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1.
Recent work with the freshwater zooplankton Daphnia has suggested that the quality of its algal prey can have a significant effect on its demographic rates and life-history patterns. Predator-prey theory linking food quantity and food quality predicts that a single system should be able to display two distinct patterns of population dynamics. One pattern is predicted to have high herbivore and low algal biomass dynamics (high HBD), whereas the other is predicted to have low herbivore and high algal biomass dynamics (low HBD). Despite these predictions and the stoichiometric evidence that many phytoplankton communities may have poor access to food of quality, there have been few tests of whether a dynamic predator-prey system can display both of these distinct patterns. Here we report, to the authors' knowledge, the first evidence for two dynamical patterns, as predicted by theory, in a single predator-prey system. We show that the high HBD is a result of food quantity effects and that the low HBD is a result of food quality effects, which are maintained by phosphorus limitation in the predator. These results provide an important link between the known effects of nutrient limitation in herbivores and the significance of prey quality in predator-prey population dynamics in natural zooplankton communities.  相似文献   

2.
Norman Owen‐Smith 《Oikos》2015,124(11):1417-1426
Simple models coupling the dynamics of single predators to single prey populations tend to generate oscillatory dynamics of both predator and prey, or extirpation of the prey followed by that of the predator. In reality, such oscillatory dynamics may be counteracted by prey refugia or by opportunities for prey switching by the predator in multi‐prey assemblages. How these mechanisms operate depends on relative prey vulnerability, a factor ignored in simple interactive models. I outline how compositional, temporal, demographic and spatial heterogeneities help explain the contrasting effects of top predators on large herbivore abundance and population dynamics in species‐rich African savanna ecosystems compared with less species‐diverse northern temperate or subarctic ecosystems. Demographically, mortality inflicted by predation depends on the relative size and life history stage of the prey. Because all animals eventually die and are consumed by various carnivores, the additive component of the mortality inflicted is somewhat less than the predation rate. Prey vulnerability varies annually and seasonally, and between day and night. Spatial variation in the risk of predation depends on vegetation cover as well as on the availability of food resources. During times of food shortage, herbivores become prompted to occupy more risky habitats retaining more food. Predator concentrations dependent on the abundance of primary prey species may restrict the occurrence of other potential prey species less resistant to predation. The presence of multiple herbivore species of similar size in African savannas allows the top predator, the lion, to shift its prey selection flexibly dependent on changing prey vulnerability. Hence top–down and bottom–up influences on herbivore populations are intrinsically entangled. Models coupling the population dynamics of predators and prey need to accommodate the changing influences of prey demography, temporal variation in environmental conditions, and spatial variation in the relative vulnerability of alternative prey species to predation. Synthesis While re‐established predators have had major impacts on prey populations in northern temperate regions, multiple large herbivore species typically coexist along with diverse carnivores in African savanna ecosystems. In order to explain these contrasting outcomes, certain functional heterogeneities must be recognised, including relative vulnerability of alternative prey, temporal variation in the risk of predation, demographic differences in susceptibility to predation, and spatial contrasts in exposure to predation. Food shortfalls prompt herbivores to exploit more risky habitats, meaning that top–down and bottom–up influences on prey populations are intrinsically entangled. Models coupling the interactive dynamics of predator and prey populations need to incorporate these varying influences on relative prey vulnerability.  相似文献   

3.
The mineral and biochemical food quality of prey may limit predator production. This well‐studied direct bottom–up effect is especially prominent for herbivore–plant interactions. Low‐quality prey species, particularly when defended, are generally considered to be less prone to predator‐driven extinction. Undefended high‐quality prey species sustain high predator production thereby potentially increasing their own extinction risk. The food quality of primary producers is highly species‐specific. In communities of competing prey species, predators thus may supplement their diets of low‐quality prey with high‐quality prey, leading to indirect horizontal interactions between prey species of different food quality. We explore how these predator‐mediated indirect interactions affect species coexistence in a general predator–prey model that is parametrized for an experimental algae– rotifer system. To cover a broad range of three essential functional traits that shape many plant–herbivore interactions we consider differences in 1) the food quality of the prey species, 2) their competitive ability for nutrient uptake and 3) their defence against predation. As expected, low food quality of prey can, similarly to defence, provide protection against extinction by predation. Counterintuitively, our simulations demonstrate that being of high food quality also prevents extinction of that prey species and additionally promotes coexistence with a competing, low‐quality prey. The persistence of the high‐quality prey enables a high conversion efficiency and control of the low‐quality prey by the predator and allows for re‐allocation of nutrients to the high‐quality competitor. Our results show that high food quality is not necessarily detrimental for a prey species but instead can protect against extinction and promote species richness and functional biodiversity.  相似文献   

