Better the devil you know: avian predators find variation in prey toxicity aversive

Toxic prey that signal their defences to predators using conspicuous warning signals are called ‘aposematic’. Predators learn about the toxic content of aposematic prey and reduce their attacks on them. However, through regulating their toxin intake, predators will include aposematic prey in their diets when the benefits of gaining the nutrients they contain outweigh the costs of ingesting the prey''s toxins. Predators face a problem when managing their toxin intake: prey sharing the same warning signal often vary in their toxicities. Given that predators should avoid uncertainty when managing their toxin intake, we tested whether European starlings (Sturnus vulgaris) preferred to eat fixed-defence prey (where all prey contained a 2% quinine solution) to mixed-defence prey (where half the prey contained a 4% quinine solution and the other half contained only water). Our results support the idea that predators should be more ‘risk-averse’ when foraging on variably defended prey and suggest that variation in toxicity levels could be a form of defence.     

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.     

Do earthworms help to sustain the slug predator Pterostichus melanarius (Coleoptera: Carabidae) within crops? Investigations using monoclonal antibodies

Earthworms provide a major potential source of alternative food for polyphagous predators, such as carabid beetles, that are natural enemies of slugs, aphids and other agricultural pests. Non-pest prey may foster larger numbers of natural enemies, which then help to control pests, or alternatively may help to divert the predators away from pest control. An earthworm-specific monoclonal antibody was developed to study carabid-earthworm interactions in the field and assess the role of earthworms as alternative prey. The antibody could identify as little at 7 ng of earthworm protein in an ELISA, and could detect earthworm remains in the foregut of the carabid beetle Pterostichus melanarius for 64 h after consumption. Thirty-six per cent of field-collected beetles contained earthworm remains. Quantities of earthworm proteins in the beetle foreguts were negatively related to total foregut biomass, suggesting that earthworm consumption increased as total prey availability declined. There was also a negative relationship between foregut biomass and beetle numbers, but both quantities and concentrations of earthworm proteins in beetle foreguts were positively related to beetle numbers. This suggests that as beetle activity-density increased, total prey availability declined, or, as prey availability declined, beetles spent more time searching. In these circumstances, beetles fed to a greater extent on earthworms, an acceptable but nonpreferred food item. Earthworms may, therefore, provide an ideal alternative prey for P. melanarius, helping to sustain it when pest numbers are low but allowing it to perform a 'lying-in-wait' strategy, ready to switch back to feeding on pests when they become available.     

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.     

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.     

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.     

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.     

Aposematism and gregariousness: the combined effect of group size and coloration on signal repellence

Many aposematic species have evolved an aggregated lifestyle, and one possible advantage of grouping in warningly coloured prey is that it makes the aposematic signal more effective by generating a greater aversion in predators. Here we investigate the effect of prey group size on predator behaviour, both when prey are aposematic and when they are not aposematic, to separate the effects of warning coloration and prey novelty. Naive domestic chicks (Gallus gallus domesticus) were presented with either solitary or groups of 3, 9 or 27 live larvae of the aposematic bug Tropidothorax leucopterus. Other naive chicks were presented with larvae of the non-aposematic bug Graptostethus servus either solitary or in groups of 27. Attack probability decreased with increasing group size of aposematic prey, both when birds were naive and when they had prior experience, whereas prey gregariousness did not affect the initial attack probability on the G. servus larvae. In a separate experiment, groups of mealworms were shown to be even more attractive than solitary mealworms to naive chicks. We conclude that the aversiveness of prey grouping in this study can be explained as increased signal repellence of specific prey coloration, in this case a classical warning coloration. These experiments thus support the idea of gregariousness increasing the signalling effect of warning coloration.     

