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Some predators depend heavily on one or a few prey types, and others have exceptionally broad diets. It is unclear how this diet variation arises. Here, we demonstrate a strong link between diet species richness and Shannon entropy of prey frequencies (a diet diversity spectrum) for a globally distributed group of apex predators-raptors. For many raptors, diet entropy is consistent with random sampling expectations given a lognormal distribution of abundances among prey species. Yet most species-rich diets often approach the maximum possible diet entropy, indicating an unexpected level of diet evenness that is not predicted by theory. Positioning along this diet diversity spectrum is linked to evolutionary history, the types of prey that are acceptable and the role of raptors as food web integrators through cross-habitat sampling. These results suggest that raptors may have a highly stabilizing effect on terrestrial food webs and play an important role in maintaining biodiversity.

To understand how global warming will impact biodiversity, we need to pay attention to those species with higher vulnerability. However, to assess vulnerability, we also need to consider the thermoregulatory mechanisms, body size, and thermal tolerance of species. Studies addressing thermal tolerance on small ectotherms have mostly focused on insects, while other arthropods, such as arachnids remain understudied. Here, we quantified the physiological thermal sensitivity of the pseudoscorpion Dactylochelifer silvestris using a respirometry setup with a ramping temperature increase. Overall, we found that D. silvestris has a much lower metabolic rate than other organisms of similar size. As expected, metabolic rate increased with body size, with adults having larger metabolic rates, but the overall metabolic scaling exponent was low. Both the temperature at which metabolism peaked and the critical thermal maxima were high (>44°C) and comparable to those of other arachnids. The activation energy, which characterizes the rising portion of the thermal sensitivity curve, was 0.66 eV, consistent with predictions for insects and other taxa in general. Heat tolerances and activation energy did not differ across life stages. We conclude that D. silvestris has low metabolic rates and a high thermal tolerance, which would likely influence how all stages and sexes of this species could endure climate change., (© The Author(s) 2024. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology.)

Understanding and predicting population responses to climate change is a crucial challenge. A key component of population responses to climate change are cases in which focal biological rates (e.g., population growth rates) change in response to climate change due to non-compensatory effects of changes in the underlying components (e.g., birth and death rates) determining the focal rates. We refer to these responses as non-compensatory climate change effects. As differential responses of biological rates to climate change have been documented in a variety of systems and arise at multiple levels of organization within and across species, non-compensatory effects may be nearly ubiquitous. Yet, how non-compensatory climate change responses combine and scale to influence the demographics of populations is often unclear and requires mapping them to the birth and death rates underlying population change. We provide a flexible framework for incorporating non-compensatory changes in upstream rates within and among species and mapping their consequences for additional downstream rates across scales to their eventual effects on population growth rates. Throughout, we provide specific examples and potential applications of the framework. We hope this framework helps to enhance our understanding of and unify research on population responses to climate change., (© 2024 The Author(s). Global Change Biology published by John Wiley & Sons Ltd.)

Functional responses describe foraging rates across prey densities and underlie many fundamental ecological processes. Most functional response knowledge comes from simplified lab experiments, but we do not know whether these experiments accurately represent foraging in nature. In addition, the difficulty of conducting multispecies functional response experiments means that it is unclear whether interaction strengths are weakened in the presence of multiple prey types. We developed a novel method to estimate wild predators' foraging rates from metabarcoding data and use this method to present functional responses for wild wolf spiders foraging on 27 prey families. These field functional responses were considerably reduced compared to lab functional responses. We further find that foraging is sometimes increased in the presence of other prey types, contrary to expectations. Our novel method for estimating field foraging rates will allow researchers to determine functional responses for wild predators and address long-standing questions about foraging in nature., (© 2024 The Authors. Ecology Letters published by John Wiley & Sons Ltd.)

Chemotaxis is widespread across many taxa and often aids resource acquisition or predator avoidance. Species interactions can modify the degree of movement facilitated by chemotaxis. In this study, we investigated the influence of symbionts on Paramecium bursaria's chemotactic behavior toward chloroviruses. To achieve this, we performed choice experiments using chlorovirus and control candidate attractors (virus stabilization buffer and pond water). We quantified the movement of Paramecia grown with or without algal and viral symbionts toward each attractor. All Paramecia showed some chemotaxis toward viruses, but cells without algae and viruses showed the most movement toward viruses. Thus, the endosymbiotic algae (zoochlorellae) appeared to alter the movement of Paramecia toward chloroviruses, but it was not clear that ectosymbiotic viruses (chlorovirus) also had this effect. The change in behavior was consistent with a change in swimming speed, but a change in attraction remains possible. The potential costs and benefits of chemotactic movement toward chloroviruses for either the Paramecia hosts or its symbionts remain unclear., (© 2023. The Author(s), under exclusive licence to Springer Science+Business Media, LLC, part of Springer Nature.)

The effects of warming on ecological communities emerge from a range of potentially asymmetric impacts on individual physiology and development. Understanding these responses, however, is limited by our ability to connect mechanisms or emergent patterns across the many processes that drive variation in demography. Further complicating this understanding is the gain or loss of predators to many communities, which may interact with changes in temperature to drive community change. Here we conducted a factorial warming and predation experiment to test generalized predictions about responses to warming. We used microcosms with a range of protists, rotifers, and a gastrotrich, with and without the predator Actinosphaerium , to assess changes in diversity, body size, function, and composition in response to warming. We find that community respiration and predator:prey biovolume ratios peak at intermediate temperatures, while species richness declined with temperature. We also found that overall biomass increased with species richness, driven by the effect of temperature on richness. There was little evidence of an interaction between predation and temperature change, likely because the predator was mostly limited to the intermediate temperatures. Overall, our results suggest that general predictions about community change are still challenging to make but may benefit by considering multiple dimensions of community patterns in an integrated way., (© 2023 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

Viruses can have large effects on the ecological communities in which they occur. Much of this impact comes from the mortality of host cells, which simultaneously alters microbial community composition and causes the release of matter that can be used by other organisms. However, recent studies indicate that viruses may be even more deeply integrated into the functioning of ecological communities than their effect on nutrient cycling suggests. In particular, chloroviruses, which infect chlorella-like green algae that typically occur as endosymbionts, participate in three types of interactions with other species. Chlororviruses (i) can lure ciliates from a distance, using them as a vector; (ii) depend on predators for access to their hosts; and (iii) get consumed as a food source by, at least, a variety of protists. Therefore, chloroviruses both depend on and influence the spatial structures of communities as well as the flows of energy through those communities, driven by predator-prey interactions. The emergence of these interactions are an eco-evolutionary puzzle, given the interdependence of these species and the many costs and benefits that these interactions generate., Competing Interests: The authors declare no conflict of interest.

