Temperature and prey capture: opposite relationships in two predator taxa

Abstract 1. All other things equal, predator capture rates are expected to depend on encounter rate with prey, prey escape capability (including prey defences), and on predator agility. Ectotherm predators and their prey both respond to increasing temperature by increased activity, i.e. predators increase their search area and prey may enhance their escape capability. This means that, as temperature changes, the ability of a predator to catch prey will decrease, increase, or remain unchanged depending on the relative effect of temperature on predator and prey. Their responses may further be differentially moulded by light conditions depending on whether the predator is diurnally or nocturnally active. It was hypothesised that flying Diptera are vulnerable to carabid beetles only at low temperatures and over the full temperature range for spiders because carabids, in contrast to spiders, are not built to catch swiftly moving prey. 2. The first experiment examined the spontaneous locomotor activity of the predators and of fruit flies at different temperatures (5, 10, 15, 20, 25, and 30 °C) and light conditions (light, dark). A second experiment examined the effect of temperature and light on the predation rate of two carabid beetles (Pterostichus versicolor and Calathus fuscipes) and two spiders (Clubiona phragmitis and Pardosa prativaga) using fruit flies (Drosophila melanogaster) as prey. 3. All four predators and the fruit fly increased their locomotory activity at higher temperatures. Activity of the carabid beetles peaked at intermediate temperatures; spiders and fruit flies were most active at the highest temperatures. Predation rate of the spiders increased with temperature whereas the beetles caught flies only at low temperatures (5 and 10 °C). 4. Diurnal variation in temperature may bring different prey groups within the set of potential prey at different times of the day or at different seasons. The ability of many carabid beetles to forage at low temperatures may have nutritional benefits and increases the diversity of interactions in terrestrial food webs.     

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.     

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.

     

A chance field observation and pilor laboratory studies of predation of the New Zealand flatworm by the larvae and adults of carabid and staphylinid beetles

A chance finding in an Edinburgh allotment was made of a stained carabid beetle larva that had eaten Artioposthiu triungulata previously stained red and released into the allotment during a dispersal experiment. In a pilot laboratory study the flatworm was eaten by larval and adult carabid and staphylinid beetles whereas seven other carnivorous species from five arthropod families did not. In another feeding experiment a carabid and a staphylinid larva each ate several specimens of the flatworm and gained tissue weight over 14 days. The conclusion drawn from these preliminary studies and the field observation is that certain carabid and staphylinid beetles are potential predators of A. triangulata.     

Staphylinidae and Carabidae overwintering in wheat and sown wildflower areas of different age

Species richness and abundance of staphylinid and carabid beetles overwintering in winter wheat fields and 1- to 3-year-old wildflower areas were investigated during 2000/2001 on 16 study sites in Switzerland. Abundance and species richness of overwintering staphylinids significantly increased with successional age of the wildflower areas and were always higher in older wildflower areas than in winter wheat. A similar but less distinct pattern was observed for the abundance and species richness of carabid beetles. The influence of habitat parameters (vegetation cover, fine sand content, organic matter, pH, soil pore volume, surrounding landscape structure, habitat area) on the staphylinid and carabid assemblages based on the number of individuals per species and site was analysed using canonical correspondence analysis. Vegetation cover was the most significant parameter significantly characterizing both staphylinid and carabid assemblages. The amount of vegetation cover explained 15.7% of the variance, fine sand content accounted for 13.3% and surrounding landscape structure for 10.9% of the variance in the staphylinid assemblage. In the carabid assemblage, vegetation cover was the only significant factor, explaining 24.7% of the variance. This study showed for the first time that the significance of wildflower areas as a reservoir for hibernation for generalist predatory beetles increases with progressing successional age.     

