Influence of within‐field position and adjoining habitat on carabid beetle assemblages in winter wheat

• 1 The influence of within‐field position and adjoining habitat on carabid beetles was studied in 20 winter wheat fields in ten different Swiss agricultural landscapes. In each landscape, two winter wheat fields (one with adjoining sown wildflower area and one with adjoining grassy margin) were investigated.
• 2 Carabid beetles were caught in pitfall traps 3 and 30 m from the edge in each of the 20 wheat fields. Significantly more individuals were found in the centres (30‐m position) than at the edges (3‐m position). Conversely, species richness was significantly higher at the field edges than in the centres.
• 3 Of the ten most abundant species, Poecilus cupreus, Agonum muelleri and Pterostichus melanarius were significantly more abundant in the field centres than at the edges. Harpalus rufipes was significantly more abundant in the fields adjoining sown wildflower areas than in the fields adjoining grassy margins.
• 4 In conclusion, the response of carabid beetles to within‐field position and adjoining habitats was species specific. This needs to be taken into account in habitat management for biodiversity conservation and pest control.

     

Dispersal of a carabid beetle in farmland is driven by habitat‐specific motility and preference at habitat interfaces

Carabid beetles are common predators of pest insects and weed seeds in agricultural systems. Understanding their dispersal across farmland is important for designing farms and landscapes that support pest and weed biological control. Little is known, however, about the effect of farmland habitat discontinuities on dispersal behaviour and the resulting redistribution of these beetles. We released 1,985 well‐fed and 1,680 food‐deprived individuals of the predatory carabid beetle Pterostichus melanarius (Illiger) (Coleoptera: Carabidae) on a farm in Wageningen, The Netherlands. We recaptured 23.6% of those beetles over a period of 23 days in 2010. The farmland comprised agricultural fields with various crop species and tillage, separated by strips of perennial vegetation. We developed discrete Fokker‐Planck diffusion models to describe dispersal based on motility (m² day^?1) and preferential behaviour at habitat interfaces. We used model selection and Akaike’s information criterion to determine whether movement patterns were driven by variation in motility between habitats, preferential behaviour at habitat interfaces, or both. Model selection revealed differences in motility among habitats and gave strong support for preferential behaviour at habitat interfaces. Behaviour at interfaces between crop and perennial vegetation was asymmetric, with beetles preferentially moving towards the crop. Furthermore, beetles had lower motility in perennial strips than in arable fields. Also between arable habitats movement was asymmetric, with beetles preferentially moving towards the habitat in which motility was lowest. Neither crop type nor tillage explained differences in motility between crop habitats. Recapture data representing dispersal patterns of beetles were best described by a model that accounted for differences in motility between farmland habitats and preferential behaviour at habitat interfaces. Motility in farmland and behaviour at interfaces can also be estimated for other organisms and farmland habitats to support design of farmland conducive to natural pest suppression. Landscape design for early recruitment of carabids into arable fields should take into account the quantity and quality of resource habitats in the landscape, their proximity to crop fields, movement rates, and the possibility of movement responses at interfaces between landscape elements.     

Behavioral responses of three species of ground beetle (Coleoptera: Carabidae) to olfactory cues associated with prey and habitat

The responses ofPterostichus melanarius Illiger,Harpalus rufipes DeGeer, andNebria brevicollis Fabricius (Coleoptera: Carabidae) to olfactory cues of prey and habitat were studied in a four-arm continuous-airflow olfactometer. The process was semiautomated using time-lapse image analysis by microcomputer. The primary constituent of the aphid alarm pheromone, (E)--farnesene (EBF), was synthesized and tested for a kairomonal role in prey detection by carabids. In addition, individual beetles were exposed to odors from live aphids, live collembolans, and a crude extract of wheat. All three beetle species showed evidence of olfaction.P. melanarius responded to all the odors except collembolans,H. rufipes responded to EBF and wheat, andN. brevicollis to collembolans. The use of a defence allomone as a prey-finding kairomone by certain carabid species has implications for pest management. Manipulation of predator chemical ecology by the inclusion of behavior-modifying compounds in a crop spray mix with reduced amounts of insecticide may allow for efficient aphid control with less environmental contamination.     

