Phytoplankton responses to repeated pulse perturbations imposed on a trend of increasing eutrophication

While eutrophication remains one of the main pressures acting on freshwater ecosystems, the prevalence of anthropogenic and nature‐induced stochastic pulse perturbations is predicted to increase due to climate change. Despite all our knowledge on the effects of eutrophication and stochastic events operating in isolation, we know little about how eutrophication may affect the response and recovery of aquatic ecosystems to pulse perturbations. There are multiple ways in which eutrophication and pulse perturbations may interact to induce potentially synergic changes in the system, for instance, by increasing the amount of nutrients released after a pulse perturbation. Here, we performed a controlled press and pulse perturbation experiment using mesocosms filled with natural lake water to address how eutrophication modulates the phytoplankton response to sequential mortality pulse perturbations; and what is the combined effect of press and pulse perturbations on the resistance and resilience of the phytoplankton community. Our experiment showed that eutrophication increased the absolute scale of the chlorophyll‐a response to pulse perturbations but did not change the proportion of the response relative to its pre‐event condition (resistance). Moreover, the capacity of the community to recover from pulse perturbations was significantly affected by the cumulative effect of sequential pulse perturbations but not by eutrophication itself. By the end of the experiment, some mesocosms could not recover from pulse perturbations, irrespective of the trophic state induced by the press perturbation. While not resisting or recovering any less from pulse perturbations, phytoplankton communities from eutrophying systems showed chlorophyll‐a levels much higher than non‐eutrophying ones. This implies that the higher absolute response to stochastic pulse perturbations in a eutrophying system may increase the already significant risks for water quality (e.g., algal blooms in drinking water supplies), even if the relative scale of the response to pulse perturbations between eutrophying and non‐eutrophying systems remains the same.     

Decomposing multiple dimensions of stability in global change experiments

Ecological stability is the central framework to understand an ecosystem's ability to absorb or recover from environmental change. Recent modelling and conceptual work suggests that stability is a multidimensional construct comprising different response aspects. Using two freshwater mesocosm experiments as case studies, we show how the response to single perturbations can be decomposed in different stability aspects (resistance, resilience, recovery, temporal stability) for both ecosystem functions and community composition. We find that extended community recovery is tightly connected to a nearly complete recovery of the function (biomass production), whereas systems with incomplete recovery of the species composition ranged widely in their biomass compared to controls. Moreover, recovery was most complete when either resistance or resilience was high, the latter associated with low temporal stability around the recovery trend. In summary, no single aspect of stability was sufficient to reflect the overall stability of the system.     

Species richness facilitates ecosystem resilience in aquatic food webs

1. Many studies indicate that biodiversity in ecosystems affects stability, either by promoting temporal stability of ecosystem attributes or by enhancing ecosystem resistance and resilience to perturbation. The effects on temporal stability are reasonably well understood and documented but effects on resistance and resilience are not. 2. Here, we report results from an aquatic mesocosm experiment in which we manipulated the species richness and composition of aquatic food webs (macrophytes, macro‐herbivores and invertebrate predators), imposed a pulse disturbance (acidification), and monitored the resistance (initial response) and resilience (recovery) of ecosystem productivity and respiration. 3. We found that species‐rich macroinvertebrate communities had higher resilience of whole‐ecosystem respiration, but were not more resistant to perturbations. We also found that resilience and resistance were unaffected by species composition, despite the strong role composition is known to play in determining mean levels of function in these communities. 4. Biodiversity’s effects on resilience were probably mediated through complex pathways affecting phytoplankton and microbial communities (e.g. via changes in nutrient regeneration, grazing or compositional changes) rather than through simpler effects (e.g. insurance effects, enhanced facilitation) although these simpler mechanisms probably played minor roles in enhancing respiration resilience. 5. Current mechanisms for understanding biodiversity’s effects on ecosystem stability have been developed primarily in the context of single‐trophic level communities. These mechanisms may be overly simplistic for understanding the consequences of species richness on ecosystem stability in complex, multi‐trophic food webs where additional factors such as indirect effects and highly variable life‐history traits of species may also be important.     

