Local interactions between predators and prey call into question commonly used functional responses

Commonly used functional response models (Holling’s type I and type II models) assume that the encounter rate of a predator increases linearly with prey density, provided that the predator is searching for prey. In other other words, aN (a is the baseline encounter rate and N is prey density) describes the encounter rate. This study examined whether the models are adequate when predators and prey interact locally by using a spatially explicit individual based model because local interactions affect the spatial distribution of predators and prey, which also affects the encounter rate. Predators were assumed to possess a spatial perception range that influenced their foraging behavior (e.g., if a prey is in the perception range, the predator moves towards the prey). The effect of antipredator behavior by prey was also examined. The results suggest that prey and predator densities as well as handling time affect the baseline rate (i.e., parameter a) as opposed to the common assumption that the parameter is constant. The nature of model deviations depended on both the antipredator behavior and the predators’ perception range. Understanding these deviations is important as they qualitatively affect community dynamics.     

Predation-affected spatial pattern changes in a prey population

Predation-affected spatial pattern changes in a prey population were studied. The spatial pattern of a population of Galleria mellonella is changed by its predator, Podisus maculiventris. The pattern is affected by the frequency of attack by the predator (attack ability), the homogeneity of the attack ability within a predator population and the mobility of the predator. A mathematical model incorporating these three factors was constructed, and several computer simulations were conducted by changing the parameter values. A natural enemy population having high and homogeneous attack ability and high mobility effectively kills prey individuals.     

The influence of intraguild predation on prey suppression and prey release: a meta-analysis

Intraguild predation (IGP) occurs when one predator species consumes another predator species with whom it also competes for shared prey. One question of interest to ecologists is whether multiple predator species suppress prey populations more than a single predator species, and whether this result varies with the presence of IGP. We conducted a meta-analysis to examine this question, and others, regarding the effects of IGP on prey suppression. When predators can potentially consume one another (mutual IGP), prey suppression is greater in the presence of one predator species than in the presence of multiple predator species; however, this result was not found for assemblages with unidirectional or no IGP. With unidirectional IGP, intermediate predators were generally more effective than the top predator at suppressing the shared prey, in agreement with IGP theory. Adding a top predator to an assemblage generally caused prey to be released from predation, while adding an intermediate predator caused prey populations to be suppressed. However, the effects of adding a top or intermediate predator depended on the effectiveness of these predators when they were alone. Effects of IGP varied across different ecosystems (e.g., lentic, lotic, marine, terrestrial invertebrate, and terrestrial vertebrate), with the strongest patterns being driven by terrestrial invertebrates. Finally, although IGP theory is based on equilibrium conditions, data from short-term experiments can inform us about systems that are dominated by transient dynamics. Moreover, short-term experiments may be connected in some way to equilibrium models if the predator and prey densities used in experiments approximate the equilibrium densities in nature.     

Using the functional response of a consumer to predict biotic resistance to invasive prey

Predators sometimes provide biotic resistance against invasions by nonnative prey. Understanding and predicting the strength of biotic resistance remains a key challenge in invasion biology. A predator's functional response to nonnative prey may predict whether a predator can provide biotic resistance against nonnative prey at different prey densities. Surprisingly, functional responses have not been used to make quantitative predictions about biotic resistance. We parameterized the functional response of signal crayfish (Pacifastacus leniusculus) to invasive New Zealand mud snails (Potamopyrgus antipodarum; NZMS) and used this functional response and a simple model of NZMS population growth to predict the probability of biotic resistance at different predator and prey densities. Signal crayfish were effective predators of NZMS, consuming more than 900 NZMS per predator in a 12-h period, and Bayesian model fitting indicated their consumption rate followed a type 3 functional response to NZMS density. Based on this functional response and associated parameter uncertainty, we predict that NZMS will be able to invade new systems at low crayfish densities (< 0.2 crayfish/m2) regardless of NZMS density. At intermediate to high crayfish densities (> 0.2 crayfish/m2), we predict that low densities of NZMS will be able to establish in new communities; however, once NZMS reach a threshold density of -2000 NZMS/m2, predation by crayfish will drive negative NZMS population growth. Further, at very high densities, NZMS overwhelm predation by crayfish and invade. Thus, interacting thresholds of propagule pressure and predator densities define the probability of biotic resistance. Quantifying the shape and uncertainty of predator functional responses to nonnative prey may help predict the outcomes of invasions.     

