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.     

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.     

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.     

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.     

Detection of secondary predation by PCR analyses of the gut contents of invertebrate generalist predators

Predation by generalist predators is difficult to study in the field because of the complex effects of positive and negative interactions within and between predator species and guilds. Predation can be monitored by molecular means, through identification of prey DNA within predators. However, polymerase chain reaction (PCR) amplification of prey DNA from predators cannot discriminate between primary and secondary predation (hyperpredation), in which one predator feeds on another that has recently eaten the target prey. Here we quantify, for the first time, the potential error caused by detection of prey DNA following secondary predation, using an aphid-spider-carabid model. First, the aphid Sitobion avenae was fed to the spider Tenuiphantes tenuis and the carabid Pterostichus melanarius, and the postconsumption detection periods, for prey DNA within predators, were calculated. Aphids were then fed to spiders and the spiders to carabids. Aphid DNA was detected in the predators using primers that amplified 245- and 110-bp fragments of the mitochondrial cytochrome oxidase I gene. Fragment size and predator sex had no significant effect on detection periods. Secondary predation could be detected for up to 8 h, when carabids fed on spiders immediately after the latter had consumed aphids. Beetles tested positive up to 4 h after eating spiders that had digested their aphid prey for 4 h. Clearly, the extreme sensitivity of PCR makes detection of secondary predation more likely, and the only reliable answer in future may be to use PCR to identify, in parallel, instances of intraguild predation.     

Noninvasive molecular methods to identify live scarab larvae: an example of sympatric pest and nonpest species in New Zealand

Despite the negative impact that many scarab larvae have on agro-ecosystems, very little attention has been paid to their taxonomy. Their often extremely similar morphological characteristics have probably contributed to this impediment, which has also meant that they are very difficult to identify in the field. Molecular methods can overcome this challenge and are particularly useful for the identification of larvae to enable management of pest species occurring sympatrically with nonpest species. However, the invasive collection of DNA samples for such molecular methods is not compatible with subsequent behavioural, developmental or fitness studies. Two noninvasive DNA sampling and DNA analysis methods suitable for the identification of larvae from closely related scarab species were developed here. Using the frass and larval exuviae as sources of DNA, field-collected larvae of Costelytra zealandica (White) and Costelytra brunneum (Broun) (Scarabaeidae: Melolonthinae) were identified by multiplex PCR based on the difference in size of the resulting PCR products. This study also showed that small quantities of frass can be used reliably even 7 days after excretion. This stability of the DNA is of major importance in ecological studies where timeframes rarely allow daily monitoring. The approach developed here is readily transferable to the study of any holometabolous insect species for which morphological identification of larval stages is difficult.     

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.     

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.     

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.     

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

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Additive effects of predator diversity on pest control caused by few interactions among predator species

1. Studies of the impact of predator diversity on biological pest control have shown idiosyncratic results. This is often assumed to be as a result of differences among systems in the importance of predator–predator interactions such as facilitation and intraguild predation. The frequency of such interactions may be altered by prey availability and structural complexity. A direct assessment of interactions among predators is needed for a better understanding of the mechanisms affecting prey abundance by complex predator communities. 2. In a field cage experiment, the effect of increased predator diversity (single species vs. three‐species assemblage) and the presence of weeds (providing structural complexity) on the biological control of cereal aphids were tested and the mechanisms involved were investigated using molecular gut content analysis. 3. The impact of the three‐predator species assemblages of aphid populations was found to be similar to those of the single‐predator species treatments, and the presence or absence of weeds did not alter the patterns observed. This suggests that both predator facilitation and intraguild predation were absent or weak in this system, or that these interactions had counteracting effects on prey suppression. Molecular gut content analysis of predators provided little evidence for the latter hypothesis: predator facilitation was not detected and intraguild predation occurred at a low frequency. 4. The present study suggests additive effects of predators and, therefore, that predator diversity per se neither strengthens nor weakens the biological control of aphids in this system.     

农田生态系统植物多样性对害虫种群数量的影响

着重分析植物多样性影响害虫发生为害及种群数量的生态学机制,综合评述了关于这种机制的两种主要假说,即天敌假说和资源集中假说．同时总结了植物多样性增大和减少对害虫控制的有利和不利因素．研究表明农田生态系统中植物多样性的增大在多数情况下能导致某些害虫种群数量的下降,但是目前很难就不同栖境中所有类型的害虫形成一般性的结论     

Observation of epigeic predators and predation on artificial prey in a cereal field

Predation activity of different epigeic predators on artificially supplied prey was studied in a cereal field during several 24 h periods from spring to early summer 1993. Carabids were the most frequently observed predators. Among themPoecilus cupreus was the most voracious species during day-time, andAgonum muelleri during night-time. Predation rates were highest between 10.00 and 18.00 h and lowest between 02.00 and 10.00 h. Linear regressions showed significantly positive relations between temperature and predation rates, and between predation and activity density of epigeic predators. The latter relation was stronger when only activity density of predators which were observed to consume baits was included. Quadrat sampling and pitfall sampling showed that with the former only a small proportion of bait-consuming predators were caught whereas with the latter method this proportion was high.     

Preliminary experiments on the olfactory responses of Pterostichus melanarius Illiger (Coleoptera: Carabidae) to intact plants

1 In two successive years, the olfactory responses of Pterostichus melanarius Illiger (Coleoptera: Carabidae) to intact cabbage and white clover plants were studied, using a four‐arm olfactometer. 2 The first set of experiments tested the response to the odours of cabbage, white clover and the two plants grown together against potting compost; in the second and third sets of experiments, white clover and cabbage were tested, respectively. 3 The results indicate that female P. melanarius individuals are arrested by the combined odours of cabbage and white clover, spending significantly more time in the cabbage with white clover odour field than in either the cabbage, white clover or control odour fields. 4 Similar results were not found for males. 5 Cabbage plants alone did not elicit significant responses in either sex. 6 White clover alone did not elicit significant responses by females and the responses of males were not uniform in the two successive years.     