4.
Quantitative approaches to predator–prey interactions are central to understanding the structure of food webs and their dynamics. Different predatory strategies may influence the occurrence and strength of trophic interactions likely affecting the rates and magnitudes of energy and nutrient transfer between trophic levels and stoichiometry of predator–prey interactions. Here, we used spider–prey interactions as a model system to investigate whether different spider web architectures—orb, tangle, and sheet‐tangle—affect the composition and diet breadth of spiders and whether these, in turn, influence stoichiometric relationships between spiders and their prey. Our results showed that web architecture partially affects the richness and composition of the prey captured by spiders. Tangle‐web spiders were specialists, capturing a restricted subset of the prey community (primarily Diptera), whereas orb and sheet‐tangle web spiders were generalists, capturing a broader range of prey types. We also observed elemental imbalances between spiders and their prey. In general, spiders had higher requirements for both nitrogen (N) and phosphorus (P) than those provided by their prey even after accounting for prey biomass. Larger P imbalances for tangle‐web spiders than for orb and sheet‐tangle web spiders suggest that trophic specialization may impose strong elemental constraints for these predators unless they display behavioral or physiological mechanisms to cope with nutrient limitation. Our findings suggest that integrating quantitative analysis of species interactions with elemental stoichiometry can help to better understand the occurrence of stoichiometric imbalances in predator–prey interactions.  相似文献   

5.
Nutrient stoichiometric ratios are primary driving factors of planktonic food web dynamics. Ecological stoichiometry theory postulates the elemental ratios of consumer species to be homeostatic, while primary-producer stoichiometry may vary with ambient nutrient availability. The notion of phytoplankton intracellular storage is far from novel, but remains largely unexplored in modeling studies of population dynamics. We constructed a seasonally-unforced, zero-dimensional, nutrient–phytoplankton–zooplankton–detritus (NPZD) model that considers dynamic phytoplankton phosphorus reserves and quasi-dynamic zooplankton stoichiometry. A generic food quality term is used to express seston biochemical composition, ingestibility, and digestibility. We examined the sensitivity of the planktonic food web patterns to light and nutrient availability, zooplankton mortality, and detritus food quality as well as to phytoplankton intracellular storage and zooplankton stoichiometry. Our results reinforce earlier findings that high quality seston exerts a stabilizing effect on food web dynamics. However, we also found that the combination of low algal and high detritus food quality with high zooplankton mortality yielded limit cycles and multiple steady states, suggesting that the heterogeneity characterizing seston nutritional quality may have more complicated ecological ramifications. Our numerical experiments identify resource competition strategies related to nutrient transport rates and internal nutrient quotas that may be beneficial for phytoplankton to persevere in resource-limiting habitats. We also highlight the importance of the interplay between optimal stoichiometry and the factors controlling homeostatic rigidity in zooplankton. In particular, our predictions show that the predominance of phosphorus-rich and tightly-homeostatic herbivores in nutrient-enriched environments with low seston food quality can potentially result in high phytoplankton abundance, high phytoplankton-to-zooplankton ratios, and acceleration of oscillatory dynamics. Generally, our modeling study emphasizes the impact of both intracellular/somatic storage and food quality on prey–predator interactions, pinpointing an important aspect of food web dynamics usually neglected by the contemporary modeling studies.  相似文献   

