One of the most pervasive concepts in the study of community assembly is the metaphor of the environmental filter, which refers to abiotic factors that prevent the establishment or persistence of species in a particular location. The metaphor has its origins in the study of community change during succession and in plant community dynamics, although it has gained considerable attention recently as part of a surge of interest in functional trait and phylogenetic‐based approaches to the study of communities. While the filtering metaphor has clear utility in some circumstances, it has been challenging to reconcile the environmental filtering concept with recent developments in ecological theory related to species coexistence. These advances suggest that the evidence used in many studies to assess environmental filtering is insufficient to distinguish filtering from the outcome of biotic interactions. We re‐examine the environmental filtering metaphor from the perspective of coexistence theory. In an effort to move the discussion forward, we present a simple framework for considering the role of the environment in shaping community membership, review the literature to document the evidence typically used in environmental filtering studies and highlight research challenges to address in coming years. The current usage of the environmental filtering term in empirical studies likely overstates the role abiotic tolerances play in shaping community structure. We recommend that the term ‘environmental filtering’ only be used to refer to cases where the abiotic environment prevents establishment or persistence in the absence of biotic interactions, although only 15% of the studies in our review presented such evidence. Finally, we urge community ecologists to consider additional mechanisms aside from environmental filtering by which the abiotic environment can shape community pattern. Lay Summary
The effects of the present biodiversity crisis have been largely focused on the loss of species. However, a missed component of biodiversity loss that often accompanies or even precedes species disappearance is the extinction of ecological interactions. Here, we propose a novel model that (i) relates the diversity of both species and interactions along a gradient of environmental deterioration and (ii) explores how the rate of loss of ecological functions, and consequently of ecosystem services, can be accelerated or restrained depending on how the rate of species loss covaries with the rate of interactions loss. We find that the loss of species and interactions are decoupled, such that ecological interactions are often lost at a higher rate. This implies that the loss of ecological interactions may occur well before species disappearance, affecting species functionality and ecosystems services at a faster rate than species extinctions. We provide a number of empirical case studies illustrating these points. Our approach emphasizes the importance of focusing on species interactions as the major biodiversity component from which the ‘health’ of ecosystems depends. Lay Summary
The relationship between stomatal conductance (g s ) and leaf water potential (Ψ l ) is key to the understanding of plant function under changing climate. The variability among tree species gave rise to selection towards either of two contrasting water management types: isohydric or anisohydric. This study explores the variability of g s to Ψ l across tree species. Curves of g s (Ψ l ) were collected from the scientific literature for 70 woody plant species. The data set is comprised of angiosperm and gymnosperm species from all major forest biomes. The hypothesis that curves from different tree species diverge between isohydric and anisohydric behaviours was tested. Species‐specific curves formed a continuum, rather than dichotomy between isohydric and anisohydric, as confirmed by distribution models. Alternatively, the water potential at 50% of the maximum g s (Ψg s 50) was used to quantitatively compare between species. A major difference emerged among xylem anatomy classes whereby ring‐porous species had higher absolute g s at Ψ l < −2 MP a than diffuse‐porous and coniferous species. A positive, linear correlation was shown between Ψg s 50 and Ψ l at 50% loss of xylem conductivity. The results suggest that stomatal sensitivity to leaf water potential strongly relates to xylem characteristics. The use of Ψg s 50 offers a quantitative alternative to the current, yet biased, distinction between isohydric and anisohydric species. Lay Summary
The subdiscipline of 'community phylogenetics' is rapidly growing and influencing thinking regarding community assembly. In particular, phylogenetic dispersion of co-occurring species within a community is commonly used as a proxy to identify which community assembly processes may have structured a particular community: phylogenetic clustering as a proxy for abiotic assembly, that is habitat filtering, and phylogenetic overdispersion as a proxy for biotic assembly, notably competition. We challenge this approach by highlighting (typically) implicit assumptions that are, in reality, only weakly supported, including (i) phylogenetic dispersion reflects trait dispersion; (ii) a given ecological function can be performed only by a single trait state or combination of trait states; (iii) trait similarity causes enhanced competition; (iv) competition causes species exclusion; (v) communities are at equilibrium with processes of assembly having been completed; (vi) assembly through habitat filtering decreases in importance if assembly through competition increases, such that the relative balance of the two can be thus quantified by a single parameter; and (vii) observed phylogenetic dispersion is driven predominantly by local and present-day processes. Moreover, technical sophistication of the phylogenetic-patterns-as-proxy approach trades off against sophistication in alternative, potentially more pertinent approaches to directly observe or manipulate assembly processes. Despite concerns about using phylogenetic dispersion as a proxy for community assembly processes, we suggest there are underappreciated benefits of quantifying the phylogenetic structure of communities, including (i) understanding how coexistence leads to the macroevolutionary diversification of habitat lineage-pools (i.e. phylogenetic-patterns-as-result approach); and (ii) understanding the macroevolutionary contingency of habitat lineage-pools and how it affects present-day species coexistence in local communities (i.e. phylogenetic-patterns-as-cause approach). We conclude that phylogenetic patterns may be little useful as proxy of community assembly. However, such patterns can prove useful to identify and test novel hypotheses on (i) how local coexistence may control macroevolution of the habitat lineage-pool, for example through competition among close relatives triggering displacement and diversification of characters, and (ii) how macroevolution within the habitat lineage-pool may control local coexistence of related species, for example through origin of close relatives that can potentially enter in competition. 2015 British Ecological Society.
