Species interactions and plant polyploidy

Polyploidy is a common mode of speciation that can have far‐reaching consequences for plant ecology and evolution. Because polyploidy can induce an array of phenotypic changes, there can be cascading effects on interactions with other species. These interactions, in turn, can have reciprocal effects on polyploid plants, potentially impacting their establishment and persistence. Although there is a wealth of information on the genetic and phenotypic effects of polyploidy, the study of species interactions in polyploid plants remains a comparatively young field. Here we reviewed the available evidence for how polyploidy may impact many types of species interactions that range from mutualism to antagonism. Specifically, we focused on three main questions: (1) Does polyploidy directly cause the formation of novel interactions not experienced by diploids, or does it create an opportunity for natural selection to then form novel interactions? (2) Does polyploidy cause consistent, predictable changes in species interactions vs. the evolution of idiosyncratic differences? (3) Does polyploidy lead to greater evolvability in species interactions? From the scarce evidence available, we found that novel interactions are rare but that polyploidy can induce changes in pollinator, herbivore, and pathogen interactions. Although further tests are needed, it is likely that selection following whole‐genome duplication is important in all types of species interaction and that there are circumstances in which polyploidy can enhance the evolvability of interactions with other species.     

Friend or foe: inter‐specific interactions and conflicts of interest within the family

1. Interactions between species can vary from mutually beneficial to evolutionarily neutral to antagonistic, even when the same two species are involved. Similarly, social interactions between members of the same species can lie on a spectrum from conflict to cooperation. 2. The aim of the present study was to investigate whether variation in the two types of social behaviour are interconnected. Is the fitness of the various classes of social partner within species (such as parent and offspring, or male and female) differently affected by interactions with a second species? Moreover, can inter‐specific interactions influence the outcome of social interactions within species? 3. The present experiments focus on the interactions between the burying beetle Nicrophorus vespilloides Herbst and the phoretic mite Poecilochirus carabi G. Canestrini & R. Canestrini. The approach was to measure the fitness of burying beetle mothers, fathers, and offspring after reproduction, which took place either in the presence or absence of mites. 4. We found that male, female, and larval burying beetles derive contrasting fitness costs and benefits from their interactions with the mite, despite sharing a common family environment. From the mite's perspective, its relationship with the burying beetle can, therefore, be simultaneously antagonistic, neutral, and possibly even mutualistic, depending on the particular family member involved. We also found that mites can potentially change the outcome of evolutionary conflicts within the family. 5. We conclude that inter‐specific interactions can explain some of the variation in social interactions seen within species. It is further suggested that intra‐specific interactions might contribute to variation in the outcome of interactions between species.     

Contrasting effects of biotic interactions on richness and distribution of vascular plants,bryophytes and lichens in an arctic–alpine landscape

Biotic interactions may strongly affect the distribution of individual species and the resulting patterns of species richness. However, the impacts can vary depending on the species or taxa examined, suggesting that the influences of interactions on species distributions and diversity are not always straightforward and can be taxon-contingent. The aim of this study was therefore to examine how the importance of biotic interactions varies within a community. We incorporated three biotic predictors (cover of the dominant vascular species) into two correlative species richness modelling frameworks to predict spatial variation in the number of vascular plants, bryophytes and lichens in arctic–alpine Fennoscandia, in N Europe. In addition, predictions based on single-species distribution models were used to determine the nature of the impact (negative vs. positive outcome) of the three dominant species on individual vascular plant, bryophyte and lichen species. Our results suggest that biotic variables can be as important as abiotic variables, but their relative contributions in explaining the richness of sub-dominant species vary among dominant species, species group and the modelling framework implemented. Similarly, the impacts of biotic interactions on individual species varied among the three species groups and dominant species, with the observed patterns partly reflecting species’ biogeographic range. Our study provides additional support for the importance of biotic interactions in modifying arctic–alpine biodiversity patterns and highlights that the impacts of interactions are not constant across taxa or biotic drivers. The influence of biotic interactions, including the taxon contingency and range-based impacts, should therefore be accounted for when developing biodiversity forecasts.     

