Trait-mediated Effects Research Articles

Parasite susceptibility in an amphibian host is modified by salinization and predators.

Secondary salinization represents a global threat to freshwater ecosystems. Salts, such as NaCl, can be toxic to freshwater organisms and may also modify the outcome of species interactions (e.g. host-parasite interactions). In nature, hosts and their parasites are embedded in complex communities where they face anthropogenic and biotic (i.e. predators) stressors that influence host-parasite interactions. As human populations grow, considering how anthropogenic and natural stressors interact to shape host-parasite interactions will become increasingly important. We conducted two experiments investigating: (1) the effects of NaCl on tadpole susceptibility to trematodes and (2) whether density- and trait-mediated effects of a parasite-predator (i.e. damselfly) and a host-predator (i.e. dragonfly), respectively, modify the effects of NaCl on susceptibility to trematode infection. In the first experiment, we exposed tadpoles to three concentrations of NaCl and measured parasite infection in tadpoles. In the second experiment, we conducted a 2 (tadpoles exposed to 0 g L-1 NaCl vs. 1 g L-1 NaCl) x 4 (no predator, free-ranging parasite-predator (damselfly), non-lethal host-predator (dragonfly kairomone), and free-ranging parasite-predator+dragonfly kairomone) factorial experiment. In the absence of predators, exposure to NaCl increased parasite infection. Of the predator treatments, NaCl only caused an increase in parasite infection in the presence of the parasite-predator. However, direct consumption of trematodes caused a reduction in overall infection in the parasite-predator treatment. In the dragonfly kairomone treatment, a reduction in tadpole movement (i.e. trematode avoidance behavior) led to an increase in overall infection. In the parasite-predator+dragonfly kairomone treatment, antagonistic effects of the parasite-predator (reduction in trematode abundance) and dragonfly kairomone (reduction in parasite avoidance behavior) resulted in intermediate parasite infection. Collectively, these findings demonstrate that NaCl can increase amphibian susceptibility to parasites, and underscores the importance of considering predator-mediated interactions in understanding how contaminants influence host-parasite interactions.

Cattle Metabolic Fingerprinting to Predict Acute Stress Response Post-Wolf Encounters

Author(s): Valerio, Azzurra; Wielgus, Robert; Valerio, Mariacristina; Casadei, Luca | Abstract: As wolves recolonize across their former range in western North America, encounters between livestock and wolves are expected to increase in frequency. Understanding the physiological state of the prey, as a response to stress imposed by the presence of a predator (trait-mediated effects), will help with predicting the total effect of predators on their prey beyond direct consumption (density-mediated effects). Fecal glucocorticoid (GCM) is widely used to measure stress response, but provides inconclusive results, particularly when applied to a finer spatio-temporal variation in predation risk (i.e., wolf-livestock encounters). Since the impact of external stressors (i.e., wolf encounters) on the body influences ultimately the physical and metabolic state of the animal, we purpose to investigate: 1) if the fecal metabolome extracted from cattle fecal samples reflects changes in the GCM levels, and 2) if the cattle metabolome can better predict stress response than GCM post-wolf encounter. We first performed a controlled stress experiment on five captive cows in a pasture. We also conducted a field study in Washington where we fit GPS collars equipped with proximity sensors on two wolves in two separate packs, and on 40 range cows in four different livestock herds. When a wolf and a cow equipped with proximity sensors are within 128 meters from one another, the GPS collars send a real-time message that allows us to assess the physiological state of the prey at a finer spatial-temporal scale. In both studies, we collected fecal samples from GPS radio-collared cattle and compared their metabolome to GCM levels. Our results clearly show that the metabolomic analysis of the cattle gut microbiome can better predict acute stress response than the GCM, in both the controlled and free-ranging environment.