Niche differentiation and intraguild predation (IGP) can allow ecologically similar species to coexist, although it is unclear which coexistence mechanism predominates in consumer communities. Until now, a limited ability to quantify diets from metabarcoding data has precluded the use of sequencing data to determine the relative importance of these mechanisms. Here, we pair a recent metabarcoding quantification approach with stable isotope analysis to examine diet composition in a wolf spider community. We compare the prevalence of resource partitioning and IGP in these spiders and test whether factors that influence foraging performance, including individual identity, morphology, prey community and environmental conditions, can explain variation in diet composition and IGP. Extensive IGP is likely the primary coexistence mechanism in this community, and other factors to which foraging variation is often attributed do not explain diet composition and IGP here. Rather, IGP increases as prey diversity decreases. Foragers are driven to IGP where resource niches are limited. We highlight the need to examine how drivers of predator-prey interaction strengths translate into foraging in natural systems., (© 2023 The Authors. Journal of Animal Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society.)

Viruses impact host cells and have indirect effects on ecosystem processes. Plankton such as ciliates can reduce the abundance of virions in water, but whether virus consumption translates into demographic consequences for the grazers is unknown. Here, we show that small protists not only can consume viruses they also can grow and divide given only viruses to eat. Moreover, the ciliate Halteria sp. foraging on chloroviruses displays dynamics and interaction parameters that are similar to other microbial trophic interactions. These results suggest that the effect of viruses on ecosystems extends beyond (and in contrast to) the viral shunt by redirecting energy up food chains.

Stochastic processes such as genetic drift may hinder adaptation, but the effect of such stochasticity on evolution via its effect on ecological dynamics is poorly understood. Here we evaluate patterns of adaptation in a population subject to variation in demographic stochasticity. We show that stochasticity can alter population dynamics and lead to evolutionary outcomes that are not predicted by classic eco-evolutionary modeling approaches. We also show, however, that these outcomes are governed by nonequilibrium evolutionary attractors-these are maxima in lifetime reproductive success when stochasticity keeps the ecological system away from the deterministic equilibrium. These NEEAs alter the path of evolution but are not visible through the equilibrium lens that underlies much evolutionary theory. Our results reveal that considering population processes during transient periods can greatly improve our understanding of the path and pace of evolution., (© 2022 The Authors. Ecology published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

Functional responses, the relationships between consumer foraging rate and resource (prey) density, provide key insights into consumer-resource interactions while also being a major driver of population dynamics and food web structure. We present a global database of 2598 standardized functional responses and parameters extracted from the published literature. We refit the functional responses with a Type II model using standardized methods and report the fitted parameters along with data on experimental conditions, consumer and resource taxonomy and type, as well as the habitat and dimensionality of the foraging interaction. The consumer and resource species covered here are taxonomically diverse, from protozoans filtering algae to wasps parasitizing moth larvae to wolves hunting moose. The FoRAGE (Functional Responses from Around the Globe in all Ecosystems) database (doi: 10.5063/DB807S) is a living data set that will be updated periodically as new functional responses are published. Data are released under a CC-By-NC-SA license, and credit should be given to this paper when referring to this specific version of the data release., (© 2022 The Authors. Ecology © 2022 The Ecological Society of America.)

Functional responses are central to predator-prey dynamics and describe how predation varies with prey abundance. Functional responses often are measured without regard to prey size (i.e., body mass) or the temperature dependence of feeding rates. However, variation in prey size within populations is ubiquitous, and predation rates are often both size and temperature-dependent. Here, we assessed functional responses of larvae and adult Harmonia axyridis on the 1st, 2nd, and 3rd instars of the prey Spodoptera litura across a range of temperatures (i.e., 15, 20, 25, 30, and 35°C). The type and parameters of the functional responses were determined using logistic regression and fitted to the Roger's random predator equation. The magnitude of predation varied with the predator and prey stage, but prey predation increased with warming and predator age. Predation by the female and 4th instar of H. axyridis on the 1st instar of prey was greater, followed by the 2nd and 3rd instar of prey S. litura . No predation occurred on the larger prey for the 1st, 2nd, and 3rd instars of H. axyridis . The larvae and adult H. axyridis produced a type II (hyperbolic) functional response curve across all temperatures and the three prey types they consumed. Space clearance rates, handling time, and maximum predation rates of H. axyridis changed with temperature and prey size, increasing with temperature and decreasing with prey size, suggesting more predation will occur on younger prey. This study indicates an interactive role of temperature and prey/predator size in shaping functional responses, which might complicate the planning of effective biocontrol strategies against this serious pest., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2022 Islam, Shah, Güncan, DeLong and Zhou.)

