Trait-mediated Effects Research Articles

Predator-induced prey dispersal can cause hump-shaped density-area relationships in prey populations.

Predation can both reduce prey abundance directly (through density-dependent effects) and indirectly through prey trait-mediated effects. Over the years, many studies have focused on describing the density-area relationship (DAR). However, the mechanisms responsible for the DAR are not well understood. Loss and fragmentation of habitats, owing to human activities, creates landscape-level spatial heterogeneity wherein patches of varying size, isolation and quality are separated by a human-modified "matrix" of varying degrees of hostility and has been a primary driver of species extinctions and declining biodiversity. How matrix hostility in combination with trait-mediated effects influence DAR, minimum patch size, and species coexistence remains an open question. In this paper, we employ a theoretical spatially explicit predator-prey population model built upon the reaction-diffusion framework to explore effects of predator-induced emigration (trait-mediated emigration) and matrix hostility on DAR, minimum patch size, and species coexistence. Our results show that when trait-mediated response strength is sufficiently strong, ranges of patch size emerge where a nonlinear hump-shaped prey DAR is predicted and other ranges where coexistence is not possible. In a conservation perspective, DAR is crucial not only in deciding whether we should have one large habitat patch or several-small (SLOSS), but for understanding the minimum patch size that can support a viable population. Our study lends more credence to the possibility that predators can alter prey DAR through predator-induced prey dispersal.

Trait- and density-mediated effects of predation on fecundity and fertility of Aedes aegypti (Diptera: Culicidae) mosquitoes.

Prey populations that encounter predators experience density-mediated effects through lethality and associated numerical changes in the population. Prey also exhibit trait-mediated effects through nonlethal alterations in phenotypic traits associated with exposure to predators. Immature stages of mosquitoes commonly co-occur in habitats along with predators, a natural source of mortality and potential biocontrol agent. Toxorhynchites rutilus Coquillett 1896 is a natural source of mortality with potential as a biological control agent. Previous studies have shown that predation and the mere presence of Tx. rutilus (predator cues) can alter the life-history traits of Aedes aegypti (L. 1762). In addition to observed reductions in recruitment of adults (lethality), exposure to Tx. rutilus without consumption resulted in adult Ae. aegypti females with altered growth and reduced lifespan. To determine the influence of predation on the reproductive biology of Ae. aegypti, we tested the hypothesis that predation, or exposure to predator cues, will compromise the reproduction of adult survivors through reductions in fecundity (egg batch size) and fertility (hatch rate). We observed that for both female and male Ae. aegypti, survival to adulthood was the lowest and development time the shortest in treatments containing prey removal effects, attributable to predation and treatment manipulations of density reduction. There were effects of Ae. aegypti weight, but not predation treatments, on fecundity and fertility. Results suggest that predator-mediated effects on Ae. aegypti derive from lethal effects due to consumption and alterations in other phenotypic traits of survivors, including development, weight, lifespan of adults, and population growth, but not reproductive parameters measured here.

Landscape structure influences grassland productivity through plant functional diversity

Agriculture intensification led to the gradual destruction of semi-natural habitats and landscape simplification in the North-Western Europe. All these profound changes affected plant assemblages, especially in semi-natural ecosystems such as grasslands. They could have negative effects on the plant diversity and the related ecosystem functions, such as primary productivity. At the landscape scale this biodiversity-productivity relationship still have to be tested in real-world systems. It could have important economic implications concerning fodder production. We used a large-scale sampling design (30 landscape windows – 1 ×1 km) to quantify landscape structure variables (semi-natural habitat amounts and landscape heterogeneity), grassland plant taxonomic diversity and functional diversity calculated from multidimensional functional spaces (functional richness, evenness, dispersion), and grassland productivity (mean, temporal stability, spatial variability) derived from satellite image time-series. Taxonomic dissimilarity, functional evenness and functional dispersion of grassland plant assemblages were related to semi-natural habitat amount. Functional evenness and dispersion decreased with grassland amount whereas taxonomic dissimilarity and functional evenness increased with hedgerow amount. Only taxonomic and functional richness depended on landscape heterogeneity, and with a positive relationship. Grassland plant assemblages with a higher functional evenness presented a higher and more stable mean productivity. In addition to the direct effect of landscape heterogeneity on plant productivity, we demonstrated plant trait-mediated effect of landscape structure on mean and temporal stability of grassland productivity at the landscape scale. Landscape management, especially through the conservation of hedgerows can help maintain high and resilient production of fodder in the future as a result of their positive effects on plant functional diversity.