Seagrass-bivalve facilitative interactions: Trait-mediated effects along an environmental gradient

Facilitative interactions are important forces in shaping community structure and function, and understanding how they respond to environmental changes has become an increasing concern in ecology. Lucinid bivalves play a significant role in seagrass meadows, through a mutualism in which the seagrass provides habitat and oxygen via the roots, while the bivalves and their associated bacteria eliminate sulfides from pore water, improving thus plant performance. In this study, we evaluated how this mutualism is modified along a gradient of organic matter content in the sediment, in a coastal bay dominated by Cymodocea nodosa meadows. We used a correlative approach, seeking statistical association between sediment organic matter content, lucinid abundance, and plant traits. Lucinid abundance was higher in vegetated that in bare areas. In vegetated areas, lucinid abundance decreased as organic matter content in the sediment increased, decrease seemingly associated to modification in plant traits, i.e. root abundance and morphology. In organic-rich sediments, roots are less abundant and less branched, reducing the potential habitat for lucinids and suggesting a weakening of the interaction. This finding contributes to our understanding of how facilitative interactions can be modified along human disturbance gradients, and how disturbances can reduce resilience of seagrasses through this modification.

Rapid evolution of hosts begets species diversity at the cost of intraspecific diversity

Ecosystems are complex food webs in which multiple species interact and ecological and evolutionary processes continuously shape populations and communities. Previous studies on eco-evolutionary dynamics have shown that the presence of intraspecific diversity affects community structure and function, and that eco-evolutionary feedback dynamics can be an important driver for its maintenance. Within communities, feedbacks are, however, often indirect, and they can feed back over many generations. Here, we studied eco-evolutionary feedbacks in evolving communities over many generations and compared two-species systems (virus-host and prey-predator) with a more complex three-species system (virus-host-predator). Both indirect density- and trait-mediated effects drove the dynamics in the complex system, where host-virus coevolution facilitated coexistence of predator and virus, and where coexistence, in return, lowered intraspecific diversity of the host population. Furthermore, ecological and evolutionary dynamics were significantly altered in the three-species system compared with the two-species systems. We found that the predator slowed host-virus coevolution in the complex system and that the virus' effect on the overall population dynamics was negligible when the three species coexisted. Overall, we show that a detailed understanding of the mechanism driving eco-evolutionary feedback dynamics is necessary for explaining trait and species diversity in communities, even in communities with only three species.

Experimental Assessment in Nature of the Ecological Effects of a Specialist Parasite

Parasites can have important effects on the structure and composition of natural biological communities, either directly by influencing host fecundity and survival (i.e., density-mediated effects) or indirectly by influencing host traits such as behavior, life-history, morphology, and physiology (i.e., trait-mediated effects). Yet few studies have explored how these effects play out in the wild, as opposed to simplified and controlled laboratory or mesocosm settings. We addressed this information gap by translocating Gyrodactylus ectoparasites of the Trinidadian guppy into previously Gyrodactylus-free guppy populations in two rivers (Marianne and Paria) in the Northern Mountain Range in Trinidad. We then measured phenotypic and demographic changes in the guppy host and its competitor, Rivulus hartii, and compared these changes to guppies and Rivulus in control reaches just upstream. In the Marianne, where guppies invest more in reproduction (i.e., greater size and number of embryos), the introduction of G...

Bottom-up and top-down effects of tree species diversity on leaf insect herbivory.

The diversity of plant neighbors commonly results in direct, bottom‐up effects on herbivore ability to locate their host, and in indirect effects on herbivores involving changes in plant traits and a top‐down control by their enemies. Yet, the relative contribution of bottom‐up and top‐down forces remains poorly understood. We also lack knowledge on the effect of abiotic constraints such as summer drought on the strength and direction of these effects. We measured leaf damage on pedunculate oak (Quercus robur), alone or associated with birch, pine or both in a long‐term tree diversity experiment (ORPHEE), where half of the plots were irrigated while the other half remained without irrigation and received only rainfall. We tested three mechanisms likely to explain the effects of oak neighbors on herbivory: (1) Direct bottom‐up effects of heterospecific neighbors on oak accessibility to herbivores, (2) indirect bottom‐up effects of neighbors on the expression of leaf traits, and (3) top‐down control of herbivores by predators. Insect herbivory increased during the growth season but was independent of neighbor identity and irrigation. Specific leaf area, leaf toughness, and thickness varied with neighbor identity while leaf dry matter content or C:N ratio did not. When summarized in a principal component analysis (PCA), neighbor identity explained 87% of variability in leaf traits. PCA axes partially predicted herbivory. Despite greater rates of attack on dummy caterpillars in irrigated plots, avian predation, and insect herbivory remained unrelated. Our study suggests that neighbor identity can indirectly influence insect herbivory in mixed forests by modifying leaf traits. However, we found only partial evidence for these trait‐mediated effects and suggest that more attention should be paid to some unmeasured plant traits such as secondary metabolites, including volatile organic compounds, to better anticipate the effects of climate change on plant‐insect interactions in the future.