Incorporating consumer–resource spatial interactions in reserve design

The persistence of species in reserves depends in large part on the persistence of functional ecological interactions. Despite their importance, however, ecological interactions have not yet been explicitly incorporated into conservation prioritization methods. We develop here a general method for incorporating consumer–resource interactions into spatial reserve design. This method protects spatial consumer–resource interactions by protecting areas that maintain the connectivity between the distribution of consumers and resources. We illustrate our method with a conservation planning case study of a mammalian predator, American marten (Martes americana), and its two primary prey species, Red-backed vole (Clethrionomys rutilus) and Deer mouse (Peromyscus maniculatus). The conservation goal was to identify a reserve for marten that comprised 12% of a forest management unit in the boreal forest in Québec, Canada. We compared reserves developed using analysis variants that utilized different levels of information about predator and prey habitat distributions, species-specific connectivity requirements, and interaction connectivity requirements. The inclusion of consumer–resource interactions in reserve-selection resulted in spatially aggregated reserves that maintained local habitat quality for the species. This spatial aggregation was not induced by applying a qualitative penalty for the boundary length of the reserve, but rather was a direct consequence of modelling the spatial needs of the interacting consumer and resources. Our method for maintaining connectivity between consumers and their resources within reserves can be applied even under the most extreme cases of either complete spatial overlap or complete spatial segregation of consumer–resource distributions. The method has been made available via public software.     

Predation affects the susceptibility of hard clam Meretrix lusoria to Hg-stressed birnavirus

Predator–prey interaction in aquatic ecosystem is one of the simplest drivers affecting the species population dynamics. Predation controls are recognized as important aspects of ecosystem husbandry and management. In this paper we investigated how predation control cause an increase in host growth in the abundance of hard clam (Meretrix lusoria) populations subject to mercury (Hg)-stressed birnavirus. Here we linked predator–prey relationships with a bioenergetic matrix population model (MPM) associated with a susceptible–infectious–mortality (SIM) model based on a host–pathogen–predator framework to quantify the predator effects on population dynamics of disease in hard clam populations. Our results indicated that relative high predation rates could promote the hard clam abundances in relation to predators that selectively captured the infected hard clam, by which the disease transmission was suppressed. The results also demonstrated that predator-induced modifications in host behavior could have potential negative or positive effects on host growth depending on relative species density and resource dynamics. The most immediate implication of this study for the management of aquatic ecosystem is that, beyond the potential for causing a growth in abundance, predation might provoke greater predictability in aquatic ecosystem species populations and thereby increase the safety of ecosystem production from stochastic environmental events.     

Associations between the abundance of piscivorous fishes and their prey on coral reefs: implications for prey-fish mortality

Few studies have examined predator-prey relationships in diverse communities such as those found on coral reefs. Here we examined patterns of spatial and temporal association between the local abundance of predator and prey fishes at Lizard Island on the Great Barrier Reef, Australia. We predicted that the nature of this association would have implications for patterns of prey-fish mortality. Strong positive relationships between prey and piscivore abundance were found throughout the study. Greater densities of predators and of prey were found on patch-reef habitats, compared with contiguous reef-slope habitats. Declines in prey-fish abundance on patch reefs were density-dependent and correlated with the densities of predators. The relative roles of recruitment and piscivore movement in determining patterns of predator and prey abundance were assessed from surveys of recruit densities and an intensive programme of tagging two species of rock-cod, Cephalopholis cyanostigma and C. boenak (Serranidae), over 2 years. Patterns of recruitment explained little of the variation in the abundance and distribution of piscivorous fish. If movement explains large-scale patterns of distribution, this was not evident from the tagging study. The two rock-cod species were highly sedentary, with individuals on patch reefs seldom moving among reefs. Individuals on reef slopes were also highly site-attached, although they moved greater distances than those on patch reefs. Although the mechanisms responsible remain to be determined, this study demonstrated strong associations between the abundance of piscivorous fish and their prey on coral reefs. This relationship appeared to be an important factor in producing density-dependent declines in the abundance of prey. Received: 30 April 2000 / Accepted: 22 September 2000     

