Warming and top-down control of stage-structured prey: Linking theory to patterns in natural systems.

Warming has broad and often nonlinear impacts on organismal physiology and traits, allowing it to impact species interactions like predation through a variety of pathways that may be difficult to predict. Predictions are commonly based on short-term experiments and models, and these studies often yield conflicting results depending on the environmental context, spatiotemporal scale, and the predator and prey species considered. Thus, the accuracy of predicted changes in interaction strength, and their importance to the broader ecosystems they take place in, remain unclear. Here, we attempted to link one such set of predictions generated using theory, modeling, and controlled experiments to patterns in the natural abundance of prey across a broad thermal gradient. To do so, we first predicted how warming would impact a stage-structured predator-prey interaction in riverine rock pools between Pantala spp. dragonfly nymph predators and Aedes atropalpus mosquito larval prey. We then described temperature variation across a set of hundreds of riverine rock pools (n = 775) and leveraged this natural gradient to look for evidence for or against our model's predictions. Our model's predictions suggested that warming should weaken predator control of mosquito larval prey by accelerating their development and shrinking the window of time during which aquatic dragonfly nymphs could consume them. This was consistent with data collected in rock pool ecosystems, where the negative effects of dragonfly nymph predators on mosquito larval abundance were weaker in warmer pools. Our findings provide additional evidence to substantiate our model-derived predictions while emphasizing the importance of assessing similar predictions using natural gradients of temperature whenever possible.

Incorporating nonlinearity with generalized functional responses to simulate multiple predator effects.

Predicting the combined effects of predators on shared prey has long been a focus of community ecology, yet quantitative predictions often fail. Failure to account for nonlinearity is one reason for this. Moreover, prey depletion in multiple predator effects (MPE) studies generates biased predictions in applications of common experimental and quantitative frameworks. Here, we explore additional sources of bias stemming from nonlinearities in prey predation risk. We show that in order to avoid bias, predictions about the combined effects of independent predators must account for nonlinear size-dependent risk for prey as well as changes in prey risk driven by nonlinear predator functional responses and depletion. Historical failure to account for biases introduced by well-known nonlinear processes that affect predation risk suggest that we may need to reevaluate the general conclusions that have been drawn about the ubiquity of emergent MPEs over the past three decades.

Phenology of Rock Pool Mosquitoes in the Southern Appalachian Mountains: Surveys Reveal Apparent Winter Hatching of Aedes japonicus and the Potential For Asymmetrical Stage-Specific Interactions.

The North American rock pool mosquito, Aedes atropalpus, has reportedly decreased in abundance following the introduction of Ae. japonicus japonicus to the USA, but the specific mechanisms responsible for the reduction remain unclear. Thus, there is a need for field studies to improve our knowledge of natural rock pool systems where both species co-occur. We sampled rock pool invertebrates over a 12-month period along the Chattooga River at a high-elevation site (728 m) near Cashiers, NC, and at a lower-elevation site (361 m) near Clayton, GA. We identified 12 orders of macroinvertebrates representing at least 19 families and 5 mosquito species. Aedes j. japonicus was present year-round at both sites. We observed overwintering Ae. j. japonicus larvae in pools with water temperatures as cold as 3°C and detected apparent winter egg hatching in water below 10°C. Aedes atropalpus was rarely encountered at the high-elevation site but was highly abundant in the summer months at the low-elevation site. Late-stage Ae. j. japonicus larvae inhabited pools in March 2019 when Ae. atropalpus first appeared in the same pools, creating the potential for asymmetrical stage-specific interactions. Our observations provide evidence of overwintering and early hatching of Ae. j. japonicus in the southeastern climate. Further study of the importance of stage-dependent competition and winter egg hatching of diapausing Ae. j. japonicus eggs is warranted.

Aquatic thermal conditions predict the presence of native and invasive rock pool Aedes (Diptera: Culicidae) in the southern Appalachians, U.S.A.

