When phenology matters: age–size truncation alters population response to trophic mismatch

Climate-induced shifts in the timing of life-history events are a worldwide phenomenon, and these shifts can de-synchronize species interactions such as predator–prey relationships. In order to understand the ecological implications of altered seasonality, we need to consider how shifts in phenology interact with other agents of environmental change such as exploitation and disease spread, which commonly act to erode the demographic structure of wild populations. Using long-term observational data on the phenology and dynamics of a model predator–prey system (fish and zooplankton in Windermere, UK), we show that age–size truncation of the predator population alters the consequences of phenological mismatch for offspring survival and population abundance. Specifically, age–size truncation reduces intraspecific density regulation due to competition and cannibalism, and thereby amplifies the population sensitivity to climate-induced predator–prey asynchrony, which increases variability in predator abundance. High population variability poses major ecological and economic challenges as it can diminish sustainable harvest rates and increase the risk of population collapse. Our results stress the importance of maintaining within-population age–size diversity in order to buffer populations against phenological asynchrony, and highlight the need to consider interactive effects of environmental impacts if we are to understand and project complex ecological outcomes.     

Examining predator–prey body size,trophic level and body mass across marine and terrestrial mammals

Predator–prey relationships and trophic levels are indicators of community structure, and are important for monitoring ecosystem changes. Mammals colonized the marine environment on seven separate occasions, which resulted in differences in species'' physiology, morphology and behaviour. It is likely that these changes have had a major effect upon predator–prey relationships and trophic position; however, the effect of environment is yet to be clarified. We compiled a dataset, based on the literature, to explore the relationship between body mass, trophic level and predator–prey ratio across terrestrial (n = 51) and marine (n = 56) mammals. We did not find the expected positive relationship between trophic level and body mass, but we did find that marine carnivores sit 1.3 trophic levels higher than terrestrial carnivores. Also, marine mammals are largely carnivorous and have significantly larger predator–prey ratios compared with their terrestrial counterparts. We propose that primary productivity, and its availability, is important for mammalian trophic structure and body size. Also, energy flow and community structure in the marine environment are influenced by differences in energy efficiency and increased food web stability. Enhancing our knowledge of feeding ecology in mammals has the potential to provide insights into the structure and functioning of marine and terrestrial communities.     

Prey and predator density‐dependent interactions under different water volumes

Predation is a critical ecological process that directly and indirectly mediates population stabilities, as well as ecosystem structure and function. The strength of interactions between predators and prey may be mediated by multiple density dependences concerning numbers of predators and prey. In temporary wetland ecosystems in particular, fluctuating water volumes may alter predation rates through differing search space and prey encounter rates. Using a functional response approach, we examined the influence of predator and prey densities on interaction strengths of the temporary pond specialist copepod Lovenula raynerae preying on cladoceran prey, Daphnia pulex, under contrasting water volumes. Further, using a population dynamic modeling approach, we quantified multiple predator effects across differences in prey density and water volume. Predators exhibited type II functional responses under both water volumes, with significant antagonistic multiple predator effects (i.e., antagonisms) exhibited overall. The strengths of antagonistic interactions were, however, enhanced under reduced water volumes and at intermediate prey densities. These findings indicate important biotic and abiotic contexts that mediate predator–prey dynamics, whereby multiple predator effects are contingent on both prey density and search area characteristics. In particular, reduced search areas (i.e., water volumes) under intermediate prey densities could enhance antagonisms by heightening predator–predator interference effects.     

