The complementarity between ecological infrastructure types benefits natural enemies and pollinators in a Mediterranean vineyard agroecosystem

Under the current scenario of biodiversity loss across agroecosystems, the preservation of ecological infrastructures is crucial to maintain ecosystem functions and deliver ecosystem services such as pollination or biological control of pests, which largely determine agricultural productivity and food security. The role that ecological infrastructures such as hedges or strips have in enhancing functionally important groups, including predators, parasitoids and pollinators has been reasonably well investigated in temperate areas. However, it is much less documented in agroecosystems from the Mediterranean basin, even though this area has been identified as a biodiversity hotspot for conservation priorities. In a study developed during 2014 in a traditional vineyard agroecosystem in Northern Spain using a combination of different sampling methods, we analysed beneficial arthropod assemblages across four types of linear ecological infrastructures: woodland hedges, rosaceous hedges, grass strips and flower strips. We investigated the value of these elements as a source of predators, parasitoids and pollinators. We show that ecological infrastructures across this Mediterranean vineyard agroecosystem support functionally important groups and that the abundance, richness and distribution of families of each functional group is determined by features characterising the different types of those infrastructures. Ecological infrastructures with a suitable level of structural diversity constitute an important reservoir of predators. On the other hand, parasitoids and pollinators were limited by the availability of hosts, pollen and nectar resources. Flower strips, less complex infrastructures that offer a high amount of floral resources, benefited communities of both parasitoids and pollinators. Because of the variability in habitat preference found among the different functionally important taxa, we highlight the importance of preserving different types of ecological infrastructures across vineyard agroecosystems to maintain well‐structured assemblages of beneficial arthropods and maximise underlying services.     

Biodiversity as both a cause and consequence of resource availability: a study of reciprocal causality in a predator-prey system

1. One of the oldest questions in ecology is how species diversity in any given trophic level is related to the availability of essential resources that limit biomass (e.g. water, nutrients, light or prey). Researchers have tried to understand this relationship by focusing either on how diversity is influenced by the availability of resources, or alternatively, how resource abundance is influenced by species diversity. These contrasting perspectives have led to a seeming paradox '... is species diversity the cause or the consequence of resources that limit community biomass?' 2. Here we present results of an experiment that show it is possible for species diversity and resource density to exhibit reciprocal causal relationships in the same ecological system. Using a guild of ladybeetle predators and their aphid prey, we manipulated the number of predator species in field enclosures to examine how predator diversity impacts prey population size. At the same time, we manipulated the abundance of aphid prey in discrete habitat patches within each enclosure to determine how smaller-scale spatial variation in resource abundance affects the number of co-occurring predator species. 3. We found that the number of ladybeetle species added to enclosures had a significant impact on aphid population dynamics because interference competition among the predators reduced per capita rates of predation and, in turn, the overall efficiency of the predator guild. At the same time, spatial variation in aphid abundance among smaller habitat patches generated variation in the observed richness of ladybeetles because more species occurred in patches where predators aggregated in response to high aphid density. 4. The results of our experiment demonstrate that it is possible for species diversity to simultaneously be a cause and a consequence of resource density in the same ecological system, and they shed light on how this might occur for groups of mobile consumers that exhibit rapid responses to spatial and temporal variation in their prey.     

Anthropogenic impacts on tropical forest biodiversity: a network structure and ecosystem functioning perspective

Huge areas of diverse tropical forest are lost or degraded every year with dramatic consequences for biodiversity. Deforestation and fragmentation, over-exploitation, invasive species and climate change are the main drivers of tropical forest biodiversity loss. Most studies investigating these threats have focused on changes in species richness or species diversity. However, if we are to understand the absolute and long-term effects of anthropogenic impacts on tropical forests, we should also consider the interactions between species, how those species are organized in networks, and the function that those species perform. I discuss our current knowledge of network structure and ecosystem functioning, highlighting empirical examples of their response to anthropogenic impacts. I consider the future prospects for tropical forest biodiversity, focusing on biodiversity and ecosystem functioning in secondary forest. Finally, I propose directions for future research to help us better understand the effects of anthropogenic impacts on tropical forest biodiversity.     

