Bifurcations and hydra effects in Bazykin's predator-prey model.

We consider Bazykin's model to address harvesting induced stability exchanges through bifurcation analysis. We examine the existence of hydra effects and analyze the stock pattern under predator harvesting. Prey harvesting cannot produce hydra effects in our model, whereas predator harvesting may cause multiple hydra effects. Our study reveals that type II response function and mutual interference among predators jointly induce multiple hydra effects and bistability. Bifurcations such as single Hopf-bifurcation, multiple Hopf-bifurcations and multiple saddle-node bifurcations appear for increasing harvesting rate on the predators. However, over-exploitation of the predators cannot generate any such bifurcation in our study. In simulations, the maximum sustainable yield (MSY) exists at a globally stable state. When predator is culled under increasing effort, basin of attraction of the equilibrium corresponding to the higher predator stock gets expanded, which alternatively is in favor of stock benefit for predators. The ecological theory developed in this study might be useful to understand conservation policy and fishery management.

Stability switching and hydra effect in a predator-prey metapopulation model.

A metapopulation model is investigated to explore how the spatial heterogeneity affects predator-prey interactions. A Rosenzweig-MacArthur (RM) predator-prey model with dispersal of both the prey and predator is formulated. We propose such a system as a well mixed spatial model. Here, partially mixed spatial models are defined in which the dispersal of only one of the communities (prey or predator) is considered. In our study, the spatial heterogeneity is induced by dissimilar (unbalanced) dispersal rates between the patches. A large difference between the predator dispersal rates may stabilize the unstable positive equilibrium of the model. The existence of two ecological phenomena are found under independent harvesting strategy: stability switching and hydra effect. When prey or predator is harvested in a heterogenious environment, a positive stable steady state becomes unstable with increasing the harvesting effort, and a further increase in the effort leads to a stable equilibrium. Thus, a stability switching happens. Furthermore, the predator biomass (at stable state) in both the patches (and hence total predator stock) increases when the patch with a higher predator density is harvested; resulting a hydra effect. These two phenomena do not occur in the non-spatial RM model. Hence, spatial heterogeneity induces stability switching and hydra effect.

Balancing maximum sustainable yield and ecological resilience in an exploited two-predator one-prey system.

In this paper, we consider a two-predator one-prey system to determine the feedback of exploitation in individual as well as joint population levels. As balancing yield with resilience is highly essential for the conservation of species in the marine ecosystem, here we measure both the maximum sustainable yield (MSY) and resilience simultaneously. Then we investigate both the trade-offs and synergies among maximum yield, conservation, and resilience that emerge from different harvesting plans. It is found that for single species harvesting, a prey species-oriented system is capable of producing more yield in compare to any predator-oriented system but for resilience, a prey species-oriented system is far behind the others. In the case of joint harvesting of all the species, it is observed that the first predator-oriented system has a better ability to absorb the disturbances than the other cases. The correlation between yield and resilience at the MSY level is studied in all the cases. It is further observed that the increase of intraspecific competition in the predator decreases the risk of sustainability. In this way, this study may be helpful for fishery management to fulfill their goals without affecting the ecosystem's health in the long run.

Hydra effects in stable food chain models.

In this paper, we explore the occurrence of the hydra effect in food chains, a popular research theme in the current decade. The hydra effect, one of the paradoxical results in theoretical and applied ecology refers to the fact where increasing mortality rate on a population enhances its own stock. The main focus is to propose a dynamical system model of food chain showing a stable steady state and estimate the variation of stock of targeted species with increasing mortality. In our model, the per capita growth rate of any predator trophic level does not depend upon its density. The prey-predator model incorporating such a feature for predator growth is referred to as 'pure predator system' (see Sieber and Hilker (2012), J. Math. Biol. (2012) 64: 341-360, Journal of Mathematical Biology). Keeping the above feature in mind, we study a Rosenweig-MacArthur food chain model with logistic prey growth and Holling type II functional responses. It is shown that hydra effect at stable state appears on (a) prey in a four-trophic system, (b) first predator in a five-trophic system, and (c) prey and second predator in a six-trophic system. Xiao and Cao (2009) (Mathematical and Computer Modelling 50 (2009) 360-379) established that limit cycle may be observed due to harvesting in a system with the ratio-dependent prey-predator system (example of a non "non-pure predator system"). Therefore, if harvesting causes instability on some range of mortality rate, the hydra effect cannot occur at a stable state. Some results show that the unique stable steady state in our model remains stable under harvesting of either trophic level. As a whole, our investigations have some contribution in understanding population interactions, fishery management and biological pest control tactic.

Managing yield and resilience in a harvested tri-trophic food chain model.

In this article, we compare the two ecological services known as yield and resilience, for a tri-trophic food chain model consisting of a prey, an intermediate predator and a top predator. For this comparison process, we use both analytical and numerical techniques. It is shown that a variety of patterns are possible based on the intensity of efforts distributed among different trophic levels. Thus we may suggest that fishing down the food chain, as suggested by Pauly et al. (1998) is not bound to happen. Our analysis also shows that balancing the harvest between prey, intermediate predator and top predator could give more yield and stabilizing the ecosystem, than the selective harvesting of any one species. This balanced harvesting may not be a win-win situation for yield and resilience, but it could be a most favourable strategy to balance them. This research would help to correlate resilience with yield and determines the desirable selection of two policies, resilience maximizing yield or maximum sustainable yield to safeguard ecological communities.

Harvesting induced stability and instability in a tri-trophic food chain.

