Spatial and temporal changes in group dynamics and range use enable anti-predator responses in African buffalo

The reintroduction of large predators provides a framework to investigate responses by prey species to predators. Considerable research has been directed at the impact that reintroduced wolves (Canis lupus) have on cervids, and to a lesser degree, bovids, in northern temperate regions. Generally, these impacts alter feeding, activity, and ranging behavior, or combinations of these. However, there are few studies on the response of African bovids to reintroduced predators, and thus, there is limited data to compare responses by tropical and temperate ungulates to predator reintroductions. Using the reintroduction of lion (Panthera leo) into the Addo Elephant National Park (AENP) Main Camp Section, South Africa, we show that Cape buffalo (Syncerus caffer) responses differ from northern temperate ungulates. Following lion reintroduction, buffalo herds amalgamated into larger, more defendable units; this corresponded with an increase in the survival of juvenile buffalo. Current habitat preference of buffalo breeding herds is for open habitats, especially during the night and morning, when lion are active. The increase in group size and habitat preference countered initial high levels of predation on juvenile buffalo, resulting in a return in the proportion of juveniles in breeding herds to pre-lion levels. Our results show that buffalo responses to reintroduced large predators in southern Africa differ to those of northern temperate bovids or cervids in the face of wolf predation. We predict that the nature of the prey response to predator reintroduction is likely to reflect the trade-off between the predator selection and hunting strategy of predators against the life history and foraging strategies of each prey species.     

Does the risk of encountering lions influence African herbivore behaviour at waterholes?

A central question in the study of predator–prey relationships is to what extent prey behaviour is determined by avoidance of predators. Here, we test whether the long-term risk of encountering lions and the presence of lions in the vicinity influence the behaviour of large African herbivores at waterholes through avoidance of high-risk areas, increases in group size, changes in temporal niche or changes in the time spent in waterhole areas. In Hwange National Park, Zimbabwe, we monitored waterholes to study the behaviour of nine herbivore species under different risks of encountering lions. We radio-collared 26 lions in the study area which provided the opportunity to monitor whether lions were present during observation sessions and to map longer-term seasonal landscapes of risk of encountering lions. Our results show that the preferred prey species for lions (buffalo, kudu and giraffe) avoided risky waterholes. Group size increased as encounter risk increased for only two species (wildebeest and zebra), but this effect was not strong. Interestingly, buffalo avoided the hours of the day which are dangerous when the long-term and short-term risks of encountering lions were high, and all species showed avoidance of waterhole use at night times when lions were in the vicinity. This illustrates well how prey can make temporal adjustments to avoid dangerous periods coinciding with predator hunting. Additionally, many herbivores spent more time accessing water to drink when the long- and short-term risks of encountering lions were high, and they showed longer potential drinking time when the long-term risk of encountering lions was high, suggesting higher levels of vigilance. This study illustrates the diversity of behavioural adjustments to the risk of encountering a predator and how prey respond differently to temporal variations in this risk.     

Validity of the Prey‐Trap Hypothesis for Carnivore‐Ungulate Interactions at Wildlife‐Crossing Structures

Abstract: Wildlife‐exclusion fencing and wildlife‐crossing structures (e.g., underpasses and overpasses) are becoming increasingly common features of highway projects around the world. The prey‐trap hypothesis posits that predators exploit crossing structures to detect and capture prey. The hypothesis predicts that predation events occur closer to a highway after the construction of fences and crossing structures and that prey species’ use of crossings increases the probability that predators will attack prey. We examined interactions between ungulates and large carnivores at 28 wildlife crossing structures along 45 km of the Trans‐Canada Highway in Banff National Park, Alberta. We obtained long‐term records of locations where ungulates were killed (kill sites) before and after crossing structures were built. We also placed remote, motion‐triggered cameras at two crossing structures to monitor predator behavior following ungulate passage through the structure. The proximity of ungulate kill sites to the highway was similar before and after construction of fencing and crossing structures. We found only five kill sites near crossing structures after more than 32,000 visits over 13 years. We found no evidence that predator behavior at crossing structures is affected by prey movement. Our results suggest that interactions between large mammals and their prey at wildlife‐crossing structures in Banff National Park are not explained by the prey‐trap hypothesis.     