1. Indices quantifying the functional aspect of biodiversity are essential in understanding relationships between biodiversity, ecosystem functioning and environmental constraints. Many indices of functional diversity have been published but we lack consensus about what indices quantify, how redundant they are and which ones are recommended. 2. This study aims to build a typology of functional diversity indices from artificial data sets encompassing various community structures (different assembly rules, various species richness levels) and to identify a set of independent indices able to discriminate community assembly rules. 3. Our results confirm that indices can be divided into three main categories, each of these corresponding to one aspect of functional diversity: functional richness, functional evenness and functional divergence. Most published indices are highly correlated and quantify functional richness while quadratic entropy (Q) represents a mix between functional richness and functional divergence. Conversely, two indices (FEve and FDiv respectively quantifying functional evenness and functional divergence) are rather independent to all the others. The power analysis revealed that some indices efficiently detect assembly rules while others performed poorly. 4. To accurately assess functional diversity and establish its relationships with ecosystem functioning and environmental constraints, we recommend investigating each functional component separately with the appropriate index. Guidelines are provided to help choosing appropriate indices given the issue being investigated. 5. This study demonstrates that functional diversity indices have the potential to reveal the processes that structure biological communities. Combined with complementary methods (phylogenetic and taxonomie diversity), the multifaceted framework of functional diversity will help improve our understanding of how biodiversity interacts with ecosystem processes and environmental constraints.
1. Terrestrial ectotherms are likely to face increased periods of heat stress as mean temperatures and temperature variability increase over the next few decades. Here, we consider the extent to which changes in upper thermal limits, through plasticity or evolution, might be constrained, and we survey insect and reptile data to identify groups likely to be particularly susceptible to thermal stress. 2. Plastic changes increase thermal limits in many terrestrial ectotherms, but tend to have less effect on upper limits than lower limits. 3. Although comparisons across insect species have normally not taken into account the potential for plastic responses, mid-latitude species seem most prone to experience heat stress now and into the future, consistent with data from lizards and other groups. 4. Evolutionary adaptive potential has only been measured for some species; there is likely to be genetic variation for heat responses in populations, but selection and heritability experiments suggest that upper thermal limits may not increase much. 5. Although related species can differ by several degrees in their upper thermal limits, there is strong phylogenetic signal for upper limits. If these reflect evolutionary constraints, substantial molecular changes may be required to increase upper thermal limits. 6. Findings point to many terrestrial ectotherms having a limited potential to change their thermal limits particularly within the context of an average predicted temperature increase of 2—4 °C for mid-latitude populations over the next few decades.
