Interspecific infanticide deters predators

It is well known that young, small predator stages are vulnerable to predation by conspecifics, intra-guild competitors or hyperpredators. It is less known that prey can also kill vulnerable predator stages that present no danger to the prey. Since adult predators are expected to avoid places where their offspring would run a high predation risk, this opens the way for potential prey to deter dangerous predator stages by killing vulnerable predator stages. We present an example of such a complex predator–prey interaction. We show that (1) the vulnerable stage of an omnivorous arthropod prey discriminates between eggs of a harmless predator species and eggs of a dangerous species, killing more eggs of the latter; (2) prey suffer a minor predation risk from newly hatched predators; (3) adult predators avoid ovipositing near killed predator eggs, and (4) vulnerable prey near killed predator eggs experience an almost fourfold reduction of predation. Hence, by attacking the vulnerable stage of their predator, prey deter adult predators and thus reduce their own predation risk. This provides a novel explanation for the killing of vulnerable stages of predators by prey and adds a new dimension to anti-predator behaviour.     

Transient dynamics limit the effectiveness of keystone predation in bringing about coexistence

We analyze the transient dynamics of simple models of keystone predation, in which a predator preferentially consumes the dominant of two (or more) competing prey species. We show that coexistence is unlikely in many systems characterized both by successful invasion of either prey species into the food web that lacks it and by a stable equilibrium with high densities of all species. Invasion of the predator-resistant consumer species often causes the resident, more vulnerable prey to crash to such low densities that extinction would occur for many realistic population sizes. Subsequent transient cycles may entail very low densities of the predator or of the initially successful invader, which may also preclude coexistence of finite populations. Factors causing particularly low minimum densities during the transient cycles include biotic limiting resources for the prey, limited resource partitioning between the prey, a highly efficient predator with relatively slow dynamics, and a vulnerable prey whose population dynamics are rapid relative to the less vulnerable prey. Under these conditions, coexistence of competing prey via keystone predation often requires that the prey's competitive or antipredator characteristics fall within very narrow ranges. Similar transient crashes are likely to occur in other food webs and food web models.     

Top–down limits on prey populations may be more severe in larger prey species,despite having fewer predators

Variation in the vulnerability of herbivore prey to predation is linked to body size, yet whether this relationship is size‐nested or size‐partitioned remains debated. If size‐partitioned, predators would be focused on prey within their preferred prey size range. If size‐nested, smaller prey species should become increasingly more vulnerable because increasingly more predators are capable of catching them. Yet, whether either of these strategies manifests in top–down prey population limitation would depend both on the number of potential predator species as well as the total mortality imposed. Here we use a rare ecosystem scale ‘natural experiment’ comparing prey population dynamics between a period of intense predator persecution and hence low predator densities and a period of active predator protection and population recovery. We use three decades of data on herbivore abundance and distribution to test the role of predation as a mechanism of population limitation among prey species that vary widely in body size. Notably, we test this within one of the few remaining systems where a near‐full suite of megaherbivores occur in high density and are thus able to include a thirtyfold range in herbivore body size gradient. We test whether top–down limitation on prey species of particular body size leads to compositional shifts in the mammalian herbivore community. Our results support both size‐nested and size‐partitioning predation but suggest that the relative top–down limiting impact on prey populations may be more severe for intermediate sized species, despite having fewer predators than small species. In addition we show that the gradual recovery of predator populations shifted the herbivore community assemblage towards large‐bodied species and has led to a community that is strongly dominated by large herbivore biomass.     

Keeping the herds healthy and alert: implications of predator control for infectious disease

Predator control programmes are generally implemented in an attempt to increase prey population sizes. However, predator removal could prove harmful to prey populations that are regulated primarily by parasitic infections rather than by predation. We develop models for microparasitic and macroparasitic infection that specify the conditions where predator removal will (a) increase the incidence of parasitic infection, (b) reduce the number of healthy individuals in the prey population and (c) decrease the overall size of the prey population. In general, predator removal is more likely to be harmful when the parasite is highly virulent, macroparasites are highly aggregated in their prey, hosts are long‐lived and the predators select infected prey.     

