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Functional redundancy, the potential for the functional role of one species to be fulfilled by another, is a key determinant of ecosystem viability. Scavenging transfers huge amount of energy through ecosystems and is, therefore, crucial for ecosystem viability and healthy ecosystem functioning. Despite this, relatively few studies have examined functional redundancy in scavenger communities. Moreover, the results of these studies are mixed and confined to a very limited range of habitat types and taxonomic groups. This study attempts to address this knowledge gap by conducting a field experiment in an undisturbed natural environment assessing functional roles and redundancy in vertebrate and invertebrate scavenging communities in a South African savanna. We used a large-scale field experiment to suppress ants in four 1 ha plots in a South African savanna and paired each with a control plot. We distributed three types of small food bait: carbohydrate, protein and seed, across the plots and excluded vertebrates from half the baits using cages. Using this combination of ant suppression and vertebrate exclusion, allowed us explore the contribution of non-ant invertebrates, ants and vertebrates in scavenging and also to determine whether either ants or vertebrates were able to compensate for the loss of one another. In this study, we found the invertebrate community carried out a larger proportion of overall scavenging services than vertebrates. Moreover, although scavenging was reduced when either invertebrates or vertebrates were absent, the presence of invertebrates better mitigated the functional loss of vertebrates than did the presence of vertebrates against the functional loss of invertebrates. There is a commonly held assumption that the functional role of vertebrate scavengers exceeds that of invertebrate scavengers; our results suggest that this is not true for small scavenging resources. Our study highlights the importance of invertebrates for securing healthy ecosystem functioning both now and into the future. We also build upon many previous studies which show that ants can have particularly large effects on ecosystem functioning. Importantly, our study suggests that scavenging in some ecosystems may be partly resilient to changes in the scavenging community, due to the potential for functional compensation by vertebrates and ants., (© 2024 The Authors. Journal of Animal Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society.)

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 (CT max ) and the Critical Thermal Minima (CT min ) 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 CT max and CT min , 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 CT max values of the termite colonies. CT min 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., (© 2022 The Authors. Journal of Animal Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society.)

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., (© 2021 British Ecological Society.)

Ants are diverse and abundant, especially in tropical ecosystems. They are often cited as the agents of key ecological processes, but their precise contributions compared with other organisms have rarely been quantified. Through the removal of food resources from the forest floor and subsequent transport to nests, ants play an important role in the redistribution of nutrients in rainforests. This is an essential ecosystem process and a key energetic link between higher trophic levels, decomposers and primary producers. We used the removal of carbohydrate, protein and seed baits as a proxy to quantify the contribution that ants, other invertebrates and vertebrates make to the redistribution of nutrients around the forest floor, and determined to what extent there is functional redundancy across ants, other invertebrate and vertebrate groups. Using a large-scale, field-based manipulation experiment, we suppressed ants from c. 1 ha plots in a lowland tropical rainforest in Sabah, Malaysia. Using a combination of treatment and control plots, and cages to exclude vertebrates, we made food resources available to: (i) the whole foraging community, (ii) only invertebrates and (iii) only non-ant invertebrates. This allowed us to partition bait removal into that taken by vertebrates, non-ant invertebrates and ants. Additionally, we examined how the non-ant invertebrate community responded to ant exclusion. When the whole foraging community had access to food resources, we found that ants were responsible for 52% of total bait removal whilst vertebrates and non-ant invertebrates removed the remaining 48%. Where vertebrates were excluded, ants carried out 61% of invertebrate-mediated bait removal, with all other invertebrates removing the remaining 39%. Vertebrates were responsible for just 24% of bait removal and invertebrates (including ants) collectively removed the remaining 76%. There was no compensation in bait removal rate when ants and vertebrates were excluded, indicating low functional redundancy between these groups. This study is the first to quantify the contribution of ants to the removal of food resources from rainforest floors and thus nutrient redistribution. We demonstrate that ants are functionally unique in this role because no other organisms compensated to maintain bait removal rate in their absence. As such, we strengthen a growing body of evidence establishing ants as ecosystem engineers, and provide new insights into the role of ants in maintaining key ecosystem processes. In this way, we further our basic understanding of the functioning of tropical rainforest ecosystems., (© 2017 The Authors. Journal of Animal Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society.)

In a world where even documenting species declines in tropical systems is challenging enough, Klimes et al. raise the bar by addressing the deceptively simple, yet inherently complex, question of why species richness is lower in secondary forests. Using the first plot-scale inventory of arboreal ant nests, combined with an innovative rarefaction technique, they quantify the relative importance of a range of successional factors and highlight the contribution of beta diversity to the higher richness in primary forest., (© 2012 The Author. Journal of Animal Ecology © 2012 British Ecological Society.)

1. Interspecific trade-offs are thought to facilitate coexistence between species at small spatial scales. The discovery-dominance trade-off, analogous to a competition-colonisation trade-off, is considered an important structuring mechanism in ant ecology. A trade-off between species' ability to discover food resources and to dominate them may explain how so many species apparently dependent on similar resources can coexist. 2. The discovery-dominance trade-off is thought to be broken by invasive species in enemy-free space or territorial species whose activity is fuelled by domination of carbohydrate resources. It may also be mediated by factors such as temperature and habitat structure. 3. We investigate the generality and form of the discovery-dominance relationship in an experiment using habitats of contrasting complexity across three continents. In addition, to assess how widespread the discovery-dominance trade-off is, we conducted a systematic review combining all empirical studies (published and from our experiment). 4. From our own fieldwork and meta-analyses of available studies, we find surprisingly little empirical support for the trade-off, with results indicating that mean effect sizes were either not significantly different from 0 or significantly positive. The trade-off was only detected in studies with parasitoids present. Additionally, experimental data from simple and complex habitats within each continent suggest that simple habitats may facilitate both food resource discovery and dominance. 5. We conclude that the discovery-dominance trade-off is the exception, rather than the rule. Instead, these abilities were commonly correlated. Real food resources provide many axes along which partitioning may occur, and discovery-dominance trade-offs are not a prerequisite for coexistence., (© 2011 The Authors. Journal of Animal Ecology © 2011 British Ecological Society.)

1. Competition is considered a key factor structuring many communities, and has been described as the 'hallmark' of ant ecology. Dominant species are thought to play a key role structuring local ant assemblages through competitive exclusion. 2. However, while there have been many studies demonstrating competitive exclusion and consequently reduced richness at baits, it is not clear whether such regulation of 'momentary' diversity at clumped food resources can scale up to the regulation of richness at the site or assemblage level. 3. In this study, ant assemblages were sampled in three different savanna habitats in South Africa using both baiting and pitfall trapping. 4. As has been found in previous studies, there was a unimodal relationship between dominant ants and species richness at baits, with high abundances of dominant ants regulating species richness through competitive exclusion. Analysis of pitfall samples revealed strong convergence in pattern, and results from null model co-occurrence analyses supported the findings. 5. The importance of competition in structuring local ant assemblages was, however, only apparent at one of the three savanna habitats suggesting that a full range of extreme environments is needed to produce the full unimodal relationship at the assemblage level. 6. Although the relative importance of competition varied with habitat type, the study demonstrated that in some habitats, dominant ants can control species richness at the assemblage level.