Functional compensation in a savanna scavenger community.

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.

Stronger regional biosecurity is essential to prevent hundreds of harmful biological invasions.

Biological invasions often transcend political boundaries, but the capacity of countries to prevent invasions varies. How this variation in biosecurity affects the invasion risks posed to the countries involved is unclear. We aimed to improve the understanding of how the biosecurity of a country influences that of its neighbours. We developed six scenarios that describe biological invasions in regions with contiguous countries. Using data from alien species databases, socio-economic and biodiversity data and species distribution models, we determined where 86 of 100 of the world's worst invasive species are likely to invade and have a negative impact in the future. Information on the capacity of countries to prevent invasions was used to determine whether such invasions could be avoided. For the selected species, we predicted 2,523 discrete invasions, most of which would have significant negative impacts and are unlikely to be prevented. Of these invasions, approximately a third were predicted to spread from the country in which the species first establishes to neighbouring countries where they would cause significant negative impacts. Most of these invasions are unlikely to be prevented as the country of first establishment has a low capacity to prevent invasions or has little incentive to do so as there will be no impact in that country. Regional biosecurity is therefore essential to prevent future harmful biological invasions. In consequence, we propose that the need for increased regional co-operation to combat biological invasions be incorporated in global biodiversity targets.

Thermoregulatory traits combine with range shifts to alter the future of montane ant assemblages.

Predicting and understanding the biological response to future climate change is a pressing challenge for humanity. In the 21st century, many species will move into higher latitudes and higher elevations as the climate warms. In addition, the relative abundances of species within local assemblages are likely to change. Both effects have implications for how ecosystems function. Few biodiversity forecasts, however, take account of both shifting ranges and changing abundances. We provide a novel analysis predicting the potential changes to assemblage-level relative abundances in the 21st century. We use an established relationship linking ant abundance and their colour and size traits to temperature and UV-B to predict future abundance changes. We also predict future temperature driven range shifts and use these to alter the available species pool for our trait-mediated abundance predictions. We do this across three continents under a low greenhouse gas emissions scenario (RCP2.6) and a business-as-usual scenario (RCP8.5). Under RCP2.6, predicted changes to ant assemblages by 2100 are moderate. On average, species richness will increase by 26%, while species composition and relative abundance structure will be 26% and 30% different, respectively, compared with modern assemblages. Under RCP8.5, however, highland assemblages face almost a tripling of species richness and compositional and relative abundance changes of 66% and 77%. Critically, we predict that future assemblages could be reorganized in terms of which species are common and which are rare: future highland assemblages will not simply comprise upslope shifts of modern lowland assemblages. These forecasts reveal the potential for radical change to montane ant assemblages by the end of the 21st century if temperature increases continue. Our results highlight the importance of incorporating trait-environment relationships into future biodiversity predictions. Looking forward, the major challenge is to understand how ecosystem processes will respond to compositional and relative abundance changes.

Low levels of intraspecific trait variation in a keystone invertebrate group.

The trait-based approach to ecology promises to provide a mechanistic understanding of species distributions and ecosystem functioning. Typically, trait analyses focus on average species trait values and assume that intraspecific variation is small or negligible. Recent work has shown, however, that intraspecific trait variation can often contribute substantially to total trait variation. Whilst many studies have investigated intraspecific variation in plants, very few have done so for invertebrates. There is no research on the level of intraspecific trait variation in ants (Hymenoptera: Formicidae), despite the fact that there is a growing body of literature using ant morphological trait data and demonstrating that these insects play important roles in many ecosystems and food webs. Here, we investigate the intraspecific variability of four commonly used ant morphological traits from 23 species from the Maloti-Drakensberg Mountains of southern Africa. In total, we measured 1145 different individuals and made 6870 trait measurements. Intraspecific variation accounted for only 1-4% of total trait variation for each of the four traits we analysed. We found no links between intraspecific variation, phylogeny and elevation. On average, six individuals generated robust species means but under biased sampling scenarios 20 individuals were needed. The low levels of intraspecific morphological variation that we find suggest that the approach of using mean species traits is valid, in this fauna at least. Regardless, we encourage ant trait ecologists to measure greater numbers of individuals, especially across gradients, to shed further light on intraspecific variation in this functionally important group of insects.

