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1.
Intraguild predation (IGP) is a combination of competition and predation which is the most basic system in food webs that contains three species where two species that are involved in a predator/prey relationship are also competing for a shared resource or prey. We formulate two intraguild predation (IGP: resource, IG prey and IG predator) models: one has generalist predator while the other one has specialist predator. Both models have Holling-Type I functional response between resource-IG prey and resource-IG predator; Holling-Type III functional response between IG prey and IG predator. We provide sufficient conditions of the persistence and extinction of all possible scenarios for these two models, which give us a complete picture on their global dynamics. In addition, we show that both IGP models can have multiple interior equilibria under certain parameters range. These analytical results indicate that IGP model with generalist predator has “top down” regulation by comparing to IGP model with specialist predator. Our analysis and numerical simulations suggest that: (1) Both IGP models can have multiple attractors with complicated dynamical patterns; (2) Only IGP model with specialist predator can have both boundary attractor and interior attractor, i.e., whether the system has the extinction of one species or the coexistence of three species depending on initial conditions; (3) IGP model with generalist predator is prone to have coexistence of three species.  相似文献   

2.
Trophic supplements to intraguild predation   总被引:2,自引:0,他引:2  
Intraguild predation (IGP) is a dominant community module in terrestrial food webs that occurs when multiple consumers feed both on each other and on a shared prey. This specific form of omnivory is common in terrestrial communities and is of particular interest for conservation biology and biological control given its potential to disrupt management of threatened or pest species. Extensive theory exists to describe the dynamics of three-species IGP, but these models have largely overlooked the potential for other, exterior interactions, to alter the dynamics within the IGP module. We investigated how three forms of feeding outside of the IGP module by intraguild predators (i.e. trophic supplementation) affect the dynamics of the predators (both IG predator and IG prey) and their shared resource. Specifically, we examined how the provision of a constant donor-controlled resource, the availability of an alternative prey species, and predator plant-feeding affect the dynamics of IGP models. All three forms of trophic supplements modified the basic expectations of IGP theory in two important ways, and their effects were similar. First, coexistence was possible without the IG prey being a superior competitor for the original shared resource if the IG prey could effectively exploit one of the types of trophic supplements. However, supplements to the IG predator restricted the potential for coexistence. Second, supplements to the IG prey ameliorated the disruptive effects of the IG predator on the suppression of the shared resource, promoting effective control of the resource in the presence of both predators. Consideration of these three forms of trophic supplementation, all well documented in natural communities, adds substantial realism and predictive power to intraguild predation theory.  相似文献   

3.
In basic intraguild predation (IGP) systems, predators and prey also compete for a shared resource. Theory predicts that persistence of these systems is possible when intraguild prey is superior in competition and productivity is not too high. IGP often results from ontogenetic niche shifts, in which the diet of intraguild predators changes as a result of growth in body size (life-history omnivory). As a juvenile, a life-history omnivore competes with the species that becomes its prey later in life. Competition can hence limit growth of young predators, while adult predators can suppress consumers and therewith neutralize negative effects of competition. We formulate and analyze a stage-structured model that captures both basic IGP and life-history omnivory. The model predicts increasing coexistence of predators and consumers when resource use of stage-structured predators becomes more stage specific. This coexistence depends on adult predators requiring consumer biomass for reproduction and is less likely when consumers outcompete juvenile predators, in contrast to basic IGP. Therefore, coexistence occurs when predation structures the community and competition is negligible. Consequently, equilibrium patterns over productivity resemble those of three-species food chains. Life-history omnivory thus provides a mechanism that allows intraguild predators and prey to coexist over a wide range of resource productivity.  相似文献   

4.
Traditionally, productivity and disturbance have been hypothesized as important determinants of food-chain length. More recently, growing empirical evidence suggests a strong role of ecosystem size. To theoretically explore the effects of basal productivity, disturbance, and ecosystem size on food-chain length, we develop and analyze a metacommunity model of intraguild predation (IGP). The model finds that, when local IGP is weak, increasing basal productivity, weakening disturbance, and increasing ecosystem size will generally increase food-chain length. When local IGP is strong, by contrast, increasing basal productivity or weakening disturbance favors intraguild predators and hinders the coexistence of intraguild predators and intraguild prey, limiting food-chain length. In contrast, increasing ecosystem size can promote coexistence even when local IGP is strong, increasing food-chain length through inserting intraguild prey and changing the degree of omnivory by intraguild predators. Intraguild prey needs to be the superior colonizer to intraguild predators for this to occur. We discuss that these theoretical predictions appear consistent with empirical patterns.  相似文献   