Bottom‐up and trait‐mediated effects of resource quality on amphibian parasitism

Leaf litter subsidies are important resources for aquatic consumers like tadpoles and snails, causing bottom-up effects on wetland ecosystems. Recent studies have shown that variation in litter nutritional quality can be as important as litter quantity in driving these bottom-up effects. Resource subsidies likely also have indirect and trait-mediated effects on predation and parasitism, but these potential effects remain largely unexplored. We generated predictions for differential effects of litter nutrition and secondary polyphenolic compounds on tadpole (Lithobates sylvatica) exposure and susceptibility to Ribeiroia ondatrae, based on ecological stoichiometry and community-ecology theory. We predicted direct and indirect effects on key traits of the tadpole host (rates of growth, development and survival), the trematode parasite (production of the cercaria infective stages) and the parasite's snail intermediate host (growth and reproduction). To test these predictions, we conducted a large-scale mesocosm experiment using a natural gradient in the concentrations of nutrients (nitrogen) and toxic secondary compounds (polyphenolics) of nine leaf litter species. To differentiate between effects on exposure vs. susceptibility to infection, we included multiple infection experiments including one with constant per capita exposure. We found that increased litter nitrogen increased tadpole survival, and also increased cercaria production by the snail intermediate hosts, causing opposing effects on tadpole per capita exposure to trematode infection. Increased litter polyphenolics slowed tadpole development, leading to increased infection by increasing both their susceptibility to infection and the length of time they were exposed to parasites. Based on these results, recent shifts in forest composition towards more nitrogen-poor litter species should decrease trematode infection in tadpoles via density- and trait-mediated effects on the snail intermediate hosts. However, these shifts also involve increased abundance of litter species with high polyphenolic levels, which should increase trematode infection via trait-mediated effects on tadpoles. Future studies will be needed to determine the relative strength of these opposing effects in natural wetland communities. [Correction added after online publication on 5 January 2017: wording changed to 'which should increase trematode infection via trait-mediated effects on tadpoles'.].

Feeding guild of non-host community members affects host-foraging efficiency of a parasitic wasp.

Interactions between predator and prey, or parasitoid and host, are shaped by trait- and density-mediated processes involving other community members. Parasitoids that lay their eggs in herbivorous insects locate their hosts through infochemicals such as herbivore-induced plant volatiles (HIPVs) and host-produced kairomones. Hosts are frequently accompanied by non-host herbivores that are unsuitable for the parasitoid. These non-hosts may interfere with host location primarily through trait-mediated processes, by their own infochemicals, and their induction of the emission of plant volatiles. Although it is known that single non-hosts can interfere with parasitoid host location, it is still unknown whether the observed effects are due to species specific characteristics or to the feeding habits of the non-host herbivores. Here we addressed whether the feeding guild of non-host herbivores differentially affects foraging of the parasitoid Cotesia glomerata for its common host, caterpillars of Pieris brassicae feeding on Brassica oleracea plants. We used different phloem-feeding and leaf-chewing non-hosts to study their effects on host location by the parasitoid when searching for host-infested plants based on HIPVs and when searching for hosts on the plant using infochemicals. To evaluate the ultimate effect of these two phases in host location, we studied parasitism efficiency of parasitoids in small plant communities under field-tent conditions. We show that leaf-chewing non-hosts primarily affected host location through trait-mediated effects via plant volatiles, whereas phloem-feeding non-hosts exerted trait-mediated effects by affecting foraging efficiency of the parasitoid on the plant. These trait-mediated effects resulted in associational susceptibility of hosts in environments with phloem feeders and associational resistance in environments with non-host leaf chewers.