Timing and magnitude of drought impacts on carbon uptake across a grassland biome

Although drought is known to negatively impact grassland functioning, the timing and magnitude of these impacts within a growing season remain unresolved. Previous small-scale assessments indicate grasslands may only respond to drought during narrow periods within a year; however, large-scale assessments are now needed to uncover the general patterns and determinants of this timing. We combined remote sensing datasets of gross primary productivity and weather to assess the timing and magnitude of grassland responses to drought at 5 km² temporal resolution across two expansive ecoregions of the western US Great Plains biome: the C₄-dominated shortgrass steppe and the C₃-dominated northern mixed prairies. Across over 700,000 pixel-year combinations covering more than 600,000 km², we studied how the driest years between 2003–2020 altered the daily and bi-weekly dynamics of grassland carbon (C) uptake. Reductions to C uptake intensified into the early summer during drought and peaked in mid- and late June in both ecoregions. Stimulation of spring C uptake during drought was small and insufficient to compensate for losses during summer. Thus, total grassland C uptake was consistently reduced by drought across both ecoregions; however, reductions were twice as large across the more southern and warmer shortgrass steppe. Across the biome, increased summer vapor pressure deficit (VPD) was strongly linked to peak reductions in vegetation greenness during drought. Rising VPD will likely exacerbate reductions in C uptake during drought across the western US Great Plains, with these reductions greatest during the warmest months and in the warmest locations. High spatiotemporal resolution analyses of grassland response to drought over large areas provide both generalizable insights and new opportunities for basic and applied ecosystem science in these water-limited ecoregions amid climate change.     

Impact of avian and arthropod predation on lepidopteran caterpillar densities and plant productivity in an ephemeral agroecosystem

Abstract.  1. Most studies evaluating the combined impact of spiders and other predators on herbivore densities in agroecosystems have focused primarily on their trophic connections with invertebrate predators (e.g. carabids, chrysopids); however linkages among spiders and vertebrate predators may also help structure the population dynamics of insect herbivores. A field experiment was conducted to examine the impact of avian and spider predation on lepidopteran caterpillar densities and plant productivity within a Brassica agroecosystem.
2. Arthropod abundance, leaf-chewing damage, and final plant productivity associated with broccoli, Brassica oleracea L. (var. italica ), were recorded for four treatments: (1) bird present but spiders removed; (2) both birds and spiders present; (3) birds excluded, spiders present; and (4) birds and spiders both excluded.
3. Densities of Artogeia rapae L. (Lepidoptera: Pieridae) and Trichoplusia ni Hübner (Lepidoptera: Noctuidae) large caterpillars and post feeding stages were reduced significantly by bird predation. The abundance of large caterpillars was also reduced on spider-inhabited plants during early plant growth; however the assemblage of birds and spiders did not suppress caterpillar densities more significantly than either predator alone.
4. Plants protected by birds, spiders, and birds plus spiders sustained less folivory attributable to leaf chewing caterpillars than check plants. Plant productivity was also greater for predator-protected plants than check plants.
5. Although spiders and parasitoids were responsible for some of the mortality inflicted upon lepidopteran caterpillars, it was concluded that in this study system, birds are the most important natural enemies of folivores.     

Factors favouring the development and maintenance of outbreaks in an invasive leaf miner Cameraria ohridella (Lepidoptera: Gracillariidae): a life table study

1 Life tables were constructed for seven generations of the invasive horse‐chestnut leaf miner Cameraria ohridella in Switzerland, to assess the factors allowing the moth to build and maintain outbreak densities and to identify ‘gaps’ among their mortality factors that could be targeted by new control methods. The fecundity of females was calculated and the mortality factors affecting all developmental stages determined. 2 Females contained 106.6 eggs. In captivity, the mean number of eggs laid per female varied between 34 and 82 eggs, with a maximum of 184 eggs. Egg mortality was 18–75% and was mostly due to unknown factors. 3 The main mortality factors affecting larvae and pupae in the leaves were predation by birds and arthropods in all generation and leaf senescence in the autumn generation. Parasitoids killed less than 5% of the population. 4 Nearly 90% of overwintering pupae died in dead leaves, the majority of them being killed by earthworms and other leaf decomposers. As a result, the overwintering generations were the only ones showing a net rate of increase less than one. Non‐overwintering generations had net rates of increase between four and 11, allowing populations to build up from spring to autumn. 5 The persistence of high outbreak densities in Europe is explained by a combination of at least three factors: (i) high fecundity; (ii) multivoltinism; (iii) the unusually low impact of traditional natural enemies of leaf miners and particularly parasitoids. 6 Yearly variations in population densities could be at least partly explained by differences in larval mortality due to leaf senescence and intraspecific competition in the last generation in autumn.     

