Habitat choice of some field-inhabiting carabid beetles (Coleoptera: Carabidae) studied by recapture of marked individuals

ABSTRACT.

• 1 Habitat choice of some field-inhabiting carabid beetles was studied in the field, and included the adult-overwintering Bembidion lampros Herbst, Pterostichus cupreus L. and Agonum dorsale Pontoppidan, and the larval-overwintering Trechus secalis Paykull, Pterostichus melanarius Illiger, P.niger Schaller and Harpalus rufipes De Geer. These were compared to the forest-inhabiting and adult-overwintering species, Carabus nemoralis Müller.
• 2 Marked beetles were released in the centre of a circular enclosure which was placed on the edge between a cereal field and a wood. The direction of movement in this circle was compared with that in a control circle placed entirely in the cereal field.
• 3 In the field/wood circle, adult-overwinterers and the larval-overwin-terer H.rufipes choose to move into the cereal field. The majority of T.secalis. P.melanarius and C.nemoralis, however, moved into the wood, while P.niger exhibited no preference.
• 4 In the field circle, adult-overwinterers moved towards the more sun-exposed parts of the circle, whereas the movements of larval-overwin-terers were random.
• 5 Some species-specific characteristics considered important for habitat choice by these carabid beetles, and the relevance of these characteristics for their dispersal behaviour are discussed.

     

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.     

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.     

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.     

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.

     

Kleptoparasitic behaviour of Aphodius rufipes (L.) larvae in nests of Geotrupes spiniger Marsh. (Coleoptera, Scarabaeidae)

Abstract.

• 1 Third instar larvae of A.rufipes were found in short vertical shafts in the soil beneath horse dung and they entered the underground brood masses of G.spiniger when these occurred beneath the same deposit of dung.
• 2 A.rufipes larvae excavated shafts (60–80mm deep) beneath dung in cages. Just before diapause they burrowed down to the floor of the cage. Burrowing was inhibited in compact soil and in very dry soil. The feeding larvae were attracted to dung and to moisture and they readily attacked other larvae with their mandibles.
• 3 In cages which contained G. spiniger nests the A.rufipes larvae burrowed down to feed on the G.spiniger brood masses. G.spiniger eggs and larvae did not act as attractants but they were often destroyed if the A.rufipes larvae encountered them by chance.
• 4 In cages which contained G.stercorarxus nests A.fossor larvae burrowed down to feed on the brood masses if no dung was provided at the surface.
• 5 This non-obligatory parasitism arises from the normal behaviour of the larvae and is usually of trivial significance, but under certain conditions the protected environment of an underground nest may favour the survival of the invading larvae.

     

Impact of predatory carabids on below- and above-ground pests and yield in strawberry

The impact of adult carabid beetles on below- and above-ground pests and fruit yield was examined in the laboratory and a two-year strawberry field study. In the laboratory, adults of Carabus nemoralis Muller, Nebria brevicollis (F.), Pterostichus algidus LeConte, Pterostichus melanarius (Illiger), and Scaphinotus marginatus Fischer (Coleoptera: Carabidae) consumed black vine weevil, Otiorhynchus sulcatus (F.) (Coleoptera: Curculionidae) eggs, larvae and/or pupae placed on the surface. The same five carabid species showed no impact or low removal rates of O. sulcatus larvae that had burrowed into the root of potted strawberry plants. In an assay with only P. melanarius, adults consumed O. sulcatus larvae placed on the soil surface more frequently than larvae buried 1.3 or 5?cm below. In a field study, the density of adult carabids, predominantly P. melanarius, was manipulated with augmented, exclusion, and open control plots (2?×?2?m). Manipulating carabid density had no impact on the removal of sentinel O. sulcatus larvae and pupae that were buried belowground which is consistent with laboratory observations. Increasing carabid density within augmented plots led to greater removal of red clover seeds, Trifolium pratense L., placed on the soil surface in the first?year. Decreasing carabid density within exclusion plots resulted in fewer marketable fruits compared to control plots in both years. These results suggest that certain adult carabids may have limited impact belowground, and some beneficial impacts above-ground with pest control and crop protection.     

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.     

