Population demography maintains biogeographic boundaries.

Global biodiversity is organised into biogeographic regions that comprise distinct biotas. The contemporary factors maintaining differences in species composition between regions are poorly understood. Given evidence that populations with sufficient genetic variation can adapt to fill new habitats, it is surprising that more homogenisation of species assemblages across regions has not occurred. Theory suggests that expansion across biogeographic regions could be limited by reduced adaptive capacity due to demographic variation along environmental gradients, but this possibility has not been empirically explored. Using three independently curated data sets describing continental patterns of mammalian demography and population genetics, we show that populations near biogeographic boundaries have lower effective population sizes and genetic diversity, and are more genetically differentiated. These patterns are consistent with reduced adaptive capacity in areas where one biogeographic region transitions into the next. That these patterns are replicated across mammals suggests they are stable and generalisable in their contribution to long-term limits on biodiversity homogenisation. Understanding the contemporary processes that maintain compositional differences among regional biotas is crucial for our understanding of the current and future organisation of global biodiversity.

Warm and arid regions of the world are hotspots of superorganism complexity.

Biologists have long been fascinated by the processes that give rise to phenotypic complexity of organisms, yet whether there exist geographical hotspots of phenotypic complexity remains poorly explored. Phenotypic complexity can be readily observed in ant colonies, which are superorganisms with morphologically differentiated queen and worker castes analogous to the germline and soma of multicellular organisms. Several ant species have evolved 'worker polymorphism', where workers in a single colony show quantifiable differences in size and head-to-body scaling. Here, we use 256 754 occurrence points from 8990 ant species to investigate the geography of worker polymorphism. We show that arid regions of the world are the hotspots of superorganism complexity. Tropical savannahs and deserts, which are typically species-poor relative to tropical or even temperate forests, harbour the highest densities of polymorphic ants. We discuss the possible adaptive advantages that worker polymorphism provides in arid environments. Our work may provide a window into the environmental conditions that promote the emergence of highly complex phenotypes.

Temperature drives caste-specific morphological clines in ants.

The morphology of organisms relates to most aspects of their life history and autecology. As such, elucidating the drivers of morphological variation along environmental gradients might give insight into processes limiting species distributions. In eusocial organisms, the concept of morphology is more complex than in solitary organisms. Eusocial insects such as ants exhibit drastic morphological differences between reproductive and worker castes. How environmental selection operates on the morphology of each caste, and whether caste-specific selection has fitness consequences is largely unknown, but is potentially crucial to understand what limits ant species' distributions. Here we aimed to examine whether ant shape and body size covaries with climate at the scale of an entire continent, and whether such relationship might be caste specific. We used 26,472 georeferenced morphometric measurements from 2,206 individual ants belonging to 32 closely related North American species in the genus Formica to assess how ant morphology relates to geographic variation in the abiotic environment. Although precipitation and seasonality explained some of the geographic variation in morphology, temperature was the best predictor. Specifically, geographic variation in body size was positively related to temperature, meaning that ants are smaller in cold than in warm environments. Moreover, the strength of the relationship between size and temperature was stronger for the reproductive castes (i.e. queens and males) than for the worker caste. The shape of workers and males also varied along these large-scale abiotic gradients. Specifically, the relative length of workers' legs, thoraxes and antennae positively related to temperature, meaning that they had shorter appendages in cold environments. In contrast, males had smaller heads, but larger thoraxes in more seasonal environments. Overall, our results suggest that geographic variation in ambient temperature influences the morphology of ants, but that the strength of this effect is caste specific. In conclusion, whereas ant ecology has traditionally focused on workers, our study shows that considering the ecology of the reproductive castes is imperative to move forward in this field.

Ant community response to disturbance: A global synthesis.