Crossing regimes of temperature dependence in animal movement

A pressing challenge in ecology is to understand the effects of changing global temperatures on food web structure and dynamics. The stability of these complex ecological networks largely depends on how predator–prey interactions may respond to temperature changes. Because predators and prey rely on their velocities to catch food or avoid being eaten, understanding how temperatures may affect animal movement is central to this quest. Despite our efforts, we still lack a mechanistic understanding of how the effect of temperature on metabolic processes scales up to animal movement and beyond. Here, we merge a biomechanical approach, the Metabolic Theory of Ecology and empirical data to show that animal movement displays multiple regimes of temperature dependence. We also show that crossing these regimes has important consequences for population dynamics and stability, which depend on the parameters controlling predator–prey interactions. We argue that this dependence upon interaction parameters may help explain why experimental work on the temperature dependence of interaction strengths has so far yielded conflicting results. More importantly, these changes in the temperature dependence of animal movement can have consequences that go well beyond ecological interactions and affect, for example, animal communication, mating, sensory detection, and any behavioral modality dependent on the movement of limbs. Finally, by not taking into account the changes in temperature dependence reported here we might not be able to properly forecast the impact of global warming on ecological processes and propose appropriate mitigation action when needed.     

A size-constrained feeding-niche model distinguishes predation patterns between aquatic and terrestrial food webs

Understanding the formation of feeding links provides insights into processes underlying food webs. Generally, predators feed on prey within a certain body-size range, but a systematic quantification of such feeding niches is lacking. We developed a size-constrained feeding-niche (SCFN) model and parameterized it with information on both realized and non-realized feeding links in 72 aquatic and 65 terrestrial food webs. Our analyses revealed profound differences in feeding niches between aquatic and terrestrial predators and variation along a temperature gradient. Specifically, the predator–prey body-size ratio and the range in prey sizes increase with the size of aquatic predators, whereas they are nearly constant across gradients in terrestrial predator size. Overall, our SCFN model well reproduces the feeding relationships and predation architecture across 137 natural food webs (including 3878 species and 136,839 realized links). Our results illuminate the organisation of natural food webs and enables novel trait-based and environment-explicit modelling approaches.     

Rapid prey evolution and the dynamics of two-predator food webs

Traits affecting ecological interactions can evolve on the same time scale as population and community dynamics, creating the potential for feedbacks between evolutionary and ecological dynamics. Theory and experiments have shown in particular that rapid evolution of traits conferring defense against predation can radically change the qualitative dynamics of a predator–prey food chain. Here, we ask whether such dramatic effects are likely to be seen in more complex food webs having two predators rather than one, or whether the greater complexity of the ecological interactions will mask any potential impacts of rapid evolution. If one prey genotype can be well-defended against both predators, the dynamics are like those of a predator–prey food chain. But if defense traits are predator-specific and incompatible, so that each genotype is vulnerable to attack by at least one predator, then rapid evolution produces distinctive behaviors at the population level: population typically oscillate in ways very different from either the food chain or a two-predator food web without rapid prey evolution. When many prey genotypes coexist, chaotic dynamics become likely. The effects of rapid evolution can still be detected by analyzing relationships between prey abundance and predator population growth rates using methods from functional data analysis.     

Size‐based interactions across trophic levels in food webs of shallow Mediterranean lakes

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Tadpole–odonate larvae interactions: influence of body size and diel rhythm

Several studies have shown that prey and predator body size may affect the outcome of predator–prey interactions. However, few studies have taken in account the changes on predator–prey interactions over 24 h. In a tropical freshwater system I evaluated how predator and prey size, and their diel rhythm in activity influenced the interaction between Physalaemus pustulosus tadpoles and dragonfly larvae. Tadpoles of different size classes were exposed to two size classes of the dragonfly larvae Rhionaeschna spec. Feeding trials were conducted during day and night. Tadpole activity showed a diel rhythm and affected size-selective predation of the smallest dragonfly larvae, but not of the larger ones. Predator and prey size had a significant effect on the prey survivorship and prey size had a significant effect on the preference of the predator. The interaction between both factors was significant, indicating that they did not operate independently. I conclude that the predator–prey interactions between odonate larvae and anuran tadpoles were mainly affected by the size of the prey and the predator, and less by the diel activity pattern of the prey.     