Can rare positive interactions become common when large carnivores consume livestock?

Livestock populations in protected areas are viewed negatively because of their interaction with native ungulates through direct competition for food resources. However, livestock and native prey can also interact indirectly through their shared predator. Indirect interactions between two prey species occur when one prey modifies either the functional or numerical responses of a shared predator. This interaction is often manifested as negative effects (apparent competition) on one or both prey species through increased predation risk. But indirect interactions can also yield positive effects on a focal prey if the shared predator modifies its functional response toward increased consumption of an abundant and higher-quality alternative prey. Such a phenomenon between two prey species is underappreciated and overlooked in nature. Positive indirect effects can be expected to occur in livestock-dominated wildlife reserves containing large carnivores. We searched for such positive effects in Acacia-Zizhypus forests of India's Gir sanctuary where livestock (Bubalus bubalis and Bos indicus) and a coexisting native prey (chital deer, Axis axis) are consumed by Asiatic lions (Panthera leo persica). Chital vigilance was higher in areas with low livestock density than in areas with high livestock density. This positive indirect effect occurred because lion predation rates on livestock were twice as great where livestock were abundant than where livestock density was low. Positive indirect interactions mediated by shared predators may be more common than generally thought with rather major consequences for ecological understanding and conservation. We encourage further studies to understand outcomes of indirect interactions on long-term predator-prey dynamics in livestock-dominated protected areas.     

Antipredator responses of koomal (Trichosurus vulpecula hypoleucus) against introduced and native predators

Antipredator behavior studies generally assess prey responses to single predator species although most real systems contain multiple species. In multi-predator environments prey ideally use antipredator responses that are effective against all predator species, although responses may only be effective against one predator and counterproductive for another. Multi-predator systems may also include introduced predators that the prey did not co-evolve with, so the prey may either fail to recognize their threat (level 1 naiveté), use ineffective responses (level 2 naiveté) or succumb to their superior hunting ability (level 3 naiveté). We analyzed microhabitat selection of an Australian marsupial (koomal, Trichosurus vulpecula hypoleucus) when faced with spatiotemporal differences in the activity/density levels of one native (chuditch, Dasyurus geoffroii) and two introduced predators (red fox, Vulpes vulpes; feral cat, Felis catus). From this, we inferred whether koomal recognized introduced predators as a threat, and whether they minimized predation risk by either staying close to trees and/or using open or dense microhabitats. Koomal remained close to escape trees regardless of the predator species present, or activity/density levels, suggesting koomal employ this behavior as a first line of defense. Koomal shifted to dense cover only under high risk scenarios (i.e., with multiple predator species present at high densities). When predation risk was low, koomal used open microhabitats, which likely provided benefits not associated with predator avoidance. Koomal did not exhibit level 1 naiveté, although further studies are required to determine if they exhibit higher levels of naiveté (2–3) against foxes and cats.     

Modelling predation by transient leopard seals for an ecosystem-based management of Southern Ocean fisheries