Predicting species distribution and abundance responses to climate change: why it is essential to include biotic interactions across trophic levels

Current predictions on species responses to climate change strongly rely on projecting altered environmental conditions on species distributions. However, it is increasingly acknowledged that climate change also influences species interactions. We review and synthesize literature information on biotic interactions and use it to argue that the abundance of species and the direction of selection during climate change vary depending on how their trophic interactions become disrupted. Plant abundance can be controlled by aboveground and belowground multitrophic level interactions with herbivores, pathogens, symbionts and their enemies. We discuss how these interactions may alter during climate change and the resulting species range shifts. We suggest conceptual analogies between species responses to climate warming and exotic species introduced in new ranges. There are also important differences: the herbivores, pathogens and mutualistic symbionts of range-expanding species and their enemies may co-migrate, and the continuous gene flow under climate warming can make adaptation in the expansion zone of range expanders different from that of cross-continental exotic species. We conclude that under climate change, results of altered species interactions may vary, ranging from species becoming rare to disproportionately abundant. Taking these possibilities into account will provide a new perspective on predicting species distribution under climate change.     

Multiple stressors on biotic interactions: how climate change and alien species interact to affect pollination

Global change may substantially affect biodiversity and ecosystem functioning but little is known about its effects on essential biotic interactions. Since different environmental drivers rarely act in isolation it is important to consider interactive effects. Here, we focus on how two key drivers of anthropogenic environmental change, climate change and the introduction of alien species, affect plant–pollinator interactions. Based on a literature survey we identify climatically sensitive aspects of species interactions, assess potential effects of climate change on these mechanisms, and derive hypotheses that may form the basis of future research. We find that both climate change and alien species will ultimately lead to the creation of novel communities. In these communities certain interactions may no longer occur while there will also be potential for the emergence of new relationships. Alien species can both partly compensate for the often negative effects of climate change but also amplify them in some cases. Since potential positive effects are often restricted to generalist interactions among species, climate change and alien species in combination can result in significant threats to more specialist interactions involving native species.     

The architecture of weighted mutualistic networks

Several ecosystem services directly depend on mutualistic interactions. In species rich communities, these interactions can be studied using network theory. Current knowledge of mutualistic networks is based mainly on binary links; however, little is known about the role played by the weights of the interactions between species. What new information can be extracted by analyzing weighted mutualistic networks? In performing an exhaustive analysis of the topological properties of 29 weighted mutualistic networks, our results show that the generalist species, defined as those with a larger number of interactions in a network, also have the strongest interactions. Though most interactions of generalists are with specialists, the strongest interactions occur between generalists. As a result and by defining binary and weighted clustering coefficients for bipartite networks, we demonstrate that generalists form strongly‐interconnected groups of species. The existence of these strong clusters reinforces the idea that generalist species govern the coevolution of the whole community.     

Interactions between non-identical prion proteins

Prion formation involves the conversion of soluble proteins into an infectious amyloid form. This process is highly specific, with prion aggregates templating the conversion of identical proteins. However, in some cases non-identical prion proteins can interact to promote or inhibit prion formation or propagation. These interactions affect both the efficiency with which prion diseases are transmitted across species and the normal physiology of yeast prion formation and propagation. Here we examine two types of heterologous prion interactions: interactions between related proteins from different species (the species barrier) and interactions between unrelated prion proteins within a single species. Interestingly, although very subtle changes in protein sequence can significantly reduce or eliminate cross-species prion transmission, in Saccharomyces cerevisiae completely unrelated prion proteins can interact to affect prion formation and propagation.     

Stability of a diamond-shaped module with multiple interaction types

Indirect interactions among species emerge from the complexity of ecological networks and can strongly affect the response of communities to disturbances. To determine these indirect interactions and understand better community dynamics, ecologists focused on the interactions within small sets of species or modules. Thanks to their analytical tractability, modules bring insights on the mechanisms occurring in complex interaction networks. So far, most studies have considered modules with a single type of interaction although numerous species are involved in mutualistic and antagonistic interactions simultaneously. In this study, we analyse the dynamics of a diamond-shaped module with multiple interaction types: two resource species sharing a mutualist and a consumer. We describe the different types of indirect interaction occurring between the resource species and the conditions for a stable coexistence of all species. We show that the nature of indirect interactions between resource species (i.e. apparent facilitation, competition or antagonism), as well as stable coexistence, depend on the species generalism and asymmetry of interactions, or in other words, on the distribution of interaction strengths among species. We further unveil that a balance between mutualistic and antagonistic interactions at the level of resource species favours stable coexistence, and that species are more likely to coexist stably if there is apparent facilitation between the two resource species rather than apparent competition. Our results echo existing knowledge on the trophic diamond-shaped module, and confirm that our understanding of communities combining different interaction types can gain from module analyses.     