In Focus: Andersen, A. N. (2019). Responses of ant communities to disturbance: Five principles for understanding the disturbance dynamics of a globally dominant faunal group. Journal of Animal Ecology 88, 350-362. Disturbance is a key driver of ecosystem dynamics. Whereas plant community responses to disturbance are relatively well understood, the same does not hold for animals. With rapid changes affecting our world's ecosystems, predicting the response of important ecological groups to ongoing disturbance should be a focus. In particular, ants are ecosystem engineers that create habitats for other organisms and have a crucial role to play in nutrient cycling. Nevertheless, our understanding of ant community response to disturbance is, at best, fragmented. Moreover, how ant communities respond to disturbance on a global scale appears highly idiosyncratic. The perspective article by Andersen (Journal of Animal Ecology 88, 350-362.) proposes five general principles that can help elucidate ant community response to disturbance. Specifically, this synthesis deepens our understanding of how contemporary disturbances, ecological processes and the evolutionary and biogeographic history of lineages interact to influence ant community structure.

A global database of ant species abundances.

What forces structure ecological assemblages? A key limitation to general insights about assemblage structure is the availability of data that are collected at a small spatial grain (local assemblages) and a large spatial extent (global coverage). Here, we present published and unpublished data from 51 ,388 ant abundance and occurrence records of more than 2,693 species and 7,953 morphospecies from local assemblages collected at 4,212 locations around the world. Ants were selected because they are diverse and abundant globally, comprise a large fraction of animal biomass in most terrestrial communities, and are key contributors to a range of ecosystem functions. Data were collected between 1949 and 2014, and include, for each geo-referenced sampling site, both the identity of the ants collected and details of sampling design, habitat type, and degree of disturbance. The aim of compiling this data set was to provide comprehensive species abundance data in order to test relationships between assemblage structure and environmental and biogeographic factors. Data were collected using a variety of standardized methods, such as pitfall and Winkler traps, and will be valuable for studies investigating large-scale forces structuring local assemblages. Understanding such relationships is particularly critical under current rates of global change. We encourage authors holding additional data on systematically collected ant assemblages, especially those in dry and cold, and remote areas, to contact us and contribute their data to this growing data set.

Process-Based Species Pools Reveal the Hidden Signature of Biotic Interactions Amid the Influence of Temperature Filtering.

A persistent challenge in ecology is to tease apart the influence of multiple processes acting simultaneously and interacting in complex ways to shape the structure of species assemblages. We implement a heuristic approach that relies on explicitly defining species pools and permits assessment of the relative influence of the main processes thought to shape assemblage structure: environmental filtering, dispersal limitations, and biotic interactions. We illustrate our approach using data on the assemblage composition and geographic distribution of hummingbirds, a comprehensive phylogeny and morphological traits. The implementation of several process-based species pool definitions in null models suggests that temperature-but not precipitation or dispersal limitation-acts as the main regional filter of assemblage structure. Incorporating this environmental filter directly into the definition of assemblage-specific species pools revealed an otherwise hidden pattern of phylogenetic evenness, indicating that biotic interactions might further influence hummingbird assemblage structure. Such hidden patterns of assemblage structure call for a reexamination of a multitude of phylogenetic- and trait-based studies that did not explicitly consider potentially important processes in their definition of the species pool. Our heuristic approach provides a transparent way to explore patterns and refine interpretations of the underlying causes of assemblage structure.

Climate mediates the effects of disturbance on ant assemblage structure.

Many studies have focused on the impacts of climate change on biological assemblages, yet little is known about how climate interacts with other major anthropogenic influences on biodiversity, such as habitat disturbance. Using a unique global database of 1128 local ant assemblages, we examined whether climate mediates the effects of habitat disturbance on assemblage structure at a global scale. Species richness and evenness were associated positively with temperature, and negatively with disturbance. However, the interaction among temperature, precipitation and disturbance shaped species richness and evenness. The effect was manifested through a failure of species richness to increase substantially with temperature in transformed habitats at low precipitation. At low precipitation levels, evenness increased with temperature in undisturbed sites, peaked at medium temperatures in disturbed sites and remained low in transformed sites. In warmer climates with lower rainfall, the effects of increasing disturbance on species richness and evenness were akin to decreases in temperature of up to 9°C. Anthropogenic disturbance and ongoing climate change may interact in complicated ways to shape the structure of assemblages, with hot, arid environments likely to be at greatest risk.

Linking environmental filtering and disequilibrium to biogeography with a community climate framework.