6.
In traditional models of predator–prey population dynamics, it is usually assumed that consumed prey are immediately removed from the population. However, in plant–herbivore interactions, damaged plants are generally alive after attacks by herbivores. This can result in successive or simultaneous attacks by multiple predators on a single prey item (here, the term prey is expanded to include plants). We constructed a mathematical model with two time scales, taking into account predation processes within a generation, considering post‐predation survival and the modularity of prey. We assumed that a prey item can be divided into modules and that it can be fed on by multiple predators or parasitized by multiple parasites at the same time. The model includes two essential factors: the modularity of prey for predators (n) and the detaching/attaching ratio of predators to prey (ε). Based on the formulae, we revealed a general property of realistic dynamics in plant–herbivore and host–parasite interactions. The analysis showed that the model could be approximated by models with the type I, type II or Beddington–DeAngelis functional responses by taking appropriate limits to the situations. When modularity is low or the detaching/attaching ratio is high, population dynamics tend to be stabilized. These stabilizing effects may be related to interference competition among predator individuals or increases in free prey modules and free predator individuals. In the limit of high modularity, the ratio of the attached prey modules to the total prey modules becomes negligible and the dynamics tend to be destabilized. However, if quantity and quality of prey modules are negatively correlated, the equilibrium is likely to be stabilized at high modularity as long as it remains feasible. These results suggest that considering post‐predation survival and modularity of prey is crucial to understand predator–prey interactions.  相似文献   

7.
Variation in the vulnerability of herbivore prey to predation is linked to body size, yet whether this relationship is size‐nested or size‐partitioned remains debated. If size‐partitioned, predators would be focused on prey within their preferred prey size range. If size‐nested, smaller prey species should become increasingly more vulnerable because increasingly more predators are capable of catching them. Yet, whether either of these strategies manifests in top–down prey population limitation would depend both on the number of potential predator species as well as the total mortality imposed. Here we use a rare ecosystem scale ‘natural experiment’ comparing prey population dynamics between a period of intense predator persecution and hence low predator densities and a period of active predator protection and population recovery. We use three decades of data on herbivore abundance and distribution to test the role of predation as a mechanism of population limitation among prey species that vary widely in body size. Notably, we test this within one of the few remaining systems where a near‐full suite of megaherbivores occur in high density and are thus able to include a thirtyfold range in herbivore body size gradient. We test whether top–down limitation on prey species of particular body size leads to compositional shifts in the mammalian herbivore community. Our results support both size‐nested and size‐partitioning predation but suggest that the relative top–down limiting impact on prey populations may be more severe for intermediate sized species, despite having fewer predators than small species. In addition we show that the gradual recovery of predator populations shifted the herbivore community assemblage towards large‐bodied species and has led to a community that is strongly dominated by large herbivore biomass.  相似文献   

8.
Predictions on the consequences of the rapidly increasing atmospheric CO2 levels and associated climate warming for population dynamics, ecological community structure and ecosystem functioning depend on mechanistic energetic models of temperature effects on populations and their interactions. However, such mechanistic approaches combining warming effects on metabolic (energy loss of organisms) and feeding rates (energy gain by organisms) remain a key, yet elusive, goal. Aiming to fill this void, we studied the metabolic rates and functional responses of three differently sized, predatory ground beetles on one mobile and one more resident prey species across a temperature gradient (5, 10, 15, 20, 25 and 30 °C). Synthesizing metabolic and functional‐response theory, we develop novel mechanistic predictions how predator–prey interaction strengths (i.e., functional responses) should respond to warming. Corroborating prior theory, warming caused strong increases in metabolism and decreases in handling time. Consistent with our novel model, we found increases in predator attack rates on a mobile prey, whereas attack rates on a mostly resident prey remained constant across the temperature gradient. Together, these results provide critically important information that environmental warming generally increases the direct short‐term per capita interaction strengths between predators and their prey as described by functional‐response models. Nevertheless, the several fold stronger increase in metabolism with warming caused decreases in energetic efficiencies (ratio of per capita feeding rate to metabolic rate) for all predator–prey interactions. This implies that warming of natural ecosystems may dampen predator–prey oscillations thus stabilizing their dynamics. The severe long‐term implications; however, include predator starvation due to energetic inefficiency despite abundant resources.  相似文献   

