Asexual reproduction changes predator population dynamics in a life predator–prey system

Many organisms display oscillations in population size. Theory predicts that these fluctuations can be generated by predator–prey interactions, and empirical studies using life model systems, such as a rotifer-algae community consisting of Brachionus calyciflorus as predator and Chlorella vulgaris as prey, have been successfully used for studying such dynamics. B. calyciflorus is a cyclical parthenogen (CP) and clones often differ in their sexual propensity, that is, the degree to which they engage into sexual or asexual (clonal) reproduction. Since sexual propensities can affect growth rates and population sizes, we hypothesized that this might also affect population oscillations. Here, we studied the dynamical behaviour of B. calyciflorus clones representing either CPs (regularly inducing sex) or obligate parthenogens (OPs). We found that the amplitudes of population cycles to be increased in OPs at low nutrient levels. Several other population dynamic parameters seemed unaffected. This suggests that reproductive mode might be an important additional variable to be considered in future studies of population oscillations.     

What is an apex predator?

Large ‘apex’ predators influence ecosystems in profound ways, by limiting the density of their prey and controlling smaller ‘mesopredators’. The loss of apex predators from much of their range has lead to a global outbreak of mesopredators, a process known as ‘mesopredator release’ that increases predation pressure and diminishes biodiversity. While the classifications apex‐ and meso‐predator are fundamental to current ecological thinking, their definition has remained ambiguous. Trophic cascades theory has shown the importance of predation as a limit to population size for a variety of taxa (top–down control). The largest of predators however are unlikely to be limited in this fashion, and their densities are commonly assumed to be determined by the availability of their prey (bottom–up control). However, bottom–up regulation of apex predators is contradicted by many studies, particularly of non‐hunted populations. We offer an alternative view that apex predators are distinguishable by a capacity to limit their own population densities (self‐regulation). We tested this idea using a set of life‐history traits that could contribute to self‐regulation in the Carnivora, and found that an upper limit body mass of 34 kg (corresponding with an average mass of 13–16 kg) marks a transition between extrinsically‐ and self‐regulated carnivores. Small carnivores share fast reproductive rates and development and higher densities. Large carnivores share slow reproductive rates and development, extended parental care, sparsely populated territories, and a propensity towards infanticide, reproductive suppression, alloparental care and cooperative hunting. We discuss how the expression of traits that contribute to self‐regulation (e.g. reproductive suppression) depends on social stability, and highlight the importance of studying predator–prey dynamics in the absence of predator persecution. Self‐regulation in large carnivores may ensure that the largest and the fiercest do not overexploit their resources.     

Effects of the probability of a predator catching prey on predator–prey system stability

To understand the effect of the probability of a predator catching prey, P_catch, on the stability of the predator–prey system, a spatially explicit lattice model consisting of predators, prey, and grass was constructed. The predators and prey randomly move on the lattice space, and the grass grows according to its growth probability. When a predator encounters prey, the predator eats the prey in accordance with the probability P_catch. When a prey encounters grass, the prey eats the grass. The predator and prey give birth to offspring according to a birth probability after eating prey or grass, respectively. When a predator or prey is initially introduced or newly born, its health state is set at a high given value. This health state decreases by one with every time step. When the state of an animal decreases to less than zero, the individual dies and is removed from the system. Population densities for predator and prey fluctuated significantly according to P_catch. System stability was characterized by the standard deviation ? of the fluctuation. The simulation results showed that ? for predators increased with an increase of P_catch; ? for prey reached a maximum at P_catch = 0.4; and ? for grass fluctuated little regardless of P_catch. These results were due to the tradeoff between P_catch and the predator–prey encounter rate, which represents the degree of interaction between predator and prey and the average population density, respectively.     

Geometric optimization for prey–predator strategies

This paper investigates several strategies for prey and predator in both bounded and unbounded domains, assuming they have the same speed. The work describes how the prey should move to escape from the predator and how predator should move to catch the prey. The approach is agent-based and explicitly tracks movement of individuals as prey and predator. We show that the prey escapes one or two competing predators, while might be caught in the case of three predators. The paper also describes a strategy for finding a well camouflaged static prey which emits signals.     