Non-equilibrium dynamics in the form of oscillations or chaos is often found to be a natural phenomenon in complex ecological systems. In this paper, we first analyze a tri-trophic food chain, which is an extension of the Rosenzweig-MacArthur di-trophic food chain. We then explore the impact of harvesting individual trophic levels to answer the following questions : a) when a non-equilibrium dynamics persists, b) whether it can locally be stabilized to a steady state, c) when the system switches from a stable steady state to a non-equilibrium dynamics and d) whether the Maximum Sustainable Yield (MSY) always exists when the top predator is harvested. It is shown that searching for a general theory to unify the harvesting induced stability must take into account the number of trophic levels and the degree of species enrichment, the outcomes that cannot be obtained from the earlier reports on prey-predator models. We also identify the situation where harvesting induces instability switching: the non-equilibrium state enters into a stable steady-state and then, upon more intensive harvesting, the steady-state again loses its stability. One of the new and important results is also that the MSY may not exist for harvesting the top predator. In general, our results contribute to biological conservation theory, fishery and ecosystem biodiversity management.

Biological conservation through marine protected areas in the presence of alternative stable states.

This article addresses how depleted stock can be restored by creation of marine reserve and species mobility when alternative stable states persist in a marine ecosystem. To understand the role of a marine protected area, we develop a two-patch version of an originally single-patch model. In the two-patch model, we prove that some of the locally stable equilibria are not stable equilibria from an ecological viewpoint. Similarly, some unstable equilibria determined classically from the mathematical model are no longer equilibria. It is shown that increasing reserve size may produce three alternative stable states in the presence of harvesting. Dynamic solutions have a tendency to reach an upper stable state from a lower stable state when reserve size is increased, but the opposite phenomenon (i.e., shifting to a lower stable state from an upper one) never occurs. This suggests that MPAs always have a positive effect in stock conservation even when alternative stable states inherently persist in marine ecosystems.

Relationship between exploitation, oscillation, MSY and extinction.

We give answers to two important problems arising in current fisheries: (i) how maximum sustainable yield (MSY) policy is influenced by the initial population level, and (ii) how harvesting, oscillation and MSY are related to each other in prey-predator systems. To examine the impact of initial population on exploitation, we analyze a single species model with strong Allee effect. It is found that even when the MSY exists, the dynamic solution may not converge to the equilibrium stock if the initial population level is higher but near the critical threshold level. In a prey-predator system with Allee effect in the prey species, the initial population does not have such important impact neither on MSY nor on maximum sustainable total yield (MSTY). However, harvesting the top predator may cause extinction of all species if odd number of trophic levels exist in the ecosystem. With regard to the second problem, we study two prey-predator models and establish that increasing harvesting effort either on prey, predator or both prey and predator destroys previously existing oscillation. Moreover, equilibrium stock both at MSY and MSTY level is stable. We also discuss the validity of found results to other prey-predator systems.

Maximum sustainable yield and species extinction in a prey-predator system: some new results.

Though the maximum sustainable yield (MSY) approach has been legally adopted for the management of world fisheries, it does not provide any guarantee against from species extinction in multispecies communities. In the present article, we describe the appropriateness of the MSY policy in a Holling-Tanner prey-predator system with different types of functional responses. It is observed that for both type I and type II functional responses, harvesting of either prey or predator species at the MSY level is a sustainable fishing policy. In the case of combined harvesting, both the species coexist at the maximum sustainable total yield (MSTY) level if the biotic potential of the prey species is greater than a threshold value. Further, increase of the biotic potential beyond the threshold value affects the persistence of the system.

Possible ecosystem impacts of applying maximum sustainable yield policy in food chain models.

This paper describes the possible impacts of maximum sustainable yield (MSY) and maximum sustainable total yield (MSTY) policy in ecosystems. In general it is observed that exploitation at MSY (of single species) or MSTY (of multispecies) level may cause the extinction of several species. In particular, for traditional prey-predator system, fishing under combined harvesting effort at MSTY (if it exists) level may be a sustainable policy, but if MSTY does not exist then it is due to the extinction of the predator species only. In generalist prey-predator system, harvesting of any one of the species at MSY level is always a sustainable policy, but harvesting of both the species at MSTY level may or may not be a sustainable policy. In addition, we have also investigated the MSY and MSTY policy in a traditional tri-trophic and four trophic food chain models.

Sustainability and economic consequences of creating marine protected areas in multispecies multiactivity context.

The present study deals with harvesting of prey species in the presence of predator in a multispecies marine fishery. The total habitat is divided into two patches: one is reserve area where fishing is completely banned and other zone is called fishing area where only prey is exploited. We assume that the prey fish possesses heterogeneous intrinsic growth rate with uniform carrying capacity where as predator has constant intrinsic growth rate with prey dependent carrying capacity. The analytical conditions are derived to prevent the species extinction for larger employed effort in single (only prey) species fishery. Optimal equilibrium premium are presented for both monospecies and multispecies fishery for all degree of protection. Increasing standing stock (ISS) and protected standing stock (PSS) are measured in the presence of prey-predator interaction.

Sustainability and optimal control of an exploited prey predator system through provision of alternative food to predator.

In the present paper, we develop a simple two species prey-predator model in which the predator is partially coupled with alternative prey. The aim is to study the consequences of providing additional food to the predator as well as the effects of harvesting efforts applied to both the species. It is observed that the provision of alternative food to predator is not always beneficial to the system. A complete picture of the long run dynamics of the system is discussed based on the effort pair as control parameters. Optimal augmentations of prey and predator biomass at final time have been investigated by optimal control theory. Also the short and large time effects of the application of optimal control have been discussed. Finally, some numerical illustrations are given to verify our analytical results with the help of different sets of parameters.