Molecular gut content analysis indicates the inter‐ and intra‐guild predation patterns of spiders in conventionally managed vegetable fields

Inter‐ and intra‐guild interactions are important in the coexistence of predators and their prey, especially in highly disturbed vegetable cropping systems with sporadic food resources. Assessing the dietary range of a predator taxon characterized by diverse foraging behavior using conventional approaches, such as visual observation and conventional molecular approaches for prey detection, has serious logistical problems. In this study, we assessed the prey compositions and compare the dietary spectrum of a functionally diverge group of predators—spiders—to characterize their trophic interactions and assess biological control potential in Brassica vegetable fields. We used high‐throughput sequencing (HTS) and biotic interaction networks to precisely annotate the predation spectrum and highlight the predator–predator and predator–prey interactions. The prey taxa in the gut of all spider families were mainly enriched with insects (including dipterans, coleopterans, orthopterans, hemipterans, and lepidopterans) with lower proportions of arachnids (such as Araneae) along with a wide range of other prey factions. Despite the generalist foraging behavior of spiders, the community structure analysis and interaction networks highlighted the overrepresentation of particular prey taxa in the gut of each spider family, as well as showing the extent of interfamily predation by spiders. Identifying the diverse trophic niche proportions underpins the importance of spiders as predators of pests in highly disturbed agroecosystems. More specifically, combining HTS with advanced ecological community analysis reveals the preferences and biological control potential of particular spider taxa (such as Salticidae against lepidopterans and Pisauridae against dipterans), and so provides a valuable evidence base for targeted conservation biological control efforts in complex trophic networks.     

Influence of intra‐ and interspecific variation in predator–prey body size ratios on trophic interaction strengths

1. Predation is a pervasive force that structures food webs and directly influences ecosystem functioning. The relative body sizes of predators and prey may be an important determinant of interaction strengths. However, studies quantifying the combined influence of intra‐ and interspecific variation in predator–prey body size ratios are lacking.
2. We use a comparative functional response approach to examine interaction strengths between three size classes of invasive bluegill and largemouth bass toward three scaled size classes of their tilapia prey. We then quantify the influence of intra‐ and interspecific predator–prey body mass ratios on the scaling of attack rates and handling times.
3. Type II functional responses were displayed by both predators across all predator and prey size classes. Largemouth bass consumed more than bluegill at small and intermediate predator size classes, while large predators of both species were more similar. Small prey were most vulnerable overall; however, differential attack rates among prey were emergent across predator sizes. For both bluegill and largemouth bass, small predators exhibited higher attack rates toward small and intermediate prey sizes, while larger predators exhibited greater attack rates toward large prey. Conversely, handling times increased with prey size, with small bluegill exhibiting particularly low feeding rates toward medium–large prey types. Attack rates for both predators peaked unimodally at intermediate predator–prey body mass ratios, while handling times generally shortened across increasing body mass ratios.
4. We thus demonstrate effects of body size ratios on predator–prey interaction strengths between key fish species, with attack rates and handling times dependent on the relative sizes of predator–prey participants.
5. Considerations for intra‐ and interspecific body size ratio effects are critical for predicting the strengths of interactions within ecosystems and may drive differential ecological impacts among invasive species as size ratios shift.

     

Trophic cascades alter eco-evolutionary dynamics and body size evolution

Trait evolution in predator–prey systems can feed back to the dynamics of interacting species as well as cascade to impact the dynamics of indirectly linked species (eco-evolutionary trophic cascades; EETCs). A key mediator of trophic cascades is body mass, as it both strongly influences and evolves in response to predator–prey interactions. Here, we use Gillespie eco-evolutionary models to explore EETCs resulting from top predator loss and mediated by body mass evolution. Our four-trophic-level food chain model uses allometric scaling to link body mass to different functions (ecological pleiotropy) and is realistically parameterized from the FORAGE database to mimic the parameter space of a typical freshwater system. To track real-time changes in selective pressures, we also calculated fitness gradients for each trophic level. As predicted, top predator loss generated alternating shifts in abundance across trophic levels, and, depending on the nature and strength in changes to fitness gradients, also altered trajectories of body mass evolution. Although more distantly linked, changes in the abundance of top predators still affected the eco-evolutionary dynamics of the basal producers, in part because of their relatively short generation times. Overall, our results suggest that impacts on top predators can set off transient EETCs with the potential for widespread indirect impacts on food webs.     

