Predators can influence the host-parasite dynamics of their prey via nonconsumptive effects.

Ecological communities are partly structured by indirect interactions, where one species can indirectly affect another by altering its interactions with a third species. In the absence of direct predation, nonconsumptive effects of predators on prey have important implications for subsequent community interactions. To better understand these interactions, we used a Daphnia-parasite-predator cue system to evaluate if predation risk affects Daphnia responses to a parasite. We investigated the effects of predator cues on two aspects of host-parasite interactions (susceptibility to infection and infection intensity), and whether or not these effects differed between sexes. Our results show that changes in response to predator cues caused an increase in the prevalence and intensity of parasite infections in female predator-exposed Daphnia. Importantly, the magnitude of infection risk depended on how long Daphnia were exposed to the cues. Additionally, heavily infected Daphnia that were constantly exposed to cues produced relatively more offspring. While males were ~5× less likely to become infected compared to females, we were unable to detect effects of predator cues on male Daphnia-parasite interactions. In sum, predators, prey, and their parasites can form complex subnetworks in food webs, necessitating a nuanced understanding of how nonconsumptive effects may mediate these interactions.

Gender underlies the formation of STEM research groups.

Research groups are the cornerstone of scientific research, yet little is known about how these groups are formed and how their organization is influenced by the gender of the research group leader. This represents an important gap in our understanding of the processes shaping gender structure within universities and the academic fields they represent. Here, we report the results of an email survey sent to department chairs and discipline-specific listservs. We received responses from 275 female and 175 male research group leaders. Most respondents were biologists (n = 328) but psychology (n = 27), chemistry (n = 16), physics (n = 32), and mathematics (n = 30) were also relatively well represented. We found that men were self-reported as overrepresented in research groups in the physical sciences, particularly at later career stages. Within biology, male and female group leaders reported supervising a disproportionate number of same-gender trainees (students and postdoctoral fellows), particularly early in their careers. These self-reported patterns were driven primarily by gender-based differences in the pool of students applying to their research groups, while gender differences in acceptance rates played a seemingly smaller role. We discuss the implications of our results for women continuing into the professoriate and for the recruitment of young scientists into research groups.

Ecological and Evolutionary Stochasticity Shape Natural Selection.

The distribution of biodiversity depends on the combined and interactive effects of ecological and evolutionary processes. The joint contribution of these processes has focused almost exclusively on deterministic effects, even though mechanisms that increase the importance of random ecological processes are expected to also increase the importance of random evolutionary processes. Here we manipulate the sizes of old field fragments to generate correlated sampling effects for a focal population (a gall maker) and its enemy community. Traits and communities were more variable in smaller patches. However, because of the preference of some enemies for some trait values (gall sizes), random variation in population mean trait values exacerbated differences in community composition. The random distribution of traits and interactions created predictable but highly variable patterns of natural selection. Our study highlights how stochastic processes can affect ecological and evolutionary processes structuring the strength and direction of selection locally and at larger scales.

Ecological rigidity and the hardness of selection in the wild.

Hutchinson's ecological theater and evolutionary play is a classical view of evolutionary ecology-ecology provides a template in which evolution occurs. An opposing view is that ecological and evolutionary changes are like two actors on a stage, intertwined by density and frequency dependence. These opposing views correspond to hard and soft selection, respectively. Although often presented as diametrically opposed, both types of selection can occur simultaneously, yet we largely lack knowledge of the relative importance of hard versus soft selection in the wild. I use a dataset of 3000 individual gall makers from 15 wild local populations over 5 years to investigate the hardness of selection. I show that enemy attack consistently favors some gall sizes over others (hard selection) but that these biases can be fine-tuned by density and frequency dependence (soft selection). As a result, selection is hard and soft in roughly equal measures, but the importance of each type varies as species interactions shift. I conclude that eco-evolutionary dynamics should occur when a mix of hard and soft selection acts on a population. This work contributes to the rapprochement of disparate views of evolutionary ecology-ecology is neither a rigid theater nor a flexible actor, but instead embodies components of both.

Abundance and traits link predator ontogeny to prey communities.

