The diversity of social complexity in termites.

Sociality underpins major evolutionary transitions and significantly influences the structure and function of complex ecosystems. Social insects, seen as the pinnacle of sociality, have traits like obligate sterility that are considered 'master traits', used as single phenotypic measures of this complexity. However, evidence is mounting that completely aligning both phenotypic and evolutionary social complexity, and having obligate sterility central to both, is erroneous. We hypothesize that obligate and functional sterility are insufficient in explaining the diversity of phenotypic social complexity in social insects. To test this, we explore the relative importance of these sterility traits in an understudied but diverse taxon: the termites. We compile the largest termite social complexity dataset to date, using specimen and literature data. We find that although functional and obligate sterility explain a significant proportion of variance, neither trait is an adequate singular proxy for the phenotypic social complexity of termites. Further, we show both traits have only a weak association with the other social complexity traits within termites. These findings have ramifications for our general comprehension of the frameworks of phenotypic and evolutionary social complexity, and their relationship with sterility.

Forest disturbance increases functional diversity but decreases phylogenetic diversity of an arboreal tropical ant community.

Tropical rainforest trees host a diverse arthropod fauna that can be characterised by their functional diversity (FD) and phylogenetic diversity (PD). Human disturbance degrades tropical forests, often coinciding with species invasion and altered assembly that leads to a decrease in FD and PD. Tree canopies are thought to be particularly vulnerable, but rarely investigated. Here, we studied the effects of forest disturbance on an ecologically important invertebrate group, the ants, in a lowland rainforest in New Guinea. We compared an early successional disturbed plot (secondary forest) to an old-growth plot (primary forest) by exhaustively sampling their ant communities in a total of 852 trees. We expected that for each tree community (1) disturbance would decrease FD and PD in tree-dwelling ants, mediated through species invasion. (2) Disturbance would decrease ant trait variation due to a more homogeneous environment. (3) The main drivers behind these changes would be different contributions of true tree-nesting species and visiting species. We calculated FD and PD based on a species-level phylogeny and 10 ecomorphological traits. Furthermore, we assessed by data exclusion the influence of species, which were not nesting in individual trees (visitors) or only nesting species (nesters), and of non-native species on FD and PD. Primary forests had higher ant species richness and PD than secondary forest. However, we consistently found increased FD in secondary forest. This pattern was robust even if we decoupled functional and phylogenetic signals, or if non-native ant species were excluded from the data. Visitors did not contribute strongly to FD, but they increased PD and their community weighted trait means often varied from nesters. Moreover, all community-weighted trait means changed after forest disturbance. Our finding of contradictory FD and PD patterns highlights the importance of integrative measures of diversity. Our results indicate that the tree community trait diversity is not negatively affected, but possibly even enhanced by disturbance. Therefore, the functional diversity of arboreal ants is relatively robust when compared between old-growth and young trees. However, further study with higher plot-replication is necessary to solidify and generalise our findings.

Assessing the healthiness of menus of all out-of-home food outlets and its socioeconomic patterns in Great Britain.

Food environment research predominantly focuses on the spatial distribution of out-of-home food outlets. However, the healthiness of food choices available within these outlets has been understudied, largely due to resource constraints. In this study, we propose an innovative, low-resource approach to characterise the healthiness of out-of-home food outlets at scale. Menu healthiness scores were calculated for food outlets on JustEat, and a deep learning model was trained to predict these scores for all physical out-of-home outlets in Great Britain, based on outlet names. Our findings highlight the "double burden" of the unhealthy food environment in deprived areas where there tend to be more out-of-home food outlets, and these outlets tend to be less healthy. This methodological advancement provides a nuanced understanding of out-of-home food environments, with potential for automation and broad geographic application.

A simple method to account for thermal boundary layers during the estimation of CTmax in small ectotherms.