Diet choice and predator—prey dynamics

Summary We compare the dynamics of predator-prey systems with specialist predators or adaptive generalist predators that base diet choice on energy-maximizing criteria. Adaptive predator behaviour leads to functional responses that are influenced by the relative abundance of alternate prey. This results in the per capita predation risk being positively density-dependent near points of diet expansion. For a small set of parameter values, systems with adaptive predators can be locally stable whereas systems with specialist predators would be unstable. This occurs mainly when alternate prey have low enough profitability that predators cannot sustain themselves indefinitely when feeding on alternate prey. Local stability of systems with adaptive predator behaviour is inversely related to the goodness of fit to optimal diet choice criteria. Hence, typical patterns of partial prey preference are more stabilizing than perfect optimal diet selection. Locally stable systems with adaptive predators are often globally unstable, converging on limit cycles for many initial population densities. The small range of parameter combinations and initial population densities leading to stable equilibria suggest that adaptive diet selection is unlikely to be a ubiquitous stabilizing factor in trophic interactions.     

Fading out of vole and predator cycles?

Northern voles and lemmings are famous for their spectacular multiannual population cycles with high amplitudes. Such cyclic vole populations in Scandinavia have shown an unexpected and marked long-term decline in density since the early 1970s, particularly with a marked shift to lower spring densities in the early 1980s. The vole decline, mainly characterized by a strongly decreased rate of change in numbers over winter, is associated with an increased occurrence of mild and wet winters brought about by a recent change in the North Atlantic Oscillation. This has led to a decrease in winter stability and has shortened the period with protective snow cover, the latter considered as an important prerequisite for the occurrence of multiannual, high-amplitude cycles in vole populations. Although the vole decline is predicted to be negative for predators' reproduction and abundance, empirical data showing this are rare. Here we show that the dynamics of a predator-prey system (Tengmalm's owl, Aegolius funereus, and voles), have in recent years gradually changed from 3-4 yr, high-amplitude cycles towards more or less annual fluctuations only.     

Viable populations in a prey–predator system

 Lotka–Volterra equations are considered a dynamical game, where the phenotypes of the predator and of the prey can vary. This differs from the usual procedure of specifying as a priori laws according to which strategies are supposed to change. The question at stake is the survival of each of the species, instead of the maximization of a given pay-off by each player, as it is commonly discussed in games. The predator needs the prey, while the prey can survive without the predator. These obvious and simplistic constraints are enough to shape the regulation of the system: notably, the largest closed set of initial conditions can be delineated, from which there exists at least one evolutionary path where the population can avoid extinction forever. To these so-called viable trajectories, viable strategies are associated, respectively for the prey or for the predator. A coexistence set can then be defined. Within this set and outside the boundary, strategies can vary arbitrarily within given bounds while remaining viable, whereas on the boundary, only specific strategies can guarantee the viability of the system. Thus, the largest set can be determined, outside of which strategies will never be flexible enough to avoid extinction. Received 2 May 1995; received in revised form 15 August 1995     

A nonlocal kinetic model for predator–prey interactions

We extend the aggregation model from Fetecau (2011) by adding a field of vision to individuals and by including a second species. The two species, assumed to have a predator–prey relationship, have dynamics governed by nonlocal kinetic equations that include advection and turning. The latter is the main mechanism for aggregation and orientation, which results from interactions among individuals of the same species as well as predator–prey relationships. We illustrate numerically a diverse set of predator–prey behaviors that can be captured by this model. We show that a prey’s escape outcome depends on the social interactions between its group members, the prey’s field of vision and the sophistication of the predator’s hunting strategies.     

Red queen dynamics in specific predator–prey systems

The dynamics of a predator–prey system are studied, with a comparison of discrete and continuous strategy spaces. For a \(2 \times 2\) system, the average strategies used in the discrete and continuous case are shown to be the same. It is further shown that the inclusion of constant prey switching in the discrete case can have a stabilising effect and reduce the number of available predator types through extinction.     

Are multiple predator effects directed by prey availability?

