Plant litter strengthens positive biodiversity–ecosystem functioning relationships over time

Plant biodiversity–productivity relationships become stronger over time in grasslands, forests, and agroecosystems. Plant shoot and root litter is important in mediating these positive relationships, yet the functional role of plant litter remains overlooked in long-term experiments. We propose that plant litter strengthens biodiversity–ecosystem functioning relationships over time in four ways by providing decomposing detritus that releases nitrogen (N) over time for uptake by existing and succeeding plants, enhancing overall soil fertility, changing soil community composition, and reducing the impact of residue-borne pathogens and pests. We bring new insights into how diversity–productivity relationships may change over time and suggest that the diversification of crop residue retention through increased residue diversity from plant mixtures will improve the sustainability of food production systems.     

Soil Functional Operating Range Linked to Microbial Biodiversity and Community Composition Using Denitrifiers as Model Guild

Soil microorganisms are key players in biogeochemical cycles. Yet, there is no consistent view on the significance of microbial biodiversity for soil ecosystem functioning. According to the insurance hypothesis, declines in ecosystem functioning due to reduced biodiversity are more likely to occur under fluctuating, extreme or rapidly changing environmental conditions. Here, we compare the functional operating range, a new concept defined as the complete range of environmental conditions under which soil microbial communities are able to maintain their functions, between four naturally assembled soil communities from a long-term fertilization experiment. A functional trait approach was adopted with denitrifiers involved in nitrogen cycling as our model soil community. Using short-term temperature and salt gradients, we show that the functional operating range was broader and process rates were higher when the soil community was phylogenetically more diverse. However, key bacterial genotypes played an important role for maintaining denitrification as an ecosystem functioning under certain conditions.     

Biodiversity of key-stone phylotypes determines crop production in a 4-decade fertilization experiment

Cropping systems have fertilized soils for decades with undetermined consequences for the productivity and functioning of terrestrial ecosystems. One of the critical unknowns is the role of soil biodiversity in controlling crop production after decades of fertilization. This knowledge gap limits our capacity to assess how changes in soil biodiversity could alter crop production and soil health in changing environments. Here, we used multitrophic ecological networks to investigate the importance of soil biodiversity, in particular, the biodiversity of key-stone taxa in controlling soil functioning and wheat production in a 35-year field fertilization experiment. We found strong and positive associations between soil functional genes, crop production and the biodiversity of key-stone phylotypes; soils supporting a larger number of key-stone nematode, bacteria and fungi phylotypes yielded the highest wheat production. These key-stone phylotypes were also positively associated with plant growth (phototrophic bacteria, nitrogen fixers) and multiple functional genes related to nutrient cycling. The retrieved information on the genomes clustered with key-stone bacterial phylotypes indicated that the key-stone taxa had higher gene copies of oxidoreductases (participating most biogeochemical cycles of ecosystems and linking to microbial energetics) and 71 essential functional genes associated with carbon, nitrogen, phosphorus, and sulfur cycling. Altogether, our work highlights the fundamental role of the biodiversity of key-stone phylotypes in maintaining soil functioning and crop production after several decades of fertilization, and provides a list of key-stone phylotypes linking to crop production and soil nutrient cycling, which could give science-based guidance for sustainable food production.Subject terms: Microbial ecology, Microbial ecology, Biogeochemistry     

Microbiological regulation of the biogeochemical nitrogen cycle

Most nitrogen transformations in soil are carried out by micro-organisms. An understanding of the microbiological processes is thus necessary in order for us to devise management practices in agricultural ecosystems, which will optimize plant root uptake of nitrogen and minimize nitrogen losses from the systems. Some aspects of the individual microbiological processes in the nitrogen cycle are discussed and their importance for an efficient management of agroecosystems. In soil various groups of organisms compete for available inorganic nitrogen and quantitative data are needed on the uptake kinetics for these various groups in order to be able to assess their competitive ability under different conditions. The influence of abiotic factors such as oxygen concentration, inorganic nitrogen concentration and pH is discussed in relation to the different processes. The importance of acetylene as a tool in nitrogen cycling studies is discussed briefly.     