Effects of Bt maize-fed prey on the generalist predator <Emphasis Type="Italic">Poecilus cupreus</Emphasis> L. (Coleoptera: Carabidae)

We investigated the effects of transgenic maize (Zea mays) expressing Bacillus thuringienses toxin (Bt maize) on larval and adult Poecilus cupreus carabid beetles in laboratory studies. In no-choice trials, neonate P. cupreus larvae were fed exclusively with Spodoptera littoralis caterpillars, which had been raised on Bt maize. S. littoralis raised on conventional maize or high quality Calliphora sp. pupae were fed to the beetle larvae in two control treatments. Bt-maize-fed caterpillar prey increased mortality to 100 within 40days. The experiment was repeated with 10-day-old beetle larvae. Bt treatment resulted in fewer pupae than in both controls, and in a higher mortality than in the Calliphora control. S. littoralis was suitable as exclusive prey in no-choice tests, at least for 40days, although prey quality seemed to be low compared to Calliphora pupae. The observed effects are most likely indirect effects due to further reduced nutritional prey quality. However, direct effects cannot be excluded. In the second part of the study, exposure of P. cupreus to Bt intoxicated prey was examined in paired-choice tests. Adult beetles were offered a choice between different prey conditions (frozen and thawed, freshly killed or living), prey types (S. littoralis caterpillars, Calliphorasp. pupae, cereal aphids) and prey treatments (raised on Bt or conventional maize). Living prey was preferred to frozen and dead prey. Caterpillars were only preferred to fly pupae and aphids when living. Prey treatment seemed to be least important for prey selection. The tests showed that P. cupreus ingested caterpillars readily and there was no evidence of them avoiding Bt containing prey, which means exposure in the field could occur. The presented protocols are a first step towards ecological risk assessment for carabid beetles.     

Influence of ground beetles (Carabidae) as biological agent to control of the Mediterranean fruit fly pupae,Ceratitis capitata,in Iranian citrus orchards

Damage by Mediterranean fruit fly, Ceratitis capitata (Wiedemann) (Diptera: Tephritidae) in the northern Iranian citrus orchards has been led to spray by different chemical pesticides. The present study was conducted in the citrus orchard in Mazandaran province of Iran where with the exception of mineral oil treatments for scale insects, had not been sprayed with synthetic insecticides for six years. The purpose of this study was initially to investigate faun and seasonal activity of predaceous ground carabid beetles in the citrus orchards. Additionally, their role as predators of Ceratitis capitata (Wiedemann) pupae was evaluated under laboratory and field conditions. Predation in laboratory was evaluated by no choice feeding experiments while predation under field conditions was evaluated by molecular detection of the pest DNA in the gut contents of field-collected Carabids. A total of 3437 beetles of ten carabid species were collected from a non-chemical treatment citrus orchard. The results of the seasonal density fluctuation indicated increasing population in spring reaching the highest in September, simultaneously with the peak of fruit fly damage, and thereafter decreasing during winter when the pest was overwintering. However, three species, Harpalus rufipes, Broscus karelinii and Brachinus brevicollis were most abundant in the field collection but the results of no-choice experiments in the laboratory indicated H. rufipes, B. karelinii and Pterostichus macer were the most voracious predators on pupae. Results of gut content analysis of the voracious species indicated the positive testing for pest DNA in 79.16 % of H. rufipes, 76.92 % of B. karelinii and 75 % of P. macer of analyzed beetles. The results of the study strongly indicated that in citrus orchards where intensive spraying is eliminated, natural enemies such as Carabid beetles could play an important role in reducing pests such as medfly.     

Predator interference limits fly egg biological control by a guild of ground-active beetles

We examined whether predator interference could prevent effective conservation biological control of Delia spp. flies, important pests of cole crops, by an assemblage of carabid and staphylinid beetles. In laboratory feeding trials we found that the smaller (<1 cm) beetle species common at our site readily ate dipteran eggs, while the most common large carabid species, Pterostichus melanarius, rarely did. However, P. melanarius did eat several of the smaller beetle species. We conducted two field experiments where we manipulated immigration rates of the ground predator guild and then measured predation on fly eggs. Predation rates were consistently higher in cages where predators were added at ambient densities, compared to cages where ground predators were removed. However, in the second field experiment, when we quadrupled predator immigration rates neither beetle activity-density nor predation rate increased. High immigration rate plots had a higher proportion of P. melanarius in the predator community, compared to plots with beetles added at ambient densities, suggesting that P. melanarius was reducing activity-densities of the smaller beetles, perhaps through intraguild predation. Thus, tactics to improve the biological control of Delia spp. by conserving generalist predators, such as providing in- or extra-field refuges, could be thwarted if the primary predators of fly eggs, small carabids and staphylinids, are the targets of intraguild predation by also-conserved larger predators.     