Woody encroachment slows decomposition and termite activity in an African savanna.

Woody encroachment can lead to a complete switch from open habitats to dense thickets, and has the potential to greatly alter the biodiversity and ecological functioning of grassy ecosystems across the globe. Plant litter decomposition is a critical ecosystem process fundamental to nutrient cycling and global carbon dynamics, yet little is known about how woody encroachment might alter this process. We compared grass decay rates of heavily encroached areas with adjacent nonencroached open areas in a semi-arid South African savanna using litterbags that allowed or excluded invertebrates. We also assessed the effect of woody encroachment on the activity of termites- dominant decomposer organisms in savanna systems. We found a significant reduction in decomposition rates within encroached areas, with litter taking twice as long to decay compared with open savanna areas. Moreover, invertebrates were more influential on grass decomposition in open areas and termite activity was substantially lower in encroached areas, particularly during the dry season when activity levels were reduced to almost zero. Our results suggest that woody encroachment created an unfavourable environment for invertebrates, and termites in particular, leading to decreased decomposition rates in these areas. We provide the first quantification of woody encroachment altering the functioning of African savanna ecosystems through the slowing of aboveground plant decomposition. Woody encroachment is intensifying across the globe, and our results suggest that substantial changes to the carbon balance and biodiversity of grassy biomes could occur.

Seasonal variation in the relative dominance of herbivore guilds in an African savanna.

African savannas are highly seasonal with a diverse array of both mammalian and invertebrate herbivores, yet herbivory studies have focused almost exclusively on mammals. We conducted a 2-yr exclosure experiment in South Africa's Kruger National Park to measure the relative impact of these two groups of herbivores on grass removal at both highly productive patches (termite mounds) and in the less productive savanna matrix. Invertebrate and mammalian herbivory was greater on termite mounds, but the relative importance of each group changed over time. Mammalian offtake was higher than invertebrates in the dry season, but can be eclipsed by invertebrates during the wet season when this group is more active. Our results demonstrate that invertebrates play a substantial role in savanna herbivory and should not be disregarded in attempts to understand the impacts of herbivory on ecosystems.

Border control for stowaway alien species should be prioritised based on variations in establishment debt.

Border control is one of the major approaches used by countries to limit the number of organisms introduced as stowaways. However, it is not feasible to inspect all passengers, cargo and vehicles entering a country, and so efforts need to be prioritised. Here we use South Africa as a case study to assess, based on tourism and trade data and climate matching techniques, the number of stowaway species that might be introduced ('colonisation pressure') and the likelihood that once introduced, these organisms will establish ('likelihood of establishment'). These results were used to explore how the number of species that are likely to establish ('establishment debt') varies across donor regions and seasons. A simple theoretical model was then used to compare four strategies for prioritising border control inspections: no prioritisation; based on colonisation pressure; based on likelihood of establishment; and based on both colonisation pressure and likelihood of establishment. Establishment debt was greatest in southern hemisphere spring and autumn when South Africa is climatically similar to northern hemisphere countries with which there are strong, consistent trade and tourism links (i.e. colonisation pressure varied little seasonally, but likelihood of establishment did vary across the seasons). Prioritising inspections based on both colonisation pressure and the likelihood of establishment was clearly the most effective strategy, with this strategy detecting at least 6% more potential invaders than the other strategies. While there are many practical limitations to the implementation of such prioritised inspection strategies, the results highlight the importance of national and regional studies of establishment debt.

Invasion trajectory of alien trees: the role of introduction pathway and planting history.

Global change is driving a massive rearrangement of the world's biota. Trajectories of distributional shifts are shaped by species traits, the recipient environment and driving forces with many of the driving forces directly due to human activities. The relative importance of each in determining the distributions of introduced species is poorly understood. We consider 11 Australian Acacia species introduced to South Africa for different reasons (commercial forestry, dune stabilization and ornamentation) to determine how features of the introduction pathway have shaped their invasion history. Projections from species distribution models (SDMs) were developed to assess how the reason for introduction influences the similarity between climatic envelopes in native and alien ranges. A lattice model for an idealized invasion was developed to assess the relative contribution of intrinsic traits and introduction dynamics on the abundance and extent over the course of simulated invasions. SDMs show that alien populations of ornamental species in South Africa occupy substantially different climate space from their native ranges, whereas species introduced for forestry occupy a similar climate space in native and introduced ranges. This may partly explain the slow spread rates observed for some alien ornamental plants. Such mismatches are likely to become less pronounced with the current drive towards 'eco gardens' resulting in more introductions of ornamental species with a close climate match between native and newly introduced regions. The results from the lattice model showed that the conditions associated with the introduction pathway (especially introduction pressure) dominate early invasion dynamics. The placement of introduction foci in urban areas limited the extent and abundance of invasive populations. Features of introduction events appear to initially mask the influence of intrinsic species traits on invasions and help to explain the relative success of species introduced for different purposes. Introduction dynamics therefore can have long-lasting influences on the outcomes of species redistributions, and must be explicitly considered in management plans.