1. Chronic activation of the stress axis caused by long-term uncontrollable and unpredictable factors in the environment has been regarded as causing maladaptive and/or pathological effects, both by those studying animals in the laboratory and in nature. While pathology may apply to the former, I argue that it does not apply to the latter. 2. Our thinking on the role of chronic stress in animals in nature has been heavily influenced by biomedical research, but much less so by the ecological and evolutionary context within which animals actually function. I argue that when such stressors occur (e.g. periods of high predation risk, food limitation, prolonged severe weather, social conflict, etc.), although the animal may be chronically stressed, its responses are adaptive and continue to promote fitness. 3. Chronic stressors in nature can be subdivided into whether they are reactive (direct physiological challenges threatening homeostasis and not requiring cognitive processing — for example, food limitation) or anticipatory (perceived to be threatening and requiring cognitive processing — for example, high predation risk). For anticipatory stressors, their impact on the animal should not be based on their absolute duration (they may be acute), but rather by the duration of their physiological consequences. 4. The anticipatory stressor of persistent high predation risk does not elicit chronic stress in all prey classes. Cyclic snowshoe hare and arctic ground squirrels exhibit evidence of chronic stress when predator numbers are high, but cyclic vole and noncyclic elk populations do not. I suggest that chronic stress has evolved to benefit the fitness of the former and not the later, with the key factors being lifespan and life history. I propose that chronic stress evolves in a species only if it is adaptive.
1. The role of phenotypic plasticity in evolution has historically been a contentious issue because of debate over whether plasticity shields genotypes from selection or generates novel opportunities for selection to act. Because plasticity encompasses diverse adaptive and non-adaptive responses to environmental variation, no single conceptual framework adequately predicts the diverse roles of plasticity in evolutionary change. 2. Different types of phenotypic plasticity can uniquely contribute to adaptive evolution when populas are faced with new or altered environments. Adaptive plasticity should promote establishment and persistence in a new environment, but depending on how close the plastic response is to the new favoured phenotypic optimum dictates whether directional selection will cause adaptive divergence between populations. Further, non-adaptive plasticity in response to stressful environments can result in a mean phenotypic response being further away from the favoured optimum or alternatively increase the variance around the mean due to the expression of cryptic genetic variation. The expression of cryptic genetic variation can facilitate adaptive evolution if by chance it results in a fitter phenotype. 3. We conclude that adaptive plasticity that places populations close enough to a new phenotypic optimum for directional selection to act is the only plasticity that predictably enhances fitness and is most likely to facilitate adaptive evolution on ecological timescales in new environments. However, this type of plasticity is likely to be the product of past selection on variation that may have been initially non-adaptive. 4. We end with suggestions on how future empirical studies can be designed to better test the importance of different kinds of plasticity to adaptive evolution.
1. A growing number of ecologists are turning to the enumeration of white blood cells from blood smears (leukocyte profiles) to assess stress in animals. There has been some inconsistency and controversy in the ecological literature, however, regarding their interpretation. The inconsistencies may stem partly from a lack of information regarding how stress affects leukocytes in different taxa, and partly from a failure on the part of researchers in one discipline to consult potentially informative literature from another. 2. Here, we seek to address both issues by reviewing the literature on the leukocyte response to stress, spanning the taxa of mammals (including humans), birds, amphibians, reptiles and fish. 3. We show that much of the early literature points to a close link between leukocyte profiles and glucocorticoid levels. Specifically, these hormones act to increase the number and percentage of neutrophils (heterophils in birds and reptiles), while decreasing the number and percentage of lymphocytes. This phenomenon is seen in all five vertebrate taxa in response to either natural stressors or exogenous administration of stress hormones. For the ecologist, therefore, high ratios of heterophils or neutrophils to lymphocytes ('H: L' or 'N: L' ratios) in blood samples reliably indicate high glucocorticoid levels. Furthermore, this close relationship between stress hormones and N: L or H: L ratios needs to be highlighted more prominently in haematological assessments of stress, as it aids the interpretation of results. 4. As with hormone assays, there are challenges to overcome in the use of leukocytes profiles to assess levels of stress; however, there are also advantages to this approach, and we outline each. Given the universal and consistent nature of the haematological response to stress, plus the overwhelming evidence from the veterinary, biomedical and ecological literature reviewed here, we conclude that this method can provide a reliable assessment of stress in all vertebrate taxa.
