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The presence or absence of sex can have a strong influence on the processes whereby species arise. Yet, the mechanistic underpinnings of this influence are poorly understood. To gain insights into the mechanisms whereby the reproductive mode may influence ecological diversification, we investigate how natural selection, genetic mixing, and the reproductive mode interact and how this interaction affects the evolutionary dynamics of diversifying lineages. To do so, we analyze models of ecological diversification for sexual and asexual lineages, in which diversification is driven by intraspecific resource competition. We find that the reproductive mode strongly influences the diversification rate and, thus, the ensuing diversity of a lineage. Our results reveal that ecologically-based selection is stronger in asexual lineages because asexual organisms have a higher reproductive potential than sexual ones. This promotes faster diversification in asexual lineages. However, a small amount of genetic mixing accelerates the trait expansion process in sexual lineages, overturning the effect of ecologically-based selection alone and enabling a faster niche occupancy than asexual lineages. As a consequence, sexual lineages can occupy more ecological niches, eventually resulting in higher diversity. This suggests that sexual reproduction may be widespread among species because it increases the rate of diversification. [ABSTRACT FROM AUTHOR]

Understanding how biotic interactions shape ecosystems and impact their functioning, resilience and biodiversity has been a sustained research priority in ecology. Yet, traditional assessments of ecological complexity typically focus on species-species interactions that mediate a particular function (e.g., pollination), overlooking both the synergistic effect that multiple functions might develop as well as the resulting species-function participation patterns that emerge in ecosystems that harbor multiple ecological functions. Here we propose a mathematical framework that integrates various types of biotic interactions observed between different species. Its application to recently collected data of an islet ecosystem—reporting 1537 interactions between 691 plants, animals and fungi across six different functions (pollination, herbivory, seed dispersal, decomposition, nutrient uptake, and fungal pathogenicity)—unveils a non-random, nested structure in the way plant species participate across different functions. The framework further allows us to identify a ranking of species and functions, where woody shrubs and fungal decomposition emerge as keystone actors whose removal have a larger-than-random effect on secondary extinctions. The dual insight—from species and functional perspectives—offered by the framework opens the door to a richer quantification of ecosystem complexity and to better calibrate the influence of multifunctionality on ecosystem functioning and biodiversity. Studies of species interactions tend to focus on single ecological functions. Here, the authors show that plant species tend to participate across different ecological functions in a non-random, nested structure, and some species and functions emerge as unexpected keystone actors of the multifunctional ecosystem. [ABSTRACT FROM AUTHOR]

Biodiversity can increase in both high- and low-connected landscapes. However, we lack predictions related to biodiversity dynamics when accounting for the temporal heterogeneity in the connections among the habitats of a landscape. Here, we study the relationship between fluctuations in landscape connectivity and biodiversity dynamics at local and regional scales. We contrast predictions about species richness between landscapes with and without fluctuations in connectivity. Our results show that local (α) and regional (γ) richness can increase together in dynamic landscapes characterized by periodic connectivity, clarifying empirical findings of high biodiversity in both low and high-connected landscapes. Our results also suggest that fluctuations in connectivity increase the overall number of species coexisting in dynamic landscapes when compared with static landscapes with no fluctuations in connectivity. Extending metacommunity theory, by including fluctuations in landscape connectivity, can thus provide new testable predictions about species diversity across broad spatiotemporal scales in rapidly changing landscapes. [ABSTRACT FROM AUTHOR]

Cross‐species communication, where signals are sent by one species and perceived by others, is one of the most intriguing types of communication that functionally links different species to form complex ecological networks. Global change and human activity can affect communication by increasing fluctuations in species composition and phenology, altering signal profiles and intensity, and introducing noise. So far, most studies on cross‐species communication have focused on a few specific species isolated from ecological communities. Scaling up investigations of cross‐species communication to the community level is currently hampered by a lack of conceptual and practical methodologies. Here, we propose an interdisciplinary framework based on information theory to investigate mechanisms shaping cross‐species communication at the community level. We use plants and insects, the cornerstones of most ecosystems, as a showcase and focus on chemical communication as the key communication channel. We first introduce some basic concepts of information theory, then we illustrate information patterns in plant–insect chemical communication, followed by a further exploration of how to integrate information theory into ecological and evolutionary processes to form testable mechanistic hypotheses. We conclude by highlighting the importance of community‐level information as a means to better understand the maintenance and workings of ecological systems, especially during rapid global change. [ABSTRACT FROM AUTHOR]