Individual base model of predator-prey system shows predator dominant dynamics

Individual base model of predator-prey system is constructed. Both predator and prey species have age structure and cohorts of early reproductive age have competitive advantage. The model has linear functional response in predation behavior and includes the effect of interference among predators and delay of population growth from resource intake, not by functional response but by calculation procedure. Each foraging action is calculated successively and surplus or scarce of acquired resources is interpreted into population size through individual birth and death. This model shows that biomass of prey killed by predator is dependent on demand of predator and that heterogeneity in predator population is essential in persistency and stability of predator-prey system. Heterogeneity of predator makes predator individuals of less competing ability die rapidly. Rapid death of weak individuals causes rapid decrease of total demand of predator and that makes enough room for survived predators. Therefore, the biomass of killed prey is dependent on predator's demand. As young or infant population of predator are the more vulnerable to shortage of prey, and when many of them cannot survive to reproductive age, they can stabilize the system by wasting excessive prey with only temporal numerical increase of predator population.     

Sacrificial males: the potential role of copulation and predation in contributing to copepod sex‐skewed ratios

Predation is thought to play a selective role in the emergence of behavioural traits in prey. Differences in behaviour between prey demographics may, therefore, be driven by predation with select components of the population being less vulnerable to predators. While under controlled conditions prey demography has been shown to have consequences for predation success, investigations linking these implications to natural prey population demographics are scarce. Here we assess predator–prey dynamics between notonectid predators (backswimmers) and Lovenula raynerae (Copepoda), key faunal groups in temperate ephemeral pond ecosystems. Using a combination of field and experimental approaches we test for the development and mechanism of predation‐induced sex‐skewed ratios. A natural population of L. raynerae was tracked over time in relation to their predator (notonectid) and prey (Cladocera) numbers. In the laboratory, L. raynerae sex ratios were also assessed over time but in the absence of predation pressure. Predation success and prey performance experiments evaluating differences between L. raynerae male, female, gravid female and copulating pairs exposed to notonectid predation were then examined. Under natural conditions, a female dominated copepod population developed over time and was correlated to predation pressure, while under predator‐free conditions non sex‐skewed prey population demographics persisted. Predator–prey laboratory trials showed no difference in vulnerability and escape performance for male, female and gravid female copepods, but pairs in copula were significantly more vulnerable to predation. This vulnerability was not shared by both sexes, with only female copepods ultimately escaping from successful predation on a mating pair. These results suggest that contact periods during copula may contribute to the development of sex‐skewed copepod ratios over time in ecosystems dominated by hexapod predators. This is discussed within the context of vertebrate and invertebrate predation and how these dissimilar types of predation are likely to have acted as selective pressures for copepod mating systems.     

A review of predation as a limiting factor for bird populations in mesopredator‐rich landscapes: a case study of the UK

The impact of increasing vertebrate predator numbers on bird populations is widely debated among the general public, game managers and conservationists across Europe. However, there are few systematic reviews of whether predation limits the population sizes of European bird species. Views on the impacts of predation are particularly polarised in the UK, probably because the UK has a globally exceptional culture of intensive, high‐yield gamebird management where predator removal is the norm. In addition, most apex predators have been exterminated or much depleted in numbers, contributing to a widely held perception that the UK has high numbers of mesopredators. This has resulted in many high‐quality studies of mesopredator impacts over several decades. Here we present results from a systematic review of predator trends and abundance, and assess whether predation limits the population sizes of 90 bird species in the UK. Our results confirm that the generalist predators Red Fox (Vulpes vulpes) and Crows (Corvus corone and C. cornix) occur at high densities in the UK compared with other European countries. In addition, some avian and mammalian predators have increased numerically in the UK during recent decades. Despite these high and increasing densities of predators, we found little evidence that predation limits populations of pigeons, woodpeckers and passerines, whereas evidence suggests that ground‐nesting seabirds, waders and gamebirds can be limited by predation. Using life‐history characteristics of prey species, we found that mainly long‐lived species with high adult survival and late onset of breeding were limited by predation. Single‐brooded species were also more likely to be limited by predation than multi‐brooded species. Predators that depredate prey species during all life stages (i.e. from nest to adult stages) limited prey numbers more than predators that depredated only specific life stages (e.g. solely during the nest phase). The Red Fox and non‐native mammals (e.g. the American Mink Neovison vison) were frequently identified as numerically limiting their prey species. Our review has identified predator–prey interactions that are particularly likely to result in population declines of prey species. In the short term, traditional predator‐management techniques (e.g. lethal control or fencing to reduce predation by a small number of predator species) could be used to protect these vulnerable species. However, as these techniques are costly and time‐consuming, we advocate that future research should identify land‐use practices and landscape configurations that would reduce predator numbers and predation rates.     