5.
The probability of individuals being targeted as prey often decreases as they grow in size. Such size‐dependent predation risk is very common in systems with intraguild predation (IGP), i.e. when predatory species interact through predation and competition. Theory on IGP predicts that community composition depends on productivity. When recently testing this prediction using a terrestrial experimental system consisting of two phytoseiid mite species, Iphiseius degenerans as the IG‐predator and Neoseiulus cucumeris as the IG‐prey, and pollen (Typha latifolia) as the shared resource, we could not find the predicted community shift. Instead, we observed that IG‐prey excluded IG‐predators when the initial IG‐prey/IG‐predator ratio was high, whereas the opposite held when the initial ratio was low, which is also not predicted by theory. We therefore hypothesized that the existence of vulnerable and invulnerable stages in the two populations could be an important driver of the community composition. To test this, we first demonstrate that IG‐prey adults indeed attacked IG‐predator juveniles in the presence of the shared resource. Second, we show that the invasion capacity of IG‐predators at high productivity levels indeed depended on the structure of resident IG‐prey populations. Third, we further confirmed our hypothesis by mimicking successive invasion events of IG‐predators into an established population of IG‐prey at high productivity levels, which consistently failed. Our results show that the interplay between stage structure of populations and reciprocal intraguild predation is decisive at determining the species composition of communities with intraguild predation.  相似文献   

6.
Although ecological theory exists to predict dynamics in communities with intraguild predation (IGP), few empirical tests have examined this theory. IGP theory, in particular, predicts that when two competitors interact via IGP, with increasing resource productivity: (1) the IG predator will increase in abundance as the IG prey declines, and (2) increasing dominance of the IG predator will cause resource density to increase. Here, we provide a first test of these predictions in a field community consisting of a scale insect and its two specialist parasitoids, Aphytis melinus (the IG predator) and Encarsia perniciosi (the IG prey). The shared resource, California red scale, is a pest of citrus, and its productivity varies across a threefold range among citrus cultivars. We examined both absolute and relative densities of parasitoids along this natural gradient of scale productivity in three citrus cultivars (orange, grapefruit and lemon). Although both parasitoid species were found in all three cultivars, their abundances reflected those predicted by IGP theory: the IG prey species dominated at low productivity and the IG predator dominated at high productivity. This relationship was caused by an increase in Aphytis density with productivity. In addition, the density of scale increased with the dominance of the IG predator. These results from a field system demonstrate the important dynamic outcomes for food webs with IGP.  相似文献   

7.
It is well‐known that prey species often face trade‐offs between defense against predation and competitiveness, enabling predator‐mediated coexistence. However, we lack an understanding of how the large variety of different defense traits with different competition costs affects coexistence and population dynamics. Our study focusses on two general defense mechanisms, that is, pre‐attack (e.g., camouflage) and post‐attack defenses (e.g., weaponry) that act at different phases of the predator—prey interaction. We consider a food web model with one predator, two prey types and one resource. One prey type is undefended, while the other one is pre‐ or post‐attack defended paying costs either by a higher half‐saturation constant for resource uptake or a lower maximum growth rate. We show that post‐attack defenses promote prey coexistence and stabilize the population dynamics more strongly than pre‐attack defenses by interfering with the predator's functional response: Because the predator spends time handling “noncrackable” prey, the undefended prey is indirectly facilitated. A high half‐saturation constant as defense costs promotes coexistence more and stabilizes the dynamics less than a low maximum growth rate. The former imposes high costs at low resource concentrations but allows for temporally high growth rates at predator‐induced resource peaks preventing the extinction of the defended prey. We evaluate the effects of the different defense mechanisms and costs on coexistence under different enrichment levels in order to vary the importance of bottom‐up and top‐down control of the prey community.  相似文献   