Correctly quantifying the impacts of rare apex marine predators is essential to ecosystem-based approaches to fisheries management, where harvesting must be sustainable for targeted species and their dependent predators. This requires modelling the uncertainty in such processes as predator life history, seasonal abundance and movement, size-based predation, energetic requirements, and prey vulnerability. We combined these uncertainties to evaluate the predatory impact of transient leopard seals on a community of mesopredators (seals and penguins) and their prey at South Georgia, and assess the implications for an ecosystem-based management. The mesopredators are highly dependent on Antarctic krill and icefish, which are targeted by regional fisheries. We used a state-space formulation to combine (1) a mark-recapture open-population model and individual identification data to assess seasonally variable leopard seal arrival and departure dates, numbers, and residency times; (2) a size-based bioenergetic model; and (3) a size-based prey choice model from a diet analysis. Our models indicated that prey choice and consumption reflected seasonal changes in leopard seal population size and structure, size-selective predation and prey vulnerability. A population of 104 (90–125) leopard seals, of which 64% were juveniles, consumed less than 2% of the Antarctic fur seal pup production of the area (50% of total ingested energy, IE), but ca. 12–16% of the local gentoo penguin population (20% IE). Antarctic krill (28% IE) were the only observed food of leopard seal pups and supplemented the diet of older individuals. Direct impacts on krill and fish were negligible, but the “escapement” due to leopard seal predation on fur seal pups and penguins could be significant for the mackerel icefish fishery at South Georgia. These results suggest that: (1) rare apex predators like leopard seals may control, and may depend on, populations of mesopredators dependent on prey species targeted by fisheries; and (2) predatory impacts and community control may vary throughout the predator's geographic range, and differ across ecosystems and management areas, depending on the seasonal abundance of the prey and the predator's dispersal movements. This understanding is important to integrate the predator needs as natural mortality of its prey in models to set prey catch limits for fisheries. Reliable estimates of the variability of these needs are essential for a precautionary interpretation in the context of an ecosystem-based management.     

Recognising one’s enemies: a functional approach to risk assessment by prey

Little has been done to compare the relative importance of various mechanisms through which prey assess the potential risk from natural enemies. We used predator-naive spider mites (Tetranychus urticae, Tetranychidae) to (1) compare the responses of prey to chemical cues from enemy and non-enemy species and (2) investigate the source of these cues. In the laboratory, we observed the distribution of T. urticae in response to cues from nine mite species, including (1) predators of spider mites, (2) predators/parasites of other animals, and (3) fungivores/pollen-feeders. When given a choice over 24 h, spider mites foraged and oviposited in fewer numbers on leaf discs that were previously exposed to predatory or parasitic mites (including species incapable of attacking spider mites) than on clean leaf discs (unexposed to mites). Interestingly, previous exposure of leaf arenas to fungivores and pollen-feeders had no significant effect on spider mite distribution. We then observed the response of T. urticae to cues from two species of predator that had been reared on a diet of either spider mites or pollen. T. urticae showed stronger avoidance of leaf discs that were previously exposed to spider-mite-fed predators than of discs exposed to pollen-fed predators. Nevertheless, for one predator species (Amblyseius andersoni), T. urticae still preferred to forage and oviposit on clean (unexposed) discs than on discs exposed to pollen-fed predators. Protein-derived metabolic wastes of predatory or parasitic mites may provide a general cue about potential predation risk for T. urticae. However, T. urticae also avoided areas exposed to pollen-fed predators, suggesting there may be other sources of enemy recognition by the spider mites. We discuss the ecological and evolutionary mechanisms that may influence the scope of information through which animals assess predation risk. Received: 11 January 1999 / Received in revised form: 25 October 1999 / Accepted: 20 November 1999     

Trade-offs between maternal foraging and fawn predation risk in an income breeder

The choice of neonatal hiding place is critical for ungulates adopting hiding anti-predator strategies, but the consequences of different decisions have rarely been evaluated with respect to offspring survival. First, we investigated how landscape-scale choices made by roe deer fawns and their mothers affected predation risk by red foxes in a forest–farmland mosaic in southeastern Norway. After, we examined the effect of site-specific characteristics and behaviour (i.e. visibility, mother–fawn distance and abundance of the predator’s main prey item—small rodents) on predation risk. The study of habitat use, selection and habitat-specific mortality revealed that roe deer utilised the landscape matrix in a functional way, with different habitats used for feeding, providing maternal care and as refugia from predation. Mothers faced a trade-off between foraging and offspring survival. At the landscape-scale decisions were primarily determined by maternal energetic constraints and only secondarily by risk avoidance. Indeed, forage-rich habitats were strongly selected notwithstanding the exceptionally high densities of rodents which increased fawn predation. At fine spatial scales, a high visibility of the mother was the major factor determining predation risk; however, mothers adjusted their behaviour to the level of risk at the bed site to minimise predation. Fawns selected both landscape-scale refugia and concealed bed sites, but failure to segregate from the main prey of red foxes led to higher predation. This study provides evidence for the occurrence of spatial heterogeneity in predation risk and shows that energetically stressed individuals can tackle the foraging-safety trade-off by adopting scale-dependent anti-predator responses.     