Changing vulnerability to predation related to season and sex in an African ungulate assemblage

The consequences of predation for prey population dynamics depend on the extent to which this mortality is predisposed by malnutrition or senescence, or additive in the sense that animals that would otherwise not have died at that time were killed. In places lacking effective predators, few adult ungulates die during the summer or wet season months when food is plentifully available. Hence the seasonal distribution of predator kills as well as the age and sex classes of the prey indicates the extent to which malnutrition contributes to mortality as well as other influences on vulnerability. Using records of animal deaths assembled over 35 years in South Africa's Kruger National Park, these patterns were investigated for 12 ungulate species forming the prey of lions, and for three other large predators with respect to one prey species. Buffalo, kudu and giraffe were more strongly represented in kills made during the late dry season, while wildebeest and zebra made relatively greater contributions during the wet season. Impala, waterbuck, warthog and rarer antelope species became more prominent in kills during transitional periods between seasons. Five prey species showed an elevation in representation of males in lion kills during the mating season, as well as impala for all predator species. Females were more prominently represented in kills during the time of late gestation and parturition for three prey species. Hence reproductive activities as well as changing vegetation cover and food resources affected vulnerability to predation. Shifts in susceptibility to predation over the seasonal cycle corresponded with rainfall‐related variation in the annual representation of these ungulate species in lion kills. The availability of vulnerable prey species, age and sex classes at different stages of the seasonal cycle helps maintain a high abundance of lions. These factors contribute to the strong additive impact that predation has had on the abundance of some of these ungulate populations.     

Evolutionary Consequences of Predation for Pathogens in Prey

This article investigates the impact of predation on the coexistence and competitive exclusion of pathogen strains in the prey. Two types of predator are considered—a generalist and a specialist. For each type of predator, we assume that the predator can discriminate among susceptible and infected with each strain prey. The two strains will competitively exclude each other in the absence of predation with the strain with the larger reproduction number persisting. If a generalist predator preys discriminantly and the disease is fatal, then depending on the predation level, a switch in the dominant pathogen may occur. Thus, for some predation levels, the first strain may persist while for other predation levels the second strain may persist. Furthermore, a specialist predator preying discriminantly may mediate the coexistence of the two strains. Although in most cases increasing predation reduces the disease load in the prey, when predation leads to coexistence, it may also lead to increase in the disease load.     

Effect of disease-selective predation on prey infected by contact and external sources

We propose and analyze a simple mathematical model for susceptible prey (S)–infected prey (I)–predator (P) interaction, where the susceptible prey population (S) is infected directly from external sources as well as through contact with infected class (I) and the predator completely avoids consuming the infected prey. The model is analyzed to obtain different thresholds of the key parameters under which the system exhibits stability around the biologically feasible equilibria. Through numerical simulations we display the effects of external infection and the infection through contact on the system dynamics in the absence as well as in the presence of the predator. We compare the system dynamics when infection occurs only through contact, with that when it occurs through contact and external sources. Our analysis demonstrates that under a disease-selective predation, stability and oscillations of the system is determined by two key parameters: the external infection rate and the force of infection through contact. Due to the introduction of external infection, the predator and the prey population show limit-cycle oscillations over a range parametric values. We suggest that while predicting the dynamics of such an eco-epidemiological system, the modes of infection and the infection rates might be carefully investigated.     