We present a framework to measure the strength of environmental filtering and disequilibrium of the species composition of a local community across time, relative to past, current, and future climates. We demonstrate the framework by measuring the impact of climate change on New World forests, integrating data for climate niches of more than 14000 species, community composition of 471 New World forest plots, and observed climate across the most recent glacial-interglacial interval. We show that a majority of communities have species compositions that are strongly filtered and are more in equilibrium with current climate than random samples from the regional pool. Variation in the level of current community disequilibrium can be predicted from Last Glacial Maximum climate and will increase with near-future climate change.

Niche filtering rather than partitioning shapes the structure of temperate forest ant communities.

An ever-increasing number of studies use tools from community phylogenetics to infer the processes underlying the assembly of communities. However, very few studies simultaneously use experimental approaches to characterize the ecological niches of species and directly assess the importance of these structuring processes. In this study, we developed an experimental approach for quantifying the use of four types of food resources and three habitat templets in temperate forest ant assemblages. We then used null models to assess whether niches overlapped more or less than expected by chance. Finally, we integrated comparative phylogenetic methods with experimental data on niche use to assess the degree of phylogenetic signal in several key components of the niche. We found that niche filtering, rather than partitioning, was the predominant structuring force. Niche filtering resulted from conservatism in habitat niches in evolutionary time and limitations in the availability of food resources in ecological time. Our study thus supports the idea that similarities in niches among species, rather than the differences, drive the assembly of ant communities.

Heterogeneous dispersal networks to improve biodiversity science.

Dispersal has a key role in shaping spatial patterns of biodiversity, yet its spatial heterogeneity is often overlooked in biodiversity analyses and management strategies. Properly parameterised heterogeneous dispersal networks capture the complex interplay between landscape structure and species-specific dispersal capacities. However, this heterogeneity is recurrently neglected when studying the processes underlying biodiversity variation. To address this gap, we introduce a conceptual framework detailing the fundamental processes driving dispersal heterogeneity and its effects on biodiversity dynamics. We propose methods to parameterise heterogeneous dispersal networks, facilitating their integration into commonly used quantitative frameworks for biodiversity analyses. By considering the architecture of heterogeneous dispersal networks, we demonstrate their critical role in guiding biodiversity management strategies.

Marine fishes experiencing high-velocity range shifts may not be climate change winners.

Climate change is driving the global redistribution of species. A common assumption is that rapid range shifts occur in tandem with overall stable or positive abundance trends throughout the range and thus these species may be considered as climate change 'winners'. However, although establishing the link between range shift velocities and population trends is crucial for predicting climate change impacts it has not been empirically tested. Using 2,572 estimates of changes in marine fish abundance spread across the world's oceans, we show that poleward range shifts are not necessarily associated with positive population trends. Species experiencing high-velocity range shifts seem to experience local population declines irrespective of the position throughout the species range. High range shift velocities of 17 km yr-1 are associated with a 50% decrease in population sizes over a period of 10 yr, which is dramatic compared to the overall stable population trends in non-shifting species. This pattern, however, mostly occurs in populations located in the poleward, colder, portion of the species range. The lack of a positive association between poleward range shift velocities and population trends at the coldest portion of the range contrasts with the view that rapid range shifts safeguard against local population declines. Instead, our work suggests that marine fishes experiencing rapid range shifts could be more vulnerable to climatic change and therefore should be carefully assessed for conservation status.

The evolution of plasticity at geographic range edges.

Phenotypic plasticity enables rapid responses to environmental change, and could facilitate range shifts in response to climate change. What drives the evolution of plasticity at range edges, and the capacity of range-edge individuals to be plastic, remain unclear. Here, we propose that accurately predicting when plasticity itself evolves or mediates adaptive evolution at expanding range edges requires integrating knowledge on the demography and evolution of edge populations. Our synthesis shows that: (i) the demography of edge populations can amplify or attenuate responses to selection for plasticity through diverse pathways, and (ii) demographic effects on plasticity are modified by the stability of range edges. Our spatially explicit synthesis for plasticity has the potential to improve predictions for range shifts with climate change.

