Multitrophic diversity effects of network degradation

Predicting the functional consequences of biodiversity loss in realistic, multitrophic communities remains a challenge. No existing biodiversity–ecosystem function study to date has simultaneously incorporated information on species traits, network topology, and extinction across multiple trophic levels, while all three factors are independently understood as critical drivers of post‐extinction network structure and function. We fill this gap by comparing the functional consequences of simulated species loss both within (monotrophic) and across (bitrophic) trophic levels, in an ecological interaction network estimated from spatially explicit field data on tropical fecal detritus producer and consumers (mammals and dung beetles). We simulated trait‐ordered beetle and mammal extinction separately (monotrophic extinction) and the coextinction of beetles following mammal loss (bitrophic extinction), according to network structure. We also compared the diversity effects of bitrophic extinction models using a standard monotrophic function (the daily production or consumption of fecal detritus) and a unique bitrophic functional metric (the proportion of daily detritus production that is consumed). We found similar mono‐ and bitrophic diversity effects, regardless of which species traits were used to drive extinctions, yet divergent predictions when different measures of function were used. The inclusion of information on network structure had little apparent effect on the qualitative relationship between diversity and function. These results contribute to our growing understanding of the functional consequences of biodiversity from real systems and underscore the importance of species traits and realistic functional metrics to assessments of the ecosystem impacts of network degradation through species loss.     

A ratio-dependent food chain model and its applications to biological control

While biological controls have been successfully and frequently implemented by nature and human, plausible mathematical models are yet to be found to explain the often observed deterministic extinctions of both pest and control agent in such processes. In this paper we study a three trophic level food chain model with ratio-dependent Michaelis-Menten type functional responses. We shall show that this model is rich in boundary dynamics and is capable of generating such extinction dynamics. Two trophic level Michaelis-Menten type ratio-dependent predator-prey system was globally and systematically analyzed in details recently. A distinct and realistic feature of ratio-dependence is its capability of producing the extinction of prey species, and hence the collapse of the system. Another distinctive feature of this model is that its dynamical outcomes may depend on initial populations levels. Theses features, if preserved in a three trophic food chain model, make it appealing for modelling certain biological control processes (where prey is a plant species, middle predator as a pest, and top predator as a biological control agent) where the simultaneous extinctions of pest and control agent is the hallmark of their successes and are usually dependent on the amount of control agent. Our results indicate that this extinction dynamics and sensitivity to initial population levels are not only preserved, but also enriched in the three trophic level food chain model. Specifically, we provide partial answers to questions such as: under what scenarios a potential biological control may be successful, and when it may fail. We also study the questions such as what conditions ensure the coexistence of all the three species in the forms of a stable steady state and limit cycle, respectively. A multiple attractor scenario is found.     

Pattern of functional extinctions in ecological networks with a variety of interaction types

There is a strong trend of declining populations in many species of both animals and plants. Dwindling numbers of species can eventually lead to their functional extinction. Functional, or ecological, extinction occurs when a species becomes too rare to fulfill its ecological, interactive role in the ecosystem, leading to true (numerical) extinction of other depending species. Recent theoretical work on food webs suggests that the frequency of functional extinction might be surprisingly high. However, little is known about the risk of functional species extinctions in networks with other types of interactions than trophic ones. Here, we explore the frequency of functional extinctions in model ecological networks having different proportions of antagonistic and mutualistic links. Furthermore, we investigate the topological relationship between functionally and numerically extinct species. We find that (1) the frequency of functional extinctions is higher in networks containing a mixture of antagonistic and mutualistic interactions than in networks with only one type of interaction, (2) increased mortality rate of species having both mutualistic and antagonistic links is more likely to lead to extinction of another species than to extinction of the species itself compared to species having only mutualistic or antagonistic links, and (3) trophic distance (shortest path) between functionally and numerically extinct species is, on average, longer than one, indicating the importance of indirect effects. These results generalize the findings of an earlier study on food webs, demonstrating the potential importance of functional extinction in a variety of ecological network types.     