The native rock pool mosquito, Aedes atropalpus (Coquillett), and the invasive Aedes japonicus (Theobald) have been found in many types of artificial and natural containers throughout North America. Little is known about the ecology of these two species in habitats where they co-occur, although multiple investigators have reported the decline of the native species concurrent with the introduction and spread of the invasive species. Here we report the results of riverine rock pool collections (n=503) in the southern Appalachian Mountains between 2009-2015. Surface water temperatures strongly predicted the presence of each species across a broad range of observed temperatures (11-39.8° C). For every unit of increase in temperature (°C) the odds of collecting Ae. atropalpus larvae increased by 0.34 while the odds of collecting Ae. japonicus larvae decreased by 0.28. No Ae. japonicus larvae or pupae were collected at temperatures greater than 36° C; however, immature Ae. atropalpus were found in rock pools with temperatures up to 39.8° C. In contrast, Ae. japonicus were highly abundant in cooler rock pools (<17° C) where Ae. atropalpus were infrequent or absent. Our findings suggest that in spite of the successful invasion by Ae. japonicus, Ae. atropalpus remains well established in the southern Appalachian Mountains. Given the strong correlation of temperature with the presence of the two species and the contrasting absence of each species at observed temperature extremes, the role of thermal conditions should be carefully explored in the context of other ecological factors likely influencing the range and abundance of these mosquitoes.

Size-dependent functional response of Xenopus laevis feeding on mosquito larvae.

Predators can play an important role in regulating prey abundance and diversity, determining food web structure and function, and contributing to important ecosystem services, including the regulation of agricultural pests and disease vectors. Thus, the ability to predict predator impact on prey is an important goal in ecology. Often, predators of the same species are assumed to be functionally equivalent, despite considerable individual variation in predator traits known to be important for shaping predator-prey interactions, like body size. This assumption may greatly oversimplify our understanding of within-species functional diversity and undermine our ability to predict predator effects on prey. Here, we examine the degree to which predator-prey interactions are functionally homogenous across a natural range of predator body sizes. Specifically, we quantify the size-dependence of the functional response of African clawed frogs (Xenopus laevis) preying on mosquito larvae (Culex pipiens). Three size classes of predators, small (15-30 mm snout-vent length), medium (50-60 mm) and large (105-120 mm), were presented with five densities of prey to determine functional response type and to estimate search efficiency and handling time parameters generated from the models. The results of mesocosm experiments showed that type of functional response of X. laevis changed with size: small predators exhibited a Type II response, while medium and large predators exhibited Type III responses. Functional response data showed an inversely proportional relationship between predator attack rate and predator size. Small and medium predators had highest and lowest handling time, respectively. The change in functional response with the size of predator suggests that predators with overlapping cohorts may have a dynamic impact on prey populations. Therefore, predicting the functional response of a single size-matched predator in an experiment may misrepresent the predator's potential impact on a prey population.

Variability in life-history switch points across and within populations explained by Adaptive Dynamics.

Understanding the factors that shape the timing of life-history switch points (SPs; e.g. hatching, metamorphosis and maturation) is a fundamental question in evolutionary ecology. Previous studies examining this question from a fitness optimization perspective have advanced our understanding of why the timing of life-history transitions may vary across populations and environments. However, in nature we also often observe variability among individuals within populations. Optimization theory, which typically predicts a single optimal SP under physiological and environmental constraints for a given environment, cannot explain this variability. Here, we re-examine the evolution of a single life-history SP between juvenile and adult stages from an Adaptive Dynamics (AD) perspective, which explicitly considers the feedback between the dynamics of population and the evolution of life-history strategy. The AD model, although simple in structure, exhibits a diverse range of evolutionary scenarios depending upon demographic and environmental conditions, including the loss of the juvenile stage, a single optimal SP, alternative optimal SPs depending on the initial phenotype, and sympatric coexistence of two SP phenotypes under disruptive selection. Such predictions are consistent with previous optimization approaches in predicting life-history SP variability across environments and between populations, and in addition they also explain within-population variability by sympatric disruptive selection. Thus, our model can be used as a theoretical tool for understanding life-history variability across environments and, especially, within species in the same environment.