A trait-based framework for dung beetle functional ecology.

Traits are key for understanding the environmental responses and ecological roles of organisms. Trait approaches to functional ecology are well established for plants, whereas consistent frameworks for animal groups are less developed. Here we suggest a framework for the study of the functional ecology of animals from a trait-based response-effect approach, using dung beetles as model system. Dung beetles are a key group of decomposers that are important for many ecosystem processes. The lack of a trait-based framework tailored to this group has limited the use of traits in dung beetle functional ecology. We review which dung beetle traits respond to the environment and affect ecosystem processes, covering the wide range of spatial, temporal and biological scales at which they are involved. Dung beetles show trait-based responses to variation in temperature, water, soil properties, trophic resources, light, vegetation structure, competition, predation and parasitism. Dung beetles' influence on ecosystem processes includes trait-mediated effects on nutrient cycling, bioturbation, plant growth, seed dispersal, other dung-based organisms and parasite transmission, as well as some cases of pollination and predation. We identify 66 dung beetle traits that are either response or effect traits, or both, pertaining to six main categories: morphology, feeding, reproduction, physiology, activity and movement. Several traits pertain to more than one category, in particular dung relocation behaviour during nesting or feeding. We also identify 136 trait-response and 77 trait-effect relationships in dung beetles. No response to environmental stressors nor effect over ecological processes were related with traits of a single category. This highlights the interrelationship between the traits shaping body-plans, the multi-functionality of traits, and their role linking responses to the environment and effects on the ecosystem. Despite current developments in dung beetle functional ecology, many knowledge gaps remain, and there are biases towards certain traits, functions, taxonomic groups and regions. Our framework provides the foundations for the thorough development of trait-based dung beetle ecology. It also serves as an example framework for other taxa.

Dogs suppress a pivotal function in the food webs of sandy beaches

Domestic dogs are the most abundant carnivore globally and have demonstrable negative impacts to wildlife; yet, little evidence regarding their functional roles in natural food webs exists. Adding dogs to food webs may result in a net loss (via suppression of naturally occurring species), net gain (via mesopredator release), or no change (via functional replacement) to ecosystem function. Scavenging is a pivotal function in ecosystems, particularly those that are energetically supported by carrion. Dogs also scavenge on animal carcasses, but whether scavenging by dogs influences the structural and functional properties of food webs remains unclear. Here we used camera traps baited with carrion to test the effect of dogs on the composition and diversity of the vertebrate scavenger guild, as well as carrion detection and consumption rates. We conducted this work in sandy beach ecosystems, which rely on the import of marine organic matter (i.e. stranding of dead marine animals). Diversity of the scavenger community was similar on beaches without dogs. Dogs increased the time it took for carcasses to be detected and decreased the proportion of carrion consumed. This ‘dog suppression effect’ on scavenging was stronger for nocturnal mammalian scavengers, presumably being driven by indirect trait-mediated effects, which raises further questions about the broader ecological consequences of domestic dogs in natural systems.

Primary and secondary host selection by Ips typographus depends on Norway spruce crown characteristics and phenolic-based defenses

Climate change is expected to intensify bark beetle population outbreaks in forests globally, affecting biodiversity and trajectories of change. Aspects of individual tree resistance remain poorly quantified, particularly with regard to the role of phenolic compounds, hindering robust predictions of forest response to future conditions. In 2003, we conducted a mechanical wounding experiment in a Norway spruce forest that coincided with an outbreak of the bark beetle, Ips typographus. We collected phloem samples from 97 trees and monitored tree survival for 5 months. Using high-performance liquid chromatography, we quantified induced changes in the concentrations of phenolics. Classification and regression tools were used to evaluate relationships between phenolic production and bark beetle resistance, in the context of other survival factors. The proximity of beetle source populations was a principal determinant of survival. Proxy measures of tree vigor, such as crown defoliation, mediated tree resistance. Controlling for these factors, synthesis of catechin was found to exponentially increase tree survival probability. However, even resistant trees were susceptible in late season due to high insect population growth. Our results show that incorporating trait-mediated effects improves predictions of survival. Using an integrated analytical approach, we demonstrate that phenolics play a direct role in tree defense to herbivory.