Species interactions lie at the heart of many theories of macroevolution, from adaptive radiation to the Red Queen. Although some theories describe the imprint that interactions will have over long timescales, we are still missing a comprehensive understanding of the effects of interactions on macroevolution. Current research shows strong evidence for the impact of interactions on macroevolutionary patterns of trait evolution and diversification, yet many macroevolutionary studies have only a tenuous relationship to ecological studies of interactions over shorter timescales. We review current research in this area, highlighting approaches that explicitly model species interactions and connect them to broad‐scale macroevolutionary patterns. We also suggest that progress has been made by taking an integrative interdisciplinary look at individual clades. We focus on African cichlids as a case study of how this approach can be fruitful. Overall, although the evidence for species interactions shaping macroevolution is strong, further work using integrative and model‐based approaches is needed to spur progress towards understanding the complex dynamics that structure communities over time and space. [ABSTRACT FROM AUTHOR]

Theoretical models pertaining to feedbacks between ecological and evolutionary processes are prevalent in multiple biological fields. An integrative overview is currently lacking, due to little crosstalk between the fields and the use of different methodological approaches.Here, we review a wide range of models of eco‐evolutionary feedbacks and highlight their underlying assumptions. We discuss models where feedbacks occur both within and between hierarchical levels of ecosystems, including populations, communities and abiotic environments, and consider feedbacks across spatial scales.Identifying the commonalities among feedback models, and the underlying assumptions, helps us better understand the mechanistic basis of eco‐evolutionary feedbacks. Eco‐evolutionary feedbacks can be readily modelled by coupling demographic and evolutionary formalisms. We provide an overview of these approaches and suggest future integrative modelling avenues.Our overview highlights that eco‐evolutionary feedbacks have been incorporated in theoretical work for nearly a century. Yet, this work does not always include the notion of rapid evolution or concurrent ecological and evolutionary time scales. We show the importance of density‐ and frequency‐dependent selection for feedbacks, as well as the importance of dispersal as a central linking trait between ecology and evolution in a spatial context. A plain language summary is available for this article. Plain Language Summary [ABSTRACT FROM AUTHOR]

ABSTRACT: Knowledge of species composition and their interactions, in the form of interaction networks, is required to understand processes shaping their distribution over time and space. As such, comparing ecological networks along environmental gradients represents a promising new research avenue to understand the organization of life. Variation in the position and intensity of links within networks along environmental gradients may be driven by turnover in species composition, by variation in species abundances and by abiotic influences on species interactions. While investigating changes in species composition has a long tradition, so far only a limited number of studies have examined changes in species interactions between networks, often with differing approaches. Here, we review studies investigating variation in network structures along environmental gradients, highlighting how methodological decisions about standardization can influence their conclusions. Due to their complexity, variation among ecological networks is frequently studied using properties that summarize the distribution or topology of interactions such as number of links, connectance, or modularity. These properties can either be compared directly or using a procedure of standardization. While measures of network structure can be directly related to changes along environmental gradients, standardization is frequently used to facilitate interpretation of variation in network properties by controlling for some co‐variables, or via null models. Null models allow comparing the deviation of empirical networks from random expectations and are expected to provide a more mechanistic understanding of the factors shaping ecological networks when they are coupled with functional traits. As an illustration, we compare approaches to quantify the role of trait matching in driving the structure of plant–hummingbird mutualistic networks, i.e. a direct comparison, standardized by null models and hypothesis‐based metaweb. Overall, our analysis warns against a comparison of studies that rely on distinct forms of standardization, as they are likely to highlight different signals. Fostering a better understanding of the analytical tools available and the signal they detect will help produce deeper insights into how and why ecological networks vary along environmental gradients. [ABSTRACT FROM AUTHOR]