冻融对温带土壤可溶性氮库、氮转化过程及细菌群落多样性的影响

土壤冻融会影响土壤氮的有效性。氮的转化与土壤微生物密不可分,而土壤冻融对温带土壤细菌群落的影响并不十分清楚。假设: 冻融影响细菌群落结构多样性及其组成,从而改变土壤可溶氮含量和氮转化过程。为了验证这一假设,本研究设计了不同冻融循环次数(分别为6次和15次循环)的培养试验,并以2 ℃恒温培养作为对照。结果表明: 随着冻融循环次数的增加,可溶性全氮、可溶性无机氮、微生物生物量氮和净氮矿化率均显著降低。冻融循环次数对细菌α多样性(包括Chao1和Shannon指数)无显著影响,恒温培养的培养周期数与细菌的α多样性呈显著正相关。冻融处理显著影响细菌群落功能和组成,但冻融循环次数对细菌群落结构的影响较小。偏冗余分析表明,冻融处理下细菌群落结构和功能多样性与土壤可溶氮含量和氮转化过程密切相关。     

氮沉降对杉木人工林土壤可溶性有机质数量和结构的影响

氮沉降作为现在乃至未来气候变化的趋势之一,其可能深刻影响土壤可溶性有机质的数量和结构。选取我国中亚热带杉木人工林不同深度土壤（0-10 cm和10-20 cm）进行氮沉降试验,利用光谱技术研究氮沉降对土壤可溶性有机质数量和结构的影响。试验设对照（CT,0 kg hm^-2 a^-1）、高氮（HN：80 kg hm^-2 a^-1）、低氮（LN：40 kg hm^-2 a^-1）3种处理。结果表明：（1）在0-10 cm和10-20 cm土层,HN、LN处理的土壤可溶性有机碳和可溶性有机氮含量显著高于CT。（2）在0-10 cm和10-20 cm土层,1月时HN、LN处理的芳香性指数和腐殖化程度都显著高于CT,而4月时HN、LN处理的芳香性指数和腐殖化程度都显著低于CT。除了氮含量的直接影响外,RDA （冗余分析）表明,两土层中土壤含水量、pH和土壤有机碳是驱动氮沉降对土壤DOM数量和结构的重要环境因子。因此,氮沉降对土壤DOM的影响是复杂的,未来尤其应该注重氮沉降对生态系统影响的季节模式。     

Agroecosystem resilience is modified by management system via plant–soil feedbacks

Designing resilient cropping systems is essential to sustain agricultural production in the face of changing environmental and social pressures. However, the extent to which changes in farm management systems could alter resistance and resilience is largely unknown, especially in response to climate change. Plant and soil microbial community interactions are a vital component of functioning and resilient agroecosystems. The aim of our study was to use winter wheat (Triticum aestivum L.) and pea (Pisum sativum L.) plant–soil feedbacks (i.e. plant species-specific effects on soil biota and their impacts on subsequent plant growth) as a metric of system resilience and resistance to climate variability in three different farming management systems: 1) a chemical no-till system, 2) an USDA-certified organic system reliant on tillage and 3) an USDA-certified organic system that included sheep grazing with the overall goal of minimizing tillage intensity. Climate conditions soil experienced were ambient, warmer, and warmer and drier and were manipulated in the field using open-top chamber and rain-out shelters. Plant–soil feedbacks were negative for wheat and positive for pea but varied among farming management systems but were less sensitive to climate conditions. Plant–soil feedbacks were lower in magnitude in the tilled organic system indicating more resistance to the accumulation of pathogenic soil microbiota resulting from repeated cropping of wheat. However, recovery was lower when the crop was pea in the tilled organic indicating slower recovery and less resilience. Results indicate that while increases in crop diversity may promote more resilient agroecosystems, farming management will affect agroecosystem resilience.     

Crop rotational diversity enhances belowground communities and functions in an agroecosystem

Biodiversity loss, an important consequence of agricultural intensification, can lead to reductions in agroecosystem functions and services. Increasing crop diversity through rotation may alleviate these negative consequences by restoring positive aboveground–belowground interactions. Positive impacts of aboveground biodiversity on belowground communities and processes have primarily been observed in natural systems. Here, we test for the effects of increased diversity in an agroecosystem, where plant diversity is increased over time through crop rotation. As crop diversity increased from one to five species, distinct soil microbial communities were related to increases in soil aggregation, organic carbon, total nitrogen, microbial activity and decreases in the carbon‐to‐nitrogen acquiring enzyme activity ratio. This study indicates positive biodiversity–function relationships in agroecosystems, driven by interactions between rotational and microbial diversity. By increasing the quantity, quality and chemical diversity of residues, high diversity rotations can sustain soil biological communities, with positive effects on soil organic matter and soil fertility.     