As temperatures rise, understanding how ectotherms will become impacted by thermal stress is of critical importance. In this context, many researchers quantify critical temperatures - these are the upper (CTmax) and lower (CTmin) thermal limits at which organisms can no longer function. Most studies estimate CTs using bath-based methods where organisms are submerged within a set thermal environment. Plate-based methods (i.e. hot plates), however, offer huge opportunity for automation and are readily available in many lab settings. Plates, however, generate a unidirectional thermal boundary layer above their surface which means that the temperatures experienced by organisms of different sizes is different. This boundary layer effect can bias estimates of critical temperatures. Here, we test the hypothesis that biases in critical temperature estimation on hot plates are driven by organism height. We also quantify the composition of the boundary layer in order to correct for these biases. We assayed four differently sized species of UK ants for their CTmax in dry baths (with no boundary layer) and on hot plates (with a boundary layer). We found that hot plates overestimated the CTmax values of the different ants, and that this overestimate was larger for taller species. By statistically modelling the thickness of the thermal boundary layer, and combining with estimates of species height, we were able to correct this overestimation and eliminate methodological differences. Our study provides two main findings. First, we provide evidence that organism height is positively related to the bias present in plate-based estimates of CTmax. Second, we show that a relatively simple statistical model can correct for this bias. By using simple corrections for boundary layer effects, as we have done here, researchers could open up a new possibility space in the design and implementation of thermal tolerance assays using plates rather than restrictive dry or water baths.

dsSurvival 2.0: privacy enhancing survival curves for survival models in the federated DataSHIELD analysis system.

OBJECTIVE: Survival models are used extensively in biomedical sciences, where they allow the investigation of the effect of exposures on health outcomes. It is desirable to use diverse data sets in survival analyses, because this offers increased statistical power and generalisability of results. However, there are often challenges with bringing data together in one location or following an analysis plan and sharing results. DataSHIELD is an analysis platform that helps users to overcome these ethical, governance and process difficulties. It allows users to analyse data remotely, using functions that are built to restrict access to the detailed data items (federated analysis). Previous works have provided survival modelling functionality in DataSHIELD (dsSurvival package), but there is a requirement to provide functions that offer privacy enhancing survival curves that retain useful information. RESULTS: We introduce an enhanced version of the dsSurvival package which offers privacy enhancing survival curves for DataSHIELD. Different methods for enhancing privacy were evaluated for their effectiveness in enhancing privacy while maintaining utility. We demonstrated how our selected method could enhance privacy in different scenarios using real survival data. The details of how DataSHIELD can be used to generate survival curves can be found in the associated tutorial.

Ecological strategies of (pl)ants: Towards a world-wide worker economic spectrum for ants.

Current global challenges call for a rigorously predictive ecology. Our understanding of ecological strategies, imputed through suites of measurable functional traits, comes from decades of work that largely focussed on plants. However, a key question is whether plant ecological strategies resemble those of other organisms.Among animals, ants have long been recognised to possess similarities with plants: as (largely) central place foragers. For example, individual ant workers play similar foraging roles to plant leaves and roots and are similarly expendable. Frameworks that aim to understand plant ecological strategies through key functional traits, such as the 'leaf economics spectrum', offer the potential for significant parallels with ant ecological strategies.Here, we explore these parallels across several proposed ecological strategy dimensions, including an 'economic spectrum', propagule size-number trade-offs, apparency-defence trade-offs, resource acquisition trade-offs and stress-tolerance trade-offs. We also highlight where ecological strategies may differ between plants and ants. Furthermore, we consider how these strategies play out among the different modules of eusocial organisms, where selective forces act on the worker and reproductive castes, as well as the colony.Finally, we suggest future directions for ecological strategy research, including highlighting the availability of data and traits that may be more difficult to measure, but should receive more attention in future to better understand the ecological strategies of ants. The unique biology of eusocial organisms provides an unrivalled opportunity to bridge the gap in our understanding of ecological strategies in plants and animals and we hope that this perspective will ignite further interest. Read the free Plain Language Summary for this article on the Journal blog.