Evolution of resistance to transgenic crops: interactions between insect movement and field distribution

The refuge strategy is designed to delay evolution of pest resistance to transgenic crops producing Bacillus thuringiensis Berliner (Bt) toxins. Movement of insects between Bt crops and refuges of non-Bt crops is essential for the refuge strategy because it increases chances that resistant adults mate with susceptible adults from refuges. Conclusions about optimal levels of movement for delaying resistance are not consistent among previous modeling studies. To clarify the effects of movement on resistance evolution, we analyzed simulations of a spatially explicit model based partly on the interaction of pink bollworm, Pectinophora gossypiella (Saunders), with Bt cotton. We examined resistance evolution as a function of insect movement under 12 sets of assumptions about the relative abundance of Bt cotton (50 and 75%), temporal distribution of Bt cotton and refuge fields (fixed, partial rotation, and full rotation), and spatial distribution of fields (random and uniform). The results show that interactions among the relative abundance and distribution of refuges and Bt cotton fields can alter the effects of movement on resistance evolution. The results also suggest that differences in conclusions among previous studies can be explained by differences in assumptions about the relative abundance and distribution of refuges and Bt crop fields. With fixed field locations and all Bt cotton fields adjacent to at least one refuge, resistance evolved slowest with low movement. However, low movement and fixed field locations favored rapid resistance evolution when some Bt crop fields were isolated from refuges. When refuges and Bt cotton fields were rotated to the opposite crop type each year, resistance evolved fastest with low movement. Nonrecessive inheritance of resistance caused rapid resistanceevolution regardless of movement rate. Confirming previous reports, results described here show that resistance can be delayed effectively by fixing field locations and distributing refuges uniformly to ensure that Bt crop fields are not isolated from refuges. However, rotating fields provided better insect control and reduced the need for insecticide sprays.     

土壤微生物群落抵抗力和恢复力研究进展

人类活动的不断加剧使得土壤生态系统承受着环境干扰压力。土壤微生物受到环境干扰的响应程度（抵抗力）及恢复至原来状态的能力（恢复力）决定着土壤生态系统的可持续性。梳理和总结了土壤微生物群落对环境干扰的抵抗力和恢复力方面的研究进展。首先,在介绍土壤微生物群落抵抗力和恢复力概念的基础上,阐述了通过评估微生物群落的结构和功能的变化来系统表征抵抗力和恢复力;随后,分析了最近十年（2012-2021年）有关文献,发现土壤微生物群落的结构和（或）功能在环境干扰后的恢复力总体较弱,但耕作、有机物料添加和轮作等农田管理措施下的响应趋势表现出一定的规律性;继而,从个体水平的休眠和胁迫忍耐、种群水平的生存策略、群落水平的多样性和相互作用以及生态系统水平的历史遗留效应等方面分析了土壤微生物群落抵抗力和恢复力的维持机制;最后,从功能性状、多功能性和植物-土壤微生物整体性对未来研究做出了展望,以期为构建土壤健康评价体系及预测环境干扰对土壤功能的影响提供科学依据。     

Drought refuges,spatial scale and recolonisation by invertebrates in non‐perennial streams

