Intermediate Predator Research Articles

Dynamics of a Stochastic Three‐Species Food Web Model with Omnivory and Ratio‐Dependent Functional Response

This paper is concerned with a stochastic three‐species food web model with omnivory which is defined as feeding on more than one trophic level. The model involves a prey, an intermediate predator, and an omnivorous top predator. First, by the stochastic comparison theorem, we show that there is a unique global positive solution to the model. Next, we investigate the asymptotic pathwise behavior of the model. Then, we conclude that the model is persistent in mean and extinct and discuss the stochastic persistence of the model. Further, by constructing a suitable Lyapunov function, we establish sufficient conditions for the existence of an ergodic stationary distribution to the model. Then, we present the application of the main results in some special models. Finally, we introduce some numerical simulations to support the main results obtained. The results in this paper generalize and improve the previous related results.

Effects of Turbidity and Acidity on Predator-Prey Interactions

Predator-prey interactions are influenced by environmental condition changes, such as increased turbidity and acidification caused by human disturbance. These anthropogenic factors can affect trophic interactions from pisciovorous fish, invertivorous fish, and shredder invertebrates to leaf litter leaching. In this study, we investigated effects of increased turbidity and acidification on predator-prey interaction through a four-level detritus-based food chain, which are top predator (pisciovorous fish), intermediate predator (invertivorous fish), shredder invertebrates, and allochthonous leaf litter. The experiment had a top predator-snakehead (Channa argus), an intermediate predator-black carp (Mylopharyngodon piceus), shredder prey-a freshwater crustacean isopods (Asellus sp.), and allochthonous leaf litter sakura (Cerasus sp.). The pisciovorous snakehead was caged, providing non-lethal predation effect on black carp. The effects of turbidity condition changes with different Nephelometric Turbidity Unit (high level: 60 NTU, and low: 10 NTU) and acidic condition changes (weakly acidic: pH 6.0 and normal: pH 7.5) on leaf litter weight loss. The experiment measured shredder density change and leaf litter weight change that was caused by both leaf litter leaching and shredder’s foraging processes. Results indicated that: the high turbid treatment (60 NTU) reduced the black carp’s antipredator defense to snakehead, so that the black carp’s foraging caused the high mortality of isopod shredders, which reduced leaf litter weight loss. By contrast, the weakly acidic treatment (pH=6.0) did not influence black carp’s top-predator avoidance, but induced the loss of predator avoidance of isopod Asellus sp. shredders that caused isopod high mortality. The acidic treatment did not influence litter weight change. Our study highlights that increased turbidity can influence predator-prey interactions and affect leaf litter weight change that caused by both shredder foraging and litter leaching processes in freshwater systems. The increased acidic condition also had a significant effect on predator-prey interaction. With intensified urbanization and increased climate change, such anthropogenic impacts with multiple stressors on predator-prey interaction should be further investigated for understanding their ecological mechanisms.

Potential for managing life history diversity in a commercially exploited intermediate predator, the goldsinny wrasse (Ctenolabrus rupestris)

Potential for managing life history diversity in a commercially exploited intermediate predator, the goldsinny wrasse (Ctenolabrus rupestris)

Potential for managing life history diversity in a commercially exploited intermediate predator, the goldsinny wrasse (Ctenolabrus rupestris)

Abstract Small-bodied wrasse species are important for structuring coastal marine ecosystems but are also increasingly harvested as parasite cleaners on farmed salmon. Identifying management regulations that will support long-term sustainability of wrasse fisheries is challenging, because there is still limited knowledge about the impacts of fisheries on the demography of these intermediate predators in their natural environments. To this end, we studied individual growth histories of goldsinny wrasse (Ctenolabrus rupestris) at a fine spatial scale across replicated marine protected areas (MPAs) and areas open to commercial harvesting on the Norwegian coast. The MPAs were established 1–7 years prior to our sampling. We detected significant fine-scale spatial variation in wrasse asymptotic body size, but found no consistent difference between MPAs and fished areas. Male wrasses reached larger asymptotic body sizes than females, whereas fyke nets captured individuals with larger asymptotic body sizes compared with baited traps. These are the two commonly used gear types in wrasse fisheries. An extended use of baited traps, along with slot-size limits, could therefore aid in protecting large-growing phenotypes such as nest-guarding males.

Distribution Patterns of Odonate Assemblages in Relation to Environmental Variables in Streams of South Korea.