Tillage differentially affects the capture rate of pitfall traps for three species of carabid beetles

The influence of tillage, gender, and microclimate on capture rates of pitfall traps for the beetles Poecilus scitulus LeConte, Poecilus lucublandus (Say), and Pterostichus melanarius Illiger (Coleoptera: Carabidae) were assessed in mark–release–recapture experiments in spring pea and spring wheat. Experiments were conducted during June, July, and August of 2003 in the Palouse region of northern Idaho, USA. Rates of capture in pitfall traps for the three carabid species were differentially affected by crop‐tillage systems. Capture rates for P. scitulus and P. lucublandus were higher in no‐till (NT) than in conventional tillage (CT) peas, whereas capture rates for P. scitulus and Pt. melanarius were higher in CT than in NT wheat. Ground‐level temperatures and relative humidity (r.h.) differed little among tillage systems. Nevertheless, capture rates were generally positively correlated with ground‐level temperature and negatively correlated with r.h., with correlations more often significant in NT than in CT systems. The response of the thermophilic Poecilus spp. to temperature provides a possible mechanistic explanation for capture rate patterns in legumes during June, but not in other months for peas or any month in wheat during the experiments. Movement impedance due to residue could explain lower capture rates of P. scitulus and Pt. melanarius in NT than in CT spring wheat. These results suggest that researchers using pitfall trapping for carabid populations should take into account potential capture biases their treatments can introduce.     

Augmentation of beneficial arthropods by strip-management

The effect of strip-management on surface activity, movements and activity density of abundant carabid beetles during two vegetation periods in a cereal field was investigated using mark-recapture techniques. Significantly higher recapture rates, indicating higher activity, were found in the strip-managed area than in the control area, especially in Poecilus cupreus and also in Carabus granulatus and Pterostichus melanarius. Several observations led to the conclusion that this higher activity is generally due to a prolongation of the reproductive period in the strip-managed area. Significantly higher activity densities were found for P. cupreus, Pterostichus anthracinus, C. granulatus and Pt. melanarius in the strip-managed area than in a bordering control area. P. cupreus is greatly attracted to the strips during its migrations, as can be deduced from the high percentage of movements that contact the strips. Also, significantly more marked individuals moved from the control to the strip-managed area than vice versa. Pt. melanarius and Pt. anthracinus show less preference for the strips in their movements than P. cupreus. However, significantly more individuals of Pt. melanarius also moved from the control to the strip-managed area. C. granulatus, on the other hand, kept mainly to the cereal areas. After harvest only Pterostichus niger and Harpalus rufipes distinctly preferred the strips. The generally marked degree of attraction exercised by strip-management on carabid beetles is discussed.     

Movements of carabid beetles (Coleoptera: Carabidae) inhabiting cereal fields: a field tracing study

Summary The individual movements of four carabid species were investigated by tracing beetles in the field using a portable radar system (harmonic radar). The field-inhabiting a Pterostichus melanarius Illiger, P. niger Schaller and Harpalus rufipes De Geer were traced in cereal fields. The movements of P. melanarius Mnd P. niger were also tracked in a wood and then compared with the movements of the forest-species Carabus nemoralis Müller. P. melanarius and P. niger displayed directed movements in cereal fields, although P. niger moved faster than P. melanarius. The movement pattern of H. rufipes in the field, and C. nemoralis and P. malanarius in the wood can be described as a c correlated random walk. In cereal fields, P. melanarius burrowed actively into the soil, and P. niger searched out crevices. In contrast, none of these two carabids burrowed in the forest soil, whereas C. nemoralis did. No statistically significant difference was observed between the movements of hungry and satiated beetles, although starved beetles tended to disperse shorter distances per move and turned back more often (especially P. niger). The significance of the various movement patterns exhibited by the species studied for their reproduction and survival on arable land is discussed.     