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.

Strong influence of regional species pools on continent-wide structuring of local communities.

There is a long tradition in ecology of evaluating the relative contribution of the regional species pool and local interactions on the structure of local communities. Similarly, a growing number of studies assess the phylogenetic structure of communities, relative to that in the regional species pool, to examine the interplay between broad-scale evolutionary and fine-scale ecological processes. Finally, a renewed interest in the influence of species source pools on communities has shown that the definition of the source pool influences interpretations of patterns of community structure. We use a continent-wide dataset of local ant communities and implement ecologically explicit source pool definitions to examine the relative importance of regional species pools and local interactions for shaping community structure. Then we assess which factors underlie systematic variation in the structure of communities along climatic gradients. We find that the average phylogenetic relatedness of species in ant communities decreases from tropical to temperate regions, but the strength of this relationship depends on the level of ecological realism in the definition of source pools. We conclude that the evolution of climatic niches influences the phylogenetic structure of regional source pools and that the influence of regional source pools on local community structure is strong.

The importance of eco-evolutionary dynamics for predicting and managing insect range shifts.

Evolutionary change impacts the rate at which insect pests, pollinators, or disease vectors expand or contract their geographic ranges. Although evolutionary changes, and their ecological feedbacks, strongly affect these risks and associated ecological and economic consequences, they are often underappreciated in management efforts. Greater rigor and scope in study design, coupled with innovative technologies and approaches, facilitates our understanding of the causes and consequences of eco-evolutionary dynamics in insect range shifts. Future efforts need to ensure that forecasts allow for demographic and evolutionary change and that management strategies will maximize (or minimize) the adaptive potential of range-shifting insects, with benefits for biodiversity and ecosystem services.

Canopy and litter ant assemblages share similar climate-species density relationships.

Tropical forest canopies house most of the globe's diversity, yet little is known about global patterns and drivers of canopy diversity. Here, we present models of ant species density, using climate, abundance and habitat (i.e. canopy versus litter) as predictors. Ant species density is positively associated with temperature and precipitation, and negatively (or non-significantly) associated with two metrics of seasonality, precipitation seasonality and temperature range. Ant species density was significantly higher in canopy samples, but this difference disappeared once abundance was considered. Thus, apparent differences in species density between canopy and litter samples are probably owing to differences in abundance-diversity relationships, and not differences in climate-diversity relationships. Thus, it appears that canopy and litter ant assemblages share a common abundance-diversity relationship influenced by similar but not identical climatic drivers.

The Odonata of Quebec: Specimen data from seven collections.

BACKGROUND: The Odonata, dragonflies and damselflies, constitute one of the more charismatic and better-studied orders of insects. The approximately 6,000 extant species on Earth can be variously found on all continents, except Antarctica. A relatively stable taxonomy, a relative ease of species identification and an aquatic immature stage has made the Odonata a taxon of interest in documenting the symptoms of global environmental change, especially at higher latitudes. The Odonata fauna of the north-temperate Canadian province of Quebec includes 150 species, many of which are at the northern limits of their geographic distribution. NEW INFORMATION: Quebec hosts multiple entomological specimen depositories, including seven publicly-accessible research collections. One of these, the University of Montreal's Ouellet-Robert Entomological Collection, houses an exceptionally large collection of Odonata. An initial specimen data capture project for this collection gathered 31,595 Quebec Odonata occurrence records, but several Quebec species were missing and geographic coverage was biased towards the Montreal region. To complement this dataset, we undertook to digitise the Odonata records of six other public research collections. They are, in order of Quebec Odonata collection size, the Laval University Entomological Collection, McGill University's Lyman Entomological Museum, the Insectarium of Montreal Research Collection, the Quebec Government's Insect Collection, Bishop's University's Insect Collection and the Laurentian Forestry Centre's René-Martineau Insectarium. Of the 40,447 total specimen occurrence records, 36,951 are identified to the species level, including 137 of the 150 species officially-recorded in Quebec and 2 non-nominotypical subspecies. We here summarise the data and highlight the strengths and weaknesses of the datasets. The complete dataset is available with this publication (Suppl. material 1), whereas the specimen data associated with each collection are available as Darwin Core archives at Canadensys.net and will be updated as appropriate.