Antagonistic/synergistic interactions among predators foraging on the same prey have been assumed to play a major role in shaping community structure. Studies in systems with multiple predator species have shown that the strength of these interactions may not be predictable and is largely dependent on individual behavioural traits, species density and habitat complexity. Although the association of prey consumption and satiation of a foraging predator has long been recognized, there has been relatively little research on how prey availability affects multiple predators’ effects. In this work, we present a framework to investigate the variation in two coexisting/competing predators’ effects on prey risk as affected by the prey availability rate. Functional responses by each predator species were first studied in single-predator treatments. Then, the intra- and inter-specific competition was investigated by employing additive and substitutative experimental designs to highlight the nature of multiple effects. Intra- and interspecific interactions were found to be similar and there was risk reduction, and risk enhancement for the prey at intermediate and high levels, respectively, according to the multiplicative risk model (MRM). The results indicated that when similar predators are concerned, the outcomes of MRM may vary according to the functional response curve of these predators. Thus, studies involving a wide range of prey densities are required to explore the nature of interactions. Moreover, this kind of experimental data can contribute to unravelling complexities in theoretical approaches by earlier studies and ultimately promote understanding the effect of multiple predators on prey population regulation.     

Predation by a water mite (Piona exigua) on enclosed populations of zooplankton

Short term, replicated experimental alteration of densities of a predatory water mite Piona exigua Viets, inside 0.45–0.475 m³ litre enclosures, revealed little evidence of the effects of predation on the number and relative abundance of the enclosed zooplankton species. Predation rates more closely approximated those estimated from single prey functional response experiments in the second experimental period (December) than in the first (March). In December Daphnia was the only susceptible taxon present in large numbers, whereas in March, Ceriodaphnia and Chydorus were also present. This result is consistent with laboratory findings that predation rates are lowered in the presence of more than one prey type.The difficulty of obtaining evidence for significant effects of these planktonic predators is in part due to changes in the preferred prey species in the diet of Piona depending on stage and sex of the mite and to aspects of experimental design. The wide variability between replicate enclosures at each predator density reduced the power of the statistical analyses used to test the null hypothesis. Enclosures with no predators are necessary to investigate the effects of enclosure on the zooplankton prey, since these effects may outweigh those due to predator consumption.     

Evaluating predation pressure on green treefrog larvae across a habitat gradient

The effect of a predator on the abundance of a prey species depends upon the predators abundance and its ability to capture that prey. The objectives of this research were to evaluate the community structure of predators of green treefrog (Hyla cinerea) tadpoles across habitat types and evaluate the effectiveness of individual predators on H. cinerea tadpoles. Correspondence and cluster analyses of predator frequencies across 23 aquatic habitats indicated that the majority of variance in predator communities was due to a division between permanent and temporary habitats. Experimental work demonstrated that survival of the smallest H. cinerea tadpoles was significantly lower than survival of medium and large tadpoles with the most effective predators, indicating that H. cinerea tadpoles attain a refuge from predation at larger body sizes. We combined the effectiveness of predators in experiments with the abundance of each predator species from the predator community survey to demonstrate that predation pressure on H. cinerea tadpoles is higher in temporary ponds. This pattern may explain in part why this species generally breeds successfully only in permanent habitats. It also confirms that discussions about an increasing gradient of predation pressure from temporary to permanent aquatic habitats should be restricted to individual prey species for which such a gradient has been demonstrated.     

Multiple intraguild predators reduce mortality risk of a mutual agricultural pest prey in simple,but not in complex,experimental settings

Multiple predator species that coexist with each other and their mutual prey can have combined effects on prey mortality that are similar to the sum of each predator's individual impact (linear effects), greater than the sum of each predator's individual impact (risk enhancement), or less than the sum of each predator's individual impact (risk reduction). Understanding multiple predator effects is important to determine the impact of predators on pest prey in agroecosystems. If two predators share the same broad spatial domain and hunting mode and engage in intraguild predation, then their combination is expected to result in risk reduction for a mutual prey. We tested this hypothesis using both additive and replacement experimental designs on two species of generalist wolf spider predators (Tasmanicosa leuckartii and Hogna crispipes) that hunt in the same domain, and a mutual insect prey (cotton bollworm Helicoverpa armigera). We used two types of enclosures: a small simple laboratory enclosure, and a larger more complex cotton plant enclosure. We found that in the small simple laboratory enclosures, the presence of two spiders led to risk reduction of Helicoverpa larva mortality as expected, but in larger more complex cotton plant enclosures the presence of both species resulted in linear effects rather than risk reduction on Helicoverpa mortality. Furthermore, intraguild predation did not change multiple predator effects in laboratory or plant enclosures. This study has implications for managing arthropod predators in agroecosystems; contrary to predictions of ecological frameworks, coexistence of predators that share the same hunting mode and hunting domain may not lead to risk reduction on a mutual prey in more complex environments, where encounters among predators can be lower. Conservation of multiple predators of a single guild can play an essential role on biological control of insect pests.     

