The Visually Mediated Escape Response in Fish: Predicting Prey Responsiveness and the Locomotor Behaviour of Predators and Prey

The outcome of predator-prey encounters is determined by a number of factors related to the locomotor and sensory performance of the animals. Escape responses can be triggered visually, i.e. by the magnifying retinal image of an approaching object (i.e. a predator), called the looming effect, and calculated as the rate of change of the angle subtended by the predator frontal profile as seen by the prey. A threshold of looming angle (ALT, the Apparent Looming Threshold) determines the reaction distance of a startled fish, which is proportional to the attack speed of the predator and its apparent frontal profile. Optimal tactics for predator attacks as well as consideration on their functional morphology are discussed in relation to ALT. Predator optimal attack speeds depend on predator morphology as well as the prey ALT. Predictions on the scaling of ALT suggest that ALT may increase (i.e. implying a decrease in reaction distance) with prey size in cases in which predator attack speeds are high (i.e. > 4 L/s in a 1-m long predator), while it may be relatively independent of prey size when predators attack at lower speeds. The issue of scaling of ALT is discussed using examples from field and laboratory studies. While the timing of the escape is a crucial issue for avoiding being preyed upon, the direction of escape manoeuvres may also determine the success of the escape. A simple theoretical framework for optimal escape trajectories is presented here and compared with existing data on escape trajectories of fish reacting to startling stimuli.     

Previous and Present Diets of Mite Predators Affect Antipredator Behaviour of Whitefly Prey

Predator diet is known to influence antipredator behaviour in prey. Yet, it is not clear how antipredator behaviour is affected by diet changes of the predator. We studied the effect of previous and present diet of a predatory mite Typhlodromips swirskii on the antipredator response of its prey, the whitefly Bemisia tabaci. An earlier study showed that adult female whiteflies that had experienced predators, had learned to avoid ovipositing on plants with predators whose previous and present diet consisted of whitefly eggs and immatures. Here, we investigate whether adult whiteflies also avoid plants with predators whose present and/or previous diet consisted of a non-whitefly food source. Adult whiteflies were found not to avoid plants with predators whose present diet consisted of pollen and whose previous diet had consisted of either pollen or whitefly eggs and larvae. They did avoid plants with predators whose present diet consisted of whiteflies and whose previous diet had consisted of pollen, but to a lesser extent than when previous and present diet consisted of whiteflies. In a choice experiment, whiteflies discriminated between plants with predators whose present diet consisted of whiteflies, but that differed in previous diet. Our results show that both previous and present diets of predators are important in eliciting antipredator behaviour.     

Prey Vulnerability Limits Top-Down Control and Alters Reciprocal Feedbacks in a Subsidized Model Food Web

Resource subsidies increase the productivity of recipient food webs and can affect ecosystem dynamics. Subsidies of prey often support elevated predator biomass which may intensify top-down control and reduce the flow of reciprocal subsidies into adjacent ecosystems. However, top-down control in subsidized food webs may be limited if primary consumers posses morphological or behavioral traits that limit vulnerability to predation. In forested streams, terrestrial prey support high predator biomass creating the potential for strong top-down control, however armored primary consumers often dominate the invertebrate assemblage. Using empirically based simulation models, we tested the response of stream food webs to variations in subsidy magnitude, prey vulnerability, and the presence of two top predators. While terrestrial prey inputs increased predator biomass (+12%), the presence of armored primary consumers inhibited top-down control, and diverted most aquatic energy (∼75%) into the riparian forest through aquatic insect emergence. Food webs without armored invertebrates experienced strong trophic cascades, resulting in higher algal (∼50%) and detrital (∼1600%) biomass, and reduced insect emergence (−90%). These results suggest prey vulnerability can mediate food web responses to subsidies, and that top-down control can be arrested even when predator-invulnerable consumers are uncommon (20%) regardless of the level of subsidy.     