1. If resistance traits drive recolonisation after drought, then drought refuges should contribute strongly to assemblage composition within streams. If resilience traits drive recolonisation, macroinvertebrates emerging from refuges may disperse widely, colonising many streams. To determine whether the contribution of drought refuges to macroinvertebrate recolonisation in non‐perennial streams was mostly local (within stream) or broader scale (across streams), we measured the association between the composition of invertebrate assemblages in different types of in‐stream drought refuge and the assemblage composition of streams when flow resumed. 2. We sampled 16 streams of varying hydrological regime on the western side of the Victoria Range in the Grampians National Park, Victoria, Australia. Drought refuges (perennial pools, dry sediment, damp sediment, seeps, patches of leaf litter, beneath stones) were identified and sampled during autumn. Most taxa were found in perennial pools; few taxa were found aestivating beneath stones or having desiccation‐resistant stages in dry sediment. Perennial pools and perennially flowing reaches were the refuges that harboured the greatest diversity of macroinvertebrate taxa. 3. Streams were sampled again during spring. Assemblage composition of non‐perennial reaches in spring was unrelated to composition in nearby refuges in the previous autumn. In contrast, assemblage composition in perennial reaches during spring was strongly correlated with composition during autumn. Therefore, drought refuges did not directly influence assemblage composition locally within non‐perennial streams. Rather, both perennially flowing reaches and perennial pools acted as drought refuges across the broader landscape. Resilience traits are likely to drive recolonisation in these streams. 4. Monitoring of drought refuges in a particular stream will therefore not predict species composition when flow resumes. Drought refuges are likely to sustain biodiversity over larger spatial scales such as groups of streams or whole drainage networks. Consequently, stream networks will need to be managed as entities rather than as single waterways and the focus of drought refuge protection should be on perennial pools and reaches.     

Role of Spatial and Temporal Refuges in the Evolution of Pest Resistance to Toxic Crops

Toxic plants have been used for years in agriculture to control major crop pests. However, the continuous exposure of targeted pests to toxins dramatically increases the rate of resistance evolution (Gassman et al. in Annu Rev Entomol 54:147–163, 2009a; Tabashnik et al. Nat Biotechnol 26:199–202, 2008). To prevent or delay resistance, non toxic host plants can be used as refuges. Our study considers spatial and temporal refuges that are respectively implemented concurrently or alternatively a toxic crop. A conceptual model based on impulsive differential equations is proposed to describe the dynamics of the susceptible and resistant pest populations over time. The mathematical study enlightens threshold values of the proportion of the spatial refuge and key parameters that should help to understand evolution of pest resistance to toxic crop.     

Co‐adaptation impacts the robustness of predator–prey dynamics against perturbations

Global change threatens the maintenance of ecosystem functions that are shaped by the persistence and dynamics of populations. It has been shown that the persistence of species increases if they possess larger trait adaptability. Here, we investigate whether trait adaptability also affects the robustness of population dynamics of interacting species and thereby shapes the reliability of ecosystem functions that are driven by these dynamics. We model co‐adaptation in a predator–prey system as changes to predator offense and prey defense due to evolution or phenotypic plasticity. We investigate how trait adaptation affects the robustness of population dynamics against press perturbations to environmental parameters and against pulse perturbations targeting species abundances and their trait values. Robustness of population dynamics is characterized by resilience, elasticity, and resistance. In addition to employing established measures for resilience and elasticity against pulse perturbations (extinction probability and return time), we propose the warping distance as a new measure for resistance against press perturbations, which compares the shapes and amplitudes of pre‐ and post‐perturbation population dynamics. As expected, we find that the robustness of population dynamics depends on the speed of adaptation, but in nontrivial ways. Elasticity increases with speed of adaptation as the system returns more rapidly to the pre‐perturbation state. Resilience, in turn, is enhanced by intermediate speeds of adaptation, as here trait adaptation dampens biomass oscillations. The resistance of population dynamics strongly depends on the target of the press perturbation, preventing a simple relationship with the adaptation speed. In general, we find that low robustness often coincides with high amplitudes of population dynamics. Hence, amplitudes may indicate the robustness against perturbations also in other natural systems with similar dynamics. Our findings show that besides counteracting extinctions, trait adaptation indeed strongly affects the robustness of population dynamics against press and pulse perturbations.     