Prey-mediated effects of the protease inhibitor aprotinin on the predatory carabid beetle Nebria brevicollis

To investigate the potential non-target impacts of transgenic pest-resistant plants, prey-mediated impacts of a protease inhibitor (PI) on the predatory carabid, Nebria brevicollis, were investigated. The PI used was aprotinin, a serine PI of mammalian origin with insecticidal properties when incorporated in artificial diet or expressed in transgenic plants. Field-collected N. brevicollis adults, kept at 23 °C, 16:8 L:D, were fed, over their pre-aestivation activity period of 24 days, with Helicoverpa armigera larvae reared on an artificial diet containing 0.5% (w:w, fresh mass) aprotinin. These larvae contained 22.62 μg aprotinin/g insect. Control prey was reared on diet without aprotinin. Beetle survival and body mass were unaffected by prey type. Beetles consuming PI-fed prey lost significantly more mass than the control beetles during two periods of mass loss, but gained significantly more mass during the final period of mass gain. This was not due to differences in amounts of prey supplied or consumed. The final mass gain coincided with increased consumption of PI-prey. Female beetles were significantly heavier than males, but we found no consistent gender-based differences in response to PI-prey. At the end of the experiment, body mass of all beetles was similar to field-collected ones (approximately 55 mg). All experimental beetles had significantly lower activities of digestive cysteine proteases and the serine proteases chymotrypsin and trypsin than field-collected ones. Beetles consuming PI-fed prey had significantly lower levels of trypsin and higher levels of chymotrypsin and elastase than the control beetles.     

Morphometry of eyes, antennae and wings in three species of Siagona(Coleoptera, Carabidae)

In carabid beetles, physiological and behavioural characteristics reflect specific habitat demands and there is a strong correlation between body form and habit in species with different life style. In this study, we compared the morphometry and compound eye characteristics of three species of the genus Siagona: Siagona jenissoni, Siagona dejeani and Siagona europaea. These carabids have a stenotopic lifestyle in Mediterranean clayey soils, inhabiting the ground fissure system formed during the dry season. All species have a Mediterranean distribution and are nocturnal olfactory hunters, and are strict ant predators. For morphometric measurements, we considered body length (mm), wing length (mm), antenna length (mm), head width (mm), trochanter length (mm), number of ommatidia, eye surface area (mm2), ommatidia density (number of ommatidia/mm2 of eye surface area), head height (mm), thorax height (mm) and abdomen height (mm). The data revealed intersexual and interspecific differences. The three species differ in relative length of the antennae, density and number of ommatidia and relative trochanter length. Significant differences occurred in wing sizes, which are well developed in Siagona europaea, the only species capable of flight. When eye size is compared with other ground beetles of various lifestyles, Siagona shows pronounced "microphthalmy" an adaptation to subterranean life in clayey crevices of tropical and subtropical climates with a marked dry season.     

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.     

Dietary conservatism may facilitate the initial evolution of aposematism

It has generally been assumed that warningly coloured organisms pay a cost associated with their increased visibility, because naïve predators notice and eat them. This cost is offset by their enhanced protection from educated predators who associate the colour pattern with unprofitability. However, some studies have suggested that avoidance of novel prey by avian predators ("dietary conservatism") can actually place novel colour morphs at a selective advantage over familiar ones, even when they are highly conspicuous. To test this idea, we experimentally simulated the appearance of a single novel-coloured mutant in small populations (20 individuals) of palatable artificial prey. The colour morph frequencies in each "generation" were determined by the relative survival of the previous generation under predation by birds. We used wild-caught European robins Erithacus rubecula foraging on pastry "prey" of different colours. The aim was to test whether prey selection by predators prevented or facilitated the novel colour morph persisting in the prey population over successive generations. We found that the novel colour morph quickly increased to fixation in 14/40 prey "populations", and at least once each in 8 of the 10 birds tested. Novel mutants of the classic aposematic colours (red and yellow) reached fixation most frequently, but even the green and blue novel morphs both increased to fixation in 2/40 trials. Novel colours reached fixation significantly faster than could be accounted for by drift, indicating active avoidance by the birds. These results suggest that a novel colour morph arising in a prey population can persist and increase under the selective pressure imposed by predators, even to the local exclusion of the original morph, despite being fully palatable. The consequences of this finding are discussed in relation to receiver psychology, the evolution of aposematism and the existence of polymorphism in Müllerian mimics.     