Symbiotic microbes have become increasingly recognized to mediate interactions between natural enemies and their hosts. The ecologies of these symbioses, however, are poorly understood in many systems, and a predictive framework is needed to guide future studies. To achieve this, we focus on heritable, defensive microbes of insects. Our review of laboratory‐based studies identifies diverse bacterial species that have independently evolved to protect a range of insects against parasitoids, parasites, predators and pathogens. Although defensive mechanisms are typically unknown, some involve toxins or the upregulation of host immunity. Despite substantial benefits of infection in the presence of natural enemies, the protective symbionts of insects are often found at intermediate levels in natural populations. Using a host‐centred population genetics approach made possible by the host restriction and cytoplasmic inheritance of these microbes, we propose that balancing selection plays a major role in symbiont maintenance, with protective benefits in the presence of enemies and infection costs in their absence. Other mediating factors are likely to be important, including temperature, superinfections and transmission dynamics. While few studies have provided evidence for defence in the field, several studies have shown symbiont infection frequencies to be dynamic, varying across temporal and spatial gradients and food–plant associations. Newly presented data from our pea aphid research reveal that temporal shifts in defensive symbiont prevalence can be quite rapid, with Hamiltonella defensa showing 10–20% shifts around a seasonal average of c . 50%. Such findings contrast with more unidirectional changes seen in laboratory population cages, suggesting temporal changes in the costs and benefits of symbionts in the field. To frame future research on defensive symbiont ecology, we briefly consider a range of studies needed to test laboratory‐ and field‐derived predictions on defensive symbiosis. Included are investigations of defensive mechanisms, symbiont‐driven co‐evolution and community‐level effects. We also consider the need for more thorough and highly resolved molecular diagnostics of natural infections, laboratory studies on functional differences between symbiont strains and species and studies on the relative costs and benefits of defenders in nature. The emerging theme of symbiont‐mediated defence across eukaryotes suggests that knowledge of the functional mechanisms behind protection and natural symbiont dynamics could be key to understanding many of the world's antagonistic species interactions. Thus, the development of insects as a model for such studies holds promise for these organisms and beyond. Lay Summary
1. Body condition is a major concept in ecology addressed in countless studies, and a variety of non-destructive methods are used to estimate the condition of individuals based on the relationship between body mass M and measures of length L. There is currently no consensus about the most appropriate condition index (CI) method, and various traditions have been established within subdisciplines in which ecologists tend to apply that method used previously by their peers. 2. Here, we present a reappraisal of six conventional CI methods: Fulton's index (K = M/L³), Quételet's index (BMI = M/L²), Relative condition (K n , computed as the observed individual mass divided by the predicted mass M i * = a L i b where a and b are determined by ordinary least squares (OLS) regression of M against L), Relative mass (W r , where a and b above are determined from a reference population), the Residual index (R i , the residuals from an OLS regression of M against L) and ANCOVA. We compare the performance of these methods with that of the Scaled mass index, a novel method which was previously shown to perform better than R i as a predictor of fat and other body components [J. Peig & A.J. Green (2009) Oikos, 118, 1883]. 3. To be reliable, a CI method must successfully account for the changing relationship between M and L as body size changes and growth occurs (i.e. for the scaling relationship between M and L). Using data from three species of small mammals we show that, unlike the Scaled mass index, all six conventional methods fail to do this, and as a result they consistently lead to significant differences in CIs between age classes and sex that are a mere consequence of changes in body size. The Scaled mass index was also particularly successful at detecting changes in CI resulting from high levels of contaminants.
1. Many aspects of the social environment affect hypothalamic-pituitary-adrenal (HPA) axis function and increase circulating glucocorticoid concentrations. In this review, we examine the relationships between the social environment and the function of the HPA axis in vertebrates. 2. First, we explore the effects of the social environment on glucocorticoid secretion in territorial (primarily non-social) species, with an emphasis on the effects of variation in population density, as modified by environmental factors such as predation risk and food availability. In general, high population density or frequent territorial intrusions are associated with increased glucocorticoid secretion in a wide range of taxa, including mammals, birds, fish and reptiles, although there is considerable variability across species. 3. Second, we consider the effects of social interactions and dominance rank on glucocorticoid secretion in social species, mostly in birds and mammals. We review studies that have detected an association between social status and glucocorticoid levels — sometimes with higher glucocorticoid levels in low-ranking individuals, and sometimes with higher glucocorticoid levels in dominant individuals. The relationship between dominance and glucocorticoid levels varies among species, populations and years, in a manner that depends on the stability of the social hierarchy, environmental conditions, the type of breeding system, and the manner in which high rank is obtained and maintained. 4. Finally, we discuss the concept of allostasis and consider interactions between social effects and other environmental factors, noting that there is relatively little research on these interactions to date. For both non-social and social species, we identify priorities of future research. These priorities include more complete descriptions of HPA function that move beyond measurements of basal glucocorticoid concentrations (which will generally require field experiments), to studies that examine organizational effects of social stressors, that directly test the relationship between HPA function and fitness, and that examine how glucocorticoid responses affect population dynamics. 5. Although several lines of evidence suggest that glucocorticoid responses can affect the fitness of individuals and therefore can alter the dynamics of populations, the effect of glucocorticoid responses on population dynamics remains essentially unstudied.