Egg clutch dehydration induces early hatching in red-eyed treefrogs, Agalychnis callidryas.

Terrestrial eggs have evolved repeatedly in tropical anurans exposing embryos to the new threat of dehydration. Red-eyed treefrogs, Agalychnis callidryas, lay eggs on plants over water. Maternally provided water allows shaded eggs in humid sites to develop to hatching without rainfall, but unshaded eggs and those in less humid sites can die from dehydration. Hatching responses of amphibian eggs to dry conditions are known from two lineages with independent origins of terrestrial eggs. Here, we experimentally tested for dehydration-induced early hatching in another lineage (Agalychnis callidryas, Phyllomedusidae), representing a third independent origin of terrestrial eggs. We also investigated how dehydration affected egg and clutch structure, and egg mortality. We collected clutches from a pond in Gamboa, Panama, and randomly allocated them to wet or dry treatments at age 1 day. Embryos hatched earlier from dry clutches than from wet clutches, accelerating hatching by ∼11%. Clutch thickness and egg diameter were affected by dehydration, diverging between treatments over time. Meanwhile, mortality in dry clutches was six-fold higher than in control clutches. With this study, early hatching responses to escape mortality from egg dehydration are now known from three anuran lineages with independent origins of terrestrial eggs, suggesting they may be widespread. Further studies are needed to understand how terrestrial amphibian eggs can respond to, or will be affected by, rapid changes in climate over the next decades.

Predator diversity reduces habitat colonization by mosquitoes and midges.

Changes in predator diversity via extinction and invasion are increasingly widespread and can have important ecological and socio-economic consequences. Anticipating and managing these consequences requires understanding how predators shape ecological communities. Previous predator biodiversity research has focused on post-colonization processes. However, predators can also shape communities by altering patterns of prey habitat selection during colonization. The sensitivity of this non-consumptive top down mechanism to changes in predator diversity is largely unexamined. To address this gap, we examined patterns of dipteran oviposition habitat selection in experimental aquatic habitats in response to varied predator species richness while holding predator abundance constant. Caged predators were used in order to disentangle behavioural oviposition responses to predator cues from potential post-oviposition consumption of eggs and larvae. We hypothesized that because increases in predator richness often result in greater prey mortality than would be predicted from independent effects of predators, prey should avoid predator-rich habitats during colonization. Consistent with this hypothesis, predator-rich habitats received 48% fewer dipteran eggs than predicted, including 60% fewer mosquito eggs and 38% fewer midge eggs. Our findings highlight the potentially important links between predator biodiversity, prey habitat selection and the ecosystem service of pest regulation.

Bioinsecticide and leaf litter combination increases oviposition and reduces adult recruitment to create an effective ovitrap for Culex mosquitoes.

Mosquito egg traps, aquatic habitats baited with oviposition attractant and insecticide, are important tools for surveillance and control efforts in integrated vector management programs. The bioinsecticide Bacillus thuringiensis subsp. israelensis (Bti) is increasingly used as an environmentally friendly alternative to chemical insecticides and the combination of Bti with a simple oviposition attractant like leaf litter to create an effective egg trap seems appealing. However, previous research suggests that Bti may itself alter oviposition, and that leaf litter may dramatically reduce Bti toxicity. Here we present results from field experiment designed to link the effects of litter and Bti on mosquito oviposition habitat selection and post-colonization survival to production of adult mosquitoes. Tripling litter increased Culex spp. oviposition nearly nine-fold, while Bti had no effect on oviposition. Neither factor altered egg survival, thus larval abundance reflected the effects of litter on oviposition. Both Bti and litter reduced larval survival by ∼60%. We found no evidence that increased litter reduced Bti toxicity. Adult production was dependent upon both litter and Bti. In the absence of Bti, effects of litter on oviposition translated into three-fold more adults. However, in the presence of Bti, initial increases in oviposition were erased by the combined negative effects of Bti and litter on post-colonization survival. Thus, our study provides field evidence that combined litter and Bti application creates an effective ovitrap. This combined treatment had the highest oviposition and the lowest survival, and thus removed the greatest number of mosquitoes from the landscape.