Local prey vulnerability increases with multiple attacks by a predator

Theory and empirical evidence suggest that predator activity makes prey more wary and less vulnerable to predation. However if at least some prey in the population are energetically or spatially constrained, then predators may eventually increase local prey vulnerability because of the cumulative costs of anti‐predation behaviour. We tested whether repeated attacks by a predator might increase prey vulnerability in a system where redshanks on a saltmarsh are attacked regularly by sparrowhawks from adjacent woodland. Cumulative attack number led to a reduction in redshank numbers and flock size (but had no effect on how close redshanks fed to predator‐concealing cover) because some redshanks moved to safer but less profitable habitats, leaving smaller flocks on the saltmarsh. This effect held even though numbers of redshank on the saltmarsh increased with time of day. As a result of the change in flock size, predicted attack‐success increased up to 1.6‐fold for the sparrowhawk, while individual risk of capture for the redshank increased up to 4.5‐fold among those individuals remaining on the saltmarsh. The effect did not arise simply because hawks were more likely to attack smaller flocks because attack rate was not dependent on flock size or abundance. Our data demonstrate that when some individual prey are constrained in their ability to feed on alternative, safer foraging sites, their vulnerability to predation increases as predator attacks accumulate, although those, presumably better quality individuals that leave the immediate risky area will have lower vulnerability, so that the mean vulnerability across the entire population may not have changed substantially. This suggests that the selective benefits of multiple low‐cost attacks by predators on prey could potentially lead to 1) locally heightened trait‐mediated interactions, 2) locally reduced interference among competing predators, and 3) the evolution of active prey manipulation by predators.     

The impact of mortality on predator population size and stability in systems with stage-structured prey

The relationships between a predator population's mortality rate and its population size and stability are investigated for several simple predator-prey models with stage-structured prey populations. Several alternative models are considered; these differ in their assumptions about the nature of density dependence in the prey's population growth; the nature of stage-transitions; and the stage-selectivity of the predator. Instability occurs at high, rather than low predator mortality rates in most models with highly stage-selective predation; this is the opposite of the effect of mortality on stability in models with homogeneous prey populations. Stage-selective predation also increases the range of parameters that lead to a stable equilibrium. The results suggest that it may be common for a stable predator population to increase in abundance as its own mortality rate increases in stable systems, provided that the predator has a saturating functional response. Sufficiently strong density dependence in the prey generally reverses this outcome, and results in a decrease in predator population size with increasing predator mortality rate. Stability is decreased when the juvenile stage has a fixed duration, but population increases with increasing mortality are still observed in large areas of stable parameter space. This raises two coupled questions which are as yet unanswered; (1) do such increases in population size with higher mortality actually occur in nature; and (2) if not, what prevents them from occurring? Stage-structured prey and stage-related predation can also reverse the 'paradox of enrichment', leading to stability rather than instability when prey growth is increased.     

The role of virus infection in a simple phytoplankton zooplankton system

Many planktonic species show spectacular bursts ("blooms") in population density. Though viral infections are known to cause behavioural and other changes in phytoplankton and other aquatic species, yet their role in regulating the phytoplankton population is still far from being understood. To study the role of viral diseases in the planktonic species, we model the phytoplankton-zooplankton system as a prey-predator system. Here the prey (phytoplankton) species is infected with a viral disease that divides the prey population into susceptible and infected classes, with the infected prey being more vulnerable to predation by the predator (zooplankton). The dynamical behaviour of the system is investigated from the point of view of stability and persistence both analytically and numerically. The model shows that infection can be sustained only above a threshold of force of infection, and, there exists a range in the infection rate where this system shows "bloom"-like stable limit cycle oscillations. The time series of natural "blooms" with different types of irregular oscillations can arise in this model simply from a biologically realistic feature, i.e., by the random variation of the epidemiological parameter (rate of infection) in the infected prey population. The difference in mean strength of infection alone can lead to the different types of patterns observed in natural planktonic blooms.     