Testing the context dependence of ant nutrient preference across habitat strata and trophic levels in Neotropical biomes.

Animals are integrated into the wider ecosystem via their foraging and behavior. The compensation hypothesis predicts that animals target their foraging efforts (i) toward nutrients that are scarce in the environment and (ii) toward nutrients that are not present in the usual diet of species, which varies across trophic levels. Understanding how foraging for resources varies locally, such as across habitat strata, and trophic levels will help to elucidate the links between the local environment and communities to the ecological functions that animals mediate. We examined whether the relative resource use of ants varies consistently along a habitat strata gradient and across trophic levels across Neotropical biomes. We placed 4500 baited tubes, each containing one of five liquid resources (sugar, amino acid, lipid, sodium, and distilled water) in one of three habitat strata (subterranean, epigaeic, and arboreal) across 60 transects in Amazon, Atlantic Forest, Caatinga, Cerrado, Pampa, and Pantanal biomes. We assessed the relative resource use of all ants across the habitat strata and among two different trophic groups across biomes. The relative preference for sugar increased from subterranean to arboreal strata in all biomes, while the relative preference for lipids decreased at this gradient in five biomes. We also found that in general sugar-consuming ants foraged more for sugar and less for lipids than predatory ants across biomes. Conversely, we found no consistency across biomes in nutrient preference of amino acid and sodium across habitat strata or trophic levels. Overall, our results indicate sugar limitation in the arboreal stratum and lipid limitation on the ground across biomes and that the trophic level of ants strongly determines their foraging efforts-possibly because ants try to fix their dietary nutrient imbalances. Hence, our findings suggest strong local niche partitioning of sugar and lipid use across habitat strata and trophic levels and that other large spatial scale processes influence the local amino acid and sodium dynamics.

Os animais se integram nos ecossistemas pelos seus esforços e comportamento de forrageio. A hipótese da compensação prevê que os animais direcionam seus esforços de forrageio para (i) nutrientes que são escassos no ambiente e (ii) para nutrientes que não estão presentes em sua dieta, que variam entre os níveis tróficos. Assim, entender como a busca de recursos varia localmente entre os estratos de habitat e entre níveis tróficos, ajudará a entender as conexões entre o ambiente e suas comunidades com as funções ecológicas mediadas pelos animais. Avaliamos se o uso relativo de recursos das formigas varia consistentemente em um gradiente de estratos de habitat e em diferentes níveis tróficos nos biomas Neotropicais. Disponibilizamos um total de 4.500 tubos contendo um de cinco recursos líquidos (açúcar, aminoácido, lipídio, sódio e água destilada) em três estratos de habitat (subterrâneo, epigéico e arbóreo) em 60 transectos distribuídos nos biomas Amazônia, Mata Atlântica, Caatinga, Cerrado, Pampa e Pantanal. Avaliamos o uso relativo de recursos de todas as formigas ao longo dos estratos de habitat e entre dois grupos tróficos diferentes nos biomas. A preferência relativa por açúcar aumentou do estrato subterrâneo para o arbóreo em todos os biomas, enquanto a preferência relativa por lipídios diminuiu neste gradiente em cinco biomas. Também descobrimos que, em geral, as formigas consumidoras de açúcar procuram mais açúcar e menos lipídios do que as formigas predadoras nos biomas. Por outro lado, não encontramos consistência entre biomas na preferência nutricional de aminoácidos e sódio nos estratos de habitat ou níveis tróficos. No geral, nossos resultados indicam limitação de açúcar no estrato arbóreo e limitação de lipídios no solo em todos os biomas e que o nível trófico das formigas determina fortemente seus esforços de forrageamento - possivelmente porque as formigas tentam corrigir seus desequilíbrios de nutrientes na dieta. Portanto, nossas descobertas sugerem forte partição de nicho local de uso de açúcar e lipídios em estratos de habitat e níveis tróficos e que outros processos de grande escala espacial influenciam a dinâmica local de aminoácidos e sódio.