Temporal Links in Daily Activity Patterns between Coral Reef Predators and Their Prey

Few studies have documented the activity patterns of both predators and their common prey over 24 h diel cycles. This study documents the temporal periodicity of two common resident predators of juvenile reef fishes, Cephalopholis cyanostigma (rockcod) and Pseudochromis fuscus (dottyback) and compares these to the activity and foraging pattern of a common prey species, juvenile Pomacentrus moluccensis (lemon damselfish). Detailed observations of activity in the field and using 24 h infrared video in the laboratory revealed that the two predators had very different activity patterns. C. cyanostigma was active over the whole 24 h period, with a peak in feeding strikes at dusk and increased activity at both dawn and dusk, while P. fuscus was not active at night and had its highest strike rates at midday. The activity and foraging pattern of P. moluccensis directly opposes that of C. cyanostigma with individuals reducing strike rate and intraspecific aggression at both dawn and dusk, and reducing distance from shelter and boldness at dusk only. Juveniles examined were just outside the size-selection window of P. fuscus. We suggest that the relatively predictable diel behaviour of coral reef predators results from physiological factors such as visual sensory abilities, circadian rhythmicity, variation in hunting profitability, and predation risk at different times of the day. Our study suggests that the diel periodicity of P. moluccensis behaviour may represent a response to increased predation risk at times when both the ability to efficiently capture food and visually detect predators is reduced.     

Food Web Effects of Prey Size Refugia: Variable Interactions and Alternative Stable Equilibria

Predators can have highly variable effects on the abundance and composition of food webs, ranging from strong to weak effects of top predators. Typical food web models assume that individual prey are identical in their susceptibility to predators throughout their lives, but many prey species become less vulnerable to predators through ontogeny. A simple set of models is explored where prey must pass through a vulnerable stage prior to achieving a predator-invulnerable size refuge. As productivity of the environment increases, the proportional impact of predators decreases because more individuals become and remain in the invulnerable adult stage. The addition of a competitor prey species that can not achieve size refuge results in contrasting outcomes. At low productivity, the small species wins in competition, and the system is strongly consumer controlled. At high productivity, the large species wins due to the presence of predators, and the system becomes less consumer controlled. At intermediate productivity, either the small or the large species can win depending on initial conditions, and the system can be either strongly or weakly consumer controlled. Such alternative stable equilibria derived from models with prey size refugia may help to explain many natural situations.     

Resource Management in the Orb-Weaving Spider Zygiella x-notata: I. Influence of Spider Behaviour and Available Prey

Management of resource exploitation by the orb-weaving spider Zygiella x-notata was investigated in relation to available prey. Prey capture is studied according to prey type (cricket or fly), and the spider's behaviour (waiting without prey or consuming other prey), at the moment when a prey item was placed onto the web. Characteristics of the spider's exploitation of the previous item (feeding time, and quantities ingested) were analysed according to the type and quantity of prey presented, (1 or 2 items at 1-h intervals). Capture time did not vary in relation to prey type, nor spider activity. However, duration and quantities of certain behavioural components (biting and wrapping) did vary according to prey type and the spider's behaviour, at the moment of presentation of prey. Whatever the type of prey received, if the first item consumed was a cricket, the spiders reduced time spent feeding on this prey without any decrease in the quantities ingested.     

Parasites Affect Food Web Structure Primarily through Increased Diversity and Complexity