Persistent and context-dependent effects of the larval feeding environment on post-metamorphic performance through the adult stage

One of the central issues in ecology is the identification of processes affecting the population structure and dynamics of species with complex life cycles. In such species, variation in both the number of larvae that enter a population and their phenotype are important drivers of survival and growth after metamorphosis. Larval experience can have strong effects on key postmetamorphic traits, but the temporal scale of such 'trait-mediated effects' may be short, and their magnitude may depend on the environment experienced after metamorphosis. We used an intertidal barnacle to study the long-term consequences of trait-mediated effects under different postmetamorphic conditions by manipulating larval food concentration and monitoring patterns of survival and growth in juveniles at 2 intertidal levels over a 5 mo period. In 2 replicated experiments, higher food levels resulted in increased body size, mass and reserves (measured from elemental composition) in the settling larval stage and increased body size of newly metamorphosed juveniles. In Expt 1, high food concentration reduced juvenile mortality at low intertidal levels, while on the upper intertidal, mortality was high for all larval food concentrations. By contrast, in Expt 2, low larval food concentration decreased juvenile survival at both shore levels. When present, effects were established early (Weeks 1 or 2) and persisted for over 10 wk in Expt 1 and 22 wk in Expt 2. Interactive effects of the larval and juvenile environments can have important implications for population size: trait-mediated effects may persist for long periods, helping to explain patterns of adult abundance. (Less)

Disease Ecology Meets Ecosystem Science

Growing evidence indicates that parasites—when considered—can play influential roles in ecosystem structure and function, highlighting the need to integrate disease ecology and ecosystem science. To strengthen links between these traditionally disparate fields, we identified mechanisms through which parasites can affect ecosystems and used empirical literature searches to explore how commonly such mechanisms have been documented, the ecosystem properties affected, and the types of ecosystems in which they occur. Our results indicate that ecosystem-disease research has remained consistently rare, comprising less than 2% of disease ecology publications. Existing studies from terrestrial, freshwater, and marine systems, however, demonstrate that parasites can strongly affect (1) biogeochemical cycles of water, carbon, nutrients, and trace elements, (2) fluxes of biomass and energy, and (3) temporal ecosystem dynamics including disturbance, succession, and stability. Mechanistically, most studies have demonstrated density-mediated indirect effects, rather than trait-mediated effects, or direct effects of parasites, although whether this is representative remains unclear. Looking forward, we highlight the importance of applying traits-based approaches to predict when parasites are most likely to exert ecosystem-level effects. Future research should include efforts to extend host–parasite studies across levels of ecological organization, large-scale manipulations to experimentally quantify ecosystem roles of parasites, and the integration of parasites and disease into models of ecosystem functioning.

Natural and experimental tests of trophic cascades: gray wolves and white-tailed deer in a Great Lakes forest.

Herbivores can be major drivers of environmental change, altering plant community structure and changing biodiversity through the amount and species of plants consumed. If natural predators can reduce herbivore numbers and/or alter herbivore foraging behavior, then predators may reduce herbivory on sensitive plants, and a trophic cascade will emerge. We have investigated whether gray wolves (Canis lupus) generate such trophic cascades by reducing white-tailed deer (Odocoileus virginianus) herbivory on saplings and rare forbs in a northern mesic forest (Land O' Lakes, WI). Our investigation used an experimental system of deer exclosures in areas of high and low wolf use that allowed us to examine the role that wolf predation may play in reducing deer herbivory through direct reduction in deer numbers or indirectly through changing deer behavior. We found that in areas of high wolf use, deer were 62 % less dense, visit duration was reduced by 82 %, and percentage of time spent foraging was reduced by 43 %; in addition, the proportion of saplings browsed was nearly sevenfold less. Average maple (Acer spp.) sapling height and forb species richness increased 137 and 117 % in areas of high versus low wolf use, respectively. The results of the exclosure experiments revealed that the negative impacts of deer on sapling growth and forb species richness became negligible in high wolf use areas. We conclude that wolves are likely generating trophic cascades which benefit maples and rare forbs through trait-mediated effects on deer herbivory, not through direct predation kills.