Monitoring the Nutrient Composition of Food Prepared Out-of-Home in the United Kingdom: Database Development and Case Study.

BACKGROUND: Hand transcribing nutrient composition data from websites requires extensive human resources and is prone to error. As a result, there are limited nutrient composition data on food prepared out of the home in the United Kingdom. Such data are crucial for understanding and monitoring the out-of-home food environment, which aids policy making. Automated data collection from publicly available sources offers a potential low-resource solution to address this gap. OBJECTIVE: In this paper, we describe the first UK longitudinal nutritional database of food prepared out of the home, MenuTracker. As large chains will be required to display calorie information on their UK menus from April 2022, we also aimed to identify which chains reported their nutritional information online in November 2021. In a case study to demonstrate the utility of MenuTracker, we estimated the proportions of menu items exceeding recommended energy and nutrient intake (eg, >600 kcal per meal). METHODS: We have collated nutrient composition data of menu items sold by large chain restaurants quarterly since March 2021. Large chains were defined as those with 250 employees or more (those covered by the new calorie labeling policy) or belonging to the top 100 restaurants based on sales volume. We developed scripts in Python to automate the data collection process from business websites. Various techniques were used to harvest web data and extract data from nutritional tables in PDF format. RESULTS: Automated Python programs reduced approximately 85% of manual work, totaling 500 hours saved for each wave of data collection. As of January 2022, MenuTracker has 76,405 records from 88 large out-of-home food chains at 4 different time points (ie, March, June, September, and December) in 2021. In constructing the database, we found that one-quarter (24.5%, 256/1043) of large chains, which are likely to be subject to the United Kingdom's calorie menu labeling regulations, provided their nutritional information online in November 2021. Across these chains, 24.7% (16,391/66,295) of menu items exceeded the UK government's recommendation of a maximum of 600 kcal for a single meal. Comparable figures were 46.4% (29,411/63,416) for saturated fat, 34.7% (21,964/63,388) for total fat, 17.6% (11,260/64,051) for carbohydrates, 17.8% (11,434/64,059) for sugar, and 35.2% (22,588/64,086) for salt. Furthermore, 0.7% to 7.1% of the menu items exceeded the maximum daily recommended intake for these nutrients. CONCLUSIONS: MenuTracker is a valuable resource that harnesses the power of data science techniques to use publicly available data online. Researchers, policy makers, and consumers can use MenuTracker to understand and assess foods available from out-of-home food outlets. The methods used in development are available online and can be used to establish similar databases elsewhere.

dsSurvival: Privacy preserving survival models for federated individual patient meta-analysis in DataSHIELD.

OBJECTIVE: Achieving sufficient statistical power in a survival analysis usually requires large amounts of data from different sites. Sensitivity of individual-level data, ethical and practical considerations regarding data sharing across institutions could be a potential challenge for achieving this added power. Hence we implemented a federated meta-analysis approach of survival models in DataSHIELD, where only anonymous aggregated data are shared across institutions, while simultaneously allowing for exploratory, interactive modelling. In this case, meta-analysis techniques to combine analysis results from each site are a solution, but an analytic workflow involving local analysis undertaken at individual studies hinders exploration. Thus, the aim is to provide a framework for performing meta-analysis of Cox regression models across institutions without manual analysis steps for the data providers. RESULTS: We introduce a package (dsSurvival) which allows privacy preserving meta-analysis of survival models, including the calculation of hazard ratios. Our tool can be of great use in biomedical research where there is a need for building survival models and there are privacy concerns about sharing data.

dsSynthetic: synthetic data generation for the DataSHIELD federated analysis system.