Parasites of spiders: Their impacts on host behavior and ecology

Parasites are some of the most abundant, diverse, and ecologically important organisms on the planet. Similarly, spiders are diverse, abundant, and play important roles in many terrestrial ecosystems. It is unfortunate that our understanding of the parasites that affect spiders is so underdeveloped relative to similar fields (e.g., parasites of insects). With this review, we describe characteristics of the major groups known to parasitize spiders and illustrate the ways in which spider biology presents unique challenges and opportunities for their parasites. Particularly promising avenues of future research include testing how parasites alter their spider hosts' behavior and ecology through density-dependent and trait-mediated effects. We close by providing future directions and testable hypotheses at the forefront of spider-parasite research.

Trait-based filtering mediates the effects of realistic biodiversity losses on ecosystem functioning

Biodiversity losses are a major driver of global changes in ecosystem functioning. While most studies of the relationship between biodiversity and ecosystem functioning have examined randomized species losses, trait-based filtering associated with species-specific vulnerability to drivers of diversity loss can strongly influence how ecosystem functioning responds to declining biodiversity. Moreover, the responses of ecosystem functioning to diversity loss may be mediated by environmental variability interacting with the suite of traits remaining in depauperate communities. We do not yet understand how communities resulting from realistic diversity losses (filtered by response traits) influence ecosystem functioning (via effect traits of the remaining community), especially under variable environmental conditions. Here, we directly test how realistic and randomized plant diversity losses influence productivity and invasion resistance across multiple years in a California grassland. Compared with communities based on randomized diversity losses, communities resulting from realistic (drought-driven) species losses had higher invasion resistance under climatic conditions that matched the trait-based filtering they experienced. However, productivity declined more with realistic than with randomized species losses across all years, regardless of climatic conditions. Functional response traits aligned with effect traits for productivity but not for invasion resistance. Our findings illustrate that the effects of biodiversity losses depend not only on the identities of lost species but also on how the traits of remaining species interact with varying environmental conditions. Understanding the consequences of biodiversity change requires studies that evaluate trait-mediated effects of species losses and incorporate the increasingly variable climatic conditions that future communities are expected to experience.

Soil engineering by ants facilitates plant compensation for large herbivore removal of aboveground biomass.

The interplay between top-down and bottom-up processes determines ecosystem productivity. Yet, the factors that mediate the balance between these opposing forces remain poorly understood. Furthering this challenge, complex and often cryptic factors like ecosystem engineering and trait-mediated interactions may play major roles in mediating the outcomes of top-down and bottom-up interactions. In semiarid grasslands of northeastern China, we conducted a large-scale, three-year experiment to evaluate how soil engineering by ants and plasticity in plants independently and jointly influenced the top-down effects of grazing by a ubiquitous herbivore (cattle) on aboveground standing biomass of the dominant perennial grass, Leymus chinensis. Herbivory had strong top-down effects, reducing L. chinensis AB by 25% relative to baseline levels without cattle or ants. In contrast, soil engineering by ants facilitated weak bottom-up effects in the absence of herbivory. However, in the presence of herbivory, soil engineering effects were strong enough to fully offset herbivore removal of aboveground biomass. This outcome was mediated by L. chinensis's plasticity in reallocating growth from below- to aboveground biomass, a result linked to additive effects of engineers and herbivores increasing soil N availability and engineering effects improving soil structure. Soil engineering increased soil N by 12%, promoting aboveground biomass. Herbivores increased soil N by 13% via defecation, but this increase failed to offset their reductions in aboveground biomass in isolation. However, when combined, engineers and herbivores increased soil N by 26% and engineers improved soil bulk density, facilitating L. chinensis to shift resource allocations from below- to aboveground biomass sufficiently to fully offset herbivore suppression of aboveground biomass. Our results demonstrate that soil engineering and trait-mediated effects of plant plasticity can strongly mediate the outcome of top-down and bottom-up interactions. These cryptic but perhaps ubiquitous processes may help to explain the long-debated phenomenon of plant compensatory responses to large grazers.