Optimal diet theory: behavior of a starved predatory snail

Summary The tenets of optimal foraging theory are used to contrast the behavior of the predatory snail Acantina spirata when feeding on the barnacles Balanus glandula and Chthamalus fissus under conditions of satiation and starvation. As predicted in optimal diet models, A. spirata is less selective (ratio of attack frequency on a prey species to number of individuals available) when the higher ranking prey has low abundance. When given a choice, starved snails attack both barnacle species equally, whereas satiated individuals preferentially attack B. glandula, the more profitable prey (ash-free dry weight of barnacles ingested per unit handling time). Under starvation conditions, equal attack frequency does not result in equal prey species consumption because Acanthina spirata is more successful at attacking C. fissus than B. glandula.The assumption of constant prey encounter rates in optimal diet models is not met when A. spirata goes from a state of satiation to starvation. The encounter rate on B. glandula is lowered due to a decrease in attack success. A loss of feeding skills in starved A. spirata is responsible for the greater difficulty snails have in gaining access through the opercular plates of B. glandula.Behavioral changes in A. spirata as snails pass from satiation to hunger translate into an energetic disadvantage during feeding for hungry snails for two reasons. First, higher prey handling times result in a decreased rate of biomass intake. Second, alteration in the relative attack frequency between barnacle species, combined with a decrease in attack success on the more profitable prey leads to more frequent ingestion of the less profitable prey.     

Predation by the Carabid Beetles Pterostichus melanarius and Poecilus cupreus on Slugs and Slug Eggs

Predation on slugs, Arion lusitanicus (Mabille) and Deroceras reticulatum (Müller), and their eggs by the carabid beetles Pterostichus melanarius (Illiger) and Poecilus cupreus (L.) was investigated in the laboratory. Slugs of different size and slug eggs were offered to the beetles in petri dishes with and without several alternative prey. Pterostichus melanarius destroyed eggs of D. reticulatum and A. lusitanicus, with a clear preference for D. reticulatum eggs. The availability of some alternative prey types adversely affected feeding on D. reticulatum eggs by P. cupreus and feeding on A. lusitanicus eggs by P. melanarius. Yet consumption of D. reticulatum eggs by P. melanarius was not significantly influenced by any alternative prey. Only P. melanarius killed small D. reticulatum in the presence of alternative prey. This suggests that P. melanarius and P. cupreus may have the potential to reduce slug populations in the field by destroying slug eggs and, in the case of P. melanarius, killing freshly hatched slugs.     

Monoclonal antibodies reveal changes in predator efficiency with prey spatial pattern

Spatially explicit predator–prey interactions can alter the predatory potential of natural enemies augmented through conservation biological control. To test hypotheses regarding such interactions and predatory efficiency, we used a combination of molecular techniques and mark–release–recapture to study the foraging behaviour of a generalist carabid predator, Poecilus cupreus , in response to spatial patterns of its cereal aphid prey ( Metapolophium dirhodum and Sitobion avenae ). Beetle and aphid numbers were measured across two grids of sampling locations, within which aphid spatial pattern had been manipulated to generate patchy and more homogenous distributions. Aphid consumption was measured by enzyme-linked immunosorbent assays (ELISA) of beetle gut contents, using an aphid-specific monoclonal antibody. Movement and distribution patterns suggest that P. cupreus does not aggregate at, nor instigate prey-taxis within, aphid patches. However, more than two-thirds of the 2169 P. cupreus tested by ELISA had consumed aphids and the proportion of beetles containing aphid proteins was positively related to aphid density. Against expectation, the proportion of predators feeding on aphids was greatest where prey were homogenously distributed, and this was attributed to the loss of partial refuges for prey in aphid patches. The functional value of this type of uniform foraging strategy is ideally suited to early colonization of the crop habitat, when aphid numbers are low, before populations build up and form strong spatial patterns.     