Diet, fecundity and egg size in some polyphagous predatory carabid beetles

Egg size was measured at different rates of egg laying in three polyphagous carabid species, known to be useful predators of cereal aphids; the small Bembidion lampros Herbst and the medium-sized Pterostichus cupreus L. and P. melanarius Illiger. Variations in fecundity, as well as the ability of the medium-sized species to also build up fat reserves, were obtained when beetles were subjected to different dietary regimes consisting of aphids, or foods with a lower or higher protein content. Egg size was found to be dependent on the rate of egg laying within a species. A diet of cereal aphids appeared to be adequate for egg production in these polyphagous carabids, but female P. cupreus were unable to build up fat reserves when they ingested aphids contaminated with the aphicide pirimicarb. Beetles were able to devote resources to more and larger eggs (B. lampros), or to larger eggs and/or fat reserves (P. melanarius/P. cupreus) when given access to a carbohydrate-rich food with low protein content. The highest rate of egg laying was obtained when female P. cupreus and P. melanarius were given a more varied diet at frequent intervals; including regular shifts between unsprayed aphids, carbohydrate-rich food and protein-rich maggots. Within the varied diet treatment a negative relationship was obtained between egg size and egg number among similar-sized individuals of P. cupreus and P. melanarius; females producing the largest number also laid the smallest eggs. Egg size affected larval survival, since first instars hatching from large eggs were found to survive longer than those hatching from small eggs. The influence of differences in food intake on reproduction, maintenance metabolism, and survival of fieldinhabiting carabids is discussed.     

Isolating the components of activity-density for the carabid beetle Pterostichus melanarius in farmland

The activity and density of the carabid beetle, Pterostichus melanarius, were studied over 10 weeks in a continuous mark-recapture experiment using a grid of pitfall traps spanning a hedgerow and extending approximately 30 m into two cereal fields; 1777 beetles were individually marked. The recapture rate was approximately 60% and 40% for males and females, respectively. Activity-density rose and fell four times between early June and mid August. Jolly-Seber estimates of density showed population density increasing to a single peak in late July. The mean population density in late July and August was relatively stable at approximately 0.26 m⁻². Activity varied over the 10 weeks and was significantly higher during August than in June or July. The daily displacement distance frequencies, calculated from 750 male and 485 female recaptures of individually marked beetles, were distributed exponentially. Mean displacement distances were approximately 2.6 m day⁻¹ during June and July, and 5.3 m day⁻¹ during August. A diffusion model provided diffusion coefficients of 23.7 and 27.9 m² day⁻¹ for female and male beetles, respectively. The spatial distribution of P. melanarius was aggregated in patches. Spatial analysis by distance indices showed the spatial distribution of counts between successive periods of activity-density to be significantly associated. Approximately 5.75% of recaptures were from releases on the opposite side of the hedgerow. Approximately 20% of recaptures were from releases in opposite halves of the grid within the same field. The hedgerow acted as a significant barrier to dispersal between fields, with implications for the metapopulation structure of the species. Received: 26 January 1998 / Accepted: 1 April 1998     

Predation by Pterostichus melanarius (Illiger) (Coleoptera: Carabidae) on immature Rhagoletis mendax Curran (Diptera: Tephritidae) in semi-field and field conditions

Practices that enhance abundance and diversity of generalist predators are often employed with the objective of improving biological control of insect pests. Ground beetles and other predators can prey on blueberry maggot, an important pest of blueberries, when mature larvae pupate in the ground. We conducted mesocosm and field experiments to determine if Pterostichus melanarius, a common predatory ground beetle, lowers maggot numbers in compost mulch or when predator and alternative prey abundances are manipulated. At background (field) densities of alternative prey, increasing densities of P. melanarius did not significantly reduce pest numbers in mesocosms containing compost or soil. When alternative prey were removed from compost, beetles reduced pest numbers by up to 35%. In field experiments, maggot numbers were higher when beetles and other predators were excluded from soil plots, but beetle exclusion had no effect in compost plots where both predator and alternative prey numbers were high. Our results indicate that there can be some reduction of blueberry maggot by P. melanarius and other potential predators when there are few alternative prey. However, despite attracting large numbers of predators compost mulch did not lead to a significant reduction in blueberry maggot; in fact, the high abundance of alternative food associated with compost appeared to interfere with beetle predation on blueberry maggot.     

