Trophic interactions and abiotic factors drive functional and phylogenetic structure of vertebrate herbivore communities across the Arctic tundra biome

Communities are assembled from species that evolve or colonise a given geographic region, and persist in the face of abiotic conditions and interactions with other species. The evolutionary and colonisation histories of communities are characterised by phylogenetic diversity, while functional diversity is indicative of abiotic and biotic conditions. The relationship between functional and phylogenetic diversity infers whether species functional traits are divergent (differing between related species) or convergent (similar among distantly related species). Biotic interactions and abiotic conditions are known to influence macroecological patterns in species richness, but how functional and phylogenetic diversity of guilds vary with biotic factors, and the relative importance of biotic drivers in relation to geographic and abiotic drivers is unknown. In this study, we test whether geographic, abiotic or biotic factors drive biome‐scale spatial patterns of functional and phylogenetic diversity and functional convergence in vertebrate herbivores across the Arctic tundra biome. We found that functional and phylogenetic diversity both peaked in the western North American Arctic, and that spatial patterns in both were best predicted by trophic interactions, namely vegetation productivity and predator diversity, as well as climatic severity. Our results show that both bottom–up and top–down trophic interactions, as well as winter temperatures, drive the functional and phylogenetic structure of Arctic vertebrate herbivore assemblages. This has implications for changing Arctic ecosystems; under future warming and northward movement of predators potential increases in phylogenetic and functional diversity in vertebrate herbivores may occur. Our study thus demonstrates that trophic interactions can determine large‐scale functional and phylogenetic diversity just as strongly as abiotic conditions.     

Climate and food diversity as drivers of mammal diversity in Inner Mongolia

Traditionally, geographical distribution of biodiversity is assumed to be codetermined by multiple factors, for example, temperature, precipitation, environmental heterogeneity, and biotic interactions. However, few studies have simultaneously compared the relative roles of these factors in shaping the mammal diversity patterns for different feeding groups, that is, herbivores, insectivores, and carnivores. In this study, we assessed the relations between mammal diversity and current climate (mean annual temperature and precipitation), altitudinal range as well as mammal's food diversity in Inner Mongolia. Our results showed that the species richness for the three feeding guilds of mammals consistently increased with their food diversity, that is, species richness of plants, insects, and rodents. Mammal diversity also significantly decreased with mean annual temperature and precipitation. Random Forest models indicated that climate and food diversity were always included in the combinations of variables most associated with mammal diversity. Our findings suggest that while climate is an important predictor of large scale distribution of mammal diversity, biotic interactions, that is, food diversity, could also play important roles.     

Assessing the latitudinal gradient in herbivory

Plant–herbivore interactions occur in all ecosystems and provide a major avenue for energy flow to higher trophic levels. A long‐standing hypothesis to explain the latitudinal gradient in species diversity proposes that the relatively stable and frost‐free climate of the tropics should lead to more intense biotic interactions in tropical compared with temperate environments, giving rise to a greater diversity of plants and herbivores. Herbivory rates have been compared across latitudes to test this biotic interactions hypothesis, with herbivory typically being measured from observable leaf damage. However, we argue that a measure of percentage leaf damage alone does not straightforwardly reflect the cost of herbivory to the plant, and on its own does not constitute an appropriate test of the biotic interactions hypothesis. For a given amount of herbivory, the impact of herbivory is dependent upon many factors, such as the construction cost of the leaf, the growth and replacement rates and leaf life span. We investigate the latitudinal gradient in herbivory by analysing a large dataset of herbivory rates for 452 tree species and separating the species into those with short and long leaf life spans. We show that annual herbivory rates tend to be greater at lower latitudes for evergreen species (which have long‐lived leaves), but no trend in herbivory rate with latitude was found for species with short leaf life spans. Phylogenetic least squares regression assuming Ornstein‐Uhlenbeck processes also showed a negative effect of latitude on herbivory rate for evergreen trees, but we caution that viewing herbivory as a species trait is problematic. An integrative approach that incorporates leaf life span, as well as the costs of investment in growth and potential costs of losing leaf tissue, is needed to further our understanding of the ecological and evolutionary dynamics of herbivory.     