Ratio-dependent response of a temperate Australian estuarine system to sustained nitrogen loading

Classical resource- and the less studied ratio-dependent models of predator–prey relationships provide divergent predictions as to the sustained ecological effects of bottom-up forcing. While resource-dependent models, which consider only instantaneous prey density in modelling predator responses, predict community responses that are dependent on the number of trophic levels in a system, ratio-dependent models, which consider the number of prey per consumer, predict proportional increase in each level irrespective of chain length. The two models are only subtly different for systems with two or three trophic levels but in the case of four trophic levels, predict opposite effects of enrichment on primary producers. Despite the poor discriminatory power of tests of the models in systems with two or three trophic levels, field tests in estuarine and marine systems with four trophic levels have been notably absent. Sampling of phytoplankton, macroinvertebrates, invertebrate-feeding fishes, piscivorous fishes in Kooloonbung Creek, Hastings River estuary, eastern Australia, subject to over 20 years of sewage discharge, revealed increased abundances in all four trophic levels at the disturbed location relative to control sites. Increased abundance of phytoplankton at the disturbed site was counter to the predictions of resource-dependent models, which posit a reduction in the first trophic level in response to enrichment. By contrast, the increase in abundance of this first trophic level and the proportionality of increases in abundances of each of the four trophic groups to nitrogen loading provided strong support for ratio dependency. This first evidence of ratio dependence in an estuarine system with four trophic levels not only demonstrates the applicability of ecological theory which seeks to simplify the complexity of systems, but has implications for management. Although large nutrient inputs frequently induce mortality of invertebrates and fish, we have shown that smaller inputs may in fact enhance biomass of all trophic levels.     

基于定性网络模型评价生态调控情景对海洋牧场生态系统的影响——以獐子岛海洋牧场为例

在海洋牧场建设过程中,生态调控方式实施后对海洋牧场生态系统的影响通常难以预测,这对海洋牧场的生态安全和高质量发展提出了严峻挑战。为此,建立了一种基于定性网络模型(Qualitative network model, QNM)的海洋牧场生态系统模拟评价方法,并以獐子岛海洋牧场近岸增殖海域作为研究区域,构建以增殖目标种为核心的定性网络模型,模拟评估海洋牧场3种不同类型的生态调控情景(增殖目标种、移除捕食者、海藻场修复)及其复合条件下,牧场群落范围内的响应,分析海洋牧场生态调控策略与生物功能群变化之间潜在关系。结果显示:目标种增殖(仿刺参和虾夷扇贝)产生的上行效应导致其捕食者呈现积极响应,产生的下行效应导致其它底栖动物、浮游植物和有机碎屑等功能群呈现消极响应,移除捕食者海星产生的下行效应导致虾夷扇贝呈现积极响应,表明在增殖区清除敌害生物的重要性,海藻场修复对整个群落有明显的积极影响,体现了海藻场在养护近岸生态系统的重要生态意义。研究表明:QNM可有效识别生态系统潜在的营养级联效应,评估生物功能群的响应,基于QNM的海洋牧场生态调控模拟评价方法,突破了定量食物网模型在数据有限系统中使用的局限性,可为海洋牧场建设的生态调控策略制定提供科学参考。     

Conceptualising the interactive effects of climate change and biological invasions on subarctic freshwater fish