OBJECTIVE: Platforms such as DataSHIELD allow users to analyse sensitive data remotely, without having full access to the detailed data items (federated analysis). While this feature helps to overcome difficulties with data sharing, it can make it challenging to write code without full visibility of the data. One solution is to generate realistic, non-disclosive synthetic data that can be transferred to the analyst so they can perfect their code without the access limitation. When this process is complete, they can run the code on the real data. RESULTS: We have created a package in DataSHIELD (dsSynthetic) which allows generation of realistic synthetic data, building on existing packages. In our paper and accompanying tutorial we demonstrate how the use of synthetic data generated with our package can help DataSHIELD users with tasks such as writing analysis scripts and harmonising data to common scales and measures.

Associations between exploratory dietary patterns and incident type 2 diabetes: a federated meta-analysis of individual participant data from 25 cohort studies.

PURPOSE: In several studies, exploratory dietary patterns (DP), derived by principal component analysis, were inversely or positively associated with incident type 2 diabetes (T2D). However, findings remained study-specific, inconsistent and rarely replicated. This study aimed to investigate the associations between DPs and T2D in multiple cohorts across the world. METHODS: This federated meta-analysis of individual participant data was based on 25 prospective cohort studies from 5 continents including a total of 390,664 participants with a follow-up for T2D (3.8-25.0 years). After data harmonization across cohorts we evaluated 15 previously identified T2D-related DPs for association with incident T2D estimating pooled incidence rate ratios (IRR) and confidence intervals (CI) by Piecewise Poisson regression and random-effects meta-analysis. RESULTS: 29,386 participants developed T2D during follow-up. Five DPs, characterized by higher intake of red meat, processed meat, French fries and refined grains, were associated with higher incidence of T2D. The strongest association was observed for a DP comprising these food groups besides others (IRRpooled per 1 SD = 1.104, 95% CI 1.059-1.151). Although heterogeneity was present (I2 = 85%), IRR exceeded 1 in 18 of the 20 meta-analyzed studies. Original DPs associated with lower T2D risk were not confirmed. Instead, a healthy DP (HDP1) was associated with higher T2D risk (IRRpooled per 1 SD = 1.057, 95% CI 1.027-1.088). CONCLUSION: Our findings from various cohorts revealed positive associations for several DPs, characterized by higher intake of red meat, processed meat, French fries and refined grains, adding to the evidence-base that links DPs to higher T2D risk. However, no inverse DP-T2D associations were confirmed.

The response of ants to climate change.

Ants (Hymenoptera: Formicidae) are one of the most dominant terrestrial organisms worldwide. They are hugely abundant, both in terms of sheer numbers and biomass, on every continent except Antarctica and are deeply embedded within a diversity of ecological networks and processes. Ants are also eusocial and colonial organisms-their lifecycle is built on the labor of sterile worker ants who support a small number of reproductive individuals. Given the climatic changes that our planet faces, we need to understand how various important taxonomic groups will respond; this includes the ants. In this review, we synthesize the available literature to tackle this question. The answer is complicated. The ant literature has focused on temperature, and we broadly understand the ways in which thermal changes may affect ant colonies, populations, and communities. In general, we expect that species living in the Tropics, and in thermally variable microhabitats, such as the canopy and leaf litter environments, will be negatively impacted by rising temperatures. Species living in the temperate zones and those able to thermally buffer their nests in the soil or behaviorally avoid higher temperatures, however, are likely to be unaffected or may even benefit from a changed climate. How ants will respond to changes to other abiotic drivers associated with climate change is largely unknown, as is the detail on how altered ant populations and communities will ramify through their wider ecological networks. We discuss how eusociality may allow ants to adapt to, or tolerate, climate change in ways that solitary organisms cannot and we identify key geographic and phylogenetic hotspots of climate vulnerability and resistance. We finish by emphasizing the key research questions that we need to address moving forward so that we may fully appreciate how this critical insect group will respond to the ongoing climate crisis.

Socioeconomic inequalities in food outlet access through an online food delivery service in England: A cross-sectional descriptive analysis.

