Manipulating carabid beetle abundance alters prey removal rates in corn fields

Generalist natural enemies such as carabid beetles have the potential to maintain a variety of pests below outbreak levels in annual crops. To assess the relationship between carabid beetle abundance and field rates of prey removal, we created plots surrounded by different boundaries that selectively affected dispersal of edaphic arthropods, primarily carabids. Three treatments were established: (1) naturally occurring communities, (2) augmented communities using ingress boundaries, and (3) reduced communities using egress boundaries. Selective boundaries altered carabid communities with minimal habitat alteration and without use of insecticides. Three times during the growing season, a fixed number of onion fly pupae were placed in plots to evaluate the impact of carabid abundance on predation rates. A combination of vertebrate and invertebrate exclosures allowed us to evaluate prey removal by invertebrates alone. In comparison to the no boundary treatment, carabids increased 54.2% and decreased 83.1% in plots surrounded by ingress and egress boundaries respectively. Predation rates were positively correlated with carabid abundance (r² = 0.70, p < 0.0001). Significantly more pupae were removed from exclosures allowing access to invertebrates alone than from total exclosures, suggesting that invertebrates represented an important group of predators. Laboratory trials tested the feeding potential of the four most abundant carabid species and showed that they readily consumed onion fly pupae, supporting our hypothesis that carabids were the main predators in field tests. This study corroborates and extends previous observations of the importance of carabid beetles as generalist predators of insect pests in agricultural fields.     

Trophic links between functional groups of arable plants and beetles are stable at a national scale

1. There is an urgent need to accurately model how environmental change affects the wide-scale functioning of ecosystems, but advances are hindered by a lack of knowledge of how trophic levels are linked across space. It is unclear which theoretical approach to take to improve modelling of such interactions, but evidence is gathering that linking species responses to their functional traits can increase understanding of ecosystem dynamics. Currently, there are no quantitative studies testing how this approach might improve models of multiple, trophically interacting species, at wide spatial scales. 2. Arable weeds play a foundational role in linking food webs, providing resources for many taxa, including carabid beetles that feed on their seeds and weed-associated invertebrate prey. Here, we model associations between weeds and carabids across farmland in Great Britain (GB), to test the hypothesis that wide-scale trophic links between these groups are structured by their species functional traits. 3. A network of c. 250 arable fields, covering four crops and most lowland areas of GB, was sampled for weed, carabid and invertebrate taxa over 3 years. Data sets of these groups were closely matched in time and space, and each contained numerous species with a range of eco-physiological traits. The consistency of trophic linkages between multiple taxa sharing functional traits was tested within multivariate and log-linear models. 4. Robust links were established between the functional traits of taxa and their trophic interactions. Autumn-germinating, small-seeded weeds were associated with smaller, spring-breeding carabids, more specialised in seed feeding, whereas spring-germinating, large-seeded weeds were associated with a range of larger, autumn-breeding omnivorous carabids. These relationships were strong and dynamic, being independent of changes in invertebrate food resources and consistent across sample dates, crops and regions of GB. 5. We conclude that, in at least one system of interacting taxa, functional traits can be used to predict consistent, wide-scale trophic links. This conceptual approach is useful for assessing how perturbations affecting lower trophic levels are ramified throughout ecosystems and could be used to assess how environmental change affects a wider range of secondary consumers.     

