between carbon and biodiversity in temperate forest may require stand‐scale management that prioritizes either biodiversity or carbon in order to maximize co‐benefits at broader scales. Importantly, this contrasts with tropical forests, where climate and biodiversity objectives can be integrated at the stand scale, thus highlighting the need for context‐specificity when managing for multiple objectives. Accounting for critical change‐points of target taxa can help to deal with this specificity, by defining a safe operating space to manipulate carbon while avoiding biodiversity losses.     

Sixty years of community change in the prairie–savanna–forest mosaic of Wisconsin

Biodiversity loss is a global concern, and maintaining habitat complexity in naturally patchy landscapes can help retain regional diversity. A mosaic of prairie, savanna, and forest historically occurred across central North America but currently is highly fragmented due to human land conversion. It is unclear how each habitat type now contributes to regional diversity. Using legacy data, we resurveyed savanna plant communities originally surveyed in the 1950s to compare change in savannas to that in remnant forests and prairies. Savanna community structure and composition changed substantially over the past 60 years. Tree canopy density nearly doubled and many prairie and savanna specialist species were replaced by forest and non‐native species. All three habitats gained and lost many species since the 1950s, resulting in large changes in community composition from local colonizations and extinctions. Across all three habitats, regional species extinctions matched that of regional colonization resulting in no net change in regional species richness. Synthesis—Despite considerable species turnover within savannas, many species remain within the broader prairie–savanna–forest mosaic. Both regional extinctions and colonizations were high over the past 60 years, and maintaining the presence of all three community types—prairie, savanna and forest—on the landscape is critical to maintaining regional biodiversity.     

Water availability drives aboveground biomass and bird richness in forest restoration plantings to achieve carbon and biodiversity cobenefits

To combat global warming and biodiversity loss, we require effective forest restoration that encourages recovery of species diversity and ecosystem function to deliver essential ecosystem services, such as biomass accumulation. Further, understanding how and where to undertake restoration to achieve carbon sequestration and biodiversity conservation would provide an opportunity to finance ecosystem restoration under carbon markets. We surveyed 30 native mixed‐species plantings in subtropical forests and woodlands in Australia and used structural equation modeling to determine vegetation, soil, and climate variables most likely driving aboveground biomass accrual and bird richness and investigate the relationships between plant diversity, aboveground biomass accrual, and bird diversity. We focussed on woodland and forest‐dependent birds, and functional groups at risk of decline (insectivorous, understorey‐nesting, and small‐bodied birds). We found that mean moisture availability strongly limits aboveground biomass accrual and bird richness in restoration plantings, indicating potential synergies in choosing sites for carbon and biodiversity purposes. Counter to theory, woody plant richness was a poor direct predictor of aboveground biomass accrual, but was indirectly related via significant, positive effects of stand density. We also found no direct relationship between aboveground biomass accrual and bird richness, likely because of the strong effects of moisture availability on both variables. Instead, moisture availability and patch size strongly and positively influenced the richness of woodland and forest‐dependent birds. For understorey‐nesting birds, however, shrub cover and patch size predicted richness. Stand age or area of native vegetation surrounding the patch did not influence bird richness. Our results suggest that in subtropical biomes, planting larger patches to higher densities, ideally using a diversity of trees and shrubs (characteristics of ecological plantings) in more mesic locations will enhance the provision of carbon and biodiversity cobenefits. Further, ecological plantings will aid the rapid recovery of woodland and forest bird richness, with comparable aboveground biomass accrual to less diverse forestry plantations.     

Local native plant diversity responds to habitat loss and fragmentation over different time spans and spatial scales

Separating the threats that habitat loss and habitat fragmentation pose to biodiversity is challenging because these processes usually occur simultaneously. Additionally, their importance may be underestimated due to time-delayed extinction. In central Texas savannas, woody plant encroachment reduces the amount of habitat available to herbaceous species while fragmenting remaining habitat. We examined the relationships between present species richness and present and past habitat amount and fragmentation (measured as fractal dimension) using a series of aerial photographs taken over nearly 60 years. We show that woody plant encroachment, a common phenomenon in savannas worldwide, reduces the diversity of herbaceous vegetation through both habitat loss and fragmentation. Habitat loss has the strongest impact on species richness over short time spans and small spatial scales in these savannas. Habitat fragmentation, however, has the strongest impact over longer time spans and larger spatial scales and generates long-term extinction debts. We also demonstrate that examining habitat loss and habitat fragmentation across different time periods and at different spatial scales is essential for understanding their joint and individual effects on plant community composition.     