Climate change and species invasions represent key threats to global biodiversity. Subarctic freshwaters are sentinels for understanding both stressors because the effects of climate change are disproportionately strong at high latitudes and invasion of temperate species is prevalent. Here, we summarize the environmental effects of climate change and illustrate the ecological responses of freshwater fishes to these effects, spanning individual, population, community and ecosystem levels. Climate change is modifying hydrological cycles across atmospheric, terrestrial and aquatic components of subarctic ecosystems, causing increases in ambient water temperature and nutrient availability. These changes affect the individual behavior, habitat use, growth and metabolism, alter population spawning and recruitment dynamics, leading to changes in species abundance and distribution, modify food web structure, trophic interactions and energy flow within communities and change the sources, quantity and quality of energy and nutrients in ecosystems. Increases in temperature and its variability in aquatic environments underpin many ecological responses; however, altered hydrological regimes, increasing nutrient inputs and shortened ice cover are also important drivers of climate change effects and likely contribute to context‐dependent responses. Species invasions are a complex aspect of the ecology of climate change because the phenomena of invasion are both an effect and a driver of the ecological consequences of climate change. Using subarctic freshwaters as an example, we illustrate how climate change can alter three distinct aspects of species invasions: (1) the vulnerability of ecosystems to be invaded, (2) the potential for species to spread and invade new habitats, and (3) the subsequent ecological effects of invaders. We identify three fundamental knowledge gaps focused on the need to determine (1) how environmental and landscape characteristics influence the ecological impact of climate change, (2) the separate and combined effects of climate and non‐native invading species and (3) the underlying ecological processes or mechanisms responsible for changes in patterns of biodiversity.     

Complexity of multitrophic interactions in a grassland ecosystem depends on plant species diversity

1.?We studied the theoretical prediction that a loss of plant species richness has a strong impact on community interactions among all trophic levels and tested whether decreased plant species diversity results in a less complex structure and reduced interactions in ecological networks. 2.?Using plant species-specific biomass and arthropod abundance data from experimental grassland plots (Jena Experiment), we constructed multitrophic functional group interaction webs to compare communities based on 4 and 16 plant species. 427 insect and spider species were classified into 13 functional groups. These functional groups represent the nodes of ecological networks. Direct and indirect interactions among them were assessed using partial Mantel tests. Interaction web complexity was quantified using three measures of network structure: connectance, interaction diversity and interaction strength. 3.?Compared with high plant diversity plots, interaction webs based on low plant diversity plots showed reduced complexity in terms of total connectance, interaction diversity and mean interaction strength. Plant diversity effects obviously cascade up the food web and modify interactions across all trophic levels. The strongest effects occurred in interactions between adjacent trophic levels (i.e. predominantly trophic interactions), while significant interactions among plant and carnivore functional groups, as well as horizontal interactions (i.e. interactions between functional groups of the same trophic level), showed rather inconsistent responses and were generally rarer. 4.?Reduced interaction diversity has the potential to decrease and destabilize ecosystem processes. Therefore, we conclude that the loss of basal producer species leads to more simple structured, less and more loosely connected species assemblages, which in turn are very likely to decrease ecosystem functioning, community robustness and tolerance to disturbance. Our results suggest that the functioning of the entire ecological community is critically linked to the diversity of its component plants species.     

The form of direct interspecific competition modifies secondary extinction patterns in multi‐trophic food webs

Forcibly removing species from ecosystems has important consequences for the remaining assemblage, leading to changes in community structure, ecosystem functioning and secondary (cascading) extinctions. One key question that has arisen from single‐ and multi‐trophic ecosystem models is whether the secondary extinctions that occur within competitive communities (guilds) are also important in multi‐trophic ecosystems? The loss of consumer–resource links obviously causes secondary extinction of specialist consumers (topological extinctions), but the importance of secondary extinctions in multi‐trophic food webs driven by direct competitive exclusion remains unknown. Here I disentangle the effects of extinctions driven by basal competitive exclusion from those caused by trophic interactions in a multi‐trophic ecosystem (basal producers, intermediate and top consumers). I compared food webs where basal species either show diffuse (all species compete with each other identically: no within guild extinctions following primary extinction) or asymmetric competition (unequal interspecific competition: within guild extinctions are possible). Basal competitive exclusion drives extra extinction cascades across all trophic levels, with the effect amplified in larger ecosystems, though varying connectance has little impact on results. Secondary extinction patterns based on the relative abundance of the species lost in the primary extinction differ qualitatively between diffuse and asymmetric competition. Removing asymmetric basal species with low (high) abundance triggers fewer (more) secondary extinctions throughout the whole food web than removing diffuse basal species. Rare asymmetric competitors experience less pressure from consumers compared to rare diffuse competitors. Simulations revealed that diffuse basal species are never involved in extinction cascades, regardless of the trophic level of a primary extinction, while asymmetric competitors were. This work highlights important qualitative differences in extinction patterns that arise when different assumptions are made about the form of direct competition in multi‐trophic food webs.     