Function and abundances shape species interactions and thus ecological communities. While communities are often summarized as the mean function of each species, intraspecific variation in traits and thus function is an important driver of community composition. Ontogeny is a common source of intraspecific variation, but while age-related functional changes can alter species interactions, so too can the effects of those functions on the density of the focal organism. For instance, ontogenetic variation can trigger higher levels of cannibalism, reducing abundances and altering interspecific interactions. I manipulate ontogenetic variation in damselfly larvae to show that intraspecific variation can impact communities through two distinct mechanisms. First, within-species differences affect population sizes, and thus indirectly shape communities (indirect effect). In particular, ontogenetic variation resulted in smaller damselfly populations, likely because of increased cannibalism rates, and thus ontogenetically diverse populations had a smaller total effect on their prey. Second, trait variation can affect communities by creating differences in the strength of per capita species interactions (direct effect). In this case, damselfly populations with greater age variation had smaller per capita effects on prey communities. I conclude that ontogeny of a single species can directly and indirectly shape community composition.

Urbanization reshapes a food web.

Cities represent humanity's most intense impact on our planet, with more than half of all humans now residing in urban areas. Indeed, urbanization has well-understood impacts on both individual species and general patterns of biodiversity. However, species do not exist in isolation, but are instead members of complex interaction networks that shape patterns of diversity and influence ecosystem services. Despite the importance of species interaction for creating patterns of diversity, we do not understand how urbanization alters these interactions. Here, we investigate how an interaction network (food web) is reshaped by urbanization. We show that, consistent with theory, cities tend to support less diverse ecological communities, and rare species that interact with few species are particularly sensitive to urbanization. As a result, remnant urban food webs tend to have more interactions per species and greater connectance, creating more integrated interaction networks. We discuss the implications of this food web reshaping for ecological stability, eco-evolutionary dynamics, and the joining of interaction networks and conservation planning. The role of cities in reshaping interaction networks provides an interesting study of food web (dis)assembly, while also shedding light on new approaches to applied conservation issues.

Individual and Population Differences Shape Species Interactions and Natural Selection.

Trait variation is central to our understanding of species interactions, and trait variation arising within species is increasingly recognized as an important component of community ecology. Ecologists generally consider intraspecific variation either among or within populations, yet these differences can interact to create patterns of species interactions. These differences can also affect species interactions by altering processes occurring at distinct scales. Specifically, intraspecific variation may shape species interactions simply by shifting a population's position along a trait-function map or by shifting the relationship between traits and their ecological function. I test these ideas by manipulating within- and among-population intraspecific variation in wild populations of a gall-forming insect before quantifying species interactions and phenotypic selection. Within- and among-population differences in gall size interact to affect attack rates by an enemy community, but among-population differences were far more consequential. Intraspecific differences shaped species interactions by both shifting the position of populations along the trait-function map and altering the relationship between traits and their function, with ultimate consequences for patterns of natural selection. I suggest that intraspecific variation can affect communities and natural selection by acting through individual- and population-level mechanisms.

Trait variation across biological scales shapes community structure and ecosystem function.

Trait variation underlies our understanding of the patterns and importance of biodiversity, yet we have a poor understanding of how variation at different levels of biological organization structures communities and ecosystems. Here, we use a mesocosm experiment to test for the effects of a larval dragonfly functional trait on community and ecosystem dynamics by creating artificial populations to mirror within- and between-population trait variation observed in our study area. Specifically, we manipulate variation in activity rate, a key functional trait shaping food webs, across three levels of biological organization: within-populations (differences in trait variation in a population), among-populations (differences in population mean trait values), and among-species (species-level differences of co-occurring dragonflies). We show that differences in activity rate alter prey communities, trophic cascades, and multiple ecosystem processes. However, trait variation among populations had much larger effects than differences between co-occurring species or even the presence of a predator, whereas within-population variation had a relatively minor impact. Interestingly, combined with earlier work in the same system, our study suggests that the relative importance of species vs. individual level differences for ecosystem functioning will depend on the spatial scale considered. Ecological processes, including biodiversity-ecosystem-functioning relationships, cannot be understood without accounting for trait variation across biological scales of organization, including at fine scales.