The feeding behavior of Beroe ovata

Patches of the neritic ctenophores Beroe ovata and Bolinopsis vitrea were observed on the edge of the Great Bahama Bank in order to determine the interaction between the predator Beroe ovata and its prey Bolinopsis vitrea. Laboratory experiments on Beroe ovata showed that it responds chemokinetically to the presence of its prey; as it swims it collides with other etenophores on which it preys. The unique use of macrocilia as cutting implements aids the predator in removing tissue from its prey, yielding maximum gain from each encounter. By means of direct observations while diving, data on horizontal and vertical patchiness, swimming speeds, abundance, and feeding behavior were collected, and used to assess the impact of populations of the predator on its prey. Together, these two animal populations form an ecological feed-back system which affects other portions of the planktonic community.     

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

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

The influence of initial age structure on predator–prey interaction

Initial age structure influences the growth of a prey population and the outcome of the predator–prey interaction. In order to quantify that influence, we employed a simple numerical model using experimental data from the system Tetranychus urticae–Phytoseiulus persimilis. Four major points were drawn from the results: (1) A population created by young females grows much faster than a population created by the same number of females but distributed among the stable age structure. Final number of individuals after a few generations is then much higher than what a plant could support. Consequently, a stable age structure is probably never achieved under these conditions; (2) In the presence of a predator, such a population can persist for a sufficiently long time to overexploit its host plant and to produce enough individuals to allow dispersal; (3) The impact of the predator on the prey population is drastically different according to its own age structure at the beginning of the interaction; and (4) Predators disturb the prey age structure during the course of interactions and thus maintain the prey growth potential at a high level. These points constitute an important adaptation that determine the persistence of the prey and the predator at a metapopulation level. They bring a new insight on the adaptive characters of young female dispersal.     

Geostatistical analysis on the spatial pattern of Pinus tabulaeformis population at different habitats in the Lingkong Mountain北大核心CSCD

In order to reveal the distribution pattern and spatial correlation of Pinus tabulaeformis population, this research used geostatistical theory and methods to examine the distribution pattern and spatial correlation of P. tabulaeformis population in different habitats of the Lingkong Mountain. The results showed that the semivariograms well fitted the exponential model for P. tabulaeformis populations at the mountain ridge, and the spherical model for those in slow slopes and valley areas. All three populations showed aggregated pattern. The spatial heterogeneity of the habitats of ridge, slow slope and valley was mostly due to the spatial autocorrelation in the ranges of 11.16 m, 9.99 m and 4.74 m, respectively. The populations at the ridges and gentle slopes showed greater sill values and determinant coefficient but lower fractal dimension than the pines in valleys, indicating that the first two habitats are characterized by higher degree of spatial heterogeneity and more significant spatial patterns, which is not found in the valley populations. Therefore the variations in the spatial distribution patterns of P. tabulaeformis populations is likely resulted from both the intrinsic biological characteristics and the environmental factors.     

Predator and prey space use: dragonflies and tadpoles in an interactive game

Predator and prey spatial distributions have important population and community level consequences. However, little is known either theoretically or empirically about behavioral mechanisms that underlie the spatial patterns that emerge when predators and prey freely interact. We examined the joint space use and behavioral rules governing movement of freely interacting groups of odonate (dragonfly) predators and two size classes of anuran (tadpole) prey in arenas containing two patches with different levels of the prey's resource. Predator and prey movement and space use was quantified both when they were apart and together. When apart from predators, large tadpoles strongly preferred the high resource patch. When apart from prey, dragonflies weakly preferred the high resource patch. When together, large prey shifted to a uniform distribution, while predators strongly preferred the high resource patch. These patterns qualitatively fit the predictions of several three trophic level, ideal free distribution models. In contrast, the space use of small prey and predators did not deviate from uniform. Three measures of joint space use (spatial correlations, overlap, and co-occurrence) concurred in suggesting that prey avoidance of predators was more important than predator attraction to prey in determining overall spatial patterns. To gain additional insight into behavioral mechanisms, we used a model selection approach to identify behavioral movement rules that can potentially explain the observed, emergent patterns of space use. Prey were more likely to leave patches with more predators and more conspecific competitors; resources had relatively weak effects on prey movements. In contrast, predators were more likely to leave patches with low resources (that they do not consume) and more competing predators; prey had relatively little effect on predator movements. These results highlight the importance of investigating freely interacting predators and prey, the potential for simple game theory models to predict joint spatial distributions, and the utility of using model choice methods to identify potential key factors that govern movement.     