Correction to: The biogeochemical consequences of litter transformation by insect herbivory in the Subarctic: a microcosm simulation experiment

Northern forest ecosystems are projected to experience warmer growing seasons and increased soil freeze–thaw cycles in winter over the next century. Past studies show that warmer soils in the growing season enhance nitrogen uptake by plants, while soil freezing in winter reduces plant uptake and ecosystem retention of nitrogen, yet the combined effects of these changes on plant root capacity to take up nitrogen are unknown. We conducted a 2-year (2014–2015) experiment at Hubbard Brook Experimental Forest in New Hampshire, USA to characterize the response of root damage, nitrogen uptake capacity, and soil solution nitrogen to growing season warming combined with soil freeze–thaw cycles in winter. Winter freeze–thaw cycles damaged roots, reduced nitrogen uptake capacity by 42%, and increased soil solution ammonium in the early growing season (May–June). During the peak growing season (July), root nitrogen uptake capacity was reduced 40% by warming alone and 49% by warming combined with freeze–thaw cycles. These results indicate the projected combination of colder soils in winter and warmer soils in the snow-free season will alter root function by reducing root nitrogen uptake capacity and lead to transient increases of nitrogen in soil solution during the early growing season, with the potential to alter root competition for soil nitrogen and seasonal patterns of soil nitrogen availability. We conclude that considering interactive effects of changes in climate during winter and the snow-free season is essential for accurate determination of the response of nitrogen cycling in the northern hardwood forest to climate change.     

Quantification of Key Genes Steering the Microbial Nitrogen Cycle in the Rhizosphere of Sorghum Cultivars in Tropical Agroecosystems

The effect of agricultural management practices on geochemical cycles in moderate ecosystems is by far better understood than in semiarid regions, where fertilizer availability and climatic conditions are less favorable. We studied the impact of different fertilizer regimens in an agricultural long-term observatory in Burkina Faso at three different plant development stages (early leaf development, flowering, and senescence) of sorghum cultivars. Using real-time PCR, we investigated functional microbial communities involved in key processes of the nitrogen cycle (nitrogen fixation, ammonia oxidation, and denitrification) in the rhizosphere. The results indicate that fertilizer treatments and plant development stages combined with environmental factors affected the abundance of the targeted functional genes in the rhizosphere. While nitrogen-fixing populations dominated the investigated communities when organic fertilizers (manure and straw) were applied, their numbers were comparatively reduced in urea-treated plots. In contrast, ammonia-oxidizing bacteria (AOB) increased not only in absolute numbers but also in relation to the other bacterial groups investigated in the urea-amended plots. Ammonia-oxidizing archaea exhibited higher numbers compared to AOB independent of fertilizer application. Similarly, denitrifiers were also more abundant in the urea-treated plots. Our data imply as well that, more than in moderate regions, water availability might shape microbial communities in the rhizosphere, since low gene abundance data were obtained for all tested genes at the flowering stage, when water availability was very limited.Land degradation is one of the most serious threats to food production on the African continent. Soil erosion, nutrient depletion, low organic matter content, and unfavorable pH values are some of the reasons for a deficient soil fertility (30), mainly in Central African countries. Combined with high variability and irregular distribution of rainfall, these factors contribute to negative nutrient balances. For example, 4.4 million tons of nitrogen (N) are lost per year in African soils, but only 0.8 million tons are reapplied by fertilization (12, 34). Since nitrogen is a key nutrient determining the productivity of agroecosystems (7, 11, 43), it is of central importance to optimize the nitrogen balance in these countries, mainly by steering the genetic resources of soil microbes in a way that losses of applied nitrogen are minimized and biological nitrogen fixation is increased. The aim should be to obtain a highly efficient nitrogen turnover, with leaching of nitrate and losses of gaseous products such as nitrous oxide (N₂O) or dinitrogen (N₂) as low as possible.Despite the importance of this issue, not much data are available on microbial community structure and function related to the nitrogen cycle in agroecosystems of Central Africa, and scenarios from moderate climatic regions cannot simply be transferred to tropical agroecosystems. Furthermore, the few studies published thus far only investigated effects of agricultural management on a single process of the nitrogen turnover (18, 20, 32) and ignore the fact that nitrogen turnover is a network of closely interlinked processes.Therefore, we sought to investigate the effects of different fertilizer regimens on multiple transformation processes within the nitrogen cycle in agroecosystems from semiarid areas in Central Africa. We investigated nitrogen dynamics on a full-cycle approach, including the most important steps in well-aerated agricultural soils (nitrification, denitrification, and nitrogen fixation). We hypothesized that each fertilizer regimen results in typical abundance pattern of the functional populations. While in moderate agroecosystems the plant (respectively, the plant development stage and the plant performance) plays an important role in shaping microbial community structure and function in the rhizosphere, we further postulated that nitrogen turnover and the corresponding populations are also influenced by the availability of water in semiarid soils. To test these hypotheses on a molecular basis, we quantified bacterial genes encoding the nitrogenase reductase (nifH), ammonia monooxygenase (amoA), and nitrite reductase (nirK and nirS), as well as archaeal amoA genes, by real-time PCR and linked the data to soil properties and grain yield.     