1. In this essay I summarize current trends in the evolutionary ecology of plant defence, while advocating for approaches that integrate community ecology with specific tests of classic evolutionary hypotheses. Several conclusions emerge. 2. The microevolution of defence is perhaps best studied by reciprocal transplant experiments of differentiated plant populations while simultaneously manipulating the presence of the herbivore(s) hypothesized to be the agent(s) of natural selection. 3. Although there is continued interest in the costs of defence, I argue that some empirical approaches to estimating costs (e.g. genetic engineering) may provide limited insight into evolutionary processes. 4. Essentially all plants employ several different lines of defence against herbivory. It is thus time to abandon searching for single silver bullet traits and the simple trade-off model (where traits are arbitrarily expected to negatively covary across genotypes or species). We still know very little about which trait combinations are most effective and have repeatedly evolved together. Thus, some of our prominent theories (e.g. a predicted trade-off between direct and indirect defence) need to be revised. 5. Studies of the macroevolution of plant defence are enjoying renewed interest due to available phylogenies and analytical methods. Although general trends are not currently surmisable, we will soon have strong case studies evaluating both biotic and abiotic drivers of convergent evolution in defence strategies and the role of defence evolution in the adaptive radiation of plant lineages. 6. The evolution of specificity is proposed as a final frontier in understanding complexity in plant-herbivore interactions. Although it is abundantly clear that plants can deploy highly specific defensive responses that are differentially perceived by herbivore species, how such responses evolve and are physiologically regulated remains an important gap. Relatively straightforward methodologies are now available to close the loop between plant perception of herbivory, hormonal responses, and production of defensive end-products across genotype or species.
Trait‐based approaches are increasingly being used to test mechanisms underlying species assemblages and biotic interactions across a wide range of organisms including terrestrial arthropods and to investigate consequences for ecosystem processes. Such an approach relies on the standardized measurement of functional traits that can be applied across taxa and regions. Currently, however, unified methods of trait measurements are lacking for terrestrial arthropods and related macroinvertebrates (terrestrial invertebrates hereafter). Here, we present a comprehensive review and detailed protocol for a set of 29 traits known to be sensitive to global stressors and to affect ecosystem processes and services. We give recommendations how to measure these traits under standardized conditions across various terrestrial invertebrate taxonomic groups. We provide considerations and approaches that apply to almost all traits described, such as the selection of species and individuals needed for the measurements, the importance of intraspecific trait variability, how many populations or communities to sample and over which spatial scales. The approaches outlined here provide a means to improve the reliability and predictive power of functional traits to explain community assembly, species diversity patterns and ecosystem processes and services within and across taxa and trophic levels, allowing comparison of studies and running meta‐analyses across regions and ecosystems. This handbook is a crucial first step towards standardizing trait methodology across the most studied terrestrial invertebrate groups, and the protocols are aimed to balance general applicability and requirements for special cases or particular taxa. Therefore, we envision this handbook as a common platform to which researchers can further provide methodological input for additional special cases. A lay summary is available for this article. Lay Summary
1. Oxidative stress is usually defined as an imbalance arising when the rate of production of reactive oxygen species (ROS) exceeds the capacity of the antioxidant defence and repair mechanisms, leading to oxidative damage to biomolecules, but the concept can be expanded to include the disruption of reduction : oxidation (redox) reactions involved in cellular signalling. In this review, we consider how the need to circumvent oxidation may shape the phenotypes of organisms throughout their life and that of their offspring, underpinning a diverse range of life‐history trade‐offs. 2. A recent explosion of interest in this field has shown that both ROS production and the capacity of animals to deal with it change from early development through to adulthood, and vary with environmental conditions and lifestyle. Oxidative stress may both stimulate and be caused by reproduction, although direct evidence of either process is surprisingly weak. Many forms of secondary sexual traits may signal the individual’s oxidative balance to potential mates, but the underlying mechanisms are still debated. 3. Germline cells may be especially vulnerable to oxidative stress, leading to transgenerational effects on offspring viability and possible consequences for the evolution of mate choice. 4. Both antioxidant defences and the ability to repair oxidative damage tend to decline with old age, contributing to cellular and whole organism senescence. This increasing vulnerability to oxidative stress, although little studied, appears especially marked in sexually selected traits. 5. Challenges for the future include the incorporation of longitudinal approaches into experiments that analyse oxidative balance over an individual’s lifetime (preferably under near‐natural conditions), the exploration of the genetic basis for trade‐offs involving oxidative stress, the assimilation of current redox signalling knowledge, and the study of the consequences of heritable oxidative damage to germline DNA.