Feeding ecology of a size-structured predator population,the South American sun-star Heliaster helianthus

Patterns of feeding in a population of Heliaster helianthus (Lamarck), a common and dominant species of starfish indigenous to the Pacific South American coast, were investigated in an intertidal habitat in central Peru from October 1986 to April 1987. The H. helianthus population comprised individuals of 3.5 to 30.2 cm body size (diameter) with two modal size classes. The number of rays ranged between 18 and 40, and individuals with 31 to 33 rays accounted for ca. 42% of the total population. There was a higher rate of increase in ray number with body size amongst small individuals(<13.0 cm diam). H. helianthus is capable of feeding on more than one prey item at a time (average of 5.6 to 13.2 prey items handled, with several predators observed to hold >100), and both the number of prey individuals captured and the total prey biomass were significantly correlated with predator size. Amongst a total of 1132 feeding observations, the largest number of predators (an average of 85.4% of those feeding) were preying on the mussel Semimytilus algosus whilst another mussel, Perumytilus purpuratus, ranked second with 21,9% of predators feeding. The proportion of S. algosus in the diet increased from 65.4% in the smallest predator size-group (10.9 cm diam) to 91.2% in the largest (19.0 cm). In contrast, P. purpuratus and barnacles were more highly represented in the diet of small H. helianthus. The smallest size-group (10.9 cm) had low dietary overlap with larger sizes and less specialized prey utilization. Two geographically separated populations of H. helianthus in Peru and Chile showed contrasting patterns of prey utilization. S. algosus and P. purpuratus comprised 85.5 and 6.5% by number in the diet of the Peruvian population, respectively, whilst corresponding figures for the Chilean population were 8.3 and 60.5%, with barnacles attaining a higher share (22.6%). However, the total number of prey individuals per feeding predator was almost the same in Peru and Chile, with 10.0 and 10.7 individuals, respectively. H. helianthus individuals of different sizes occupy slightly different microhabitats within the intertidal area, which, coupled with differential spatial distribution of prey species, results in the predator population being able to utilize a wide range of resources.     

Risk vs. reward: how predators and prey respond to aging olfactory cues

Many animals use olfaction to find food and avoid predators, and must negotiate environments containing odors of varying compositions, strengths, and ages to distinguish useful cues from background noise. Temporal variation in odor cues (i.e., “freshness”) seems an obvious way that animals could distinguish cues, yet there is little experimental evidence for this phenomenon. Fresh cues provide a more reliable indicator of donor presence than aged cues, but we hypothesize that the benefits of responding to aged cues depend on whether the cue indicates the proximity of a predator or a potential meal. As prey cannot remain eternally risk averse in response to predator odor, we predict that antipredator responses should diminish as predator cues age. In contrast, animals searching for food should investigate aged prey cues if investigation costs are sufficiently low and the potential benefit (a meal) sufficiently high; thus, we predict that predators will maintain interest in aged prey cues. We tested these ideas using free-ranging rats (Rattus spp.) in two separate experiments; firstly assessing giving-up densities in the presence of predator odor, and secondly examining investigation rates of prey odors. As predicted, giving-up densities dropped once predator odor had aged, but investigation rates remained similar for aged and fresh prey odor. Thus, rats used temporal variation in odor cues to evaluate the cost–benefit relationship of responding to predator and prey odors. We suggest that the ecological significance of variable cue age needs more research and should be considered when interpreting behavioral responses to olfactory information.     