食饵庇护所对斑块环境下Leslie-Gower捕食系统的影响

建立了一个显式含有空间庇护所的两斑块Leslie-Gower捕食者-食饵系统。假设只有食饵种群在斑块间以常数迁移率迁移,且在每个斑块上食饵间的迁移比局部捕食者-食饵相互作用发生的时间尺度要快。利用两个时间尺度,可以构建用来描述所有斑块总的食饵和捕食者密度的综合系统。数学分析表明,在一定条件下,存在唯一的正平衡点,并且此平衡点全局稳定。进一步,捕食者的数量随着食饵庇护所数量增加而降低;在一定条件下,食饵的数量随着食饵庇护所数量增加先增加后降低,在足够强的庇护所强度下,两物种出现灭绝。对比以往研究,利用显式含有和隐含空间庇护所的数学模型所得结论不一致,这意味着在研究庇护所对捕食系统种群动态影响时,空间结构可能起着重要作用。     

Quantifying the impact of ecological memory on the dynamics of interacting communities

Ecological memory refers to the influence of past events on the response of an ecosystem to exogenous or endogenous changes. Memory has been widely recognized as a key contributor to the dynamics of ecosystems and other complex systems, yet quantitative community models often ignore memory and its implications.Recent modeling studies have shown how interactions between community members can lead to the emergence of resilience and multistability under environmental perturbations. We demonstrate how memory can be introduced in such models using the framework of fractional calculus. We study how the dynamics of a well-characterized interaction model is affected by gradual increases in ecological memory under varying initial conditions, perturbations, and stochasticity.Our results highlight the implications of memory on several key aspects of community dynamics. In general, memory introduces inertia into the dynamics. This favors species coexistence under perturbation, enhances system resistance to state shifts, mitigates hysteresis, and can affect system resilience both ways depending on the time scale considered. Memory also promotes long transient dynamics, such as long-standing oscillations and delayed regime shifts, and contributes to the emergence and persistence of alternative stable states. Our study highlights the fundamental role of memory in communities, and provides quantitative tools to introduce it in ecological models and analyse its impact under varying conditions.     

The role of refugia for fishes during drought: a review and synthesis

• 1 Drought is a natural disturbance of aquatic ecosystems and can be a major factor in structuring aquatic communities. For individuals, populations and communities to persist in disturbed environments, they must have refuge from disturbance or disturbance must be minimal. Refugia convey spatial and temporal resistance or resilience in the face of disturbance, but the role of refugia in aquatic systems remains poorly understood. 2. We review available literature on aquatic refugia for fishes in order to synthesise current knowledge and provide suggestions for needed research. Our objectives were to clarify definitions of disturbance and refugia in the context of drought in aquatic systems, review how refuge habitats influence fish community structure, and consider the potential impact of refugia on fish population and community dynamics during drought. 3. Droughts cause a decrease in surface area/volume and an increase in extremes of physical and chemical water quality parameters. These conditions are linked with biotic interactions that structure the community of fishes residing in low‐flow or dry season refugia by increasing mortality rates, decreasing birth rates and/or increasing migration rates. Many aquatic organisms seek refuge from disturbance and/or have adaptations (e.g. physiological tolerance) that provide refuge. 4. Drought in aquatic systems leads to shifts in refugia spacing and connectance at multiple spatial and temporal scales. Refuge size, disturbance intensity, and mobility of organisms is predicted to play a large role in population persistence. We expect that refuge habitats will experience net immigration during drying and net emigration after rewetting, with the opposite occurring in surrounding habitat patches. Population dynamics of fishes using refugia during drought are best modelled by modified source‐sink dynamics, but dynamics are likely to change with spatial scale.

     

Red imported fire ants (<Emphasis Type="Italic">Solenopsis invicta</Emphasis>) and seasonality influence community refuge use

Refuges are fundamental to animal ecology as refuge availability affects many levels of biological organization—from the behavior and physiology of individuals to the interspecific dynamics of a community. Although frequently studied in the context of predator–prey interactions, refuges may also mediate interspecific competition between native and invasive taxa given the role of refuges as a valuable resource. Because interspecific interactions (e.g., competition and predation) can be modulated by temporal and biotic (e.g., trophic level) factors, we used a manipulative approach to investigate community-wide refuge-use patterns in the context of two important ecological factors: invasive species and seasonality. We surveyed refuge (artificial cover object) use of ants and vertebrates in a forest community for 2 years, and we systematically suppressed an established invasive species (red imported fire ant, Solenopsis invicta) to examine its impact on community refuge use. Native Camponotus ants appeared to co-exist and share refuges with S. invicta, but we found evidence for a negative effect of S. invicta on vertebrate refuge use that was also influenced by season. Vertebrates were more abundant under refuges undergoing suppression of S. invicta, and they were less abundant under refuges during the fall (the season characterized by the highest occupancy of refuges by S. invicta). Thus, researchers must continue to examine the entire community and to incorporate the effects of season when assessing the impact of invasive species (e.g., at our site, a survey conducted only in the summer or only on native ants would have indicated a negligible effect of S. invicta on community refuge use).     