植草带宽度对蜘蛛和步甲群落动态及比邻玉米田害虫发生的影响

农田边界植草带是农田系统各斑块间生物物种迁移和群落演替、物质和能量交换的主要区域,对维持农业生物多样性及提高农田生产力具有重要意义。基于作物-害虫-天敌互作有机整体对植草带宽度的响应,选取不同宽度植草带(2 m、4 m、6 m)及其比邻农田并以对应宽度作物边界作为对照,采用地表陷阱法对蜘蛛和步甲进行取样调查,探究植草带宽度对蜘蛛和步甲种群结构、迁移扩散规律以及对农田害虫控制潜力的影响。结果表明:植草带内蜘蛛和步甲个体数均显著高于对照条带,4 m、6 m植草带的蜘蛛和步甲多度显著高于2 m植草带,且物种丰富度和多度显著高于其对照条带。此外,不同宽度植草带蜘蛛和步甲群落存在较高的异质性,蜘蛛和步甲群落结构差异显著;在4 m和6 m植草带及其比邻玉米田内,蜘蛛和步甲多度在植草带和植草带与比邻玉米田交界处显著高于农田内部,且随距离的增加,蜘蛛物种多度呈显著降低趋势,植草带宽度和距离共同决定蜘蛛和步甲的空间分布;Monte Carlo检验结果表明,植草带宽度、植被Margalef指数对农田中蜘蛛和步甲群落结构的影响均达到显著水平,蜘蛛主要优势科和步甲主要优势种主要分布在6 m植草带比邻玉米田内;4 m、6 m植草带比邻的玉米田中蜘蛛和步甲对玉米害虫的益害比显著高于2 m植草带比邻的玉米田,且4 m、6 m植草带显著增加了玉米籽粒的百粒重及比邻农田的玉米总产量。本研究对植草带涵养捕食性天敌预防害虫发生的最适宽度进行探索,综合考虑植草带耕地占用、天敌涵养和生物控害潜力等因素,确定4 m宽度植草带是具有推广价值的农田边界模式。     

Monitoring carabid indicators could reveal environmental impacts of genetically modified maize

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Impact of Bt maize pollen (MON810) on lepidopteran larvae living on accompanying weeds

Environmental risks of Bt maize, particularly pollen drift from Bt maize, were assessed for nontarget lepidopteran larvae in maize field margins. In our experimental approach, we carried out 3-year field trials on 6 ha total. Three treatments were used in a randomized block design with eight replications resulting in 24 plots: (i) near-isogenic control variety without insecticide (control), (ii) near-isogenic control variety with chemical insecticide (Baytroid) and (iii) Bt maize expressing the recombinant toxin. We established a weed strip (20 x 1 m) in every plot consisting of a Chenopodium album (goosefoot)/Sinapis alba (mustard) mixture. In these strips we measured diversity and abundance of lepidopteran larvae during maize bloom and pollen shed. C. album hosted five species but all in very low densities; therefore data were not suitable for statistical analysis. S. alba hosted nine species in total. Most abundant were Plutella xylostella and Pieris rapae. For these species no differences were detected between the Bt treatment and the control, but the chemical insecticide treatment reduced larval abundance significantly. Conclusions regarding experimental methodology and results are discussed in regard to environmental risk assessment and monitoring of genetically modified organisms.     

A field experiment on the effectiveness of spiders and carabid beetles as biocontrol agents in soybean

1 Spiders and carabid beetles are abundant generalist predators that prey upon insect pests of soybean. A field experiment was conducted to determine the impact of spiders and carabids on soybean yield. Prior to planting, three 7 × 7 m plots were fenced in order to reduce spider and carabid immigration. Carabids that emerged within the plots were not removed, but spiders that ballooned into these predator‐reduction plots or that entered by climbing the fence were removed by pitfall trapping and searching the vegetation. Three unmanipulated, unfenced plots served as the control treatment. 2 Densities of spiders on soybean vegetation, and activity‐densities of spiders and carabids determined by pitfall trapping, were c. 75% lower in the spider‐carabid reduction treatment than in control plots. Despite clear differences between treatments in numbers and activity of these major generalist predators, the weight of soybeans harvested did not differ between control and spider‐carabid reduction plots. 3 Paralleling the absence of an effect of predator reduction on soybean yield was the absence of any significant difference between treatments in densities of whiteflies (Aleyrodidae), leafhoppers (Cicadellidae), thrips (Thysanoptera), Lepidoptera larvae and herbivorous Coleoptera. 4 Our experiment provides no evidence that spiders and carabid beetles at ambient densities affect soybean yield. Low populations of pest species or low predation pressure on soybean pests by spiders and carabids at the ambient densities of this experiment could be responsible for this result.     