Predator functional responses describe predator feeding rates and are central to predator-prey theory. Originally defined as the relationship between predator feeding rates and prey densities, it is now well known that functional responses are shaped by a multitude of factors. However, much of our knowledge about how these factors influence functional responses is based on laboratory studies that are generally logistically constrained to examining only a few factors simultaneously and that have unclear links to the conditions organisms experience in the field. We apply an observational approach for measuring functional responses to understand how sex/stage differences, temperature and predator densities interact to influence the functional response of zebra jumping spiders on midges under natural conditions. We used field surveys of jumping spiders to infer their feeding rates and examine the relationships between feeding rates, sex/stage, midge density, predator density and temperature using generalized additive models. We then used the relationships supported by the models to fit parametric functional responses to the data. We find that feeding rates of zebra jumping spiders follow some expectations from previous laboratory studies such as increasing feeding rates with body size and decreasing feeding rates with predator densities. However, in contrast to previous results, our results also show a lack of temperature response in spider feeding rates and differential decreases in the feeding rates of females and juveniles with densities of different spider sexes/stages. Our results illustrate the multidimensional nature of functional responses in natural settings and reveal how factors influencing functional responses can interact with one another through behaviour and morphology. Further studies investigating the influence of multiple mechanisms on predator functional responses under field conditions will increase our understanding of the drivers of predator-prey interaction strengths and their consequences for communities and ecosystems., (© 2022 The Authors. Journal of Animal Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society.)

Understanding how phenotypes emerge from genotypes is a foundational goal in biology. As challenging as this task is when considering cellular life, it is further complicated in the case of viruses. During replication, a virus as a discrete entity (the virion) disappears and manifests itself as a metabolic amalgam between the virus and the host (the virocell). Identifying traits that unambiguously constitute a virus's phenotype is straightforward for the virion, less so for the virocell. Here, we present a framework for categorizing virus phenotypes that encompasses both virion and virocell stages and considers functional and performance traits of viruses in the context of fitness. Such an integrated view of virus phenotype is necessary for comprehensive interpretation of viral genome sequences and will advance our understanding of viral evolution and ecology., (© 2021. Springer Nature Limited.)

AbstractA scientific understanding of the biological world arises when ideas about how nature works are formalized, tested, refined, and then tested again. Although the benefits of feedback between theoretical and empirical research are widely acknowledged by ecologists, this link is still not as strong as it could be in ecological research. This is in part because theory, particularly when expressed mathematically, can feel inaccessible to empiricists who may have little formal training in advanced math. To address this persistent barrier, we provide a general and accessible guide that covers the basic, step-by-step process of how to approach, understand, and use ecological theory in empirical work. We first give an overview of how and why mathematical theory is created, then outline four specific ways to use both mathematical and verbal theory to motivate empirical work, and finally present a practical tool kit for reading and understanding the mathematical aspects of ecological theory. We hope that empowering empiricists to embrace theory in their work will help move the field closer to a full integration of theoretical and empirical research.

Chloroviruses are large viruses that replicate in chlorella-like green algae and normally exist as mutualistic endosymbionts (referred to as zoochlorellae) in protists such as Paramecium bursaria . Chlorovirus populations rise and fall in indigenous waters through time; however, the factors involved in these virus fluctuations are still under investigation. Chloroviruses attach to the surface of P. bursaria but cannot infect their zoochlorellae hosts because the viruses cannot reach the zoochlorellae as long as they are in the symbiotic phase. Predators of P. bursaria , such as copepods and didinia, can bring chloroviruses into contact with zoochlorellae by disrupting the paramecia, which results in an increase in virus titers in microcosm experiments. Here, we report that another predator of P. bursaria , Bursaria truncatella , can also increase chlorovirus titers. After two days of foraging on P. bursaria , B. truncatella increased infectious chlorovirus abundance about 20 times above the controls. Shorter term foraging (3 h) resulted in a small increase of chlorovirus titers over the controls and more foraging generated more chloroviruses. Considering that B. truncatella does not release viable zoochlorellae either during foraging or through fecal pellets, where zoochlorellae could be infected by chlorovirus, we suggest a third pathway of predator virus catalysis. By engulfing the entire protist and digesting it slowly, virus replication can occur within the predator and some of the virus is passed out through a waste vacuole. These results provide additional support for the hypothesis that predators of P. bursaria are important drivers of chlorovirus population sizes and dynamics.

How and if organisms can adapt to changing temperatures has drastic consequences for the natural world. Thermal adaptation involves finding a match between temperatures permitting growth and the expected temperature distribution of the environment. However, if and how this match is achieved, and how tightly linked species change together, is poorly understood. Paramecium bursaria is a ciliate that has a tight physiological interaction with endosymbiotic green algae (zoochlorellae). We subjected a wild population of P. bursaria to a cold and warm climate (20 and 32℃) for ∼300 generations. We then measured the thermal performance curve (TPC) for intrinsic rate of growth (r max ) for these evolved lines across temperatures. We also evaluated number and size of the zoochlorellae populations within paramecia cells. TPCs for warm-adapted populations were shallower and broader than TPCs of cold-adapted populations, indicating that the warm populations adapted by moving along a thermal generalist/specialist trade off rather than right-shifting the TPC. Zoochlorellae populations within cold-adapted paramecia had fewer and larger zoochlorellae than hot-adapted paramecia, indicating phenotypic shifts in the endosymbiont accompany thermal adaptation in the host. Our results provide new and novel insight into how species involved in complex interactions will be affected by continuing increasing global temperatures., (© 2021 The Authors. Evolution © 2021 The Society for the Study of Evolution.)

Predator functional responses describe predator feeding rates and are central to predator-prey theory. Ecologists have measured thousands of predator functional responses using the same basic experimental method. However, this design is ill-suited to address many current questions regarding functional responses. We derive a new experimental design and statistical analysis that quantifies functional responses using the times between a predators' feeding events requiring only one or a few trials. We examine the feasibility of the experimental method and analysis using simulations to assess the ability of the statistical model to estimate functional response parameters and perform a proof-of-concept experiment estimating the functional responses of two individual jumping spiders. Our simulations show that the statistical method reliably estimates functional response parameters. Our proof-of-concept experiment illustrates that the method provides reasonable estimates of functional response parameters. By virtue of the fewer number of trials required to measure a functional response, the method derived here promises to expand the questions that can be addressed using functional response experiments and the systems in which they can be measured. Thus, we hope that our method will refine our understanding of functional responses and predator-prey interactions more generally., (© 2021 by the Ecological Society of America.)

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.

