Notes on two species of the subgenus Lyrothorax Chaudoir (Coleoptera: Carabidae; genus Pterostichus), Pterostichus amagisanus Tanaka and Ishida and Pterostichus fujitai Tanaka and Ishida

Two species of the subgenus Lyrothorax Chaudoir (Coleoptera: Carabidae; genus Pterostichus ), Pterostichus amagisanus Tanaka and Ishida and Pterostichus fujitai Tanaka and Ishida, were revised based on the male endophallus (inner sac everted from aedeagus). P. amagisanus was newly recorded based on a single male from Kyushu, southwest Japan, far from its known distribution (Honshu; the Fuji-Hakone-Izu volcano area), although additional materials are necessary to confirm this record. Despite a highly disjunct distribution, no conspicuous difference was recognized in either external or genital characters between the materials from Honshu and Kyushu. The nominal species P. fujitai was separated into two species, P. fujitai (Honshu) and Pterostichus eoyoritomus sp. nov. (Shikoku; type locality: Mount Jingayama); these two species have some significant differences in the endophallic structures. Character states in male genitalia suggest a sister relationship between P. eoyoritomus sp. nov. and Pterostichus yoritomus Bates.     

Low-vacuum electron microscopy of carabid chemoreceptors: a new tool for the identification of live and valuable museum specimens

High-vacuum scanning electron microscopy, following coating of specimens with gold, produces high quality images that have proved invaluable for the study of insect sensilla. Unfortunately, the technique is essentially destructive, and cannot be used on live or valuable museum specimens. In particular, high-vacuum scanning usually causes the collapse of the tips of the palps, interfering with any examination of the sensilla in this area. A new low-vacuum technique is described that avoids these problems. Insect cuticle does not need to be coated with gold, thus avoiding damage to important specimens. Examples are given of scans of the palp tips of live carabid beetles, anaesthetised with CO₂. It was shown that the technique could consistently display these tips in their natural convex state. In all, four types of sensilla were identified by low-vacuum scans of the maxillary palps, and four further types on the terminal segment of the antennae, plus glandular openings. The antennae revealed a type of sensilla that has not previously been described on carabids. These sensilla showed clear structural differences between the two species studied, Pterostichus melanarius Illiger and P. niger Schaller (Coleoptera: Carabidae), and can be used as a diagnostic character for both fresh and dried specimens. The low-vacuum technique can be recommended for examining valuable type specimens without risk of damage.     

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.     

Habitat characteristics of grassland Pterostichus species (Coleoptera, Carabidae)

Abstract.

• 1 Pterostichus species were sampled on ninety-two sites distributed throughout north-east England using pitfall trapping.
• 2 The incidence of each species was related to measured site environmental variables using logistic regression.
• 3 Four species groups were identified on the basis of size. The extent of overlap in habitat types between species in each group varied. Two large species overlapped considerably whilst the smaller species showed different responses to one or more environmental variable.
• 4 Possible causes of the differences in habitat overlap between Pterostichus species are discussed.

     

Multiplex reactions for the molecular detection of predation on pest and nonpest invertebrates in agroecosystems

Species- and group-specific PCR primers were developed to study predation on pest and nonpest invertebrate species by generalist carabid predators in agroecosystems. To ensure the amplification of degraded DNA in predator gut samples, amplicons were designed to be less than 300 bp. Specificity of primers was assessed by cross-amplification against a panel of target and nontarget invertebrate species. The new primers were combined with previously published primers for slugs and collembolla in multiplex reactions to simultaneously screen each predator for the presence of multiple prey. All prey species were detected in a screen of the gut contents of field-caught predators.     

Wing-dimorphism and population expansion of Pterostichus melanarius (Illiger, 1798) at small and large scales in central Alberta, Canada (Coleoptera, Carabidae, Pterostichini)

A study spanning ten years revealed changes in wing-morph ratios corroborating the hypothesis that the wing-dimorphic introduced carabid, Pterostichus melanarius Ill.,is spreading through flight, from the city of Edmonton, Canada and establishing populations in natural aspen forest of more rural areas 45-50 km to the East. Comparison of wing-morph ratios between Pterostichus melanarius and the native wing dimorphic species Agonum retractum LeConte suggests that the spatial variation in ratios for Pterostichus melanarius does not reflect underlying environmental variation, but instead the action of selective forces on this wing-dimorphic species. About ten years after its earliest detection in some rural sites the frequency of macropterous individuals in Pterostichus melanarius has decreased c. five-fold, but it is still above the level seen in European populations in which the two wing-morphs are thought to exist in equilibrium. Pterostichus melanarius is expanding its range in native aspen forest much faster than three other introduced species Clivina fossor L.), Carabus granulatus O.F. Müllerand Clivina fossor L also encountered in this study. The two Carabus species are flightless, but Carabus fossor can be dimorphic. Although these four non-native ground beetle species comprise >85% of the carabids collected at sites in urban Edmonton, activity-density of native carabids was similar across the urban-rural gradient, suggesting little direct impact of introduced species on the local abundance of native species. In a second study conducted at a smaller scale near George Lake, Alberta, macropterous individuals of Pterostichus melanarius have penetrated furthest and most rapidly into native aspen forest. Furthermore, the percentage of micropterous individuals has increased markedly in areas first colonized a decade previously. Overall, these studies support the idea that macropterous beetles in wing-d dimorphic species are important vanguards for early colonization of unexploited territory, but that flightless individuals replace the flying morph relatively rapidly once populations are established.     