Patch use and vigilance by sympatric lemmings in predator and competitor-driven landscapes of fear

Prey living in risky environments can adopt a variety of behavioral tactics to reduce predation risk. In systems where predators regulate prey abundance, it is reasonable to assume that differential patterns of habitat use by prey species represent adaptive responses to spatial variation in predation. However, patterns of habitat use also reflect interspecific competition over habitat. Collared (Dicrostonyx groenlandicus) and brown (Lemmus trimucronatus) lemmings represent such a system and possess distinct upland tundra versus mesic meadow habitat preferences consistent with interspecific competition. Yet, we do not know whether this habitat preference might also reflect differences in predation risk or whether the two species differ in their behavioral tactics used to avoid predation. We performed experiments where we manipulated putative predation risk perceived by lemmings by increasing protective cover in upland and meadow habitats while we recorded lemming activity and behavior. Both lemming species preferentially used cover more than open patches, but Dicrostonyx was more vigilant than Lemmus. Both species also constrained their activity to protective patches in upland and meadow habitats, but during different periods of the day. Use of cover and vigilance were independent of habitat, suggesting that both species live in a fearsome but flattened landscape of fear at Walker Bay (Nunavut, Canada), and that their habitat preference is a consequence of competition rather than predation risk. Future studies aiming to map the contours of fear in multi-prey–predator systems should consider how predation and competition interact to modify prey species’ habitat preference, patch use, and vigilance.     

Habitat structure affects intraguild predation

Intraguild predation is thought to be ubiquitous in natural food webs. Yet, theory on intraguild predation predicts the intraguild prey to persist only under limited conditions. This gap between theory and empirical observations needs scrutiny. One reason might be that theory has focused on equilibrium dynamics and a limited set of species (usually three) that interact in well-mixed populations in unstructured habitats, and these assumptions will often not hold in natural systems. In this review, we focus on the effects of habitat structure on intraguild predation. Habitat structure could reduce encounter rates between predators and prey and could create refuges for prey. In both cases, habitat structure could reduce the strength of intraguild interactions, thereby facilitating species coexistence. A meta-analysis of studies on manipulation of habitat structure shows that intraguild prey indeed suffer less from intraguild predation in structured habitats. This was further confirmed by a meta-analysis in which studies on intraguild predation were classified according to habitat structure. Intraguild predation reduced densities of the intraguild prey significantly more in habitats with little structure than in habitats rich in structure. The effect of intraguild predation on the shared prey was negative, and not significantly affected by habitat structure. We conclude that habitat structure may increase persistence of the intraguild prey by decreasing the strength of the interaction between intraguild predator and intraguild prey.     

Antipredator responses to invasive lionfish, Pterois volitans: interspecific differences in cue utilization by two coral reef gobies

Understanding prey response to predators and their utilization of sensory cues to assess local predation risk is crucial in determining how predator avoidance strategies affect population demographics. This study examined the antipredator behaviors of two ecologically similar species of Caribbean coral reef fish, Coryphopterus glaucofraenum and Gnatholepis thompsoni, and characterized their responses to different reef predators. In laboratory assays, the two species of gobies were exposed to predator visual cues (native Nassau grouper predator vs. invasive lionfish predator), damage-released chemical cues from gobies, and combinations of these, along with appropriate controls. Behavioral responses indicate that the two prey species differ in their utilization of visual and chemical cues. Visual cues from predators were decisive for both species’ responses, demonstrating their relative importance in the sensory hierarchy, whereas damage-released cues were a source of information only for C. glaucofraenum. Both prey species could distinguish between native and invasive predators and subsequently altered their antipredator responses.     