Dietary metabarcoding-the process of taxonomic identification of food species from DNA in consumer guts or faeces-has been rapidly adopted by ecologists to gain insights into biocontrol, invasive species and the structure of food webs. However, an outstanding issue with metabarcoding is the semi-quantitative nature of the data it provides: because metabarcoding is likely to produce false negatives for some prey more often than for other prey, we cannot infer relative frequencies of prey in the diet. To correct for this, we can adjust detected prey frequencies using DNA detectability half-lives unique to each predator-prey combination. Because the feeding experiments required to deduce these half-lives are time- and resource-intensive, our ability to weight the frequency of observations using their detectability has thus far been limited to systems with just a few prey. Here, we present a meta-analysis of 24 spider prey DNA half-lives and show that these half-lives are predictable given predator and prey mass, predator family, digestion temperature and DNA amplicon length. We further provide a new technique for weighting observations with half-lives, which allows not just for the ranking of prey in the diet, but reveals the proportion of the diet each prey comprises. Lastly, we apply this method to published dietary metabarcoding data to calculate half-lives and proportion of the predator's diet for 35 prey families, demonstrating that this technique can generate improved understanding of diets in real, diverse systems., (© 2020 John Wiley & Sons Ltd.)

AbstractBody mass-based links between predator and prey are fundamental to the architecture of food webs. These links determine who eats whom across trophic levels and strongly influence the population abundance, flow of energy, and stability properties of natural communities. Body mass links scale up to create predator-prey mass relationships across species, but the origin of these relationships is unclear. Here I show that predator-prey mass relationships are consistent with the idea that body mass evolves to maximize a dependable supply of resource uptake. I used a global database of ~2,100 predator-prey links and a mechanistic optimization model to correctly predict the slope of the predator-prey mass scaling relationships across species generally and for nine taxonomic subsets. The model also predicted cross-group variation in the heights of the body mass relationships, providing an integrated explanation for mass relationships and their variation across taxa. The results suggest that natural selection on body mass at the local scale is detectable in ecological organization at the macro scale.

Background: Predicting the effects of climate warming on the dynamics of ecological systems requires understanding how temperature influences birth rates, death rates and the strength of species interactions. The temperature dependance of these processes-which are the underlying mechanisms of ecological dynamics-is often thought to be exponential or unimodal, generally supported by short-term experiments. However, ecological dynamics unfold over many generations. Our goal was to empirically document shifts in predator-prey cycles over the full range of temperatures that can possibly support a predator-prey system and then to uncover the effect of temperature on the underlying mechanisms driving those changes., Methods: We measured the population dynamics of the Didinium-Paramecium predator-prey system across a wide range of temperatures to reveal systematic changes in the dynamics of the system. We then used ordinary differential equation fitting to estimate parameters of a model describing the dynamics, and used these estimates to assess the long-term temperature dependance of all the underlying mechanisms., Results: We found that predator-prey cycles shrank in state space from colder to hotter temperatures and that both cycle period and amplitude varied with temperature. Model parameters showed mostly unimodal responses to temperature, with one parameter (predator mortality) increasing monotonically with temperature and one parameter (predator conversion efficiency) invariant with temperature. Our results indicate that temperature can have profound, systematic effects on ecological dynamics, and these can arise through diverse and simultaneous changes in multiple underlying mechanisms. Predicting the effects of temperature on ecological dynamics may require additional investigation into how the underlying drivers of population dynamics respond to temperature beyond a short-term, acute response., Competing Interests: The authors declare that they have no competing interests., (© 2020 DeLong and Lyon.)

Functional responses describe how consumer foraging rates change with resource density. Despite extensive research looking at the factors underlying foraging interactions, there remains ongoing controversy about how temperature and body size control the functional response parameters space clearance (or attack) rate and handling time. Here, we investigate the effects of temperature, consumer mass, and resource mass using the largest compilation of functional responses yet assembled. This compilation contains 2,083 functional response curves covering a wide range of foragers and prey types, environmental conditions, and habitats. After accounting for experimental arena size, dimensionality of the foraging interaction, and consumer taxon, we find that both space clearance rate and handling time are optimized at intermediate temperatures (a unimodal rather than monotonic response), suggesting that the response to global climate change depends on the location of the consumer's current temperature relative to the optimum. We further confirm that functional responses are higher and steeper for large consumers and small resources, and models using consumer and resource masses separately outperformed models using consumer:resource mass ratios, suggesting that consumer and resource body mass act independently to set interaction strengths. Lastly, we show that the extent to which foraging is affected by temperature or mass depends on the taxonomic identity of the consumer and the dimensionality of the consumer-resource interaction. We thus argue that although overall body size and temperature effects can be identified, they are not universal, and therefore food web and community modeling approaches could be improved by considering taxonomic identity along with body size and unimodal temperature effects., (© 2020 by the Ecological Society of America.)

The effects of climate change-such as increased temperature variability and novel predators-rarely happen in isolation, but it is unclear how organisms cope with multiple stressors simultaneously. To explore this, we grew replicate Paramecium caudatum populations in either constant or variable temperatures and exposed half to predation. We then fit thermal performance curves (TPCs) of intrinsic growth rate ( r max ) for each replicate population ( N  = 12) across seven temperatures (10°C-38°C). TPCs of P. caudatum exposed to both temperature variability and predation responded only to one or the other (but not both), resulting in unpredictable outcomes. These changes in TPCs were accompanied by changes in cell morphology. Although cell volume was conserved across treatments, cells became narrower in response to temperature variability and rounder in response to predation. Our findings suggest that predation and temperature variability produce conflicting pressures on both thermal performance and cell morphology. Lastly, we found a strong correlation between changes in cell morphology and TPC parameters in response to predation, suggesting that responses to opposing selective pressures could be constrained by trade-offs. Our results shed new light on how environmental and ecological pressures interact to elicit changes in characteristics at both the individual and population levels. We further suggest that morphological responses to interactive environmental forces may modulate population-level responses, making prediction of long-term responses to environmental change challenging., (© 2020 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

Predicting how organisms respond to climate change requires that we understand the temperature dependence of fitness in relevant ecological contexts (e.g., with or without predation risk). Predation risk often induces changes to life history traits that are themselves temperature dependent. We explore how perceived predation risk and temperature interact to determine fitness (indicated by the intrinsic rate of increase, r) through changes to its underlying components (net reproductive rate, generation time, and survival) in Daphnia magna. We exposed Daphnia to predation cues from dragonfly naiads early, late, or throughout their ontogeny. Predation risk increased r differentially across temperatures and depending on the timing of exposure to predation cues. The timing of predation risk likewise altered the temperature-dependent response of T and R 0 . Daphnia at hotter temperatures responded to predation risk by increasing r through a combination of increased R 0 and decreased T that together countered an increase in mortality rate. However, only D. magna that experienced predation cues early in ontogeny showed elevated r at colder temperatures. These results highlight the fact that phenotypically plastic responses of life history traits to predation risk can be strongly temperature dependent.