Time‐scale dependency of host plant biomass‐ and trait‐mediated indirect effects of deer herbivory on a swallowtail butterfly

Despite recent attempts to quantify the relative strength of density- and trait-mediated indirect effects, rarely has the issue been properly addressed at the population level. Most research is based on short-term small-scale experiments in which behavioural and/or physiological responses prevail. Here, we estimated the time-scales during which density- and trait-mediated effects manifest, as well as the strength of these effects, using an interaction chain with three organisms (deer-plant-butterfly). A hierarchical Bayesian model was performed by using a long-term data set of deer density in the Boso Peninsula, central Japan (where local densities differ spatially and temporally) as well as densities of the swallowtail butterfly Byasa alcinous and its host plant Aristolochia kaempferi. The time-scale effect of deer on plant quantity and quality was estimated according to the degree of carry-over effects. The negative influence on leaf density showed a temporal saturation pattern over the long term, while the positive influence on leaf quality due to resprouting of leaves after deer browsing showed no clear temporal trend. The net indirect effect changed from positive to negative with time, with the negative density-mediated effect becoming prominent in the long term. Our novel approach is widely applicable in assessing the dynamic impacts of wildlife if the spatio-temporal variability of expansion and/or invasion history is known.

Non‐consumptive effects of a top‐predator decrease the strength of the trophic cascade in a four‐level terrestrial food web

The fear of predators can strongly impact food web dynamics and ecosystem functioning through effects on herbivores morphology, physiology or behaviour. While non‐consumptive predator effects have been mostly studied in three‐level food chains, we lack evidence for the propagation of non‐consumptive indirect effects of apex predators in four level food‐webs, notably in terrestrial ecosystems. In experimental mesocosms, we manipulated a four‐level food chain including top‐predator cues (snakes), mesopredators (lizards), herbivores (crickets), and primary producers (plants). The strength of the trophic cascade induced by mesopredators through the consumption of herbivores decreased in the presence of top‐predator cues. Specifically, primary production was higher in mesocosms where mesopredators were present relative to mesocosms with herbivores only, and this difference was reduced in presence of top‐predator cues, probably through a trait‐mediated effect on lizard foraging. Our study demonstrates that non‐consumptive effects of predation risk can cascade down to affect both herbivores and plants in a four‐level terrestrial food chain and emphasises the need to quantify the importance of such indirect effects in natural communities.

Local and landscape management drive trait-mediated biodiversity of nine taxa on small grassland fragments

Aim Biodiversity across the globe is heavily eroded by intensified management at local and landscape scales. Species communities of calcareous grasslands, which are among Europe's most diverse habitats, are severely threatened by the cessation of appropriate traditional management, loss of habitat connectivity and simplification of the surrounding landscape. However, our understanding of these often interrelated factors remains limited, in particular for trait-mediated responses across taxa. Here, we test the independent effects of local management (grazing, mowing and abandonment), habitat connectivity (measured by a connectivity index) and landscape complexity (indicated by the percentage of arable land) on nine taxa: plants, butterflies, bees, grasshoppers, hoverflies, spiders, true bugs, rove beetles and leafhoppers on small semi-natural calcareous grassland remnants (< 1 ha). Location Central Germany. Methods We use a joint analysis across taxa to identify general and trait-mediated responses (body size and Red List status) in species richness, abundance and community composition. Results We identified three key drivers of local diversity patterns: First, an increasing proportion of arable land from 10% to 80% led to a 29% loss of overall species richness. Second, despite differences between taxa, increasing habitat connectivity generally enhanced species richness. Connectivity effects were more accentuated in the large species per taxon, which can be expected to be good dispersers. Finally, grazing reduced species richness and abundance much more than annual mowing or short-term abandonment (5–15 years), in particular for red-listed species. We attribute this to plant resource removal through overgrazing and trampling. Main conclusions For the conservation management of small calcareous grasslands, we advocate an alternating strategy of mowing or lenient grazing and short-term abandonment, prioritizing connected fragments surrounded by diverse landscapes. Despite taxon-specific responses, our study across nine taxa demonstrates universal, trait-mediated effects of management, landscape complexity and connectivity on local biodiversity in fragmented communities.