Risk assessment based on indirect predation cues: revisiting fine-grained variation.

To adaptively express inducible defenses, prey must gauge risk based on indirect cues of predation. However, the information contained in indirect cues that enable prey to fine-tune their phenotypes to variation in risk is still unclear. In aquatic systems, research has focused on cue concentration as the key variable driving threat-sensitive responses to risk. However, while risk is measured as individuals killed per time, cue concentration may vary with either the number or biomass killed. Alternatively, fine-grained variation in cue, that is, frequency of cue pulses irrespective of concentration, may provide a more reliable signal of risk. Here, we present results from laboratory experiments that examine the relationship between red-eyed treefrog tadpole growth and total cue, cue per pulse, and cue pulse frequency. We also reanalyze an earlier study that examined the effect of fine-grained variation in predator cues on wood frog tadpole growth. Both studies show growth declines with increasing cue pulse frequency, even though individual pulses in high-frequency treatments contained very little cue. This result suggests that counter to earlier conclusions, tadpoles are using fine-grained variation in cue arising from the number of predation events to assess and respond to predation risk, as predicted by consumer-resource theory.

Consequences of life history switch point plasticity for juvenile morphology and locomotion in the Túngara frog.

Many animals with complex life cycles can cope with environmental uncertainty by altering the timing of life history switch points through plasticity. Pond hydroperiod has important consequences for the fitness of aquatic organisms and many taxa alter the timing of life history switch points in response to habitat desiccation. For example, larval amphibians can metamorphose early to escape drying ponds. Such plasticity may induce variation in size and morphology of juveniles which can result in carry-over effects on jumping performance. To investigate the carry-over effects of metamorphic plasticity to pond drying, we studied the Túngara frog, Physalaemus pustulosus, a tropical anuran that breeds in highly ephemeral habitats. We conducted an outdoor field mesocosm experiment in which we manipulated water depth and desiccation and measured time and size at metamorphosis, tibiofibula length and jumping performance. We also conducted a complimentary laboratory experiment in which we manipulated resources, water depth and desiccation. In the field experiment, metamorphs from dry-down treatments emerged earlier, but at a similar size to metamorphs from constant depth treatments. In the laboratory experiment, metamorphs from the low depth and dry-down treatments emerged earlier and smaller. In both experiments, frogs from dry-down treatments had relatively shorter legs, which negatively impacted their absolute jumping performance. In contrast, reductions in resources delayed and reduced size at metamorphosis, but had no negative effect on jumping performance. To place these results in a broader context, we review past studies on carry-over effects of the larval environment on jumping performance. Reductions in mass and limb length generally resulted in lower jumping performance across juvenile anurans tested to date. Understanding the consequences of plasticity on size, morphology and performance can elucidate the linkages between life stages.

Putting µ/g in a new light: plasticity in life history switch points reflects fine-scale adaptive responses.