Sexual size dimorphism (SSD) often results in dramatic differences in body size between females and males. Despite its ecological importance, little is known about the relationship between developmental, physiological, and energetic mechanisms underlying SSD. We take an integrative approach to understand the relationship between developmental trajectories, metabolism, and environmental conditions resulting in extreme female-biased SSD in the crab spider Mecaphesa celer (Thomisidae). We tested for sexual differences in growth trajectories, as well as in the energetics of growth, hypothesizing that female M. celer have lower metabolic rates than males or higher energy assimilation. We also hypothesized that the environment in which spiderlings develop influences the degree of SSD of a population. We tracked growth and resting metabolic rates of female and male spiderlings throughout their ontogeny and quantified the adult size of individuals raised in a combination of two diet and two temperature treatments. We show that M. celer's SSD results from differences in the shape of female and male growth trajectories. While female and male resting metabolic rates did not differ, diet, temperature, and their interaction influenced body size through an interactive effect with sex, with females being more sensitive to the environment than males. We demonstrate that the shape of the growth curve is an important but often overlooked determinant of SSD and that females may achieve larger sizes through a combination of high food ingestion and low activity levels. Our results highlight the need for new models of SSD based on ontogeny, ecology, and behavior.

Laboratory measurements of physiological and demographic tolerances are important in understanding the impact of climate change on species diversity; however, it has been recognized that forecasts based solely on these laboratory estimates overestimate risk by omitting the capacity for species to utilize microclimatic variation via behavioral adjustments in activity patterns or habitat choice. The complex, and often context-dependent nature, of microclimate utilization has been an impediment to the advancement of general predictive models. Here, we overcome this impediment and estimate the potential impact of warming on the fitness of ectotherms using a benefit/cost trade-off derived from the simple and broadly documented thermal performance curve and a generalized cost function. Our framework reveals that, for certain environments, the cost of behavioral thermoregulation can be reduced as warming occurs, enabling behavioral buffering (e.g., the capacity for behavior to ameliorate detrimental impacts) and "behavioral rescue" from extinction in extreme cases. By applying our framework to operative temperature and physiological data collected at an extremely fine spatial scale in an African lizard, we show that new behavioral opportunities may emerge. Finally, we explore large-scale geographic differences in the impact of behavior on climate-impact projections using a global dataset of 38 insect species. These multiple lines of inference indicate that understanding the existing relationship between thermal characteristics (e.g., spatial configuration, spatial heterogeneity, and modal temperature) is essential for improving estimates of extinction risk., (© 2019 John Wiley & Sons Ltd.)

Species-area relationships (SAR) and biodiversity-ecosystem function (BEF) relationships are central patterns in community ecology. Although research on both patterns often invokes mechanisms of community assembly, both SARs and BEFs are generally treated as separate phenomena. Here we link the two by creating an experimental SAR in microcosm communities and show that greater species richness in larger areas is accompanied by greater ecosystem function. We then explore mechanisms of community assembly by determining whether rare, large, or high-biomass species are more likely to persist in the larger microcosms. Our results indicate that larger areas harbor more rare species of a wider range of body sizes and have higher functional diversity, implying that the addition of niche space that supports rare species underlies the effect of area on species richness and function. Our results suggest that the preservation of large areas is a potentially useful way of maximizing the provisioning of ecosystem services through the maintenance of biodiversity.

Chloroviruses exist in aquatic systems around the planet and they infect certain eukaryotic green algae that are mutualistic endosymbionts in a variety of protists and metazoans. Natural chlorovirus populations are seasonally dynamic, but the precise temporal changes in these populations and the mechanisms that underlie them have heretofore been unclear. We recently reported the novel concept that predator/prey-mediated virus activation regulates chlorovirus population dynamics, and in the current study, we demonstrate virus-packaged chemotactic modulation of prey behavior. IMPORTANCE Viruses have not previously been reported to act as chemotactic/chemoattractive agents. Rather, viruses as extracellular entities are generally viewed as non-metabolically active spore-like agents that await further infection events upon collision with appropriate host cells. That a virus might actively contribute to its fate via chemotaxis and change the behavior of an organism independent of infection is unprecedented., (Copyright © 2019 American Society for Microbiology.)

Invading predators can negatively affect naïve prey populations due to a lack of evolved defenses. Many species therefore may be at risk of extinction due to overexploitation by exotic predators. Yet the strong selective effect of predation might drive evolution of imperiled prey toward more resistant forms, potentially allowing the prey to persist. We evaluated the potential for evolutionary rescue in an imperiled prey using Gillespie eco-evolutionary models (GEMs). We focused on a system parameterized for protists where changes in prey body size may influence intrinsic rate of population growth, space clearance rate (initial slope of the functional response), and the energetic benefit to predators. Our results show that the likelihood of rescue depends on (a) whether multiple parameters connected to the same evolving trait (i.e., ecological pleiotropy) combine to magnify selection, (b) whether the evolving trait causes negative indirect effects on the predator population by altering the energy gain per prey, (c) whether heritable trait variation is sufficient to foster rapid evolution, and (d) whether prey abundances are stable enough to avoid very rapid extinction. We also show that when evolution fosters rescue by increasing the prey equilibrium abundance, invasive predator populations also can be rescued, potentially leading to additional negative effects on other species. Thus, ecological pleiotropy, indirect effects, and system dynamics may be important factors influencing the potential for evolutionary rescue for both imperiled prey and invading predators. These results suggest that bolstering trait variation may be key to fostering evolutionary rescue, but also that the myriad direct and indirect effects of trait change could either make rescue outcomes unpredictable or, if they occur, cause rescue to have side effects such as bolstering the populations of invasive species.