Dissecting the role of transitivity and intransitivity on coexistence in competing species networks

It is well established that intransitively assembled interaction networks can support the coexistence of competing species, while transitively assembled (hierarchical) networks are prone to species loss through competitive exclusion. However, as the number of species grows, the complexity of ecological interaction networks grows disproportionately, and species can get involved simultaneously in transitive and intransitive groups of interactions. In such complex networks, the effects of intransitivity on species persistence are not straightforward. Dissecting networks into intransitive/transitive components can help us to understand the complex role that intransitivity may play in supporting species diversity. We show through simulations that those species participating in the largest group of intransitive interactions (the core of the network) have high probabilities of persisting in the long term. However, participation in a group of intransitive interactions other than the core does not always improve persistence. Likewise, participating in transitive interactions does not always decrease persistence because certain species (the satellites) transitively linked to the core have also a high persistence probability. Therefore, when networks contain transitive and intransitive structures, as it can be expected in real ecological networks, the existence of a large intransitive core of species can have a disproportionate positive effect on species richness.     

How do species interactions affect species distribution models?

One of the most promising recent advances in biogeography has been the increased interest and understanding of species distribution models – estimates of the probability that a species is present given environmental data. Unfortunately, such analyses ignore many aspects of ecology, and so are difficult to interpret. In particular, we know that species interactions have a profound influence on distributions, but it is not usually possible to incorporate this knowledge into species distribution models. What is needed is a rigorous understanding of how unmeasured biotic interactions affect the inferences generated by species distribution models. To fill this gap, we develop a general mathematical approach that uses probability theory to determine how unmeasured biotic interactions affect inferences from species distribution models. Using this approach, we reanalyze one of the most important classes of mechanistic models of competition: models of consumer resource dynamics. We determine how measurements of one aspect of the environment – a single environmental variable – can be used to estimate the probability that an environment is suitable with species distribution models. We show that species distribution models, which ignore numerous facets of consumer resource dynamics such as the presence of a competitor or the dynamics of depletable resources, can furnish useful predictions for the probability that an environment is suitable in some circumstances. These results provide a rigorous link between complex mechanistic models of species interactions and species distribution models. In so doing they demonstrate that unmeasured biotic interactions can have strong and counterintuitive consequences on species distribution models.     

Interspecific interactions and range limits: contrasts among interaction types

There is a great deal of interest in the effects of biotic interactions on geographic distributions. Nature contains many different types of biotic interactions (notably mutualism, commensalism, predation, amensalism, and competition), and it is difficult to compare the effects of multiple interaction types on species’ distributions. To resolve this problem, we analyze a general, flexible model of pairwise biotic interactions that can describe all interaction types. In the absence of strong positive feedback, a species’ ability to be present depends on its ability to increase in numbers when it is rare and the species it is interacting with is at equilibrium. This insight leads to counterintuitive conclusions. Notably, we often predict the same range limit when the focal species experiences competition, predation, or amensalism. Similarly, we often predict the same range margin or when the species experiences mutualism, commensalism, or benefits from prey. In the presence of strong positive density-dependent feedback, different species interactions produce different range limits in our model. In all cases, the abiotic environment can indirectly influence the impact of biotic interactions on range limits. We illustrate the implications of this observation by analyzing a stress gradient where biotic interactions are harmful in benign environments but beneficial in stressful environments. Our results emphasize the need to consider the effects of all biotic interactions on species’ range limits and provide a systematic comparison of when biotic interactions affect distributions.     

Nested species interactions promote feasibility over stability during the assembly of a pollinator community

The foundational concepts behind the persistence of ecological communities have been based on two ecological properties: dynamical stability and feasibility. The former is typically regarded as the capacity of a community to return to an original equilibrium state after a perturbation in species abundances and is usually linked to the strength of interspecific interactions. The latter is the capacity to sustain positive abundances on all its constituent species and is linked to both interspecific interactions and species demographic characteristics. Over the last 40 years, theoretical research in ecology has emphasized the search for conditions leading to the dynamical stability of ecological communities, while the conditions leading to feasibility have been overlooked. However, thus far, we have no evidence of whether species interactions are more conditioned by the community''s need to be stable or feasible. Here, we introduce novel quantitative methods and use empirical data to investigate the consequences of species interactions on the dynamical stability and feasibility of mutualistic communities. First, we demonstrate that the more nested the species interactions in a community are, the lower the mutualistic strength that the community can tolerate without losing dynamical stability. Second, we show that high feasibility in a community can be reached either with high mutualistic strength or with highly nested species interactions. Third, we find that during the assembly process of a seasonal pollinator community located at The Zackenberg Research Station (northeastern Greenland), a high feasibility is reached through the nested species interactions established between newcomer and resident species. Our findings imply that nested mutualistic communities promote feasibility over stability, which may suggest that the former can be key for community persistence.     