Disturbance and the role of refuges in mediterranean climate streams

Refuges protect plant and animal populations from disturbance. Knowledge of refuges from disturbance in mediterranean climate rivers (med-rivers) has increased the last decade. We review disturbance processes and their relationship to refuges in streams in mediterranean climate regions (med-regions). Med-river fauna show high endemicity and their populations are often exposed to disturbance; hence the critical importance of refuges during (both seasonal and supraseasonal) disturbances. Disturbance pressures are increasing in med-regions, in particular from climatic change, salinisation, sedimentation, water extraction, hydropower generation, supraseasonal drought, and wildfire. Med-rivers show annual cycles of constrained precipitation and predictable seasonal drying, causing the biota to depend on seasonal refuges, in particular, those that are spatially predictable. This creates a spatial and temporal mosaic of inundation that determines habitat extent and refuge function. Refuges of sufficient size and duration to maintain populations, such as perennially flowing reaches, sustain biodiversity and may harbour relict populations, particularly during increasing aridification, where little other suitable habitat remains in landscapes. Therefore, disturbances that threaten perennial flows potentially cascade disproportionately to reduce regional scale biodiversity in med-regions. Conservation approaches for med-river systems need to conserve both refuges and refuge connectivity, reduce the impact of anthropogenic disturbances and sustain predictable, seasonal flow patterns.     

The role of refuges in the persistence of Australian dryland mammals

Irruptive population dynamics are characteristic of a wide range of fauna in the world's arid (dryland) regions. Recent evidence indicates that regional persistence of irruptive species, particularly small mammals, during the extensive dry periods of unpredictable length that occur between resource pulses in drylands occurs as a result of the presence of refuge habitats or refuge patches into which populations contract during dry (bust) periods. These small dry‐period populations act as a source of animals when recolonisation of the surrounding habitat occurs during and after subsequent resource pulses (booms). The refuges used by irruptive dryland fauna differ in temporal and spatial scale from the refugia to which species contract in response to changing climate. Refuges of dryland fauna operate over timescales of months and years, whereas refugia operate on timescales of millennia over which evolutionary divergence may occur. Protection and management of refuge patches and refuge habitats should be a priority for the conservation of dryland‐dwelling fauna. This urgency is driven by recognition that disturbance to refuges can lead to the extinction of local populations and, if disturbance is widespread, entire species. Despite the apparent significance of dryland refuges for conservation management, these sites remain poorly understood ecologically. Here, we synthesise available information on the refuges of dryland‐dwelling fauna, using Australian mammals as a case study to provide focus, and document a research agenda for increasing this knowledge base. We develop a typology of refuges that recognises two main types of refuge: fixed and shifting. We outline a suite of models of fixed refuges on the basis of stability in occupancy between and within successive bust phases of population cycles. To illustrate the breadth of refuge types we provide case studies of refuge use in three species of dryland mammal: plains mouse (Pseudomys australis), central rock‐rat (Zyzomys pedunculatus), and spinifex hopping‐mouse (Notomys alexis). We suggest that future research should focus on understanding the species‐specific nature of refuge use and the spatial ecology of refuges with a focus on connectivity and potential metapopulation dynamics. Assessing refuge quality and understanding the threats to high‐quality refuge patches and habitat should also be a priority. To facilitate this understanding we develop a three‐step methodology for determining species‐specific refuge location and habitat attributes. This review is necessarily focussed on dryland mammals in continental Australia where most refuge‐based research has been undertaken. The applicability of the refuge concept and the importance of refuges for dryland fauna conservation elsewhere in the world should be investigated. We predict that refuge‐using mammals will be widespread particularly among dryland areas with unpredictable rainfall patterns.     