Life history theory predicts that organisms with complex life cycles should transition between life stages when the ratio of growth rate (g) to risk of mortality (µ) in the current stage falls below that in the subsequent stage. Empirical support for this idea has been mixed. Implicit in both theory and empirical work is that the risk of mortality in the subsequent stage is unknown. However, some embryos and larvae of both vertebrates and invertebrates assess cues of post-transition predation risk and alter the timing of hatching or metamorphosis accordingly. Furthermore, although life history switch points of prey have traditionally been treated as discrete shifts in morphology or habitat, for many organisms they are continuous transitional periods within which the timing of specific developmental and behavioral events can be plastic. We studied red-eyed treefrogs (Agalychnis callidryas), which detect predators of both larvae and metamorphs, to test if plastic changes during the process of metamorphosis could reconcile the mismatch between life history theory and empirical data and if plasticity in an earlier stage transition (hatching) would affect plasticity at a subsequent stage transition (metamorphosis). We reared tadpoles from hatching until metamorphosis in a full-factorial cross of two hatching ages (early- vs. late-hatched) and the presence or absence of free-roaming predators of larvae (giant water bugs) and metamorphs (fishing spiders). Hatching age affected the times from oviposition to tail resorption and from hatching to emergence onto land, but did not alter responses to predators or developmental stage at emergence. Tadpoles did not alter their age at emergence or tail resorption in response to larval or metamorph predators, despite the fact that predators reduced tadpole density by ~30%. However, developmental stage at emergence and time needed to complete metamorphosis in the terrestrial environment were plastic and consistent with predictions of the "minimize µ/g" framework. Our results demonstrate that likely adaptive changes in life history transitions occur at previously unappreciated timescales. Consideration of plasticity in the developmental timing of ecologically important events within metamorphosis, rather than treating it as a discrete switch point, may help to reconcile inconsistencies between empirical studies of predator effects and expectations of long-standing ecological theory.

Consequences of induced hatching plasticity depend on predator community.

Many prey species face trade-offs in the timing of life history switch points like hatching and metamorphosis. Costs associated with transitioning early depend on the biotic and abiotic conditions found in the subsequent life stage. The red-eyed treefrog, Agalychnis callidryas, faces risks from predators in multiple, successive life stages, and can hatch early in response to mortality threats at the egg stage. Here we tested how the consequences of life history plasticity, specifically early hatching in response to terrestrial egg predators, depend on the assemblage of aquatic larval predators. We predicted that diverse predator assemblages would impose lower total predation pressure than the most effective single predator species and might thereby reduce the costs of hatching early. We then conducted a mesocosm experiment where we crossed hatchling phenotype (early vs. normal hatching) with five larval-predator environments (no predators, either waterbugs, dragonflies, or mosquitofish singly, or all three predator species together). The consequences of hatching early varied across predator treatments, and tended to disappear through time in some predation treatments, notably the waterbug and diverse predator assemblages. We demonstrate that the fitness costs of life history plasticity in an early life stage depend critically on the predator community composition in the next stage.

Behavioral plasticity mitigates risk across environments and predators during anuran metamorphosis.

Most animals metamorphose, changing morphology, physiology, behavior and ecological interactions. Size- and habitat-dependent mortality risk is thought to affect the evolution and plastic expression of metamorphic timing, and high predation during the morphological transition is posited as a critical selective force shaping complex life cycles. Nonetheless, empirical data on how risk changes across metamorphosis and stage-specific habitats, or how that varies with size, are rare. We examined predator-prey interactions of red-eyed treefrogs, Agalychnis callidryas, with an aquatic predator (giant water bug, Belostoma) and a semi-terrestrial predator (fishing spider, Thaumasia) across metamorphosis. We manipulated tadpole density to generate variation in metamorph size and conducted predation trials at multiple developmental stages. We quantified how frog behavior (activity) changes across metamorphic development, habitats, and predator presence or absence. In aquatic trials with water bugs, frog mortality increased with forelimb emergence, as hypothesized. In semi-terrestrial trials, contrary to predictions, predation by spiders increased, not decreased, with tail resorption. In neither case did frog size affect mortality. Frogs reduced activity upon forelimb emergence in the water, and further with emergence into air, then increased activity with tail resorption. Longer-tailed metamorphs were captured more often in spider attacks, but attacked less, as most attacks followed prey movements. Metamorphs behaviorally compensated for poor escape performance more effectively on land than in water, thus emergence timing may critically affect mortality. The developmental timing of the ecological transition between environments that select for different larval and juvenile phenotypes is an important, neglected variable in studies of complex life cycles.