The aggregative numerical response of polyphagous predators to aphids in cereal fields: attraction to what?

We tested in a field experiment two hypotheses for why polyphagous predators aggregate at concentrations of aphids: 1) because they are attracted directly to aphids as prey, or 2) because they are attracted to alternative prey that aggregate around the honeydew produced by aphids. Small plots were established in the field with two experimental treatments, aphid addition and honey spraying, and a watersprayed control, each replicated 10 times. Arthropods were recorded by pitfall traps and sticky traps in each plot. Diptera were significantly more abundant in the honey plots. Of the predators, Agonum dorsale, “All carabids” and Philonthus sp. were most abundant in the honey plots; Tachyporus spp. and carabid and staphylinid larvae were most abundant in the aphid addition plots. It is suggested that these results reflect differences among the predators in their ability to tolerate and utilise aphids as food.     

Plant versus prey resources: Influence on omnivore behavior and herbivore suppression

Conservation biological control tactics, such as beetle banks, that increase habitat complexity generally increase epigeal predator abundance. Habitat complexity also increases alternative food which can attract and sustain predators but may reduce predation of target pests. Our goal was to determine how alternative food from different trophic levels (fly pupae and seeds) affects behavior and biological control efficacy of omnivorous carabid beetles. Seed subsidies increased omnivorous carabid abundance more than pupae by increasing aggregation and reducing emigration. Laboratory experiment demonstrated that both omnivorous carabid species preferred seeds and pupae over cutworms. However, in field cages seeds but not pupae resulted in greater cutworm damage to corn seedlings. Our results indicate that omnivorous carabids have a stronger behavioral response to seeds than prey such that only seeds influence aggregation, emigration, and crop damage. Interestingly, whereas seeds increased omnivorous carabid abundance, pupae had no affect on carnivore abundance. Thus, carabid guild composition is skewed in favor of omnivores when seed density increases. An important finding was that the effect of seeds on behavior, predation, and crop damage was consistent among replicate carabid species suggesting our results pertain to other omnivorous species in resource diverse habitats.Our results provide insight into the mechanisms underlying the unpredictable benefit of conservation biological control tactics that alter habitat complexity.     

The Structure of Ground Beetle Assemblages (Coleoptera: Carabidae) at Fruit Falls of Melastomataceae Trees in a Brazilian Terra Firme Rain Forest1

Although most Carabidae are recognized as predacious, recent studies have shown that some tropical rain forest Carabidae use the small seeds of fallen fruit from some tree species as a key food resource (particularly figs [Moraceae] but also those from other families). Assemblages of carabid beetles (Coleoptera: Carabidae) at fruit falls of the Melastomataceae tree genera, Bellucia, Loreya, and Miconia, were studied in a terra firme rain forest near Manaus (Amazonia, Brazil) from February 1992 to August 1996. Of 5483 specimens of 14 carabid species collected at 53 fruit falls, 98.2 percent of the collected beetles represented the seed-feeding genus Notiobia. Four of the 7 Notiobia species were found to reproduce at Melastomataceae fruit falls. The most abundant species, N. glabrata, was found to be a Melastomataceae seed generalise while N. maxima seemed to prefer the seeds of Bellucia, and N. aulica and Af. umbrifera the seeds of Miconia. The presence of larvae indicated reproduction of N. glabrata at fruit falls of all three tree genera and reproduction of N. maxima at those of Bellucia. Fruit falls of different species and genera of Melastomataceae appeared to occur at different times of die year and not continuously throughout the year. The seed-feeding Notiobia species, which specialize on Melastomataceae, used fruit falls of figs as migratory “stepping stones” during periods when there were no Melastomataceae fruit falls. The comparable suite of fig seed-feeding specialist species of Notiobia did not appear to do the reverse and seemed to avoid Melastomataceae fruit falls. The carabid assemblage at fig fruit falls was more diverse than the assemblage at those of Melastomataceae. Among the Melastomataceae, carabid species richness was highest at Miconia fruit falls and lowest at Loreya fruit falls. The three most common predatory species caught at Ficus fruit falls were also caught at those of Melastomataceae and appeared to reproduce at both fruit fall types. The proportion of the carabid assemblage accounted by predatory species was much lower at Melastomataceae (1.8%) than at fig fruit falls (8%).     