Odonata species are sensitive to environmental changes, particularly those caused by humans, and provide valuable ecosystem services as intermediate predators in food webs. We aimed: (i) to investigate the distribution patterns of Odonata in streams on a nationwide scale across South Korea; (ii) to evaluate the relationships between the distribution patterns of odonates and their environmental conditions; and (iii) to identify indicator species and the most significant environmental factors affecting their distributions. Samples were collected from 965 sampling sites in streams across South Korea. We also measured 34 environmental variables grouped into six categories: geography, meteorology, land use, substrate composition, hydrology, and physicochemistry. A total of 83 taxa belonging to 10 families of Odonata were recorded in the dataset. Among them, eight species displayed high abundances and incidences. Self-organizing map (SOM) classified sampling sites into seven clusters (A–G) which could be divided into two distinct groups (A–C and D–G) according to the similarities of their odonate assemblages. Clusters A–C were characterized by members of the suborder Anisoptera, whereas clusters D–G were characterized by the suborder Zygoptera. Non-metric multidimensional scaling (NMDS) identified forest (%), altitude, and cobble (%) in substrata as the most influential environmental factors determining odonate assemblage compositions. Our results emphasize the importance of habitat heterogeneity by demonstrating its effect on odonate assemblages.

Intermediate predator na\xefvet\xe9 and sex-skewed vulnerability predict the impact of an invasive higher predator

The spread of invasive species continues to reduce biodiversity across all regions and habitat types globally. However, invader impact prediction can be nebulous, and approaches often fail to integrate coupled direct and indirect invader effects. Here, we examine the ecological impacts of an invasive higher predator on lower trophic groups, further developing methodologies to more holistically quantify invader impact. We employ functional response (FR, resource use under different densities) and prey switching experiments to examine the trait- and density-mediated impacts of the invasive mosquitofish Gambusia affinis on an endemic intermediate predator Lovenula raynerae (Copepoda). Lovenula raynerae effectively consumed larval mosquitoes, but was naïve to mosquitofish cues, with attack rates and handling times of the intermediate predator unaffected by mosquitofish cue-treated water. Mosquitofish did not switch between male and female prey, consistently displaying a strong preference for female copepods. We thus demonstrate a lack of risk-reduction activity in the presence of invasive fish by L. raynerae and, in turn, high susceptibility of such intermediate trophic groups to invader impact. Further, we show that mosquitofish demonstrate sex-skewed predator selectivity towards intermediate predators of mosquito larvae, which may affect predator population demographics and, perversely, increase disease vector proliferations. We advocate the utility of FRs and prey switching combined to holistically quantify invasive species impact potential on native organisms at multiple trophic levels.

Interactions among predators and plant specificity protect herbivores from top predators.

The worldwide loss of top predators from natural and agricultural systems has heightened the need to understand how important they are in controlling herbivore abundance. The effect of top predators on herbivore species is likely to depend on (1) the importance of the consumption of intermediate predators by top predators (intra-guild predation; IGP), but also on (2) plant specificity by herbivores, because specialists may defend themselves better (enemy-free space; EFS). Insectivorous birds, as top predators, are generally known to effectively control herbivorous insects, despite also consuming intermediate predators such as spiders, but how this effect varies among herbivore species in relation to the cascading effects of IGP and EFS is not known. To explore this, we excluded birds from natural fynbos vegetation in South Africa using large netted cages and recorded changes in abundance relative to control plots for 199 plant-dwelling intermediate predator and 341 herbivore morpho-species that varied in their estimated plant specificity. We found a strong negative effect of birds on the total abundance of all intermediate predators, with especially clear effects on spiders (strong IGP). In contrast with previous studies, which document a negative effect of birds on herbivores, we found an overall neutral effect of birds on herbivore abundance, but the effect varied among species: some species were negatively affected by birds, suggesting that they were mainly consumed by birds, whereas others, likely released from spiders by IGP, were positively affected. Some species were also effectively neutrally affected by birds. These tended to be more specialized to plants compared to the other species, which may imply that some plant specialists benefited from protection provided by EFS from both birds and spiders. These results suggest that the response of herbivore species to top predators may depend on cascading effects of interactions among predators and on their degree of plant specificity.