Host-range evolution in Aphidius parasitoids: fidelity, virulence and fitness trade-offs on an ancestral host

The diversity of parasitic insects remains one of the most conspicuous patterns on the planet. The principal factor thought to contribute to differentiation of populations and ultimately speciation is the intimate relationship parasites share with hosts and the potential for disruptive selection associated with using different host species. Traits that generate this diversity have been an intensely debated topic of central importance to the evolution of specialization and maintenance of ecological diversity. A fundamental hypothesis surrounding the evolution of specialization is that no single genotype is uniformly superior in all environments. This "trade-off" hypothesis suggests that negative fitness correlations can lead to specialization on different hosts as alternative stable strategies. In this study we demonstrate a trade-off in the ability of the parasitoid, Aphidius ervi, to maintain a high level of fitness on an ancestral and novel host, which suggests a genetic basis for host utilization that may limit host-range expansion in parasitoids. Furthermore, behavioral evidence suggests mechanisms that could promote specialization through induced host fidelity. Results are discussed in the context of host-affiliated ecological selection as a potential source driving diversification in parasitoid communities and the influence of host species heterogeneity on population differentiation and local adaptation.     

On‐farm evaluation of a fall‐seeded rye cover crop for suppression of soybean aphid (Hemiptera: Aphididae) on soybean

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The effectiveness of flower strips and hedgerows on pest control,pollination services and crop yield: a quantitative synthesis

Floral plantings are promoted to foster ecological intensification of agriculture through provisioning of ecosystem services. However, a comprehensive assessment of the effectiveness of different floral plantings, their characteristics and consequences for crop yield is lacking. Here we quantified the impacts of flower strips and hedgerows on pest control (18 studies) and pollination services (17 studies) in adjacent crops in North America, Europe and New Zealand. Flower strips, but not hedgerows, enhanced pest control services in adjacent fields by 16% on average. However, effects on crop pollination and yield were more variable. Our synthesis identifies several important drivers of variability in effectiveness of plantings: pollination services declined exponentially with distance from plantings, and perennial and older flower strips with higher flowering plant diversity enhanced pollination more effectively. These findings provide promising pathways to optimise floral plantings to more effectively contribute to ecosystem service delivery and ecological intensification of agriculture in the future.     

中国生物多样性核心监测指标遥感产品体系构建与思考

生物多样性的稳定维持关乎人类生存发展与地球健康。生物多样性核心监测指标(Essential Biodiversity Variables, EBVs)旨在结合地面调查与遥感技术, 为大尺度、长时间序列的生物多样性监测提供新的解决方案。然而, 目前学界仍然缺乏一套国家尺度标准化EBVs遥感监测产品数据集, 以进行生物多样性评估。本研究旨在对中国生物多样性核心监测指标遥感产品进行体系构建与思考, 首先综述了目前EBVs的遥感研究概况, 并根据EBVs研究文献的数量进行调研分析; 同时, 本文在已有遥感生物多样性产品优先标准的基础上, 添加了“可重复性”的新标准, 并据此构建了中国EBVs遥感产品体系与监测数据集的指标清单, 最终对中国EBVs遥感研究存在的问题进行思考与讨论。本研究可为中国的生物多样性遥感监测提供科学依据, 有望为中国生物多样性政策的制定提供支撑。     

Molecular identification of predation by carabid beetles on exotic and native slugs in a strawberry agroecosystem

Generalist predators have the capacity to exert significant pressure on prey populations. However, integrating them into biological control programs relies on a detailed understanding of their foraging behavior and the levels of trophic connectedness with pest species. Carabid beetles are important predators of slugs, pests of agricultural, floricultural and horticultural crops worldwide, but these interactions have been rarely studied outside the Western Palearctic ecozone. Diagnostic molecular gut-content analysis was used to examine the strength of trophic pathways between a community of carabid beetles and two slug species, the exotic Deroceras reticulatum and native Deroceras laeve, in strawberry agroecosystems. Strawberries were grown according to standard horticultural practices for central Kentucky, following traditional bare ground planting or with the addition of detrital subsidies, to quantify the impact of habitat management on the abundance of pests and the strength of these trophic pathways. Following laboratory characterization of species-specific molecular markers targeting both Deroceras species, carabid beetles collected from a strawberry agroecosystem were screened for slug DNA. Field collections revealed important food web pathways existed between Harpalus pensylvanicus and D. reticulatum, with 7.2% screening positive for these prey yet none screening positive for D. laeve. In contrast, Chlaenius tricolor was found to feed on both slug species in the field, with 16% screening positive for both Deroceras. Despite below normal rainfall limiting slug densities in the field, the results presented here reveal the potential importance of carabid beetles in slug population dynamics in the Nearctic.     