Trophic cascades in rocky shore tide pools: distinguishing lethal and nonlethal effects

The effects of predators on the density of their prey can have positive indirect effects on the abundance of the preys resource via a trophic cascade. This concept has strongly influenced contemporary views of how communities are structured. However, predators also can transmit indirect effects by inducing changes in prey traits. We show that the mere presence of predator risk cues can initiate a trophic cascade in rocky shore tide pools. In large (mean surface area =9 m²), natural tide pools, we manipulated crab density and their foraging ability to examine the relative importance of lethal (density-mediated) and non-lethal (trait-mediated) predator effects to algal community development. We found that perceived predation risk reduced snail density as much as the direct predation treatment, showing that green crab predation was not an important factor regulating local snail density. Instead, snail emigration away from resident crabs appears to be the most important factor regulating local snail density. As a result, the abundance of ephemeral green algae was similar in the predation risk and direct predation treatments, suggesting that the consumption of snails by crabs plays a minimal role in mediating the trophic cascade. Increased attention to trait-mediated effects that are transmitted by predator-induced changes in prey behavior may change our view of how predators exert their strong influence on community structure.     

Predation risk of typical ovoid and ‘winged’ morphs of Euplotes (Protozoa,Ciliophora)

Freshwater species of the genus Euplotes (Protozoa, Ciliophora) change their morphology in the presence of some of their predators. The ciliates develop extended lateral wings as well as dorsal and ventral projections which make engulfment by predators more difficult. In a series of laboratory experiments ingestion rates of four protozoan predators, the ciliates Lembadion bullinum, Dileptus anser, Stylonychia mytilus and Urostyla grandis, and one metazoan predator, the turbellarian Stenostomum sphagnetorum, on three species of Euplotes (E. octocarinatus, E. patella and E. aediculatus) were determined. It was calculated that the probability of rejection by a predator changed from 1:1 for ovoid morphs of Euplotes to about 2:1–20:1 for winged morphs of Euplotes, dependent on the prey and predator species that were combined. The nutritional condition of the prey also had some influence. In mixed-species cultures of prey and predators, transformed cells of E. octocarinatus survived for several months.     

Density-dependent effects of multiple predators sharing a common prey in an endophytic habitat

Multiple predator species feeding on a common prey can lead to higher or lower predation than would be expected by simply combining their individual effects. Such emergent multiple predator effects may be especially prevalent if predators share feeding habitat. Despite the prevalence of endophagous insects, no studies have examined how multiple predators sharing an endophytic habitat affect prey or predator reproduction. We investigated density-dependent predation of Thanasimus dubius (Coleoptera: Cleridae) and Platysoma cylindrica (Coleoptera: Histeridae) on a bark beetle prey, Ips pini (Coleoptera: Scolytidae), in a laboratory assay. I. pini utilize aggregation pheromones to group-colonize and reproduce within the stems of conifers. T. dubius and P. cylindrica exploit these aggregation pheromones to arrive simultaneously with the herbivore. Adult T. dubius prey exophytically, while P. cylindrica adults enter and prey within the bark beetle galleries. Larvae of both predators prey endophytically. We used a multiple regression analysis, which avoids confounding predator composition with density, to examine the effects of varying predator densities alone and in combination on herbivore establishment, herbivore reproduction, and predator reproduction. Predators reduced colonization success by both sexes, and decreased I. pini reproduction on a per male and per female basis. The combined effects of these predators did not enhance or reduce prey establishment or reproduction in unexpected manners, and these predators were entirely substitutable. The herbivores net replacement rate was never reduced significantly below one at prey and predator densities emulating field conditions. Similar numbers of each predator species emerged from the logs, but predator reproduction suffered from high intraspecific interference. The net replacement rate of P. cylindrica was not affected by conspecifics or T. dubius. In contrast, the net replacement rate of T. dubius decreased with the presence of conspecifics or P. cylindrica. Combinations of both predators led to an emergent effect, a slightly increased net replacement rate of T. dubius. This may have been due to predation by larval T. dubius on pupal P. cylindrica, as P. cylindrica develops more rapidly than T. dubius within this shared habitat.     