Emergent effects of ground beetles size diversity on the strength of prey suppression

相似文献   

Experimental evidence of bark beetle adaptation to a fungal symbiont

The importance of symbiotic microbes to insects cannot be overstated; however, we have a poor understanding of the evolutionary processes that shape most insect–microbe interactions. Many bark beetle (Coleoptera: Curculionidae, Scolytinae) species are involved in what have been described as obligate mutualisms with symbiotic fungi. Beetles benefit through supplementing their nutrient‐poor diet with fungi and the fungi benefit through gaining transportation to resources. However, only a few beetle–fungal symbioses have been experimentally manipulated to test whether the relationship is obligate. Furthermore, none have tested for adaptation of beetles to their specific symbionts, one of the requirements for coevolution. We experimentally manipulated the western pine beetle–fungus symbiosis to determine whether the beetle is obligately dependent upon fungi and to test for fine‐scale adaptation of the beetle to one of its symbiotic fungi, Entomocorticium sp. B. We reared beetles from a single population with either a natal isolate of E. sp. B (isolated from the same population from which the beetles originated), a non‐natal isolate (a genetically divergent isolate from a geographically distant beetle population), or with no fungi. We found that fungi were crucial for the successful development of western pine beetles. We also found no significant difference in the effects of the natal and non‐natal isolate on beetle fitness parameters. However, brood adult beetles failed to incorporate the non‐natal fungus into their fungal transport structure (mycangium) indicating adaption by the beetle to particular genotypes of symbiotic fungi. Our results suggest that beetle–fungus mutualisms and symbiont fidelity may be maintained via an undescribed recognition mechanism of the beetles for particular symbionts that may promote particular associations through time.     

Forest edge contrasts have a predictable effect on the spatial distribution of carabid beetles in urban forests

Fragmented urban forest remnants are characterised by sharp edges and are bordered by various land-use types, which may have a considerable effect on the fauna and flora at forest edges, and into forest interiors. To investigate the effects of differentially contrasting edges (low vs. intermediate vs. high) on carabid beetle assemblages in urban boreal forests, we placed pitfall traps along a gradient from 6?m into three matrix types (secondary forest vs. grassland vs. asphalt) up to 60?m into urban forest patches in the cities of Vantaa and Helsinki, southern Finland. Individual species and carabid beetle assemblages were strongly affected by edge contrasts and distance from the forest edge. The strongest effect on individual species was caused by high contrasting edges: generalist and open-habitat species were favoured or not affected while forest specialists were affected negatively. Effects of the abundances of potential prey and competitors on the carabid beetles were also evaluated. Forest and moisture-associated carabid species were negatively to neutrally associated with springtail abundances while generalist and open habitat, and dryness associated species were more positively related to springtail abundances (a potential food source). In terms of potential competitors, forest and moisture-associated carabid species were negatively and/or neutrally affected by ant and wood ant numbers, while generalist and open-habitat species were neutrally to positively associated with these taxa. It appears that carabid beetle habitat associations are more important in the responses of these beetles across edges of different contrast than are the prey and competitor numbers collected there. We recommend the creation of “soft” or low-contrast urban edges if the aim of urban management is to protect forest carabids in cities.     

基于MaxEnt模型的草原步甲属潜在地理分布研究

步甲(Carabid beetle)是草原生态系统重要的昆虫类群之一,对草原生态系统环境监测及维护生物多样性起着重要的作用。基于步甲属(雕步甲和长叶步甲)在宁夏草原131个有效调查记录和19个环境因子,利用最大熵模型(Maximum Entropy Model,简称MaxEnt)预测步甲属潜在空间地理分布,运用受试者工作特征曲线(ROC曲线)检测预测模型精度,刀切法(Jackknife)筛选影响步甲属潜在地理分布的主要生态因子。结果表明：(1) MaxEnt模型预测结果 ROC曲线AUC值分别为0.981、0.914,模型预测结果的可信度较高;(2) 19个环境因子中,对步甲属的潜在空间分布预测贡献率较高的环境因子为最冷月份最低气温、年均降水量、最干季均温、年平均气温;(3)雕步甲主要分布在温性荒漠草原及温性草甸草原的北部,长叶步甲主要分布在温性草甸草原。以上研究为步甲调查和保护提供参考依据。     

Reactions of passerine birds to aposematic and non-aposematic firebugs (Pyrrhocoris apterus ; Heteroptera)

