Colonially breeding seabirds: Predators or prey?

The evolution of aggregation in seabirds is usually attributed to predation pressure, although many authors have argued for the role of foraging constraints to be considered. Discriminating between factors that result in group living and characteristics arising from group living has been regarded as an insoluble problem; yet it is possible to test the predictions of the different models proposed to explain colonial nesting. The resulting explanation suggests that seabird aggregations have evolved in order to utilize a dispersed and unpredictable food supply. As such, colonial nesting leads to increased vulnerability to predation, rather than being a strategy adopted to combat it.     

Non-webbuilding spiders: prey specialists or generalists?

Summary Feeding experiments were performed with seven species of non-webbuilding spiders and a variety of prey taxa. Some species were generally polyphagous whereas other spiders restricted their prey to a few groups. At one end of the spectrum of prey specialization the thomisid Misumena vatia is limited to a few taxa of possible prey (Table 1). The literature of prey records of non-webbuilding spiders is reviewed (Table 2) with special emphasis on oligophagous or monophagous spiders. Monophagous spiders are generally rare and have specialized on only a few prey taxa: social insects (ants, bees, termites) and spiders.     

Predator–prey encounter rates in freshwater piscivores: effects of prey density and water transparency

One of the most fundamental components of predator–prey models is encounter rate, modelled as the product of prey density and search efficiency. Encounter rates have, however, rarely been measured in empirical studies. In this study, we used a video system approach to estimate how encounter rates between piscivorous fish that use a sit-and-wait foraging strategy and their prey depend on prey density and environmental factors such as turbidity. We first manipulated prey density in a controlled pool and field enclosure experiments where environmental factors were held constant. In a correlative study of 15 freshwater lakes we then estimated encounter rates in natural habitats and related the results to both prey fish density and environmental factors. We found the expected positive dependence of individual encounter rates on prey density in our pool and enclosure experiments, whereas the relation between school encounter rate and prey density was less clear. In the field survey, encounter rates did not correlate with prey density but instead correlated positively with water transparency. Water transparency decreases with increasing prey density along the productivity gradient and will reduce prey detection distance and thus predator search efficiency. Therefore, visual predator–prey encounter rates do not increase, and may even decrease, with increasing productivity despite increasing prey densities.     

Predator–prey system with strong Allee effect in prey

Global bifurcation analysis of a class of general predator–prey models with a strong Allee effect in prey population is given in details. We show the existence of a point-to-point heteroclinic orbit loop, consider the Hopf bifurcation, and prove the existence/uniqueness and the nonexistence of limit cycle for appropriate range of parameters. For a unique parameter value, a threshold curve separates the overexploitation and coexistence (successful invasion of predator) regions of initial conditions. Our rigorous results justify some recent ecological observations, and practical ecological examples are used to demonstrate our theoretical work.     

Effects of rapid prey evolution on predator–prey cycles

We study the qualitative properties of population cycles in a predator-prey system where genetic variability allows contemporary rapid evolution of the prey. Previous numerical studies have found that prey evolution in response to changing predation risk can have major quantitative and qualitative effects on predator-prey cycles, including: (1) large increases in cycle period, (2) changes in phase relations (so that predator and prey are cycling exactly out of phase, rather than the classical quarter-period phase lag), and (3) "cryptic" cycles in which total prey density remains nearly constant while predator density and prey traits cycle. Here we focus on a chemostat model motivated by our experimental system (Fussmann et al. in Science 290:1358-1360, 2000; Yoshida et al. in Proc roy Soc Lond B 424:303-306, 2003) with algae (prey) and rotifers (predators), in which the prey exhibit rapid evolution in their level of defense against predation. We show that the effects of rapid prey evolution are robust and general, and furthermore that they occur in a specific but biologically relevant region of parameter space: when traits that greatly reduce predation risk are relatively cheap (in terms of reductions in other fitness components), when there is coexistence between the two prey types and the predator, and when the interaction between predators and undefended prey alone would produce cycles. Because defense has been shown to be inexpensive, even cost-free, in a number of systems (Andersson et al. in Curr Opin Microbiol 2:489-493, 1999: Gagneux et al. in Science 312:1944-1946, 2006; Yoshida et al. in Proc Roy Soc Lond B 271:1947-1953, 2004), our discoveries may well be reproduced in other model systems, and in nature. Finally, some of our key results are extended to a general model in which functional forms for the predation rate and prey birth rate are not specified.     