Introductions do not compensate for functional and phylogenetic losses following extinctions in insular bird assemblages

The ratio of species extinctions to introductions has been comparable for many insular assemblages, suggesting that introductions could have ‘compensated’ for extinctions. However, the capacity for introduced species to replace ecological roles and evolutionary history lost following extinction is unclear. We investigated changes in bird functional and phylogenetic diversity in the wake of extinctions and introductions across a sample of 32 islands worldwide. We found that extinct and introduced species have comparable functional and phylogenetic alpha diversity. However, this was distributed at different positions in functional space and in the phylogeny, indicating a ‘false compensation’. Introduced and extinct species did not have equivalent functional roles nor belong to similar lineages. This makes it unlikely that novel island biotas composed of introduced taxa will be able to maintain ecological roles and represent the evolutionary histories of pre‐disturbance assemblages and highlights the importance of evaluating changes in alpha and beta diversity concurrently.     

Diversity indices for ecological networks: a unifying framework using Hill numbers

Describing how ecological interactions change over space and time and how they are shaped by environmental conditions is crucial to understand and predict ecosystem trajectories. However, it requires having an appropriate framework to measure network diversity locally, regionally and between samples (α‐, γ‐ and β‐diversity). Here, we propose a unifying framework that builds on Hill numbers and accounts both for the probabilistic nature of biotic interactions and the abundances of species or groups. We emphasise the importance of analysing network diversity across different species aggregation levels (e.g. from species to trophic groups) to get a better understanding of network structure. We illustrate our framework with a simulation experiment and an empirical analysis using a global food‐web database. We discuss further usages of the framework and show how it responds to recent calls on comparing ecological networks and analysing their variation across environmental gradients and time.     

The tri‐trophic niche concept and adaptive radiation of phytophagous insects

A conceptual divide exists between ecological and evolutionary approaches to understanding adaptive radiation, although the phenomenon is inherently both ecological and evolutionary. This divide is evident in studies of phytophagous insects, a highly diverse group that has been frequently investigated with the implicit or explicit goal of understanding its diversity. Whereas ecological studies of phytophagous insects increasingly recognize the importance of tri‐trophic interactions as determinants of niche dimensions such as host‐plant associations, evolutionary studies typically neglect the third trophic level. Here we attempt to reconcile ecological and evolutionary approaches through the concept of the ecological niche. We specifically present a tri‐trophic niche concept as a foil to the traditional bi‐trophic niche concept for phytophagous insects. We argue that these niche concepts have different implications for understanding herbivore community structure, population divergence, and evolutionary diversification. To this end, we offer contrasting empirical predictions of bi‐ and tri‐trophic niche concepts for patterns of community structure, the process of population divergence, and patterns of evolutionary diversification of phytophagous insects.     

From species distributions to meta‐communities

The extent that biotic interactions and dispersal influence species ranges and diversity patterns across scales remains an open question. Answering this question requires framing an analysis on the frontier between species distribution modelling (SDM), which ignores biotic interactions and dispersal limitation, and community ecology, which provides specific predictions on community and meta‐community structure and resulting diversity patterns such as species richness and functional diversity. Using both empirical and simulated datasets, we tested whether predicted occurrences from fine‐resolution SDMs provide good estimates of community structure and diversity patterns at resolutions ranging from a resolution typical of studies within reserves (250 m) to that typical of a regional biodiversity study (5 km). For both datasets, we show that the imprint of biotic interactions and dispersal limitation quickly vanishes when spatial resolution is reduced, which demonstrates the value of SDMs for tracking the imprint of community assembly processes across scales.     