Comparative research on food web structure has revealed generalities in trophic organization, produced simple models, and allowed assessment of robustness to species loss. These studies have mostly focused on free-living species. Recent research has suggested that inclusion of parasites alters structure. We assess whether such changes in network structure result from unique roles and traits of parasites or from changes to diversity and complexity. We analyzed seven highly resolved food webs that include metazoan parasite data. Our analyses show that adding parasites usually increases link density and connectance (simple measures of complexity), particularly when including concomitant links (links from predators to parasites of their prey). However, we clarify prior claims that parasites “dominate” food web links. Although parasites can be involved in a majority of links, in most cases classic predation links outnumber classic parasitism links. Regarding network structure, observed changes in degree distributions, 14 commonly studied metrics, and link probabilities are consistent with scale-dependent changes in structure associated with changes in diversity and complexity. Parasite and free-living species thus have similar effects on these aspects of structure. However, two changes point to unique roles of parasites. First, adding parasites and concomitant links strongly alters the frequency of most motifs of interactions among three taxa, reflecting parasites'' roles as resources for predators of their hosts, driven by trophic intimacy with their hosts. Second, compared to free-living consumers, many parasites'' feeding niches appear broader and less contiguous, which may reflect complex life cycles and small body sizes. This study provides new insights about generic versus unique impacts of parasites on food web structure, extends the generality of food web theory, gives a more rigorous framework for assessing the impact of any species on trophic organization, identifies limitations of current food web models, and provides direction for future structural and dynamical models.     

Predator Diversity Effects in an Exotic Freshwater Food Web

Cascading trophic interactions are often defined as the indirect effects of a predator on primary producers through the effect of the predator on herbivores. These effects can be both direct through removal of herbivores [density-mediated indirect interactions (DMIIs)] or indirect through changes in the behavior of the herbivores [trait-mediated indirect interactions (TMIIs)]. How the relative importance of these two indirect interactions varies with predator diversity remains poorly understood. We tested the effect of predator diversity on both TMIIs and DMIIs on phytoplankton using two competitive invasive dreissenid mussel species (zebra mussel and quagga mussel) as the herbivores and combinations of one, two or all three species of the predators pumpkinseed sunfish, round goby, and rusty crayfish. Predators had either direct access to mussels and induced both TMII and DMII, or no direct access and induced only TMII through the presence of risk cues. In both sets of treatments, the predators induced a trophic cascade which resulted in more phytoplankton remaining with predators present than with only mussels present. The trophic cascade was weaker in three-predator and two-predator treatments than in one-predator treatments when predators had direct access to dreissenids (DMIIs and TMIIs). Crayfish had higher cascading effects on phytoplankton than both pumpkinseed and round goby. Increased predator diversity decreased the strength of DMIIs but had no effect on the strength of TMIIs. The strength of TMIIs was higher with zebra than quagga mussels. Our study suggests that inter-specific interference among predators in multi-species treatments weakens the consumptive cascading effects of predation on lower trophic levels whereas the importance of predator diversity on trait mediated effects depends on predator identity.     

Landuse Change in Savannas Disproportionately Reduces Functional Diversity of Invertebrate Predators at the Highest Trophic Levels: Spiders as an Example

Predators play a disproportionately positive role in ensuring integrity of food webs, influencing ecological processes and services upon which humans rely. Predators tend to be amongst the first species to be affected by anthropogenic disturbance, however. Spiders impact invertebrate population dynamics and stabilise food webs in natural and agricultural systems (potentially mitigating against crop pests and reduced yields). Africa’s savannas are undergoing continent-wide conversion from low-density rangelands to villages and croplands, as human populations burgeon. Despite limited research, and evidence of deleterious impacts to biodiversity, African savannas are earmarked by prominent international organisations for conversion to cropland. Given the key role of spiders in food webs, they can have beneficial impacts in agroecosystems. Furthermore, functional diversity (FD) reflects ecosystem pattern and processes better than species diversity, so we evaluated impacts of large-scale landuse change on both species richness and FD. We surveyed spiders using pitfall traps at 42 sites (14 replicates each in rangeland, cropland, and villages) in South African savannas, investigating effects of landuse, season, and habitat variables on spider species diversity and FD. Species richness was lowest in villages. FD was lowest in cropland, however, with reduced representation of traits associated with hunting of larger invertebrates. Furthermore, there were fewer specialists in croplands. These findings suggest that even when cropland does not impact species diversity, loss of FD can still occur. As savanna systems transform, impacts on invertebrate population dynamics may increase the possibility of a breakdown in pest control in natural and agricultural systems, given changes in FD of invertebrate predators.     