Although life histories are shaped by temperature and predation, their joint influence on the interdependence of life-history traits is poorly understood. Shifts in one life-history trait often necessitate shifts in another-structured in some cases by trade-offs-leading to differing life-history strategies among environments. The offspring size-number trade-off connects three traits whereby a constant reproductive allocation ( R ) constrains how the number ( O ) and size ( S ) of offspring change. Increasing temperature and size-independent predation decrease size at and time to reproduction which can lower R through reduced time for resource accrual or size-constrained fecundity. We investigated how O, S, and R in a clonal population of Daphnia magna change across their first three clutches with temperature and size-independent predation risk. Early in ontogeny, increased temperature moved O and S along a trade-off curve (constant R ) toward fewer larger offspring. Later in ontogeny, increased temperature reduced R in the no-predator treatment through disproportionate decreases in O relative to S . In the predation treatment, R likewise decreased at warmer temperatures but to a lesser degree and more readily traded off S for O whereby the third clutch showed a constant allocation strategy of O versus S with decreasing R . Ontogenetic shifts in S and O rotated in a counterclockwise fashion as temperature increased and more drastically under risk of predation. These results show that predation risk can alter the temperature dependence of traits and their interactions through trade-offs.

The sensitivity of metabolic rate to temperature constrains the climate in which ectotherms can function, yet the temperature dependence of metabolic rate may evolve in response to biotic and abiotic factors. We compiled a dataset on the temperature dependence of metabolic rate for heterotrophic ectotherms from studies that show a peak in metabolic rate at an optimal temperature (i.e. that describe the thermal performance curve for metabolic rate). We found that peak metabolic rates were lower in aquatic than terrestrial habitats and increased with body mass, latitude and the optimal temperature. In addition, the optimal temperature decreased with latitude. These results support competing hypotheses about metabolic rate adaptation, with hotter being better in the tropics but colder being better towards the poles. Moreover, our results suggest that the temperature dependence of metabolic rate is more complex than previously suggested., (© 2018 The Author(s).)

Many chloroviruses replicate in endosymbiotic zoochlorellae that are protected from infection by their symbiotic host. To reach the high virus concentrations that often occur in natural systems, a mechanism is needed to release zoochlorellae from their hosts. We demonstrate that the ciliate predator Didinium nasutum foraging on zoochlorellae-bearing Paramecium bursaria can release live zoochlorellae from the ruptured prey cell that can then be infected by chloroviruses. The catalysis process is very effective, yielding roughly 95% of the theoretical infectious virus yield as determined by sonication of P. bursaria. Chlorovirus activation is more effective with smaller Didinia, as larger Didinia typically consume entire P. bursaria cells without rupturing them, precluding the release of zoochlorellae. We also show that the timing of Chlorovirus growth is tightly linked to the predator-prey cycle between Didinium and Paramecium, with the most rapid increase in chloroviruses temporally linked to the peak foraging rate of Didinium, supporting the idea that predator-prey cycles can drive cycles of Chlorovirus abundance.

Stability contributes to the persistence of ecological communities, yet the interactions among different stabilizing forces are poorly understood. We assembled mesocosms with an algal resource and one to eight different clones of the consumer Daphnia ambigua and tracked algal and Daphnia abundances through time. We then fitted coupled ordinary differential equations (ODEs) to the consumer-resource time series. We show that variation in different components of stability (local stability and the magnitude of population fluctuations) across mesocosms arises through variation in life history traits and the functional processes represented by ODE model parameters. Local stability was enhanced by increased algal growth rate and Daphnia mortality and foraging rate. Population fluctuations were dampened by high Daphnia conversion efficiency and lower interaction strengths, low algal growth rate, high Daphnia death rate, and low Daphnia foraging. These results indicate that (1) stability in consumer-resource systems may arise through the net effect of multiple related stabilizing pathways and (2) different aspects of stability can vary independently and may respond in opposite directions to the same forces., (© 2017 by the Ecological Society of America.)

Predator-prey interactions play a crucial role in structuring food webs, and the functional response is one way to measure the strength of this interaction. Here, we examine how predator and prey body size affects the functional response of a generalist predator-damselfly nymphs-feeding on three prey types: copepods, Daphnia, and Chydorus. Our results suggest that consumption of copepods is independent of predator body size, while increased predator body size is associated with an increased space clearance rate for Daphnia and a reduced space clearance rate for Chydorus. When considered together, foraging rates on Daphnia and Chydorus (both cladocerans) are consistent with a hump-shaped functional response, with peak foraging rates occurring at an intermediate predator-prey size ratio. Thus, although most food web theory assumes allometric predator-prey links or peaked functional responses at intermediate predator-prey size ratios, our results suggest that both relationships may occur in food webs, in addition to size-independent functional responses.

Food webs (i.e., networks of species and their feeding interactions) share multiple structural features across ecosystems. The factors explaining such similarities are still debated, and the role played by most organismal traits and their intraspecific variation is unknown. Here, we assess how variation in traits controlling predator-prey interactions (e.g., body size) affects food web structure. We show that larger phenotypic variation increases connectivity among predators and their prey as well as total food intake rate. For predators able to eat only a few species (i.e., specialists), low phenotypic variation maximizes intake rates, while the opposite is true for consumers with broader diets (i.e., generalists). We also show that variation sets predator trophic level by determining interaction strengths with prey at different trophic levels. Merging these results, we make two general predictions about the structure of food webs: ( i ) trophic level should increase with predator connectivity, and ( ii ) interaction strengths should decrease with prey trophic level. We confirm these predictions empirically using a global dataset of well-resolved food webs. Our results provide understanding of the processes structuring food webs that include functional traits and their naturally occurring variation., Competing Interests: The authors declare no conflict of interest., (Published under the PNAS license.)