An inventory of native-alien populations in South Africa.

Species can be both native and alien to a given administrative region. Here we present the first consolidated inventory of these 'native-alien populations' for South Africa, and provide an overview of the data it contains. To gather data, literature searches were performed and experts were consulted both directly and via an on-line survey. Putative native-alien populations were then scored based on a newly developed protocol. The final inventory contains information on 77 native species from 49 families across nine classes that have formed 132 native-alien populations across the terrestrial, freshwater, and marine environments. The phenomenon is rare when compared to the prevalence of related phenomena, such as alien species introduced from other countries (2033 alien species in South Africa), but is under-reported. However, they pose a specific problem for regulators and managers and their importance will likely increase with global change. These data will be integrated with an existing alien species list and, we hope, will provide a useful foundation to address the issue. We encourage those working on biodiversity to contribute more records.

Scaling artificial light at night and disease vector interactions into socio-ecological systems: a conceptual appraisal.

There is burgeoning interest in how artificial light at night (ALAN) interacts with disease vectors, particularly mosquitoes. ALAN can alter mosquito behaviour and biting propensity, and so must alter disease transfer rates. However, most studies to date have been laboratory-based, and it remains unclear how ALAN modulates disease vector risk. Here, we identify five priorities to assess how artificial light can influence disease vectors in socio-ecological systems. These are to (i) clarify the mechanistic role of artificial light on mosquitoes, (ii) determine how ALAN interacts with other drivers of global change to influence vector disease dynamics across species, (iii) determine how ALAN interacts with other vector suppression strategies, (iv) measure and quantify the impact of ALAN at scales relevant for vectors, and (v) overcome the political and social barriers in implementing it as a novel vector suppression strategy. These priorities must be addressed to evaluate the costs and benefits of employing appropriate ALAN regimes in complex socio-ecological systems if it is to reduce disease burdens, especially in the developing world. This article is part of the theme issue 'Light pollution in complex ecological systems'.

Co-existence Between Two Leaf-Feeding Biological Control Agents of Lantana camara Alters Their Herbivory Under Semi-field Conditions.

Interaction between two biological control agents released against Lantana camara L. (sensu lato) (Verbenaceae) was studied in replicated semi-field plots. Caged plants under semi-field conditions were inoculated with Uroplata girardi Pic (Coleoptera: Chrysomelidae) and Ophiomyia camarae Spencer (Diptera: Agromyzidae), either alone or in combination, to investigate the extent to which co-infestation of the two agents affects the reproductive capacity and growth of their host. At the end of the trial, both single and combined attacks by the two agents had no effect on stem diameter, stem height, and canopy width. However, uncaged control plants were heavily attacked by Teleonemia scrupulosa Stål (Hemiptera: Tingidae), and therefore became significantly shorter than all the caged plants in all the treatments. When confined alone, feeding damage by O. camarae resulted in higher reduction of fruit and flower biomass relative to that caused by U. girardi alone. However, when confined alone, U. girardi caused higher reductions in leaf density and fruit biomass than when combined with O. camarae. Single attack by O. camarae caused higher reduction in flower biomass than simultaneous attack by both agents. Above-ground biomass of all single and combined treatment plants were significantly lower than those of the caged control plants. Uncaged control plants exposed to heavy attack by T. scrupulosa did not produce flowers and fruits, and their above-ground biomass was significantly lower than those of caged control plants. Overall, the study showed that simultaneous attack by the two herbivores alters their herbivory, thereby affecting reproductive capacity and growth of their host.

Prioritising surveillance for alien organisms transported as stowaways on ships travelling to South Africa.