昆明昭觉林蛙可利用食物资源季节变化

对昆明呼马山昭觉林蛙可利用食物资源研究表明:环境中可利用食物资源以节肢动物门昆虫纲和蛛形纲动物为主,其中昆虫纲主要由双翅目、直翅目、鞘翅目、同翅目、半翅目、膜翅目和鳞翅目等组成。可利用食物资源组成和数量都有着明显的季节变化。影响可利用食物资源季节变化的主要因素是气候。选择种类丰富和满足其生理需求适宜性较大的食物是昭觉林蛙利用资源的两种方式。     

Invertebrate community patterns in Mediterranean temporary wetlands along hydroperiod and salinity gradients

1. Temporary aquatic habitats often are inhabited by a unique fauna and flora and contribute significantly to regional diversity. Temporary wetlands around the world are disappearing rapidly. The individual and interacting impacts of factors influencing community structure and dynamics in temporary wetlands are not always well known.
2. Camargue wetlands are mainly characterized by variable salinity and hydroperiod. The individual and combined impacts of these local factors, together with regional variables, on invertebrate communities remain unknown. We therefore characterized and sampled invertebrates in 30 temporary wetlands along salinity and hydroperiod gradients in the Camargue (Southern France) 3, 5 and 7 months after inundation.
3. Over the three sampling occasions, a total of 17 cladoceran species and 49 macroinvertebrate taxa were identified. Hydroperiod and salinity were the most important variables explaining variation in taxonomic composition and can be considered key factors shaping the invertebrate communities in Camargue wetlands. The impact on taxon richness was significantly positive for hydroperiod but significantly negative for salinity. Regional factors had no significant effect on the structure of the studied invertebrate communities, suggesting that dispersal was not limiting and that species sorting was the most important structuring process.
4. The results of this study suggest that the combined and interacting effects of salinization and hydrological modification of Mediterranean temporary wetlands (due to water management, climate change, etc.) can result in reduced diversity in large numbers of Mediterranean wetlands and induce a considerable decline in regional diversity of aquatic invertebrates.     

Demographic Consequences of Predators on Prey: Trait and Density Mediated Effects on Mosquito Larvae in Containers

Predators may affect prey population growth and community diversity through density mediated lethal and trait mediated non-lethal effects that influence phenotypic traits of prey. We tested experimentally the roles of thinning the density of prey (lethality) in the absence of predator cues and density and trait mediated effects (lethality + intimidation) of predatory midge Corethrella appendiculata on competing native and invasive mosquito prey. Predator-mediated reductions in prey and density reductions in the absence of C. appendiculata resulted in lower percent survivorship to adulthood and estimates of the finite rate of increase (λ′) for invasive mosquito Aedes albopictus relative to that of controls. In most instances, thinning the density of prey in the absence, but not in the presence, of C. appendiculata cues resulted in lower survivorship to adulthood and λ′ for native mosquito Aedes triseriatus relative to that of controls. Together, these results suggested trait mediated effects of C. appendiculata specific to each species of mosquito prey. Release from intraspecific competition attributable to density reductions in the absence, but not in the presence, of C. appendiculata enhanced growth and lengthened adult lifespan relative to that of controls for A. albopictus but not A. triseriatus. These results show the importance of predator-mediated density and trait mediated effects on phenotypic traits and populations of invasive and native mosquitoes. Species-specific differences in the phenotypic responses of prey may be due, in part, to longer evolutionary history of C. appendiculata with A. triseriatus than A. albopictus.     

