Canards and mixed-mode oscillations in a forest pest model

We consider a three-variable forest pest model, proposed by Rinaldi & Muratori (1992) [Rinaldi, S., Muratori, S., 1992. Limit cycles in slow–fast forest–pest models. Theor. Popul. Biol. 41, 26–43]. The model allows relaxation oscillations where long pest-free periods are interspersed with outbreaks of high pest concentration. For small values of the timescale of the young trees, the model can be reduced to a two-dimensional model. By a geometrical analysis we identify a canard explosion in the reduced model, that is, a change over a narrow parameter interval from outbreak dynamics to small oscillations around an endemic state. For larger values of the timescale of the young trees the two-dimensional approximation breaks down, and a broader parameter interval with mixed-mode oscillations appear, replacing the simple canard explosion. The analysis only relies on simple and generic properties of the model, and is expected to be applicable in a larger class of multiple timescale dynamical models.     

Links between plant community composition,soil organic matter quality and microbial communities in contrasting tundra habitats

Plant communities, soil organic matter and microbial communities are predicted to be interlinked and to exhibit concordant patterns along major environmental gradients. We investigated the relationships between plant functional type composition, soil organic matter quality and decomposer community composition, and how these are related to major environmental variation in non-acid and acid soils derived from calcareous versus siliceous bedrocks, respectively. We analysed vegetation, organic matter and microbial community compositions from five non-acidic and five acidic heath sites in alpine tundra in northern Europe. Sequential organic matter fractionation was used to characterize organic matter quality and phospholipid fatty acid analysis to detect major variation in decomposer communities. Non-acidic and acidic heaths differed substantially in vegetation composition, and these disparities were associated with congruent shifts in soil organic matter and microbial communities. A high proportion of forbs in the vegetation was positively associated with low C:N and high soluble N:phenolics ratios in soil organic matter, and a high proportion of bacteria in the microbial community. On the contrary, dwarf shrub-rich vegetation was associated with high C:N and low soluble N:phenolics ratios, and a high proportion of fungi in the microbial community. Our study demonstrates a strong link between the plant community composition, soil organic matter quality, and microbial community composition, and that differences in one compartment are paralleled by changes in others. Variation in the forb-shrub gradient of vegetation may largely dictate variations in the chemical quality of organic matter and decomposer communities in tundra ecosystems. Soil pH, through its direct and indirect effects on plant and microbial communities, seems to function as an ultimate environmental driver that gives rise to and amplifies the interactions between above- and belowground systems. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Acidification and alkalinization of soils

Acidification or alkalinization of soils occurs through H⁺ transfer processes involving vegetation, soil solution and soil minerals. A permanent change in the acid neutralizing capacity of the inorganic soil fraction (ANC_(s)),i.e. soil acidification (ΔANC<0) or soil alkalinization (ΔANC>0), results from an irreversible H⁺ flux. This irreversible H⁺ flux can be caused either by direct proton addition or depletion, by different mobility of components of the ANC_(s) or by a permanent change in redox conditions. The contributions of (a) acidic atmospheric deposition, (b) nitrogen transformations, (c) deprotonation of CO₂ and of organic acids and protonation of their conjugate bases, (d) assimilation of cations and anions by the vegetation, (e) weathering or reverse weathering of minerals and (f) stream output to changes in the ANC_(s) are illustrated by means of H⁺ budgets for actual soils and watersheds.     

Using mean first passage times to quantify equilibrium resilience in perturbed intraguild predation systems

Regime shift inducibility depends on equilibrium resilience, which depends on species interactions. When species interactions include intraguild predation (IGP), integrated pest management may induce regime shifts because enhancing the abundance of intraguild predators simultaneously increases competition with, and predation on, invasive prey. To explore the dynamical consequences of such manipulations, we use a bistable, deterministic IGP model with stochastic removals that perturb invader density from the high-density equilibrium. We quantify the combined effects of IGP and such perturbations in terms of mean first passage times (MFPTs) to target invader densities such as thresholds between regimes. Analytical MFPTs compare favorably with those generated by Monte Carlo numerical solutions of the stochastically perturbed IGP model. MFPTs can therefore usefully quantify equilibrium resilience in terms of perturbation schedules. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Evaluation of pest control efficiencies for different banker plant systems with a simple predator–prey model