Predator diversity, intraguild predation, and indirect effects drive parasite transmission

Humans are altering biodiversity globally and infectious diseases are on the rise; thus, there is interest in understanding how changes to biodiversity affect disease. Here, we explore how predator diversity shapes parasite transmission. In a mesocosm experiment that manipulated predator (larval dragonflies and damselflies) density and diversity, non-intraguild (non-IG) predators that only consume free-living cercariae (parasitic trematodes) reduced metacercarial infections in tadpoles, whereas intraguild (IG) predators that consume both parasites and tadpole hosts did not. This likely occurred because IG predators reduced tadpole densities and anticercarial behaviors, increasing per capita exposure rates of the surviving tadpoles (i.e., via density- and trait-mediated effects) despite the consumption of parasites. A mathematical model demonstrated that non-IG predators reduce macroparasite infections, but IG predation weakens this "dilution effect" and can even amplify parasite burdens. Consistent with the experiment and model, a wetland survey revealed that the diversity of IG predators was unrelated to metacercarial burdens in amphibians, but the diversity of non-IG predators was negatively correlated with infections. These results are strikingly similar to generalities that have emerged from the predator diversity-pest biocontrol literature, suggesting that there may be general mechanisms for pest control and that biocontrol research might inform disease management and vice versa. In summary, we identified a general trait of predators--where they fall on an IG predation continuum--that predicts their ability to reduce infections and possibly pests in general. Consequently, managing assemblages of predators represents an underused tool for the management of human and wildlife diseases and pest populations.

Complex dynamics of a predator–prey–parasite system: An interplay among infection rate, predator's reproductive gain and preference

Parasites are considered as an important factor in regulating their host populations through trait-mediated effects. On the other hand, predation becomes particularly interesting in host–parasite systems because predation can significantly alter the abundance of parasites and their host population. The combined effects of parasites and predator on host population and community structure therefore may have larger effect. Different field experiments confirm that predators consume disproportionately large number of infected prey in comparison to their susceptible counterpart. There are also substantial evidences that predator has the ability to distinguish prey that have been infected by a parasite and avoid such prey to reduce fitness cost. In this paper we study the predator–prey dynamics, where the prey species is infected by some parasites and predators consume both the susceptible and infected prey with some preference. We demonstrate that complexity in such systems largely depends on the predator's selectivity, force of infection and predator's reproductive gain. If the force of infection and predator's reproductive gain are low, parasites and predators both go to extinction whatever be the predator's preference. The story may be totally different in the opposite case. Survival of species in stable, oscillatory or chaotic states, and their extinction largely depend on the predator's preference. The system may also show two coexistence equilibrium points for some parameter values. The equilibrium with lower susceptible prey density is always stable and the equilibrium with higher susceptible prey density is always unstable. These results suggest that understanding the consequences of predator's selectivity or preference may be crucial for community structure involving parasites.

Trait‐mediated effects between predators: ant chemical cues induce spider dispersal

AbstractPredators can induce changes in prey phenotype such as dispersal, activity and foraging rate. Such trait‐mediated effects (TMEs) can strongly affect prey populations and generate trophic cascades, rivaling the importance of predation in communities. However, the relevance of TME on intraguild interactions has rarely been addressed. Ants and spiders are widespread generalist predators in terrestrial habitats. Ants influence arthropod assemblages and disrupt top‐down effects of spiders on herbivores by killing spiders and/or by inducing spider emigration. Here, we examined whether ants induce dispersal behaviour in spiders. We tested the effect of chemical cues of two ant species (Lasius niger, Formica clara) on the walking activity and the propensity for silk‐based dispersal of spiders. Silk‐based dispersal of the web‐builder Phylloneta impressa increased by 80% with exposure to Lasius cues, whereas dispersal of the hunting spider Xysticus more than doubled when confronted with cues of both Lasius and Formica. In addition, Xysticus individuals showed a marked increase in walking activity when exposed to Formica but not Lasius cues. Our results show for the first time that perceived predation risk influences spider dispersal. The strong effect of ant chemical cues on spider dispersal demonstrates that TMEs contribute to the impact of ants on arthropod communities.