9.
Climate change will alter the distribution of rainfall, with potential consequences for the hydrological dynamics of aquatic habitats. Hydrological stability can be an important determinant of diversity in temporary aquatic habitats, affecting species persistence and the importance of predation on community dynamics. As such, prey are not only affected by drought‐induced mortality but also the risk of predation [a non‐consumptive effect (NCE)] and actual consumption by predators [a consumptive effect (CE)]. Climate‐induced changes in rainfall may directly, or via altered hydrological stability, affect predator–prey interactions and their cascading effects on the food web, but this has rarely been explored, especially in natural food webs. To address this question, we performed a field experiment using tank bromeliads and their aquatic food web, composed of predatory damselfly larvae, macroinvertebrate prey and bacteria. We manipulated the presence and consumption ability of damselfly larvae under three rainfall scenarios (ambient, few large rainfall events and several small rainfall events), recorded the hydrological dynamics within bromeliads and examined the effects on macroinvertebrate colonization, nutrient cycling and bacterial biomass and turnover. Despite our large perturbations of rainfall, rainfall scenario had no effect on the hydrological dynamics of bromeliads. As a result, macroinvertebrate colonization and nutrient cycling depended on the hydrological stability of bromeliads, with no direct effect of rainfall or predation. In contrast, rainfall scenario determined the direction of the indirect effects of predators on bacteria, driven by both predator CEs and NCEs. These results suggest that rainfall and the hydrological stability of bromeliads had indirect effects on the food web through changes in the CEs and NCEs of predators. We suggest that future studies should consider the importance of the variability in hydrological dynamics among habitats as well as the biological mechanisms underlying the ecological responses to climate change.  相似文献   

10.
Intraspecific variation is central to our understanding of evolution and population ecology, yet its consequences for community ecology are poorly understood. Animal personality – consistent individual differences in suites of behaviours – may be particularly important for trophic dynamics, where predator personality can determine activity rates and patterns of attack. We used mesocosms with aquatic food webs in which the top predator (dragonfly nymphs) varied in activity and subsequent attack rates on zooplankton, and tested the effects of predator personality. We found support for four hypotheses: (1) active predators disproportionately reduce the abundance of prey, (2) active predators select for predator‐resistant prey species, (3) active predators strengthen trophic cascades (increase phytoplankton abundance) and (4) active predators are more likely to cannibalise one another, weakening all other trends when at high densities. These results suggest that intraspecific variation in predator personality is an important determinant of prey abundance, community composition and trophic cascades.  相似文献   

11.
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.  相似文献   

12.
When the nutrient content of food is limited, herbivores often increase their feeding rates. Such an increase in the feeding rate is called ‘compensatory feeding’. Although it has a number of implications for herbivore population and plant–forager dynamics, the compensatory feeding is not yet functionally formulated especially in relation with ecological stoichiometry. Therefore, we constructed a simple mathematical model by incorporating the optimal feeding rate into the type II functional response to maximize a forager's growth rate under constraints of carbon or nutritionally important element like phosphorus (P). We used the planktonic herbivore Daphnia as a model herbivore. The model revealed that the optimal feeding rate increased by using excess carbon when relative P content of food was less than a certain level, which is known as the threshold elemental ratio. This level changed with the change of food abundance. It also showed that whether or not foragers should exhibit compensatory feeding depends on their stoichiometric characteristics and digestive traits, and also on the assimilability of a given food. These findings are helpful to test the feeding conditions under which compensatory feeding is advantageous for a given animal. Our model can be easily incorporated into forager population dynamics and prey‐consumer interaction models because the optimal feeding rate can be analytically given.  相似文献   