The banker plant system has been introduced for the biological control of various pest species in Japanese greenhouses. With the banker plant system, non-crop plants infested with a host insect (a non-commercial crop pest) are placed in the greenhouse to provide alternative resources for the parasitoids or predators. We want to evaluate the effectiveness for controlling pests on the crop in a quantitative way by immigrating predators from the banker plant. Therefore, we developed a simple model for the interaction of the pest and predator in the crop and included the banker plant only as a source for predators. For three different pest-predator systems we parameterised the model and used these models to predict under what conditions biological control in a banker plant system is successful. We defined successful as keeping the pest below the economic injury level of the crop estimated from damage analysis. Because the crop is mostly grown during a period that lasts less than a year our analysis should not only focus on the equilibrium dynamics. In contrast, it should also focus on the transient dynamics. Our main analytical result, from the equilibrium analysis, is that for successful control the maximum lifetime consumption of immigrating predators should exceed the daily prey growth at half the value of the maximum consumption rate. For practical purpose this translates into the fact that the immigration of predators at a low initial pest density is crucial for successful control.     

Models for the spread of universally fatal diseases

In the formulation of models of S-I-R type for the spread of communicable diseases it is necessary to distinguish between diseases with recovery with full immunity and diseases with permanent removal by death. We consider models which include nonlinear population dynamics with permanent removal. The principal result is that the stability of endemic equilibrium may depend on the population dynamics and on the distribution of infective periods; sustained oscillations are possible in some cases.     

Oscillations in a patchy environment disease model

For a single patch SIRS model with a period of immunity of fixed length, recruitment-death demographics, disease related deaths and mass action incidence, the basic reproduction number R(0) is identified. It is shown that the disease-free equilibrium is globally asymptotically stable if R(0)<1. For R(0)>1, local stability of the endemic equilibrium and Hopf bifurcation analysis about this equilibrium are carried out. Moreover, a practical numerical approach to locate the bifurcation values for a characteristic equation with delay-dependent coefficients is provided. For a two patch SIRS model with travel, it is shown that there are several threshold quantities determining its dynamic behavior and that travel can reduce oscillations in both patches; travel may enhance oscillations in both patches; or travel can switch oscillations from one patch to another.     

五大连池新期火山熔岩台地不同植被类型土壤微生物群落的功能多样性

土壤微生物群落是陆地生态系统的重要生物活性成分,其结构和功能多样性直接影响到系统的碳、氮等生态过程,微生物群落功能多样性与地上植被类型变化密切相关,开展植被类型对土壤微生物群落功能多样性的影响研究具有重要意义。以五大连池新期火山熔岩台地苔藓、草本、灌丛、矮曲林、针阔混交林5种典型植被类型为对象,利用BIOLOG微孔板法研究不同演替阶段植被类型土壤微生物群落功能多样性特征。结果表明：不同植被类型土壤微生物群落功能多样性存在显著差异。平均颜色变化率（AWCD）随培养时间延长而逐渐增加,大小顺序为：苔藓 > 针阔混交林 > 矮曲林 > 草本 > 灌丛。灌丛土壤微生物多样性指数与其他植被类型间差异显著。主成分分析结果表明,主成分1和主成分2分别能解释变量方差的56.24%和29.59%,不同植被类型下土壤微生物的碳源利用格局差异主要是由氨基酸类和带磷基糖类引起,二者合计解释总变异量的47.51%。冗余分析表明,速效磷、铵态氮、C：N和pH对微生物功能多样性具有显著的影响,羧酸类、氨基酸类、酯类和胺类的降解更易受到环境因素的影响。研究结果为进一步探讨植被类型与土壤微生物之间在植被演替过程中的关系提供参考。     

Microbial biomass,respiration and diversity in ultramafic soils of West Dome,New Zealand