Climate or diet? The importance of biotic interactions in determining species range size

Aim

Species geographical range sizes play a crucial role in determining species vulnerability to extinction. Although several mechanisms affect range sizes, the number of biotic interactions and species climatic tolerance are often thought to play discernible roles, defining two dimensions of the Hutchinsonian niche. Yet, the relative importance of the trophic and the climatic niche for determining species range sizes is largely unknown.

Location

Central and northern Europe.

Time period

Present.

Major taxa studied

Gall-inducing sawflies and their parasitoids.

Methods

We use data documenting the spatial distributions and biotic interactions of 96 herbivore species, and their 125 parasitoids, across Europe and analyse the relationship between species range size and the climatic and trophic dimensions of the niche. We then compare the observed relationships with null expectations based on species occupancy to understand whether the relationships observed are an inevitable consequence of species range size or if they contain information about the importance of each dimension of the niche on species range size.

Results

We find that both niche dimensions are positively correlated with species range size, with larger ranges being associated with wider climatic tolerances and larger numbers of interactions. However, diet breadth appears to more strongly limit species range size. Species with larger ranges have more interactions locally and they are also able to interact with a larger diversity of species across sites (i.e. higher β-diversity), resulting in a larger number of interactions at continental scales.

Main conclusions

We show for the first time how different aspects of species diet niches are related to their range size. Our study offers new insight into the importance of biotic interactions in determining species spatial distributions, which is critical for improving understanding and predictions of species vulnerability to extinction under the current rates of global environmental change.     

Phylogenetic beta diversity: linking ecological and evolutionary processes across space in time

A key challenge in ecological research is to integrate data from different scales to evaluate the ecological and evolutionary mechanisms that influence current patterns of biological diversity. We build on recent attempts to incorporate phylogenetic information into traditional diversity analyses and on existing research on beta diversity and phylogenetic community ecology. Phylogenetic beta diversity (phylobetadiversity) measures the phylogenetic distance among communities and as such allows us to connect local processes, such as biotic interactions and environmental filtering, with more regional processes including trait evolution and speciation. When combined with traditional measures of beta diversity, environmental gradient analyses or ecological niche modelling, phylobetadiversity can provide significant and novel insights into the mechanisms underlying current patterns of biological diversity.     

Adaptive genetic diversity of dominant species contributes to species co-existence and community assembly

The synthesis of evolutionary biology and community ecology aims to understand how genetic variation within one species can shape community properties and how the ecological properties of a community can drive the evolution of a species. A rarely explored aspect is whether the interaction of genetic variation and community properties depends on the species'' ecological role. Here we investigated the interactions among environmental factors, species diversity, and the within-species genetic diversity of species with different ecological roles. Using high-throughput DNA sequencing, we genotyped a canopy-dominant tree species, Parashorea chinensis, and an understory-abundant species, Pittosporopsis kerrii, from fifteen plots in Xishuangbanna tropical seasonal rainforest and estimated their adaptive, neutral and total genetic diversity; we also surveyed species diversity and assayed key soil nutrients. Structural equation modelling revealed that soil nitrogen availability created an opposing effect in species diversity and adaptive genetic diversity of the canopy-dominant Pa. chinensis. The increased adaptive genetic diversity of Pa. chinensis led to greater species diversity by promoting co-existence. Increased species diversity reduced the adaptive genetic diversity of the dominant understory species, Pi. kerrii, which was promoted by the adaptive genetic diversity of the canopy-dominant Pa. chinensis. However, such relationships were absent when neutral genetic diversity or total genetic diversity were used in the model. Our results demonstrated the important ecological interaction between adaptive genetic diversity and species diversity, but the pattern of the interaction depends on the identity of the species. Our results highlight the significant ecological role of dominant species in competitive interactions and regulation of community structure.     