8.
Many plants employ indirect defenses against herbivores; often plants provide a shelter or nutritional resource to predators, increasing predator abundance, and lessening herbivory to the plant. Often, predators on the same plant represent different life stages and different species. In these situations intraguild predation (IGP) may occur and may decrease the efficacy of that defense. Recently, several sticky plants have been found to increase indirect defense by provisioning predatory insects with entrapped insects (hereafter: carrion). We conducted observational studies and feeding trials with herbivores and predators on two sticky, insect‐entrapping asters, Hemizonia congesta and Madia elegans, to construct food webs for these species and determine the prevalence of IGP in these carrion‐provisioning systems. In both systems, intraguild predation was the most common interaction observed. To determine whether IGP was driven by resource abundance, whether it reduced efficacy of this indirect defense and whether stickiness or predator attraction was induced by damage, we performed field manipulations on H. congesta. Carrion supplementation led to an increase in predator abundance and IGP. IGP was asymmetric within the predator guild: assassin bugs and spiders preyed on small stilt bugs but not vice versa. Despite increased IGP, carrion provisions decreased the abundance of the two most common herbivores (a weevil and a mealybug). Overall seed set was driven by plant size, but number of seeds produced per fruit significantly increased with increasing carrion, likely because of the reduction in the density of a seed‐feeding weevil. Observationally and experimentally, we found that carrion‐mediated indirect defense of tarweeds led to much intraguild predation, though predators effectively reduced herbivore abundance despite the increase in IGP.  相似文献   

9.
Intra‐guild predation (IGP) – where a top predator (IGPred) consumes both a basal resource and a competitor for that resource (IGPrey) – has become a fundamental part of understanding species interactions and community dynamics. IGP communities composed of intraguild predators and prey have been well studied; however, we know less about IGP communities composed of predators, pathogens, and resources. Resource quality plays an important role in community dynamics and may influence IGP dynamics as well. We conducted a meta‐analysis on predator–pathogen–resource communities to determine whether resource quality mediated by the pathogen affected predator life‐history traits and if these effects met the theoretical constraints of IGP communities. To do this, we summarized results from studies that investigated the use of predators and pathogens to control insect pests. In these systems, the predators are the IGPred and pathogens are the IGPrey. We found that consumer longevity, fecundity, and survival decreased by 26%, 31% and 13% respectively, when predators consumed pathogen‐infected prey, making the infected prey a low quality resource. Predators also significantly preferred healthy prey over infected prey. When we divided consumers by enemy type, strict predators (e.g. wolf spiders) had no preference while parasitoids preferred healthy prey. Our results suggest that communities containing parasitoids and pathogens may rarely exhibit intraguild predation; whereas, communities composed of strict predators and pathogens are more likely dominated by IGP dynamics. In these latter communities, the consumption of low and high quality resources suggests that IGP communities composed of strict predators, pathogens and prey should naturally persist, supporting IGP theory. Synthesis We investigated how consuming pathogen‐infected prey influence important life‐history parameters of insect predators. Pathogens are used in a variety of biocontrol programs, especially to control crop pests. We found that true predators (i.e. wolf spiders) have no preference for healthy or infected prey and have reduced fecundity, survival and longevity consuming infected prey. However, parasitoids avoided infected prey when possible. In biocontrol programs with multiple control agents, parasitoids and pathogens would do a better job controlling pests as predators would reduce the amount of pathogen available and have reduced fitness from consuming infected prey. However, theory suggests that true predators, prey and pathogens may coexist long term.  相似文献   

10.
Occurrence of cannibalism and inferior competitive ability of predators compared to their prey have been suggested to promote coexistence in size-structured intraguild predation (IGP) systems. The intrinsic size-structure of fish provides the necessary prerequisites to test whether the above mechanisms are general features of species interactions in fish communities where IGP is common. We first experimentally tested whether Arctic char (Salvelinus alpinus) were more efficient as a cannibal than as an interspecific predator on the prey fish ninespine stickleback (Pungitius pungitius) and whether ninespine stickleback were a more efficient competitor on the shared zooplankton prey than its predator, Arctic char. Secondly, we performed a literature survey to evaluate if piscivores in general are more efficient as cannibals than as interspecific predators and whether piscivores are inferior competitors on shared resources compared to their prey fish species. Both controlled pool experiments and outdoor pond experiments showed that char imposed a higher mortality on YOY char than on ninespine sticklebacks, suggesting that piscivorous char is a more efficient cannibal than interspecific predator. Estimates of size dependent attack rates on zooplankton further showed a consistently higher attack rate of ninespine sticklebacks compared to similar sized char on zooplankton, suggesting that ninespine stickleback is a more efficient competitor than char on zooplankton resources. The literature survey showed that piscivorous top consumers generally selected conspecifics over interspecific prey, and that prey species are competitively superior compared to juvenile piscivorous species in the zooplankton niche. We suggest that the observed selectivity for cannibal prey over interspecific prey and the competitive advantage of prey species over juvenile piscivores are common features in fish communities and that the observed selectivity for cannibalism over interspecific prey has the potential to mediate coexistence in size structured intraguild predation systems.  相似文献   

