A laboratory study on temperature relations of egg production and development in two related species of carabid beetle

Summary Egg production and development of the carabid beetles Notiophilus biguttatus F. and Notiophilus rufipes Curtis were studied in relation to temperature. Adult beetles and larvae were fed springtails of the species Orchesella cincta (Linné). The rate of egg production, at the different temperatures did not differ between the species. As observed in N. biguttatus egg production at a daily fluctuating temperature regime is high compared to egg production at the constant temperatures. Food supply not only has a strong effect on egg production but also on the relation of egg production with temperature, as shown in N. biguttatus. Development times in N. rufipes are longer than in N. biguttatus particularly at low temperatures. The temperature during development has a distinct effect on adult body size. As observed in N. biguttatus body size and fecundity show a strong positive correlation.     

Hunger, partial consumption of prey and prey size preference in a carabid beetle

ABSTRACT.

• 1 Partial consumption of prey and prey size preference were studied in females of the carabid Notiophilus biguttatus F., using different size-classes of the springtail Orchesella cincta (L.) as prey.
• 2 Time to ingest a prey increases disproportionately with prey size, mainly as a consequence of satiation.
• 3 During consumption of a prey the predator's rate of ingestion decreases, partly because of diminishing returns from the prey over feeding time.
• 4 The hypothesis that the diminishing returns from the prey induce partial consumption was refuted.
• 5 Partial consumption in the beetle is due to gut limitation; its occurrence depends on prey size.
• 6 Average weight of prey remains in four prey size classes were close to weights expected from average intercatch intervals and estimates of hunger.
• 7 Prey choice depends on level of food deprivation.
• 8 Partial consumption of prey, prey size preference and profitability of prey in relation to hunger of the beetle are discussed.

     

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.     

Respiratory metabolism in two species of carabid beetle from the sub-Antarctic island of South Georgia

The influence of feeding state on cold-adapted metabolism was investigated in the adults of two carabid beetles, Trechisibus antarcticus and Oopterus soledadinus (Coleoptera: Carabidae), which have been introduced to sub-Antarctic South Georgia. The metabolic rates in both fed and starved O. soledadinus and T. antarcticus were determined at eight temperatures ranging from 0 to 35°C, using a Servomex 570A oxygen analyser. There was no significant difference in the metabolic rates between the fed and starved animals of each species. In T. antarcticus this ranged from 0.28 to 3.84 ml O₂ g⁻¹ h⁻¹, and in O. soledadinus from 0.19 to 2.80 ml O₂ g⁻¹ h⁻¹ at 0 and 35°C, respectively. In each of the four experimental groups there was a strong positive correlation between metabolic rate and temperature, with the highest increase occurring between 0 and 5°C. In contrast, the metabolic rate was significantly negatively correlated with initial live weight of the beetles at most temperatures. The results are discussed comparatively with other species and against a background of the ecology of the two carabids at South Georgia. Received: 26 August 1996 / Accepted: 3 February 1997     

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

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

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

     

The effects of vegetation types and microhabitats on carabid beetle community composition in cool temperate Japan

The effects of vegetation types and environmental factors on carabid beetle (Coleoptera: Carabidae) communities were studied. Carabid beetles were collected using pitfall traps (total 2844 trapping days) and seven microenvironmental factors were measured in four vegetation types: grassland, natural evergreen coniferous forest (Pinus densiflora), deciduous broad-leaved natural forest (Quercus crispula, Betula platyphylla, Alnus japonica, or Fagus crenata), and deciduous coniferous plantation (Larix kaempferi) in cool temperate Japan. These four vegetation types provided a novel comparison between natural forests and plantations because the vast majority of related studies have investigated only deciduous broad-leaved natural forests and evergreen coniferous plantations. PERMANOVA indicated that vegetation types affected carabid community composition. Ordination plots showed that community composition differed greatly between grassland and forest vegetation types, but that community composition in the plantation forest overlapped with that of natural forest types. Characteristics differentiating the grassland included a high proportion of winged species and a low mean carabid body weight. Among the examined environmental factors, litter depth, soil water content, and depth of the soil A-horizon had large effects on carabid communities. These results suggest that the effect of afforestation on carabid communities in cool temperate Japan might be insignificant compared with the effects of cover types (deciduous vs. evergreen) and microenvironmental factors.     