植物边缘种群遗传多样性研究进展

边缘种群指地理分布边缘可检测到的一定数量的同种个体集合, 准确评价其遗传多样性对于理解第四纪冰期后气候变化对物种边缘扩展或收缩、遗传资源保护与利用以及物种形成等有重要意义。该文探讨了维持植物边缘种群遗传多样性的进化机制, 分析交配系统对物种边缘及其遗传多样性的影响, 比较了边缘与中心种群遗传多样性的差异及其形成的生态与进化过程, 并探讨了边缘种群遗传多样性与其所在的群落物种多样性的关系及理论基础。该文提出今后研究的重点是应用全基因组序列或转录组基因序列研究前缘-后缘种群之间或边缘-中心种群之间的适应性差异, 边缘种群与所在群落其他物种之间相互作用的分子机制, 深入解析边缘种群对环境的适应及边缘种群遗传多样性与群落物种多样性关系的生态与进化过程。     

Biodiversity inhibits species' evolutionary responses to changing environments

Despite growing interplay between ecological and evolutionary studies, the question of how biodiversity influences evolutionary dynamics within species remains understudied. Here, using a classical model of phenotypic evolution in species occupying a patchy environment, but introducing global change affecting patch conditions, we show that biodiversity can inhibit species' evolution during global change. The presence of several species increases the chance that one or more species are pre-adapted to new conditions, which restricts the ecological opportunity for evolutionary responses in all the species. Consequently, environmental change tends to select for changes in species abundances rather than for changing phenotypes within each species. The buffering effects of species diversity that we describe might be one important but neglected explanation for widely observed niche conservatism in natural systems. Furthermore, the results show that attempts to understand biotic responses to environmental change need to consider both ecological and evolutionary processes in a realistically diverse setting.     

A roadmap to plant functional island biogeography

Island biogeography is the study of the spatio-temporal distribution of species, communities, assemblages or ecosystems on islands and other isolated habitats. Island diversity is structured by five classes of process: dispersal, establishment, biotic interactions, extinction and evolution. Classical approaches in island biogeography focused on species richness as the deterministic outcome of these processes. This has proved fruitful, but species traits can potentially offer new biological insights into the processes by which island life assembles and why some species perform better at colonising and persisting on islands. Functional traits refer to morphological and phenological characteristics of an organism or species that can be linked to its ecological strategy and that scale up from individual plants to properties of communities and ecosystems. A baseline hypothesis is for traits and ecological strategies of island species to show similar patterns as a matched mainland environment. However, strong dispersal, environmental and biotic-interaction filters as well as stochasticity associated with insularity modify this baseline. Clades that do colonise often embark on distinct ecological and evolutionary pathways, some because of distinctive evolutionary forces on islands, and some because of the opportunities offered by freedom from competitors or herbivores or the absence of mutualists. Functional traits are expected to be shaped by these processes. Here, we review and discuss the potential for integrating functional traits into island biogeography. While we focus on plants, the general considerations and concepts may be extended to other groups of organisms. We evaluate how functional traits on islands relate to core principles of species dispersal, establishment, extinction, reproduction, biotic interactions, evolution and conservation. We formulate existing knowledge as 33 working hypotheses. Some of these are grounded on firm empirical evidence, others provide opportunities for future research. We organise our hypotheses under five overarching sections. Section A focuses on plant functional traits enabling species dispersal to islands. Section B discusses how traits help to predict species establishment, successional trajectories and natural extinctions on islands. Section C reviews how traits indicate species biotic interactions and reproduction strategies and which traits promote intra-island dispersal. Section D discusses how evolution on islands leads to predictable changes in trait values and which traits are most susceptible to change. Section E debates how functional ecology can be used to study multiple drivers of global change on islands and to formulate effective conservation measures. Islands have a justified reputation as research models. They illuminate the forces operating within mainland communities by showing what happens when those forces are released or changed. We believe that the lens of functional ecology can shed more light on these forces than research approaches that do not consider functional differences among species.     