Ultramafic soils are colonised by plant communities adapted to naturally elevated heavy metal content but it is not known whether soil microbial communities are similarly adapted to heavy metals. We measured microbial properties of six ultramafic soils that ranged in heavy metal content to test whether microbial diversity would decrease and respiratory quotient (microbial respiration:biomass) increase due to the stress imposed by increasing metal content. Soil samples were collected from beneath Nothofagus solandri var. cliffortioides tall forest, tall Leptospermum scoparium shrubland, open Leptospermum scoparium shrubland, an open Leptospermum scoparium shrubland with the rare ultramafic endemic Celmisia spedenii, a mixed divaricate shrubland, and a red tussock (Chionochloa rubra) grassland on the Dun Mountain Ophiolite belt, New Zealand. Samples were analysed for catabolic evenness using the catabolic response profile technique, microbial biomass, microbial respiration, and soil properties (pH, total carbon, total nitrogen, magnesium and total or extractable chromium and nickel). The sites differed in base saturation, pH and concentrations of metals, particularly magnesium, chromium and nickel, properties that are a major determinant of the plant communities that develop. Microbial biomass and respiration, catabolic evenness (range of 19.1 to 22.7) and the respiratory quotient were not correlated to any of the measured soil chemical properties. Factor analysis of the respiratory responses showed that the microbial communities under each vegetation type were distinct. The second factor extracted was correlated to total carbon (r ²=0.62, P<0.01), basal respiration (r ²=0.55, P<0.01) and microbial biomass (r ²=0.65, P<0.01). Increasing metals concentrations had no direct impact on microbial diversity, biomass, respiration or community energetics. However, we suggest that metal concentrations may have exerted an indirect effect on the structure of the microbial communities through control of the vegetation community and litter inputs of carbon to the soil.     

Short-term effects on sheep pastureland due to grazing abandonment in a Western Mediterranean island ecosystem: A multidisciplinary approach

Changes of pasture communities consequent to management practices resulting from land abandonment considerably affect the structure and function of the ecosystem. This study analyses the consequences of grazing abandonment in terms of plant and soil microbial diversity and fertility, on a Mediterranean upland sheep pasture, over a short period (five years). Grazing was experimentally excluded by fencing ten 10×10 m permanent plots within an area that had supported grazing until 2000, by 0.23 sheep ha^?1. Plant and soil microbial communities and physicochemical parameters were monitored within the fenced and unfenced control plots, during three sampling times from 2000 (before the fencing) to 2005. Grazing cessation notably altered the floral composition, with an average dissimilarity of 96.7% between the vegetation communities, over five years. No significant change occurred in the control plots that were grazed throughout the sampling period. This work highlighted that, over a short term, the structural change in the specific plant composition affected only the grass species, confirming that grazing favours the small-sized species over the annual species. Further, it was evident that species groups of conservational and phytogeographic interest, like the endemic and Mediterranean-Atlantic species, tended to disappear with pasture abandonment and were substituted by more widespread species throughout the Mediterranean or even the world. Pasture abandonment was accompanied by an increase of soil pH and a decrease in soil organic matter and soil nitrogen. The microbial parameters recorded at three different sampling times revealed a substantial effect of the plant community, or the time of grazing abandonment, on soil microbial abundance and diversity. Considerable importance is given to the consequences of pasture abandonment on the conservation of plant and microbial diversity and on soil fertility.     

喀斯特土壤固氮微生物群落与植被、土壤的关系

固氮微生物作为土壤中重要的功能微生物群之一,其与地上植物群落、土壤环境之间的关系尚不清楚。在桂西北的环江县、都安县和大化县选取喀斯特典型植被类型(草丛、灌丛、次生林)建立样方,通过植被调查、测定土壤理化性质和构建克隆文库的方法,研究了土壤固氮微生物群落的结构与组成,分析了固氮微生物群落与植物群落、土壤理化性质之间的关系。结果表明:研究区典型植被类型土壤中的优势固氮微生物为慢生根瘤菌。Mantel相关性分析表明植物群落与固氮微生物群落显著相关(r=0.6116,P=0.011);结合PCo A分析和Venn图可看出,植物群落组成与结构越相似,土壤固氮微生物群落结构与组成也越相似。CCA分析前两轴的解释率之和仅为22.72%,其中总氮、有效态钾、有效态钙对固氮微生物群落的影响显著,这表明本研究涵盖的土壤理化性质指标并不能完全解释固氮微生物群落的变异,需要补充更多的土壤数据进行更深入的研究。由此可见,在喀斯特生态恢复过程中,不仅要关注地上植被群落的恢复与重建,同时也应重视地下功能微生物群落的恢复与重建。     