Heathland management effects on carabid beetle communities: the relationship between bare ground patch size and carabid biodiversity

Carabid beetles were sampled by pitfall trapping on Brentmoor Heath, Surrey, UK during the summers of 2009 and 2010 to determine the effects of bare ground patch size on beetle abundance, richness and diversity. Four patch sizes were investigated: 1, 4, 25 and 100?m² as well as the adjacent mown and unmown areas. A range of environmental parameters relating to soil characteristics, stones and nearby vegetation were measured at each patch and control habitat in order to distinguish the effect of patch size. Results show that bare ground is a valuable habitat for carabids, but that the response of their abundance, richness and diversity to bare ground patch size depends on a large number of environmental variables on and around the patches. When all variables are taken into account, smaller patches appear to benefit carabid abundance, richness and diversity. In the presence of Molinia caerulea at the patch edges, however, larger patches were more beneficial. Given this dependence on environmental variables, the perfect patch size for conservation of biodiversity is likely to be site specific and the best approach may be to use a variety of patch sizes at a range of successional stages.     

Prey-mediated effects of transgenic canola on a beneficial, non-target, carabid beetle

Transgenic plants producing insecticidal proteins from Bacillus thuringiensis (Bt) can control some major insect pests and reduce reliance on sprayed insecticides. However, large scale adoption of this technology has raised concerns about potential negative effects, including evolution of pest resistance to Bt toxins, transgene flow from Bt crops to other plants, and harm to non-target beneficial organisms. Furthermore, concern has also been expressed over the effects this technology may have on biodiversity in general. Ecologically relevant risk assessment is therefore required (Risk = Hazard × Exposure). Transgenic plants that produce Bt toxins to kill insect pests could harm beneficial predators. This might occur directly by transmission of toxin via prey, or indirectly by toxin-induced reduction in prey quality (Hazard). To test these hypotheses, we determined the effects of Bt-producing canola on a predatory ground beetle (Pterostichus madidus) fed larvae of diamondback moth (Plutella xylostella) that were either susceptible or resistant to the Bt toxin. Survival, weight gain, and adult reproductive fitness did not differ between beetles fed prey reared on Bt-producing plants and those fed prey from control plants. Furthermore, while Bt-resistant prey was shown to deliver high levels of toxin to the beetle when they were consumed, no significant impact upon the beetle was observed. Subsequent investigation showed that in choice tests (Exposure), starved and partially satiated female beetles avoided Bt-fed susceptible prey, but not Bt-fed resistant prey. However, in the rare cases when starved females initially selected Bt-fed susceptible prey, they rapidly rejected them after beginning to feed. This prey type was shown to provide sufficient nutrition to support reproduction in the bioassay suggesting that Bt-fed susceptible prey is acceptable in the absence of alternative prey, however adults possess a discrimination ability based on prey quality. These results suggest that the direct effects of Bt-producing canola on predator life history was minimal, and that predators’ behavioural preferences may mitigate negative indirect effects of reduced quality of prey caused by consumption of Bt-producing plants. The results presented here therefore suggest that cultivation of Bt canola may lead to conservation of non-target predatory and scavenging organisms beneficial in pest control, such as carabids, and may therefore provide more sustainable agricultural systems than current practices. In addition, minimal impacts on beneficial carabids in agro-ecosystems suggest that Bt canola crops are likely to be compatible with integrated pest management (IPM) systems.     

The life histories and population dynamics of two carabid species on a Dutch heathland

Summary We deal with the causes of the synchronously fluctuating numbers of subpopulations of the carabid species Calathus melanocephalus as compared with the asynchronously fluctuating numbers of subpopulations of the carabid Pterostichus versicolor. Both species continuously occupy a large heath area, Dwingelder Veld (1600 ha), in The Netherlands, and are studied there in the same localities with the same methods. Of the adults of C. melanocephalus, 90% do not cover more than 2 ha during the entire reproductive season, while 90% of adults of P. versicolor cover no more than 12 ha. In C. melanocephalus egg production in the field is usually similar to that under optimal feeding conditions in the laboratory, but in P. versicolor egg production seems to be much lower in the field. In the field 70–80% of the eggs most probably are killed by eelworms, followed by more than 90% mortality among the remaining larvae. Comparing mortality of developmental stages in laboratory experiments with that in field experiments in enclosures, it appears that mortality of larvae is not density-dependent, even when density in the experiments is much higher than it ever is in the field. Larval mortality mainly results from the poor ability of the larvae to find prey, even when in field experiments prey density is increased far above natural densities. We discuss why these poor prey-finding abilities are not improved by natural selection. In the spring breeder P. versicolor differences between localities both in abiotic factors, soil moisture and surface temperature, and biotic factors, reactions of prey species to abiotic factors, in spring and summer when the larvae are maturing contribute to the asynchronous fluctuations of numbers between subpopulations. In the autumn breeder C. melanocephalus possible differences in biotic factors between sites are outnumbered by the effects of winters with a higher or lower than normal amount of precipitation respectively. During a wet winter mortality among the larvae is much higher than during a dry winter. As these winter conditions are similar over large areas (many km²) the fluctuations of numbers between subpopulations are synchronous.Communication No. 443 of The Biological Station, Wijster     

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.     