Growth and mortality of the oak processionary moth, Thaumetopoea processionea, on two oak species: direct and trait-mediated effects of host and neighbour species

The presence of heterospecific neighbours can affect the amount of damage caused by insect pests to focal plants. However, how plant neighbours influence herbivore performance is poorly understood, in particular when they are closely related. We tested the independent and interactive effects of tree species identity and tree neighbour type (conspecific vs. heterospecific) on the performance of a major oak pest, the oak processionary moth larvae (OPM, Thaumetopoea processionea L.) fed on Quercus robur L. and Q. petraea Liebl. We performed a factorial greenhouse experiment in which we grew two oak saplings per pot, either from the same species or from both species. We quantified growth and mortality of OPM larvae, leaf phenolic compounds, C:N ratio and bud phenology. OPM larvae performed significantly better on Q. petraea than on Q. robur , regardless of plant neighbour type. Phenolic compounds and C:N, but not phenology, differed between oak species and neighbour species identity. Only bud phenology had a significant effect on OPM performance, which was better when young larvae had access to recently unfolded leaves, regardless of oak species and neighbour identity. Although oak neighbour identity altered the expression of leaf traits, this effect had no measurable consequences on OPM performance. Our results showed that OPM performed better when feeding on Q. petraea than on Q. robur . We also found that oak neighbours altered the expression of leaf traits associated with defence, but that this effect did not cascade onto OPM performance. These findings should be replicated in the field in order to improve forest management strategies against OPM attacks.

Ecological traits and the spatial structure of competitive coexistence among carnivores.

Competition is a widespread interaction among carnivores, ultimately manifested through one or more dimensions of the species' ecological niche. One of the most explicit manifestations of competitive interactions regards spatial displacement. Its interpretation under a theoretical context provides an important tool to deepen our understanding of biological systems and communities, but also for wildlife management and conservation. We used Bayesian multispecies occupancy models on camera-trapping data from multiple sites in Southwestern Europe (SWE) to investigate competitive interactions within a carnivore guild, and to evaluate how species' ecological traits are shaping coexistence patterns. Seventeen out of 26 pairwise interactions departed from a hypothesis of independent occurrence, with spatial association being twice as frequent as avoidance. Association behaviors were only detected among mesocarnivores, while avoidance mainly involved mesocarnivores avoiding the apex predator (n=4) and mesocarnivore-only interactions (n=2). Body mass ratios, defined as the dominant over the subordinate species body mass, revealed an important negative effect ( ) on co-occurrence probability, and support that spatially explicit competitive interactions are mostly expressed by larger species able to dominate over smaller ones, with a threshold in body mass ratios of ~4, above which local-scale intraguild coexistence is unlikely. We found a weak relationship between pairwise trophic niche overlap and the probability of coexistence ( ), suggesting that competition for feeding resources may not be a key driver of competition, at least at the scale of our analysis. Despite local-scale avoidance, regional-scale coexistence appears to be maintained by the spatial structuring of the competitive environment. We provide evidence that SWE ecosystems consist of spatially structured competitive environments, and propose that coexistence among near-sized species is likely achieved through the interplay of "facultative" and "behavioral" character displacements. Factors influencing carnivore coexistence likely include context-dependent density and trait-mediated effects, which should be carefully considered for a sound understanding of the mechanisms regulating these communities.

Does parasitism influence sediment stability? Evaluation of trait-mediated effects of the trematode Bucephalus minimus on the key role of cockles Cerastoderma edule in sediment erosion dynamics