Choosing among alternative refuges: Distances and directions

Many prey flee to refuges to escape from approaching predators, but little is known about how they select one among many refuges available. The problem of choice among alternative refuges has not been modeled previously, but a recent model that predicts flight initiation distance (FID = predator–prey distance when escape starts) for a prey fleeing to a refuge provides a basis for predicting which refuge should be chosen. Because fleeing is costly, prey should choose to flee to the refuge permitting the shortest FID. The model predicts that the more distant of two refuges can be favored if it is not too far and if the prey's trajectory to the farther refuge is more away from the predator than the direction to the nearer refuge. The difference in predicted FID between the farther and nearer refuges increases curvilinearly as the interpath angle for the farther refuge increases. The difference in predicted FID between the farther and nearer refuges increases linearly as the distance to the farther refuge increases. An isocline describing where nearer and farther refuges are equally favored shows a negative curvilinear relationship between interpath angle and prey distance to the farther refuge. In the region below the isocline, the farther refuge is favored, whereas above the isocline the prey should flee to the nearer refuge.     

Associational refuges among corals mediate impacts of a crown-of-thorns starfish Acanthaster planci outbreak Indirect positive interactions in communities

Interactions among coral populations can moderate the impact of coral predator outbreaks, enhancing community resilience and recovery. This study used predator-exclusion cages and neighbour removals in a field experiment to test how indirect interactions between populations of three coral taxa, Acropora, Pocillopora, and Porites, influenced their survival during an outbreak of the crown-of-thorns starfish, Acanthaster planci, in Moorea, French Polynesia. High densities of corals enhanced survival by generating associational refuges: physical structures that impeded Acanthaster and protected corals, and by simple density-dependent prey dilution that reduced predation rates. Acanthaster showed feeding preferences, resulting in varying intensities of predation on corals, which (1) influenced the type and strength of the associational refuge among corals and (2) resulted in significant loss of the competitive dominants to the benefit of the competitive inferiors. The result was a set of indirect positive interactions (IPIs) that prevented Acanthaster from eradicating Acropora and may have enhanced Porites, a relatively weak competitor among corals. IPIs probably play a key role in many ecosystems, especially in coral reefs in which corals act as engineer species, to reduce impacts of perturbations and enhance community resilience. This study illustrates the importance of IPIs in community regulation with a new conceptual model.     

Resilience of terrestrial and aquatic fauna to historical and future wildfire regimes in western North America

Wildfires in many western North American forests are becoming more frequent, larger, and severe, with changed seasonal patterns. In response, coniferous forest ecosystems will transition toward dominance by fire‐adapted hardwoods, shrubs, meadows, and grasslands, which may benefit some faunal communities, but not others. We describe factors that limit and promote faunal resilience to shifting wildfire regimes for terrestrial and aquatic ecosystems. We highlight the potential value of interspersed nonforest patches to terrestrial wildlife. Similarly, we review watershed thresholds and factors that control the resilience of aquatic ecosystems to wildfire, mediated by thermal changes and chemical, debris, and sediment loadings. We present a 2‐dimensional life history framework to describe temporal and spatial life history traits that species use to resist wildfire effects or to recover after wildfire disturbance at a metapopulation scale. The role of fire refuge is explored for metapopulations of species. In aquatic systems, recovery of assemblages postfire may be faster for smaller fires where unburned tributary basins or instream structures provide refuge from debris and sediment flows. We envision that more‐frequent, lower‐severity fires will favor opportunistic species and that less‐frequent high‐severity fires will favor better competitors. Along the spatial dimension, we hypothesize that fire regimes that are predictable and generate burned patches in close proximity to refuge will favor species that move to refuges and later recolonize, whereas fire regimes that tend to generate less‐severely burned patches may favor species that shelter in place. Looking beyond the trees to forest fauna, we consider mitigation options to enhance resilience and buy time for species facing a no‐analog future.     