Online food delivery services facilitate 'online' access to food outlets selling food prepared away-from-home. Online food outlet access has not previously been investigated in England or across an entire country. Systematic differences in online food outlet access could exacerbate existing health inequalities, which is a public health concern. However, this is not known. Across postcode districts in England (n = 2118), we identified and described the number of food outlets and unique cuisine types accessible online from the market leader (Just Eat). We investigated associations with area-level deprivation using adjusted negative binomial regression models. We also compared the number of food outlets accessible online with the number physically accessible in the neighbourhood (1600m Euclidean buffers of postcode district geographic centroids) and investigated associations with deprivation using an adjusted general linear model. For each outcome, we predicted means and 95% confidence intervals. In November 2019, 29,232 food outlets were registered to accept orders online. Overall, the median number of food outlets accessible online per postcode district was 63.5 (IQR; 16.0-156.0). For the number of food outlets accessible online as a percentage of the number accessible within the neighbourhood, the median was 63.4% (IQR; 35.6-96.5). Analysis using negative binomial regression showed that online food outlet access was highest in the most deprived postcode districts (n = 106.1; 95% CI: 91.9, 120.3). The number of food outlets accessible online as a percentage of those accessible within the neighbourhood was highest in the least deprived postcode districts (n = 86.2%; 95% CI: 78.6, 93.7). In England, online food outlet access is socioeconomically patterned. Further research is required to understand how online food outlet access is related to using online food delivery services.

Associations of Total Legume, Pulse, and Soy Consumption with Incident Type 2 Diabetes: Federated Meta-Analysis of 27 Studies from Diverse World Regions.

BACKGROUND: The consumption of legumes is promoted as part of a healthy diet in many countries but associations of total and types of legume consumption with type 2 diabetes (T2D) are not well established. Analyses across diverse populations are lacking despite the availability of unpublished legume consumption data in prospective cohort studies. OBJECTIVE: To examine the prospective associations of total and types of legume intake with the risk of incident T2D. METHODS: Meta-analyses of associations between total legume, pulse, and soy consumption and T2D were conducted using a federated approach without physical data-pooling. Prospective cohorts were included if legume exposure and T2D outcome data were available and the cohort investigators agreed to participate. We estimated incidence rate ratios (IRRs) and CIs of associations using individual participant data including ≤42,473 incident cases among 807,785 adults without diabetes in 27 cohorts across the Americas, Eastern Mediterranean, Europe, and Western Pacific. Random-effects meta-analysis was used to combine effect estimates and estimate heterogeneity. RESULTS: Median total legume intake ranged from 0-140 g/d across cohorts. We observed a weak positive association between total legume consumption and T2D (IRR = 1.02, 95% CI: 1.01 to 1.04) per 20 g/d higher intake, with moderately high heterogeneity (I2 = 74%). Analysis by region showed no evidence of associations in the Americas, Eastern Mediterranean, and Western Pacific. The positive association in Europe (IRR = 1.05, 95% CI: 1.01 to 1.10, I2 = 82%) was mainly driven by studies from Germany, UK, and Sweden. No evidence of associations was observed for the consumption of pulses or soy. CONCLUSIONS: These findings suggest no evidence of an association of legume intakes with T2D in several world regions. The positive association observed in some European studies warrants further investigation relating to overall dietary contexts in which legumes are consumed, including accompanying foods which may be positively associated with T2D.

Automatic classification of takeaway food outlet cuisine type using machine (deep) learning.