Microsatellite DNA markers for the ground beetle Pterostichus oblongopunctatus F.

Six microsatellite DNA loci were isolated from the ground beetle Pterostichus oblongopunctatus F. (Coleoptera, Carabidae) to study population structure. Each locus was polymorphic, with the number of alleles ranging from three to six. Observed heterozygosity varied between 0.20 and 0.67. All six loci were tested for amplification in the closely related P. quadrifoveolatus and they were shown to be polymorphic. The primers developed were used in multiplex polymerase chain reactions.     

Prey choice by carabid beetles feeding on an earthworm community analysed using species‐ and lineage‐specific PCR primers

The carabid beetle Pterostichus melanarius is a major natural enemy of pests, such as aphids and slugs in agricultural systems. Earthworms are a dominant non‐pest component of the diet of P. melanarius which help sustain the beetles during periods when the pest population is low or absent. In this study we wanted to test whether this predator exercises prey choice among different earthworm species or ecological groups. High levels of genetic diversity within morphological species of earthworm necessitated the development of primers that were specific not just to species but lineages and sub‐lineages within species as well. Gut samples from beetles were analysed using multiplex‐PCR and fluorescent‐labelled primers. Calibratory feeding trials were undertaken to calculate median detection times for prey DNA following ingestion. Extensive testing demonstrated that the primers were species‐specific, that detection periods were negatively related to amplicon size and that meal size had a highly significant effect on detection periods. Monte Carlo simulations showed that, in general, worms were being predated in proportion to their densities in the field with little evidence of prey choice, other than probable avoidance of the larger, deep‐living species. There was no evidence that epigeic species were being taken preferentially in comparison with endogeic species. There was also no evidence that defensive secretions by Allolobophora chlorotica reduced predation pressure on this species by P. melanarius. We concluded that any management system that increases earthworm densities generally, regardless of component species, is likely to be optimal for increasing numbers of this beneficial beetle predator.     

Taxonomic studies of Pterostichus ishikawai Nemoto (Coleoptera: Carabidae)

A macrocephalic pterostichine carabid, Pterostichus ishikawai Nemoto, endemic to the Korean Peninsula, is taxonomically revised based mainly on the membranous parts of the genitalia (male endophallus and female vagina). Two species are separated from P. ishikawai (type locality: Mount Taebaegsan) and newly described: P. ishikawaioides (type locality: Mount Sobaeksan) and P. jiricola (type locality: Mount Jirisan). Since comparative male genital morphology does not support monophyly for these three species, the two novel species should be treated as distinct species, not subspecies of P. ishikawai.     

A method for rapidly mass laser-marking individually coded ground beetles (Coleoptera: Carabidae) in the field

Abstract.  1. A laser-marking technique is described that may be used to rapidly mass-mark and individually code insects in the field. In this study, ground beetles were marked but the method is applicable to a wide range of organisms with a hard exoskeleton or shell.
2. In a field-scale capture–recapture study conducted in winter wheat, 8266 beetles were coded. Individuals marked in the first week of the trial and recaptured 2 months later retained their codes, which were therefore considered permanent.
3. Assemblages were dominated numerically by Pterostichus cupreus , which formed more than 98% of the total captures. Of the 8046 released P. cupreus , 2269 were recaptured. As an illustration of the use of these data, the POPAN parameterisation of the Jolly–Seber model was used to estimate a whole-field population of 17 237 individuals, equating to a density of 0.46 beetles m⁻².     

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.     

Modelling the dynamic spatio-temporal response of predators to transient prey patches in the field

The spatio-temporal dynamics of two aphid species ( Metopolophium dirhodum and Sitobion avenae ) and a generalist predator ( Pterostichus melanarius ) were observed in a field-scale study using a grid of 256 sampling locations with a 12-m spacing. Using Spatial Analysis by Distance Indices we demonstrate that populations show ephemeral spatial pattern at the field scale. We observed a positive, lagged beetle response to this aphid pattern; conversely, the aphids displayed a negative, lagged response to beetle spatial pattern. Examination of the local structure of the spatio-temporal dynamics revealed a strong response by the beetle population to aphid patches. The temporal structure of spatial associations between the species shows a strong correspondence with those from a conceptual model of predator–prey spatial interaction. The spatially coupled dynamics were sufficiently strong for the predator to have a negative effect on the intrinsic rate of increase of their prey.     