Influence of predator density on nonindependent effects of multiple predator species

Interactions between multiple predator species are frequent in natural communities and can have important implications for shared prey survival. Predator density may be an important component of these interactions between predator species, as the frequency of interactions between species is largely determined by species density. Here we experimentally examine the importance of predator density for interactions between predator species and subsequent impacts on prey. We show that aggressive interactions between the predatory shore crabs Carcinus maenas and Hemigrapsus sanguineus increased with predator density, yet did not increase as fast as negative interactions between conspecifics. At low density, interactions between conspecific and heterospecific predators had similar inhibitory impacts on predator function, whereas conspecific interference was greater than interference from heterospecifics at high predator density. Thus the impact of conspecific interference at high predator density was sufficient in itself that interactions with a second predator species had no additional impact on per capita predation. Spatial and temporal variability in predator density is a ubiquitous characteristic of natural systems that should be considered in studies of multiple predator species.     

A novel theory to explain species diversity in landscapes: positive frequency dependence and habitat suitability

Theories to explain the diversity of species have required that individual species occupy unique niches and/or vary in their response to environmental factors. Positive interactions within a species, although common in communities, have not been thought to maintain species diversity because in non-spatial models the more abundant species always outcompetes the rarer species. Here, we show, using a stochastic spatial model, that positive intraspecific interactions such as those caused by positive frequency dependence and/or priority effects, can maintain species diversity if interactions between individuals are primarily local and the habitat contains areas that cannot be colonized by any species, such as boulders or other physical obstructions. When intraspecific interactions are primarily neutral, species diversity will eventually erode to a single species. When the landscape is homogeneous (i.e. does not contain areas that cannot be colonized by any species), the presence of strong intraspecific interactions will not maintain diversity.     

Topological keystone species: measures of positional importance in food webs

The local extinction or large fluctuation in abundance of a species may seriously affect other species in the community. The effects spread through the community by direct and indirect interactions. The network perspective on ecology can help map the pathways of these effects, for food webs, the pathways of indirect trophic interactions. Indirect interactions typically decay in intensity as they spread. Therefore, there is a conceptual maximum range in topological space beyond which interactions have no effects, even though all species remain connected. Neither the local characteristics of species, nor the global characteristics of entire webs, suitably quantify this range. We therefore apply intermediate scale indices that reflect the limitations imposed by effect damping in networks. We present a complex analysis of the topological positional importance of species in the Chesapeake Bay web. This web is a carbon-flow network that represents trophic interactions. We present several different indices reflecting different properties and discuss which questions the different indices best answer. We look for the best indices for identifying the key players in ecosystem functioning. Our study contributes to the quantification of relative species importance and provides an exact and a priori determination of a class of candidate keystone species that can inform applied and conservation ecology as well as theoretical concerns.     

Modelling the niche space of desert annuals needs to include positive interactions

The niche is a necessary consideration when estimating habitable area and geographic range of a species. Modellers often examine the fundamental niche and the environmental requirements for plant species, ignoring interactions among species. In deserts, positive plant interactions are important drivers of biodiversity and structure communities through many mechanistic pathways including modifying environmental conditions. Thus, we tested the hypothesis that desert shrubs increase the geographical extent of some annual species because, through modifying the microclimate, they match the niche requirements of beneficiary species. We used the database of the Global Biodiversity Information Facility to construct MaxEnt species distribution models (SDM) with and without reported benefactor species within the Mojave Desert in California. We chose 20 annual species to be modeled including 10 species that had been previously reported in the literature as being facilitated (beneficiary) and 10 that had no record of being facilitated (unreported). Beneficiary annuals co‐occurred significantly more with benefactor shrubs than the unreported annual species. The inclusion of shrubs into SDMs significantly improved model predictability and geographic range for all the beneficiary annual species, but not for the unreported annual species. Thus, positive interactions are species specific and it is possible to determine annual species dependency on benefactor shrubs at the regional scale. The co‐occurrence of benefactor shrubs and annual species can be used as a proxy for facilitation and recent developments in SDM techniques encourage the inclusion of biotic interactions. Species distribution models should include estimates of facilitation because biotic interactions determine the niche of species and can have implications with a changing climate.     