In coastal environments, bioturbators greatly influence the physical and biogeochemical properties of sediments with consequences for central ecological processes such as erosion dynamics. In addition to their direct impact on sediment stability, bioturbators can have an impact on sediment erosion processes by modulating the growth of microphytobenthic organisms that stabilise the surface layer of sediments. The direct and indirect influences of bioturbators on sediment dynamics depend on the magnitude of their activity and inherently on their physiological state. Bioturbators are infected by various parasites, which have a substantial impact on their physiology and behaviour. However, the knock-on effects of parasites on key ecosystem functions like sediment dynamics remain poorly studied. We conducted flume experiments to investigate the indirect influence of the trematode Bucephalus minimus parasitising the common cockle Cerastoderma edule on the dynamics of sandy sediments enriched or not with microphytobenthos (MPB). Cockles modified bed roughness, sediment surface erodibility and hence destabilised sandy sediments. In sediments not enriched with MPB, both unparasitised and parasitised organisms had a similar impact on the stability of sandy sediments. In contrast, parasitism slightly reduced the destabilisation effect of cockles in MPB-enriched sediments. In the latter, parasitised cockles did not interfere with MPB growth whereas unparasitised organisms constrained the microalgae development. However, the enrichment of the surface layers of sandy sediments with MPB did not modulate the erosion dynamics of these environments. Thus, the lower destabilisation effect of parasitised cockles was not here linked to a stabilisation effect of MPB. When standardised for length, parasitised cockles were lighter than unparasitised organisms. Weakened cockles may have had a lower bioturbation potential than unparasitised conspecifics. If so, the influence parasitised cockles had on sediment erodibility and sediment roughness may have been reduced. The absence of a parasitism effect on the dynamics of MPB-unenriched sediments remains nonetheless unclear.

Prey adaptation along a competition-defense tradeoff cryptically shifts trophic cascades from density- to trait-mediated.

Trophic cascades have become a dominant paradigm in ecology, yet considerable debate remains about the relative strength of density- (consumptive) and trait-mediated (non-consumptive) effects in trophic cascades. This debate may, in part, be resolved by considering prey experience, which shapes prey traits (through genetic and plastic change) and influences prey survival (and therefore density). Here, we investigate the cascading role of prey experience through the addition of mosquitofish (Gambusia affinis) from predator-experienced or predator-naïve sources to mesocosms containing piscivorous largemouth bass (Micropterus salmoides), zooplankton, and phytoplankton. These two sources were positioned along a competition-defense tradeoff. Results show that predator-naïve mosquitofish suffered higher depredation rates, which drove a density-mediated cascade, whereas predator-experienced mosquitofish exhibited higher survival but fed less, which drove a trait-mediated cascade. Both cascades were similar in strength, leading to indistinguishable top-down effects on lower trophic levels. Therefore, the accumulation of prey experience with predators can cryptically shift cascade mechanisms from density- to trait-mediated.

Host density and parasitoid presence interact and shape the outcome of a tritrophic interaction on seeds of wild lima bean

The interaction between the seed beetle Zabrotes subfasciatus and its parasitoid Stenocorse bruchivora, was investigated on seeds of two populations of wild lima bean, Phaseolus lunatus. By manipulating the number of beetle larvae per seed and the presence of parasitoids, we determined how factors related to beetle larvae density, the seed in which they feed and the parasitoid, may interact and affect host and parasitoid survival. Results showed that an increase in larval beetle density had a negative impact on beetle performance. This effect cascaded up to parasitoids, high larval density strongly reduced parasitoid emergence. Also, parasitoid presence resulted in faster beetle development and lower female weight. An interactive effect between larval host density and parasitoid presence affected the number of insects that emerged from the seeds. Beetle performance was better in the bean population with the largest seeds, while parasitoid emergence was the lowest in these seeds. This study shows that the impact of parasitoids on seed beetles is contingent on the interaction between density-mediated (direct mortality) and trait-mediated (e.g. non-consumptive) effects. Indirect trait-mediated effects of natural enemies are likely prevalent across insect communities, understanding their role in driving host-parasitoid interactions can have important implications for biological control.

Habitat degradation and predators have independent trait-mediated effects on prey

Coral reefs are degrading globally leading to a catastrophic loss of biodiversity. While shifts in the species composition of communities have been well documented associated with habitat change, the mechanisms that underlie change are often poorly understood. Our study experimentally examines the effects of coral degradation on the trait-mediated effects of predators on the morphology, behaviour and performance of a juvenile coral reef fish. Juvenile damselfish were exposed to predators or controls (omnivore or nothing) in seawater that had flowed over either live or dead-degraded coral over a 45d period. No interaction between water source and predator exposure was found. However, fish exposed to degraded water had larger false eyespots relative to the size of their true eyes, and were more active, both of which may lead to a survival advantage. Non-consumptive effects of predators on prey occurred regardless of water source and included longer and deeper bodies, large false eyespots that may distract predator strikes away from the vulnerable head region, and shorter latencies in their response to a simulated predator strike. Research underscores that phenotypic plasticity may assist fishes in coping with habitat degradation and promote greater resilience to habitat change than may otherwise be predicted.

