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Global climate change is predicted to increase the frequency of droughts, with major impacts on tropical savannas. It has been suggested that during drought, increased soil moisture and nutrients on termite mounds could benefit plants but it is unclear how such benefits could cascade to affect insect communities.Here, we describe the effects of drought on vegetation structure, the cascading implications for invertebrates and how termite mounds influence such effects. We compared how changes in grass biomass affected grasshopper and ant diversity on and off Macrotermes mounds before (2012) and during a drought (2016) at two locations that experienced large variation in drought severity (Skukuza and Pretoriuskop) in the Kruger National Park, South Africa.The 2013–2016 drought was not ubiquitous across the study site, with rainfall decreasing at Skukuza and being above average at Pretoriuskop. However, grass biomass declined at both locations. Grasshopper abundance decreased at droughted Skukuza both on and off mounds but decreased on mounds and increased off mounds at non‐droughted Pretoriuskop. Ant abundance and species richness increased at Skukuza but remained the same on mounds and decreased off mounds at Pretoriuskop.Our results demonstrate the spatially extensive effects of drought. Despite above average rainfall in 2016 at Pretoriuskop, grass biomass decreased, likely due to an influx of large mammalian herbivores from drought‐affected areas. This decrease in grass biomass cascaded to affect grasshoppers and ants, further illustrating the effects of drought on invertebrates in adjoining areas with higher rainfall. Our grasshopper results also suggest that increased drought in savannas will contribute to overall declines in insect abundance. Moreover, our recorded increase in ant abundance was primarily in the form of increases in dominant species, illustrating how drought‐induced shifts in relative abundance will likely influence ecosystem structure and function.Our study highlights the phenomenon of spill‐over drought effects and suggests rather than mitigating drought, termite mounds can instead become the focus for more intense grazing, with important consequences for insect communities. [ABSTRACT FROM AUTHOR]

Grazing lawn and flammable-tussock grass communities are contrasting resource pools for mammalian grazers in terms of forage quantity and quality. Drought events fundamentally alter forage availability within these communities and therefore should alter herbivore use with repercussions for the recovery and functioning of ecosystems after drought. During and after an intense El Niño drought (2014–2017) in Kruger National Park, South Africa, we addressed two questions: (1) how does herbivore use of different grass types change during a drought and (2) how do these changes affect grass productivity post-drought? We monitored grazer use of three different grass communities (lawn, tussock and burned-tussock) at a landscape scale and measured primary productivity monthly during and post-drought. For the first drought year, grazer numbers were highest on grazing lawn communities. This pattern continued into the second dry growing season, until herbivores finally left the study area. Both lawns and tussock grasslands recovered rapidly after the first good rainfall (productivity > 150 g m−2 per month). However, grazers did not return to feed on the same patches they had frequented pre-drought resulting in grazing lawn grasses self-shading and senescing. Longer droughts have the potential to decouple grazers and grazing lawns with negative impacts on lawn productivity and persistence that could drive the loss of lawns in savanna landscapes and impact mesoherbivore populations. It is clear from our results that grazer effects need to be incorporated into drought frameworks to understand the consequences of droughts for grassland function. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Abstract: Forests and grassy vegetation (savannas and grasslands) are alternative ecosystem states in many tropical landscapes. Relative to forests the grassy ecosystems are poorly known and poorly conserved, partly because they were thought to be products of forest clearance. However many grasslands have proved to be ancient. Commensurate with their antiquity, grassy biomes have distinct suites of plant and animal species that contribute a large fraction of the diversity of forest–grassland mosaics. Grasslands differ strikingly from forests in their ecology and in the nature of threats to their future. Here we highlight the high biodiversity value of grassy biomes and, in contrast to tropical forests, we illustrate the importance of fire in maintaining these systems. We discuss the major threats to, and consequences for, biodiversity in these regions including land clearance and elevated CO2-driven forest expansion. Finally we focus on the difficulties of grassland restoration. A new approach to understanding and conserving grassy ecosystems, free from cultural prejudices of the past, is long overdue. [ABSTRACT FROM AUTHOR]

