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

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

Variable prey development time suppresses predator–prey cycles and enhances stability

Although theoretical models have demonstrated that predator–prey population dynamics can depend critically on age (stage) structure and the duration and variability in development times of different life stages, experimental support for this theory is non‐existent. We conducted an experiment with a host–parasitoid system to test the prediction that increased variability in the development time of the vulnerable host stage can promote interaction stability. Host–parasitoid microcosms were subjected to two treatments: Normal and High variance in the duration of the vulnerable host stage. In control and Normal‐variance microcosms, hosts and parasitoids exhibited distinct population cycles. In contrast, insect abundances were 18–24% less variable in High‐ than Normal‐variance microcosms. More significantly, periodicity in host–parasitoid population dynamics disappeared in the High‐variance microcosms. Simulation models confirmed that stability in High‐variance microcosms was sufficient to prevent extinction. We conclude that developmental variability is critical to predator–prey population dynamics and could be exploited in pest‐management programs.     

The arthropod predators of ant-mimetic and aposematic prey: a serological analysis

Abstract.

• 1 ELISA (enzyme-linked immunosorbant assay) was used to identify the arthropod predators of two common herbivores of a desert lupine: the aposematic plant bug Lopidea nigridea and the ant-mimetic plant bug Coquillettia insignis.
• 2 Despite the fact that the prey are closely related, the two antisera were sufficiently specific to distinguish between the antigen against which they were formed and the antigen of the other species: both antisera were 3 times as reactive against their homologous antigen as they were against their heterologous antigen.
• 3 In tests on gut contents of field-collected predators, ELISA results were generally consistent with laboratory no-choice data for the five most common arthropod species on lupine: there was a strong correlation between attack rates for each predator-prey pair as measured in the laboratory, and per cent positive reactions in ELISA as measured using field-collected predators. In particular, L.nigridea antiserum was shown to be significantly less reactive against the crab spider Xysticus montanensis when compared to C.insignis antiserum, paralleling results of laboratory no-choice experiments.
• 4 The usefulness of serological analysis as a tool for arthropod trophic link identification is discussed.

     

Bifurcation and stability analysis of a temperature-dependent mite predator-prey interaction model incorporating a prey refuge

The non-linear behavior of a differential equations-based predator-prey model, incorporating a spatial refuge protecting a consant proportion of prey and with temperature-dependent parameters chosen appropriately for a mite interaction on fruit trees, is examined using the numerical bifurcation code AUTO 86. The most significant result of this analysis is the existence of a temperature interval in which increasing the amount of refuge dynamically destabilizes the system; and on part of this interval the interaction is less likely to persist in that predator and prey minimum population densities are lower than when no refuge is available. It is also shown that increasing the amount of refuge can lead to population outbreaks due to the presence of multiple stable states. The ecological implications of a refuge are discussed with respect to the biological control of mite pests.     

Anurans as prey: an exploratory analysis and size relationships between predators and their prey

The vertebrate predators of post-metamorphic anurans were quantified and the predator–prey relationship was investigated by analysing the relative size of invertebrate predators and anurans. More than 100 vertebrate predators were identified (in more than 200 reports) and classified as opportunistic, convenience, temporary specialized and specialized predators. Invertebrate predators were classified as solitary non-venomous, venomous and social foragers according to 333 reviewed reports. Each of these categories of invertebrate predators was compared with the relative size of the anurans, showing an increase in the relative size of the prey when predators used special predatory tactics. The number of species and the number of families of anurans that were preyed upon did not vary with the size of the predator, suggesting that prey selection was not arbitrary and that energetic constraints must be involved in this choice. The relatively low predation pressure upon brachycephalids was related to the presence of some defensive strategies of its species. This compounding review can be used as the foundation for future advances in vertebrate predator–prey interactions.     

The interplay between predator's prey preference and environmental heterogeneity in food web long-term stability

Prey preference of a predator is commonly used in models to analyze the timely issue of the relation among food web structure, diversity and stability. Given the variety of these terms in ecological literature, this work shows that domains of stability and species coexistence in food webs can be significantly altered by the chosen structure of predator's prey preference and environmental heterogeneity. Such results may bear upon issues in applied ecology, e.g., species conservation, biological control. More generally, they may serve as a caution with respect to the robustness of some results of food web theory.     