Analyzing trophic ecosystem functions with the interaction functional space

Quantifying the vulnerability of ecosystems to global change requires a better understanding of how trophic ecosystem functions emerge. So far, trophic ecosystem functions have been studied from the perspective of either functional diversity or network ecology. To integrate these two perspectives, we propose the interaction functional space (IFS) a conceptual framework to simultaneously analyze the effects of traits and interactions on trophic functions. We exemplify the added value of our framework for seed dispersal and wood decomposition and show how species interactions influence the relationship between functional trait diversity and trophic functions. We propose future applications for a range of functions where the IFS can help to elucidate mechanisms underpinning trophic functions and facilitate understanding of functional changes in ecosystems amidst global change.     

The importance of ecological memory for trophic rewilding as an ecosystem restoration approach

Increasing human pressure on strongly defaunated ecosystems is characteristic of the Anthropocene and calls for proactive restoration approaches that promote self‐sustaining, functioning ecosystems. However, the suitability of novel restoration concepts such as trophic rewilding is still under discussion given fragmentary empirical data and limited theory development. Here, we develop a theoretical framework that integrates the concept of ‘ecological memory’ into trophic rewilding. The ecological memory of an ecosystem is defined as an ecosystem's accumulated abiotic and biotic material and information legacies from past dynamics. By summarising existing knowledge about the ecological effects of megafauna extinction and rewilding across a large range of spatial and temporal scales, we identify two key drivers of ecosystem responses to trophic rewilding: (i) impact potential of (re)introduced megafauna, and (ii) ecological memory characterising the focal ecosystem. The impact potential of (re)introduced megafauna species can be estimated from species properties such as lifetime per capita engineering capacity, population density, home range size and niche overlap with resident species. The importance of ecological memory characterising the focal ecosystem depends on (i) the absolute time since megafauna loss, (ii) the speed of abiotic and biotic turnover, (iii) the strength of species interactions characterising the focal ecosystem, and (iv) the compensatory capacity of surrounding source ecosystems. These properties related to the focal and surrounding ecosystems mediate material and information legacies (its ecological memory) and modulate the net ecosystem impact of (re)introduced megafauna species. We provide practical advice about how to quantify all these properties while highlighting the strong link between ecological memory and historically contingent ecosystem trajectories. With this newly established ecological memory–rewilding framework, we hope to guide future empirical studies that investigate the ecological effects of trophic rewilding and other ecosystem‐restoration approaches. The proposed integrated conceptual framework should also assist managers and decision makers to anticipate the possible trajectories of ecosystem dynamics after restoration actions and to weigh plausible alternatives. This will help practitioners to develop adaptive management strategies for trophic rewilding that could facilitate sustainable management of functioning ecosystems in an increasingly human‐dominated world.     

Do asynchronies in extinction debt affect the structure of trophic networks? A case study of antagonistic butterfly larvae–plant networks