Host traits and competitive ability jointly structure disease dynamics and community assembly.

Parasitism and competition are both ubiquitous interactions in ecological communities. The ability of host species to interact directly via competition and indirectly through shared parasites suggests that host traits related to competition and parasitism are likely important in structuring communities and disease dynamics. Specifically, those host traits affecting competition and those mediating parasitism are often correlated either because of trade-offs (in resource acquisition or resource allocation) or condition dependence, yet the consequences of these trait relationships for community and epidemiological dynamics are poorly understood. We conducted a literature review of parasite-related host traits-competitive ability relationships. We found that transmission-competitive ability relationships were most often reported, and that superior competitors exhibited elevated transmission relative to their less-competitive counterparts in nearly 80% of the cases. We also found a significant number of virulence-competitive ability and parasite shedding-competitive ability relationships. We investigated these links by altering the relationship between host competitive ability and three parasite-related traits (transmission, virulence and parasite shedding rates) in a simple model, incorporating competitive asymmetries in a multi-host community. We show that these relationships can lead to a range of different communities. For example, depending on the strength and direction of these distinct trait relationships, we observed communities with anywhere from high parasite prevalence to complete parasite extinction, and either one, two or the maximum of three host species coexisting. Our results suggest that parasite-competitive ability relationships may be common in nature, that further integration of these relationships can produce novel and unexpected community and disease dynamics, and that generalizations may allow for the prediction of how parasitism and competition jointly structure disease and diversity in natural communities.

Indirect Interactions Shape Selection in a Multispecies Food Web.

Species do not live, interact, or evolve in isolation but are instead members of complex ecological communities. In ecological terms, complex multispecies interactions can be understood by considering indirect effects that are mediated by changes in traits and abundances of intermediate species. Interestingly, traits and abundances are also central to our understanding of phenotypic selection, suggesting that indirect effects may be extended to understand evolution in complex communities. Here we explore indirect ecological effects and their evolutionary corollary in a well-understood food web comprising a plant, its herbivores, and enemies that select for opposite defensive phenotypes in one of the herbivores. We show that ecological indirect interactions are mediated by changes to both the traits and the abundances of intermediate species and that these changes ultimately reduce enemy attack and weaken selection. We discuss the generality of the link between indirect effects and selection. We go on to argue that local adaptation and eco-evolutionary feedback may be less likely in complex multispecies food webs than in simpler food chains (e.g., coevolution). Overall, considering selection in complex interaction networks can facilitate the rapprochement of community ecology and evolution.

Sexual dimorphism in a top predator (Notophthalmus viridescens) drives aquatic prey community assembly.

Intraspecific variation can have important consequences for the structure and function of ecological communities, and serves to link community ecology to evolutionary processes. Differences between the sexes are an overwhelmingly common form of intraspecific variation, but its community-level consequences have never been experimentally investigated. Here, we manipulate the sex ratio of a sexually dimorphic predacious newt in aquatic mesocosms, then track their impact on prey communities. Female and male newts preferentially forage in the benthic and pelagic zones, respectively, causing corresponding reductions in prey abundances in those habitats. Sex ratio differences also explained a large proportion (33%) of differences in the composition of entire pond communities. Ultimately, we demonstrate the impact of known patterns of sexual dimorphism in a predator on its prey, uncovering overlooked links between evolutionary adaptation and the structure of contemporary communities. Given the extreme prevalence of sexual dimorphism, we argue that the independent evolution of the sexes will often have important consequences for ecological communities.

Ontogeny and Consistent Individual Differences Mediate Trophic Interactions.