Effects of predation and habitat structure on the abundance and population structure of the rock shrimp Rhynchocinetes typus (Caridea) on temperate rocky reefs

Human disturbances, such as overfishing, may disrupt predator–prey interactions and modify food webs. Underwater surveys were carried out at six shallow-water reef barrens in temperate waters of northern-central Chile from October to December 2010 to describe the effects of predation, habitat complexity (low, medium and high) and refuge availability on the abundance and population structure of the rock shrimp Rhynchocinetes typus (Rhynchocinetidae), an important mesoconsumer on subtidal hard substrata. Three sites were within managed (restricted access) areas for fishermen, and three were unmanaged (open-access). Field observations and tethering experiments were conducted to examine the relationship between fish and shrimp abundances, and the relative predation rates on shrimps. Direct effects of predation on R. typus body-size distribution were examined from shrimps collected in the field and fish stomachs. The presence and the abundance of R. typus increased with habitat reef complexity and refuge availability. Shrimp abundance was negatively related to fish abundance in managed areas, but not in open-access areas, where shrimp densities were the highest. Also, predation rates and body-size distribution of shrimps were unrelated, although fish consumed more large shrimps than should be expected from their distribution in the field. R. typus occurred most often in shelters with wide openings, offering limited protection against predators, but providing potential aggregation sites for shrimps. Overall, direct effects of predation on shrimp densities and population structure were weak, but indirect effects on shrimp distribution within reefs appear to have been mediated through behavioural responses. Our study highlights the need to assess both numerical and behavioural responses of prey to determine the effects of predator loss on mesoconsumer populations.     

Cost-Effective Suppression and Eradication of Invasive Predators

Abstract: Introduced predators can have pronounced effects on naïve prey species; thus, predator control is often essential for conservation of threatened native species. Complete eradication of the predator, although desirable, may be elusive in budget‐limited situations, whereas predator suppression is more feasible and may still achieve conservation goals. We used a stochastic predator–prey model based on a Lotka‐Volterra system to investigate the cost‐effectiveness of predator control to achieve prey conservation. We compared five control strategies: immediate eradication, removal of a constant number of predators (fixed‐number control), removal of a constant proportion of predators (fixed‐rate control), removal of predators that exceed a predetermined threshold (upper‐trigger harvest), and removal of predators whenever their population falls below a lower predetermined threshold (lower‐trigger harvest). We looked at the performance of these strategies when managers could always remove the full number of predators targeted by each strategy, subject to budget availability. Under this assumption immediate eradication reduced the threat to the prey population the most. We then examined the effect of reduced management success in meeting removal targets, assuming removal is more difficult at low predator densities. In this case there was a pronounced reduction in performance of the immediate eradication, fixed‐number, and lower‐trigger strategies. Although immediate eradication still yielded the highest expected minimum prey population size, upper‐trigger harvest yielded the lowest probability of prey extinction and the greatest return on investment (as measured by improvement in expected minimum population size per amount spent). Upper‐trigger harvest was relatively successful because it operated when predator density was highest, which is when predator removal targets can be more easily met and the effect of predators on the prey is most damaging. This suggests that controlling predators only when they are most abundant is the “best” strategy when financial resources are limited and eradication is unlikely.     

How population dynamics shape the functional response in a one-predator-two-prey system

The type III functional response has historically been associated with switching predators; when there is a choice of prey the predator favors the more abundant prey type. Although this functional response has been found in experiments where both prey densities are manipulated, in real world studies the type II functional response is more commonly found. In modeling, the type III functional response is often used in systems where the second prey type is, implicitly, assumed to be constant. Here we define a functional response that takes into account both prey densities. This causes the functional response to show both type II and type III behavior, dependent on the interaction between the two prey densities. If we take into account population dynamics, we find a type II functional response in most cases, because predation regulates the relative prey densities. This explains why type III functional responses are found in experiments where both prey densities are manipulated, but type II functional responses occur when the feedback of population dynamics on the functional response is important. Furthermore, the results show that switching can have a stabilizing or destabilizing effect and can even lead to predator extinction.     