Dispersal and the underlying movement behaviour are processes of pivotal importance for understanding and predicting metapopulation and metacommunity dynamics. Generally, dispersal decisions are condition-dependent and rely on information in the broad sense, like the presence of conspecifics. However, studies on metacommunities that include interspecific interactions generally disregard condition-dependence. Therefore, it remains unclear whether and how dispersal in metacommunities is condition-dependent and whether rules derived from single-species contexts can be scaled up to (meta)communities. Using experimental protist metacommunities, we show how dispersal and movement depend on and are adjusted by the strength of interspecific interactions. We found that the predicting movement and dispersal in metacommunities requires knowledge on behavioural responses to intra- and interspecific interaction strengths. Consequently, metacommunity dynamics inferred directly from single-species metapopulations without taking interspecific interactions into account are likely flawed. Our work identifies the significance of condition-dependence for understanding metacommunity dynamics, stability and the coexistence and distribution of species. [ABSTRACT FROM AUTHOR]

The determinants that shape the distribution of diversity of life on Earth have been long discussed and many mechanisms underlying its formation have been proposed. Yet connecting the biogeography of hot and cold spots of diversification and current biodiversity patterns to the microevolutionary processes remains largely unexplored. Here, we combine a landscape genetics model based on demographic stochasticity with a speciation model that can be interpreted as a model of the evolution of premating incompatibility or assortative mating to map diversification rates in a spatial context. We show that landscape structure and the intensity and directionality of gene flow strongly influence the formation of hot and cold spots and its connection to patterns in species richness. Specifically, hot and cold spots form in landscapes in which gene flow is sufficiently strongly structured that the metacommunity nearly breaks up into several disconnected metacommunities. In such a landscape structure, speciation hot spots originate in the center or in the periphery of the landscape depending on whether the direction of gene flow is from the periphery to the center or viceversa, respectively. However, for any given level of gene flow intensity, diversification rates are approximately twice higher in the center than in the periphery of the landscape. These results suggest that sinks may form diversification hot spots with higher probability than sources, in particular, those sinks surrounded by highly diversified sources in different locations of the landscape. Joining mechanistically microevolutionary and macroevolutionary processes on landscapes present many fascinating challenges and opportunities to connect the biogeography of diversification with biodiversity dynamics. [ABSTRACT FROM AUTHOR]

Studies of food webs suggest that limited nonrandom dispersal can play an important role in structuring food webs. It is not clear, however, whether density-dependent dispersal fits empirical patterns of food webs better than density-independent dispersal. Here, we study a spatially distributed food web, using a series of population-dispersal models that contrast density-independent and density-dependent dispersal in landscapes where sampled sites are either homogeneously or heterogeneously distributed. These models are fitted to empirical data, allowing us to infer mechanisms that are consistent with the data. Our results show that models with density-dependent dispersal fit the α, β, and γ tritrophic richness observed in empirical data best. Our results also show that density-dependent dispersal leads to a critical distance threshold beyond which site similarity (i.e., α tritrophic richness) starts to decrease much faster. Such a threshold can also be detected in the empirical data. In contrast, models with density-independent dispersal do not predict such a threshold. Moreover, preferential dispersal from more centrally located sites to peripheral sites does not provide a better fit to empirical data when compared with symmetric dispersal between sites. Our results suggest that nonrandom dispersal in heterogeneous landscapes is an important driver that shapes local and regional richness (i.e., a and g tritrophic richness, respectively) as well as the distance-decay relationship (i.e., β tritrophic richness) in food webs. [ABSTRACT FROM AUTHOR]

Most empirical and theoretical studies have shown that sex increases the rate of evolution, although evidence of sex constraining genomic and epigenetic variation and slowing down evolution also exists. Faster rates with sex have been attributed to new gene combinations, removal of deleterious mutations, and adaptation to heterogeneous environments. Slower rates with sex have been attributed to removal of major genetic rearrangements, the cost of finding a mate, vulnerability to predation, and exposure to sexually transmitted diseases. Whether sex speeds or slows evolution, the connection between reproductive mode, the evolutionary rate, and species diversity remains largely unexplored. Here we present a spatially explicit model of ecological and evolutionary dynamics based on DNA sequence change to study the connection between mutation, speciation, and the resulting biodiversity in sexual and asexual populations. We show that faster speciation can decrease the abundance of newly formed species and thus decrease long-term biodiversity. In this way, sex can reduce diversity relative to asexual populations, because it leads to a higher rate of production of new species, but with lower abundances. Our results show that reproductive mode and the mechanisms underlying it can alter the link between mutation, evolutionary rate, speciation and biodiversity and we suggest that a high rate of evolution may not be required to yield high biodiversity. [ABSTRACT FROM AUTHOR]