1. Although secondary metabolites are recognized as fundamental to the defence of plants against insect and mammalian herbivores, their relative importance compared to other potential defensive plant traits (e.g. physical resistance, gross morphology, life-history, primary chemistry and physiology) are not well understood. 2. We conducted a meta-analysis to answer the question: What types of genetically variable plant traits most strongly predict resistance against herbivores? We performed a comprehensive literature search and obtained 499 separate measurements of the strength of covariation (measured as genetic correlations) between plant traits and herbivore susceptibility – these were extracted from 72 studies involving 19 plant families. 3. Surprisingly, we found no overall association between the concentrations of secondary metabolites and herbivore susceptibility – plant traits other than secondary metabolites most strongly predicted herbivore susceptibility. Specifically, genetic variation in life-history traits (e.g. flowering time, growth rate) consistently exhibited the strongest genetic correlations with susceptibility. Genetic variation in gross morphological traits (e.g. no. branches, plant size) and physical resistance traits (e.g. latex, trichomes) were also frequently correlated with variation in herbivore susceptibility, but these relationships depended on attributes of the herbivores (e.g. feeding guild) and plants (e.g. longevity). 4. These results call into question the conventional wisdom that secondary metabolites are the most important anti-herbivore defence of plants. We propose the hypothesis that herbivores select most strongly on genetic variation in life-history, morphological and physical resistance traits, but the greater pleiotropic effects of genes controlling these traits impose strong constraints on their evolution. Meanwhile, secondary metabolites could have evolved to be important defensive mechanisms not because they have the largest effect on herbivores, but because the constraints on their evolution are the weakest.
1. Glucocorticoids hormones (GCs) are intuitively important for mediation of age-dependent vertebrate life-history transitions through their effects on ontogeny alongside underpinning variation in life-history traits and trade-offs in vertebrates. These concepts largely derive from the ability of GCs to alter energy allocation, physiology and behaviour that influences key life-history traits involving age-specific life-history transitions, reproduction and survival. 2. Studies across vertebrates have shown that the neuroendocrine stress axis plays a role in the developmental processes that lead up to age-specific early life-history transitions. While environmental sensitivity of the stress axis allows for it to modulate the timing of these transitions within species, little is known as to how variation in stress axis function has been adapted to produce interspecific variation in the timing of life-history transitions. 3. Our assessment of the literature confirms that of previous reviews that there is only equivocal evidence for correlative or direct functional relationships between GCs and variation in reproduction and survival. We conclude that the relationships between GCs and life-history traits are complex and general patterns cannot be easily discerned with current research approaches and experimental designs. 4. We identify several future research directions including: (i) integration of proximate and ultimate measures, including longitudinal studies that measure effects of GCs on more than one life-history trait or in multiple environmental contexts, to test explicit hypotheses about how GCs and life-history variation are related and (ii) the measurement of additional factors that modulate the effects of GCs on life-history traits (e.g. GC receptors and binding protein levels) to better infer neurendocrine stress axis actions. 5. Conceptual models of HPA/I axis actions, such as allostatic load and reactive scope, to some extent explicitly predict the role of GCs in a life-history context, but are descriptive in nature. We propose that GC effects on life-history transitions, survival probabilities and fecundity can be modelled in existing quantitative demographic frameworks to improve our understanding of how GC variation influences life-history evolution and GC-mediated effects on population dynamics