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.     

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.     

Long-termed changes in ground beetle (Coleoptera: Carabidae) assemblages in a field treated by organic fertilizers

In 2001–2006, ground beetles were pitfall-trapped in a temperate lowland area of South Slovakia in an experimental field divided in five plots fertilized by four different doses of manure or biosludge (25 t stable manure ha⁻¹, 50 t biosludge ha⁻¹, 50 t stable manure ha⁻¹, 100 t biosludge ha⁻¹ and without fertilization). The field was subsequently sown by spring barley, sugar beat, maize, sunflower, sugar beat and maize. The ground beetle assemblage consisted of 31 species, but only five species predominated: Pseudoophonus rufipes representing 82.6% of individuals and five species (Poecilus cupreus, Carabus scheidleri, Calathus fuscipes, Trechus quadristriatus, Pterostichus melanarius, Anchomenus dorsalis, Dolichus halensis) representing together 14.5% of individuals. Pseudophonus rufipes represented 81.7% of dry biomass and three species (Carabus scheidleri, Poecilus cupreus Pterostichus melanarius) 15.9% of biomass. There was no significant influence of organic fertilizing on assemblage structure. During the investigation, the number of individuals and their biomass increased in all plots until 2003 and than dropped to the starting values. The culmination of 2003 was preceded by a warmer and more humid season in 2002. After a cold and dry season of 2003 abundance decreased approximately to starting values. Simultaneously, the local maxima and minima of occurrence of ground beetles in individual plots shifted independently on the doses of organic material. At the same time, number of occurring species slightly decreased. The observed changes obviously represent part of long-termed fluctuations in wider surroundings.     

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

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Mandible wear in the carabid beetle Pterostichus melanarius in relation to diet and burrowing behaviour

The effects of food texture and burrowing on mandible wear in the predatory carabid beetle Pterostichus melanarius Illiger were investigated by feeding adults soft or hard food in jars with or without soil. Both the incisor cusp (mandible tip) and cutting surface of the mandibles erode more in response to the combination of hard food and burrowing than to hard food alone. Females were found to erode their mandibles more than males in the hard-food-only treatment, whereas males wore down their mandibles more than females in the soil-only treatment. Female P. melanarius collected in a cereal field contained signficantly more solid food items compared with males. Differences in behaviour apparently resulted in differential wear patterns between the sexes.It is shown that the process of mandible wear in P. melanarius will result in worn mandibles in their first year as adults. However, mandible size of field-sampled females were found to be best correlated with females (of similar age) in the experiment exhibiting extensive wear (foraged hard food and burrowed in soil). The mandibles of field-sampled males were similar to those of males in the hard-food-only and the soil-only treatments, which showed only slight to moderate wear. The influence of mandible wear on fecundity and survival in P. melanarius as well as in other carabid species is discussed.

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Usure des mandibules du carabe, Pterostichus melanarius Illiger, en fonction de l'alimentation et du comportement de fouissage

Résumé L'analyse de l'influence de la texture des aliments et du fouissage, sur l'usure des mandibules a été effectuée en nourrissant P. melanarius avec des aliments de différentes duretés dans des récipients contenant ou non du sol. Les aliments durs seuls usent moins le bord incisif (extrémité de la mandibule) et la surface coupante, que leur association avec le fouissage. En présence exclusive d'aliments durs, les femelles avaient érodé plus vite leurs mandibules que les mâles, tandis que chez ceux-ci le fouissage seul les avaient usé plus vite que chez les femelles. Des femelles récoltées dans un champ de céréales contenaient plus d'aliments durs que les mâles. Les différences de comportement avaient provoqué apparemment des types d'usure différents suivant les sexes.Les adultes usent leurs mandibules au cours de leur première année de vie imaginale. Cependant, la taille des mandibules des femelles échantillonnées dans la nature correspondait mieux à celle des femelles (de même âge) provenant des expériences ayant provoqué une forte usure (aliment dur et fouissage du sol). Les mandibules des mâles échantillonnés présentaient seulement une usure modérée et ressemblaient à celles des mâles ayant consommé soit des aliments durs, soit foui le sol. La discussion a traité des relations entre l'usure des mandibules, la fécondité et la longévité de P. melanarius et d'autres espèces de carabes.