Resource conversion: a generalizable mechanism for resource-mediated positive species interactions

Positive species interactions are ubiquitous in natural communities, but the mechanisms through which they operate are poorly understood. One proposed mechanism is resource conversion – the conversion by a benefactor species of a resource from a resource state that is inaccessible to a potential beneficiary species into a resource state that is accessible. Such conversion often occurs as a byproduct of resource consumption, and sometimes in exchange for non-resource benefits to the benefactor species. At least five known classes of interactions, including both facilitative and mutualistic ones, may be classified as resource conversion interactions. We formulated a generalizable mathematical model for resource conversion interactions and examined two model variants that represent processing chain and nurse plant interactions. We examined the conditions under which these conformed to the stress-gradient hypothesis (SGH), which predicts increased interaction benefits in more stressful environments. These yielded four key insights: 1) resource conversion interactions can be positive (towards the resource recipient) only when facilitator-mediated resource conversion is more efficient than the baseline, spontaneous, facilitator-independent resource conversion; 2) the sign of resource conversion interaction outcomes never switches (e.g. from net positive to net negative) with changing levels of resource availability, when all other parameters are kept constant; 3) processing chain interactions at equilibrium can never be positive in a manner that conforms to the SGH; 4) nurse plant interactions can be positive and conform to the SGH, although the manner in which they do depends largely on how resource stress is defined, and the environmental supply rate of surface soil moisture. The first two insights are likely to be generalizable across all resource conversion interactions. The general agreement of the model with empirical studies suggest that resource conversion is the mechanism underlying the aforementioned interactions, and an ecologically meaningful way of classifying these previously unassociated positive species interactions.     

Theoretical and empirical aspects of gene-culture coevolution

Cultural transmission can be roughly defined as the transfer of information between individuals by social learning (see below). Genes used to be, and for most species still are, the only means available for the accurate transfer of information across generations. In species where cultural transmission has developed, notably the human, interactions can occur between the two inheritance systems. Gene-culture coevolution refers to the evolutionary phenomena that arise from these interactions. As we shall see, these interactions can take various forms.     

Sex,competition and mimicry: an eco-evolutionary model reveals unexpected impacts of ecological interactions on the evolution of phenotypes in sympatry

Phenotypic evolution in sympatric species can be strongly impacted by species interactions, either mutualistic or antagonistic. Heterospecific reproductive behaviours between sympatric species have been shown to favour phenotypic divergence of traits used as sexual cues. Those traits may also be involved in local adaptation or in other types of species interactions and, as a result, undergo complex evolutions across sympatric species. Here we focus on mimicry and study how reproductive interference may impair phenotypic convergence between species with various levels of defence. We use a deterministic model assuming two sympatric species where individuals can display two different warning colour patterns. This eco-evolutionary model explores how ecological interactions shape phenotypic evolution within sympatric species. We investigate the effect of 1) the opposing density-dependent selections exerted on colour patterns by predation and reproductive behaviour and 2) the impact of relative species and phenotype abundances on the fitness costs faced by each individual depending on their species and phenotype. Our model shows that reproductive interference may limit the convergent effect of mimetic interactions and may promote phenotypic divergence between Müllerian mimics. The divergent and convergent evolution of traits also strongly depends on the relative species and phenotype abundances and levels of trophic competition, highlighting how the eco-evolutionary feedbacks between phenotypic evolution and species abundances may result in strikingly different evolutionary routes.     

The rate of environmental fluctuations shapes ecological dynamics in a two‐species microbial system

Species interactions change when the external conditions change. How these changes affect microbial community properties is an open question. We address this question using a two‐species consortium in which species interactions change from exploitation to competition depending on the carbon source provided. We built a mathematical model and calibrated it using single‐species growth measurements. This model predicted that low frequencies of change between carbon sources lead to species loss, while intermediate and high frequencies of change maintained both species. We experimentally confirmed these predictions by growing co‐cultures in fluctuating environments. These findings complement more established concepts of a diversity peak at intermediate disturbance frequencies. They also provide a mechanistic understanding for how the dynamics at the community level emerges from single‐species behaviours and interspecific interactions. Our findings suggest that changes in species interactions can profoundly impact the ecological dynamics and properties of microbial systems.