The anti-predator function of ‘eyespots’ on camouflaged and conspicuous prey

Animals utilise various strategies to reduce the risk of predation, including camouflage, warning colours and mimicry, and many of these protective signals promote avoidance behaviour in predators. For example, various species possess paired circular ‘eyespots’, which startle or intimidate predators, preventing or halting an attack. However, little is known of how the efficacy of such signals relates to the context in which they are found, and no studies have tested the relative effectiveness of anti-predator signals when on otherwise camouflaged and conspicuous prey. We find that the protective value of conspicuous wing spots, placed on artificial moth-like targets presented to wild birds in the field, is strongly affected by the attributes of the prey ‘animal’ on which they are found. Wing spots reduced predation when on conspicuous prey but were rendered ineffective when on otherwise camouflaged targets, indeed they increased the risk of predation compared to non-marked camouflaged controls. These results demonstrate how different anti-predator strategies may interact, and that protective signals can switch from being beneficial to costly under different contexts.     

Cryptic biodiversity effects: importance of functional redundancy revealed through addition of food web complexity

Interactions between predators and the degree of functional redundancy among multiple predator species may determine whether herbivores experience increased or decreased predation risk. Specialist parasites can modify predator behavior, yet rarely have cascading effects on multiple predator species and prey been evaluated. We examined influences of specialist phorid parasites (Pseudacteon spp.) on three predatory ant species and herbivores in a coffee agroecosystem. Specifically, we examined whether changes in ant richness affected fruit damage by the coffee berry borer (Hypothenemus hampei) and whether phorids altered multi-predator effects. Each ant species reduced borer damage, and without phorids, increasing predator richness did not further decrease borer damage. However, with phorids, activity of one ant species was reduced, indicating that the presence of multiple ant species was necessary to limit borer damage. In addition, phorid presence revealed synergistic effects of multiple ant species, not observed without the presence of this parasite. Thus, a trait-mediated cascade resulting from a parasite-induced predator behavioral change revealed the importance of functional redundancy, predator diversity, and food web complexity for control of this important pest.     

Novel features of an inducible defense system in larval tree frogs (Hyla chrysoscelis)

Organisms in aquatic ecosystems must often tolerate variable environmental conditions, including an uncertain risk of predation. Individuals that can maintain plastic defenses against predation will increase their survival when predators are present, but will not incur the costs of these defenses when the risk of predation is low and the defense is not induced. Larvae of the pond-breeding anuran Hyla chrysoscelis develop a conspicuous phenotype in the presence of predators consisting of a brightly colored tail and a deeper tail fin. In this study, I attempted to identify the source of the chemical signal that induces this defensive morphology in this species. I tested whether metabolites alone, originating from the prey but passing through the predator, were able to induce the same morphological response as the combination of alarm signals released directly by attacked conspecifics, and metabolites. I used morphometric and tail conspicuousness data to assess tadpole response to the perceived risk of predation by larval odonate predators (Anax junius). I also tested whether this inducing cue could be recognized across species by measuring the morphological response of H. chrysoscelis tadpoles exposed to cues emitted when tadpoles of a closely related genus (Pseudacris crucifer) were consumed. Tadpoles exhibited a clean graded response of both overall shape and tail morphology in response to all cues, corresponding to their relative reliability as indicators of a risk of predation. H. chrysoscelis tadpoles were also able to respond to cues emitted when tadpoles of a closely related genus were consumed by predators. These results illustrate that tadpoles of this species are able to respond to metabolites alone without alarm signals, and that interspecific chemical communication is a primary mechanism for predator avoidance in this inducible defense system.     