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.     

Wild flower seed predation by Pterostichus madidus (Carabidae: Coleoptera)

The objective of the study was to investigate the ability of Pterostichus madidus, one of the most abundant carabid beetles in the United Kingdom, to consume wildflower seeds. The plant species chosen are commonly included in wildflower seed mixes sown to enhance the biodiversity of arable field margins, and hence any seed predation could potentially reduce successful seedling establishment. The study combined cafeteria style laboratory feeding experiments with predation studies in an established field margin. In laboratory studies the Pterostichus madidus consumed both unimbibed and imbibed seeds of Centaurea scabiosa, Galium verum, Leucanthemum vulgare and Primula veris. Seed consumption was still evident when Drosophila pupae were also supplied, although there was significant preference for the pupae. Placement of seeds in the field showed high levels of seed consumption, although part of this was attributed to slugs. Carabid beetles may have a significant role as wild flower seed predators and thus may affect establishment of re‐seeded flower‐rich meadows.     

Diversity and abundance of the coleopteran fauna from organic and conventional management systems in southern England

1 Studies of the epigeal coleopteran fauna on five pairs of organic and conventional farms were carried out between May and July 1994 in southern England using pitfall trapping. A total of 27 749 individuals and 140 species were identified. Overall, abundance of Coleoptera was greatest on organically managed farms. 2 Ground beetles (Coleoptera: Carabidae) and staphylinid beetles (Col. Staphylinidae) formed 79.7% and 16.7%, respectively, of the total catch. Pterostichus melanarius was the dominant carabid among the 45 species captured and significantly higher numbers were found in organic farms. Tachinus signatus was the most common of 44 staphylinid species, and was significantly more abundant on conventional farms. 3 For carabids, the log‐series α diversity index was higher on conventional farms but was not statistically different from that calculated for organic farms. The α index was identical for staphylinid species from organic and conventional farms. Diversity was significantly higher from conventional farms when data were combined for all of the recorded coleopteran species. 4 From this study, it appears that the main effect of farming practice is to influence the overall abundance and dominance of particular species, and the lower diversity of organic farms is a consequence of the large increase in dominance of a single species, namely P. melanarius.     

Predation potential of the earwig Euborellia annulipes on fruit fly larvae and trophic interactions with the parasitoid Diachasmimorpha longicaudata

The earwig Euborellia annulipes (Lucas) (Dermaptera: Anisolabididae), a generalist predator, has been observed in fruits infested with fruit fly larvae, which are frequently parasitized by parasitoid wasps. Neither the capacity of earwigs to predate on fruit flies nor intraguild interactions between earwigs and fruit fly parasitoids have been investigated. Here, we studied in laboratory conditions the predation on the fruit fly Ceratitis capitata (Wiedemann) (Diptera: Tephritidae) by the earwig E. annulipes, and whether parasitism of fruit fly larvae by the parasitoid wasp Diachasmimorpha longicaudata (Ashmead) (Hymenoptera: Braconidae) influences predation by the earwig. We evaluated the predation capacity, functional response and prey preference of E. annulipes for parasitized and non-parasitized fruit fly larvae in choice and no-choice tests. We found that earwigs prey on second- and third-instar larvae and pupae of C. capitata and consumed larger numbers of second-instar larvae, followed by third-instar larvae and pupae. Females prey on larger numbers of fruit flies than did males, regardless of the prey developmental stage, but both sexes exhibited a type II functional response. Interestingly, males killed but did not consume fruit fly larvae more than did females. In no-choice tests, earwig females consumed equal numbers of parasitized and non-parasitized fruit fly larvae. However, in choice tests, the females avoided feeding on parasitized larvae. Subsequent tests with hexane-washed parasitized and non-parasitized larvae showed that putative chemical markings left on fruit flies by parasitoids did not drive the earwig preference towards non-parasitized larvae. These findings suggest that E. annulipes is a potential biological control agent for C. capitata, and that, because the earwig avoids consuming larvae parasitized by D. longicaudata, a combination of the two natural enemies could have an additive effect on pest mortality.     