Healthy but smaller herds: Predators reduce pathogen transmission in an amphibian assemblage.

Predators and pathogens are fundamental components of ecological communities that have the potential to influence each other via their interactions with victims and to initiate density- and trait-mediated effects, including trophic cascades. Despite this, experimental tests of the healthy herds hypothesis, wherein predators influence pathogen transmission, are rare. Moreover, no studies have separated effects mediated by density vs. traits. Using a semi-natural mesocosm experiment, we investigated the interactive effects of predatory dragonfly larvae (caged or lethal [free-ranging]) and a viral pathogen, ranavirus, on larval amphibians (grey treefrogs and northern leopard frogs). We determined the influence of predators on ranavirus transmission and the relative importance of density- and trait-mediated effects on observed patterns. Lethal predators reduced ranavirus infection prevalence by 57%-83% compared to no-predator and caged-predator treatments. The healthy herds effect was more strongly associated with reductions in tadpole density than behavioural responses to predators. We also assessed whether ranavirus altered the responses of tadpoles to predators. In the absence of virus, tadpoles reduced activity levels and developed deeper tails in the presence of predators. However, there was no evidence that virus presence or infection altered responses to predators. Finally, we compared the magnitude of trophic cascades initiated by individual and combined natural enemies. Lethal predators initiated a trophic cascade by reducing tadpole density, but caged predators and ranavirus did not. The absence of a virus-induced trophic cascade is ostensibly the consequence of limited virus-induced mortality and the ability of infected individuals to continue interacting within the community. Our results provide support for the healthy herds hypothesis in amphibian communities. We uniquely demonstrate that density-mediated effects of predators outweigh trait-mediated effects in driving this pattern. Moreover, this study was one of the first to directly compare trophic cascades caused by predators and pathogens. Our results underscore the importance of examining the interactions between predators and pathogens in ecology.

The demographic and life-history costs of fear: Trait-mediated effects of threat of predation on Aedes triseriatus.

Predators alter prey populations via direct lethality (density‐mediated effects), but in many taxa, the indirect nonlethal threat of predation may be almost as strong an effect, altering phenotypically plastic traits such as prey morphology, behavior, and life history (trait‐mediated effects). There are costs to antipredator defenses and the strength of prey responses to cues of predation likely depends on both the perceived level of risk and food availability.The goal of this study was to test the hypothesis that the costs of nonlethal trait‐mediated interactions impacting larvae can have carryover effects that alter life‐history traits, adult characteristics, and ultimately population dynamics.The effects of Toxorhynchites rutilus kairomones and chemical alarm cues on Aedes triseriatus were assessed in a two‐level factorial design manipulating nutrient level (low or high) and chemical cues of predation (present or absent).Nonlethal chemical cues of predation significantly decreased female survivorship and significantly decreased female size. Females emerged as adults significantly earlier when exposed to predation cues when there was high nutrient availability. When raised in the low nutrient treatment and exposed to predator cues, adult females had 2.1 times the hazard of death compared to high nutrient‐no predator cues. Females raised in the high nutrient and predator cue treatment blood fed sooner than did females from other combinations.Fear of predation can substantially alter prey life‐history traits and behavior, which can cascade into dramatic population, community, and ecosystem effects. Exposure to predator cues significantly decreased the estimated cohort rate of increase, potentially altering the expected population density of the next generation.

How Do Trait-Mediated Non-lethal Effects of Predation Affect Population-Level Performance of Mosquitoes?