11.
Predation is a major selective force for the evolution of behavioural characteristics of prey. Predation among consumers competing for food is termed intraguild predation (IGP). From the perspective of individual prey, IGP differs from classical predation in the likelihood of occurrence because IG prey is usually more rarely encountered and less profitable because it is more difficult to handle than classical prey. It is not known whether IGP is a sufficiently strong force to evolve interspecific threat sensitivity in antipredation behaviours, as is known from classical predation, and if so whether such behaviours are innate or learned. We examined interspecific threat sensitivity in antipredation in a guild of predatory mite species differing in adaptation to the shared spider mite prey (i.e. Phytoseiulus persimilis, Neoseiulus californicus and Amblyseius andersoni). We first ranked the players in this guild according to the IGP risk posed to each other: A. andersoni was the strongest IG predator; P. persimilis was the weakest. Then, we assessed the influence of relative IGP risk and experience on maternal strategies to reduce offspring IGP risk: A. andersoni was insensitive to IGP risk. Threat sensitivity in oviposition site selection was induced by experience in P. persimilis but occurred independently of experience in N. californicus. Irrespective of experience, P. persimilis laid fewer eggs in choice situations with the high- rather than low-risk IG predator. Our study suggests that, similar to classical predation, IGP may select for sophisticated innate and learned interspecific threat-sensitive antipredation responses. We argue that such responses may promote the coexistence of IG predators and prey.  相似文献   

12.
Understanding the conditions that facilitate top predator effects upon mesopredators and prey is critical for predicting where these effects will be significant. Intraguild predation (IGP) and the ecology of fear are hypotheses used to describe the effects of top predators upon mesopredators and prey species, but make different assumptions about organismal space use. The IGP hypothesis predicts that mesopredator resource acquisition and risk are positively correlated, creating a fitness deficit. But if shared prey also avoid a top predator, then mesopredators may not have to choose between risk and reward. Prey life history may be a critical predictor of how shared prey respond to predation and may mediate mesopredator suppression. We used hierarchical models of species distribution and abundance to test expectations of IGP using two separate triangular relationships between a large carnivore, smaller intraguild carnivore, and shared mammalian prey with different life histories. Following IGP, we expected that a larger carnivore would suppress a smaller carnivore if the shared prey species did not spatially avoid the large carnivore at broad scales. If prey were fearful over broad scales, we expected less evidence of mesopredator suppression. We tested these theoretical hypotheses using remote camera detections across a large spatial extent. Lagomorphs did not appear to avoid coyotes, and fox detection probability was lower as coyote abundance increased. In contrast, white‐tailed deer appeared to avoid areas of increased wolf use, and coyote detection probability was not reduced at sites where wolves occurred. These findings suggest that mesopredator suppression by larger carnivores may depend upon the behavior of shared prey, specifically the spatial scale at which they perceive risk. We further discuss how extrinsic environmental factors may contribute to mesopredator suppression.  相似文献   

13.
In this work, a simple Lotka-Volterra model of intraguild predation with three species is analysed, searching for the effect of the top predator on the coexistence with its prey-competitor species. Apart from the well-known result that the intraguild prey must be superior in the competition for the shared prey in order to make coexistence possible, the magnitude of intraguild predation and the form by which the intraguild predator makes use of the intraguild prey have important consequences upon the dynamics, extending or restricting the possibilities of coexistence. These results are easily obtained by nullcline analysis. Also, some interesting results are obtained for the same model but including saturating functional response.  相似文献   