Habitat complexity reduces prey vulnerability: An experimental analysis using aquatic insect predators and immature dipteran prey

The effects of alternative prey and structural complexity of habitat on the selection of mosquito larvae by aquatic insect predators were evaluated in the laboratory. The water bugs Anisops bouvieri, Diplonychus (= Sphaerodema) rusticus, and D. annulatus, and the odonate nymphs, Ceriagrion coromandelianum and Brachydiplax chalybea chalybea, selected mosquito larvae based on their abundance relative to chironomid larvae and on the levels of habitat complexity. The effect of one prey species on the other was asymmetrical, as indicated through prey selectivity values. Compared to open habitat, the presence of macrophytes reduced the vulnerability of mosquito larvae while the effect was reverse in the presence of sediments. When both sediment and macrophytes were present in habitats, all the predators except D. annulatus consumed more mosquito larvae than chironomid larvae. The clearance rate, an indicator of predatory efficiency, varied among the predator species and habitat types. The results suggest that the outcome of the interactions between insect predators and mosquito immatures was context-dependent and that it was mediated by the presence of alternative controphic species and the habitat complexity.     

The motile escape response of a sessile prey: A sponge-scallop mutualism

The association of the sponges Myxilla incrustans (Esper) and Mycale adhaerens (Lambe) with the scallops Chlamys hastata hericia Gould and C. rubida (Hines) is shown to be a mutualism, which protects the sponges from predatory sponge-rasping dorid nudibranchs and the scallops from predatory starfish. The sponge is protected by scallop motility (also shown for the Suberites ficus-hermit crab association). The sponge helps to protect the scallops by altering the surface texture of the shell so increasing the efficacy of the swimming escape response by decreasing the adhesive abilities of asteroid tube-feet. The sponge also provides tactile camouflage against certain predatory starfish. There was no evidence that either component chemically deceived or repelled the predators of the other component. Predation pressure on both components of the association appears to be the major force leading to the mutualism.     

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.     

Temperature,predator–prey interaction strength and population stability

Warming could strongly stabilize or destabilize populations and food webs by changing the interaction strengths between predators and their prey. Predicting the consequences of warming requires understanding how temperature affects ingestion (energy gain) and metabolism (energy loss). Here, we studied the temperature dependence of metabolism and ingestion in laboratory experiments with terrestrial arthropods (beetles and spiders). From this data, we calculated ingestion efficiencies (ingestion/metabolism) and per capita interaction strengths in the short and long term. Additionally, we investigated if and how body mass changes these temperature dependencies. For both predator groups, warming increased metabolic rates substantially, whereas temperature effects on ingestion rates were weak. Accordingly, the ingestion efficiency (the ratio of ingestion to metabolism) decreased in all treatments. This result has two possible consequences: on the one hand, it suggests that warming of natural ecosystems could increase intrinsic population stability, meaning less fluctuations in population density; on the other hand, decreasing ingestion efficiencies may also lead to higher extinction risks because of starvation. Additionally, predicted long‐term per capita interaction strengths decreased with warming, which suggests an increase in perturbation stability of populations, i.e., a higher probability of returning to the same equilibrium density after a small perturbation. Together, these results suggest that warming has complex and potentially profound effects on predator–prey interactions and food‐web stability.     

The myth of constant predator: prey ratios

Apparent constancy in the ratio of predator species to prey species has been offered as evidence that ecological communities are structured by interspecific interactions. If significantly different from random expectation, this effect would be one of the few sound pieces of evidence for community structure. The evidence was re-evaluated by using the data from previous studies to form species pools, and forming simulated communities by drawing species at random from these pools (with replacement). Using a correlation coefficient (number of predator species versus number of prey species), and also the statistic used by the original workers (where different), the observed predator:prey correlation was compared to that for the random communities. In five studies, the observed predator:prey ratio was not significantly different from random expectation. In the only two studies where there was significant departure from the null model, it was with more variation in the ratio than expected on a random basis. It is concluded that there is as yet no evidence for near-constant predator:prey ratios.     

The dynamics of prey choice in fish: the importance of prey size and satiation

Over a number of decades the process of prey choice has been investigated using fishes as model predators. Using fishes for the model has allowed the proximate factors that determine how a mobile predator finds and chooses to eat the prey encountered within a variable 3‐D environment to be estimated. During prey choice a number of constraints exist, in particular most fish predators will eat their prey whole thus their jaws and gut create functional limitations once a prey has been attacked. By considering the relationship between the size of the prey and the predator's feeding apparatus and feeding motivation this study explores the link between mechanistic studies and theoretical, optimal foraging based predictions. How the prediction of prey choices made by the fish following prey encounter can be reconciled with what is likely to be found in the fish's stomach is discussed. This study uses a progression of empirical examples to illustrate how the limits of functional constraints and prey choice at different stages of motivation to feed can be taken into account to improve predictions of predator prey choice.     