13.
Akihiko Mougi  Kinya Nishimura 《Oikos》2008,117(11):1732-1740
Destabilization of one predator–one prey systems with an increase in nutrient input has been viewed as a paradox. We report that enrichment can damp population cycles by a food‐web structure that balances inflexible and flexible interaction links (i.e. specialist and generalist predators). We modeled six predator–prey systems involving three or four species in which the predators practice optimal foraging based on prey profitability determined by handling time. In all models, the balance of interaction links simultaneously decreased the amplitude of population oscillations and increased the minimum density with increasing enrichment, leading to a potential theoretical resolution of the paradox of enrichment in non‐equilibrium dynamics. The stabilization mechanism was common to all of the models. Important previous studies on the stability of food webs have also demonstrated that a balance of interaction strengths stabilizes systems, suggesting a general rule of ecosystem stability.  相似文献   

14.
Modeling under the framework of ecological stoichiometric allows the investigation of the effects of food quality on food web population dynamics. Recent discoveries in ecological stoichiometry suggest that grazer dynamics are affected by insufficient food nutrient content (low phosphorus (P)/carbon (C) ratio) as well as excess food nutrient content (high P:C). This phenomenon is known as the “stoichiometric knife edge.” While previous models have captured this phenomenon, they do not explicitly track P in the producer or in the media that supports the producer, which brings questions to the validity of their predictions. Here, we extend a Lotka–Volterra-type stoichiometric model by mechanistically deriving and tracking P in the producer and free P in the environment in order to investigate the growth response of Daphnia to algae of varying P:C ratios. Bifurcation analysis and numerical simulations of the full model, that explicitly tracks phosphorus, lead to quantitative different predictions than previous models that neglect to track free nutrients. The full model shows that the fate of the grazer population can be very sensitive to excess nutrient concentrations. Dynamical free nutrient pool seems to induce extreme grazer population density changes when total nutrient is in an intermediate range.  相似文献   

15.
The Lotka–Volterra model is the most commonly used framework to describe the dynamics of ecological systems in which two species interact, one as a predator and the other as prey. Theoretical ecologists have since built on variants of these equations, frequently applying them to model the dynamics of algal-herbivore interactions in aquatic systems. In this study, we augment a Lotka–Volterra system by introducing a bioenergetically-explicit, ecophysiological model to examine how variations in resource allocation affect zooplankton growth and subsequently phytoplankton dynamics. Ingested material within a zooplankter's gut is separated into distinct internal congener pools that are used to support physiological processes occurring in a hierarchical direction: neurological functions, energetics, osmoregulatory maintenance, waste management, and finally growth. Consistent with the predictions of the “stoichiometric knife edge” theory, our analysis suggests that a balanced algal congener composition is required to optimize zooplankton internal congener saturations, resulting in a maximal allocation of energy to growth. In examining the advantages rendered by different strategies of minimum and optimum somatic quotas when experiencing phosphorus-enrichment conditions, we show that herbivores with narrow homeostatic bounds and animals with low minimum quotas (or depletion specialists) achieve optimal performance first. Our analysis also predicts patterns of multiple stable equilibria in which the same environmental conditions can be characterized by dramatically different prey-to-predator ratios. Importantly, abrupt shifts from one state to another can be induced not only by short-term variations in food abundance but also by variations in the nutritional quality of the prey. Our predictions have profound implications for connecting microscopic processes with macroscopic patterns and offer new insights into the multitude of factors that modulate food web dynamics.  相似文献   

16.
Jason T. Hoverman  Rick A. Relyea 《Oikos》2012,121(8):1219-1230
Despite the amount of research on the inducible defenses of prey against predators, our understanding of the long‐term significance of non‐lethal predators on prey phenotypes, prey population dynamics, and community structure has rarely been explored. Our objectives were to assess the effects of predators on prey defenses, prey population dynamics, and the relative magnitude of density‐ versus trait‐mediated indirect interactions (DMIIs and TMIIs) over multiple prey generations. Using a freshwater snail and three common snail predators, we constructed a series of community treatments with pond mesocosms that manipulated trophic structure, the identity of the top predator, and whether predators were caged or uncaged. We quantified snail phenotypes, snail population size, and resource abundance over multiple snail generations. We found that snails were expressing inducible defenses in our system although the magnitude of the responses varied over time and across predator species. Despite the expression of inducible defenses, caged predators did not reduce snail population size. There also was no evidence of TMIIs throughout the experiment suggesting that TMIIs have a minimal role in the long‐term structure of our communities. The absence of TMIIs was largely driven by the lack of predator‐induced reductions in resource consumption and the lack of consistent reductions in population size with predator cues. In contrast, we detected strong DMIIs associated with lethal predators suggesting that DMIIs are the dominant long‐term mechanism influencing community structure. Our results demonstrate that although predators can have significant effects on prey phenotypes and sometimes cause short‐term TMIIs, there may be few long‐term consequences of these responses on population dynamics and indirect interactions, at least within simple food webs. Research directed towards addressing the long‐term consequences of predator–prey interactions within communities will help to reveal whether the conclusions and predictions generated from short‐term experiments are applicable over ecological and evolutionary timescales.  相似文献   