Movement of carabids from grassy strips to crop land in organic agriculture

The colonisation process of carabid beetles from field margins to crop fields under organic agriculture was investigated in 2010 and 2011 in northern Germany in order to find the supporting effects of grassy strips in the agricultural landscape. Displacement and speed of Pterostichus melanarius, Carabus auratus and Poecilus spp. were higher in the crops than in the grassy strips, except for the speed of C. auratus. Among the four species C. auratus, P. melanarius, Pterostichus niger, and Poecilus spp., C. auratus revealed the highest displacement and speed, whereas no significant differences were found between the other three species. P. melanarius showed higher displacement and speed in wheat than in red-clover or pumpkin. Speed and displacement of P. melanarius differed between spring and autumn generations in wheat or red-clover. When they were released at a junction of arable crop and field strips, C. auratus moved indifferently to either arable field or field margins/strips, whereas P. melanarius moved directly to crop centres. At the field margin, Poecilus spp. preferred to move along a grassy strip, while it moved irrespective from the grassy strips in a central site. C. nemoralis and P. niger preferred to move within the grassy strips. Our studies reveal that grassy strips do not affect the dispersal of typical arable field species; however, the species demanding grass vegetation can only overcome the barriers of arable fields either by grassy strips alone or by grassy strips with an adjacent hedge.     

Movement Behaviour of the Carabid Beetle Pterostichus melanarius in Crops and at a Habitat Interface Explains Patterns of Population Redistribution in the Field

Animals may respond to habitat quality and habitat edges and these responses may affect their distribution between habitats. We studied the movement behaviour of a ground-dwelling generalist predator, the carabid beetle Pterostichus melanarius (Illiger). We performed a mark-recapture experiment in two adjacent habitats; a large plot with oilseed radish (Raphanus sativus) and a plot with rye (Secale cereale). We used model selection to identify a minimal model representing the mark-recapture data, and determine whether habitat-specific motility and boundary behaviour affected population redistribution. We determined movement characteristics of P. melanarius in laboratory arenas with the same plant species using video recording. Both the field and arena results showed preference behaviour of P. melanarius at the habitat interface. In the field, significantly more beetles moved from rye to oilseed radish than from radish to rye. In the arena, habitat entry was more frequent into oilseed radish than into rye. In the field, movement was best described by a Fokker-Planck diffusion model that contained preference behaviour at the interface and did not account for habitat specific motility. Likewise, motility calculated from movement data using the Patlak model was not different between habitats in the arena studies. Motility (m² d⁻¹) calculated from behavioural data resulted in estimates that were similar to those determined in the field. Thus individual behaviour explained population redistribution in the field qualitatively as well as quantitatively. The findings provide a basis for evaluating movement within and across habitats in complex agricultural landscapes with multiple habitats and habitat interfaces.     

Pitfall trap catches of and aphid predation byPterostichus melanarius andPterostichus madidus in insecticide treated and untreated potatoes

Aphids were countedin situ and carabid beetle populations assessed by pitfall trapping betwen 1983–1985 in potato fields in Scotland treated with demeton-S-methyl (DSM) or untreated.Macrosiphum euphorbiae (Thomas) (Homoptera: Aphididae) was the most abundant aphid. Nineteen species of carabid beetles were trapped but the fauna was dominated byPterostichus melanarius (Ill.) andPterostichus madidus (Fabr.) (Coleoptera: Carabidae). Pitfall trap catches of these carabids were variable but generally lower in sprayed than unsprayed plots for a few days after DSM application and higher a week or two later. Of 1800P. melanarius and 910P. madidus dissected, 14.4 per cent and 30.5 per cent respectively, contained aphid remains. The proportion that had eaten aphids often was higher in sprayed than unsprayed plots immediately after treatment. In additional experiments, aphids treated with DSM fell from plants; four to five per cent of the theoretically applied dose of DSM reached the ground in a closed-canopy potato crop; and no mortality occurred whenPterostichus spp. were exposed to DSM in a field bioassay. It is proposed that observed patterns in trap catch ofPterostichus spp. around times of pesticide application may be due in part to aphids falling to the ground after treatment. Beetles that eat these fallen aphids may be less hungry, less active and less likely to be trapped. Later increases in catches may be attributed to hungry, more active beetles, a consequence of prey removal by the insecticide. Although this hypothesis requires further research, it is clear that the effects of insecticide treatment on carabid populations cannot be measured by pitfall trapping alone.     