Analysis of a competitive prey–predator system with a prey refuge

Gauss's competitive exclusive principle states that two competing species having analogous environment cannot usually occupy the same space at a time but in order to exploit their common environment in a different manner, they can co-exist only when they are active in different times. On the other hand, several studies on predators in various natural and laboratory situations have shown that competitive coexistence can result from predation in a way by resisting any one prey species from becoming sufficiently abundant to outcompete other species such that the predator makes the coexistence possible. It has also been shown that the use of refuges by a fraction of the prey population exerts a stabilizing effect in the interacting population dynamics. Further, the field surveys in the Sundarban mangrove ecosystem reveal that two detritivorous fishes, viz. Liza parsia and Liza tade (prey population) coexist in nature with the presence of the predator fish population, viz. Lates calcarifer by using refuges.     

Does prey preference change as a result of prey species being presented together? Analysis of prey selection by the predatory mite Typhlodromus pyri (Acarina: Phytoseiidae)

Summary Prey-selection behaviour of the phytoseiid mite Typhlodromus pyri Scheuten was analysed with a Markovtype model of feeding-state dynamics and feeding-state dependent searching behaviour (Sabelis 1981, 1986, 1989; Metz and Van Batenburg 1985a, b). All behavioural characteristics of the predator which are independent of the feeding state were represented by one parameter. The remaining feeding-state dependent characteristics were represented by a function of the feeding state, with one parameter. The best parameter values to describe a predator-prey interaction were determined by fitting the model to the predation rates in monocultures. Under the assumption that the parameter values are not dependent on the composition of prey species supply, the diet of the predators in mixed cultures was predicted from parameters estimated in monoculture experiments.Two prey types, apple rust mite (Aculus schlechtendali (Nalepa)) adults and European red spider mite (Panonychus ulmi (Koch)) larvae were studied. A large discrepancy was observed between calculated and experimentally determined predation rates of T. pyri in mixed cultures: the predators actually killed 3–7 times more P. ulmi larvae than was predicted by the model.The large difference between observed and predicted predation rates in mixed cultures cannot be explained by changes in the behaviour of the prey species as a result of being together. Therefore, it seems likely that the prey selection behaviour of the predator was different when prey species were presented together than when presented singly. Apparently the predatory mite T. pyri prefers P. ulmi to S. schlechtendali.     

Predator–prey interactions and changing environments: who benefits?

While aquatic environments have long been thought to be more moderate environments than their terrestrial cousins, environmental data demonstrate that for some systems this is not so. Numerous important environmental parameters can fluctuate dramatically, notably dissolved oxygen, turbidity and temperature. The roles of dissolved oxygen and turbidity on predator-prey interactions have been discussed in detail elsewhere within this issue and will be considered only briefly here. Here, we will focus primarily on the role of temperature and its potential impact upon predator-prey interactions. Two key properties are of particular note. For temperate aquatic ecosystems, all piscine and invertebrate piscivores and their prey are ectothermic. They will therefore be subject to energetic demands that are significantly affected by environmental temperature. Furthermore, the physical properties of water, particularly its high thermal conductivity, mean that thermal microenvironments will not exist so that fine-scale habitat movements will not be an option for dealing with changing water temperature in lentic environments. Unfortunately, there has been little experimental analysis of the role of temperature on such predator-prey interactions, so we will instead focus on theoretical work, indicating that potential implications associated with thermal change are unlikely to be straightforward and may present a greater threat to predators than to their prey. Specifically, we demonstrate that changes in the thermal environment can result in a net benefit to cold-adapted species through the mechanism of predator-prey interactions.     

Effect of prey type and inorganic turbidity on littoral predator–prey interactions in a shallow lake: an experimental approach

Predation often represents the prevailing process shaping aquatic ecosystems. As foraging and antipredatory behaviour frequently relate to vision, turbidity may often impair the interactions between the predator and its prey, depending on prey type and source and level of turbidity. We studied the effect of inorganic turbidity (0–30 NTU) on the effectiveness of fish feeding on two types of prey in different habitats: free-swimming cladoceran (Daphnia pulex) in open water and plant-associated cladoceran (Sida crystallina) attached to Nuphar lutea leaves. For the planktivore, we used vision-oriented perch (Perca fluviatilis) common in the littoral zone of temperate lakes. In our study, increasing inorganic turbidity did not appear to initiate any significant change in the feeding efficiency of perch on free-swimming Daphnia pulex. However, we saw a markedly different feeding efficiency when perch targeted plant-attached Sida crystallina. Our results substantiate that floating-leaved macrophytes in turbid lakes may provide a favourable habitat for plant-attached cladocerans.     