The effects of prey size on diet differentiation of seven passerine species at two spring stopover sites

Prey size was evaluated for seven passerine trans-Saharan migrant species at two spring stopover sites in Sardinia, Italy. The species considered were Pied Flycatcher Ficedula hypoleuca, Spotted Flycatcher Muscicapa striata, Redstart Phoenicurus phoenicurus, Garden Warbler Sylvia borin, Whitethroat Sylvia communis, Willow Warbler Phylloscopus trochilus and Wood Warbler Phylloscopus sibilatrix. The analysis was made for three prominent prey types: beetles (Coleoptera), ants (Hymenoptera: Formicidae) and “flying prey” (wasps and bees [Hymenoptera, excluding ants] and flies, Diptera, pooled), The prey size distribution in the diets of some species was very similar to that caught in our insect traps, showing that our estimates of availability are, at least partially, relevant. On the whole, diets deviated from food available in a species consistent way. The size distributions of “flying prey” differed between the two sites but were almost constant in the birds' diets. In contrast, size distributions of available beetles and ants at the sites were similar but were different in the birds' diets. Different feeding behaviour of the birds, in terms of physiological constraints during migratory stopovers, is discussed. Paired species comparisons show that the diets of most species differ significantly in the distribution of the size classes of at least some prey types. The conclusions drawn from the comparisons of the size distributions of all prey items collected from a bird species in one season are very similar to the conclusions drawn from comparisons based on the presence/absence of a size class per faecal sample. We compared the similarities of diet with and without using information on prey size. With the exception of the Pied Flycatcher and the Redstart at one of the study sites, size information did not add to diet segregation. Certain size classes within prey types tended to be common in the diet of these migrant passerines. However, specialization on certain size classes within broad taxonomic categories was not evident.     

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

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

Soil temperature and flooding effects on two species of citrus

Summary Rough lemon (Citrus jambhiri Lush.) and sour orange (C. aurantium L.) seedlings were grown at constant soil temperatures of 16, 24, and 33 C for 3 months. Shoot and root growth of rough lemon was greatest at 33 C while growth of sour orange was greatest at 24 C. There were no significant effects of soil temperature on shoot: root ratio, leaf water potential or stomatal conductance. The hydraulic conductivity of intact root systems of both species was highest when seedlings were grown at 16 C. Thus, acclimation through greater root conductivity at low soil temperature may have compensated for decreased root growth at 16 C and negated effects of soil temperature on plant water relations. Half the plants growing at each soil temperature were subsequently flooded. Within 1 week, the soil redox potential (Eh) dropped below zero mV, reaching a minimum Eh of –250mV after 3 weeks of flooded conditions. Flooded plants exhibited lower root conductivity, a cessation of shoot growth, lower leaf water potentials, lower stomatal conductances, and visual sloughing of fibrous roots. Decreases in root conductivity in response to flooding were large enough to account for the observed decreases in stomatal conductance.Florida Agricultural Experiment Stations Journal Series No. 4080.     

Asynchronous temporal variation among sites in condition of two carabid species

Abstract.  1. It is generally assumed that generalist predatory carabids are food limited, and the degree of food limitation can vary in space and time, but there are only a few studies that investigate whether or not temporal variation in insect condition is synchronised among sites.
2. In this paper temporal and spatial variation in four fitness-related traits – body mass corrected for size, body size, fat content, and fecundity – of two carabid species, Pterostichus cupreus (L.) and Pterostichus melanarius (Ill.), differing in overwintering strategy, lifespan, diet, and habitat breadth were studied.
3. Interactions between temporal and spatial variation for the fitness-related traits were found for both species. Temporal changes in food limitation were not synchronised among sites, and fitness-related traits were not generally higher on some sites than on others.
4. There was an effect of study year on body mass, fat content, and fecundity for both species, suggesting that temporal environmental variation over large areas is more important than the spatial environmental variation within a year.
5. Within a time period, most different fitness-related traits, except body size, were positively correlated to each other among individuals of the same species. The fitness-related traits of females and males of the same species were generally positively correlated. However, despite the fact that the food resources of the two species overlap, there was no correlation in fitness-related traits between the two species.     