转植酸酶基因玉米对步甲群落动态的影响

本研究以陷阱法研究转植酸酶基因玉米及其亲本玉米在整个生长季节对地表步甲类群的影响。每个玉米品系设置6块样地, 两个玉米品系交替排列, 每个样地设置2个采样点, 共设置24个样点, 整个玉米生育期取样8次。本研究共采集步甲标本8 012头, 隶属于11属23种, 其中黄斑青步甲Chlaenius micans个体数量分别占个体总数的87.54%, 为玉米田内的主要常见物种, 蠋步甲Dolichus halensis、 后斑青步甲Chlaenius posticalis和单齿蝼步甲Scarites terricola个体数量分别占除黄斑青步甲之外所有步甲个体总数的34.77%, 31.16%和6.21%, 这些物种组成当地玉米田内的常见物种。步甲物种多样性与常见物种个体数量随季节变化明显, 且转植酸酶基因玉米与其亲本玉米田内步甲物种多样性和常见物种个体数量的季节变化趋势相似度较高。重复测量方差分析（Repeated ANOVA）结果表明, 转植酸酶基因玉米对步甲物种丰富度、 物种多度、 香农威纳多样性指数和均匀度指数均没有显著影响; 转植酸酶基因玉米田内的后斑青步甲个体数量明显增加, 但其他常见物种没有显著变化。基于非度量多维度（NMDS）的群落结构分析表明转植酸酶基因玉米与亲本玉米田内的步甲群落结构非常相似。本文结果表明转植酸酶基因玉米的种植对步甲物种多样性及常见物种没有明显影响。     

Ground beetle (Coleoptera: Carabidae) assemblages in conventional and diversified crop rotation systems

Ground beetles (Coleoptera: Carabidae) are important in agro-ecosystems as generalist predators of invertebrate pests and weed seeds and as prey for larger animals. However, it is not well understood how cropping systems affect ground beetles. Over a 2-yr period, carabids were monitored two times per month using pitfall traps in a conventional chemical input, 2-yr, corn/soybean rotation system and a low input, 4-yr, corn/soybean/triticale-alfalfa/alfalfa rotation system. Carabid assemblages were largely dominated by a few species across all cropping treatments with Poecilus chalcites Say comprising >70% of pitfall catches in both years of study. Overall carabid activity density and species richness were higher in the low input, 4-yr rotation compared with the conventionally managed, 2-yr rotation. There were greater differences in the temporal activity density and species richness of carabids among crops than within corn and soybean treatments managed with different agrichemical inputs and soil disturbance regimes. Detrended correspondence analysis showed strong yearly variation in carabid assemblages in all cropping treatments. The increase in carabid activity density and species richness observed in the 4-yr crop rotation highlights the potential benefits of diverse crop habitats for carabids and the possibility for managing natural enemies by manipulating crop rotations.     

Life-history trait and functional diversity patterns of ground beetles and spiders along a coastal heathland successional gradient

We investigated patterns in species richness and diversity, life-history traits and functional diversity of carabid beetles and spiders along a coastal heathland successional gradient. We sampled in five successional stages on the island of Hiddensee, Germany. Species richness of carabids and spiders and Simpson diversity for spiders did not differ among stages. Functional dispersion (FDis), a measure of functional richness, was lowest in the youngest stage for both carabids and spiders. Older successional stages represent more heterogeneous habitats and offer a broader range of niches, which might enhance functional dispersion. Functional evenness (FEve) differed among successional stages for spiders, indicating an uneven distribution of species abundances in the functional trait space. Functional divergence (FDiv) did not differ among stages for either taxon. Shifts in life-history traits were noted in both arthropod groups, but not always in the same direction: Body size of carabid beetles increased towards older successional stages, but decreased for spiders. The number of flightless carabid species increased towards older habitats. Each successional stage seemingly favors species with distinct life-history traits (species sorting). We conclude that a trait-based approach including the use of functional diversity measurements (FDis) could contribute valuable information for biodiversity conservation.     

Effects of cultivation of genetically modified Bt maize on epigeic arthropods (Araneae; Carabidae)

A field study was conducted in Germany to determine the possible effects of transgenic maize cultivation on nontarget epigeic predator organisms. During the growing period of 2001-2003, the activity abundances of spiders and carabid beetles were recorded and compared in three treatments: (1) Bt-maize (Mon 810) expressing the Cry1ab protein from Bacillus thuringiensis (Berliner), (2) an isogenic variety, and (3) the isogenic variety treated with insecticide. All three treatments were replicated in eight plots. The results were evaluated using three different methods. The activity abundances of single species were statistically analyzed by confidence interval methods. In addition, the phenological behaviors of the spider and carabid beetle species were determined, and multivariate statistical evaluation of the community by principal component analysis was conducted. Significantly different activity abundances in Bt plots compared with isogenic control plots were observed both for spiders and carabid beetles during 2001. However, in 2002 and 2003, no changes in community structure were detectable in any of the treatments. The change in the first year may have been caused by the influence of a massive cornborer infestation and accompanying large changes in microclimatic factors.     