Indicators and mechanisms of stability and resilience to climatic and landscape changes in a remnant calcareous grassland

When considering the development of conservation strategies for threatened plant communities it is crucial to understand their resilience to environmental change, taking into account current decline and the occurrence of further habitat fragmentation and climatic changes. Many recent works describe resilience character and elements, but there is little focus on the metrics and indices that describe elements of stability and specific resistance or resilience over the community composition matrix. Communities with strong niche selection might be restricted to specific resistance strategies to cope with environmental changes. This would result in a community at greater risk from increasing fragmentation and climatic changes.In a 35 years survey of relic calcareous grasslands, we looked at measures to identify the resilience mechanisms for stability in the presence and abundance of species. We used techniques of partitioning of temporal beta diversity in nestedness and turnover components, analysis of functional strategy changes and dissimilarity analysis to detect changes in between⿿plot diversity and exchanges.Contrary to expectations, we observe strong resilience with different stabilizing mechanisms both at plot level and exchanges between plots. At the scale of our grassland complex, response diversity and environmental stochasticity allow for the maintenance of high biodiversity under natural perturbations and gradual human-induced environmental changes. This highlights the importance of dispersal, recruitment dynamics and microsite diversity.Community resilience is more than just the sum of species resilience strategies; adaptive management strategies need more emphasis on the variability of conditions, as this can enable or disrupt important community resilience mechanisms.     

Predicting evolution of insect resistance to transgenic crops in within-field refuge configurations,based on larval movement

The selection pressure imposed by the widespread use of transgenic technologies can lead to the evolution of insect resistance, and the availability of refuge areas that allow susceptible homozygous insects to survive is a key factor in delaying the evolution of resistance in agricultural landscapes. Different strategies to exploit refuge areas exist, but several insect-related ecological traits may directly affect the efficiency of refuges in slowing the development of resistance. Insect larval movement is one such trait that may affect the management of resistance, depending on the refuge strategy adopted. We developed a computational model to simulate how larval movement would affect the spatio-temporal dynamics of the evolution of resistance of insect pests to Bt crops, under different refuge configurations. In order to test the model, we used population data for Spodoptera frugiperda, one of the main target pests for control with Bt toxins. Simulations were run for spatial arrangements composed of three refuge configurations (seed mixture, blocks, and strips), with sizes ranging from 20% to 50% for two types of resistance (incomplete and complete) and three rates of larval movement (proportion of larvae moving per time step), equal to 0, 0.1 and 0.5. We demonstrated that with a seed mixture, in most cases the higher the rate of larval movement, the higher the proportion of resistant insects in the population in an area, regardless of the type of resistance tested. Strip configurations showed the opposite trend. In a block configuration, the number of resistant larvae was highest at an intermediate dispersal rate (0.1). We concluded that larval movement is an important variable affecting the evolution of resistance to Bt crops, but its effect depends on the type of resistance and the configuration and size of the refuge.     

Growth and refuge location in continuous,modular organisms: experimental and computer simulation studies

Summary Guilds of crustose bryozoans exhibited a range of growth outlines, varying from circular to elongate, and the stones on which they grew contained a range of refuge types, varying in size from a few mm² to a few hundred cm². These refuges, if encountered by the growing colonies of competitively subordinate species such as Electra pilosa L., enabled the latter to avoid competitive exclusion due to overgrowth mortality. Computer simulations suggested that, although modular organisms which are elongate are, in general, more likely to encounter spatial refuges than those with a different shape through vegetative growth, shape is less important than the size of the organisms themselves or the size and density of the spatial refuges. Moreover significant interactions between these 4 variables showed that they should be considered together and not in isolation from one another. Examples are given of variable growth forms in continuous, modular organisms from different taxa, including plants, and the importance of these predictive studies to refuge location by modular organisms is discussed.