Heterogeneous landscapes and the role of refuge on the population dynamics of a specialist predator and its prey

How, and where, a prey species survives predation by a specialist predator during low phases of population fluctuations or a cycle, and how the increase phase of prey population is initiated, are much-debated questions in population and theoretical ecology. The persistence of the prey species could be due mainly to habitats that act as refuges from predation and/or due to anti-predatory behaviour of individuals. We present models for the former conjecture in two (and three) habitat systems with a specialist predator and its favoured prey. The model is based on dispersal of prey between habitats with high reproductive output but high risk of predation, and less productive habitats with relatively low risk of predation. We illustrate the predictions of our model using parameters from one of the most intriguing vertebrate predator–prey systems, the multi-annual population cycles of boreal voles and their predators. We suggest that cyclic population dynamics could result from a sequence of extinction and re–colonization events. Field voles (Microtus agrestis), a key vole species in the system, can be hunted to extinction in their preferred meadow habitat, but persist in sub-optimal wet habitats where their main predator, the least weasel (Mustela nivalis nivalis) has a low hunting efficiency. Re–colonization of favourable habitats would occur after the predator population crashes. At the local scale, the model suggests that the periodicity and amplitude of population cycles can be strongly influenced by the relative availability of risky and safe habitats for the prey. Furthermore, factors like intra-guild predation may lead to reduced predation pressure on field voles in sub-optimal habitats, which would act as a refuge for voles during the low phase of their population cycles. Elasticity analysis suggested that our model is quite robust to changes in most parameters but sensitive to changes in the population dynamics of field voles in the optimal grassland habitat, and to the maximum predation rate of weasels.     

Sequential movement into coastal habitats and high spatial overlap of predator and prey suggest high predation pressure in protected areas

In theory, predators should attempt to match the distribution of their prey, and prey to avoid areas of high predation risk. However, there is a scarcity of empirical knowledge on predator and prey spatial use when both are moving freely in their natural environment. In the current study, we use information collated on a predators’ diet, its population structure, as well as predator and prey relative abundance, and track the movements of predator and prey simultaneously to compare habitat use and evaluate predation pressure. The study was conducted in elasmobranch protected areas of coastal Tasmania, Australia. The species considered were the broadnose sevengill shark Notorynchus cepedianus, the apex predator in the area, and five chondrichthyan prey species. Notorynchus cepedianus and its prey show similar seasonality in the use of these coastal areas: more abundant in warmer months and absent in winter. Predator and prey also showed high spatial overlap and similar habitat use patterns. These similar movement patterns of predator and prey combined with the additional ecological information (diet, population structure of predator, relative abundance of predator and prey) suggests that N. cepedianus move into coastal areas to exploit seasonally abundant prey. Also, while in protected areas, chondrichthyans are subjected to high predation pressure. Overall, results illustrate the value of simultaneously recording and integrating multiple types of information to explore predator–prey relationships and predation pressure.     

Retarded growth of cladoceran zooplankton in the presence of a copepod predator

Various instars of four different cladoceran species representing a wide spectrum of body size were grown at high food availability in the presence and in the absence of natural densities of an invertebrate predator, a cyclopoid copepod Acanthocyclops robustus (G.O. Sars). Daily weight increments calculated from individual weights at the end and at the beginning of each 1, 2 or 4 day experiment, showed that individual growth was more or less drastically retarded in the presence of the predator as well as when exposed to water in which the predator had been feeding. The data also showed that the effect of this invertebrate predator was more pronounced in small prey instars and small prey species that were more vulnerable to predation than large prey.     

Predation, individual variability and vertebrate population dynamics

Both predation and individual variation in life history traits influence population dynamics. Recent results from laboratory predator–prey systems suggest that differences between individuals can also influence predator–prey dynamics when different genotypes experience different predation-associated mortalities. Despite the growing number of studies in this field, there is no synthesis identifying the overall importance of the interactions between predation and individual heterogeneity and their role in shaping the dynamics of free-ranging populations of vertebrates. We aim to fill this gap with a review that examines how individual variability in prey susceptibility, in predation costs, in predator selectivity, and in predatory performance, might influence prey population dynamics. Based on this review, it is clear that (1) predation risk and costs experienced by free-ranging prey are associated with their phenotypic attributes, (2) many generalist predator populations consist of individual specialists with part of the specialization associated with their phenotypes, and (3) a complete understanding of the population dynamic consequences of predation may require information on individual variability in prey selection and prey vulnerability. Altogether, this work (1) highlights the importance of maintaining long-term, detailed studies of individuals of both predators and prey in contrasting ecological conditions, and (2) advocates for a better use of available information to account for interactive effects between predators and their prey when modelling prey population dynamics.     