Habitat loss and fragmentation affect species richness in fragmented habitats and can lead to immediate or time‐delayed species extinctions. Asynchronies in extinction and extinction debt between interacting species may have severe effects on ecological networks. However, these effects remain largely unknown. We evaluated the effects of habitat patch and landscape changes on antagonistic butterfly larvae–plant trophic networks in Mediterranean grasslands in which previous studies had shown the existence of extinction debt in plants but not in butterflies. We sampled current species richness of habitat‐specialist and generalist butterflies and vascular plants in 26 grasslands. We assessed the direct effects of historical and current patch and landscape characteristics on species richness and on butterfly larvae–plant trophic network metrics and robustness. Although positive species‐ and interactions–area relationships were found in all networks, structure and robustness was only affected by patch and landscape changes in networks involving the subset of butterfly specialists. Larger patches had more species (butterflies and host plants) and interactions but also more compartments, which decreased network connectance but increased network stability. Moreover, most likely due to the rescue effect, patch connectivity increased host‐plant species (but not butterfly) richness and total links, and network robustness in specialist networks. On the other hand, patch area loss decreased robustness in specialist butterfly larvae–plant networks and made them more prone to collapse against host plant extinctions. Finally, in all butterfly larvae–plant networks we also detected a past patch and landscape effect on network asymmetry, which indicates that there were different extinction rates and extinction debts for butterflies and host plants. We conclude that asynchronies in extinction and extinction debt in butterfly–plant networks provoked by patch and landscape changes caused changes in species richness and network links in all networks, as well as changes in network structure and robustness in specialist networks.     

The ecological drivers and consequences of wildlife trade

Wildlife trade is a key driver of extinction risk, affecting at least 24% of terrestrial vertebrates. The persistent removal of species can have profound impacts on species extinction risk and selection within populations. We draw together the first review of characteristics known to drive species use – identifying species with larger body sizes, greater abundance, increased rarity or certain morphological traits valued by consumers as being particularly prevalent in trade. We then review the ecological implications of this trade-driven selection, revealing direct effects of trade on natural selection and populations for traded species, which includes selection against desirable traits. Additionally, there exists a positive feedback loop between rarity and trade and depleted populations tend to have easy human access points, which can result in species being harvested to extinction and has the potential to alter source–sink dynamics. Wider cascading ecosystem repercussions from trade-induced declines include altered seed dispersal networks, trophic cascades, long-term compositional changes in plant communities, altered forest carbon stocks, and the introduction of harmful invasive species. Because it occurs across multiple scales with diverse drivers, wildlife trade requires multi-faceted conservation actions to maintain biodiversity and ecological function, including regulatory and enforcement approaches, bottom-up and community-based interventions, captive breeding or wildlife farming, and conservation translocations and trophic rewilding. We highlight three emergent research themes at the intersection of trade and community ecology: (1) functional impacts of trade; (2) altered provisioning of ecosystem services; and (3) prevalence of trade-dispersed diseases. Outside of the primary objective that exploitation is sustainable for traded species, we must urgently incorporate consideration of the broader consequences for other species and ecosystem processes when quantifying sustainability.     

Intraspecific diversity loss in a predator species alters prey community structure and ecosystem functions

Loss in intraspecific diversity can alter ecosystem functions, but the underlying mechanisms are still elusive, and intraspecific biodiversity–ecosystem function (iBEF) relationships have been restrained to primary producers. Here, we manipulated genetic and functional richness of a fish consumer (Phoxinus phoxinus) to test whether iBEF relationships exist in consumer species and whether they are more likely sustained by genetic or functional richness. We found that both genotypic and functional richness affected ecosystem functioning, either independently or interactively. Loss in genotypic richness reduced benthic invertebrate diversity consistently across functional richness treatments, whereas it reduced zooplankton diversity only when functional richness was high. Finally, losses in genotypic and functional richness altered functions (decomposition) through trophic cascades. We concluded that iBEF relationships lead to substantial top-down effects on entire food chains. The loss of genotypic richness impacted ecological properties as much as the loss of functional richness, probably because it sustains “cryptic” functional diversity.

Global change is expected to generate a loss of intraspecific diversity worldwide. This mesocosm study explores whether loss of genetic and functional diversity in a predator species affects community and ecosystem functioning of lower trophic levels in pond ecosystems, revealing that diversity loss in a single consumer species can impact an entire ecosystem, reducing its functionality.     