Tropical savannas and rainforests contrast in their flammability and the fire resilience of their associated species. While savanna species generally exhibit high resilience to burning, there is much debate about the fire resilience of forest-associated species, and the persistence of forest patches in a flammable savanna matrix. Where fire has been excluded, savanna tends on a trajectory towards forest, with an increase in forest-associated plants and animal species. This study tested the idea that given the high proportion of forest-associated taxa in long-unburnt savanna, the fauna of these areas would be expected to exhibit less resilience to fire than the fauna in frequently burnt savannas. The study investigated the immediate and short-term effects on ant assemblages of re-introducing fire into long-unburnt savanna in northern Australia. The ant fauna exhibited high resistance to fires, with no significant short-term change in mean abundance or species richness; instead, seasonality had a far stronger influence on overall ant activity. Fire caused dramatic declines in dominance of the patchily distributed forest-associated species Oecophylla smaragdina and Papyrius sp., but had no effect on overall dominance by open savanna species of Iridomyrmex. Dominance by Iridomyrmex pallidus declined, but this was compensated for by increases in I. reburrus, while two other species of Iridomyrmex showed no change. This indicates a high level of functional redundancy among dominant species of Iridomyrmex, which universally dominate open savanna communities, but not of dominant forest-associated species. Overall, our findings demonstrate a high degree of fire-resilience of the long-unburnt savanna ant fauna. Despite the occurrence of forest-associated species, the high proportion of savanna species persisting in this habitat means that long-unburnt savanna retains the general response characteristics of frequently burnt savanna. [ABSTRACT FROM AUTHOR]

Fire management is increasingly focusing on introducing heterogeneity in burning patterns under the assumption that “pyrodiversity begets biodiversity.” This concept has been formalized as patch mosaic burning (PMB), in which fire is manipulated to create a mosaic of patches representative of a range of fire histories to generate heterogeneity across space and time. Although PMB is an intuitively appealing concept, it has received little critical analysis. Thus we examined ecosystems where PMB has received the most attention and has been the most extensively implemented: tropical and subtropical savannas of Australia and Africa. We identified serious shortcomings of PMB: the ecological significance of different burning patterns remains unknown and details of desired fire mosaics remain unspecified. This has led to fire-management plans based on pyrodiversity rhetoric that lacks substance in terms of operational guidelines and capacity for meaningful evaluation. We also suggest that not all fire patterns are ecologically meaningful: this seems particularly true for the highly fire-prone savannas of Australia and South Africa. We argue that biodiversity-needs-pyrodiversity advocacy needs to be replaced with a more critical consideration of the levels of pyrodiversity needed for biodiversity and greater attention to operational guidelines for its implementation. [ABSTRACT FROM AUTHOR]

1. Despite the fact that fire is considered an important disturbance in savannas across the world and is used widely as a management tool in conservation areas, little is known about the effects of burning on their insect communities. 2. This study made use of a 50-year fire experiment to investigate the responses of ant assemblages to long-term burning regimes. The effects of fire frequency, season and time since fire (fuel age) were tested on epigaeic ants across three savanna vegetation types (Mopane woodland, Acacia savanna and Terminalia woodland) in Kruger National Park, South Africa. 3. There was no significant effect of burning on mean ant species richness and abundance between treatments, although there were significant differences in ant assemblage composition between burned (treatment) and unburned (control) plots. The effects of season, frequency of burn and plot age on assemblage structure were weak and often not significant. 4. Epigaeic ant assemblages in this savanna system appeared to be highly resistant and resilient to burning. The response of ants to fire was linked to changes in habitat cover and structure: the effect of fire on vegetation and ants was less pronounced in lower rainfall areas, where differences in vegetation structure between burnt and unburnt plots were less pronounced than in higher rainfall areas. 5. Synthesis and applications. The effect of fire on ant assemblages in the mid- to northern Kruger National Park depends on whether a patch burns or not, rather than the specifics of the burning treatment. Thus, conservation managers can focus concerns regarding the subtleties of fire regimes on other taxa or areas of particular concern. Journal of Applied Ecology (2004) 41, 630–642 [ABSTRACT FROM AUTHOR]

Aim: The mosaic of savannas that persists in the forest‐dominant Congo Basin is thought to be palaeoclimatic relics, but past biogeographical processes that have formed and maintained these systems are poorly understood. Here, we explored the post‐Pleistocene biogeography of Gabon's savannas using termites as biological indicators to understand historical and mechanistic factors influencing present‐day termite communities in the country's extant savannas. Location: Gabon, Central Africa. Taxon: Blattodea: Termitoidae. Methods: Using standardised transect methods, we sampled termite communities in four disjunct modern savanna areas of Gabon: the centre (Lopé), the southeast (Batéké) and the south (Mayombe North and South). Termites at Lopé were collected in three habitats (annually burned savannas, savannas with a depressed fire regime and forest). We used DNA barcoding of the COII region to identify termite species and compared abundance, species richness and community composition across areas and habitats. Results: Community composition differed greatly between Lopé and both Batéké and Mayombe savannas with Lopé being exceptionally depauperate and lacking characteristic savanna species. Within Lopé, termite abundance and diversity was highest in forests and lowest in annually burned savannas, with a gradual change in species composition across the forest–savanna gradient associated with fire history. Main Conclusions: The absence of savanna typical species in Lopé savannas challenges current assumptions that these savannas were linked to the south/southeastern savannas during the Pleistocene and suggests a different evolutionary history. Lopé savannas may instead have opened as an isolated grassland and never have been contiguous with neighbouring savannas, or were isolated soon after forest expansion began and have now lost savanna‐typical species. Furthermore, the patterns of termite community composition in fire suppressed savannas support a hypothesis of rapid change driven by fire frequency where either fire suppression or infrequent burning over 23 years has meant savannas have become ecologically much more forest‐like. [ABSTRACT FROM AUTHOR]