What determines prey selection in owls? Roles of prey traits,prey class,environmental variables,and taxonomic specialization

Ecological theory suggests that prey size should increase with predator size, but this trend may be masked by other factors affecting prey selection, such as environmental constraints or specific prey preferences of predator species. Owls are an ideal case study for exploring how predator body size affects prey selection in the presence of other factors due to the ease of analyzing their diets from owl pellets and their widespread distributions, allowing interspecific comparisons between variable habitats. Here, we analyze various dimensions of prey resource selection among owls, including prey size, taxonomy (i.e., whether or not particular taxa are favored regardless of their size), and prey traits (movement type, social structure, activity pattern, and diet). We collected pellets of five sympatric owl species (Athene noctua, Tyto alba, Asio otus, Strix aluco, and Bubo bubo) from 78 sites across the Mediterranean Levant. Prey intake was compared between sites, with various environmental variables and owl species as predictors of abundance. Despite significant environmental impacts on prey intake, some key patterns emerge among owl species studied. Owls select prey by predator body size: Larger owls tend to feed on wider ranges of prey sizes, leading to higher means. In addition, guild members show both specialization and generalism in terms of prey taxa, sometimes in contrast with the expectations of the predator–prey body size hypothesis. Our results suggest that while predator body size is an important factor in prey selection, taxon specialization by predator species also has considerable impact.     

Signaling by decorating webs: luring prey or deterring predators?

Many organisms convey false signals to mislead their prey orpredators. Some orb-weaving spiders build conspicuous structureson webs called decorations. Web decorations and spider colorationsare both suggested to be important signals involved in interactionsbetween spiders and other organisms. There are several hypothesesabout the functions of signaling by decorations, among whichprey attraction had received much support, but empirical evidenceregarding predator defense is controversial. In this study,we conducted field experiments to investigate the effects ofspider decoration and coloration on insect interception ratesof webs built by Argiope aemula and to evaluate whether presenceof decorations may decrease predation risk of spiders. Decoratedwebs with spiders present had the highest prey interceptionrate, followed by undecorated webs with spiders, and then undecoratedwebs without spiders. Such results indicated that decorationsof Argiope spiders functioned as visual lures, and so did spiders'bright body colorations. In the field, almost all wasp attackevents occurred on medium-sized spiders rather than on largeones. Moreover, medium-sized Arg. aemula on decorated webs receivedfar more attacks than those on undecorated webs. Results ofthis study thus show that the signals conveyed by decorationscan visually lure prey but at the cost of an increased predationrisk. Received 20 March 2007; revised 3 August 2007; accepted 5 August 2007.     

Do unprofitable prey evolve traits that profitable prey find difficult to exploit?

Prey that are unprofitable to attack (for example, those containing noxious chemicals) are often conspicuously patterned and move in a slower and more predictable manner than species lacking these defences. Contemporary theories suggest these traits have evolved as warning signals because they can facilitate both associative and discriminative avoidance learning in predators. However, it is unclear why these particular traits and not others have tended to evolve in unprofitable prey. Here we show using a signal detection model that unprofitable prey will evolve conspicuous colours and patterns partly because these characteristics cannot readily evolve in profitable prey without close mimicry. The stability of this signal is maintained through the costs of dishonesty in profitable prey. Indeed, unprofitable prey will sometimes evolve a conspicuous form to reduce mimetic parasitism, even in the unlikely event that this form can be more closely mimicked. This is one of the first mathematical models of the evolution of warning signals to allow for the possibility of mimicry, yet our analyses suggest it may offer a general explanation as to why warning signals take the form that they do. Warning signals and mimicry may therefore be more closely related than is currently supposed.     

Incorporating prey refuge in a prey–predator model with a Holling type I functional response: random dynamics and population outbreaks

A prey–predator discrete-time model with a Holling type I functional response is investigated by incorporating a prey refuge. It is shown that a refuge does not always stabilize prey–predator interactions. A prey refuge in some cases produces even more chaotic, random-like dynamics than without a refuge and prey population outbreaks appear. Stability analysis was performed in order to investigate the local stability of fixed points as well as the several local bifurcations they undergo. Numerical simulations such as parametric basins of attraction, bifurcation diagrams, phase plots and largest Lyapunov exponent diagrams are executed in order to illustrate the complex dynamical behavior of the system.     

Optimal pursuit times: How long should predators pursue their prey?

Data suggest that the length of time that a predator pursues a given prey is both variable and adjustable. We present an extension of the optimal diet model that incorporates adjustable pursuit times to predict how long a predator should pursue their prey. The model makes three significant contributions.

1. 1. When the density of good prey is increased, poor prey should gradually disappear from the diet.

2. 2. Depending on how the encounter rates with prey are varied, the model generates patterns of frequency-dependent and inverse frequency-dependent foraging.

3. 3. The density of poor prey can influence the proportion captured of more valuable prey.

     

Use of prey hotspots by an avian predator: purposeful unpredictability?