Parasites that change predator or prey behaviour can have keystone effects on community composition

Parasites play pivotal roles in structuring communities, often via indirect interactions with non-host species. These effects can be density-mediated (through mortality) or trait-mediated (behavioural, physiological and developmental), and may be crucial to population interactions, including biological invasions. For instance, parasitism can alter intraguild predation (IGP) between native and invasive crustaceans, reversing invasion outcomes. Here, we use mathematical models to examine how parasite-induced trait changes influence the population dynamics of hosts that interact via IGP. We show that trait-mediated indirect interactions impart keystone effects, promoting or inhibiting host coexistence. Parasites can thus have strong ecological impacts, even if they have negligible virulence, underscoring the need to consider trait-mediated effects when predicting effects of parasites on community structure in general and biological invasions in particular.

Cascading effects of predator–detritivore interactions depend on environmental context in a Tibetan alpine meadow

Studies of grazing food webs show that species traits can interact with environmental factors to determine the strength of trophic cascades, but analogous context dependencies in detrital food webs remain poorly understood. In predator-detritivore-plant interaction chains, predators are expected to indirectly suppress plant biomass by reducing the density of plant-facilitating detritivores. However, this outcome can be reversed where above-ground predators drive burrowing detritivores to lower soil levels, strengthening their plant-facilitating effects. Here, we show that these trait-mediated indirect interactions further depend on environmental context in a Tibetan alpine meadow. In our study system, undulating topography generates higher (dry soil) patches interspersed with lower (wet soil) patches. Because the ability of detritivores to form deep burrows is likely to be limited by oxygen availability in low patches (wet soil), we hypothesized that (i) burrowing detritivores would undergo a vertical habitat shift, allowing them to more effectively avoid predation, in high - but not low - patches, and (ii) this shift would transmit positive effects of predators to plants in high patches by improving conditions in the lower soil layer. We tested these hypotheses using complementary field and glasshouse experiments examining whether the cascading effects of above-ground predatory beetles (presence/absence) on the density and behaviour of tunnel-forming detritivorous beetles, soil properties, and plant growth varied with patch type (low/high). Results revealed that predatory beetles did not reduce the density of detritivores in either patch type but had context-dependent trait-mediated effects, increasing the tunnelling depth of detritivores, improving soil conditions and ultimately increasing plant biomass in the high but not low patches. This study adds to an emerging predictive framework linking predators to plants in detritus food webs, demonstrating that these indirect interactions depend not just on the relative habitat domains of predators and prey, but also on environmental conditions that can predictably constrain the behavioural response of detritivores to predation risk.

Predator cue studies reveal strong trait-mediated effects in communities despite variation in experimental designs

Nonconsumptive or trait-mediated effects of predators on their prey often outweigh density-mediated interactions where predators consume prey. For instance, predator presence can alter prey behaviour, physiology, morphology and/or development. Despite a burgeoning literature, our ability to identify general patterns in prey behavioural responses may be influenced by the inconsistent methodologies of predator cue experiments used to assess trait-mediated effects. We therefore conducted a meta-analysis to highlight variables (e.g. water type, predator husbandry, exposure time) that may influence invertebrate prey’s behavioural responses to fish predator cues. This revealed that changes in prey activity and refuge use were remarkably consistent overall, despite wide differences in experimental methodologies. Our meta-analysis shows that invertebrates altered their behaviour to predator cues of both fish that were fed the focal invertebrate and those that were fed other prey types, which suggests that invertebrates were not responding to specific diet information in the fish cues. Invertebrates also altered their behaviour regardless of predator cue addition regimes and fish satiation levels. Cue intensity and exposure time did not have significant effects on invertebrate behaviour. We also highlight that potentially confounding factors, such as parasitism, were rarely recorded in sufficient detail to assess the