Predator size and phenology shape prey survival in temporary ponds

Theoretical efforts suggest that the relative sizes of predators and their prey can shape community dynamics, the structure of food webs, and the evolution of life histories. However, much of this work has assumed static predator and prey body sizes. The timing of recruitment and the growth patterns of both predator and prey have the potential to modify the strength of predator–prey interactions. In this study, I examined how predator size dynamics in 40 temporary ponds over a 3-year period affected the survival of spotted salamander (Ambystoma maculatum) larvae. Across communities, gape-limited predator richness, but not size, was correlated with habitat duration (pond permanence). Within communities, mean gape-limited predator size diminished as the growing season progressed. This size reduction occurred because prey individuals grew into a body size refuge and because the largest of the predators left ponds by mid-season. Elevated gape-limited predation risk across time and space was predicted by the occurrence of two large predatory salamanders: marbled salamander larvae (Ambystoma opacum) and red-spotted newt adults (Notophthalmus viridescens). The presence of the largest gape-limited predator, A. opacum, predicted A. maculatum larval survival in the field. The distribution of large predatory salamanders among ponds and across time is expected to lead to differing community dynamics and to generate divergent natural selection on early growth and body size in A. maculatum. In general, a dynamic perspective on predator size often will be necessary to understand the ecology and evolution of species interactions. This will be especially true in frequently disturbed or seasonal habitats where phenology and ontogeny interact to determine body size asymmetries. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Seasonal shifts in predator body size diversity and trophic interactions in size-structured predator-prey systems

1.?Theory suggests that the relationship between predator diversity and prey suppression should depend on variation in predator traits such as body size, which strongly influences the type and strength of species interactions. Prey species often face a range of different sized predators, and the composition of body sizes of predators can vary between communities and within communities across seasons. 2.?Here, I test how variation in size structure of predator communities influences prey survival using seasonal changes in the size structure of a cannibalistic population as a model system. Laboratory and field experiments showed that although the per-capita consumption rates increased at higher predator-prey size ratios, mortality rates did not consistently increase with average size of cannibalistic predators. Instead, prey mortality peaked at the highest level of predator body size diversity. 3.?Furthermore, observed prey mortality was significantly higher than predictions from the null model that assumed no indirect interactions between predator size classes, indicating that different sized predators were not substitutable but had more than additive effects. Higher predator body size diversity therefore increased prey mortality, despite the increased potential for behavioural interference and predation among predators demonstrated in additional laboratory experiments. 4.?Thus, seasonal changes in the distribution of predator body sizes altered the strength of prey suppression not only through changes in mean predator size but also through changes in the size distribution of predators. In general, this indicates that variation (i.e. diversity) within a single trait, body size, can influence the strength of trophic interactions and emphasizes the importance of seasonal shifts in size structure of natural food webs for community dynamics.     

Microgeographic divergence of functional responses among salamanders under antagonistic selection from apex predators

A predator''s functional response determines predator–prey interactions by describing the relationship between the number of prey available and the number eaten. Its shape and parameters fundamentally govern the dynamic equilibrium of predator–prey interactions and their joint abundances. Yet, estimates of these key parameters generally assume stasis in space and time and ignore the potential for local adaptation to alter feeding responses and the stability of trophic dynamics. Here, we evaluate if functional responses diverge among populations of spotted salamander (Ambystoma maculatum) larvae that face antagonistic selection on feeding strategies based on their own risk of predation. Common garden experiments revealed that spotted salamander from ponds with varying predation risks differed in their functional responses, suggesting an evolutionary response. Applying mechanistic equations, we discovered that the combined changes in attack rates, handling times and shape of the functional response enhanced feeding rate in environments with high densities of gape-limited predators. We suggest how these parameter changes could alter community equilibria and other emergent properties of food webs. Community ecologists might often need to consider how local evolution at fine scales alters key relationships in ways that alter local diversity patterns, food web dynamics, resource gradients and community responses to disturbance.     