Impact of simulated drought on ecosystem biomass production: an experimental test in stream mesocosms

Climate models predict widespread shifts in precipitation patterns and increases in the frequency of extreme events such as droughts, but consequences for key processes in affected ecosystems remains poorly understood. A 2‐year manipulative experiment used a series of stream mesocosms to test the effect of recurrent drought disturbance on the composition and secondary production of macroinvertebrate consumer assemblages and functional groups. On average, secondary production in drought‐disturbed communities (mean 4.5 g m^?2 yr^?1) was less than half of that that in controls (mean 10.4 g m^?2 yr^?1). The effects of the drought differed among functional feeding groups, with substantial declines for detritivore shredders (by 69%) and engulfing predators (by 94%). Contrasting responses were evident among taxa within most functional feeding groups, ranging from extirpation to irruptions in the case of several small midge larvae, but production of most species was suppressed. Taxon‐specific responses were related to body mass and voltinism. The ratio of production to biomass (community P/B) increased under drought, reflecting a shift in production from large long‐lived taxa to smaller taxa with faster life cycles. This research provides some of the first experimental evidence of the profound effects that droughts can have on both the structure and functioning of aquatic ecosystems.     

Ecological impacts of invasive alien plants: a meta-analysis of their effects on species, communities and ecosystems

Biological invasions cause ecological and economic impacts across the globe. However, it is unclear whether there are strong patterns in terms of their major effects, how the vulnerability of different ecosystems varies and which ecosystem services are at greatest risk. We present a global meta-analysis of 199 articles reporting 1041 field studies that in total describe the impacts of 135 alien plant taxa on resident species, communities and ecosystems. Across studies, alien plants had a significant effect in 11 of 24 different types of impact assessed. The magnitude and direction of the impact varied both within and between different types of impact. On average, abundance and diversity of the resident species decreased in invaded sites, whereas primary production and several ecosystem processes were enhanced. While alien N-fixing species had greater impacts on N-cycling variables, they did not consistently affect other impact types. The magnitude of the impacts was not significantly different between island and mainland ecosystems. Overall, alien species impacts are heterogeneous and not unidirectional even within particular impact types. Our analysis also reveals that by the time changes in nutrient cycling are detected, major impacts on plant species and communities are likely to have already occurred.     

Impact of plant architecture and extraguild prey density on intraguild predation in an agroecosystem

Several models and experimental studies conducted in confined environments have shown that intraguild predation (IGP) can modulate population abundances and structure communities. A number of ecological and abiotic factors determine the nature and frequency of IGP. This study examined the effect of plant architecture and extraguild prey density, and their interactions, on the occurrence of IGP between two species of ladybird, Harmonia axyridis (Pallas) and Propylea quatuordecimpunctata L. (both Coleoptera: Coccinellidae). Theoretical concepts predict that IGP levels would increase with a decrease of both extraguild prey density and plant structural complexity. We conducted a factorial experiment in an open soybean field into which coccinellid larvae were introduced in experimental plots for a period of 5 days. We tested two levels of soybean aphid [Aphis glycines Matsumara (Hemiptera: Aphididae)] density, low and high (100 and 1 000 aphids per plot, respectively), and two levels of plant complexity, low (by removing half of the branches from the soybean plants) and high (by leaving plants intact). We used species‐specific molecular markers to detect the presence of P. quatuordecimpunctata in the digestive tract of H. axyridis. Molecular gut‐content analysis of H. axyridis revealed that rates of IGP were higher (20%) at low aphid density than at high aphid density (<6%). Decreased plant complexity did not impact the frequency of IGP. In accordance with existing literature, this study demonstrates that IGP is amplified at low extraguild prey density. We conclude that considering environmental factors, such as extraguild prey density, is crucial to improve our ability to predict the impact of intraguild predation on community structure and, from an applied perspective, biological control.     