Savanna tree density, herbivores, and the herbaceous community: bottom-up vs. top-down effects

Herbivores choose their habitats both to maximize forage intake and to minimize their risk of predation. For African savanna herbivores, the available habitats range in woody cover from open areas with few trees to dense, almost-closed woodlands. This variation in woody cover or density can have a number of consequences for herbaceous species composition, cover, and productivity, as well as for ease of predator detection and avoidance. Here, we consider two alternative possibilities: first, that tree density affects the herbaceous vegetation, with concomitant "bottom-up" effects on herbivore habitat preferences; or, second, that tree density affects predator visibility, mediating "top-down" effects of predators on herbivore habitat preferences. We sampled sites spanning a 10-fold range of tree densities in an Acacia drepanolobium-dominated savanna in Laikipia, Kenya, for variation in (1) herbaceous cover, composition, and species richness; (2) wild and domestic herbivore use; and (3) degree of visibility obstruction by the tree layer. We then used structural equation modeling to consider the potential influences that tree density may have on herbivores and herbaceous community properties. Tree density was associated with substantial variation in herbaceous species composition and richness. Cattle exhibited a fairly uniform use of the landscape, whereas wild herbivores, with the exception of elephants, exhibited a strong preference for areas of low tree density. Model results suggest that this was not a response to variation in herbaceous-community characteristics, but rather a response to the greater visibility associated with more open places. Elephants, in contrast, preferred areas with higher densities of trees, apparently because of greater forage availability. These results suggest that, for all but the largest species, top-down behavioral effects of predator avoidance on herbivores are mediated by tree density. This, in turn, appears to have cascading effects on the herbaceous vegetation. These results shed light on one of the major features of the "landscape of fear" in which African savanna herbivores exist.     

Different escape tactics of two vole species affect the success of the hunting predator, the least weasel

In the ongoing evolutionary arms race between predators and their prey, successful escape from the predator leads to the evolution of improved escape tactics in prey, but also predators become more effective in following and attacking the prey. Antipredatory behavior of prey is considered to be the strongest towards their most dangerous predators. However, prey species can differ both in vulnerability and efficiency of escape to a shared predator. We studied escape reactions of two vole species, the bank vole (Myodes glareolus) and the field vole (Microtus agrestis), under a simulated predation risk of the least weasel (Mustela nivalis nivalis). We conducted a laboratory experiment where a vole was given a possibility to escape from a weasel by fleeing to a horizontal tunnel or climbing the tree. Subsequently to the vole escape decision, we released a weasel to the same tunnel system to test how the weasel succeeded in following the vole. Weasel presence changed the behavior of voles as especially bank voles escaped by climbing. Instead, the majority of field voles fled into the ground-layer tunnel. The different escape tactics of the voles affected the success of the weasel, because climbing voles were less often successfully followed. We suggest that the difference in escape tactics has evolved as an adaptation to different habitats; meadow-exploiting field voles using ground-level escape while bank voles living in three-dimensional forest habitat frequently use arboreal escape tactics. This is likely to lead to different habitat-dependent vulnerabilities to predation in Microtus and Myodes vole species.     

Food webs and intraguild predation: community interactions of a native mesocarnivore

Trophic level interactions between predators create complex relationships such as intraguild predation. Theoretical research has predicted two possible paths to stability in intraguild systems: intermediate predators either outcompete higher-order predators for shared resources or select habitat based on security. The effects of intraguild predation on intermediate mammalian predators such as swift foxes (Vulpes velox) are not well understood. We examined the relationships between swift foxes and both their predators and prey, as well the effect of vegetation structure on swift fox-coyote (Canis latrans) interactions, between August 2001 and August 2004. In a natural experiment created by the Pinon Canyon Maneuver Site in southeastern Colorado, USA, we documented swift fox survival and density in a variety of landscapes and compared these parameters in relation to prey availability, coyote abundance, and vegetation structure. Swift fox density varied significantly between study sites, while survival did not. Coyote abundance was positively related to the basal prey species and vegetation structure, while swift fox density was negatively related to coyote abundance, basal prey species, and vegetation structure. Our results support the prediction that, under intraguild predation in terrestrial systems, top predator distribution matches resource availability (resource match), while intermediate predator distribution inversely matches predation risk (safety match). While predation by coyotes may be the specific cause of swift fox mortality in this system, the more general mechanism appears to be exposure to predation moderated by shrub density.     

Prey risk allocation in a grazing ecosystem.