Parasites as prey in aquatic food webs: implications for predator infection and parasite transmission

While the recent inclusion of parasites into food‐web studies has highlighted the role of parasites as consumers, there is accumulating evidence that parasites can also serve as prey for predators. Here we investigated empirical patterns of predation on parasites and their relationships with parasite transmission in eight topological food webs representing marine and freshwater ecosystems. Within each food web, we examined links in the typical predator–prey sub web as well as the predator–parasite sub web, i.e. the quadrant of the food web indicating which predators eat parasites. Most predator– parasite links represented ‘concomitant predation’ (consumption and death of a parasite along with the prey/host; 58–72%), followed by ‘trophic transmission’ (predator feeds on infected prey and becomes infected; 8–32%) and predation on free‐living parasite life‐cycle stages (4–30%). Parasite life‐cycle stages had, on average, between 4.2 and 14.2 predators. Among the food webs, as predator richness increased, the number of links exploited by trophically transmitted parasites increased at about the same rate as did the number of links where these stages serve as prey. On the whole, our analyses suggest that predation on parasites has important consequences for both predators and parasites, and food web structure. Because our analysis is solely based on topological webs, determining the strength of these interactions is a promising avenue for future research.     

Increases in disturbance and reductions in habitat size interact to suppress predator body size

Food webs are strongly size‐structured so will be vulnerable to changes in environmental factors that affect large predators. However, mechanistic understanding of environmental controls of top predator size is poorly developed. We used streams to investigate how predator body size is altered by three fundamental climate change stressors: reductions in habitat size, increases in disturbance and warmer temperatures. Using new survey data from 74 streams, we showed that habitat size and disturbance were the most important stressors influencing predator body size. A synergistic interaction between that habitat size and disturbance due to flooding meant the sizes of predatory fishes peaked in large, benign habitats and their body size decreased as habitats became either smaller or harsher. These patterns were supported by experiments indicating that habitat‐size reductions and increased flood disturbance decreased both the abundance and biomass of large predators. This research indicates that interacting climate change stressors can influence predator body size, resulting in smaller predators than would be predicted from examining an environmental factor in isolation. Thus, climate‐induced changes to key interacting environmental factors are likely to have synergistic impacts on predator body size which, because of their influence on the strength of biological interactions, will have far‐reaching effects on food‐web responses to global environmental change.     

Predator diversity and environmental change modify the strengths of trophic and nontrophic interactions

Understanding the dependence of species interaction strengths on environmental factors and species diversity is crucial to predict community dynamics and persistence in a rapidly changing world. Nontrophic (e.g. predator interference) and trophic components together determine species interaction strengths, but the effects of environmental factors on these two components remain largely unknown. This impedes our ability to fully understand the links between environmental drivers and species interactions. Here, we used a dynamical modelling framework based on measured predator functional responses to investigate the effects of predator diversity, prey density, and temperature on trophic and nontrophic interaction strengths within a freshwater food web. We found that (i) species interaction strengths cannot be predicted from trophic interactions alone, (ii) nontrophic interaction strengths vary strongly among predator assemblages, (iii) temperature has opposite effects on trophic and nontrophic interaction strengths, and (iv) trophic interaction strengths decrease with prey density, whereas the dependence of nontrophic interaction strengths on prey density is concave up. Interestingly, the qualitative impacts of temperature and prey density on the strengths of trophic and nontrophic interactions were independent of predator identity, suggesting a general pattern. Our results indicate that taking multiple environmental factors and the nonlinearity of density‐dependent species interactions into account is an important step towards a better understanding of the effects of environmental variations on complex ecological communities. The functional response approach used in this study opens new avenues for (i) the quantification of the relative importance of the trophic and nontrophic components in species interactions and (ii) a better understanding how environmental factors affect these interactions and the dynamics of ecological communities.     