14.
Based on the hypothesis that matching diets of intraguild (IG) predator and prey indicate strong food competition and thus intensify intraguild predation (IGP) as compared to non‐matching diets, we scrutinized diet‐dependent mutual IGP between the predatory mites Neoseiulus cucumeris and N. californicus. Both are natural enemies of herbivorous mites and insects and used in biological control of spider mites and thrips in various agricultural crops. Both are generalist predators that may also feed on plant‐derived substances such as pollen. Irrespective of diet (pollen or spider mites), N. cucumeris females had higher predation and oviposition rates and shorter attack latencies on IG prey than N. californicus. Predation rates on larvae were unaffected by diet but larvae from pollen‐fed mothers were a more profitable prey than those from spider‐mite fed mothers resulting in higher oviposition rates of IG predator females. Pollen‐fed protonymphs were earlier attacked by IG predator females than spider‐mite fed protonymphs. Spider mite‐fed N. californicus females attacked protonymphs earlier than did pollen‐fed N. californicus females. Overall, our study suggests that predator and prey diet may exert subtle influences on mutual IGP between bio‐control agents. Matching diets did not intensify IGP between N. californicus and N. cucumeris but predator and prey diets proximately influenced IGP through changes in behaviour and/or stoichiometry.  相似文献   

15.
Intraguild predation (IGP) is a widespread ecological phenomenon in which two consumers that share a resource also engage in a predator-prey interaction. Theory on IGP predicts the occurrence of alternative stable states, but empirical evidence of such states is scarce. This raises the question of whether alternative states are a rare phenomenon that is unlikely to be observed in nature. Here we analyze a model in which the resource exhibits logistic or chemostat dynamics and consumers have saturating (Type II) functional responses. We show that alternative states can arise for a wide range of biological scenarios and that environmental constraints can make their detection difficult. Our analysis identifies three possible combinations of alternative states: (i) IG prey or IG predator, (ii) coexistence or IG predator, and (iii) coexistence or IG prey. Bifurcation diagrams reveal that alternative states are possible over large regions of the parameter space. However, they can be limited to narrow ranges along the resource productivity axis, which may make it difficult to observe the occurrence of alternative states in communities with IGP. Microcosm experiments provide a promising avenue for detecting combinations of asymptotically stable states along a productivity gradient.  相似文献   

16.
Neoseiulus neobaraki and N. paspalivorus are amongst the most common phytoseiid predators of coconut mite, Aceria guerreronis, found in the spatial niche beneath coconut fruit bracts. Both predators may occur on the same coconut palms in Benin and Tanzania and are therefore likely to interact with each other. Here, we assessed cannibalism and intraguild predation (IGP) of the two predators in the absence and presence of their primary prey A. guerreronis. In the absence of the shared extraguild prey, A. guerreronis, N. neobaraki killed 19 larvae of N. paspalivorus per day and produced 0.36?eggs/female/day, while the latter species killed only 7 larvae of the former and produced 0.35?eggs/female/day. Presence of A. guerreronis only slightly decreased IGP by N. neobaraki but strongly decreased IGP by N. paspalivorus, which consumed 4-7 times less IG prey than N. neobaraki. Resulting predator offspring to IG prey ratios were, however, 4-5 times higher in N. paspalivorus than N. neobaraki. Overall, provision of A. guerreronis increased oviposition in both species. In the cannibalism tests, in the absence of A. guerreronis, N. neobaraki and N. paspalivorus consumed 1.8 and 1.2 conspecific larvae and produced almost no eggs. In the presence of abundant herbivorous prey, cannibalism dramatically decreased but oviposition increased in both N. neobaraki and N. paspalivorus. In summary, we conclude that (1) N. neobaraki is a much stronger intraguild predator than N. paspalivorus, (2) cannibalism is very limited in both species, and (3) both IGP and cannibalism are reduced in the presence of the common herbivorous prey with the exception of IGP by N. neobaraki, which remained at high levels despite presence of herbivorous prey. We discuss the implications of cannibalism and IGP on the population dynamics of A. guerreronis and the predators in view of their geographic and within-palm distribution patterns.  相似文献   

17.
Intraguild predation (IGP) among predatory species can influence many plant-arthropod associations. However, the relevance of IGP is poorly understood for truly omnivorous species such as those that can complete development on both animal and plant diets. Here we test the hypothesis that IGP among two omnivorous mirids is more common when extraguild food is either absent or not suitable. Laboratory experiments were performed in experimental cages in order to determine the effect of intraguild prey densities and diet availability on direction and intensity of IGP between Dicyphus tamaninii and Macrolophus caliginosus (Heteroptera: Miridae). Intraguild predation was symmetrical between the two mirid species in the absence of alternative food. Increasing densities of intraguild prey enhanced drastically the incidence of IGP. Intraguild predation was reduced when mirids were in the presence of green or red tomato fruits, but the presence of any other extraguild resources had no impact on IGP level. However, when given before the experiments, all resources with the exception of tomato leaves significantly reduced IGP. A second experiment was performed on live plants to compare the results of the previous trials with that obtained in a more natural setting. No IGP was observed when both mirid species were present on a plant. However, development of the intraguild prey (the more vulnerable stage) was hindered by the presence of the intraguild predator. The potential of such results is discussed from community ecology and biological control perspective.  相似文献   