Prey subsidy or predator cue? Direct and indirect effects of caged predators on aquatic consumers and resources.

The non-consumptive effects of predators on prey can affect prey phenotypes, potentially having important consequences for communities due to trait-mediated indirect interactions. Predicting non-consumptive effects and their impacts on communities can be difficult because predators can affect resources directly through nutrient cycling and indirectly by altering prey resource use, which can lead to complex interactions among resources and consumers. In this study we examined the effects of caged dragonfly predators on aquatic resources in the presence and absence of two focal herbivores, the tadpoles of Neotropical tree frogs Agalychnis callidryas and Dendropsophus ebraccatus. We crossed the presence/absence of caged dragonflies with four tadpole treatments: no tadpoles, each tadpole species alone, and both species together to examine interactions among tadpole composition, predator presence, and time on tadpole growth, resources, and zooplankton abundances. Predator effects on growth changed through ontogeny and was species-dependent. Predators initially reduced then dramatically increased A. callidryas growth, but had no effect on D. ebraccatus. Predators also increased the abundances of both periphyton and phytoplankton. However, there was no evidence of a trait-mediated trophic cascade (i.e., tadpole by predator interaction). Instead, nutrients from prey carcass subsidies likely played an increasingly important role in facilitating resources, and shaping tadpole growth, competitive interactions, and zooplankton abundances through time. In nutrient-poor aquatic systems the release of nutrients via the consumption of terrestrially derived prey items by aquatic predators may have important impacts on food webs by facilitating resources independent of the role of trait-mediated trophic cascades.

Costs of predator-induced phenotypic plasticity: a graphical model for predicting the contribution of nonconsumptive and consumptive effects of predators on prey.

Defensive modifications in prey traits that reduce predation risk can also have negative effects on prey fitness. Such nonconsumptive effects (NCEs) of predators are common, often quite strong, and can even dominate the net effect of predators. We develop an intuitive graphical model to identify and explore the conditions promoting strong NCEs. The model illustrates two conditions necessary and sufficient for large NCEs: (1) trait change has a large cost, and (2) the benefit of reduced predation outweighs the costs, such as reduced growth rate. A corollary condition is that potential predation in the absence of trait change must be large. In fact, the sum total of the consumptive effects (CEs) and NCEs may be any value bounded by the magnitude of the predation rate in the absence of the trait change. The model further illustrates how, depending on the effect of increased trait change on resulting costs and benefits, any combination of strong and weak NCEs and CEs is possible. The model can also be used to examine how changes in environmental factors (e.g., refuge safety) or variation among predator-prey systems (e.g., different benefits of a prey trait change) affect NCEs. Results indicate that simple rules of thumb may not apply; factors that increase the cost of trait change or that increase the degree to which an animal changes a trait, can actually cause smaller (rather than larger) NCEs. We provide examples of how this graphical model can provide important insights for empirical studies from two natural systems. Implementation of this approach will improve our understanding of how and when NCEs are expected to dominate the total effect of predators. Further, application of the models will likely promote a better linkage between experimental and theoretical studies of NCEs, and foster synthesis across systems.

Prey responses to predator chemical cues: disentangling the importance of the number and biomass of prey consumed.