Niche width collapse in a resilient top predator following ecosystem fragmentation

Much research has focused on identifying species that are susceptible to extinction following ecosystem fragmentation, yet even those species that persist in fragmented habitats may have fundamentally different ecological roles than conspecifics in unimpacted areas. Shifts in trophic role induced by fragmentation, especially of abundant top predators, could have transcendent impacts on food web architecture and stability, as well as ecosystem function. Here we use a novel measure of trophic niche width, based on stable isotope ratios, to assess effects of aquatic ecosystem fragmentation on trophic ecology of a resilient, dominant, top predator. We demonstrate collapse in trophic niche width of the predator in fragmented systems, a phenomenon related to significant reductions in diversity of potential prey taxa. Collapsed niche width reflects a homogenization of energy flow pathways to top predators, likely serving to destabilize remnant food webs and render apparently resilient top predators more susceptible to extinction through time.     

Functional responses and predator–prey models: a critique of ratio dependence

Arditi and Ginzburg (2012) propose ordinary differential equations (ODEs) with ratio-dependent functional responses as the new null model for predation, based on their earlier work on ratio-dependent food chains and a number of functional response measurements. Here, I discuss some of their claims, arguing for a flexible and problem-driven approach to predator–prey modeling. Models to understand population cycles and models to predict the effect of basal enrichment on food chains need not be the same. While ratio-dependent functional responses in ODE models might sometimes be useful as limit cases for food chains, they are not intrinsically more useful than prey-dependent models to understand the effect of a given predator on prey population dynamics—and sometimes less useful, given the small temporal scales considered in many models. “Instantism” is showed to be an invalid criticism when ODEs are interpreted as describing average trajectories of stochastic birth–death processes. Moreover, other modeling frameworks with strong ties to time series statistics, such as stochastic difference equations, should be promoted to improve the feedback loop between field and theoretical research. The main problems of current trophic ecology do not lie in a wrong null model, as ecologists have already several at their disposal. The loose connection of ODE models with empirical data and spatial/temporal scaling up of empirical measurements constitute more serious challenges to our understanding of trophic interactions and their consequences on ecosystem functioning.     

Rapid development of indirect effects in ecological networks

Indirect effects are important components of ecological and evolutionary interactions that may maintain biodiversity, enable or inhibit invasive species, and challenge ecosystem assessment and management. A central hypothesis of Network Environ Analysis (NEA), one type of ecological network analysis, is that indirect flows tend to dominate direct flows in ecosystem networks of conservative substance exchanges. However, current NEA methods assume that these ecosystems are stationary (i.e. time invariant exchange rates), which is unlikely to be true for many ecosystems for interesting time and space scales. For the work reported here, we investigated the sensitivity of the dominance of indirect effects hypothesis to the stationary modeling assumption by determining the development rate of indirect effects and flow intensity, as expressed as the number of transfer steps, in thirty‐one ecosystem models. We hypothesized that indirect effects develop rapidly in ecological networks, but that they would develop faster in biogeochemically based models than in trophically based models. In contrast, our results show that indirect effects develop rapidly in all thirty‐one models examined. In 94% of the models, indirect flows exceeded direct flows by a pathway length of 3. This indicates that ecological systems do not need to maintain a particular configuration for long for indirect effects to dominate. Thus, the dominance of indirect effects hypothesis remains plausible. We also found that biogeochemical models tended to require more of the extended path network than the trophic models to account for 50% and 95% of the total system activity, but that both types of models required more of the power series than is typically considered in engineered systems. These results succinctly illustrate the complexity of ecological systems and help explain why they are challenging to assess and manage.     