Plant richness and composition in hardwood forest understories vary along an acidic deposition and soil-chemical gradient in the northeastern United States

Aims

A century of atmospheric deposition of sulfur and nitrogen has acidified soils and undermined the health and recruitment of foundational tree species in the northeastern US. However, effects of acidic deposition on the forest understory plant communities of this region are poorly documented. We investigated how forest understory plant species composition and richness varied across gradients of acidic deposition and soil acidity in the Adirondack Mountains of New York State.

Methods

We surveyed understory vegetation and soils in hardwood forests on 20 small watersheds and built models of community composition and richness as functions of soil chemistry, nitrogen and sulfur deposition, and other environmental variables.

Results

Community composition varied significantly with gradients of acidic deposition, soil acidity, and base cation availability (63% variance explained). Several species increased with soil acidity while others decreased. Understory plant richness decreased significantly with increasing soil acidity (r?=?0.60). The best multivariate regression model to predict richness (p?<?0.001, adjusted-R²?=?0.60) reflected positive effects of pH and carbon-to-nitrogen ratio (C:N).

Conclusions

The relationship we found between understory plant communities and a soil-chemical gradient, suggests that soil acidification can reduce diversity and alter the composition of these communities in northern hardwood forests exposed to acidic deposition.

     

Asynchronous responses of soil microbial community and understory plant community to simulated nitrogen deposition in a subtropical forest

Atmospheric nitrogen (N) deposition greatly affects ecosystem processes and properties. However, few studies have simultaneously examined the responses of both the above- and belowground communities to N deposition. Here, we investigated the effects of 8 years of simulated N deposition on soil microbial communities and plant diversity in a subtropical forest. The quantities of experimental N added (g of N m⁻² year⁻¹) and treatment codes were 0 (N0, control), 6 (N1), 12 (N2), and 24 (N3). Phospholipid fatty acids (PLFAs) analysis was used to characterize the soil microbial community while plant diversity and coverage were determined in the permanent field plots. Microbial abundance was reduced by the N3 treatment, and plant species richness and coverage were reduced by both N2 and N3 treatments. Declines in plant species richness were associated with decreased abundance of arbuscular mycorrhizal fungi, increased bacterial stress index, and reduced soil pH. The plasticity of soil microbial community would be more related to the different responses among treatments when compared with plant community. These results indicate that long-term N deposition has greater effects on the understory plant community than on the soil microbial community and different conservation strategies should be considered.     

Above‐ and belowground linkages in Sphagnum peatland: climate warming affects plant‐microbial interactions

Peatlands contain approximately one third of all soil organic carbon (SOC). Warming can alter above‐ and belowground linkages that regulate soil organic carbon dynamics and C‐balance in peatlands. Here we examine the multiyear impact of in situ experimental warming on the microbial food web, vegetation, and their feedbacks with soil chemistry. We provide evidence of both positive and negative impacts of warming on specific microbial functional groups, leading to destabilization of the microbial food web. We observed a strong reduction (70%) in the biomass of top‐predators (testate amoebae) in warmed plots. Such a loss caused a shortening of microbial food chains, which in turn stimulated microbial activity, leading to slight increases in levels of nutrients and labile C in water. We further show that warming altered the regulatory role of Sphagnum‐polyphenols on microbial community structure with a potential inhibition of top predators. In addition, warming caused a decrease in Sphagnum cover and an increase in vascular plant cover. Using structural equation modelling, we show that changes in the microbial food web affected the relationships between plants, soil water chemistry, and microbial communities. These results suggest that warming will destabilize C and nutrient recycling of peatlands via changes in above‐ and belowground linkages, and therefore, the microbial food web associated with mosses will feedback positively to global warming by destabilizing the carbon cycle. This study confirms that microbial food webs thus constitute a key element in the functioning of peatland ecosystems. Their study can help understand how mosses, as ecosystem engineers, tightly regulate biogeochemical cycling and climate feedback in peatlands     