Diversity dynamics of Miocene mammals in relation to the history of tectonism and climate

Continental biodiversity gradients result not only from ecological processes, but also from evolutionary and geohistorical processes involving biotic turnover in landscape and climatic history over millions of years. Here, we investigate the evolutionary and historical contributions to the gradient of increasing species richness with topographic complexity. We analysed a dataset of 418 fossil rodent species from western North America spanning 25 to 5 Ma. We compared diversification histories between tectonically active (Intermontane West) and quiescent (Great Plains) regions. Although diversification histories differed between the two regions, species richness, origination rate and extinction rate per million years were not systematically different over the 20 Myr interval. In the tectonically active region, the greatest increase in originations coincided with a Middle Miocene episode of intensified tectonic activity and global warming. During subsequent global cooling, species richness declined in the montane region and increased on the Great Plains. These results suggest that interactions between tectonic activity and climate change stimulate diversification in mammals. The elevational diversity gradient characteristic of modern mammalian faunas was not a persistent feature over geologic time. Rather, the Miocene rodent record suggests that the elevational diversity gradient is a transient feature arising during particular episodes of Earth''s history.     

Evolutionary history and precipitation seasonality shape niche overlap in Neotropical bat–plant pollination networks

Species interactions are one dimension of a niche. Niche overlap arises when two species share an interaction partner. In pollination systems, environmental and biotic factors affect niche overlap. Here, we explored the effects of climate seasonality, plant and bat richness, morphological traits, and phylogenetic distance in shaping the niche overlap of Neotropical bat–plant pollination networks. We examined a dataset of 22 bat–plant pollination networks in the Neotropical region. We measured niche overlap in bats and plants with the Morisita-Horn index (Ĉ_H) and used a SAR model to test the relationships between niche overlap and both abiotic and biotic factors. We found a lower niche overlap among bats in communities composed of phylogenetically distant bat species. Moreover, plant and bat overlap was lower in regions with higher precipitation seasonality. Our results indicate that climate seasonality and bat evolutionary history drive niche overlap in Neotropical bat–plant pollination interactions. These findings suggest that a higher precipitation seasonality promotes the emergence of temporal modules, which reduces niche overlap, likely due to seasonal species phenologies. Furthermore, the method used to record the interactions affects the degree of niche overlap. Interactions recorded with pollen samples tend to have higher niche overlap than direct observations. The responses of morphological traits and phylogenetic distances in bat niche overlap were uncoupled, suggesting an effect of historical processes independent of morphological traits. Our findings reinforce the importance of evolutionary history and ecological processes in imprinting patterns of interaction niche overlap.     

A meta-analysis of biotic resistance to exotic plant invasions

Biotic resistance describes the ability of resident species in a community to reduce the success of exotic invasions. Although resistance is a well‐accepted phenomenon, less clear are the processes that contribute most to it, and whether those processes are strong enough to completely repel invaders. Current perceptions of strong, competition‐driven biotic resistance stem from classic ecological theory, Elton's formulation of ecological resistance, and the general acceptance of the enemies‐release hypothesis. We conducted a meta‐analysis of the plant invasions literature to quantify the contribution of resident competitors, diversity, herbivores and soil fungal communities to biotic resistance. Results indicated large negative effects of all factors except fungal communities on invader establishment and performance. Contrary to predictions derived from the natural enemies hypothesis, resident herbivores reduced invasion success as effectively as resident competitors. Although biotic resistance significantly reduced the establishment of individual invaders, we found little evidence that species interactions completely repelled invasions. We conclude that ecological interactions rarely enable communities to resist invasion, but instead constrain the abundance of invasive species once they have successfully established.     