Decomposing Predation: Testing for Parameters that Correlate with Predatory Performance by a Social Bacterium

Predator–prey interactions presumably play major roles in shaping the composition and dynamics of microbial communities. However, little is understood about the population biology of such interactions or how predation-related parameters vary or correlate across prey environments. Myxococcus xanthus is a motile soil bacterium that feeds on a broad range of other soil microbes that vary greatly in the degree to which they support M. xanthus growth. In order to decompose predator–prey interactions at the population level, we quantified five predation-related parameters during M. xanthus growth on nine phylogenetically diverse bacterial prey species. The horizontal expansion rate of swarming predator colonies fueled by prey lawns served as our measure of overall predatory performance, as it incorporates both the searching (motility) and handling (killing and consumption of prey) components of predation. Four other parameters—predator population growth rate, maximum predator yield, maximum prey kill, and overall rate of prey death—were measured from homogeneously mixed predator–prey lawns from which predator populations were not allowed to expand horizontally by swarming motility. All prey species fueled predator population growth. For some prey, predator-specific prey death was detected contemporaneously with predator population growth, whereas killing of other prey species was detected only after cessation of predator growth. All four of the alternative parameters were found to correlate significantly with predator swarm expansion rate to varying degrees, suggesting causal interrelationships among these diverse predation measures. More broadly, our results highlight the importance of examining multiple parameters for thoroughly understanding the population biology of microbial predation.     

Variation in foraging success among predators and its implications for population dynamics

The effects of the expected predation rate on population dynamics have been studied intensively, but little is known about the effects of predation rate variability (i.e., predator individuals having variable foraging success) on population dynamics. In this study, variation in foraging success among predators was quantified by observing the predation of the wolf spider Pardosa pseudoannulata on the cricket Gryllus bimaculatus in the laboratory. A population model was then developed, and the effect of foraging variability on predator–prey dynamics was examined by incorporating levels of variation comparable to those quantified in the experiment. The variability in the foraging success among spiders was greater than would be expected by chance (i.e., the random allocation of prey to predators). The foraging variation was density‐dependent; it became higher as the predator density increased. A population model that incorporates foraging variation shows that the variation influences population dynamics by affecting the numerical response of predators. In particular, the variation induces negative density‐dependent effects among predators and stabilizes predator–prey dynamics.     

Evolution of virulence driven by predator-prey interaction: Possible consequences for population dynamics

The evolution of pathogen virulence in natural populations has conventionally been considered as a result of selection caused by the interactions of the host with its pathogen(s). The host population, however, is generally embedded in complex trophic interactions with other populations in the community, in particular, intensive predation on the infected host can increase its mortality, and this can affect the course of virulence evolution. Reciprocally, in the long run, the evolution of virulence within an infected host can affect the patterns of population dynamics of a predator consuming the host (e.g. resulting in large amplitude oscillations, causing a severe drop in the population size, etc.). Surprisingly, neither the effect of predation on the evolution of virulence within a host, nor the influence of the evolution of virulence upon the consumer's dynamics has been addressed in the literature yet. In this paper, we consider a classical S-I ecoepidemiological model in which the infected host is consumed by a predator. We are particularly interested in the evolutionarily stable virulence of the pathogen in the model and its dependence upon ecologically relevant parameters. We show that predation can prominently shift the evolutionarily stable virulence towards more severe strains as compared to the same system without predation. We demonstrate that the evolution of virulence can result in a succession of dynamical regimes and can even lead to the extinction of the predator in the long run. The presence of a predator can indirectly affect the evolution within its prey since the evolutionarily stable virulence becomes a function of the prey growth rate, which would not be the case in a predator-free system. We find that the evolutionarily stable virulence largely depends on the carrying capacity K of the prey in a non-monotonous way. The model also predicts that in an eutrophic environment the shift of virulence towards evolutionarily stable benign strains can cause demographically stochastic evolutionary suicide, resulting in the extinction of both species, thus artificially maintaining severe strains of pathogen can enhance the persistence of both species.