Differentiation of spring carabid beetle assemblages between semi‐natural habitats and adjoining winter wheat

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Effects of narrow roads on the movement of carabid beetles (Coleoptera,Carabidae) in Nopporo Forest Park,Hokkaido

Forest-dwelling carabid beetles that have no flight ability were studied using mark-recapture methods in late-June to mid-October 2007. This study was done to determine the effects of narrow roads in Nopporo Forest Park, Hokkaido on carabid beetle movement and habitat use. The investigation was conducted at four sites: one site was an abandoned grassy road with a width of 3.5 m, two sites were gravel roads with widths of 3.5 and 4.5 m, and another site was an asphalt-paved road with a width of 4.5 m. A total of 3,580 individuals from six species of carabid beetles were collected using dry pit-fall traps, and recapture rates ranged from 6.1 to 36.2%. All examined roads acted as barriers against the movement of Leptocarabus arboreus ishikarinus. All roads, except the abandoned grassy road, acted as a barrier against Carabus granulatus yezoensis movement. Forest–roadside verge comparisons demonstrated that some carabid beetles avoid even narrow roadside verges. Harmful effects increase with increasing road width and both paved roads and narrow roads negatively affect the movement of carabid beetles inhabiting the bordering forest. Therefore, forest specialist beetles are influenced by a barrier effect that starts at the forest road verge, and this barrier effect may be exacerbated by vehicular traffic. Therefore, these barrier effects on carabid beetles should be considered when planning and implementing road construction and maintenance in forests.     

Tracking predator density dependence and subterranean predation by carabid larvae on slugs using monoclonal antibodies

1. Subterranean carabid larvae are more numerous than surface‐active adults, yet very little is known about their ecological significance, dietary preferences or ability to regulate populations of prey species, particularly pests. Part of the reason for this is that predator–prey interactions beneath the soil are almost impossible to observe. 2. Extensive field studies have shown that adult Pterostichus melanarius (Illiger) can affect the temporal and spatial dynamics of their slug prey. However, if larvae too are feeding on slugs, this could radically affect overall predator–prey dynamics. 3. We tested the hypotheses that P. melanarius larvae would kill and consume two slug species, Deroceras reticulatum Müller and Arion intermedius Normand, under laboratory and semi‐field conditions, and that there would be no significant difference in rates of predation on these slug species. 4. A new monoclonal antibody was developed that was capable of detecting the presence of slug proteins in the guts of P. melanarius larvae. 5. Pterostichus melanarius larvae killed both A. intermedius and D. reticulatum in the laboratory, feeding to a greater extent, and growing more rapidly, on the latter. The larvae were equally effective at reducing numbers of both slug species in a crop of wheat grown in semi‐field mini plots, but predation was affected by density‐dependent intra‐specific competition amongst the beetle larvae. 6. Future modelling of the dynamic interactions between carabids and slugs will need to take into account predation by larvae.     

The influence of weed-cover on the mortality imposed on artificial prey by predatory ground beetles in cereal fields

Summary Pitfall trapping was carried out in a field of winter wheat in the Vale of York to determine the levels of abundance of adult carabid and staphylinid beetles, (which formed the bulk of the natural predator complex) in the field. A point quadrat survey was carried out at the same time to assess the vegetation cover round each trap. Predation pressure by the beetles in the field was monitored using fruit fly (Drosophila) pupae as artificial prey. These artificial prey were attached to small cards inserted in the field adjacent to the pitfall traps.The numbers of beetles caught were found to be directly related to the frequency and density of Poa annua L. (annual meadow grass) the only abundant non-crop plant present at the time. More beetles were caught in areas where Poa was abundant than where Poa was scarce. This was probably related to the more suitable environmental factors offered by these of the wheat itself was found to be unrelated to captures.The number of fruit fly pupae taken was shown to be related to the numbers of carabid and staphylinid beetles present, and also to the type of vegetation occuring around each card.The implication of these results for the effects of predation on potential pest-outbreaks in the crop are discussed.     