Numerous evolutionary studies have sought to explain the distribution of diversity across the limbs of the tree of life. At the same time, ecological studies have sought to explain differences in diversity and relative abundance within and among ecological communities. Traditionally, these patterns have been considered separately, but models that consider processes operating at the level of individuals, such as neutral biodiversity theory (NBT), can provide a link between them. Here, we compare evolutionary dynamics across a suite of NBT models. We show that NBT can yield phylogenetic tree topologies with imbalance closely resembling empirical observations. In general, metacommunities that exhibit greater disparity in abundance are characterized by more imbalanced phylogenetic trees. However, NBT fails to capture the tempo of diversification as represented by the distribution of branching events through time. We suggest that population-level processes might therefore help explain the asymmetry of phylogenetic trees, but that tree shape might mislead estimates of evolutionary rates unless the diversification process is modeled explicitly. [ABSTRACT FROM AUTHOR]

Most empirical studies support a decline in speciation rates through time, although evidence for constant speciation rates also exists. Declining rates have been explained by invoking pre-existing niches, whereas constant rates have been attributed to non-adaptive processes such as sexual selection and mutation. Trends in speciation rate and the processes underlying it remain unclear, representing a critical information gap in understanding patterns of global diversity. Here we show that the temporal trend in the speciation rate can also be explained by frequency-dependent selection. We construct a frequency-dependent and DNA sequence-based model of speciation. We compare our model to empirical diversity patterns observed for cichlid fish and Darwin's finches, two classic systems for which speciation rates and richness data exist. Negative frequency-dependent selection predicts well both the declining speciation rate found in cichlid fish and explains their species richness. For groups like the Darwin's finches, in which speciation rates are constant and diversity is lower, speciation rate is better explained by a model without frequency-dependent selection. Our analysis shows that differences in diversity may be driven by incipient species abundance with frequency-dependent selection. Our results demonstrate that genetic-distance-based speciation and frequency-dependent selection are sufficient to explain the high diversity observed in natural systems and, importantly, predict decay through time in speciation rate in the absence of pre-existing niches. [ABSTRACT FROM AUTHOR]

Most studies on ecological networks consider only a single interaction type (e.g. competitive, predatory or mutualistic), and try to developrules for system stability based exclusively on properties of this interaction type. However, the stability of ecological networks may be more dependent on the way different interaction types are combined in real communities. To address this issue, we start by compiling an ecological network in the Doñana Biological Reserve, southern Spain, with 390 species and 798 mu-tualistic and antagonistic interactions. We characterize network structure by looking at how mutualistic and antagonistic interactions are combined across all plant species. Both the ratio of mutualistic to antagonistic interactions per plant, and the number of basic modules with an antagonistic and a mutualistic interaction are very heterogeneous across plant species, with a few plant species showing very high values for these parameters. To assess the implications of these network patterns on species diversity, we study analytically and by simulation a model of this ecological network. We find that the observed correlation between strong interaction strengths and high mutualistic to antagonistic ratios in a few plant species significantly increases community diversity. Thus, to predict the persistence of biodiversity we need to understand how interaction strength and the architecture of ecological networks with different interaction types are combined. [ABSTRACT FROM AUTHOR]

Spatio-temporal patterns of predator aggregations within their settlement areas (i.e. temporary settling zones used during dispersal or, more generally, foraging patches) were studied. By integrating the main behavioural rules of juveniles of Spanish imperial eagles Aquila adalberti during dispersal with the temporal availability of the eagles’ main prey, we have developed several individual-based models under different simulation scenarios. The results suggest that the spatially synchrony of the aggregation patterns that we observed were derived from a combination of behavioural responses of the prey and the distances between available settlement areas. These results are discussed in the context of predator–prey relationships, optimal foraging and self-organizing processes. [ABSTRACT FROM AUTHOR]