The role of biodiversity for element cycling and trophic interactions: an experimental approach in a grassland community

The focus of a new experiment, set up in Jena in spring 2002, are the effects of biodiversity on element cycles and the interaction of plant diversity with herbivores and soil fauna. The experimental design explicitly addresses criticisms provoked by previous biodiversity experiments. In particular, the choice of functional groups, the statistical separation of sampling versus complementarity effects, and testing for the effects of particular functional groups differ from previous experiments. Based on a species pool of 60 plant species common to the Central European Arrhenatherion grasslands, mixtures of one to 16 (60) species and of one to four plant functional groups were established on 90 plots (20 m × 20 m) with nested experiments. In order to test specific hypotheses 390 additional small-area plots (3.5 m × 3.5 m) were set-up. Exact replicates of all species mixtures serve to assess the variability in ecosystem responses. In a dominance experiment, the effects of interactions among nine selected highly productive species are studied. Each species is grown as monoculture replicated once.Effekte der Biodiversität auf Elementkreisläufe und Wechselwirkungen der pflanzlichen Artenvielfalt mit Bodenfauna und Herbivoren stehen im Mitttelpunkt eines neuen Experiments, das im Frühjahr 2002 in Jena eingerichtet wurde. Das Versuchsdesign berücksichtigt ausdrücklich die Kritik, die an den Aufbau früherer Biodiversitätsversuche gerichtet wurde. Die Auswahl funktioneller Gruppen von Pflanzenarten, die statistischen Möglichkeiten, die Effekte des “Sampling” gegen Komplementarität zu trennen sowie den Einfluß funktioneller Gruppen zu überprüfen, unterscheiden dieses Experiment von früheren Versuchen. Sechzig typische Pflanzenarten der zentraleuropäischen Frischwiesen (Arrhenatherion) bilden den Artenpool für den Versuch. Auf 90 Flächen wurden Artenmischungen etabliert, die 1 bis 16 (60) Arten und 1 bis 4 funktionelle Gruppen dieser Pflanzenarten enthalten. Die Versuchsparzellen haben eine Größe von 20 m × 20 m, auf denen in genesteter Anordnung verschiedene Teilexperimente durchgeführt werden. Zusätzlich wurden 390 kleine Parzellen (3.5 m × 3.5 m) angelegt, um spezifische Hypothesen zu überprüfen. Alle Arten werden hier mit je einer Wiederholung als Monokulturen kultiviert. Identische Wiederholungen aller Artenmischungen sollen deren Variabilität untersuchen. In einem Dominanz-Versuch werden die Effekte der Wechselwirkungen zwischen 9 ausgewählten hochproduktiven Arten untersucht.     

Impact of native predators and parasites on Spodoptera exigua, an introduced pest of alfalfa hay in northern California

A long-term study of Spodoptera exigua (Hübner) (Lepidoptera: Noctuidae) revealed that a complex of native predators and parasites exploit this introduced pest in alfalfa-hay fields in northern California. In field trials, predators typically fed on >50% of sentinel egg masses during a given sample interval; mean predation rate ranged from 40 to 100% and percentage predation per individual egg mass was density-independent (spatial context) for 16 of 17 sample intervals. The egg-predator guild consisted primarily of adults and nymphs of Lygus hesperus Knight & L. elisus Van Duzee (Miridae), Nabis americoferus Carayon (Nabidae), and Orius tristicolor (White) (Anthocoridae); and adults of Collops vittatus (Say) (Melyridae) and Hippodamia convergens Guerin-Meneville (Coccinellidae). In laboratory feeding trials, these species readily fed on eggs, as well as neonate larvae of S. exigua. The parasite guild associated with small- to medium-sized larvae consisted of Hyposoter exiguae (Viereck) and Pristomerus spinator (F.) (Ichneumonidae), both larval endoparasites; and Chelonus insularis Cresson (Braconidae), an egg-larval endoparasite. Total parasitization by these species in field samples ranged from 5 to 75%, and was density-independent (spatial context) for each of four sample dates. A few larvae were infected with a nuclear polyhedrosis virus but no epizootics were observed. These results are consistent with the hypothesis that native natural enemies, especially generalist predators, can maintain populations of S. exigua at levels well below the treatment threshold for this exotic pest of alfalfa grown for hay.     