Climatic drivers of hemispheric asymmetry in global patterns of ant species richness.

Although many taxa show a latitudinal gradient in richness, the relationship between latitude and species richness is often asymmetrical between the northern and southern hemispheres. Here we examine the latitudinal pattern of species richness across 1003 local ant assemblages. We find latitudinal asymmetry, with southern hemisphere sites being more diverse than northern hemisphere sites. Most of this asymmetry could be explained statistically by differences in contemporary climate. Local ant species richness was positively associated with temperature, but negatively (although weakly) associated with temperature range and precipitation. After contemporary climate was accounted for, a modest difference in diversity between hemispheres persisted, suggesting that factors other than contemporary climate contributed to the hemispherical asymmetry. The most parsimonious explanation for this remaining asymmetry is that greater climate change since the Eocene in the northern than in the southern hemisphere has led to more extinctions in the northern hemisphere with consequent effects on local ant species richness.

Invasive ants alter the phylogenetic structure of ant communities.

Invasive species displace native species and potentially alter the structure and function of ecological communities. In this study, we compared the generic composition of intact and invaded ant communities from 12 published studies and found that invasive ant species alter the phylogenetic structure of native ant communities. Intact ant communities were phylogenetically evenly dispersed, suggesting that competition structures communities. However, in the presence of an invasive ant species, these same communities were phylogenetically clustered. Phylogenetic clustering in invaded communities suggests that invasive species may act as strong environmental filters and prune the phylogenetic tree of native species in a nonrandom manner, such that only a few closely related taxa can persist in the face of a biological invasion. Taxa that were displaced by invasive ant species were evenly dispersed in the phylogeny, suggesting that diversity losses from invasive ant species are not clustered in particular lineages. Collectively, these results suggest that there is strong phylogenetic structuring in intact native ant communities, but the spread of invasive species disassembles those communities above and beyond the effect of simple reductions in diversity.

Shared mycorrhizae but distinct communities of other root-associated microbes on co-occurring native and invasive maples.

BACKGROUND: Biological invasions are major drivers of environmental change that can significantly alter ecosystem function and diversity. In plants, soil microbes play an important role in plant establishment and growth; however, relatively little is known about the role they might play in biological invasions. A first step to assess whether root microbes may be playing a role in the invasion process is to find out if invasive plants host different microbes than neighbouring native plant species. METHODS: In this study we investigated differences in root associated microbes of native sugar maple (Acer saccharum Marsh.) and exotic Norway maple (A. platanoides L.) collected from a forested reserve in eastern Canada. We used microscopy to examine root fungi and high-throughput sequencing to characterize the bacterial, fungal and arbuscular mycorrhizal communities of both maple species over one growing season. RESULTS: We found differences in root associated bacterial and fungal communities between host species. Norway maple had a higher bacterial and fungal OTU (operational taxonomic units) richness compared to sugar maple, and the indicator species analysis revealed that nine fungal OTUs and three bacterial OTUs had a significant preference for sugar maple. The dominant bacterial phyla found on the roots of both maple species were Actinobacteria and Proteobacteria. The most common fungal orders associated with the Norway maple roots (in descending order) were Helotiales, Agaricales, Pleosporales, Hypocreales, Trechisporales while the Agaricales, Pleosporales, Helotiales, Capnodiales and Hypocreales were the dominant orders present in the sugar maple roots. Dark septate fungi colonization levels were higher in the sugar maple, but no differences in arbuscular mycorrhizal fungal communities and colonization rates were detected between maple species. DISCUSSION: Our findings show that two congeneric plant species grown in close proximity can harbor distinct root microbial communities. These findings provide further support for the importance of plant species in structuring root associated microbe communities. The high colonization levels observed in Norway maple demonstrates its compatibility with arbuscular mycorrhizal fungi in the introduced range. Plant-associated microbial communities can affect host fitness and function in many ways; therefore, the observed differences suggest a possibility that biotic interactions can influence the dynamics between native and invasive species.