Responses of Acilius sulcatus (Coleoptera: Dytiscidae) to chemical cues from perch (Perca fluviatilis)

In this study we tested the hypothesis that the presence of chemical stimuli from a hungry predator would initiate anti-predator responses, while stimuli from a satiated predator would not. We used chemical stimuli released from starved perch (Perca fluviatilis) and from satiated perch (predator). As prey we used adult Acilius sulcatus (Coleoptera: Dytiscidae). The reaction of the beetles to different predator conditions was tested during daytime. We also tested the reaction to starved perch during the night. A. sulcatus activity decreased when it was exposed to stimuli released from starved perch during daytime when visibility was poor, due to the presence of artificial vegetation. There was, however, no reaction to satiated perch under the same experimental conditions. These results indicate that A. sulcatus can discriminate between chemical cues from hungry and satiated fish predators. When visibility was good and the concentration of chemical cues was constant, the beetles did not react to starved perch in the daytime, but their activity decreased at night in response to stimuli released from starved perch. Visual as well as chemical cues seem to be important for detecting a potential predator. When visibility is good, beetles seem to rely on visual stimuli, while in darkness they seem to use chemical stimuli to detect the presence of predators. Received: 4 October 1996 / Accepted: 7 February 1997     

Foraging in the ant Camponotus mus: nectar-intake rate and crop filling depend on colony starvation

The effects of colony starvation on the dynamics of nectar collection were studied in individual workers of the ant Camponotus mus. A laboratory colony was first deprived of carbohydrates for 15days, and thereafter fed daily ad libitum with diluted honey until satiation. During these two successive experimental phases, the probability of feeding, crop filling and fluid-intake rates were recorded daily for individual foragers collecting a 10% (w/w) sucrose solution. The feeding responses of individuals varied with the nutritional state of the colony. When the colony was deprived of sugar, acceptance of the sucrose solution was higher than under satiation. Feeding time increased with increasing starvation. During deprivation workers fed nearly continuously on the solution, whereas a number of feeding interruptions occurred under satiation. Crop filling also increased with increasing starvation, and showed a marked decrease when the colony was satiated. Fluid-intake rate during the deprivation phase was roughly twice that during the satiation phase. This matched well with the difference in sucking frequency recorded during ingestion in satiated and starved workers, which was also higher during starvation. Results indicate that the responsiveness of foragers, determined by the nutritional state of the colony, influenced both foraging decisions and the dynamics of fluid intake.     

Feeding responses of carabid beetles to dimethoate-contaminated prey

Abstract 1 The feeding responses of Pterostichus madidus Fab., P. melanarius Illiger and Nebria brevicollis Fab. (Coleoptera: Carabidae) to dimethoate‐contaminated prey were investigated in ‘no‐choice’ and ‘choice’ feeding tests. 2 In the no‐choice tests, starved beetles were presented with aphid prey treated with four concentrations of dimethoate. In the choice tests, treated and untreated prey were presented together and the feeding preferences of the starved beetles observed. 3 No avoidance or rejection behaviour was seen in any of the carabids in either of the tests, i.e. no discrimination of the treated and untreated prey was observed. 4 Sufficient dimethoate was consumed with the aphid prey to cause significant mortality levels in the carabids. 5 The concentrations of dimethoate used in these experiments are comparable to field exposure, so carabids feeding in treated fields and field margins could potentially suffer lethal effects via the indirect exposure route of consuming contaminated prey.