The use of space by predators in relation to their prey is a poorly understood aspect of predator-prey interactions. Classic theory suggests that predators should focus their efforts on areas of abundant prey, that is, prey hotspots, whereas game-theoretical models of predator and prey movement suggest that the distribution of predators should match that of their prey's resources. If, however, prey are spatially anchored to one location and these prey have particularly strong antipredator responses that make them difficult to capture with frequent attacks, then predators may be forced to adopt alternative movement strategies to hunt behaviorally responsive prey. We examined the movement patterns of bird-eating sharp-shinned hawks (Accipiter striatus) in an attempt to shed light on hotspot use by predators. Our results suggest that these hawks do not focus on prey hotspots such as bird feeders but instead maintain much spatial and temporal unpredictability in their movements. Hawks seldom revisited the same area, and the few frequently used areas were revisited in a manner consistent with unpredictable returns, giving prey little additional information about risk.     

A predator–prey refuge system: Evolutionary stability in ecological systems

A refuge model is developed for a single predator species and either one or two prey species where no predators are present in the prey refuge. An individual’s fitness depends on its strategy choice or ecotype (predators decide which prey species to pursue and prey decide what proportion of their time to spend in the refuge) as well as on the population sizes of all three species. It is shown that, when there is a single prey species with a refuge or two prey species with no refuge compete only indirectly (i.e. there is only apparent competition between prey species), that stable resident systems where all individuals in each species have the same ecotype cannot be destabilized by the introduction of mutant ecotypes that are initially selectively neutral. In game-theoretic terms, this means that stable monomorphic resident systems, with ecotypes given by a Nash equilibrium, are both ecologically and evolutionarily stable. However, we show that this is no longer the case when the two indirectly-competing prey species have a refuge. This illustrates theoretically that two ecological factors, that are separately stabilizing (apparent competition and refuge use), may have a combined destabilizing effect from the evolutionary perspective. These results generalize the concept of an evolutionarily stable strategy (ESS) to models in evolutionary ecology. Several biological examples of predator–prey systems are discussed from this perspective.     

How to analyse prey preference when prey density varies? A new method to discriminate between effects of gut fullness and prey type composition

Summary State-dependent changes in prey preference are among the phenomena to be expected in studies of predator behaviour. For example, the rate of attack on each prey type is well known to be affected by the state of satiation, the dynamics of which is often assumed to parallel that of gut fullness. An interesting question is whether satiation alone is the determinant of the attack rate or whether the particular mixture of prey types in the predator's direct environment has an additional influence by itself. To detect examples of the latter type the predictive method proposed by Cock (1978) may be useful. In the present paper the predictive tool is a model built on the assumption that gut fullness is the sole internal state variable determining the attack rate. It is provided with parameter estimates from observations in monocultures of each type and then used to predict predation in mixtures of prey types. When measured predation on these prey types differs from what is predicted, the model may be too simple in various respects, one of which is that predators change prey preference in response to their own sample estimates of the densities of each prey type and their (innate or sample) estimate of the profitability of each prey type in terms of reproductive success. Thus, the lack of fit of the model poses a challenging problem, for to explain it one must identify underlying causes, such as differences in prey quality with respect to scarce nutrients or noxious chemicals that need to be detoxified or rendered harmless in other ways. The predictive approach is illustrated by analysis of preference of predatory mites (Phytoseiulus persimilis Athias-Henriot and Typhlodromus occidentalis Nesbitt) with respect to various stages of development of their prey, the two-spotted spider mite (Tetranychus urticae Koch). The results show that the relation between attack rate and gut fullness might well explain prey stage preference of predatory mites when the prey stages are presented together rather than each alone. In another paper by Dicke et al. (1989) marked deviations between predicted and measured diet are reported when the predatory mite, Typhlodromus pyri Scheuten, was offered a choice between two prey species, i.e. apple rust mites and (larvae of) European red spider mites. The underlying causes are to be revealed by further research, the impetus of which is born out by use of the method proposed by Cock (1978) and extended in this paper.     

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.     

Predator migration in response to prey density: What are the consequences?

A predator-prey metapopulation model with two identical patches and only migration of the predator is investigated. Local predator-prey interaction is described by the so-called Rosenzweig-MacArthur model, while the migration term of the predator is put in a nonlinear form, which is derived by extending the Holling time budget argument to migration. In particular, a dimensionless parameter theta is introduced to quantify the migration tendency of predators while they are handling their prey, which gives rise to a family of models connecting two extremes: predators have no inclination to migrate while handling prey (theta = 0) and standard diffusion (theta = 1). Various aspects of the model, including changes in the number and the stability of equilibria and limit cycles, are investigated. We then focus on the key question: "Does spatial structure lead to a substantial damping of the violent oscillations exhibited by many predator-prey models?". It is known that the answer is "yes" if one adopts standard diffusion (theta = 1). However, we present substantial evidence that the answer is "no" if one takes theta = 0. We conclude that the migration submodel is an important constituent of a spatial predator-prey model and that the issue deserves scrutiny, both experimentally and theoretically.