Pitfall trap catches of and aphid predation byPterostichus melanarius andPterostichus madidus in insecticide treated and untreated potatoes

Aphids were countedin situ and carabid beetle populations assessed by pitfall trapping betwen 1983–1985 in potato fields in Scotland treated with demeton-S-methyl (DSM) or untreated.Macrosiphum euphorbiae (Thomas) (Homoptera: Aphididae) was the most abundant aphid. Nineteen species of carabid beetles were trapped but the fauna was dominated byPterostichus melanarius (Ill.) andPterostichus madidus (Fabr.) (Coleoptera: Carabidae). Pitfall trap catches of these carabids were variable but generally lower in sprayed than unsprayed plots for a few days after DSM application and higher a week or two later. Of 1800P. melanarius and 910P. madidus dissected, 14.4 per cent and 30.5 per cent respectively, contained aphid remains. The proportion that had eaten aphids often was higher in sprayed than unsprayed plots immediately after treatment. In additional experiments, aphids treated with DSM fell from plants; four to five per cent of the theoretically applied dose of DSM reached the ground in a closed-canopy potato crop; and no mortality occurred whenPterostichus spp. were exposed to DSM in a field bioassay. It is proposed that observed patterns in trap catch ofPterostichus spp. around times of pesticide application may be due in part to aphids falling to the ground after treatment. Beetles that eat these fallen aphids may be less hungry, less active and less likely to be trapped. Later increases in catches may be attributed to hungry, more active beetles, a consequence of prey removal by the insecticide. Although this hypothesis requires further research, it is clear that the effects of insecticide treatment on carabid populations cannot be measured by pitfall trapping alone.     

Rapid screening of invertebrate predators for multiple prey DNA targets

DNA-based techniques are providing valuable new approaches to tracking predator-prey interactions. The gut contents of invertebrate predators can be analysed using species-specific primers to amplify prey DNA to confirm trophic links. The problem is that each predator needs to be analysed with primers for the tens of potential prey available at a field site, even though the mean number of species detected in each gut may be as few as one or two. Conducting all these PCRs (polymerase chain reactions) is a lengthy process, and effectively precludes the analysis of the hundreds of predators that might be required for a meaningful ecological study. We report a rapid, more sensitive and practical approach. Multiplex PCRs, incorporating fluorescent markers, were found to be effective at amplifying degraded DNA from predators' guts and could amplify mitochondrial DNA fragments from 10+ species simultaneously without 'drop outs'. The combined PCR products were then separated by size on polyacrylamide gels on an ABI377 sequencer. New primers to detect the remains of aphids, earthworms, weevils and molluscs in the guts of carabid predators were developed and characterized. The multiplex-sequencer approach was then applied to field-caught beetles, some of which contained DNA from as many as four different prey at once. The main prey detected in the beetles proved to be earthworms and molluscs, although aphids and weevils were also consumed. The potential of this system for use in food-web research is discussed.     

The significance of facultative scavenging in generalist predator nutrition: detecting decayed prey in the guts of predators using PCR

Gut-content analyses using molecular techniques are an effective approach to quantifying predator-prey interactions. Predation is often assumed but scavenging is an equally likely route by which animal DNA enters the gut of a predator/scavenger. We used PCR (polymerase chain reaction) to detect scavenged material in predator gut homogenates. The rates at which DNA in decaying slugs (Mollusca: Pulmonata) and aphids (Homoptera: Aphididae) became undetectable were estimated. The detectability of DNA from both carrion types in the guts of the generalist predator Pterostichus melanarius (Coleoptera: Carabidae) was then determined. The effects of carrion age and weight, as well as beetle sex, on detection periods, were quantified. Laboratory trials measured prey preference of beetles between live and decaying prey. Further experiments measured, for the first time, feeding by P. melanarius on dead slugs and aphids directly in the field. In both field and laboratory, P. melanarius preferentially fed on dead prey if available, but preference changed as the prey became increasingly decayed. Disappearance rates for slug carrion in wheat fields and grasslands were estimated and P. melanarius was identified as the main scavenger. Comparison of the retention time for dead slugs in the field, with the detection period for decaying slug material in the guts of the predators, showed that PCR-based techniques are not able to distinguish between predated and scavenged food items. This could potentially lead to overestimation of the impact of predation on slugs (and other prey) by carabids. Possible implications of facultative scavenging by invertebrate predators for biocontrol and food-web research are discussed.     

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

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