Past selection impacts the strength of an aquatic trophic cascade

Abstract In complex food webs, interactions among species in different trophic levels can generate cascading indirect effects that couple top predators with primary producers, thereby affecting ecosystem functioning. Natural selection imposed by top predators on intermediate predators may play a role in shaping the strength of these trophic cascades, but this conjecture remains largely untested. To determine the effects of natural selection on the strength of trophic cascades, we conducted a two‐part experiment in a four‐level aquatic trophic system involving a top predator (fish), an intermediate predator (damselflies), herbivores (zooplankton) and primary producers (algae). We first quantified how predation by fish generated selection on damselfly activity levels after controlling for phenotypic plasticity. We then measured the indirect effects of this selection on primary production (phytoplankton biomass). In both experiments, we varied the density of predators, allowing us to elucidate both trait‐mediated and density‐mediated indirect effects. We found that as fish density increased, damselfly survivorship declined, which generated natural selection favouring less active damselflies. These results are robust after taking into account latent effects of plasticity in response to fish predator cues. The surviving damselflies likely foraged less, freeing herbivores from predation, which in turn reduced primary production. This selection‐driven trait‐mediated indirect effect was only apparent at low damselfly densities, because the consumptive effect of damselflies at high densities overwhelmed the effects of past selection. These results demonstrate how the past action of natural selection imposed by predators affects the phenotypes of prey that consume herbivores, which ultimately influences primary productivity—a selective trophic cascade. Natural selection can therefore act as a mechanism coupling ecological dynamics across trophic levels, which ultimately influences ecosystem functioning. A plain language summary is available for this article.

An analytical approach to top predator interference on the dynamics of a food chain model

In this paper, a nonlinear mathematical model is proposed and analyzed to study of top predator interference on the dynamics of a food chain model. The mathematical model is formulated using the system of non-linear ordinary differential equations. In this model, there are three state dimensionless variables, viz, size of prey population x, size of intermediate predator y and size of top predator population z. The analytical results are compared with the numerical simulation using MATLAB software and satisfactory results are noticed.

Complex dynamics of a three species prey-predator model with intraguild predation

‘Omnivory’, a widespread mechanism in interacting populations, is defined as feeding on more than one trophic levels. In this work we have considered a three species model consisting of prey, intermediate predator that predates upon prey and top predator with intraguild predation. It is well known that intraguild predation destabilizes the food webs consisting of three and more trophic levels, and induces chaotic oscillation for Holling type-I functional response when the intensity of the intraguild predation becomes low. Here we have considered Holling type-II functional response between intermediate predator and intraguild predator and other functional responses are assumed to be linear. First we investigate local stability and bifurcation analysis of all axial and boundary equilibrium points. We observe two types of coexistence of three species - a steady-state coexistence and an oscillatory coexistence. Although it is difficult to find the analytical expressions for stability and bifurcation of the interior equilibrium point, we have verified numerically that a stable limit cycle bifurcates from the interior equilibrium point and ultimately chaos occurs via successive period doubling bifurcations. Bifurcation diagrams have been drawn with respect to all the system parameters to explore the complete dynamics of the model. Consideration of saturating functional response instead of law of mass action can suppress the chaotic oscillation leading to stable coexistence of three species at their steady-state.

Dynamics Analysis of Anti-predator Model on Intermediate Predator With Ratio Dependent Functional Responses

This articel discusses a predator prey model with anti-predator on intermediate predator using ratio dependent functional responses. Dynamical analysis performed on the model includes determination of equilibrium point, stability and simulation. Three kinds of equilibrium points have been discussed, namely the extinction of prey point, the extinction of intermediate predator point and the extinction of predator point are exists under certain conditions. It can be shown that the result of numerical simulations are in accordance with analitical results

Stabilizing effect of intra-specific competition on prey-predator dynamics with intraguild predation

Multi-species population models with various types of interactions are important to understand the stable coexistence of prey and predator species and the role of various factors leading to complex dynamics and extinction of one or more species. Models of two predators sharing one prey population are analyzed by some researchers and the role of nonlinear functional responses and interaction strengths are reported to be responsible for the stable coexistence of three species. However, one-prey-two-predator models with intraguild predation have received relatively less attention from researchers. Recent literature on this topic mainly focused on the study of local and global stability properties of possible equilibria and appearance of various types of bifurcations. Intra-specific competition among predator species is an important factor for coexistence of more than one predator species competing for one shared prey. Hence it is quite essential to understand the role of intra-specific competition on the stability and bifurcation behavior. In this work we wish to demonstrate the stabilizing role of intra-specific competition among intermediate and top predators when the growth rate of prey species is adequate to support both the predator species. With the help of stability and bifurcation analysis we show the stabilizing role of intra-specific competition among predators. Moreover, strong self-regulation in intraguild predator prevents extinction of intraguild prey at high productivity and resolve the contradiction between empirical and previous theoretical results on the fate of intraguild prey at high productivity. To establish our claim we present, apart from analytical results, extensive numerical simulation results and couple of bifurcation diagrams.