The global shipping network facilitates the transportation and introduction of marine and terrestrial organisms to regions where they are not native, and some of these organisms become invasive. South Africa was used as a case study to evaluate the potential for shipping to contribute to the introduction and establishment of marine and terrestrial alien species (i.e. establishment debt) and to assess how this varies across shipping routes and seasons. As a proxy for the number of species introduced (i.e. 'colonisation pressure') shipping movement data were used to determine, for each season, the number of ships that visited South African ports from foreign ports and the number of days travelled between ports. Seasonal marine and terrestrial environmental similarity between South African and foreign ports was then used to estimate the likelihood that introduced species would establish. These data were used to determine the seasonal relative contribution of shipping routes to South Africa's marine and terrestrial establishment debt. Additionally, distribution data were used to identify marine and terrestrial species that are known to be invasive elsewhere and which might be introduced to each South African port through shipping routes that have a high relative contribution to establishment debt. Shipping routes from Asian ports, especially Singapore, have a particularly high relative contribution to South Africa's establishment debt, while among South African ports, Durban has the highest risk of being invaded. There was seasonal variation in the shipping routes that have a high relative contribution to the establishment debt of the South African ports. The presented method provides a simple way to prioritise surveillance effort and our results indicate that, for South Africa, port-specific prevention strategies should be developed, a large portion of the available resources should be allocated to Durban, and seasonal variations and their consequences for prevention strategies should be explored further.

Contrasting species and functional beta diversity in montane ant assemblages.

AIM: Beta diversity describes the variation in species composition between sites and can be used to infer why different species occupy different parts of the globe. It can be viewed in a number of ways. First, it can be partitioned into two distinct patterns: turnover and nestedness. Second, it can be investigated from either a species identity or a functional-trait point of view. We aim to document for the first time how these two aspects of beta diversity vary in response to a large environmental gradient. LOCATION: Maloti-Drakensberg Mountains, southern Africa. METHODS: We sampled ant assemblages along an extensive elevational gradient (900-3000 m a.s.l.) twice yearly for 7 years, and collected functional-trait information related to the species' dietary and habitat-structure preferences. We used recently developed methods to partition species and functional beta diversity into their turnover and nestedness components. A series of null models were used to test whether the observed beta diversity patterns differed from random expectations. RESULTS: Species beta diversity was driven by turnover, but functional beta diversity was composed of both turnover and nestedness patterns at different parts of the gradient. Null models revealed that deterministic processes were likely to be responsible for the species patterns but that the functional changes were indistinguishable from stochasticity. MAIN CONCLUSIONS: Different ant species are found with increasing elevation, but they tend to represent an increasingly nested subset of the available functional strategies. This finding is unique and narrows down the list of possible factors that control ant existence across elevation. We conclude that diet and habitat preferences have little role in structuring ant assemblages in montane environments and that some other factor must be driving the non-random patterns of species turnover. This finding also highlights the importance of distinguishing between different kinds of beta diversity.

Introduced and invasive cactus species: a global review.

Understanding which species are introduced and become invasive, and why, are central questions in invasion science. Comparative studies on model taxa have provided important insights, but much more needs to be done to unravel the context dependencies of these findings. The cactus family (Cactaceae), one of the most popular horticultural plant groups, is an interesting case study. Hundreds of cactus species have been introduced outside their native ranges; a few of them are among the most damaging invasive plant species in the world. We reviewed the drivers of introductions and invasions in the family and seek insights that can be used to minimize future risks. We compiled a list of species in the family and determined which have been recorded as invasive. We also mapped current global distributions and modelled the potential global distributions based on distribution data of known invasive taxa. Finally, we identified whether invasiveness is phylogenetically clustered for cacti and whether particular traits are correlated with invasiveness. Only 57 of the 1922 cactus species recognized in this treatment have been recorded as invasive. There are three invasion hotspots: South Africa (35 invasive species recorded), Australia (26 species) and Spain (24 species). However, there are large areas of the world with climates suitable for cacti that are at risk of future invasion-in particular, parts of China, eastern Asia and central Africa. The invasive taxa represent an interesting subset of the total species pool. There is a significant phylogenetic signal: invasive species occur in 2 of the 3 major phylogenetic clades and in 13 of the 130 genera. This phylogenetic signal is not driven by human preference, i.e. horticultural trade, but all invasive species are from 5 of the 12 cactus growth forms. Finally, invasive species tend to have significantly larger native ranges than non-invasive species, and none of the invasive species are of conservation concern in their native range. These results suggest fairly robust correlates of invasiveness that can be used for proactive management and risk assessments.