高等植物细胞质雄性不育分子机理的研究进展

从线粒体DNA、叶绿体DNA和线粒体质粒DNA方面较详细地阐述了高等植物细胞质雄性不育的分子机理及最新进展;探讨了细胞核DNA和细胞质DNA之间的相互关系。     

Habitat effects on second-order predation of the seed predator Harpalus rufipes and implications for weed seedbank management

Seed predators provide a valuable ecosystem service to farmers by reducing densities of weed seeds, and, in turn, densities of weed seedlings they must manage. The predominant invertebrate weed seed predator in Maine, USA, agroecosystems is the carabid beetle Harpalus rufipes DeGeer. Pitfall trapping has shown that H. rufipes prefers sites with vegetative cover to fallow sites, preference speculated to be driven by predator avoidance behavior. To test this hypothesis, ‘second-order predation assays’ were developed, in which live H. rufipes prey were presented to second-order predators. Field experiments were conducted to determine foremost if H. rufipes was subject to second-order predation, and secondly, whether (a) vegetative cover affords H. rufipes protection from second-order predators, and (b) high rates of second-order predation correspond with decreased invertebrate seed predation rates. Two 72-h experiments were conducted (mid August and September 2012) at crop and non-crop sites across a 28 ha diversified farm in Stillwater, ME, USA.Second-order predation was 2.8% per day. Based on images from motion-sensing cameras, H. rufipes’ predators included birds and small mammals. Neither a relationship between second-order predation and vegetative treatment, nor an empirical relationship between second-order predation and invertebrate seed predation were detected. However, a simulation model predicted that 2.8% per day second-order predation could increase the number of seeds entering the seedbank by more than 17% annually. Additionally, complex habitats supported higher rates of second-order predation than did simple habitats.     

A new taxonomic status for Iberoformica (Hymenoptera,Formicidae) based on the use of molecular markers

The subgeneric subdivision of the genus Formica is still open. In this article, we make a phylogenetic study on several species of the genus Formica and of its closely related genera, Polyergus and Proformica, using sequences of nuclear satellite DNA (stDNA) and the mitochondrial rrnL as molecular markers. Our goal was to shed light on their phylogenetic relationships and particularly on the systematic position of F. subrufa. This species was first included in the subgenus Serviformica, but afterwards a new subgenus (Iberoformica) was established to include only this species. The results show that a stDNA family previously reported in Formica species, with a repetitive unit 129 bp long, is also found in Polyergus rufescens and P. samurai but not in Proformica longiseta. This is the first case of presence of a stDNA family in two different ant genera. In F. subrufa, this stDNA is very divergent relative to those isolated in the remaining Formica species and in the genus Polyergus. The Bayesian analysis of mitochondrial rrnL sequences shows three highly supported groups: F. subrufa, the remaining Formica species studied, and the genus Polyergus, suggesting that parasites (Polyergus species) and hosts (Formica species) are closely related but not sibling species. The combined analysis of nuclear stDNA sequences and mitochondrial rrnL showed their phylogenetic congruence despite their distinct evolutionary dynamics. This analysis did not discriminate between the remaining Formica species that were not grouped according to the subgeneric classification. According to these results, it can no longer be assumed that F. subrufa belongs to the subgenus Serviformica or of the fusca species group. This differentiation was also supported by previous studies based on the morphological characters, molecular and cytogenetic data. Therefore, taking into consideration these arguments and others explained in detail in this article, we propose that the taxon Iberoformica, formerly synonymized subgenus, be raised to a genus status. This genus would be monotypic and only composed, up to the moment, by Iberoformica subrufa (= F. subrufa Roger, 1859 ).     

Rates of consumption of cereal aphids by some polyphagous predators in the laboratory

The consumption rates of nine species of Carabidae and five species of Staphylinidae which feed on the aphid Sitobion avenae (F.) were studied under conditions of surplus food in the laboratory. The mean daily consumption of aphids was between 2.5% and 45.6% of the predator species' mean body weight; the lowest values were for large Carabidae and the highest for small Staphylinidae. Separate significant positive relationships were found between the mean total weight of aphids consumed per day, and the mean weight of the adult of the carabid and staphylinid species at each temperature. A re-evaluation of published data also showed this. The slopes of the regression lines were steeper for the Staphylinidae than for the Carabidae and possible reasons are given. The regression equations could be used in simulation models to determine the maximum possible effects of these beetles on aphid populations.

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Résumé La consommation de Sitobion avenae F. par 9 espèces de Carabidae et de 5 de Staphylinidae a été examinée au laboratoire en condition de surapprovisionnement. Les expériences ont été répétées dans 4 conditions de températures variables caractérisées par des moyennes de 12°3, 16°6, 20°6 et 23°6. Les poids moyens de pucerons consommés variaient entre 2,5% et 45,6% du poids des prédateurs; les valeurs les plus faibles correspondant aux gros carabes et les plus élévées aux petits staphylins.Les différences entre sexes étaient vraisemblablement dues au dimorphisme pondéral. Une relation significative et positive entre le poids moyen de pucerons consommés et le poids moyen du coléoptère a été mise en évidence tant pour les carabes que les staphylins à toutes les températures (Fig. 1, Tableau 2). Cette relation ressort aussi d'une réévaluation des données publiées. L'utilisation de formules de régression dans des modèles de simulation permettra de prédire les effets des carabes et des staphylins sur les populations de pucerons dans ces conditions optimales.