The responses of polyphagous predators to prey spatial heterogeneity: aggregation by carabid and staphylinid beetles to their cereal aphid prey

Abstract. 1. Discrete patches of aphids were artificially created in winter wheat fields in 1982 and in 1983 using field cages.
2. Aggregation by polyphagous predators at these patches following cage removal was measured using pitfall traps in patch and control areas.
3. Several species of polyphagous beetle (Carabidae and Staphylinidae) aggregated in these patches while other species did not.
4. The consequences of aggregation by polyphagous predators to a single prey species are discussed.     

Low-vacuum electron microscopy of carabid chemoreceptors: a new tool for the identification of live and valuable museum specimens

High-vacuum scanning electron microscopy, following coating of specimens with gold, produces high quality images that have proved invaluable for the study of insect sensilla. Unfortunately, the technique is essentially destructive, and cannot be used on live or valuable museum specimens. In particular, high-vacuum scanning usually causes the collapse of the tips of the palps, interfering with any examination of the sensilla in this area. A new low-vacuum technique is described that avoids these problems. Insect cuticle does not need to be coated with gold, thus avoiding damage to important specimens. Examples are given of scans of the palp tips of live carabid beetles, anaesthetised with CO₂. It was shown that the technique could consistently display these tips in their natural convex state. In all, four types of sensilla were identified by low-vacuum scans of the maxillary palps, and four further types on the terminal segment of the antennae, plus glandular openings. The antennae revealed a type of sensilla that has not previously been described on carabids. These sensilla showed clear structural differences between the two species studied, Pterostichus melanarius Illiger and P. niger Schaller (Coleoptera: Carabidae), and can be used as a diagnostic character for both fresh and dried specimens. The low-vacuum technique can be recommended for examining valuable type specimens without risk of damage.     

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.     

Prey choice by carabid beetles feeding on an earthworm community analysed using species‐ and lineage‐specific PCR primers

The carabid beetle Pterostichus melanarius is a major natural enemy of pests, such as aphids and slugs in agricultural systems. Earthworms are a dominant non‐pest component of the diet of P. melanarius which help sustain the beetles during periods when the pest population is low or absent. In this study we wanted to test whether this predator exercises prey choice among different earthworm species or ecological groups. High levels of genetic diversity within morphological species of earthworm necessitated the development of primers that were specific not just to species but lineages and sub‐lineages within species as well. Gut samples from beetles were analysed using multiplex‐PCR and fluorescent‐labelled primers. Calibratory feeding trials were undertaken to calculate median detection times for prey DNA following ingestion. Extensive testing demonstrated that the primers were species‐specific, that detection periods were negatively related to amplicon size and that meal size had a highly significant effect on detection periods. Monte Carlo simulations showed that, in general, worms were being predated in proportion to their densities in the field with little evidence of prey choice, other than probable avoidance of the larger, deep‐living species. There was no evidence that epigeic species were being taken preferentially in comparison with endogeic species. There was also no evidence that defensive secretions by Allolobophora chlorotica reduced predation pressure on this species by P. melanarius. We concluded that any management system that increases earthworm densities generally, regardless of component species, is likely to be optimal for increasing numbers of this beneficial beetle predator.     

The use of tracers to estimate the exposure of beneficial insects to direct pesticide spraying in cereals

A fluorescent spray tracer was used to measure the profile of spray deposits through winter-wheat crop canopies and upon beneficial insects, placed in fixed positions which approximated to their natural distributions. Tracer deposition profiles were recorded on a series of occasions between G.S. 47 to G.S. 90. Deposition levels declined through the canopy on each occasion but the proportion of the spray reaching the ground varied between growth stages. Deposition rates upon insects followed the same trends but insects on the aerial parts of the plant received a higher volume of tracer per unit area than the part of the crop that they were placed upon. Insects placed on the ground received a dose equivalent to their ground area. For a given distribution of insects, these data permit direct spray exposure to be estimated. The general application of the method to a wide range of crops, to the calculation of risk indices, to the standardisation of field tests and to the development of selective spray placement methods are discussed.     