18.
The theory of intraguild predation (IGP) largely studies effects on equilibrium densities of predators and prey, while experiments mostly concern transient dynamics. We studied the effects of an intraguild (IG) predator, the bug Orius laevigatus, on the population dynamics of IG-prey, the predatory mite Phytoseiulus persimilis, and a shared prey, the phytophagous two-spotted spider mite Tetranychus urticae, as well as on the performance of cucumber plants in a greenhouse. The interaction of the predatory mite and the spider mite is highly unstable, and ends either by herbivores overexploiting the plant or predators exterminating the herbivores. We studied the effect of IGP on the transient dynamics of this system, and compared the dynamics with that predicted by a simple population-dynamical model with IGP added. Behavioural studies showed that the predatory bug and the predatory mite were both attracted to plants infested by spider mites and that the two predators did not avoid plants occupied by the other predator. Observations on foraging behaviour of the predatory bug showed that it attacks and kills large numbers of predatory mites and spider mites. The model predicts strong effects of predation and prey preference by the predatory bugs on the dynamics of predatory mites and spider mites. However, experiments in which the predatory bug was added to populations of predatory mites and spider mites had little or no effect on numbers of both mite species, and cucumber plant and fruit weight.  相似文献   

19.
1. Trophic interactions between predators and parasitoids can be described as intraguild predation (IGP) and are often asymmetric. Parasitoids (typically the IG prey) may respond to the threat of IGP by mitigating the predation risk for their offspring. 2. We used a system with a facultative predator Macrolophus caliginosus, the parasitoid Aphidius colemani, and their shared prey, the aphid Myzus persicae. We examined the functional responses of the parasitoid in the presence/absence of the predator on two host plants (aubergine and sweet pepper) with differing IGP risk. 3. Estimated model parameters such as parasitoid handling time increased on both plants where the predator was present, but impact of the predator varied with plant species. The predator, which could feed herbivorously on aubergine, had a reduced impact on parasitoid foraging on that plant. IG predator presence could reduce the searching effort of the IG prey depending on the plant, and on likely predation risk. 4. The results are discussed with regard to individual parasitoid's foraging behaviour and population stability; it is suggested that the presence of the predator can contribute to the stabilisation of host–parasitoid dynamics  相似文献   

20.
Species can adjust their traits in response to selection which may strongly influence species coexistence. Nevertheless, current theory mainly assumes distinct and time‐invariant trait values. We examined the combined effects of the range and the speed of trait adaptation on species coexistence using an innovative multispecies predator–prey model. It allows for temporal trait changes of all predator and prey species and thus simultaneous coadaptation within and among trophic levels. We show that very small or slow trait adaptation did not facilitate coexistence because the stabilizing niche differences were not sufficient to offset the fitness differences. In contrast, sufficiently large and fast trait adaptation jointly promoted stable or neutrally stable species coexistence. Continuous trait adjustments in response to selection enabled a temporally variable convergence and divergence of species traits; that is, species became temporally more similar (neutral theory) or dissimilar (niche theory) depending on the selection pressure, resulting over time in a balance between niche differences stabilizing coexistence and fitness differences promoting competitive exclusion. Furthermore, coadaptation allowed prey and predator species to cluster into different functional groups. This equalized the fitness of similar species while maintaining sufficient niche differences among functionally different species delaying or preventing competitive exclusion. In contrast to previous studies, the emergent feedback between biomass and trait dynamics enabled supersaturated coexistence for a broad range of potential trait adaptation and parameters. We conclude that accounting for trait adaptation may explain stable and supersaturated species coexistence for a broad range of environmental conditions in natural systems when the absence of such adaptive changes would preclude it. Small trait changes, coincident with those that may occur within many natural populations, greatly enlarged the number of coexisting species.  相似文献   

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