In spite of the existence of many experimental studies on the function of warning coloration in insects, little is known about the universality of reactions of different predators towards a particular warning signal. Reactions of nine passerine bird species, namely Parus major , Parus caeruleus , Aegithalos caudatus , Erithacus rubecula , Turdus merula , Sylvia atricapilla , Fringilla coelebs , Carduelis chloris and Emberiza citrinella , to the firebug Pyrrhocoris apterus wildtype (brachypterous adults) and its artificially obtained (painted) brown non-aposematic variant were compared. Most insectivorous birds (great tits, blue tits, robins and blackcaps) distinguished between aposematic and non-aposematic bugs, attacking the former less often. Partly granivorous buntings and finches did not distinguish between them, and attacked both variants equally. As all the birds were caught in the wild, the results can be interpreted in terms of the presence of a higher proportion of experienced individuals among insectivorous than among omnivorous species. Two insectivorous species differed from others. The heaviest blackbird attacked and killed aposematic as well as non-aposematic firebugs, and, in contrast, the lightest long-tailed tit avoided both variants. © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 78 , 517–525.     

Predators,Parasites and Heterospecific Aggregations in Chrysomeline Larvae

Gregariousness is a common feature in aposematic insect prey and offers the additional benefit of enhancing the effectiveness of their toxic defences. Aggregations of the aposematic larvae of two species of leaf beetles, Paropsis atomaria and Paropsisterna variicollis, occur together on the same Eucalyptus trees over spring and summer. Conventionally, the colouration of these larvae is thought to provide aposematic protection against vertebrate predators, but supporting evidence is limited. We determine whether environmental preferences contribute to the heterospecific aggregations, and the potential costs for larvae of living in heterospecific aggregations. We surveyed natural aggregations of the larvae of both species in the field and recorded environmental variables of these aggregations. This revealed that heterospecific aggregations occurred more commonly at higher leaf temperatures, and in less visually conspicuous locations. Paropsis atomaria larvae were twice as likely to be found in heterospecific aggregations than Pa. variicollis. Next, we manipulated larval aggregations in the field to investigate the survival and gregarious behaviour of larvae in heterospecific aggregations. The gregarious behaviour of both species of larvae did not differ between heterospecific and monospecific aggregations. Further, the survival of larvae did not significantly differ between heterospecific and monospecific aggregations. We suggest that the preference for P. atomaria larvae to aggregate with Pa. variicollis at higher leaf temperature results in the observed heterospecific aggregations, with a potential benefit of lowerer parasitoidism rates for P. atomaria.     

Studies of adaptation of carabid beetles (Coleoptera,carabidae) to feeding on the Colorado potato beetle, <Emphasis Type="Italic">Leptinotarsa decemlineata</Emphasis> say (Coleoptera,chrysomelidae) in Agrocenoses of potato and other crops

In 1979–2008, serological investigation of carabid beetles as predators of the Colorado potato beetle was carried out in the fields of potato and other Solanaceae crops in nine regions of Russia, Moldova, and Ukraine. The fraction of carabid beetles feeding on the pest in the potato, tomato, and egg-plant fields grows with the duration of the pest presence in the region and is proportional to its population density.     

Molecular identification of predation by carabid beetles on exotic and native slugs in a strawberry agroecosystem

Generalist predators have the capacity to exert significant pressure on prey populations. However, integrating them into biological control programs relies on a detailed understanding of their foraging behavior and the levels of trophic connectedness with pest species. Carabid beetles are important predators of slugs, pests of agricultural, floricultural and horticultural crops worldwide, but these interactions have been rarely studied outside the Western Palearctic ecozone. Diagnostic molecular gut-content analysis was used to examine the strength of trophic pathways between a community of carabid beetles and two slug species, the exotic Deroceras reticulatum and native Deroceras laeve, in strawberry agroecosystems. Strawberries were grown according to standard horticultural practices for central Kentucky, following traditional bare ground planting or with the addition of detrital subsidies, to quantify the impact of habitat management on the abundance of pests and the strength of these trophic pathways. Following laboratory characterization of species-specific molecular markers targeting both Deroceras species, carabid beetles collected from a strawberry agroecosystem were screened for slug DNA. Field collections revealed important food web pathways existed between Harpalus pensylvanicus and D. reticulatum, with 7.2% screening positive for these prey yet none screening positive for D. laeve. In contrast, Chlaenius tricolor was found to feed on both slug species in the field, with 16% screening positive for both Deroceras. Despite below normal rainfall limiting slug densities in the field, the results presented here reveal the potential importance of carabid beetles in slug population dynamics in the Nearctic.