Interference competition,not predation,explains the negative association between wood ants (Formica rufa) and abundance of ground beetles (Coleoptera: Carabidae)

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Determining numbers of active carabid beetles per unit area from pitfall-trap data

Movement of Pterostichus melanarius (Illiger) (Coleoptera: Carabidae) was studied in the laboratory, and a simulation model developed. The model was calibrated and validated using mark-release-recapture within a grid of pitfall traps in a commercial raspberry field. Rate of movement increased linearly with both temperature and starvation. The temperature threshold for movement, pooling all starvation levels was 3.8 °C ± 0.78 (s.e.). When beetles were released at various distances from a trap in the simulation model, the probability of capture during one day was about 0.13 at a release distance of 10 cm and fell rapidly with distance and temperature. At 5, 10, 15, and 20 °C, the maximum radius sampled by a trap in one day was about 8, 17, 24, and 29 m, respectively. Density was linearly related to catch, but the slope of the line and the variance of the density estimate increased rapidly at lower temperatures. Equations for determining density per m² (±s.d.) were developed from the model, incorporating temperature, number of beetles caught per trap, and number of traps.     

Implications for conservation biocontrol of spatio‐temporal relationships between carabid beetles and coleopterous pests in winter oilseed rape

1 The spatio‐temporal distributions of predatory carabid beetles (Coleoptera: Carabidae) and their potential prey, the larvae of three coleopterous pests, Meligethes aeneus (Fabricius) and Ceutorhynchus spp. [Ceutorhynchus pallidactylus (Marsham), the cabbage stem weevil, and Ceutorhynchus assimilis (Paykull), the cabbage seed weevil], were studied within a crop of winter oilseed rape. The distributions of Collembola were recorded as potential alternative prey. Insect distributions were analysed and compared using Spatial Analysis by Distance Indices. 2 Mature larvae of the pests dropped from the crop canopy to the soil for pupation in temporal succession from May to early July. Their distributions within the crop were irregular and differed with species. 3 Adults of seven species or genera of carabid were abundant and active within the crop during May and June: Nebria brevicollis (Fabricius), Anchomenus dorsalis (Pontoppidan), Loricera pilicornis (Fabricius), Amara similata (Gyllenhal), Asaphidion spp., Pterostichus madidus (Fabricius) and Pterostichus melanarius (Illiger). 4 During May, N. brevicollis was spatially associated with peak numbers of M. aeneus larvae and with Collembola. Anchomenus dorsalis was spatially associated with Ceutorhynchus spp. larvae during two peaks in the abundance of the latter in early and late June. Nebria brevicollis and A. dorsalis coincided in both time and space with larvae of the three coleopterous pests when they were most vulnerable to predation by epigeal predators and are therefore good candidates for conservation biocontrol. 5 The importance of carabid beeetles in the natural enemy complex in winter oilseed rape and their potential for biocontrol of spring and summer pests are discussed in relation to husbandry practices for the crop and its adjacent areas which could be manipulated to promote carabid survival for integrated pest management.     

Modelling the dynamic spatio-temporal response of predators to transient prey patches in the field

The spatio-temporal dynamics of two aphid species ( Metopolophium dirhodum and Sitobion avenae ) and a generalist predator ( Pterostichus melanarius ) were observed in a field-scale study using a grid of 256 sampling locations with a 12-m spacing. Using Spatial Analysis by Distance Indices we demonstrate that populations show ephemeral spatial pattern at the field scale. We observed a positive, lagged beetle response to this aphid pattern; conversely, the aphids displayed a negative, lagged response to beetle spatial pattern. Examination of the local structure of the spatio-temporal dynamics revealed a strong response by the beetle population to aphid patches. The temporal structure of spatial associations between the species shows a strong correspondence with those from a conceptual model of predator–prey spatial interaction. The spatially coupled dynamics were sufficiently strong for the predator to have a negative effect on the intrinsic rate of increase of their prey.     

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.     

Locomotory activity of a visually hunting carabid beetle in response to non-visual prey stimuli

In the predatory beetle Notiophilus biguttatus F. pursuit and attack of prey are induced by visual stimuli associated with prey movement. The study presented in this paper shows that locomotory activity in the predator is affected by prey (springtails — Collembola) previously occupying the relevant substratum. This implies that in prey detection non-visual stimuli are also used. The following effects on locomotion were observed as responses to the cues left by prey: a substantial increase in the frequency of stops and runs and a decrease in the average duration of runs. The adaptive significance of these effects is discussed.Analysis of the experiments includes an evaluation of the statistical powers of the tests used. The estimated powers were used to interpret nonsignificant results.