Functional trait diversity across trophic levels determines herbivore impact on plant community biomass

Understanding the consequences of trophic interactions for ecosystem functioning is challenging, as contrasting effects of species and functional diversity can be expected across trophic levels. We experimentally manipulated functional identity and diversity of grassland insect herbivores and tested their impact on plant community biomass. Herbivore resource acquisition traits, i.e. mandible strength and the diversity of mandibular traits, had more important effects on plant biomass than body size. Higher herbivore functional diversity increased overall impact on plant biomass due to feeding niche complementarity. Higher plant functional diversity limited biomass pre‐emption by herbivores. The functional diversity within and across trophic levels therefore regulates the impact of functionally contrasting consumers on primary producers. By experimentally manipulating the functional diversity across trophic levels, our study illustrates how trait‐based approaches constitute a promising way to tackle existing links between trophic interactions and ecosystem functioning.     

Invasive plants negatively impact native,but not exotic,animals

Despite our growing understanding of the impacts of invasive plants on ecosystem structure and function, important gaps remain, including whether native and exotic species respond differently to plant invasion. This would elucidate basic ecological interactions and inform management. We performed a meta‐analytic review of the effects of invasive plants on native and exotic resident animals. We found that invasive plants reduced the abundance of native, but not exotic, animals. This varied by animal phyla, with invasive plants reducing the abundance of native annelids and chordates, but not mollusks or arthropods. We found dissimilar impacts among “wet” and “dry” ecosystems, but not among animal trophic levels. Additionally, the impact of invasive plants increased over time, but this did not vary with animal nativity. Our review found that no studies considered resident nativity differences, and most did not identify animals to species. We call for more rigorous studies of invaded community impacts across taxa, and most importantly, explicit consideration of resident biogeographic origin. We provide an important first insight into how native and exotic species respond differently to invasion, the consequences of which may facilitate cascading trophic disruptions further exacerbating global change consequences to ecosystem structure and function.     

Population-level metrics of trophic structure based on stable isotopes and their application to invasion ecology

Biological invasions are a significant driver of human-induced global change and many ecosystems sustain sympatric invaders. Interactions occurring among these invaders have important implications for ecosystem structure and functioning, yet they are poorly understood. Here we apply newly developed metrics derived from stable isotope data to provide quantitative measures of trophic diversity within populations or species. We then use these to test the hypothesis that sympatric invaders belonging to the same functional feeding group occupy a smaller isotopic niche than their allopatric counterparts. Two introduced, globally important, benthic omnivores, Louisiana swamp crayfish (Procambarus clarkii) and carp (Cyprinus carpio), are sympatric in Lake Naivasha, Kenya. We applied our metrics to an 8-year data set encompassing the establishment of carp in the lake. We found a strong asymmetric interaction between the two invasive populations, as indicated by inverse correlations between carp abundance and measures of crayfish trophic diversity. Lack of isotopic niche overlap between carp and crayfish in the majority of years indicated a predominantly indirect interaction. We suggest that carp-induced habitat alteration reduced the diversity of crayfish prey, resulting in a reduction in the dietary niche of crayfish. Stable isotopes provide an integrated signal of diet over space and time, offering an appropriate scale for the study of population niches, but few isotope studies have retained the often insightful information revealed by variability among individuals in isotope values. Our population metrics incorporate such variation, are robust to the vagaries of sample size and are a useful additional tool to reveal subtle dietary interactions among species. Although we have demonstrated their applicability specifically using a detailed temporal dataset of species invasion in a lake, they have a wide array of potential ecological applications.     

Dynamics of a plant-herbivore-predator system with plant-toxicity

A system of ordinary differential equations is considered that models the interactions of two plant species populations, an herbivore population, and a predator population. We use a toxin-determined functional response to describe the interactions between plant species and herbivores and use a Holling Type II functional response to model the interactions between herbivores and predators. In order to study how the predators impact the succession of vegetation, we derive invasion conditions under which a plant species can invade into an environment in which another plant species is co-existing with a herbivore population with or without a predator population. These conditions provide threshold quantities for several parameters that may play a key role in the dynamics of the system. Numerical simulations are conducted to reinforce the analytical results. This model can be applied to a boreal ecosystem trophic chain to examine the possible cascading effects of predator-control actions when plant species differ in their levels of toxic defense.