Patch size determines the strength of edge effects on carabid beetle assemblages in urban remnant forests

Habitat fragmentation due to urbanization is increasing rapidly worldwide. Although patch area and edge effect are both important determinants of species diversity and the number of individuals in fragmented landscapes, studies that tested interaction between two effects were limited. Here we examined the interaction between area and edge effects on species richness and the number of individuals of carabids in highly fragmented forests in Tokyo, central Japan. We surveyed carabids in each of 26 forest patches (1.1–121.6 ha) using pitfall traps set in both edge and interior zones. First, we correlated the edge-to-interior differences of both species richness and the number of individuals with patch area. Second, we examined the interaction between patch area and distance to the edge on species richness and the number of individuals using generalized linear models. We found a significant decrease in carabid species richness and the number of individuals in edge zones. The edge-to-interior differences in both species richness and the number of individuals were positively correlated with patch area. Model selection revealed the evident interaction effects between patch area and distance to the edge: higher number of individuals was predicted in only large interior zones. Our results indicated that carabid beetle assemblages were influenced by the interaction between area and edge effects. Thus, in urban areas where small forest remnants dominate, circularizing the shape of the forest patches to maximize the core areas may be the most feasible and realistic means to preserve biodiversity.     

Feeding responses of carabid beetles to dimethoate-contaminated prey

Abstract 1 The feeding responses of Pterostichus madidus Fab., P. melanarius Illiger and Nebria brevicollis Fab. (Coleoptera: Carabidae) to dimethoate‐contaminated prey were investigated in ‘no‐choice’ and ‘choice’ feeding tests. 2 In the no‐choice tests, starved beetles were presented with aphid prey treated with four concentrations of dimethoate. In the choice tests, treated and untreated prey were presented together and the feeding preferences of the starved beetles observed. 3 No avoidance or rejection behaviour was seen in any of the carabids in either of the tests, i.e. no discrimination of the treated and untreated prey was observed. 4 Sufficient dimethoate was consumed with the aphid prey to cause significant mortality levels in the carabids. 5 The concentrations of dimethoate used in these experiments are comparable to field exposure, so carabids feeding in treated fields and field margins could potentially suffer lethal effects via the indirect exposure route of consuming contaminated prey.     

Defensive secretions of larvae of a carabid beetle

Secretions of an eversible gland on the metathorax of larvae of Chlaenius cordicollis Kirby (Coleoptera: Carabidae) are investigated by headspace analysis using solid phase microextraction followed by gas chromatography‐mass spectrometry (GC‐MS). Larvae from Manitoba, Canada and Pennsylvania, U.S.A., are sampled. Nine presumed defensive compounds are detected when the gland is everted, and this represents the first characterization of defensive secretions of larvae of a carabid beetle. With the exception of a single component (2‐methoxy‐4‐methylphenol), these compounds are distinct from those found in the defensive secretion of adult C. cordicollis. However, seven are more oxidized versions of the alkylphenolic compounds secreted by adult beetles: three hydroquinones (hydroquinone, methylhydroquinone and 2,3‐dimethylhydroquinone) and four quinones (p‐benzoquinone, toluquinone, 2,3‐dimethylquinone and ethyl‐p‐benzoquinone). An additional alkoxyphenol (2‐methoxy‐4‐ethylphenol) is also detected. Two patterns of composition are observed: in one, p‐benzoquinone and hydroquinone are undetectable and the ratio of toluquinone : 2,3‐dimethylquinone is 1 : 4.6 ± 0.6 (mean ± SE); in the other, all nine compounds are detectable and the ratio of toluquinone : 2,3‐dimethylquinone is 1 : 1.0 ± 0.2. These differences in pattern do not appear to be related to geographical source, sex or age of the larvae.     

Island biogeography of temporary wetland carabid beetle communities

Aim The study tests if island biogeography is applicable to invertebrate communities of habitat islands in the agricultural landscape that are not fragments of formerly larger habitats. Location Thirty temporary wetlands in the agricultural landscape of northeast Germany. Methods The composition and species richness of carabid beetle communities was analysed. Habitat area, isolation, the density of temporary wetlands in the landscape, land‐use intensity and the maximum duration of flooding were recorded as independent variables. Overall species richness and wetland species richness were studied in independent regression analyses. The community composition was analysed by means of a Canonical Correspondence Analysis (CCA). A partial CCA was used to analyse the effect of the distance to the edge of the field after removing impacts of other independent variables. Results The area of the habitats and various measures of isolation (mean distances = 81–240 m) did not influence species richness or wetland species richness. The community composition was mainly determined by the land‐use intensity, habitat area did not have significant effects, and the distance to the edge of the field was the only effective isolation parameter. Short‐winged species were more often affected by the distance to the edge of the field than full‐winged species. Main conclusion There is evidence that the distances between the wetlands do not provide an effective barrier to the species dispersal and, therefore, metapopulation structures including subpopulations of multiple temporary wetlands might counteract local area effects on subpopulations. Short‐winged species, however, might be more affected by isolation than full‐winged species. As carabid beetle community structure in most early successional habitats is similar, these results may be representative of many agricultural landscape habitats. Nature conservancy concepts that aim to increase habitat area and habitat connectivity have successfully been applied to fragmented late‐successional habitats. The present study indicates that such concepts do not necessarily result in higher diversity or larger populations in early successional habitats.