Plant-specific expression of antipredator behaviour by larval damselflies

Summary In this study of interactions between larval damselflies (Zygoptera: Coenagrionidae) and pumpkinseed sunfish (Lepomis gibbosus) we focus on the behaviour of the damselfly prey. First we document a prey behaviour in which damselflies use plant stems or leaves to hide from pumpkinseeds. We then test two hypotheses: (1) that damselfly hiding is a specific antipredator behaviour and (2) that hiding occurs more frequently in plant habitats where damselflies experience greater risk of predation. Since plant species growth forms can influence predation risk, our second hypothesis implies that hiding behaviour is conditional upon the type of vegetation providing habitat structure. Conditional expression of antipredator behaviour according to vegetation type may be important in littoral environments, since predator-prey interactions can occur in habitats with a wide range of macrophyte growth forms. The first hypothesis was supported by our findings that damselfly hiding increased in frequency in the presence of pumpkinseeds, that it was related to the frequency of predator approaches, and that its use reduced damselfly predation risk in high risk habitats. The second hypothesis was supported by our results that damselfly hiding rates were greater in the high risk Scirpus habitats than in the lower risk Potamogeton habitats. These results indicate that prey behaviour can influence predator-prey interactions, and that variation in plant growth form can influence prey behaviour, thus contributing to the impact of habitat structure on predator-prey dynamics.     

Effects of adaptive predatory and anti-predator behaviour in a two-prey—one-predator system

Summary Two prey populations that share a common predator can interact indirectly by causing changes in the predator's foraging behaviour. Previous work suggests that adaptive choice of prey by the predator usually has two related consequences: (i) the predation rate on a particular prey species increases with the relative and/or absolute abundance of that prey; and (ii) increases in either prey population produce a short-term increase in the fitness of the other prey (short-term indirect mutualism between prey). This paper investigates how these two consequences are changed if the prey exhibit adaptive anti-predator behaviour. In this case, the predation rate on a particular prey often decreases as the prey's density increases. The predator then usually exhibits negative switching between prey. However, the presence of adaptive antipredator behaviour does not change the short-term mutualism between prey. In this case, as a prey becomes less common, it achieves a larger growth rate by reducing its anti-predator effort. These results imply that observations of the relationship between prey density and predation rate cannot be used to infer the nature of the behavioural indirect effect between prey that share a predator.     

Food-web models that generate constant predator-prey ratios

Summary An approximately constant ratio of number of predator species/number of prey species is observed in several natural communities, although the exact value of the ratio may vary with habitat and the types of organisms in the food web. We test the hypothesis that a constant predator/prey ratio can be generated by what Holt (1977) terms apparent competition and what Jeffries and Lawton (1984) call competition for enemy-free space. We create simple, two trophic-level communities by drawing species of predators and prey at random from a species pool, simulating their interactions using Lotka-Volterra models. The simulated food webs converge over successive periods of invasion and extinction to locally stable systems with approximately constant ratios of number of predator species/number of prey species, despite varying initial conditions. As expected, predator/prey ratios take different values depending upon the biology of the simulated species. We conclude that apparent competition between prey species via shared enemies may be one mechanism whereby approximately constant predator/prey ratios are generated in natural communities.     

Prey abundance vs diet breadth in a spider test system

Summary Decisions made as to what prey types to include in the diet were analysed for two populations of the spider,Agelenopsis aperta existing under markedly different prey availability and predation levels. Potential prey types were ranked as to their relative profitabilities with respect to energy gain per handling effort and predation risk. Members of the population experiencing limited prey availability but low risk of predation to visually hunting predators exhibited a significantly higher capture attempt rate towards all prey encountered than the population for which prey were abundant but for which predation was a significant problem. Neither spider population preferentially attacked prey that exhibited higher profitability rankings. An experiment was completed that indicates thatA. aperta can discriminate between more and less profitable prey. Suggestions are made as to why the population experiencing abundant food did not exhibit a narrower diet when compared to the population existing under limited food.