Seasonal Changes in Plankton Food Web Structure and Carbon Dioxide Flux from Southern California Reservoirs

Reservoirs around the world contribute to cycling of carbon dioxide (CO₂) with the atmosphere, but there is little information on how ecosystem processes determine the absorption or emission of CO₂. Reservoirs are the most prevalent freshwater systems in the arid southwest of North America, yet it is unclear whether they sequester or release CO₂ and therefore how water impoundment impacts global carbon cycling. We sampled three reservoirs in San Diego, California, weekly for one year. We measured seasonal variation in the abundances of bacteria, phytoplankton, and zooplankton, as well as water chemistry (pH, nutrients, ions, dissolved organic carbon [DOC]), which were used to estimate partial pressure of CO₂ (pCO₂), and CO₂ flux. We found that San Diego reservoirs are most often undersaturated with CO₂ with respect to the atmosphere and are estimated to absorb on average 3.22 mmol C m^-2 day^-1. pCO₂ was highest in the winter and lower in the summer, indicating seasonal shifts in the magnitudes of photosynthesis and respiration associated with day length, temperature and water inputs. Abundances of microbes (bacteria) peaked in the winter along with pCO₂, while phytoplankton, nutrients, zooplankton and DOC were all unrelated to pCO₂. Our data indicate that reservoirs of semi-arid environments may primarily function as carbon sinks, and that carbon flux varies seasonally but is unrelated to nutrient or DOC availability, or the abundances of phytoplankton or zooplankton.     

Prey Capture Behaviour in the Social Spider Anelosimus eximius (Araneae: Theridiidae): Responses to Prey Size and Type

Some species of web building spiders use different capture tactics for different prey types. The main factors influencing the attack behaviour are the ability of the insect to escape, the risks of injury to the spiders and prey size. This study evaluated the effects of size and prey type on prey capture behaviour of the social spider Anelosimus eximius as influenced by the number of spiders attracted by prey movements that did not bite until the immobilization (bystanders) and the number of spiders that contributed to prey immobilization (catchers). We carried out a two‐factor (prey size and type) experiment offering prey belonging to four orders: Diptera, Lepidoptera, Hymenoptera and Orthoptera, in a size gradient within each prey type. Both factors influenced the number of spiders recruited as bystanders, but only prey body size influenced the number of catchers in the subduing process. The possible advantages of the presence of bystanders around the interception site are discussed.     

The Feeding Behaviour of a Sit-and-Wait Predator,Ranatra dispar (Heteroptera: Nepidae): The Combined Effect of Food Deprivation and Prey Size on the Behavioural Components of Prey Capture

Previous work has shown a significant effect of hunger on the predatory behaviour in a sit-and-wait predator Ranatra dispar, the water stick insect (Bailey 1986 a). The experiments reported here were designed to investigate the combined effect of prey size and hunger on the predatory behaviour in order to identify which behavioural components are effected. It was found that the hunger level determines whether R. dispar will initially be aroused or not but the distance at which the arousal takes place is influenced by the size of the prey. This is believed to reflect the capacity and interrelation between visual and mechanoreceptor, sensory organs. The decision to strike at a prey is, although again influenced by hunger, significantly affected by prey size. The distance of the prey when the strike takes place is affected by hunger not the size of the prey. The outcome of the strike is determined by the size of the prey, not the hunger level of the predator. This is believed to reflect the relationship between strike trajectory, leg morphology and prey size. Food deprivation affects all components of predatory behaviour of R. dispar leading up to prey capture, by increasing not only distance of response but also the number of strikes, hits, and captures per unit presentation of prey. It does not affect capture efficiency which remains at about 70 to 80 %. Food deprivation also increases the range of prey sizes that R. dispar responds to and attempts to capture. The effect of food deprivation is considered to reflect a motivational change in responsiveness to particular prey stimuli usually described as a sensitization of particular stimulus-response relations rather than the food deprivation affecting the sensory mechanisms. The predatory success in relation to size of model prey suggested a hypothetical size that could be captured, irrespective of predator motivational level, which is based primarily on the relationship between the shape of the grasping leg and size of prey.     