Ecologists use species traits to predict responses to environmental change and, ultimately, to understand the composition of biological communities. However, this ignores known and substantial intraspecific variation that can have important consequences for species interactions and community composition. This within-species variation results from two distinct sources: ontogeny and consistent individual differences. Ontogeny and consistent differences interact to produce phenotypes, but the community-level consequences of this interaction have not been studied. Using larval dragonfly communities, I investigate patterns of intraguild predation by manipulating (1) consistent individual differences in activity rate and (2) the ontogeny of the focal and interacting species. I show that activity rate is a consistent individual trait but that the effect of activity rate on intraguild predation depends on the functional role of an organism in the community (predator or prey). An organism's functional role itself varies across ontogeny of both the focal and interacting individuals. I suggest that ontogeny and consistent individual differences interact to produce intraspecific variation, with consequences for species interactions, communities, and eco-evolutionary dynamics.

Consumer-resource interactions along urbanization gradients drive natural selection.

Urbanization is an important component of global change. Urbanization affects species interactions, but the evolutionary implications are rarely studied. We investigate the evolutionary consequences of a common pattern: the loss of high trophic-level species in urban areas. Using a gall-forming fly, Eurosta solidaginis, and its natural enemies that select for opposite gall sizes, we test for patterns of enemy loss, selection, and local adaptation along five urbanization gradients. Eurosta declined in urban areas, as did predation by birds, which preferentially consume gallmakers that induce large galls. These declines were linked to changes in habitat availability, namely reduced forest cover in urban areas. Conversely, a parasitoid that attacks gallmakers that induce small galls was unaffected by urbanization. Changes in patterns of attack by birds and parasitoids resulted in stronger directional selection, but loss of stabilizing selection in urban areas, a pattern which we suggest may be general. Despite divergent selective regimes, gall size did not very systematically with urbanization, suggesting but not conclusively demonstrating that environmental differences, gene flow, or drift, may have prevented the adaptive divergence of phenotypes. We argue that the evolutionary effects of urbanization will have predictable consequences for patterns of species interactions and natural selection.

Predator macroevolution drives trophic cascades and ecosystem functioning.

Biologists now recognize that ecology can drive evolution, and that evolution in turn produces ecological patterns. I extend this thinking to include longer time scales, suggesting that macroevolutionary transitions can create phenotypic differences among species, which then have predictable impacts on species interactions, community assembly and ecosystem functioning. Repeated speciation can exacerbate these patterns by creating communities with similar phenotypes and hence ecological impacts. Here, I use several experiments to test these ideas in dragonfly larvae that occupy ponds with fish, ponds without fish, or both. I show that macroevolutionary transitions between habitats cause fishless pond species to be more active relative to fish pond specialists, reducing prey abundance, shifting prey community composition and creating stronger trophic cascades. These effects scale up to the community level with predictable consequences for ecosystem multi-functioning. I suggest that macroevolutionary history can have predictable impacts on phenotypic traits, with consequences for interacting species and ecosystems.

Physiology underlies the assembly of ecological communities.

Trait-based community ecology promises an understanding of the factors that determine species abundances and distributions across habitats. However, ecologists are often faced with large suites of potentially important traits, making generalizations across ecosystems and species difficult or even impossible. Here, we hypothesize that key traits structuring ecological communities may be causally dependent on common physiological mechanisms and that elucidating these mechanisms can help us understand the distributions of traits and species across habitats. We test this hypothesis by investigating putatively causal relationships between physiological and behavioral traits at the species and community levels in larvae of 17 species of dragonfly that co-occur at the landscape scale but segregate among lakes. We use tools borrowed from phenotypic selection analyses to show that physiological traits underlie activity rate, which has opposing effects on foraging and predator avoidance behaviors. The effect of activity on these behaviors ultimately shapes species distributions and community composition in habitats with either large-bodied fish or invertebrates as top predators. Remarkably, despite the inherent complexity of ecological communities, the expression of just two biomolecules accounts for a high proportion of the variation in behavioral traits and hence, dragonfly community composition between habitats. We suggest that causal relationships among traits can drive species distributions and community assembly.

Animal behaviour and algal camouflage jointly structure predation and selection.