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Zusammenfassung Die Verfolgung und der Angriff auf eine Beute folgt beim Laufkäfer Notiophilus biguttatus allein auf die visuelle Wahrnehmung bewegender Beutetiere. In der vorliegenden Arbeit wird gezeigt, das die Aktivität des Käfers sich ändert, wenn vorher in seiner Umgebung Beutetiere (Collembolen) anwesend waren. Diese Änderung bedeutet, dass beim Beutefang auch chemische Spuren benützt werden. Dabei wurden folgende Effekte auf die Lokomotion des Käfers festgestelt: eine beträchtliche Zunahme in der Frequenz von Stehenbleiben und Laufen und eine Abnahme der durchschnittlichen Zeitdauer der Läufe. Die adaptive Bedeutung dieser Effekte wird besprochen.Die Experimente wurden einer statistischen power-Analyse unterworfen. Die power-Analyse wurde eingesetzt zur Deutung nichtsignifikanter Ergebnisse und zum Vergleich der Empfindlichkeit verschiedener Aspekte der Lokomotion für den betreffende Reiz.

     

The significance of facultative scavenging in generalist predator nutrition: detecting decayed prey in the guts of predators using PCR

Gut-content analyses using molecular techniques are an effective approach to quantifying predator-prey interactions. Predation is often assumed but scavenging is an equally likely route by which animal DNA enters the gut of a predator/scavenger. We used PCR (polymerase chain reaction) to detect scavenged material in predator gut homogenates. The rates at which DNA in decaying slugs (Mollusca: Pulmonata) and aphids (Homoptera: Aphididae) became undetectable were estimated. The detectability of DNA from both carrion types in the guts of the generalist predator Pterostichus melanarius (Coleoptera: Carabidae) was then determined. The effects of carrion age and weight, as well as beetle sex, on detection periods, were quantified. Laboratory trials measured prey preference of beetles between live and decaying prey. Further experiments measured, for the first time, feeding by P. melanarius on dead slugs and aphids directly in the field. In both field and laboratory, P. melanarius preferentially fed on dead prey if available, but preference changed as the prey became increasingly decayed. Disappearance rates for slug carrion in wheat fields and grasslands were estimated and P. melanarius was identified as the main scavenger. Comparison of the retention time for dead slugs in the field, with the detection period for decaying slug material in the guts of the predators, showed that PCR-based techniques are not able to distinguish between predated and scavenged food items. This could potentially lead to overestimation of the impact of predation on slugs (and other prey) by carabids. Possible implications of facultative scavenging by invertebrate predators for biocontrol and food-web research are discussed.     

The effects of forestry on carabid beetles (Coleoptera: Carabidae) in boreal forests

As compared to natural forests, managed boreal forests are younger, more homogeneous in terms of tree age and species composition, and consist of smaller fragments. Here we examine the effects of such characteristics caused by forestry on carabid beetles (Coleoptera, Carabidae) in the boreal region. The main results are the following. (1) Fragmentation of forests and the size of a fragment appear not to be crucial for the survival of the majority of forest carabids, as they tend to be distributed over various successional stages, but species requiring old-growth habitats suffer. (2) For carabids there appear to be no or very few edge specialist species, and forest-open land edges appear to be effective barriers for species associated with forest or open habitat. However, generalist species easily cross the edge, and edges of forest fragments may be invaded by species from the surrounding open habitat. (3) Habitat change following clear-cutting dramatically changes the composition of carabid assemblages: species restricted to mature forests disappear and open-habitat species invade, while habitat generalists survive at least in the short term. Carabid diversity can probably best be maintained if forest management mimics natural processes, maintains natural structures and includes the natural composition of vegetation and other structural elements (such as dead wood) within the stands, provided that these forest features can be maintained and recreated through forest management practices. At a larger scale, the whole spectrum of forest types and ages (especially old-growth forests), and different successional processes (especially fire) should be maintained. These require the development and use of innovative logging methods, and the planning, implementation, and assessment of landscape-scale ecological management strategies.     