Effect of beetle size on predation of cabbage root fly eggs by ground beetles

Laboratory studies were done to determine the numbers of cabbage root fly (Delia radicum L.) (Diptera: Anthomyiidae) eggs eaten by sixty species of ground beetles (Coleoptera: Carabidae) collected from a range of habitats. For 35 species that were between 2.7 mm and 10 mm long, there was a linear relationship between the numbers of eggs eaten and beetle length. For each 1 mm increase in length above 2.7 mm, an additional 18 eggs were eaten/beetle/day. Eight further species within the above size range did not eat any fly eggs, as these species are known to be either phytophagous or to feed on moving prey. Large numbers of eggs were eaten by beetles in 13 of the 25 genera tested. Based on size, maximum numbers of eggs were eaten by 7 species of Agonum Bonelli, 5 species of Amara Bonelli and seven species of Bembidion Latreille. Only one of the 6 species of Pterostichus Bonelli tested, P. strenuus (Panz.), ate the expected numbers of eggs, the other species being too large to feed from such small prey items as cabbage root fly eggs. The numbers of beetles larger than 10 mm that ate eggs was highly variable and so 10 mm was considered the upper size limit of carabid predators of cabbage root fly eggs.     

Subsidy from the detrital food web, but not microhabitat complexity, affects the role of generalist predators in an aboveground herbivore food web

The activity and density of generalist predators, such as carabid beetles, rove beetles and spiders, may increase in response to: (1) increased availability of prey from the belowground subsystem and/or (2) enhanced complexity of aboveground vegetation. Organic farming practices support decomposer populations and enhance habitat complexity due to an increased weed density. A response by generalist predators to such below‐ or aboveground changes could affect predation rates on herbivores in the aboveground food web. We tested this hypothesis in a replicated field experiment conducted in a winter wheat field, where increased predator activity could lead to improved control of herbivorous pests. In a crossed design, we increased and lowered densities of decomposer prey, and manipulated vegetation complexity using artificial plants in order to examine the effect of structural complexity in isolation from effects of plant‐attracted additional prey. Isotomid Collembola exhibited lowest activity‐densities (AD) in plots treated with soil insecticide and had gradually increasing AD in untreated plots and plots receiving detrital subsidies. Carabid beetles and cursorial spiders did not respond to increased availability of isotomid prey, and they unexpectedly displayed higher AD in the structurally less‐complex plots. Aphid density mirrored the positive response of isotomids to detrital subsidies, suggesting that aphids benefited from reduced predation due to predators switching to abundant prey in the decomposer subsystem. The absence of a numerical response by surface‐active predators apparently strengthened this indirect effect of isotomids on aphids. Our results suggest that indirect predator‐mediated prey‐prey interactions can reduce beneficial effects of detrital subsidies on pest suppression. We further demonstrated that generalist predators may not per se benefit from structural complexity. Both results document the challenges associated with management practices that support generalist predators, as these measures may not necessarily improve herbivore suppression.     