Trophic cascades and the transient keystone concept

Apex predator reintroductions are commonly motivated by the imperative to restore populations and wider ecosystem function by precipitating trophic cascades that release basal species. Yet evidence for the existence of such cascades is often equivocal, particularly where consumptive interactions between apex and intermediate predators are weak or absent. Here, using a tri-trophic skate-crab-bivalve study-system, we find that non-consumptive interactions between apex skate and intermediate crabs cascade down to consumptive interactions between crabs and bivalves, significantly reducing bivalve mortality. However, skate only functioned as keystone where crabs foraged for bivalves in the absence of mature bivalve reef: where reef was present, bivalve mortality was not significantly different in the presence or absence of skate. By facilitating the establishment of basal species which, in turn, diminish apex-intermediate effects, the skate's keystone function is subject to negative regulation. Thus, we propose that keystone functionality can be transient with respect to environmental context. Our findings have two central implications for apex predator reintroductions and basic ecology: (i) species hitherto not considered as keystone may have the capacity to act as such transiently, and; (ii) keystones are known to regulate ecosystems, but transience implies that ecosystems can regulate keystone function.

Refuge increases food chain length: modeled impacts of littoral structure in lake food webs

Food chain length (FCL) represents a fundamental metric within ecology because it has implications for ecosystem function and responses to environmental change. Omnivory between linked food chains situated within large ecosystems can increase FCL, whereas overlap of food chains within small or spatially compressed ecosystems is generally thought to decrease FCL. Yet FCL varies widely in small ecosystems and the mechanisms underlying determinants of FCL in these systems is unclear. In small shallow lakes, littoral structure is a predictor of FCL but it is unclear whether this is due to productivity or refuge mechanisms. Here we provide evidence, using consumer resource food web modules parameterized with empirical data, that refuge in spatially compressed ecosystems has the ability on its own to increase the trophic position of top predators by increasing the biomass of top and intermediate predators across a range of common food web module structures. Our results suggest that refuge is an important driver of FCL in small ecosystems, which has implications for determining responses of these systems to environmental change.

Feeding ecology of two sympatric species of large-sized groupers (Perciformes: Epinephelidae) on Southwestern Atlantic coralline reefs

ABSTRACT Red and black groupers are large-bodied opportunistic ambush predators commonly found in Southwestern Atlantic tropical reefs. We investigated the diet of both species in order to detail ontogenetic, spatial and temporal trends, and to assess the extent of overlap in resource use between these two sympatric predators on the Abrolhos Bank, Brazil. Decapods and fishes were the main food items of Epinephelus morio while fishes were the main prey of Mycteroperca bonaci. Both diets were significantly influenced by body size and habitat, but only smaller individuals of E. morio feed almost exclusively on crustaceans. While the two groupers rely on many of the same prey types, coexistence may be facilitated by E. morio feeding more heavily on crustaceans, particularly the blackpoint sculling crab Cronius ruber, while black grouper take comparatively few crustaceans but lots of fish prey. Predators like red and black groupers could trigger indirect effects in the community and influence a large range of ecological processes, such as linkages between top and intermediate predators, and intermediate predators and their resources.

Anthropogenic and natural determinants of fish food-chain length in a midsize river system

AbstractFood-chain length (FCL) is a key factor underlying patterns of energy flow and trophic structure in food webs. Natural gradients in resource availability, ecosystem size, and disturbance are thought to govern FCL in rivers. However, anthropogenic influences on FCL, including landscape modification (e.g., land-cover change) and river regulation (e.g., dams), remain unresolved. From 2010 to 2013 we characterized fish food webs at 12 river reaches along an ∼185-km section of the 6th-order Scioto River system (Ohio, USA) to assess the effects of natural and anthropogenic factors and the potential influence of shifts in foodweb structure (e.g., diversity, top-predator body size, intermediate predator richness) on fish FCL. Stable isotope-derived estimates of FCL ranged from 3.66 to 4.35, which are comparable to global values of FCL in rivers. A model-selection approach returned a set of linear regression models that explained 38 to 74% of the variation in FCL. Local ecosystem size (i.e., channel width)...