Non-lethal, trait-mediated effects of predation impact prey behavior and life-history traits. Studying how these effects in turn influence prey demography is crucial to understand prey life-history evolution. Mosquitoes are important vectors that claim several million lives every year worldwide by transmitting a range of pathogens. Several ecological factors affect life-history traits of both larval and adult mosquitoes, creating effects that cascade to population-level consequences. Few studies have comprehensively explored the non-lethal effects of predation and its interactions with resources and competition on larval, adult, and population traits of mosquitoes. Understanding these interactions is important because the effects of predation are hypothesized to rescue prey populations from the effects of density-dependence resulting from larval competition. Aedes aegypti larvae reared at two different larval densities and subjected to three non-lethal predator treatments were monitored for survival, development time, and adult size through the larval stages to adult eclosion, and adult females were monitored for survival and reproduction through their first gonotrophic cycle. Intraspecific competition increased larval development time, yielded small-bodied adults, and reduced fecundity in individuals exposed to predatory chemical cues as larvae. Exposure to cues from a living predator affected both body size and latency to blood feed in females. Analysis of life-table traits revealed significant effects of competition on net reproductive rate (R 0) of mosquitoes. The interaction between competition and predator treatments significantly affected the cohort rate of increase (r) and the index of performance (r'). The index of performance, which estimates rate of population change based on the size-fecundity relationship, was significantly and positively correlated with r, but overestimated r slightly. Lack of significant effect of predator treatments and larval density on cohort generation time (T c) further suggests that the observed effects of treatments on r and r' were largely a consequence of the effects on R 0. Also, the significant effects of treatment combinations on larval development time, adult body size and fecundity were ultimately manifested as effects on life-table traits estimated from adult survival and reproduction.

Copepods drive large-scale trait-mediated effects in marine plankton.

Fear of predation may influence food webs more than actual predation. However, the mechanisms and magnitude of nonconsumptive predator effects are largely unknown in unicellular-dominated food webs such as marine plankton. We report a general mechanism of chemically induced predator effects in marine plankton. Copepods, the most abundant zooplankton in the oceans, imprint seawater with unique polar lipids-copepodamides-which trigger toxin production and bioluminescence in harmful dinoflagellates. We show that copepodamides also elicit defensive traits in other phytoplankton, inducing the harmful algal bloom-forming diatom Pseudo-nitzschia seriata to produce 10 times more toxins, and colony-forming diatoms to decrease colony size by half. A 1-year study in the northeast Atlantic revealed that natural copepodamide concentrations are high enough to induce harmful algal toxins and size reduction in dominant primary producers when copepods are abundant. We conclude that copepodamides will structure marine plankton toward smaller, more defended life forms on basin-wide scales.

Fished species uniformly reduced escape behaviors in response to protection

Predation is a critical ecological process that alters the structure and functioning of ecosystems through density-mediated and trait-mediated effects on lower trophic levels. Although studies have focused on harvest-driven reductions in abundances and sizes of targeted species, human harvest also alters species morphologies, life histories, and behaviors by selection, plasticity, and shifts in species interactions. Restricting harvest can recover the biomass of targeted species, but it is less clear how behavioral phenotypes recover, particularly relative to the impacts of potentially opposing pathways of human influence. We investigated the effects of protection on the behavioral traits of a marine fish assemblage, recording behavior of 1377 individual fishes of nine targeted kelp forest species across 16 California marine protected areas (MPAs) varying in age, protection level, and diver visitation. With long-term, full protection from harvest, all fish species exhibited shorter flight initiation distance (FID, or the distance at which an animal flees from an approaching threat) and longer time delays before fleeing, despite differences in trophic position, microhabitat use, and other ecological characteristics. These escape behaviors were amplified across new MPAs regardless of protection level, suggesting that recovery is slow and likely the result of differences in genetic or early-life experience among individuals in these long-lived species. Although the effects of full protection from harvest were partially offset by recovering populations of large piscivorous predators, the net effect of long-term, full protection on fish behavior was shorter FID. Additionally, all species had shorter FID at sites more frequently visited by divers, and this effect was greater in sites with long-term, full protection from fishing. To the extent that escape behavior is correlated with foraging behavior and predation rates, these results suggest that human-induced behavioral changes may affect ecosystem processes, even after abundances have recovered. If recovery of ecosystem functioning and services are the management goal, assessments should be broadened to include the recovery of functional traits (including behavior).