In-field cover crop strips support carabid communities and shape the ecological trait repartition in maize fields

1. The enhancement of pest regulation service in crops depends for a large part on the capacity of agroecological practices to increase the presence of key species or functional traits in arthropod communities within fields.
2. We investigated the effects of undestroyed strips of winter cover crops in maize fields on carabid community composition, and on the distribution of three ecological traits: diet, wing status and body size.
3. We found that the community composition and the distribution of ecological traits in the in-field cover crop strips had commonalities with both adjacent cropped areas and field margins. Some species were recorded mostly or only in the strips indicating that strips could support carabid species and help increase local diversity from the first year of establishment.
4. The activity-density of Poecilus cupreus and Pterostichus melanarius was higher in the cropped proximity of the strip, and the body size was influenced by the distance from the strip.
5. Our results suggest that carabid communities are shaped by the habitat type, but the influence of such agroecological infrastructures on communities of adjacent crops is minor beyond a distance of 10 m. However, overall species abundance was increased and thus potentially provided enhanced pest regulation.

     

How good are carabid beetles (Coleoptera,Carabidae) as indicators of invertebrate abundance and order richness?

Ground invertebrates were sampled by pitfall trapping on Brentmoor Heath, Surrey, UK during the summers of 2009 and 2010 to determine whether the abundance and species richness of carabids on bare ground patches are representative of invertebrate abundance and order richness on the same patches. Four patch sizes were investigated as well as the adjacent mown and unmown areas, and a range of environmental parameters were measured at each patch. Results show that increasing abundance and richness of carabids can give a broad indication of whether invertebrate abundance and order richness are increasing. Invertebrates in general are sensitive to a greater number of environmental variables and complex interactions than carabids are, due to the vast array of biological requirements of the large range of invertebrate species. All else being equal, smaller patches appear to benefit both carabid abundance and invertebrate abundance. Converse to the results of carabid species richness, larger patches appear to benefit invertebrate order richness. This study highlights the importance of testing the effect of habitat management strategies on a range of taxa rather than assuming the results for a bioindicator will suffice for management decisions. Whilst bioindicators may give a broad indication of effects on similar taxa, the precise nature of those effects will be determined by a number of species- and site-specific variables.     

Rubidium marking reveals different patterns of movement in four ground beetle species (Col., Carabidae) between adjacent alfalfa and maize

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Distribution of forest dwelling carabids (Coleoptera): spatial scale and the concept of communities

Associations between spatial distribution of ground-beetles (Carabidae) and environmental variables were studied over three hierarchical scales in deciduous forest in central Alberta, Canada We also examined the relationship between species abundance and distribution on several scales ranging from the local scale of our study to that of the North American temperate deciduous forest Understorey plant cover, tree cover, and occurrence of other carabids were associated with distribution of particular species at the smallest ecological scales within populations However, great differences in population sues of carabid species among five distinct sites several kilometres apart were not correlated with variation in the same environmental variables In central Alberta, abundance and extent of distribution were correlated positively among the 30 carabid species collected, and distributions of the ten species classified as 'core' species were generally aggregated at all spatial scales On the continental scale, there was a significant positive correlation between abundance and distribution for the 114 species of the entire data set, and the six species meeting the criteria of 'core' taxa on this scale, were also 'core' elements in central Alberta Further analysis of covariance of core elements of species assemblages across different taxa provides a sound empirical approach for understanding community organization     