农作物授粉生态系统服务评估——以丽江老君山地区为例

为科学评估野生传粉昆虫对农作物授粉生态系统服务能力以及空间格局,基于不同土地利用/覆被类型上各种野生传粉蜂的蜜粉源植物和其巢穴资源的可得性,并结合其传粉的飞行行为及对其传粉有一定依赖性的作物,以ArcGIS9.2为平台构建了针对每个农作物像元的生态系统传粉授粉服务能力评估模型。并在云南丽江老君山地区,以当地野生东方蜜蜂(Apiscerana Fabricius)为例进行了模型的应用。结果表明:当地野生东方蜜蜂在森林的平均生境质量指数最高,达到2.24×10-2;而在栽培植被最低,为0.37×10-2。由于研究区西部农作物区分布零散,周边天然林可提供丰富的蜜粉源和栖息地;而东部农作区集中连片,作物相单一,传粉有较大的空间障碍,且周边多为次生林,研究区农田范围内授粉服务能力空间分布呈现"西高东低"格局。     

Differential impact of adults and nymphs of a generalist predator on an exotic invasive pest demonstrated by molecular gut-content analysis

Generalist predators have the capacity to regulate herbivore populations through a variety of mechanisms, but food webs are complex and defining the strength of trophic linkages can be difficult. Molecular gut-content analysis has revolutionized our understanding of these systems. Utilizing this technology, we examined the structure of a soybean food web, identified the potential for adult and immature Orius insidiosus (Hemiptera: Anthocoridae) to suppress Aphis glycines (Hemiptera: Aphididae), and tested the hypotheses that foraging behaviour would vary between life stages, but that both adults and immatures would exert significant predation pressure upon this invasive pest. We also identified the strength of trophic pathways with two additional food items: an alternative prey item, Neohydatothrips variabilis (Thysanoptera: Thripidae), and an intraguild predator, Harmonia axyridis (Coleoptera: Coccinellidae). A. glycines constituted a greater proportion of the diet of immature O. insidiosus, but N. variabilis DNA was found in greater frequency in adults. However, both life stages were important early-season predators of this invasive pest, a phenomenon predicted as having the greatest impact on herbivore population dynamics and establishment success. No adult O. insidiosus screened positive for H. axyridis DNA, but a low proportion (2.5%) of immature individuals contained DNA of this intraguild predator, thus indicating the existence of this trophic pathway, albeit a relatively minor one in the context of biological control. Interestingly, approximately two-thirds of predators contained no detectable prey and fewer than 3% contained more than one prey item, suggesting the possibility for food limitation in the field. This research implicates O. insidiosus as a valuable natural enemy for the suppression of early-season A. glycines populations.     

Impact of elevated CO2 on the third trophic level: A predator Harmonia axyridis and a parasitoid Aphidius picipes

The effects of elevated CO₂ (550 and 750 µL/L vs. ambient CO₂) on the third trophic level, a predator Harmonia axyridis (Pallas) and a parasitoid Aphidius picipes (Nees), were studied in open-top chambers. The impact of elevated CO₂ on the growth and development of H. axyridis was either weak or nonexistent, whereas the abundance of the parasitized aphid (Sitobion avenae Fabricius) by A. picipes showed a significant increase in 550 (12.5%) and 750 (19.6%) µL/L CO₂ compared to ambient CO₂, respectively. In addition, there was a significant decrease (10%) in the emergence rate of A. picipes under 750 µL/L CO₂ compared to ambient CO₂ (P<0.05). The predator and the parasitoid both substantially suppressed aphid abundance, especially in elevated CO₂ for A. picipes. Moreover, H. axyridis and A. picipes preferred to prey on/parasitize more aphids, S. avenae, infested on 550 (9.1 and 16.9%) and 750 (23 and 25.7%) µL/L CO₂-grown wheat plants than those fed ambient CO₂-grown wheat plants. These initial results indicate that elevated CO₂ markedly changes the predation/parasitization preference by the predator/parasitoid for wheat aphids. The biocontrol efficiency of A. picipes against S. avenae can be enhanced in elevated CO₂; simultaneously, elevated CO₂ has adverse effects on the growth and development of A. picipes-parasitized S. avenae.