Trait variation can structure interactions between individuals, thus shaping selection. Although antipredator strategies are an important component of many aquatic systems, how multiple antipredator traits interact to influence consumption and selection remains contentious. Here, I use a common larval dragonfly (Epitheca canis) and its predator (Anax junius) to test for the joint effects of activity rate and algal camouflage on predation and survival selection. I found that active and poorly camouflaged Epitheca were more likely to be consumed, and thus, survival selection favoured inactive and well-camouflaged individuals. Notably, camouflage dampened selection on activity rate, likely by reducing attack rates when Epitheca encountered a predator. Correlational selection is therefore conferred by the ecological interaction of traits, rather than by opposing selection acting on linked traits. I suggest that antipredator traits with different adaptive functions can jointly structure patterns of consumption and selection.

Keystone Individuals Alter Ecological and Evolutionary Consumer-Resource Dynamics.

Intraspecific variation is central to our understanding of evolution and ecology, but these fields generally consider either the mean trait value or its variance. Alternatively, the keystone individual concept from behavioral ecology posits that a single individual with an extreme phenotype can have disproportionate and irreplaceable effects on group dynamics. Here, I generalize this concept to include nonbehavioral traits and broader ecological and evolutionary dynamics. I test for the effects of individuals with extreme phenotypes on the ecology and evolution of a gall-forming fly and its natural enemies that select for opposite gall sizes. Specifically, I introduce a putatively keystone predator-attracting individual gall-maker, hypothesizing that the presence of such an individual should (1) increase gall maker population-level mortality, (2) cause consumer communities to be dominated by species that are most attracted to the keystone individual, (3) increase selection for traits conferring defense against the most common consumer, and (4) weaken patterns of stabilizing selection. I find support for both the ecological and evolutionary consequences of single individuals with extreme phenotypes, suggesting that they can be considered keystone individuals. I discuss the generality of the keystone individual concept, suggesting likely consequences for ecology and evolution.

An invasive herbivore structures plant competitive dynamics.

Species interactions are central to our understanding of ecological communities, but may change rapidly with the introduction of invasive species. Invasive species can alter species interactions and community dynamics directly by having larger detrimental effects on some species than others, or indirectly by changing the ways in which native species compete among themselves. We tested the direct and indirect effects of an invasive aphid herbivore on a native aphid species and two host milkweed species. The invasive aphid caused a 10-fold decrease in native aphid populations, and a 30% increase in plant mortality (direct effects). The invasive aphid also increased the strength of interspecific competition between the two native plant hosts (indirect effects). By investigating the role that indirect effects play in shaping species interactions in native communities, our study highlights an understudied component of species invasions.

Cannibalism by damselflies increases with rising temperature.

Trophic interactions are likely to change under climate warming. These interactions can be altered directly by changing consumption rates, or indirectly by altering growth rates and size asymmetries among individuals that in turn affect feeding. Understanding these processes is particularly important for intraspecific interactions, as direct and indirect changes may exacerbate antagonistic interactions. We examined the effect of temperature on activity rate, growth and intraspecific size asymmetries, and how these temperature dependencies affected cannibalism in Lestes congener, a damselfly with marked intraspecific variation in size. Temperature increased activity rates and exacerbated differences in body size by increasing growth rates. Increased activity and changes in body size interacted to increase cannibalism at higher temperatures. We argue that our results are likely to be general to species with life-history stages that vary in their temperature dependencies, and that the effects of climate change on communities may depend on the temperature dependencies of intraspecific interactions.

Predator personality structures prey communities and trophic cascades.

Intraspecific variation is central to our understanding of evolution and population ecology, yet its consequences for community ecology are poorly understood. Animal personality - consistent individual differences in suites of behaviours - may be particularly important for trophic dynamics, where predator personality can determine activity rates and patterns of attack. We used mesocosms with aquatic food webs in which the top predator (dragonfly nymphs) varied in activity and subsequent attack rates on zooplankton, and tested the effects of predator personality. We found support for four hypotheses: (1) active predators disproportionately reduce the abundance of prey, (2) active predators select for predator-resistant prey species, (3) active predators strengthen trophic cascades (increase phytoplankton abundance) and (4) active predators are more likely to cannibalise one another, weakening all other trends when at high densities. These results suggest that intraspecific variation in predator personality is an important determinant of prey abundance, community composition and trophic cascades.