The ultrastructural localisation of cadmium

Summary A modified technique for the ultrastructural localisation of heavy metals is described in this paper. The method involves precipitation of heavy metals as sulphides in the tissue by using (NH₄)₂S after brief fixation in glutaraldehyde. The sulphides are, in the presence of a physical developer, then used to catalyse the reduction of silver ions into visible molecular silver. This latter step of physical development has been normally carried out after embedding and sectioning. However, when we followed this method we found that the dark metal sulphide was lost from the tissue during the embedding in epoxy resin. Hence the method was unsuitable for our porposed experiment on the ultrastructural localisation of cadmium. We subsequently modified the technique primarily by treating very thin tissue slices with the developer before dehydration and embedding, thus eliminating any problem from sulphide loss. This modified technique was used to investigate the ultrastructural localisation of cadmium in the kidneys of mice which had been exposed to 50 ppm cadmium in their drinking water for up to eight months. The molecular silver was found to be located mainly in the proximal tubule cells, either as dense clumps in apical vesicles and lysosomes or diffuse grains throughout the cytoplasm of the cells particularly in the basal region. We interpret these results as indicating that cadmium is found in the apical vesicles, lysosomes and cytoplasm of proximal tubule cells.     

生态系统稳定性定义剖析

生态系统稳定性是理论生态学的焦点问题之一 ,综述和剖析生态系统稳定性的定义 ,对已有的定义进行了改进。生态系统稳定性是不超过生态阈值的生态系统的敏感性和恢复力。在这个概念中涉及到 3个概念 :生态阈值、敏感性和恢复力 ,阈值是生态系统在改变为另一个退化 (或进化 )系统前所能承受的干扰限度 ;敏感性是生态系统受到干扰后变化的大小和与其维持原有状态的时间 ;退化生态系统的恢复力就是消除干扰后生态系统能回到原有状态的能力 ,包括恢复速度和与原有状态的相似程度。在保护生态学中 ,阈值与恢复力的定义具有广泛的应用 ,特别是生态系统受到负面的干扰后而退化 ,退化的生态系统逐步恢复的过程可以利用恢复力来测定 ;而保护的成果就是力图避免干扰超过系统的阈值而达到一个实际的演替     

Body masses,functional responses and predator–prey stability

The stability of ecological communities depends strongly on quantitative characteristics of population interactions (type‐II vs. type‐III functional responses) and the distribution of body masses across species. Until now, these two aspects have almost exclusively been treated separately leaving a substantial gap in our general understanding of food webs. We analysed a large data set of arthropod feeding rates and found that all functional‐response parameters depend on the body masses of predator and prey. Thus, we propose generalised functional responses which predict gradual shifts from type‐II predation of small predators on equally sized prey to type‐III functional‐responses of large predators on small prey. Models including these generalised functional responses predict population dynamics and persistence only depending on predator and prey body masses, and we show that these predictions are strongly supported by empirical data on forest soil food webs. These results help unravelling systematic relationships between quantitative population interactions and large‐scale community patterns.     

Prey preference in stoneflies: a comparative analysis of prey vulnerability

Summary Laboratory feeding trials were conducted with the predaceous stonefly Hesperoperla pacifica and a number of mayfly and dipteran prey species to investigate the effects of predator size, and prey size and morphology, on the predator's success. Observations under dim red light permitted estimation of encounter rate (E/min), attack propensity (A/E), capture success (C/A) and handling time (HT). For prey of a particular species and size, HT decreased log-linearly with increasing predator size. Across all prey categories, HT increased log-linearly with increasing values of the ratio prey dry wt/predator dry wt, and differences among species appeared to be small. Overall, capture success was low, but C/A was higher for dipterans than for mayflies, especially with large H. pacifica. Predator size affected C/A when prey fell within a certain size range, but was not a detectable influence with very small or very large prey. Values of A/E of near 10% typified many predatorprey combinations; however, ephemerellid mayflies suffered markedly fewer attacks, and values of A/E up to 30% were obtained with some species-size combinations. We estimated benefit to the predator first as prey wt ingested per unit time (dry wt/HT), and second by mutliplying the former term by capture success. Values increased with increasing size of the predator, and inclusion of the C/A term indicated that predators would obtain greater reward from small relative to large prey, and from dipterans relative to mayflies. Howerver, there was little evidence that attacks were biased toward more profitable prey. We compare the relative contributions of E/min, A/E and C/A to prey choice, and discuss their applicability to predation events in nature.