干湿交替条件下土壤氮素转化及其影响研究进展

对干湿交替条件下土壤氮素的矿化、损失以及对土壤团聚体结构和土壤胀缩性、土壤微生物活性和群落等的影响进行了综合评述。研究表明,干湿交替能够对土壤氮素的矿化和损失产生影响,并且干湿交替条件下土壤中氮素的转化与土壤团聚体结构、土壤胀缩性以及土壤微生物活性和群落密切相关。此外,针对目前研究中的不足,提出了今后的研究方向,认为开展干湿交替条件下土壤氮素转化与物理性质和微生物的关系,尤其是肥料氮素的微生物转化过程,为进一步开展干湿交替条件下氮素在土壤中的转化过程,明确自然条件下土壤氮素转化机制的相关研究提供参考。     

Effects of species and soil‐nitrogen availability on root system architecture traits – study on a set of weed and crop species

Better managing crop : weed competition in cropping systems while reducing both nitrogen and herbicide inputs is a real challenge that requires a better understanding of crop and weed root architecture in relation to soil‐nitrogen availability. An original approach was used which considered the parameters of a simulation model of root architecture as traits to analyse (a) the interspecific diversity of root system architecture, and (b) its response to soil‐nitrogen availability. Two greenhouse experiments were conducted using three crop and nine weed species grown at two contrasted concentrations of soil‐nitrogen availability. Plant traits were measured to characterise both overall plant growth and root architecture, with a focus on primary root emergence, root elongation and branching. The studied root traits varied among species (from a twofold to a fourfold factor, depending on the trait), validating their use as indicators to analyse the interspecific variability of root architecture. The largest interspecies differences were for two traits: ‘maximal apical root diameter’ and ‘interbranch distance’ (distance between two successive laterals on the same root). Conversely, most of the studied root traits varied little with soil‐nitrogen availability (from no variation to a 1.1‐fold factor, depending on the trait) even though soil‐nitrogen availability varied with a 17‐fold factor and impacted the overall shoot and root biomass. So, the root traits used in this article are stable whatever soil‐nitrogen availability. As they reflect processes underlying root system architecture, this low effect of nitrogen suggests that the rules governing root architecture are little affected by plant nitrogen status and soil‐nitrogen availability. We propose that the determinants of differences in root system architecture between soils with contrasted nitrogen availability mainly originate from differences in the amount of carbon allocated to and within the root system. Characterising each plant species by a combination of root traits gave insights regarding the potential species competitive ability for soil resources in agroecosystems.     

Do biotic interactions modulate ecosystem functioning along stress gradients? Insights from semi-arid plant and biological soil crust communities

Climate change will exacerbate the degree of abiotic stress experienced by semi-arid ecosystems. While abiotic stress profoundly affects biotic interactions, their potential role as modulators of ecosystem responses to climate change is largely unknown. Using plants and biological soil crusts, we tested the relative importance of facilitative–competitive interactions and other community attributes (cover, species richness and species evenness) as drivers of ecosystem functioning along stress gradients in semi-arid Mediterranean ecosystems. Biotic interactions shifted from facilitation to competition along stress gradients driven by water availability and temperature. These changes were, however, dependent on the spatial scale and the community considered. We found little evidence to suggest that biotic interactions are a major direct influence upon indicators of ecosystem functioning (soil respiration, organic carbon, water-holding capacity, compaction and the activity of enzymes related to the carbon, nitrogen and phosphorus cycles) along stress gradients. However, attributes such as cover and species richness showed a direct effect on ecosystem functioning. Our results do not agree with predictions emphasizing that the importance of plant–plant interactions will be increased under climate change in dry environments, and indicate that reductions in the cover of plant and biological soil crust communities will negatively impact ecosystems under future climatic conditions.     