Linking Morphological and Behavioural Defences: Prey Fish Detect the Morphology of Conspecifics in the Odour Signature of their Predators

Predation is a strong selective force acting on both morphology and behaviour of prey animals. While morphological defences (e.g. crypsis, presence of armours or spines or specific body morphologies) and antipredator behaviours (e.g. change in foraging or reproductive effort, or hiding and fleeing behaviours) have been widely studied separately, few studies have considered the interplay between the two. The question raised in our study is whether antipredator behaviours of a prey fish to predator odours could be influenced by the morphology of prey conspecifics in the diet of the predator. We used goldfish (Carassius auratus) as our test species; goldfish exposed to predation risk significantly increase their body depth to length ratio, which gives them a survival advantage against gape‐limited predators. We exposed shallow‐bodied and deep‐bodied goldfish to the odour of pike (Esox lucius) fed either form of goldfish. Deep‐bodied goldfish displayed lower intensity antipredator responses than shallow‐bodied ones, consistent with the hypothesis that individuals with morphological defences should exhibit less behavioural modification than those lacking such defences. Moreover, both shallow‐ and deep‐bodied goldfish displayed their strongest antipredator responses when exposed to the odour of pike fed conspecifics of their own morphology, indicating that goldfish are able to differentiate the morphology of conspecifics through predator diet cues. For a given individual, predator threat increases as the prey become more like the individual eaten, revealing a surprising level of sophistication of chemosensory assessment by prey fish.     

Nutrient Enrichment and Food Web Composition Affect Ecosystem Metabolism in an Experimental Seagrass Habitat

Background

Food web composition and resource levels can influence ecosystem properties such as productivity and elemental cycles. In particular, herbivores occupy a central place in food webs as the species richness and composition of this trophic level may simultaneously influence the transmission of resource and predator effects to higher and lower trophic levels, respectively. Yet, these interactions are poorly understood.

Methodology/Principal Findings

Using an experimental seagrass mesocosm system, we factorially manipulated water column nutrient concentrations, food chain length, and diversity of crustacean grazers to address two questions: (1) Does food web composition modulate the effects of nutrient enrichment on plant and grazer biomasses and stoichiometry? (2) Do ecosystem fluxes of dissolved oxygen and nutrients more closely reflect above-ground biomass and community structure or sediment processes? Nutrient enrichment and grazer presence generally had strong effects on biomass accumulation, stoichiometry, and ecosystem fluxes, whereas predator effects were weaker or absent. Nutrient enrichment had little effect on producer biomass or net ecosystem production but strongly increased seagrass nutrient content, ecosystem flux rates, and grazer secondary production, suggesting that enhanced production was efficiently transferred from producers to herbivores. Gross ecosystem production (oxygen evolution) correlated positively with above-ground plant biomass, whereas inorganic nutrient fluxes were unrelated to plant or grazer biomasses, suggesting dominance by sediment microbial processes. Finally, grazer richness significantly stabilized ecosystem processes, as predators decreased ecosystem production and respiration only in the zero- and one- species grazer treatments.

Conclusions/Significance

Overall, our results indicate that consumer presence and species composition strongly influence ecosystem responses to nutrient enrichment, and that increasing herbivore diversity can stabilize ecosystem flux rates in the face of perturbations.     

Predators and avian community organization: an experiment in a semi-desert grassland

Summary We provide experimental evidence that predators are a major factor organizing a community of granivorous grassland birds (mostly emberizid finches). Our focus is not on the lethal effects of predators, but on the simple idea that (i) birds will not settle where they perceive a high risk of predation, and (ii) species differ in their perception of the safety of woody vegetative cover due to differences in antipredator escape behavior. Consistent with this idea is the fact that the composition of this grassland community responds markedly to minor manipulations in the distribution of woody cover. In particular, with the addition of cover to open grasslands, species with cover-dependent escape tactics increase in abundance, while the abundance of cover-independent species decreases greatly; this decrease may reflect aggression from cover-dependent species, but evidence suggests that some cover-independent species may actively avoid cover-rich areas per se. Non-predatory effects of cover, most notably those concerning food resources and microclimate, appear unable to explain these results. Predators may influence many communities of terrestrial vertebrates via species-specific responses to cover.