17.
Predation and scavenging have been classically understood as independent processes, with predator–prey interactions and scavenger–carrion relationships occurring separately. However, the mere recognition that most predators also scavenge at variable rates, which has been traditionally ignored in food‐web and community ecology, leads to a number of emergent interaction routes linking predation and scavenging. The general goal of this review is to draw attention to the main inter‐specific interactions connecting predators (particularly, large mammalian carnivores), their live prey (mainly ungulates), vultures and carrion production in terrestrial assemblages of vertebrates. Overall, we report an intricate network of both direct (competition, facilitation) and indirect (hyperpredation, hypopredation) processes, and provide a conceptual framework for the future development of this promising topic in ecological, evolutionary and biodiversity conservation research. The classic view that scavenging does not affect the population dynamics of consumed organisms is questioned, as multiple indirect top‐down effects emerge when considering carrion and its facultative consumption by predators as fundamental and dynamic components of food webs. Stimulating although challenging research opportunities arise from the study of the interactions among living and detrital or non‐living resource pools in food webs.  相似文献   

18.
Food chain theory is one of the cornerstones of ecology, providing many of its basic predictions, such as biomass pyramids, trophic cascades and predator–prey oscillations. Yet, ninety years into this theory, the conditions under which these patterns may occur and persist in nature remain subject to debate. Rather than address each pattern in isolation, we propose that they must be understood together, calling for synthesis in a fragmented landscape of theoretical and empirical results. As a first step, we propose a minimal theory that combines the long‐standing energetic and dynamical approaches of food chains. We chart theoretical predictions on a concise map, where two main regimes emerge: across various functioning and stability metrics, one regime is characterised by pyramidal patterns and the other by cascade patterns. The axes of this map combine key physiological and ecological variables, such as metabolic rates and self‐regulation. A quantitative comparison with data sheds light on conflicting theoretical predictions and empirical puzzles, from size spectra to causes of trophic cascade strength. We conclude that drawing systematic connections between various existing approaches to food chains, and between their predictions on functioning and stability, is a crucial step in confronting this theory to real ecosystems.  相似文献   

19.
20.
To assess the effects of fluctuating prey availability on predator population dynamics and grazing impact on phytoplankton, we measured growth and grazing rates of three heterotrophic dinoflagellate species—Oxyrrhis marina, Gyrodinium dominans and Gyrodinium spirale—before and after depriving them of phytoplankton prey. All three dinoflagellate species survived long periods (> 10 d) without algal prey, coincident with decreases in predator abundance and cell size. After 1–3 wks, starvation led to a 17–57% decrease in predator cell volume and some cells became deformed and transparent. When re‐exposed to phytoplankton prey, heterotrophs ingested prey within minutes and increased cell volumes by 4–17%. At an equivalent prey concentration, continuously fed predators had ~2‐fold higher specific growth rates (0.18 to 0.55 d?1) than after starvation (?0.16 to 0.25 d?1). Maximum specific predator growth rates would be achievable only after a time lag of at least 3 d. A delay in predator growth poststarvation delays predator‐induced phytoplankton mortality when prey re‐emerges at the onset of a bloom event or in patchy prey distributions. These altered predator‐prey population dynamics have implications for the formation of phytoplankton blooms, trophic transfer rates, and potential export of carbon.  相似文献   

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