Models of population genetics of Batesian mimicry

A mathematical model has been set up to investigate the changes in frequency of a gene inducing Batesian mimicry, due to selective predation. The selective force acting on this gene is frequency dependent since the tendency of predators to eat the mimic varies with the abundance of the mimic itself relative to the appropriate distasteful model. Cases have been investigated both where the mimicry is limited to one sex only and where both sexes can be mimetic. This kind of selection can eventually lead to three different results: (a) fixation of the mimetic gene; (b) an interval of neutral equilibria; (c) a unique, nontrivial equilibrium. In the last case, both the trivial equilibria are unstable, both locally and globally, whereas the intermediate equilibrium can be stable or unstable. When this occurs, the gene frequency eventually undergoes stable oscillations. The results actually obtained depend mainly on the behavior of the predators and the abundance of the mimic relative to the model.     

Responses of soil microorganisms to resource availability in urban, desert soils

Terrestrial desert ecosystems are strongly structured by the distribution of plants, which concentrate resources and create islands of fertility relative to interplant spaces. Atmospheric nitrogen (N) deposition resulting from urbanization has the potential to change those spatial patterns via resource inputs, resulting in more homogeneous soil resource availability. We sampled soils at 12 desert remnant sites around Phoenix, Arizona along a model-predicted gradient in N deposition to determine the degree to which deposition has altered spatial patterns in soil resource availability and microbial activity. Soil microbial biomass and abundance were not influenced by atmospheric N deposition. Instead, plant islands remained strong organizers of soil microbial processes. These islands of fertility exhibited elevated pools of resources, microbial abundance, and activity relative to interspaces. In both plant islands and interspaces, soil moisture and soil N concentrations predicted microbial biomass and abundance. Following experimental wetting, carbon dioxide (CO₂) flux from soil of interspaces was positively correlated with N deposition, whereas in plant islands, soil CO₂ flux was positively correlated with soil moisture content and soil organic matter. Soil CO₂ flux in both patch types showed rapid and short-lived responses to precipitation, demonstrating the brief time scales during which soil biota may process deposited materials. Although we observed patterns consistent with N limitation of microbes in interspaces, we conclude that atmospheric N deposition likely accumulates in soils because microbes are primarily limited by water and secondarily by carbon or nitrogen. Soil microbial uptake of atmospherically deposited N likely occurs only during sparse and infrequent rainfall.     

Ecosystem recovery: heathland response to a reduction in nitrogen deposition

Atmospheric deposition of nitrogen is responsible for widespread changes in the structure and function of sensitive seminatural ecosystems. The proposed reduction in emissions of nitrogenous pollutants in Europe under the Gothenburg Protocol raises the question of whether affected ecosystems have the potential to recover to their previous condition and, if so, over what timescale. Since 1998, we have monitored the response of a lowland heathland in southern England following the cessation of a long‐term nitrogen addition experiment, and subsequent management, assessing changes in vegetation growth and chemistry, soil chemistry and the soil microbial community. Persistent effects of earlier nutrient loading on Calluna growth and phenology, and on the abundance of lichens, were apparent up to 8 years after nitrogen additions ceased, indicating the potential for long‐term effects of modest nutrient loading (up to 15.4 kg N ha^?1 yr^?1, over 7 years) on heathland ecosystems. The size and activity of the soil microbial community was elevated in former N‐treated plots, 6–8 years after additions ceased, suggesting a prolonged effect on the rate of nutrient cycling. Although habitat management in 1998 reduced nitrogen stores in plant biomass, effects on belowground nitrogen stores were small. Although some parameters (e.g. soil pH) recover pretreatment levels relatively rapidly, others (e.g. vegetation cover and microbial activity) respond much more slowly, indicating that the ecological effects of even small increases in nitrogen deposition will persist for many years after deposition inputs are reduced. Indeed, calculations suggest that the additional soil nitrogen storage associated with 7 years of experimental nitrogen inputs could sustain the observed effects on plant growth and phenology for several decades. Carry over effects on plant phenology and sensitivity to drought suggest that the persistence of vegetation responses to nitrogen deposition should be integrated into long‐term assessments of the impact of global climate change on sensitive ecosystems.     