冻融作用对陆地生态系统氮循环关键过程的影响效应及其机制

冻融作用是中、高纬度及高海拔地区土壤普遍存在的一种自然现象,是非生长季陆地生态系统氮循环的重要影响因素.冻融作用主要通过改变土壤的理化性质及生物学性状来影响氮素在土壤中的迁移与转化.目前,冻融作用对陆地生态系统氮循环各个过程影响的研究结果不尽一致,理论机制尚不明晰,研究方法也需进一步地探索与创新,因此有必要对现有成果进行梳理和分析,以更好地把握冻融作用下的氮循环过程.本文结合国内外已有研究成果,论述了冻融作用对陆地生态系统氮循环关键过程（氮矿化、固持、硝化与反硝化过程、氮淋溶及气态损失）的影响效应及其主要机制,对目前研究中存在的不足进行了剖析,并对未来研究中迫切需要关注的重点研究方向进行了探讨与展望.     

Agronomic and environmental aspects of diazotrophic bacteria in rice fields

This article provides an overview of the free-living and plant-associated nitrogen (N)-fixing bacterial communities in wet rice fields, with a focus on describing the elements affecting community assemblages in this waterlogged soil–plant system. Nitrogen is a crucial nutrient for rice yield and growth. Characteristics of the rice paddy ecosystem promote N-fertilizer losses, resulting in negative impacts on the environment. Public concerns on sustainable rice crop production and food security have accentuated interest in exploring biological supplementary nitrogen sources. Biological N-fixation is a significant source of the nitrogen in agroecosystems. The nitrogen requirement of rice crops can be partly remedied by managing and promoting the activities of N-fixing microorganisms. These changes are leading towards a cleaner approach that maintains sustainability while simultaneously improving crop production targets. The use of N-fixing microorganisms as biofertilizers and the factors driving the success of this technology in wet rice paddies are also discussed.     

不同土壤水分条件下香樟凋落叶覆盖对土壤碳氮循环的影响

凋落物作为森林生态系统碳库的重要组成部分对森林土壤碳、氮循环具有重要作用.为探讨香樟凋落叶对土壤碳、氮循环的影响,室内模拟研究了10%、20%和30% 3种土壤含水量条件下香樟凋落叶覆盖森林土壤中碳、氮元素的变化.结果表明: 3种含水量条件下香樟凋落叶覆盖均显著增加了土壤CO₂排放速率和土壤溶解性有机碳(WSOC)含量,但显著降低了土壤中硝态氮含量,表明香樟凋落叶覆盖能够增强土壤呼吸强度和碳矿化,抑制土壤硝化作用;香樟凋落叶覆盖能够显著增加10%含水量土壤中铵态氮含量,但降低了20%和30%含水量土壤铵态氮含量,表明香樟凋落叶覆盖对土壤铵态氮含量的影响与土壤含水量有关.香樟凋落叶中部分单萜烯浓度在不同土壤含水量条件下分别与土壤CO₂排放速率和铵态氮含量呈显著正相关,而与土壤WSOC和硝态氮含量呈显著负相关,说明香樟凋落叶覆盖对土壤碳、氮循环的影响可能与凋落叶中的单萜烯有关.     

Facilitative Root Interactions in Intercrops

Facilitation takes place when plants ameliorate the environment of their neighbours, and increase their growth and survival. Facilitation occurs in natural ecosystems as well as in agroecosystems. We discuss examples of facilitative root interactions in intercropped agroecosystems; including nitrogen transfer between legumes and non-leguminous plants, exploitation of the soil via mycorrhizal fungi and soil-plant processes which alter the mobilisation of plant growth resources such as through exudation of amino acids, extra-cellular enzymes, acidification, competition-induced modification of root architecture, exudation of growth stimulating substances, and biofumigation. Facilitative root interactions are most likely to be of importance in nutrient poor soils and in low-input agroecosystems due to critical interspecific competition for plant growth factors. However, studies from more intensified cropping systems using chemical and mechanical inputs also show that facilitative interactions definitely can be of significance. It is concluded that a better understanding of the mechanisms behind facilitative interactions may allow us to benefit more from these phenomena in agriculture and environmental management.     