Prey–predator dynamics with predator switching regulated by a catabolic repression control mode

Since many predators can live under certain circumstances as saprophytes or consume more than one prey, and different enzymes are generally required for each prey or nutrient digestion, the predator must be sufficiently adaptive for effective utilization of the prey mass. Control modes as induction and repression, however, act at the level of genes and cause changes in the biosynthesis rate of these enzymes. In this work, an extension of the catabolic repression control mode from the level of genes to the level of the behavior of the predator is proposed, in order to model the balanced attack of the predator on the prey. It is demonstrated that, when the prey population has the competitive advantage over the predator (in using the common substrate), the catabolic repression mechanism favors the prey population, which dominates over the predator even at low specific dilution rate values, whereas, the stable steady or periodic coexistence state is not favored. When the predator has the competitive advantage at low substrate concentrations and the prey at high substrate concentrations, the introduction of the catabolic repression mechanism in the model again favors the stable steady state of the prey, while the coexistence region is dramatically reduced. Conversely, when the prey population has the competitive advantage at low and the predator at high substrate concentrations, dominance of prey and coexistence steady state could be favored by the catabolic repression mechanism. It is concluded that the catabolic repression control favors dominance of the prey population and, under certain circumstances, coexistence of both prey and predator populations.     

Sensitivity to assumptions in models of generalist predation on a cyclic prey

Ecological theory predicts that generalist predators should damp or suppress long-term periodic fluctuations (cycles) in their prey populations and depress their average densities. However, the magnitude of these impacts is likely to vary depending on the availability of alternative prey species and the nature of ecological mechanisms driving the prey cycles. These multispecies effects can be modeled explicitly if parameterized functions relating prey consumption to prey abundance, and realistic population dynamical models for the prey, are available. These requirements are met by the interaction between the Hen Harrier (Circus cyaneus) and three of its prey species in the United Kingdom, the Meadow Pipit (Anthus pratensis), the field vole (Microtus agrestis), and the Red Grouse (Lagopus lagopus scoticus). We used this system to investigate how the availability of alternative prey and the way in which prey dynamics are modeled might affect the behavior of simple trophic networks. We generated cycles in one of the prey species (Red Grouse) in three different ways: through (1) the interaction between grouse density and macroparasites, (2) the interaction between grouse density and male grouse aggressiveness, and (3) a generic, delayed density-dependent mechanism. Our results confirm that generalist predation can damp or suppress grouse cycles, but only when the densities of alternative prey are low. They also demonstrate that diametrically opposite indirect effects between pairs of prey species can occur together in simple systems. In this case, pipits and grouse are apparent competitors, whereas voles and grouse are apparent facilitators. Finally, we found that the quantitative impacts of the predator on prey density differed among the three models of prey dynamics, and these differences were robust to uncertainty in parameter estimation and environmental stochasticity.     

Effects of density-dependent dispersal behaviours on the speed and spatial patterns of range expansion in predator-prey metapopulations

Dispersal can strongly affect the spatiotemporal dynamics of a species (its spread, spatial distribution and persistence). We investigated how two dispersal behaviours, namely prey evasion (PE) and predator pursuit (PP), affect the dynamics of a predator-prey system. PE portrays the tendency of prey avoiding predators by dispersing into adjacent patches with fewer predators, while PP describes the tendency of predators to pursue the prey by moving into patches with more prey. Based on the Beddington predation model, a spatially explicit metapopulation model was built to incorporate PE and PP. Numerical simulations were run to investigate the effects of PE and PP on the rate of spread, spatial synchrony and the persistence of populations. Results show that both PE and PP can alter spatial synchrony although PP has a weaker desynchronising effect than PE. The predator-prey system without PE and PP expanded in circular waves. The effect of PE can push the prey to distribute in a circular ring front, whereas the effect of PP can change the circular waves to anisotropic expansion. Furthermore, weak PE and PP can accelerate the spread of prey while strong and disproportionate intensities slow down the range expansion. The effects of PE and PP further enhance the population size, break down the spatial synchrony and promote the persistence of populations.