Understanding the factors that control decomposition is critical for predicting how the carbon cycle will alter with global change. Until recently, the accepted paradigm was that climate primarily drives decomposition rates, and interactions among decomposers only control variation at finer scales. Although it is now understood that biotic agents can play an important role, we know less about the importance of species interactions with a lack of field experiments at a large scale. Predation is a key ecological interaction that could influence decomposition by directly or indirectly regulating the abundances of decomposer organisms, but a comprehensive understanding of the cascading effects that predation can have on decomposition, particularly at large scales, is missing.Here we report on an experiment where we suppressed the abundance of ants, which are major predators of termites, at a hectare scale in a natural African savanna and examined how this affected the decomposition of three common substrates (wood, grass and dung).Our study revealed that ants exert considerable top‐down control on decomposition via their predation of termites: decomposition of wood, grass and dung increased by 98%, 74% and 84% with ant suppression, respectively. Suppression of ants increased termite activity and consequently resulted in increased termite‐mediated decomposition. Remarkably, for all substrates, the suppression of ants nearly doubled decomposition by termites. Additionally, for grass and dung substrates, the dominant agent of decomposition switched from microbes to termites with ant suppression.Our study highlights the critical importance of considering species interactions in decomposition studies, particularly given declines in predatory species in the Anthropocene. Read the free Plain Language Summary for this article on the Journal blog. [ABSTRACT FROM AUTHOR]

Deadwood is a large global carbon store with its store size partially determined by biotic decay. Microbial wood decay rates are known to respond to changing temperature and precipitation. Termites are also important decomposers in the tropics but are less well studied. An understanding of their climate sensitivities is needed to estimate climate change effects on wood carbon pools. Using data from 133 sites spanning six continents, we found that termite wood discovery and consumption were highly sensitive to temperature (with decay increasing >6.8 times per 10°C increase in temperature)—even more so than microbes. Termite decay effects were greatest in tropical seasonal forests, tropical savannas, and subtropical deserts. With tropicalization (i.e., warming shifts to tropical climates), termite wood decay will likely increase as termites access more of Earth’s surface. [ABSTRACT FROM AUTHOR]

The most diverse and abundant family of termites, the Termitidae, evolved in African tropical forests. They have since colonised grassy biomes such as savannas. These open environments have more extreme conditions than tropical forests, notably wider extremes of temperature and lower precipitation levels and greater temporal fluctuations (of both annual and diurnal variation). These conditions are challenging for soft‐bodied ectotherms, such as termites, to survive in, let alone become as ecologically dominant as termites have.Here, we quantified termite thermal limits to test the hypothesis that these physiological limits are wider in savanna termite species to facilitate their existence in savanna environments.We sampled termites directly from mound structures, across an environmental gradient in Ghana, ranging from wet tropical forest through to savanna. At each location, we quantified both the Critical Thermal Maxima (CTmax) and the Critical Thermal Minima (CTmin) of all the most abundant mound‐building Termitidae species in the study areas. We modelled the thermal limits in two separate mixed‐effects models against canopy cover at the mound, temperature and rainfall, as fixed effects, with sampling location as a random intercept.For both CTmax and CTmin, savanna species had significantly more extreme thermal limits than forest species. Between and within environments, areas with higher amounts of canopy cover were significantly associated with lower CTmax values of the termite colonies. CTmin was significantly positively correlated with rainfall. Temperature was retained in both models; however, it did not have a significant relationship in either. Sampling location explained a large proportion of the residual variation, suggesting there are other environmental factors that could influence termite thermal limits.Our results suggest that savanna termite species have wider thermal limits than forest species. These physiological differences, in conjunction with other behavioural adaptations, are likely to have enabled termites to cope with the more extreme environmental conditions found in savanna environments and facilitated their expansion into open tropical environments. [ABSTRACT FROM AUTHOR]