A spatial theory for characterizing predator–multiprey interactions in heterogeneous landscapes

Trophic interactions in multiprey systems can be largely determined by prey distributions. Yet, classic predator–prey models assume spatially homogeneous interactions between predators and prey. We developed a spatially informed theory that predicts how habitat heterogeneity alters the landscape-scale distribution of mortality risk of prey from predation, and hence the nature of predator interactions in multiprey systems. The theoretical model is a spatially explicit, multiprey functional response in which species-specific advection–diffusion models account for the response of individual prey to habitat edges. The model demonstrates that distinct responses of alternative prey species can alter the consequences of conspecific aggregation, from increasing safety to increasing predation risk. Observations of threatened boreal caribou, moose and grey wolf interacting over 378 181 km² of human-managed boreal forest support this principle. This empirically supported theory demonstrates how distinct responses of apparent competitors to landscape heterogeneity, including to human disturbances, can reverse density dependence in fitness correlates.     

Temperature,predator–prey interaction strength and population stability

Warming could strongly stabilize or destabilize populations and food webs by changing the interaction strengths between predators and their prey. Predicting the consequences of warming requires understanding how temperature affects ingestion (energy gain) and metabolism (energy loss). Here, we studied the temperature dependence of metabolism and ingestion in laboratory experiments with terrestrial arthropods (beetles and spiders). From this data, we calculated ingestion efficiencies (ingestion/metabolism) and per capita interaction strengths in the short and long term. Additionally, we investigated if and how body mass changes these temperature dependencies. For both predator groups, warming increased metabolic rates substantially, whereas temperature effects on ingestion rates were weak. Accordingly, the ingestion efficiency (the ratio of ingestion to metabolism) decreased in all treatments. This result has two possible consequences: on the one hand, it suggests that warming of natural ecosystems could increase intrinsic population stability, meaning less fluctuations in population density; on the other hand, decreasing ingestion efficiencies may also lead to higher extinction risks because of starvation. Additionally, predicted long‐term per capita interaction strengths decreased with warming, which suggests an increase in perturbation stability of populations, i.e., a higher probability of returning to the same equilibrium density after a small perturbation. Together, these results suggest that warming has complex and potentially profound effects on predator–prey interactions and food‐web stability.     

Habitat context influences predator interference interactions and the strength of resource partitioning

Despite increasing evidence that habitat structure can shape predator–prey interactions, few studies have examined the impact of habitat context on interactions among multiple predators and the consequences for combined foraging rates. We investigated the individual and combined effects of stone crabs (Menippe mercenaria) and knobbed whelks (Busycon carica) when foraging on two common bivalves, the hard clam (Mercenaria mercenaria) and the ribbed mussel (Geukensia demissa) in oyster reef and sand flat habitats. Because these species co-occur across these and other estuarine habitats of varying physical complexity, this system is ideal for examining how habitat context influences foraging rates and the generality of predator interactions. Consistent with results from previous studies, consumption rates of each predator in isolation from the other were higher in the sand flat than in the more structurally complex oyster reef habitat. However, consumption by the two predators when combined surprisingly did not differ between the two habitats. This counterintuitive result probably stems from the influence of habitat structure on predator–predator interactions. In the sand-flat habitat, whelks significantly reduced their consumption of their less preferred prey when crabs were present. However, the structurally more complex oyster reef habitat appeared to reduce interference interactions among predators, such that consumption rates when the predators co-occurred did not differ from predation rates when alone. In addition, both habitat context and predator–predator interactions increased resource partitioning by strengthening predator dietary selectivity. Thus, an understanding of how habitat characteristics such as physical complexity influence interactions among predators may be critical to predicting the effects of modifying predator populations on their shared prey.