There are two common approaches to food webs. On the one hand, empirical studies have described aggregate statistical measures of many-species food webs. On the other hand, theoretical studies have explored the dynamic properties of simple tri-trophic food chains (i.e., trophic modules). The question remains to what extent results based on simple modules are relevant for whole food webs. Here we bridge between these two independent research agendas by exploring the relative frequency of different trophic modules in the five most resolved food webs. While apparent competition and intraguild predation are overrepresented when compared to a suite of null models, the frequency of omnivory highly varies across communities. Inferences about the representation of modules may also depend on the null model used for statistical significance. [ABSTRACT FROM AUTHOR]

The stability of ecological communities largely depends on the strength of interactions between predators and their prey. Here we show that these interaction strengths are structured nonrandomly in a large Caribbean marine food web. Specifically, the cooccurrence of strong interactions on two consecutive levels of food chains occurs less frequently than expected by chance. Even when they occur, these strongly interacting chains are accompanied by strong omnivory more often than expected by chance. By using a food web model, we show that these interaction strength combinations reduce the likelihood of trophic cascades after the overfishing of top predators. However, fishing selectively removes predators that are overrepresented in strongly interacting chains. Hence, the potential for strong community-wide effects remains a threat. [ABSTRACT FROM AUTHOR]

Most studies of plant-animal mutualisms involve a small number of species. There is almost no information on the structural organization of species-rich mutualistic networks despite its potential importance for the maintenance of diversity. Here we analyze 52 mutualistic networks and show that they are highly nested; that is, the more specialist species interact only with proper subsets of those species interacting with the more generalists. This assembly pattern generates highly asymmetrical interactions and organizes the community cohesively around a central core of interactions. Thus, mutualistic networks are neither randomly assembled nor organized in compartments arising from tight, parallel specialization. Furthermore, nestedness increases with the complexity (number of interactions) of the network: for a given number of species, communities with more interactions are significantly more nested. Our results indicate a nonrandom pattern of community organization that may be relevant for our understanding of the organization and persistence of biodiversity. [ABSTRACT FROM AUTHOR]

Abstract Recent studies of biological networks have focused on the distribution of the number of links per node. However, the connectivity distribution does not uncover all the complexity of their topology. Here, we analyse the relation between the connectivity of a species and the average connectivity of its nearest neighbours in three of the most resolved community food webs. We compare the pattern arising with the one recently reported for protein networks and for a simple null model of a random network. Whereas two highly connected nodes are unlikely to be connected between each other in protein networks, the reverse happens in food webs. We discuss this difference in organization in relation to the robustness of biological networks to different types of perturbation. [ABSTRACT FROM AUTHOR]

In this paper we explore simple food web models to study how metacommunity structure affects species response to habitat loss. We find that patch abundances and extinction thresholds vary according to the kind of food web. Second, for intermediate species, a slight decrease in the exploration cost of the better competitor has a strong effect on the extinction threshold of the poorer competitor. When predicting extinction risk one should consider not only the amount of habitat destroyed, but also the structure of the food web in which species are embedded. Both direct and indirect interactions are critical for predicting the consequences of habitat destruction. [ABSTRACT FROM AUTHOR]

Parasites can shape the structure and function of ecosystems by influencing both the density and traits of their hosts. Such changes in ecosystems are particularly likely when the host is a predator that mediates the dynamics of trophic cascades. Here, we experimentally tested how parasite load of a small predatory fish, the threespine stickleback, can affect the occurrence and strength of trophic cascades and ecosystem functioning. In a factorial mesocosm experiment, we manipulated the density of stickleback (low vs. high), and the level of parasite load (natural vs. reduced). In addition, we used two stickleback populations from different lineages: an eastern European lineage with a more pelagic phenotype (Lake Constance) and a western European lineage with a more benthic phenotype (Lake Geneva). We found that stickleback caused trophic cascades in the pelagic but not the benthic food chain. Evidence for pelagic trophic cascades was stronger in treatments where parasite load of stickleback was reduced with an antihelmintic medication, and where fish originated from Lake Constance (i.e., the more pelagic lineage). A structural equation model revealed that differences in stickleback lineage and parasite load were most likely to impact trophic cascades via changes in the composition, rather than overall biomass, of zooplankton communities. Overall, our results provide experimental evidence that parasites of predators can influence the cascading effects of fish on lower trophic levels with consequences on ecosystem functioning. [ABSTRACT FROM AUTHOR]