To effectively balance investment in predator defenses versus other traits, organisms must accurately assess predation risk. Chemical cues caused by predation events are indicators of risk for prey in a wide variety of systems, but the relationship between how prey perceive risk in relation to the amount of prey consumed by predators is poorly understood. While per capita predation rate is often used as the metric of relative risk, studies aimed at quantifying predator-induced defenses commonly control biomass of prey consumed as the metric of risk. However, biomass consumed can change by altering either the number or size of prey consumed. In this study we determine whether phenotypic plasticity to predator chemical cues depends upon prey biomass consumed, prey number consumed, or both. We examine the growth response of red-eyed treefrog tadpoles (Agalychnis callidryas) to cues from a larval dragonfly (Anax amazili). Biomass consumed was manipulated by either increasing the number of prey while holding individual prey size constant, or by holding the number of prey constant and varying individual prey size. We address two questions. (i) Do prey reduce growth rate in response to chemical cues in a dose dependent manner? (ii) Does the magnitude of the response depend on whether prey consumption increases via number or size of prey? We find that the phenotypic response of prey is an asymptotic function of prey biomass consumed. However, the asymptotic response is higher when more prey are consumed. Our findings have important implications for evaluating past studies and how future experiments should be designed. A stronger response to predation cues generated by more individual prey deaths is consistent with models that predict prey sensitivity to per capita risk, providing a more direct link between empirical and theoretical studies which are often focused on changes in population sizes not individual biomass.

Spatial contagion drives colonization and recruitment of frogflies on clutches of red-eyed treefrogs.

Spatial contagion occurs when the perceived suitability of neighbouring habitat patches is not independent. As a result, organisms may colonize less-preferred patches near preferred patches and avoid preferred patches near non-preferred patches. Spatial contagion may thus alter colonization dynamics as well as the type and frequency of post-colonization interactions. Studies have only recently documented the phenomenon of spatial contagion and begun to examine its consequences for local recruitment. Here, we test for spatial contagion in the colonization of arboreal egg clutches of red-eyed treefrogs by a frogfly and examine the consequences of contagion for fly recruitment. In laboratory choice experiments, flies oviposit almost exclusively on clutches containing dead frog eggs. In nature, however, flies often colonize intact clutches without dead eggs. Consistent with predictions of contagion-induced oviposition, we found that flies more frequently colonize intact clutches near damaged clutches and rarely colonize intact clutches near other intact clutches. Moreover, contagion appears to benefit flies. Flies survived equally well and suffered less parasitism on clutches lacking dead eggs. This study demonstrates how reward contagion can influence colonization dynamics and suggests that colonization patterns caused by contagion may have important population- and community-level consequences.

Fluxes of terrestrial and aquatic carbon by emergent mosquitoes: a test of controls and implications for cross-ecosystem linkages.

Adult aquatic insects are a common resource for many terrestrial predators, often considered to subsidize terrestrial food webs. However, larval aquatic insects themselves consume both aquatic primary producers and allochthonous terrestrial detritus, suggesting that adults could provide aquatic subsidy and/or recycled terrestrial energy to terrestrial consumers. Understanding the source of carbon (aquatic vs. terrestrial) driving aquatic insect emergence is important for predicting magnitude of emergence and effects on recipient food web dynamics; yet direct experimental tests of factors determining source are lacking. Here, we use Culex mosquitoes in experimental pools as an exemplar to test how variation in general factors common to aquatic systems (terrestrial plant inputs and light) may alter the source and amount of energy exported to terrestrial ecosystems in adult aquatic insects that rely on terrestrial resources as larvae. We found strong sequential effects of terrestrial plant inputs and light on aquatic insect oviposition, diet, and emergence of Culex mosquitoes. Ovipositing mosquitoes laid ~3 times more egg masses in high terrestrial input pools under low light conditions. This behavior increased adult emergence from pools under low light conditions; however, high input pools (which had the highest mosquito densities) showed low emergence rates due to density-dependent mortality. Mosquito diets consisted mainly of terrestrial resources (~70-90 %). As a result, the amount of aquatic carbon exported from pools by mosquitoes during the experiment was ~18 times higher from low versus high light pools, while exports of terrestrial carbon peaked from pools receiving intermediate levels of inputs (3-6 times higher) and low light (~6 times higher). Our results suggest that understanding the interplay among terrestrial plant inputs, light availability and biotic responses of aquatic insects may be key in predicting source and magnitude of emergence, and thus the strength and effects of aquatic-terrestrial linkages in freshwater systems.