Spatially-explicit estimation of geographical representation in large-scale species distribution datasets.

Much ecological research relies on existing multispecies distribution datasets. Such datasets, however, can vary considerably in quality, extent, resolution or taxonomic coverage. We provide a framework for a spatially-explicit evaluation of geographical representation within large-scale species distribution datasets, using the comparison of an occurrence atlas with a range atlas dataset as a working example. Specifically, we compared occurrence maps for 3773 taxa from the widely-used Atlas Florae Europaeae (AFE) with digitised range maps for 2049 taxa of the lesser-known Atlas of North European Vascular Plants. We calculated the level of agreement at a 50-km spatial resolution using average latitudinal and longitudinal species range, and area of occupancy. Agreement in species distribution was calculated and mapped using Jaccard similarity index and a reduced major axis (RMA) regression analysis of species richness between the entire atlases (5221 taxa in total) and between co-occurring species (601 taxa). We found no difference in distribution ranges or in the area of occupancy frequency distribution, indicating that atlases were sufficiently overlapping for a valid comparison. The similarity index map showed high levels of agreement for central, western, and northern Europe. The RMA regression confirmed that geographical representation of AFE was low in areas with a sparse data recording history (e.g., Russia, Belarus and the Ukraine). For co-occurring species in south-eastern Europe, however, the Atlas of North European Vascular Plants showed remarkably higher richness estimations. Geographical representation of atlas data can be much more heterogeneous than often assumed. Level of agreement between datasets can be used to evaluate geographical representation within datasets. Merging atlases into a single dataset is worthwhile in spite of methodological differences, and helps to fill gaps in our knowledge of species distribution ranges. Species distribution dataset mergers, such as the one exemplified here, can serve as a baseline towards comprehensive species distribution datasets.

Interaction between Uroplata girardi (Coleoptera: Chrysomelidae) and Ophiomyia camarae (Diptera: Agromyzidae) on a shared host Lantana camara (Verbenaceae).

Multiple releases of insect agents intended to target a single plant pest species could result in competitive interactions that in turn might affect the community structure of the phytophagous insects. Two leaf-feeding biological control agents, Uroplata girardi Pic (Coleoptera: Chrysomelidae) and Ophiomyia camarae Spencer (Dipetera: Agromyzidae), were released against the weed Lantana camara L. (Verbenaceae) in South Africa in the 1970s and 2001, respectively. Since the population explosion of O. camarae in 2005, a decline of U. girardi populations had been observed in KwaZulu-Natal (KZN) humid coast, leading to speculation that negative interaction may be operating between the agents. The study therefore was conducted to determine the competitive effect of O. camarae on U. girardi. The study showed that 76% of O. camarae larval mines were formed on uninfested (clean) compared with only 24% formed on U. girardi-infested leaves, suggesting that the fly chose to lay more eggs on clean leaves. Almost the same number of U. girardi larval mines was formed on both O. camarae-infested and clean leaves, indicating that U. girardi females in this case oviposited indiscriminately on the two types of leaves. The survival of U. girardi was 53.8% when reared on clean leaves compared with only 14.6% survival on O. camarae-infested leaves. At the end of the sampling period, densities of U. girardi was over two times higher in single-species than in combined-species treatment. Releasing both agents together did not significantly affect O. camarae densities during the sampling period. In the field, O. camarae densities increased rapidly from spring to autumn, whereas those of U. girardi remained consistently low during the same period. The bias toward oviposition on clean leaves in O. camarae enables its larvae to avoid unfavorable encounters with U. girardi larvae, thus enhancing its development and survival. The apparent inability of U. girardi to distinguish between suitable and unsuitable leaves for oviposition could compromise the fitness of this beetle, and this could explain the suppression of U. girardi populations during summer when O. camarae populations begin to increase rapidly. This study provides evidence for an asymmetric interaction between two introduced agents, and therefore highlights the importance of conducting interaction studies on agents with extensive niche overlap before their release into the environment.