Carabid beetle assemblage under diversified agroecosystems

Corn (Zea mays L.) and soybean (Glycine max Merrill) were planted in monoculture and strip-intercropping plots (18.3×15.5 m each) under conventional tillage and no-tillage to create differences in habitat structure. The effects of these agronomic practices on carabid beetle species composition were measured by pitfall-trap collections during three years. Although neither the number of individuals or number of species were significantly affected by treatments, analyses of variance indicated that four of the six dominant species (75.7% of the total number of carabids captured in three years) responded to tillage treatments.Harpalus pensylvanicus DeGeer andCyclotrachelus sodalis (LeConte) were more abundant in no-tillage plots, whereasPterostichus chalcites Say andScarites substriatus Haldeman were more common in conventional tillage plots.Pterostichus chalcites was also significantly more abundant in soybean plots than in intercropping or corn plots.H. pensylvanicus, a primarily herbivorous carabid, was the most common species collected. However,P. chalcites, a predaceous carabid, was the dominant species in soybean/conventional tillage plots. The results suggest that characteristics of individual species should be carefully considered for studies of communities such as carabid beetles, which have few strongly dominant species and a wide range of resource utilization.     

Useful model organisms,indicators, or both? Ground beetles (Coleoptera,Carabidae) reflecting environmental conditions

Classic studies have successfully linked single-species abundances, life-history traits, assemblage structures and biomass of carabid beetles to past and present, human-caused environmental impacts and variation in ‘natural’ conditions. This evidence has led many to suggest carabids to function as ‘indicators’ − a term that bears multiple meanings. Here, a conservation-oriented definition for an indicator is used, carabid indicator potential from seven views is evaluated, and ways to proceed in indicator research are discussed. (1) Carabid species richness poorly indicates the richness and abundance of other taxa, which underlines the importance of using multiple taxa in environmental assessments. The ability of assemblage indices and specialist or functional-group abundances to reflect rare species and habitats should be examined in detail. (2) Experimental evidence suggests that carabids may potentially serve as keystone indicators. (3) Carabids are sensitive to human-altered abiotic conditions, such as pesticide use in agro-ecosystems and heavy metal contamination of soils. Carabids might thus reflect ecological sustainability and ‘ecosystem health’. (4) Carabid assemblages host abundant species characteristic of particular habitat types or successional stages, which makes them promising dominance indicators. (5) Carabids reflect variation in ‘natural’ conditions, but vegetation and structural features are more commonly adopted as condition indicators. Carabids nevertheless provide yet another, equally accurate, view on the structure of the environment. (6) Carabids may function as early-warning signalers, as suggested by recent studies linking climate and carabid distributions. (7) Carabids reflect natural and human-caused disturbances and management, but the usefulness of these responses for conservation purposes requires further research. In summary, European carabids appear useful model organisms and possibly indicators because they are diverse, taxonomically and ecologically well-known, efficiently reflect biotic and abiotic conditions, are relevant at multiple spatial scales, and are easy to collect in sufficiently large numbers to allow statistical analyses. The assumption that carabid responses would reflect rare environmental conditions or the responses of rare and threatened species ‒ crucial information for conservationists and managers ‒ has not yet been critically evaluated. Even if it holds, the usefulness will be context dependent: species and their populations vary, conditions vary, questions put forward vary, and assessment goals vary.     

Epigeic spiders are not affected by the genetically modified maize MON 88017

The genetically modified (GM) maize MON 88017 facilitates weed management owing to its tolerance to glyphosate, and resists western corn rootworm (WCR), Diabrotica virgifera virgifera, owing to the expression of Cry3Bb1 toxin. MON 88017 could therefore contribute to the solution of two major problems of European agriculture: continuous WCR spreading and high use of herbicides. To assess possible unwanted environmental impacts of MON 88017, we compared communities of spiders in plots planted in three successive years with this maize, its near isogenic non‐GM cultivar treated or not treated with an insecticide and two unrelated maize cultivars. Each of the five treatments was applied on five 0.5 ha plots in a 14 ha field. Spiders were collected in five pitfall traps per plot five times per year. Upon reaching the waxy ripening stage, all plants of first‐year cultivation were shredded to small pieces and ploughed into the soil in the respective plot, whereas in the 2nd and 3rd year the harvest was used for biogas production and only digestate was returned to the field. Out of 79 spider species, Pardosa agrestis, Pachygnatha degeeri and Oedothorax apicatus made up 28%, 25% and 23% of the total spider count in the 1st year of study; 2%, 8% and 84% in the 2nd; and 40%, 8% and 35% in the 3rd year. Statistical analysis did not reveal any influence of GM maize on the spider abundance and biodiversity. The abundance, and in two years also the species diversity, was insignificantly higher on the plots with GM maize than on plots with the insecticide‐treated non‐GM maize. The composition and size of spider community varied year to year, probably reflecting weather conditions and differences in field fertilization with organic matter.     