Subsidy from the detrital food web, but not microhabitat complexity, affects the role of generalist predators in an aboveground herbivore food web

The activity and density of generalist predators, such as carabid beetles, rove beetles and spiders, may increase in response to: (1) increased availability of prey from the belowground subsystem and/or (2) enhanced complexity of aboveground vegetation. Organic farming practices support decomposer populations and enhance habitat complexity due to an increased weed density. A response by generalist predators to such below‐ or aboveground changes could affect predation rates on herbivores in the aboveground food web. We tested this hypothesis in a replicated field experiment conducted in a winter wheat field, where increased predator activity could lead to improved control of herbivorous pests. In a crossed design, we increased and lowered densities of decomposer prey, and manipulated vegetation complexity using artificial plants in order to examine the effect of structural complexity in isolation from effects of plant‐attracted additional prey. Isotomid Collembola exhibited lowest activity‐densities (AD) in plots treated with soil insecticide and had gradually increasing AD in untreated plots and plots receiving detrital subsidies. Carabid beetles and cursorial spiders did not respond to increased availability of isotomid prey, and they unexpectedly displayed higher AD in the structurally less‐complex plots. Aphid density mirrored the positive response of isotomids to detrital subsidies, suggesting that aphids benefited from reduced predation due to predators switching to abundant prey in the decomposer subsystem. The absence of a numerical response by surface‐active predators apparently strengthened this indirect effect of isotomids on aphids. Our results suggest that indirect predator‐mediated prey‐prey interactions can reduce beneficial effects of detrital subsidies on pest suppression. We further demonstrated that generalist predators may not per se benefit from structural complexity. Both results document the challenges associated with management practices that support generalist predators, as these measures may not necessarily improve herbivore suppression.     

In-phase and antiphase self-oscillations in a model of two electrically coupled pacemakers

The dynamic behavior of a model of two electrically coupled oscillatory neurons was studied while the external polarizing current was varied. It was found that the system with weak coupling can demonstrate one of five stable oscillatory modes: (1) in-phase oscillations with zero phase shift; (2) antiphase oscillations with halfperiod phase shift; (3) oscillations with any fixed phase shift depending on the value of the external polarizing current; (4) both in-phase and antiphase oscillations for the same current value, where the oscillation type depends on the initial conditions; (5) both in-phase and quasiperiodic oscillations for the same current value. All of these modes were robust, and they persisted despite small variations of the oscillator parameters. We assume that similar regimes, for example antiphase oscillations, can be detected in neurophysiological experiments. Possible applications to central pattern generator models are discussed.     

Changes in species richness and composition in European acidic grasslands over the past 70 years: the contribution of cumulative atmospheric nitrogen deposition

Our study investigates the negative impact of nitrogen (N) deposition on species richness in acidic grasslands, based on a temporal comparison of vegetation data spanning a period of almost 70 years. We compiled a large data base of plots assigned to the Violion caninae grassland type, composed of managed, but unfertilized semi-natural grasslands on nutrient-poor, acidic soils. In total 1114 plots, mainly from Great Britain, the Netherlands and Germany, were compiled, dating back to 1939. Environmental site information included geographical and soil (mean Ellenberg values) variables as well as estimates of cumulative N and sulphur (S) deposition since 1939. Statistical analyses were carried out separately for the data subsets from the three regions. In all regions, the vegetation differentiation was mainly related to soil acidity and nutrient availability, as well as to the year of sampling and the cumulative amounts of N and S deposition. Plot-species richness of vascular plants and bryophytes (analysed for Great Britain only) decreased with time and analyses suggest these are affected by various factors, notably soil pH, but also latitude and cumulative N deposition. The latter explained more of the variation in species number than the year of sampling and cumulative S deposition, which supports the interpretation that the decline in species richness is mainly caused by increasing N availability and less by altered management and soil acidification. For Great Britain and Germany, cumulative N deposition showed a strong negative relationship with several biodiversity measures, especially the proportion of dicots, whereas it was positively related to the proportion of grass species. In general, our results give temporal evidence for the negative effect of N deposition on species richness in semi-natural vegetation.