The Mediterranean Sea is an invasion hotspot, with non-indigenous species suspected to be a major driver behind community changes. We used size spectra, a reliable index of food web structure, to examine how the influx of Red Sea fishes into the Mediterranean Sea has impacted the indigenous species community. This is the first attempt to use changes in the size spectra to reveal the effect of biological invasions. We used data from trawl catches along Israel's shoreline spanning 20 years to estimate changes in the community size spectra of both indigenous and non-indigenous species. We found that the relative biomass of non-indigenous species increased over the 20 years, especially for small and large species, leading to a convergence with the indigenous species size spectra. Hence, the biomass of indigenous and non-indigenous species has become identical for all size classes, suggesting similar energetic constraints and sensitivities to fishing. However, over this time period the size spectrum of indigenous species has remained remarkably constant. This suggests that the wide-scale invasion of non-indigenous species into the Mediterranean may have had little impact on the community structure of indigenous species., (© 2017 The Author(s).)

Synopsis: The mean and variance of environmental temperature are changing as a consequence of human activities. Ectotherms are sensitive to these temperature changes in the short term, typically displaying a unimodal response of most biological rates to temperature (thermal performance curves; TPCs). Many organisms, however, may acclimate or evolve in response to new temperature regimes. In particular, population growth rate TPCs (r TPCs) reflect the ability to maintain positive growth under a range of temperatures, and therefore shifts in r TPCs due to acclimation are fundamental to our understanding of how ectotherms will respond to changes in climate. Here, we derive a model for r TPCs rooted in temperature dependent metabolic rate (through enzyme kinetics and activity). We then use this model to interpret the effects of acclimation to different temperatures on r TPCs of the protist Paramecium bursaria. Intermediate acclimation temperatures generally resulted in higher upper critical thermal limits, thermal optima, maximum population growth rate, and the area under the TPC. Lower critical thermal limits increased linearly with acclimation temperature, causing a decrease in thermal breadth with increased acclimation temperature. Thus, rather than showing improved performance at the acclimation temperature, P. bursaria appeared to pay a price at all temperatures for acclimating to higher temperatures. The fits of our data to our model also suggest that changes in the structure and function of metabolic enzymes may underlie the changes in the TPCs. Specifically, our results suggest that both the delta heat capacity and delta enthalpy of formation of metabolic enzymes may have increased with acclimation. Since these two factors are correlated across acclimation temperatures, our data also suggest potential trade-offs that may constrain changes in TPCs., (© The Author 2017. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology. All rights reserved. For permissions please email: journals.permissions@oup.com.)

Ecosystems are linked through the transfer of materials and energy. Studies examining material fluxes across habitat boundaries frequently quantify unidirectional flows of nutrients and energy. However, material fluxes can be multidirectional, and we lack a conceptual framework to describe how their quantity and stoichiometry influence the net transfer of individual elements between ecosystems. Here we develop a zero net transfer isocline (ZNTI) framework that integrates the relative mass and stoichiometry of fluxes into and out of an ecosystem. We then use case studies with amphibians and salmon to elucidate how life history, ontogenetic shifts in stoichiometry, and trophic interactions shape relative fluxes of nutrients between aquatic and terrestrial ecosystems. Because they increase in both size and Ca content from ova to metamorphs, amphibian life histories strongly bias them toward net Ca export into the terrestrial environment. Because amphibian biomass, C, P, and Ca ZNTIs do not overlap, there is no value of survivorship where the net flux of biomass, C, P, and Ca are simultaneously balanced between terrestrial and aquatic habitats. The degree of iteroparity and semelparity in salmon strongly affects both the magnitude of net biomass and P flux between riverine and marine environments. While the net direction of biomass flux generally remains strongly biased toward import into the riverine system, net P flux can reach net export into the marine environment because of increasing adult breeding survival leading to reduced mass and %P of what they deposit in rivers (e.g., ova vs. whole carcasses). These examples highlight how ontogenetic shifts in body size and stoichiometry result in asymmetric fluxes of elements and biomass that can lead to simultaneous net imports and exports of different elements within the same system. Furthermore, they demonstrate how changes in life-history characteristics and stage-specific survivorship can lead to changes in net elemental transport between ecosystems.

Population dynamics may carry a signature of an ecology-evolution-ecology feedback, known as eco-evolutionary dynamics, when functionally important traits change. Given current theory, the absence of a feedback from a trait with strong links to species interactions should not occur. In a previous study with the Didinium-Paramecium predator-prey system, however, rapid and large-magnitude changes in predator cell volume occurred without any noticeable effect on the population dynamics. Here I resolve this theory-data conflict by showing that ecological pleiotropy-when a trait has more than one functional effect on an ecological process-suppresses shifts in dynamics that would arise, given the links between cell volume and the species interaction. Whether eco-evolutionary dynamics arise, therefore, depends not just on the ecology-evolution feedback but on the net effect that a trait has on different parts of the underlying interaction.

A mechanistic understanding of the response of metabolic rate to temperature is essential for understanding thermal ecology and metabolic adaptation. Although the Arrhenius equation has been used to describe the effects of temperature on reaction rates and metabolic traits, it does not adequately describe two aspects of the thermal performance curve (TPC) for metabolic rate-that metabolic rate is a unimodal function of temperature often with maximal values in the biologically relevant temperature range and that activation energies are temperature dependent. We show that the temperature dependence of metabolic rate in ectotherms is well described by an enzyme-assisted Arrhenius (EAAR) model that accounts for the temperature-dependent contribution of enzymes to decreasing the activation energy required for reactions to occur. The model is mechanistically derived using the thermodynamic rules that govern protein stability. We contrast our model with other unimodal functions that also can be used to describe the temperature dependence of metabolic rate to show how the EAAR model provides an important advance over previous work. We fit the EAAR model to metabolic rate data for a variety of taxa to demonstrate the model's utility in describing metabolic rate TPCs while revealing significant differences in thermodynamic properties across species and acclimation temperatures. Our model advances our ability to understand the metabolic and ecological consequences of increases in the mean and variance of temperature associated with global climate change. In addition, the model suggests avenues by which organisms can acclimate and adapt to changing thermal environments. Furthermore, the parameters in the EAAR model generate links between organismal level performance and underlying molecular processes that can be tested for in future work.