Intraspecific litter diversity and nitrogen deposition affect nutrient dynamics and soil respiration

Anthropogenic forces are concurrently reducing biodiversity and altering terrestrial nutrient cycles. As natural populations decline, genetic diversity within single species also declines. The consequences of intraspecific genetic loss for ecosystem functions are poorly understood, and interactions among intraspecific diversity, nitrogen deposition, and nutrient cycling are unknown. We present results from an experiment that simulated both a decline in biodiversity and an increase in nitrogen deposition. In soil microcosms, we tested effects of variation in intraspecific litter diversity and nitrogen deposition on soil respiration and nitrogen leaching. Increases in intraspecific litter diversity increased soil respiration overall, with the greatest increases in respiration occurring under high nitrogen deposition. Nitrogen deposition increased the amount of inorganic nitrogen leached, while the amount of dissolved organic nitrogen leached was correlated with initial litter chemistry (lignin concentration) and remained independent of litter diversity and nitrogen deposition treatments. Our results demonstrate the potential for losses in genetic diversity to interact with other global environmental changes to influence terrestrial nutrient cycles.     

Physiological shifts in the microbial community drive changes in enzyme activity in a perennial agroecosystem

Perennial agroecosystems have the potential to promote plant–microbial linkages by increasing the quantity of root carbon entering the soil. However, an understanding of how perennial cropping systems affect microbial communities remains incomplete. The objective of this study was to determine the potential for a fertilized perennial bioenergy cropping system to impact microbial growth and enzyme activity. Three times throughout the growing season we examined the activity of four enzymes involved in decomposition (ß-glucosidase, ß-xylosidase, cellobiohydrolase, and N-acetyl glucosaminidase) in replicated plots of an annual (corn) and perennial-based (switchgrass) cropping system. We also took simultaneous measurements of microbial biomass and potential rates of microbial respiration and net N mineralization. Microbial biomass was unaffected by cropping system. Mid-summer, however, we observed increases in enzyme activity and potential microbial respiration in the perennial system that were independent of microbial biomass, likely in response to labile carbon inputs. Further, we observed lower net N mineralization, higher microbial biomass nitrogen and higher activity of nitrogen liberating enzymes, which are indicative of a community with high nitrogen demands. Overall, our research demonstrates that perennial agroecosystems can affect the physiological capacity of the microbial community, yielding communities with greater nitrogen retention and greater rates of decomposition as a result of allocation of resources towards enzyme production and nitrogen mining. These results can inform biogeochemical models with respect to the importance of temporally dynamic changes in carbon and nitrogen availability and microbial carbon use efficiency as drivers of enzyme production.     

Temporal dynamics of natural enemy–pest interactions in a changing environment

Agroecosystems contain complex networks of interacting organisms and these interaction webs are structured by the relative timing of key biological and ecological events. Recent intensification of land management and global changes in climate threaten to desynchronize the temporal structure of interaction webs and disrupt the provisioning of ecosystem services, such as biological control by natural enemies. It is therefore critical to recognize the central role of temporal dynamics in driving predator–prey interactions in agroecosystems. Specifically, ecological dynamics in crop fields routinely behave as periodic oscillations, or cycles. Familiar examples include phenological cycles, diel activity rhythms, and crop-management cycles. The relative timing and the degree of overlap among ecological cycles determine the nature and magnitude of the ecological interactions among organisms, and ultimately determine whether ecosystem services, such as biological control, can be provided. Additionally, the ecological dynamics in many cropping systems are characterized by a pattern of frequent disturbances due to management actions such as harvest, sowing and pesticide applications. These disturbance cycles cause agroecosystems to be dominated by dispersal and repopulation dynamics. However, they also serve as selective filters that regulate which animals can persist in agroecosystems over larger temporal scales. Here, we review key concepts and examples from the literature on temporal dynamics in ecological systems, and provide a framework to guide biological control strategies for sustainable pest management in a changing world.