Predation on the slug Deroceras reticulatum by the carabid beetles Pterostichus madidus and Nebria brevicollis in the presence of alternative prey

1 Slugs are important pests in many agricultural crops and potential biological control agents are being studied as an alternative to molluscicides. Carabid beetles may be able to reduce slug populations, but their role as control agents may be influenced by the presence of alternative prey. 2 Attacks on the pest slug Deroceras reticulatum (Müller) by the carabid beetles Pterostichus madidus (Fabricius) and Nebria brevicollis (Fabricius) were investigated in the presence of alternative prey (earthworms and Calliphora fly larvae). Consumption of slug eggs and aphids was also investigated. 3 All five prey types were consumed to varying degrees during the experiments. Both beetle species showed a significant preference for Calliphora larvae over slugs. Pterostichus madidus showed a significant preference for earthworms over slugs. No preference was shown between earthworms or Calliphora larvae by P. madidus females or N. brevicollis. However, P. madidus males showed a significant preference for Calliphora larvae over worms. Pterostichus madidus showed no preference between slug eggs and aphids; N. brevicollis showed a significant preference for aphids over slug eggs. 4 The results from this study indicate that generalist beetles will often attack other prey in preferences to adult slugs. Slugs may not be preferred because of their mucus. Other prey items occur frequently in arable soils and generalist carabids may ignore slugs altogether and may only feed on them when slug density is high or other prey are unavailable.     

Consumption by carabid beetles of three cereal aphid species relative to other prey types

The cereal aphidRhopalosiphum padi has previously been found to be a low quality prey for a range of generalist arthropod predators. The aim of this study was to reveal, using food consumption experiments whether this applies to other cereal aphids. The question of whether predator feeding capacity increased when several aphid species were offered relative to a single aphid species was also addressed by measuring food consumption on a mixed aphid diet relative to single aphid diets. Food consumption by five carabid beetles of the three cereal aphid speciesRhopalosiphum padi, Sitobion avenae andMetopolophium dirhodum was determined relative to fruit fliesDrosophila melanogaster and the collembolanIsotoma anglicana. Feeding rate was measured as food consumption over 24 hour both for previously satiated and beetles starved for 7 days. Generally the largest aphid consumption was ofM. dirhodum and the lowest ofR. padi, withS. avenae in between. The mixed aphid consumption experiments did not reveal a higher feeding rate on mixed aphid diets relative to single aphid diets. The results indicate low preference forR. padi andS. avenae.     

Predation by the carabid beetles Pterostichus madidus and Nebria brevicollis is affected by size and condition of the prey slug Deroceras reticulatum

1 Slugs are important pests in many agricultural crops but molluscicides commonly used to control slugs affect non‐target organisms. Encouraging biological control may help to reduce molluscicide use, but the efficiency of potential natural enemies needs to be investigated. 2 Serological tests have shown that certain carabid species consume slugs. These techniques, however, do not distinguish between scavenging and true predation, nor do they provide information on the size or other characteristics of the prey consumed. The study reported here was undertaken to establish whether scavenging of dead slugs might be an important factor contributing to positive serological test results. 3 Both Pterostichus madidus (Fabricius) and Nebria brevicollis (Fabricius) consumed Deroceras reticulatum (Müller) under laboratory conditions. Dead slugs were scavenged in preference to injured or healthy slugs. 4 Only small, live slugs (< 0.11 g) were killed by both beetle species, which may, therefore, be incapable of killing larger slugs. 5 These generalist beetle species appeared unable to overcome the defence mucus produced by slugs. The data suggest that positive serological results from field collected beetles may reflect scavenging rather than predation on live or injured slugs.     

Differential antigen decay rates during digestion of molluscan prey by carabid predators

Carabid beetles (Coleoptera: Carabidae) were fed upon slugs (Mollusca: Pulmonata) in the laboratory, and their crop contents analysed for mollusc remains, using an enzyme-linked immunosorbent assay (ELISA) with an anti-Arion ater (L.) haemolymph antiserum. Crop weight loss and antigenic recognition of prey proteins were examined as separate variables in a series of validatory experiments. Two model predators,Abax parallelepipedus Piller and Mitterpacher andPterostichus madidus F., were fed upon two species of pest slugs,Deroceras reticulatum (Müller) andArion hortensis Férussac. The fitting of regression equations to the transformed antigenic response data allowed the ‘half-life’ and detection period to be calculated for each predator-prey combination. Following a one hour feeding interval, the half-life of the antigenic response toD. reticulatum remains was almost twice as long inP. madidus as that inA. parallelepipedus, and the detection period more than 2.5 times as long. However, covariant analysis showed that there was a significant difference between predator species in the rate at which detectability declined, but not in the rate of crop weight loss. WhenA. parallelepipedus was allowed to feed uponA. hortensis for eight hours, prey remains were still detectable at the end of the experiment, 13 days after feeding. Calibration of the differential rates of antigen decay and crop weight loss could potentially be used to calculate the size of the original meal, but only if prey species, and the time since feeding, can be determined. Potential solutions to these problems are discussed.