Multi‐scale patterns and bush encroachment in an arid savanna with a shallow soil layer

Abstract. Question: Bush encroachment (i.e. an increase in density of woody plants often unpalatable to domestic livestock) is a serious problem in many savannas and threatens the livelihood of many pastoralists. Can we derive a better understanding of the factors causing bush encroachment by investigating the scale dependency of patterns and processes in savannas? Location: An arid savanna in the Khomas Hochland, Namibia. Methods: Patterns of bush, grass, and soil nutrient distribution were surveyed on several scales along a rainfall gradient, with emphasis on intraspecific interactions within the dominant woody species, Acacia reficiens. Results: Savannas can be interpreted as patch‐dynamic systems where landscapes are composed of many patches (a few ha in size) in different states of transition between grassy and woody dominance. Conclusions: In arid savannas, this patchiness is driven both by rainfall that is highly variable in space and time and by inter‐tree competition. Within the paradigm of patch‐dynamic savannas, bush encroachment is part of a cyclical succession between open savanna and woody dominance. The conversion from a patch of open savanna to a bush‐encroached area is initiated by the spatial and temporal overlap of several (localized) rainfall events sufficient for Acacia germination and establishment. With time, growth and self‐thinning will transform the bush‐encroached area into a mature Acacia stand and eventually into open savanna again. Patchiness is sustained due to the local rarity (and patchiness) of rainfall sufficient for germination of woody plants as well as by plant‐soil interactions.     

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.     

Fire‐sensitive species dominate seed rain after fire suppression: Implications for plant community diversity and woody encroachment in the Cerrado

Woody encroachment is becoming common in tropical savannas. We studied natural seed rain and performed seed addition experiments in a Brazilian savanna that had not been burned for several decades. We found greater abundance of fire‐sensitive species in the seed rain, likely contributing to woody encroachment. Flexible fire management policies that allow for natural and prescribed fires may be required to maintain savanna diversity.     

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.     

Undertaking large‐scale forest restoration to generate ecosystem services

The global community is seeking to substantially restore the world's forest cover to improve the supply of ecosystem services. However, it is not clear what type of reforestation must be used and there is a risk that the techniques used in industrial timber plantations will become the default methodology. This is unlikely to be sufficient because of the well‐known relationship between biodiversity and ecological functioning. Restoration may be achieved through natural regeneration but this may not always occur at critical locations. Ecological restoration involving species‐rich plantings might also be used but can be difficult to implement at landscape scales. I review here the consequence of planting more limited numbers of species and the effects of this on the delivery of ecosystem services. Evidence suggests many commonly sought ecosystem services—though not all—may be generated by the modest levels of species richness provided these species have appropriate traits. The literature also shows that the alpha diversity of restored forests is not the only driver of functionality and that the location and extent of any reforestation are significant as well; beta and gamma diversity may also affect functionality but these relationships remain unclear. Encouraging the adoption of even moderately diverse plantings at landscape scales and at key locations will require policies and institutions to balance the type, location, and scale of restoration and make the necessary trade‐offs between national and local aspirations. New approaches and metrics will have to be developed to monitor and assess restoration success at these larger scales.     

Use of indigenous ecological knowledge of the Maasai pastoralists for assessing rangeland biodiversity in Tanzania

This paper incorporates the indigenous ecological knowledge (IEK) of the Maasai pastoralists and ecological methods to assess effects of grazing and cropping on rangeland biodiversity at macro‐ and micro‐landscape scales in northern Tanzania. The joint surveys with pastoralists identified indicator plant species and their associations with micro‐landscapes and livestock grazing suitability (i.e. for cattle and small ruminant grazing), while traditional calf‐pasture reserves (alalili pl. alalilia) were evaluated for preservation of rangeland biodiversity. The macro‐landscapes comprising the cool high plateau (osupuko pl. isipuki) and montane forest highland (endim) were included in the survey. At micro‐landscape scales, the osupuko was classified into uplands (orkung'u), slopes (andamata) and dry valley bottomlands (ayarata). The micro‐landscapes were assessed in terms of herbaceous plant species and woody species richness and risks of soil erosion. Biodiversity varied at both the macro‐ and micro‐landscape scales and in accordance with the land‐use types. Greater plant species diversity and less erosion risks were found in the pastoral landscapes than in the agro‐pastoral landscapes. The calf‐grazing pastures had greater herbaceous species richness than the non‐calf pastures, which in turn had more woody species. The study concludes that the indigenous systems of landscape classification provides a valuable basis for assessing rangeland biodiversity, which ecologists should incorporate into ecological surveys of the rangelands in East Africa in the future.     