It is increasingly acknowledged that plant-soil feedbacks may play an important role in driving the composition of plant communities and functioning of terrestrial ecosystems. However, the mechanistic understanding of plant-soil feedbacks, as well as their roles in natural ecosystems in proportion to other possible drivers, is still in its infancy. Such knowledge will enhance our capacity to determine the contribution of plant-soil feedback to community and ecosystem responses under global environmental change. Here, we review how plant-soil feedbacks may develop under extreme drought and precipitation events, CO2 and nitrogen enrichment, temperature increase, land use change and plant species loss vs. gain. We present a framework for opening the black box of soil' considering the responses of the various biotic components (enemies, symbionts and decomposers) of plant-soil feedback to the global environmental changes, and we discuss how to integrate these components to understand and predict the net effects of plant-soil feedbacks under the various scenarios of change. To gain an understanding of how plant-soil feedback plays out in realistic settings, we also use the framework to discuss its interaction with other drivers of plant community composition, including competition, facilitation, herbivory, and soil physical and chemical properties. We conclude that understanding the role that plant-soil feedback plays in shaping the responses of plant community composition and ecosystem processes to global environmental changes requires unravelling the individual contributions of enemies, symbionts and decomposers. These biotic factors may show different response rates and strengths, thereby resulting in different net magnitudes and directions of plant-soil feedbacks under various scenarios of global change. We also need tests of plant-soil feedback under more realistic conditions to determine its contribution to changes in patterns and processes in the field, both at ecologically and evolutionary relevant time-scales.
1. Several theories have provided a framework for understanding variation in plant defence against herbivores. Among them, the plant apparency theory and the resource availability hypothesis (RAH) have aimed to explain the patterns of defence investment and the selective pressures that have led to the variety of defensive strategies across species. Here we provide a historical review of both theories, present evidence that shaped their development and contrast their predictions. 2. We present the results of a meta-analysis of the utility of the RAH 25 years after it was proposed and compare it to apparency theory. We performed a meta-analysis of 50 studies that have examined plant growth, defences and herbivory in relation to resource availability across latitude and ontogeny. Specifically, we tested four predictions that follow the RAH: (i) species adapted to resource-rich environments have intrinsically faster growth rates than species adapted to resourcepoor environments; (ii) fast-growing species have shorter leaf lifetimes than slow-growing species; (iii) fast-growing species have lower amounts of constitutive defences than slow-growing species; and (iv) fast-growing species support higher herbivory rates than slow-growing species. 3. Our results confirm the predictions that species adapted to resource-poor environments grow inherently more slowly, invest more in constitutive defences and support lower herbivory than species from more productive habitats. Our data also showed that variation in growth rate among species better explains the differences in herbivory than variation in apparency, suggesting that the evolution of different defensive strategies across species is resource, rather than herbivore driven. We also found that the application of this theory appears robust across latitude and ontogeny, as the magnitude of the effect sizes for most of the predictions did not vary significantly between ecosystems or across ontogenic stages. 4. We conclude that the RAH has served as a valid framework for investigating the patterns of plant defences and that its applicability is quite general.
It has been suggested that diverse forests utilize canopy space more efficiently than species‐poor ones, as mixing species with complementary architectural and physiological traits allows trees to pack more densely. However, whether positive canopy packing–diversity relationships are a general feature of forests remains unclear. Using crown allometric data collected for 12 939 trees from permanent forest plots across Europe, we test (i) whether diversity promotes canopy packing across forest types and (ii) whether increased canopy packing occurs primarily through vertical stratification of tree crowns or as a result of intraspecific plasticity in crown morphology. We found that canopy packing efficiency increased markedly in response to species richness across a range of forest types and species combinations. Positive canopy packing–diversity relationships were primarily driven by the fact that trees growing in mixture had sizably larger crowns (38% on average) than those in monoculture. The ability of trees to plastically adapt the shape and size of their crowns in response to changes in local competitive environment is critical in allowing mixed‐species forests to optimize the use of canopy space. By promoting the development of denser and more structurally complex canopies, species mixing can strongly impact nutrient cycling and storage in forest ecosystems. Lay Summary