Fire is a fundamental process in savannas and is widely used for management. Pyrodiversity, variation in local fire characteristics, has been proposed as a driver of biodiversity although empirical evidence is equivocal. Using a new measure of pyrodiversity (Hempson et al.), we undertook the first continent-wide assessment of how pyrodiversity affects biodiversity in protected areas across African savannas. The influence of pyrodiversity on bird and mammal species richness varied with rainfall: strongest support for a positive effect occurred in wet savannas (> 650 mm/year), where species richness increased by 27% for mammals and 40% for birds in the most pyrodiverse regions. Range-restricted birds were most increased by pyrodiversity, suggesting the diversity of fire regimes increases the availability of rare niches. Our findings are significant because they explain the conflicting results found in previous studies of savannas. We argue that managing savanna landscapes to increase pyrodiversity is especially important in wet savannas. [ABSTRACT FROM AUTHOR]

Questions Termite mounds of the genus Macrotermes are prominent features in African savannas, forming nutrient hotspots that support greater plant diversity, which is of higher nutritional value than the surrounding savanna matrix. However, little is known about grass communities on and around mounds or how the functional importance of mounds varies across sites. As mean annual rainfall increases, savannas in southern Africa become increasingly dystrophic through increased denitrification (including pyrodenitrification) and the leaching of soil nutrients. The functional importance of mounds is concomitantly expected to increase as the difference in foliar nutrient levels between mounds and the savanna matrix increases. We tested this prediction on grass communities across a rainfall gradient to: (i) determine the degree to which grass assemblages differ between termite mounds and the savanna matrix; (ii) determine the spatial extent to which mounds influence grass communities; and (iii) investigate whether these patterns differ across savanna types. Location Kruger National Park, South Africa. Methods Grass communities were surveyed at three savanna sites differing in mean annual rainfall (550-750 mm·yr−1). Grass diversity and tissue nitrogen concentrations were measured on and off termite mounds and along transects away from mounds in order to calculate the spatial influence of termite mounds on savanna grass communities. Using termite mound densities estimated from airborne Li DAR, we up-scaled field-based results to determine the percentage of the landscape influenced by Macrotermes termite activity. Results Although species richness of grasses was lower on mounds than in the savanna matrix, the assemblage composition varied significantly, with higher nutrient concentrations in grasses located on mounds. This pattern became more distinct with increasing rainfall. The spatial extent of these nutrient-rich grasses also differed across the rainfall gradient, with a larger sphere of influence around mounds in wetter areas. Mounds distinctly altered grass communities over ca. 2% of the entire landscape. Conclusions Our results show that Macrotermes mounds are important components of savanna heterogeneity, and reveal that the functional importance of mounds increases with increasing rainfall. [ABSTRACT FROM AUTHOR]

According to the pyrodiversity paradigm, a wide range of fire regimes is required to maintain biodiversity in fire-prone landscapes. However, the requisite level of pyrodiversity has seldom been tested and may actually be very low., Here, we examine the sensitivity of tropical savanna ants to variation in fire regimes using results from a long-term fire experiment near Darwin, Australia. Six experimental fire regimes, with varying fire frequency and seasonality, have been applied to 18 one-ha plots in three replicated blocks since 2004, with ants sampled prior to experimental burning and then annually after up to 2009. Our primary focus is on the extent to which different patterns of ant richness and composition are associated with each of the six treatment regimes, or whether there is such high overlap that differences only become apparent when experimental treatments are grouped to provide strongly contrasting fire regimes., When treating each of the six fire treatments separately, we were unable to detect a significant influence of fire on any ant community response variable. We were only able to detect significant ant responses when we grouped the experimental treatments into two contrasting fire frequency classes, low (burnt at most once over the 5 years) vs. high (burnt every 1 or 2 years). Even then, these responses were only evident after 3 years of fire treatment., Our findings demonstrate that ant communities have very high resilience in relation to fire, with differences evident only between strongly contrasting regimes. Such resilience appears to be characteristic of savanna ants throughout the world., Synthesis and applications. A large range of finely tuned fire regimes is unlikely to promote regional ant diversity. Rather, only very limited pyrodiversity (a combination of frequently and infrequently burnt areas) would appear to be sufficient for maintaining diversity at a landscape scale. It is important that fire management for biodiversity conservation focuses on the demonstrated requirements of target species, rather than be based on an assumption that 'pyrodiversity begets biodiversity'. [ABSTRACT FROM AUTHOR]