Habitat diversity associated to island size and environmental filtering control the species richness of rock‐savanna plants in neotropical inselbergs

Disentangling the multiple factors controlling species diversity is a major challenge in ecology. Island biogeography and environmental filtering are two influential theories emphasizing respectively island size and isolation, and the abiotic environment, as key drivers of species richness. However, few attempts have been made to quantify their relative importance and investigate their mechanistic basis. Here, we applied structural equation modelling, a powerful method allowing test of complex hypotheses involving multiple and indirect effects, on an island‐like system of 22 French Guianan neotropical inselbergs covered with rock‐savanna. We separated the effects of size (rock‐savanna area), isolation (density of surrounding inselbergs), environmental filtering (rainfall, altitude) and dispersal filtering (forest‐matrix openness) on the species richness of all plants and of various ecological groups (terrestrial versus epiphytic, small‐scale versus large‐scale dispersal species). We showed that the species richness of all plants and terrestrial species was mainly explained by the size of rock‐savanna vegetation patches, with increasing richness associated with higher rock‐savanna area, while inselberg isolation and forest‐matrix openness had no measurable effect. This size effect was mediated by an increase in terrestrial‐habitat diversity, even after accounting for increased sampling effort. The richness of epiphytic species was mainly explained by environmental filtering, with a positive effect of rainfall and altitude, but also by a positive size effect mediated by enhanced woody‐plant species richness. Inselberg size and environmental filtering both explained the richness of small‐scale and large‐scale dispersal species, but these ecological groups responded in opposite directions to altitude and rainfall, that is positively for large‐scale and negatively for small‐scale dispersal species. Our study revealed both habitat diversity associated with island size and environmental filtering as major drivers of neotropical inselberg plant diversity and showed the importance of plant species growth form and dispersal ability to explain the relative importance of each driver.     

Multiscale Patterns of Riparian Plant Diversity and Implications for Restoration

Planning riparian restoration to resemble historic reference conditions requires an understanding of both local and regional patterns of plant species diversity. Thus, understanding species distributions at multiple spatial scales is essential to improve restoration planting success, to enhance long‐term ecosystem functioning, and to match restoration planting designs with historic biogeographic distributions. To inform restoration planning, we examined the biogeographic patterns of riparian plant diversity at local and regional scales within a major western U.S.A. drainage, California's Sacramento—San Joaquin Valley. We analyzed patterns of species richness and complementarity (β‐diversity) across two scales: the watershed scale and the floodplain scale. At the watershed scale, spatial patterns of native riparian richness were driven by herbaceous species, whereas woody species were largely cosmopolitan across the nearly 38,000 km² study area. At the floodplain scale, riparian floras reflected species richness and dissimilarity patterns related to hydrological and disturbance‐driven successional sequences. These findings reinforce the importance of concurrently evaluating both local and regional processes that promote species diversity and distribution of native riparian flora. Furthermore, as restoration activities become more prevalent across the landscape, strategies for restoration outcomes should emulate the patterns of species diversity and biogeographic distributions found at regional scales.     

Bird diversity and endemism along a land‐use gradient in Madagascar: The conservation value of vanilla agroforests

Land‐use change is the most important driver of biodiversity loss worldwide and particularly so in the tropics, where natural habitats are transformed into large‐scale monocultures or heterogeneous landscape mosaics of largely unknown conservation value. Using birds as an indicator taxon, we evaluated the conservation value of a landscape mosaic in northeastern Madagascar, a biodiversity hotspot and the center of global vanilla production. We assessed bird species richness and composition by conducting point counts across seven prevalent land‐use types (forest‐ and fallow‐derived vanilla agroforests, woody and herbaceous fallow that are part of a shifting cultivation system, rice paddy, forest fragment and contiguous old‐growth forest). We find that old‐growth forest had the highest species richness, driven by a high share of endemics. Species richness and community composition in forest‐derived vanilla agroforest were similar to forest fragment, whereas fallow‐derived vanilla agroforest was most comparable to woody fallow. The open land‐use types herbaceous fallow and rice paddy had fewest species. Across forest fragments, vanilla agroforests, and woody fallows, endemic bird species richness was positively correlated to landscape‐scale forest cover. We conclude that both fallow‐ and forest‐derived vanilla agroforests play an important but contrasting role for bird conservation: Fallow‐derived agroforests are less valuable but take fallow land out of the shifting cultivation cycle, possibly preventing further degradation. Conversely, forest‐derived agroforests contribute to forest degradation but may avoid total loss of tree cover from forest fragments. Considering the land‐use history of agroforests may thus be a promising avenue for future research beyond the case of vanilla. Abstract in Malagasay is available with online material     