Understanding the behaviorally mediated indirect effects of predators in ecosystems requires knowledge of predator-prey behavioral interactions. In predator-ungulate-plant systems, empirical research quantifying how predators affect ungulate group sizes and distribution, in the context of other influential variables, is particularly needed. The risk allocation hypothesis proposes that prey behavioral responses to predation risk depend on background frequencies of exposure to risk, and it can be used to make predictions about predator-ungulate-plant interactions. We determined non-predation variables that affect elk (Cervus elaphus) group sizes and distribution on a winter range in the Greater Yellowstone Ecosystem (GYE) using logistic and log-linear regression on surveys of 513 1-km2 areas conducted over two years. Employing model selection techniques, we evaluated risk allocation and other a priori hypotheses of elk group size and distributional responses to wolf (Canis lupus) predation risk while accounting for influential non-wolf-predation variables. We found little evidence that wolves affect elk group sizes, which were strongly influenced by habitat type and hunting by humans. Following predictions from the risk allocation hypothesis, wolves likely created a more dynamic elk distribution in areas that they frequently hunted, as elk tended to move following wolf encounters in those areas. This response should dilute elk foraging pressure on plant communities in areas where they are frequently hunted by wolves. We predict that this should decrease the spatial heterogeneity of elk impacts on grasslands in areas that wolves frequently hunt. We also predict that this should decrease browsing pressure on heavily browsed woody plant stands in certain areas, which is supported by recent research in the GYE.     

Predator hunting mode and habitat domain alter nonconsumptive effects in predator-prey interactions

Predators can affect prey populations through changes in traits that reduce predation risk. These trait changes (nonconsumptive effects, NCEs) can be energetically costly and cause reduced prey activity, growth, fecundity, and survival. The strength of nonconsumptive effects may vary with two functional characteristics of predators: hunting mode (actively hunting, sit-and-pursue, sit-and-wait) and habitat domain (the ability to pursue prey via relocation in space; can be narrow or broad). Specifically, cues from fairly stationary sit-and-wait and sit-and-pursue predators should be more indicative of imminent predation risk, and thereby evoke stronger NCEs, compared to cues from widely ranging actively hunting predators. Using a meta-analysis of 193 published papers, we found that cues from sit-and-pursue predators evoked stronger NCEs than cues from actively hunting predators. Predator habitat domain was less indicative of NCE strength, perhaps because habitat domain provides less reliable information regarding imminent risk to prey than does predator hunting mode. Given the importance of NCEs in determining the dynamics of prey communities, our findings suggest that predator characteristics may be used to predict how changing predator communities translate into changes in prey. Such knowledge may prove particularly useful given rates of local predator change due to habitat fragmentation and the introduction of novel predators.     

Trait-mediated effects on flowers: artificial spiders deceive pollinators and decrease plant fitness

Although predators can affect foraging behaviors of floral visitors, rarely is it known if these top-down effects of predators may cascade to plant fitness through trait-mediated interactions. In this study we manipulated artificial crab spiders on flowers of Rubus rosifolius to test the effects of predation risk on flower-visiting insects and strength of trait-mediated indirect effects to plant fitness. In addition, we tested which predator traits (e.g., forelimbs, abdomen) are recognized and avoided by pollinators. Total visitation rate was higher for control flowers than for flowers with an artificial crab spider. In addition, flowers with a sphere (simulating a spider abdomen) were more frequently visited than those with forelimbs or the entire spider model. Furthermore, the presence of artificial spiders decreased individual seed set by 42% and fruit biomass by 50%. Our findings indicate that pollinators, mostly bees, recognize and avoid flowers with predation risk; forelimbs seem to be the predator trait recognized and avoided by hymenopterans. Additionally, predator avoidance by pollinators resulted in pollen limitation, thereby affecting some components of plant fitness (fruit biomass and seed number). Because most pollinator species that recognized predation risk visited many other plant species, trait-mediated indirect effects of spiders cascading down to plant fitness may be a common phenomenon in the Atlantic rainforest ecosystem.