BACKGROUND AND PURPOSE: Researchers have not disaggregated neighbourhood exposure to takeaway ('fast-') food outlets by cuisine type sold, which would otherwise permit examination of differential impacts on diet, obesity and related disease. This is partly due to the substantial resource challenge of manual classification of unclassified takeaway outlets at scale. We describe the development of a new model to automatically classify takeaway food outlets, by 10 major cuisine types, based on business name alone. MATERIAL AND METHODS: We used machine (deep) learning, and specifically a Long Short Term Memory variant of a Recurrent Neural Network, to develop a predictive model trained on labelled outlets (n = 14,145), from an online takeaway food ordering platform. We validated the accuracy of predictions on unseen labelled outlets (n = 4,000) from the same source. RESULTS: Although accuracy of prediction varied by cuisine type, overall the model (or 'classifier') made a correct prediction approximately three out of four times. We demonstrated the potential of the classifier to public health researchers and for surveillance to support decision-making, through using it to characterise nearly 55,000 takeaway food outlets in England by cuisine type, for the first time. CONCLUSIONS: Although imperfect, we successfully developed a model to classify takeaway food outlets, by 10 major cuisine types, from business name alone, using innovative data science methods. We have made the model available for use elsewhere by others, including in other contexts and to characterise other types of food outlets, and for further development.

Darker ants dominate the canopy: Testing macroecological hypotheses for patterns in colour along a microclimatic gradient.

Gradients in cuticle lightness of ectotherms have been demonstrated across latitudes and elevations. Three key hypotheses have been used to explain these macroecological patterns: the thermal melanism hypothesis (TMH), the melanism-desiccation hypothesis (MDH) and the photo-protection hypothesis (PPH). Yet the broad abiotic measures, such as temperature, humidity and UV-B radiation, typically used to detect these ecogeographical patterns, are a poor indication of the microenvironment experienced by small, cursorial ectotherms like ants. We tested whether these macroecological hypotheses explaining cuticle lightness held at habitat and microclimatic level by using a vertical gradient within a tropical rainforest. We sampled 222 ant species in lowland, tropical rainforest across four vertical strata: subterranean, ground, understory and canopy. We recorded cuticle lightness, abundance and estimated body size for each species and calculated an assemblage-weighted mean for cuticle lightness and body size for each vertical stratum. Abiotic variables (air temperature, vapour pressure deficit and UV-B radiation) were recorded for each vertical stratum. We found that cuticle lightness of ant assemblages was vertically stratified: ant assemblages in the canopy and understory were twice as dark as assemblages in ground and subterranean strata. Cuticle lightness was not correlated with body size, and there was no support for the TMH. Rather, we attribute this cline in cuticle lightness to a combination of the MDH and the PPH. Our findings indicate that broad macroecological patterns can be detected at much smaller spatial scales and that microclimatic gradients can shape trait variation, specifically the cuticle lightness of ants. These results suggest that any changes to microclimate that occur due to land-use change or climate warming could drive selection of ants based on cuticle colour, altering assemblage structure and potentially ecosystem functioning.

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.

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.

Dominance-diversity relationships in ant communities differ with invasion.

The relationship between levels of dominance and species richness is highly contentious, especially in ant communities. The dominance-impoverishment rule states that high levels of dominance only occur in species-poor communities, but there appear to be many cases of high levels of dominance in highly diverse communities. The extent to which dominant species limit local richness through competitive exclusion remains unclear, but such exclusion appears more apparent for non-native rather than native dominant species. Here we perform the first global analysis of the relationship between behavioral dominance and species richness. We used data from 1,293 local assemblages of ground-dwelling ants distributed across five continents to document the generality of the dominance-impoverishment rule, and to identify the biotic and abiotic conditions under which it does and does not apply. We found that the behavioral dominance-diversity relationship varies greatly, and depends on whether dominant species are native or non-native, whether dominance is considered as occurrence or relative abundance, and on variation in mean annual temperature. There were declines in diversity with increasing dominance in invaded communities, but diversity increased with increasing dominance in native communities. These patterns occur along the global temperature gradient. However, positive and negative relationships are strongest in the hottest sites. We also found that climate regulates the degree of behavioral dominance, but differently from how it shapes species richness. Our findings imply that, despite strong competitive interactions among ants, competitive exclusion is not a major driver of local richness in native ant communities. Although the dominance-impoverishment rule applies to invaded communities, we propose an alternative dominance-diversification rule for native communities.