Eco‐evolutionary feedbacks between prey densities and linkage disequilibrium in the predator maintain diversity

Diversity occurs at multiple scales. Within a single population, there is diversity in genotypes and phenotypes. At a larger scale, within ecological communities, there is diversity in species. A number of studies have investigated how diversity at these two scales influence each other through what has been termed eco‐evolutionary feedbacks. Here we study a three‐species ecological module called apparent competition, in which the predator is evolving in a trait that determines its interaction with two prey species. Unlike previous studies on apparent competition, which employed evolutionary frameworks with very simple genetics, we study an eco‐evolutionary model in which the predator's trait is determined by two recombining diallelic loci, so that its mean and variance can evolve, as well as associations (linkage disequilibrium) between the loci. We ask how eco‐evolutionary feedbacks with these two loci affect the coexistence of the prey species and the maintenance of polymorphisms within the predator species. We uncover a novel eco‐evolutionary feedback between the prey densities and the linkage disequilibrium between the predator's loci. Through a stability analysis, we demonstrate how these feedbacks affect polymorphisms at both loci and, among others, may generate stable cycling.     

Species mtDNA genetic diversity explained by infrapopulation size in a host‐symbiont system

Understanding what shapes variation in genetic diversity among species remains a major challenge in evolutionary ecology, and it has been seldom studied in parasites and other host‐symbiont systems. Here, we studied mtDNA variation in a host‐symbiont non‐model system: 418 individual feather mites from 17 feather mite species living on 17 different passerine bird species. We explored how a surrogate of census size, the median infrapopulation size (i.e., the median number of individual parasites per infected host individual), explains mtDNA genetic diversity. Feather mite species genetic diversity was positively correlated with mean infrapopulation size, explaining 34% of the variation. As expected from the biology of feather mites, we found bottleneck signatures for most of the species studied but, in particular, three species presented extremely low mtDNA diversity values given their infrapopulation size. Their star‐like haplotype networks (in contrast with more reticulated networks for the other species) suggested that their low genetic diversity was the consequence of severe bottlenecks or selective sweeps. Our study shows for the first time that mtDNA diversity can be explained by infrapopulation sizes, and suggests that departures from this relationship could be informative of underlying ecological and evolutionary processes.     

Ecotone formation through ecological niche construction: the role of biodiversity and species interactions

Rapid changes in species composition, also known as ecotones, can result from various causes including rapid changes in environmental conditions, or physiological thresholds. The possibility that ecotones arise from ecological niche construction by ecosystem engineers has received little attention. In this study, we investigate how the diversity of ecosystem engineers, and their interactions, can give rise to ecotones. We build a spatially explicit dynamical model that couples a multispecies community and its abiotic environment. We use numerical simulations and analytical techniques to determine the biotic and abiotic conditions under which ecotone emergence is expected to occur, and the role of biodiversity therein. We show that the diversity of ecosystem engineers can lead to indirect interactions through the modification of their shared environment. These interactions, which can be either competitive or mutualistic, can lead to the emergence of discrete communities in space, separated by sharp ecotones where a high species turnover is observed. Considering biodiversity is thus critical when studying the influence of species–environment interactions on the emergence of ecotones. This is especially true for the wide range of species that have small to moderate effects on their environment. Our work highlights new mechanisms by which biodiversity loss could cause significant changes in spatial community patterns in changing environments.     

Non-random interactions within and across guilds shape the potential to coexist in multi-trophic ecological communities

Theory posits that the persistence of species in ecological communities is shaped by their interactions within and across trophic guilds. However, we lack empirical evaluations of how the structure, strength and sign of biotic interactions drive the potential to coexist in diverse multi-trophic communities. Here, we model community feasibility domains, a theoretically informed measure of multi-species coexistence probability, from grassland communities comprising more than 45 species on average from three trophic guilds (plants, pollinators and herbivores). Contrary to our hypothesis, increasing community complexity, measured either as the number of guilds or community richness, did not decrease community feasibility. Rather, we observed that high degrees of species self-regulation and niche partitioning allow for maintaining larger levels of community feasibility and higher species persistence in more diverse communities. Our results show that biotic interactions within and across guilds are not random in nature and both structures significantly contribute to maintaining multi-trophic diversity.