Determinants of woody encroachment and cover in African savannas

Savanna ecosystems are an integral part of the African landscape and sustain the livelihoods of millions of people. Woody encroachment in savannas is a widespread phenomenon but its causes are widely debated. We review the extensive literature on woody encroachment to help improve understanding of the possible causes and to highlight where and how future scientific efforts to fully understand these causes should be focused. Rainfall is the most important determinant of maximum woody cover across Africa, but fire and herbivory interact to reduce woody cover below the maximum at many locations. We postulate that woody encroachment is most likely driven by CO₂ enrichment and propose a two-system conceptual framework, whereby mechanisms of woody encroachment differ depending on whether the savanna is a wet or dry system. In dry savannas, the increased water-use efficiency in plants relaxes precipitation-driven constraints and increases woody growth. In wet savannas, the increase of carbon allocation to tree roots results in faster recovery rates after disturbance and a greater likelihood of reaching sexual maturity. Our proposed framework can be tested using a mixture of experimental and earth observational techniques. At a local level, changes in precipitation, burning regimes or herbivory could be driving woody encroachment, but are unlikely to be the explanation of this continent-wide phenomenon.     

Woody overstorey effects on soil carbon and nitrogen pools in South African savanna

Abstract Woody plant encroachment in savannas may alter carbon (C) and nitrogen (N) pools over the long‐term, which could have regional or global biogeochemical implications given the widespread encroachment observed in the vast savanna biome. Soil and litter %C and %N were surveyed across four soil types in two encroached, semi‐arid savanna landscapes in northern South Africa. Litter at sampling points with a woody component had a higher %N and lower C : N ratio than litter at solely herbaceous points. Severely encroached areas had lower C : N ratios throughout the soil profile than less encroached areas. Soil %C and %N were highly influenced by soil texture but were also influenced by the presence of a woody overstorey, which increased surface soil %C on three soil types but decreased it on the most heavily encroached soil type. Soil C sequestration may initially increase with bush encroachment but then decline if bush densities become so high as to inhibit understorey grass growth.     

Temporal changes in the dominance of tree functional traits,but no changes in species diversity during woody plant encroachment in a Brazilian savanna

Questions : Woody encroachment in savannas has been associated with changing taxonomic composition and ecosystem function. Interestingly, there is little understanding of how encroachment impacts plant functional diversity and how those changes relate to plant demography, a crucial mediator between taxonomic composition and ecosystem function. Location : Southeastern Brazil. Methods: Using a landscape scale fire suppression experiment in a diverse Brazilian savanna, we quantify how change in species composition over seven years impacted vegetative and reproductive tree functional diversity as determined by new recruits, dead and surviving trees. Results: Over seven years, tree above-ground biomass increased by 15%, while total species richness did not change. Despite minor changes, species composition remained overall similar (82%), with few species contributing significantly to plot dissimilarity over time. There were small changes in vegetative traits, where the community-weighted mean increased in maximum tree height (↑ 2.1%) and specific leaf area (↑ 5.3%), and decreased in wood density (↓ 1.3%) and bark thickness (↓ 9.4%). Changes in reproductive traits were larger than in vegetative traits, with an increase in the prevalence of monoecy (↑ 32.6%), dioecy (↑ 44.2%), large seeds (↑ 20.3%), animal-mediated seed dispersal (↑ 4.9%) and pollination by very small insects (↑ 45.5%), and a decrease in the prevalence of hermaphroditism (↓ 9%), small seeds (6.8%) and pollination by small insects (12.5%). The overall decrease in bark thickness and increase in monoecy and dioecy were mainly driven by characters of the new recruits, while the overall increase in specific leaf area (SLA) and decrease in small seeds appeared largely determined by the loss of trees possessing those traits. Conclusions: Encroachment leads to changes that are likely increasing ecosystem vulnerability to fire and drought. Further, the compositional changes observed appear to drive marked change in reproductive traits, indicating increasing dependence on animals for dispersal and reproduction. Understanding post-hoc encroachment impacts in an era of widespread pervasive encroachment is fundamental to reconciling ecosystem functions such as nutrient cycling and pollination services as there is a loss of species with open ecosystem life-history strategies. Among savannas, there remains an urgent need to understand relationships between woody cover and ecosystem function to determine thresholds in woody cover promoting resilient savanna ecosystems.     

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.