华北盐渍化改造区农田步甲群落及其动态研究:以河北省曲周县为例

1996与1997年在河北省曲周县盐渍化改造区,采用陷阱法调查了农田中步甲的物种组成与数量分布。共记录到步甲19个种,分隶于8个属。其中婪步甲属(Harpalus)在数量上占绝对优势。总体来说,本地步甲区系组成相对简单,属与物种数较少。单齿婪步甲(Harpalus simplicidens)、毛婪步甲(H.griseus)和三齿婪步甲(H.tridens)为优势物种。从时间动态上看,步甲数量呈总体上升趋势,9月份最多,10月份下降到最低。步甲数量的季节变动在不同种植模式间不同,并在一定程度上受农田景观及农事活动的影响。农田边界是生物的避难所和聚集地,对步甲有重要的保护作用,农田边界上的步甲密度普遍比相邻农田中高,月份间步甲数量的波动幅度也较小。     

Progressively excluding mammals of different body size affects community and trait structure of ground beetles

Mammalian grazing induces changes in vegetation properties in grasslands, which can affect a wide variety of other animals including many arthropods. However, the impacts may depend on the type and body size of these mammals. Furthermore, how mammals influence functional trait syndromes of arthropod communities is not well known. We progressively excluded large (e.g. red deer, chamois), medium (e.g. alpine marmot, mountain hare), and small (e.g. mice) mammals using size‐selective fences in two vegetation types (short‐ and tall‐grass vegetation) of subalpine grasslands. We then assessed how these exclusions affected the community composition and functional traits of ground beetles (Coleoptera, Carabidae), and which vegetation characteristic mediated the observed effects. Total carabid biomass, the activity densities of carabids with specific traits (i.e. small eyes, short wings), the richness of small‐eyed species and the richness of herbivorous species were significantly higher when certain mammals were excluded compared to when all mammals had access, regardless of vegetation type. Excluding large and medium mammals increased the activity density of herbivorous carabid species, but only in short‐grass vegetation. Similarly, excluding large mammals (ungulates) altered carabid species composition in the short‐, but not in the tall‐grass vegetation. All these responses were related to aboveground plant biomass, but not to plant Shannon diversity or vegetation structural heterogeneity. Our results indicate that changes in aboveground plant biomass are key drivers of mammalian grazers’ influence on carabids, suggesting that bottom–up forces are important in subalpine grassland systems. The exclusion of ungulates provoked the strongest carabid response. Our results, however, also highlight the ecological significance of smaller herbivorous mammals. Our study furthermore shows that mammalian grazing not only altered carabid community composition, but also caused community‐wide functional trait shifts, which could potentially have a wider impact on species interactions and ecosystem functioning.     

Impacts of Leaf-litter Addition on Carabids in a Conifer Plantation

As a fine-scale, manipulative model experiment leaf litter was added in plots to increase habitat heterogeneity in a 50-year-old Norway spruce plantation, established after the clear-cutting of a native beech forest, during a 2-year period in the Hungarian Mountain Range. Pitfall trap catches of carabids from leaf-litter plots were compared with those from control plots to explore the effect of leaf-litter addition. Difference in the species composition was revealed by ordination; scores of the samples of the two plot types were significantly separated along the first MDS axis. The most numerous species (Pterostichus oblongopunctatus) was significantly more abundant in the leaf-litter plots. However, there were no significant differences for the other most frequently obtained species. Habitat generalist species were the most abundant, followed by forest generalists, then forest specialists, and there were some open habitat species. Enhanced habitat heterogeneity (leaf-litter addition) in homogeneous plantations influenced the spatial distribution and composition of carabids, through altered abiotic (lower ground temperature in the leaf-litter plots) and biotic (more prey items) factors. Differences in abundance, species richness and Shannon diversity were not significant between the control and the manipulated plots, although carabid catch was higher in the leaf-litter plots during both years. Forestry practices to increase habitat heterogeneity should be considered to enhance biodiversity in managed forests.