     

Diversity and seasonal phenology of aboveground arthropods in conventional and transgenic maize crops in Central Spain

One of the major concerns regarding the release of Bt maize is its potential negative impact on non-target organisms present in this crop. In this paper, we compare the temporal phenology and community structure of the aboveground arthropods in commercial Bt maize fields in Central Spain with those of conventional maize crops, with or without an insecticide (imidacloprid) seed treatment, over a period of three years. Spiders, harvestmen, centipedes, ground beetles, rove beetles, carrion beetles, click beetles, earwigs and damsel bugs were captured in pitfall traps every year in sufficient number to provide meaningful phenological data. One predator spider and three omnivorous species of ground beetles have been consistently present in the maize fields: Pardosa occidentalis, Poecilus cupreus, Pseudophonus rufipes and Pseudophonus griseus, respectively. Rove beetles were caught to a lesser extent, with three dominant species: Acrotona aterrima, Philonthus varians and Platystethus nitens. The variability in activity–density patterns of the aboveground fauna was mainly influenced by the year, but no detrimental effects could be attributed to Bt maize. The only exception being the changes detected in rove beetles, although these differences were transitory and varied from year to year. No changes in species richness and diversity indices for spiders and ground beetles resulted from treatments. However, imidacloprid-treated maize caused a reduction in species richness of rove beetles, even though the abundance of the main species was not reduced. Our results suggest that Bt maize could be compatible with natural enemies that are common in maize fields in Spain.     

Application of harmonic radar to analyze dispersal behavior of the Japanese pine sawyer beetle,Monochamus alternatus (Coleoptera: Cerambycidae)

Monochamus alternatus (Hope) is a severe wood‐boring pest in coniferous forests and a major vector of pine wilt disease in East Asia. Harmonic radar is a powerful tool for studying the dispersal behavior of insects and it could be applied to control pine wilt disease. In this study, we validated the application of harmonic radar for analyzing the dispersal behavior of M. alternatus beetles in a field environment. We determined the wing capacities of the beetles and the effects of electronic tagging and marking on their movement, flight ability, survivorship, and food consumption in the laboratory to confirm the suitability of this technique. The detection rate and recovery rate for beetles were analyzed separately using radar on caged pine stands and in the field environment. The results showed that the minimum wing capacity of the Japanese pine sawyer was 24.9 mg, which was seven times the weight of the electronic tag (3.5 mg). Marking and tagging the beetles had no significant adverse effects on their movement, flight capacity, food consumption, and survivorship. The detection rate using the radar system and recovery rate based on visual observations of the beetles in caged pines were both 95.6%. However, in the field environment, the detection and recovery rates were only 55.6% and 37.8% after one week, respectively, and 33.3% and 7.8% after two weeks. Harmonic radar is a promising technique for studying the dispersal behavior of the Japanese pine sawyer, but its performance is not satisfactory and major improvements are required for both the radar system and electronic tags.     

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.     

The influence of different age classes on the seasonal activity and reproduction of four medium-sized carabid species inhabiting cereal fields

Reproduction and survival of four carabid species commonly occurring in cereal fields in central Sweden were investigated. The adult-overwintering Pterostichus cupreus L. bred in early summer (June), and the largest proportion of sexually active beetles were found to be in their first breeding year; only a minority were 2 (or more) years old. In contrast, the larval-overwintering P. melanarius Illiger, P. niger Schaller and Harpalus rufipes De Geer had a biennal life cycle, and mainly old beetles (2 years or older) contributed to the population's reproductive effort each year. The long prereproductive adult phase in the larval-overwintering species and the relatively short summer in central Sweden prevented many new-generation adults from breeding in their first year. Several overlapping breeding cohorts are therefore present each year, and overwintered beetles bred throughout summer and during part of the autumn. Mandible wear (combined with other characters) was used to determine the age of beetles. However, the rate of mandible wear also was found to vary, depending upon the extent to which each species burrowed in the ground. The ecological significance of (a) a long adult life and (b) the existence of several overlapping breeding cohorts for reproduction and survival of populations in cereal fields is discussed.