Impact of generalist type sexually reproductive top predator interference on the dynamics of a food chain model

Food uptake ability of higher trophic level species are more complicated and interesting due to their choice and availability of food and consequently their growth. In the present study, the effect of top predator interference on the dynamics of a tritrophic food chain model involving an intermediate and a generalist type top predator are considered. It is assumed that the interaction between the logistically growing prey and intermediate predator follows the Volterra scheme, while that between the top predator and its favorite food i.e. the intermediate predator follows Crowley–Martin type functional response. Also, it is considered that the growth of top predator is by sexual reproduction. The boundedness of the system, existence of an attracting set, local and global stability of non-negative equilibrium points and uniform persistence are established by very complicated and highly nonlinear biological considerations. With the help of empirical results from different field and laboratory experiment, it is clarified that Crowley–Martin type functional response gives better result than other predator dependent functional responses for the issue of interference among own population. Also it is clarified from literature survey that high trophic species are mostly generalist type i.e. they can survive rather than their favorite food depending upon the present environmental situation and their growth rate depends upon the mating success. Moreover, increasing the top predator interference stabilizes the system, while increasing the normalization of the residual reduction in the top predator population destabilizes the system. Different numerical examples support these ecological phenomenon.

Fragment edge and isolation affect the food web: effects on the strength of interactions among trophic guilds

Abstract Habitat loss and fragmentation are processes that may affect communities by changing species interactions. These changes occur because the strength of linkages between species is not exclusively dependent on predator and prey traits. Species interaction changes also depend on the spatial context in which they take place. We used structural equation modelling to evaluate effects of these processes at patch-scale on top-down and bottom-up controls in food webs in Atlantic Forest. The model was composed of multiple species, and trophic guilds responded differently to fragment edge and isolation. Changes in bottom-up and top-down controls were mainly related to intermediate predator interactions. Efforts to restore connectivity among fragments should help recover the equilibrium of the trophic interactions by benefiting intermediate predators.

Tri-trophic Plankton Models Revised: Importance of Space, Food Web Structure and Functional Response Parametrisation

Revealing mechanisms of efficient top-down control in eutrophic ecosystems remains a long term challenge in theoretical ecology. In this paper, we revisit the role of environmental heterogeneity, food web structure and shape of the predator functional response in persistence and stabilization of a planktonic system with high nutrient supply. We consider a 1D vertically resolved tri-trophic planktonic food web composed of a primary producer, an intermediate predator and a highly mobile top predator, such as a system of phytoplankton, microzooplankton and copepods. We explore the realistic scenario in which the top predator is omnivorous, i.e. when copepods can feed both on phytoplankton and microzooplankton. We show that the interplay between heterogeneity of the environment due to for instance, a light gradient in the water column, and trophic interaction between species can result in an efficient top-down control which would otherwise be impossible in the corresponding well-mixed system. We also find that allowing the top predator to feed on the primary producer may dramatically impede the coexistence of the three trophic levels, with only two levels generally surviving. The coexistence of all three trophic levels within a wide range of parameters becomes possible only when the top predator exhibits active food source switching behaviour. We also show the phenomenon of bistability in the considered tri-trophic food web: a small initial amount of the top predator should lead to its extinction whereas introduction of a supercritical initial amount will eventually result in establishment of the population. The demonstrated mechanism of top-down control seems to be rather generic and might be a good candidate to explain stability in some other non-planktonic tri-trophic ecosystems.

Top predators induce the evolutionary diversification of intermediate predator species

We analyze the evolutionary branching phenomenon of intermediate predator species in a tritrophic food chain model by using adaptive dynamics theory. Specifically, we consider the adaptive diversification of an intermediate predator species that feeds on a prey species and is fed upon by a top predator species. We assume that the intermediate predator׳s ability to forage on the prey can adaptively improve, but this comes at the cost of decreased defense ability against the top predator. First, we identify the general properties of trade-off relationships that lead to a continuously stable strategy or to evolutionary branching in the intermediate predator species. We find that if there is an accelerating cost near the singular strategy, then that strategy is continuously stable. In contrast, if there is a mildly decelerating cost near the singular strategy, then that strategy may be an evolutionary branching point. Second, we find that after branching has occurred, depending on the specific shape and strength of the trade-off relationship, the intermediate predator species may reach an evolutionarily stable dimorphism or one of the two resultant predator lineages goes extinct.