Virus population growth depends on contacts between viruses and their hosts. It is often unclear how sufficient contacts are made between viruses and their specific hosts to generate spikes in viral abundance. Here, we show that copepods, acting as predators, can bring aquatic viruses and their algal hosts into contact. Specifically, predation of the protist Paramecium bursaria by copepods resulted in a >100-fold increase in the number of chloroviruses in 1 d. Copepod predation can be seen as an ecological "catalyst" by increasing contacts between chloroviruses and their hosts, zoochlorellae (endosymbiotic algae that live within paramecia), thereby facilitating viral population growth. When feeding, copepods passed P. bursaria through their digestive tract only partially digested, releasing endosymbiotic algae that still supported viral reproduction and resulting in a virus population spike. A simple predator-prey model parameterized for copepods consuming protists generates cycle periods for viruses consistent with those observed in natural ponds. Food webs are replete with similar symbiotic organisms, and we suspect the predator catalyst mechanism is capable of generating blooms for other endosymbiont-targeting viruses., Competing Interests: The authors declare no conflict of interest.

Ectotherms generally demonstrate nonlinear changes in performance (e.g., movement speed, individual growth, population growth) as a function of temperature that are characterized by thermal performance curves (TPC). Predation risk elicits phenotypic and behavioral changes that likewise impact performance measures. We tested whether exposure to predation Orthocyclops modestus impacts the maximum population growth rate ( r max ) TPC of the protist Paramecium aurelia . We fit predator and non-predator exposed P. aurelia population growth rates to a function previously shown to best describe Paramecium population growth rate TPC's (Lactin-2) and compared subsequent parameter estimates between curves. For Paramecium exposed to predation risk, maximum population growth increased more rapidly as temperatures rose and decreased more rapidly as temperatures fell compared to the initial temperature. The area under each TPC curve remained approximately the same, consistent with the idea of a trade-off in performance across temperatures. Our results indicate TPCs are flexible given variation in food web context and that trophic interactions may play an important role in shaping TPCs. Furthermore, this and other studies illustrate the need for a mechanistic model of TPCs with parameters tied to biologically meaningful properties.

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., (© 2016 John Wiley & Sons Ltd.)

'Demographic transition theory' assumes that fertility decline is irreversible. This commonly held assumption is based on observations of recent and historical reductions in fertility that accompany modernization and declining mortality. The irreversibility assumption, however, is highly suspect from an evolutionary point of view, because demographic traits are at least partially influenced by genetics and are responsive to social and ecological conditions. Nonetheless, an inevitable shift from high mortality and fertility to low mortality and fertility is used as a guiding framework for projecting human population sizes into the future. This paper reviews some theoretical and empirical evidence suggesting that the assumption of irreversibility is ill-founded, at least without considerable development in theory that incorporates evolutionary and ecological processes. We offer general propositions for how fertility could increase in the future, including natural selection on high fertility variants, the difficulty of maintaining universal norms and preferences in a large, diverse and economically differentiated population, and the escalating resource demands of modernization., (© 2016 The Author(s).)

Understanding the factors that determine the strength of predator-prey interactions is essential to understanding community structure and stability. Variation in the strength of predator-prey interactions often can be attributed to predator mass and prey mass, or abiotic factors like temperature. However, even when accounting for these factors, there remains a considerable amount of unexplained variation that may be attributed to other traits. We compiled functional response data from the literature to investigate how predator mass, prey mass, prey type (taxonomic identity), temperature, and prey defenses (hard vs soft integument) contributed to the variation found in the predator-prey interactions between freshwater cyclopoid copepods and their prey. Surprisingly, our results indicate that prey identity (taxonomic group) and defenses (hard vs soft integument) are more important for generating variation in interaction strengths than body mass and temperature. This suggests that allometric functions can only take us so far when attempting to better understand variation in individual predator prey interactions, and that we must evaluate how other traits influence interaction strengths. Identifying additional factors such as prey defenses may enable us to better predict potential changes in the structure and function of planktonic and other food webs by better accounting for the variation in the interactions between generalists and their many prey types.

Heritable trait variation is a central and necessary ingredient of evolution. Trait variation also directly affects ecological processes, generating a clear link between evolutionary and ecological dynamics. Despite the changes in variation that occur through selection, drift, mutation, and recombination, current eco-evolutionary models usually fail to track how variation changes through time. Moreover, eco-evolutionary models assume fitness functions for each trait and each ecological context, which often do not have empirical validation. We introduce a new type of model, Gillespie eco-evolutionary models (GEMs), that resolves these concerns by tracking distributions of traits through time as eco-evolutionary dynamics progress. This is done by allowing change to be driven by the direct fitness consequences of model parameters within the context of the underlying ecological model, without having to assume a particular fitness function. GEMs work by adding a trait distribution component to the standard Gillespie algorithm - an approach that models stochastic systems in nature that are typically approximated through ordinary differential equations. We illustrate GEMs with the Rosenzweig-MacArthur consumer-resource model. We show not only how heritable trait variation fuels trait evolution and influences eco-evolutionary dynamics, but also how the erosion of variation through time may hinder eco-evolutionary dynamics in the long run. GEMs can be developed for any parameter in any ordinary differential equation model and, furthermore, can enable modeling of multiple interacting traits at the same time. We expect GEMs will open the door to a new direction in eco-evolutionary and evolutionary modeling by removing long-standing modeling barriers, simplifying the link between traits, fitness, and dynamics, and expanding eco-evolutionary treatment of a greater diversity of ecological interactions. These factors make GEMs much more than a modeling advance, but an important conceptual advance that bridges ecology and evolution through the central concept of heritable trait variation.