Impact of the invasive moss Campylopus introflexus on carabid beetles (Coleoptera: Carabidae) and spiders (Araneae) in acidic coastal dunes at the southern Baltic Sea

Campylopus introflexus is an invasive moss in Europe and North America that is adapted to acidic and nutrient-poor sandy soils with sparse vegetation. In habitats like acidic coastal dunes (grey dunes) it can reach high densities, build dense carpets and modify habitat conditions. While the impact of the moss invasion on the vegetation is well analyzed, there is a lack of knowledge regarding possible effects on arthropods. In the present study we analyzed the impact of Campylopus introflexus on the ground-dwelling arthropods carabid beetles and spiders, as both taxa are known to be useful indicator taxa even on a small-scale level. In 2009 we compared species composition in a) invaded, moss-rich (C. introflexus) and b) native, lichen-rich (Cladonia spp.) acidic coastal dunes by using pitfall traps. A total of 1,846 carabid beetles (39 species) and 2,682 spiders (66 species) were caught. Species richness of both taxa and activity densities of spiders were lower in invaded sites. Species assemblages of carabids and spiders differed clearly between the two habitat types and single species were displaced by the moss encroachment. Phytophagous carabid beetles, web-building spiders and wolf spiders were more abundant in native, lichen-rich sites. Shifts in species composition can be explained by differences in the vegetation structure, microclimate conditions and most likely a reduced food supply in invaded sites. By forming dense carpets and covering large areas, the moss invasion strongly alters typical arthropod assemblages of endangered and protected (EU-directive) acidic coastal dunes.     

Activity and satiation state in Pterostichus melanarius: an experiment in different agricultural habitats

1. A capture–recapture experiment was conducted to examine the effects of satiation state on the activity and foraging efficiency of the carabid beetle Pterostichus melanarius Ill., in four different habitats (a hedge, a transition zone between the hedge and the adjacent wheat crop, the margin of the wheat crop, and the centre of the wheat crop) in an agricultural ecosystem in France. 2. Prey biomass and density decreased from the centre of the hedge to the crop margin and to the centre of the crop, where prey density was low but prey biomass was moderate. Starved and satiated beetles were released in circular enclosures and recaptured using pitfall traps for 6 days. 3. Immediately after release, starved P. melanarius were significantly more active than satiated beetles, but this difference disappeared after the first day. At each recapture date, starved individuals had gained weight whereas satiated individuals had lost weight. The analysis of activity and weight variation at the end of the first day indicated that the centre of the crop seemed to be the most favourable habitat for starved individuals to find food rapidly, whereas the behaviour of satiated individuals was identical in all habitats. 4. This experiment shows that habitat use of P. melanarius is flexible. The results are discussed in the context of the classical view of foraging behaviour in carabid beetles. The activity and foraging efficiency in P. melanarius in different habitats can be explained as the result of the interaction between individual satiation state, prey availability in the habitat, and particular preferences of the species.     

Population persistence of the parasitoid fly Zaira cinerea (Fallén) (Diptera: Tachinidae) utilizing multiple host carabid beetles with different seasonality and quality

Zaira cinerea (Fallén) is a parasitoid fly (Diptera: Tachinidae) that attacks adult carabid beetles. To better understand mechanisms of population persistence in this species, we examined seasonality of host beetle abundance, the frequency of parasitism, and the timing of fly eclosion. In addition, we evaluated host quality using numbers of larvae or puparia per individual beetle as a measure of quality. The fly parasitized only large carabids (≥15 mm body length); the lengths of fly puparia reached 7.4–10.8 mm during development in beetle abdomens, and larger hosts are likely essential. Of the 18 large carabid species collected in this study, we chose two, Carabus maiyasanus Bates and Leptocarabus procerulus (Bates), because they were large and abundant (87% of total catch). The two carabids had different phonologies; C. maiyasanus was abundant from spring to summer, and its abundance dropped sharply in autumn, while L. procerulus was abundant in autumn and rare from spring to summer except July. Parasitism was observed in all the months from May to November except June, and adult flies eclosed more than once a year (in early summer, late summer, and mid‐autumn), indicating that the species is multivoltine. Host quality of L. procerulus was higher than that of C. maiyasanus. Carabus maiyasanus was mainly used as a host from spring to summer, and L. procerulus was used in autumn. Thus, adult beetles of one or both species are available over most of spring, summer, and autumn, allowing population persistence of this fly species over time.