Seasonal aspects of life cycles in Calathus melanocephalus and C. micropterus (Coleoptera, Carabidae) in the northern taiga

The seasonal dynamics of the activity and the demographic structure of Calathus melanocephalus and C. micropterus populations were studied in the northern taiga of Arkhangelsk Province. The period of adult activity lasts from early June to mid-or late September with the maximum surface activity observed in the middle of summer. In C. melanocephalus, mature individuals are recorded from early June to early September, and in C. micropterus, during the entire season. The data on seasonal changes of the demographic structure of the population demonstrate that in the northern taiga, these species probably possess a biennial life cycle with summer reproduction. The geographical variability of the demographic structure of populations and of life cycles of two species of the genus Calathus was demonstrated. Northwards from the south, the period of activity decreases and the period of reproduction increases and shifts from autumn to the middle of summer. A hypothesis was formulated, according to which in the species studied the annual life cycle in the central part of the range is replaced by the biennial cycle in the north.     

Fecundity in relation to wing-morph of three closely related species of the melanocephalus group of the genus Calathus (Coleoptera: Carabidae)

Summary In two successive years the fecundity of the carabid beetles Calathus (Neocalathus) cinctus, C. (N.) melanocephalus and C. (N.) mollis was studied in relation to wing-morph and temperature. Differences were found between the three species in both egg production and timing and length of the oviposition period. In all species the fecundity of laboratory bred beetles was significantly higher than that of females collected in the field. Long-winged females of both cinctus and melanocephalus had significantly higher egg production than short-winged females, and they also tended to produce eggs over a longer period. In mollis only the fecundity of the long-winged morph was established. The observed lower relative fitness of the short-winged morph in both cinctus and melanocephalus contradicts the supposed increase of the frequency of this morph in ageing, more or less isolated, populations of these species. The loss of long-winged genotypes, resulting from flight activities, is considered the most plausible cause of the increase of short-winged beetles in ageing populations. The higher fecundity of macropterous females makes them especially suited for (re)establishing populations.Communication No. 429 of the Biological Station WijsterPresent address and address for offprint requests: Kortenburg 31, NL-6871 ND Rentum     

Prey selection and foraging behaviour by Pterostichus cupreus L. (Col., Carabidae) under laboratory conditions

Abstract: Several authors have indicated that carabid beetles offer potential as predators of insect pests on arable crops, but this potential is reduced by their limited ability to climb the crop plants. In the current investigation an initial laboratory experiment indicated that seven species of carabid beetles commonly found in arable fields in the UK would feed on both aphid and collembolan prey. Pterostichus cupreus L. and Bembidion guttula Fabr. were found to accept both live and dead aphids and collembolans. In further laboratory investigations P. cupreus exhibited greater consumption of the cereal aphid Metopolophium dirhodum Walker than the alternative prey species Heteromurus nitidus , Templeton (Collembola, Entomobryidae) and there was some indication of preference for the aphid prey as these were often consumed first. Significantly more handling time was required for the consumption of M. dirhodum compared to H. nitidus. The treatments where P. cupreus was given a diet of aphids or collembolans prior to the prey choice experiment, did not appear to influence the number or type of prey items consumed.
Using a four-arm olfactometer P. cupreus reacted positively to the volatiles from collembolans, but gave a less clear response to those from the cereal aphid M. dirhodum . In an artificial field arena P. cupreus readily climbed barley plants in search of M. dirhodum , perhaps as a response to volatile emissions, but this behaviour changed when H. nitidus were introduced onto the substrate of the arena. Although the current results indicated that P. cupreus has potential for the control of cereal aphids, the presence of alternative prey appeared to reduce this potential and suggest that the carabid is an opportunistic feeder.     

Changes in ground beetle diversity and community composition in age structured forests (Coleoptera, Carabidae)

We examined diversity, community composition, and wing-state of Carabidae as a function of forest age in Piedmont North Carolina. Carabidae were collected monthly from 396 pitfall traps (12×33 sites) from March 2009 through February 2010, representing 5 forest age classes approximately 0, 10, 50, 85, and 150 years old. A total of 2,568 individuals, representing 30 genera and 63 species, were collected. Carabid species diversity, as estimated by six diversity indices, was significantly different between the oldest and youngest forest age classes for four of the six indices. Most carabid species were habitat generalists, occurring in all or most of the forest age classes. Carabid species composition varied across forest age classes. Seventeen carabid species were identified as potential candidates for ecological indicators of forest age. Non-metric multidimensional scaling (NMDS) showed separation among forest age classes in terms of carabid beetle community composition. The proportion of individuals capable of flight decreased significantly with forest age.     

Humidity responses and water balance of riparian species of Bembidiini (Col., Carabidae)

Abstract. 1. The humidity responses of Asaphidion pallipes (Duftschmid) and Bembidion spp. were studied at various temperatures in choice chambers.
2. Several species initially showed a hygropositive or indifferent response. Other species had a hygronegative reaction which later changed to a hygropositive one. The duration of the hygronegative reaction varied with species, age, temperature and relative humidity. B.litorale (Olivier) and B.semipunctatum (Donovan) change response when they have lost 8–9% of their initial body weight as water.
3. The survival time of species of Bembidion under dry conditions was short compared with that reported for many other carabid beetles. Newly emerged adults of B.argenteolum Ahrens that had been kept away from sand for 3 weeks had lower transpiration rates than any other species. Young adult Bembidion had lower transpiration rates than those in the breeding phase, larvae and dead adults. Transpiration rate appeared to be inversely proportional to the relative humidity whereas there was a non-linear increase in weight loss with increasing temperature.
4. The species gain most water by drinking and/or from food.
5. There was no correlation between humidity response and transpiration rate or ability to tolerate dry conditions of the species. There was, however, a correlation between behaviour/habitat affinity and humidity response. Diurnal species, living in sandy or silty, half shady-exposed habitats had an initial hygronegative response, whereas those which hide among gravel/stones, leaves or vegetation were indifferent or hygropositive.     

Molecular systematics and evolutionary history of the genus Carabus (Col. Carabidae)

Despite the number of evolutionary, ecological and conservation studies that are conducted on Carabus, the global evolutionary history of the genus remains poorly understood. Here, we analysed 7.5 kilobases of DNA sequence data (six mitochondrial and four nuclear genes) from a worldwide sample of 45% of the known subgenera (99 species and 14 subspecies). We compared the nuclear and mitochondrial phylogenies obtained from Maximum likelihood and Bayesian analyses through topological tests of congruence and dating analyses. Our results mostly corroborate the monophyly of the morphological subgroups of Carabus. However, current morphological and molecular data appear unable to accurately infer the deep branchings within the genus. We show that Carabus originated ca. 16.7-25.1Ma, approximately 25Myr later than previously estimated. Major groups of Carabus are subdivided into clades that diverged from each other in a relatively short period of time around 10Ma (6.6-14.8). This time frame suggests that the present-day distribution of Carabus subgroups may be explained by isolation resulting from Eurasian forest fragmentation brought on by Miocene climate changes and by mountain orogenesis. Finally, we highlight several conflicts between mitochondrial and nuclear topologies that may be explained by mitochondrial introgression.     

The life cycles and age of Carabus glabratus Paykull and C.problematicus Herbst (Col.: Carabidae) on moorland in northern England

Abstract. 1. The life cycles of C.glabratus and C.problematicus on moorland in northern England were studied in 1967. C.glabratus was a spring/summer breeder whilst C.problematicus was a late summer/autumn breeder and both had biennial life cycles.
2. Mandible wear and reproductive condition were used to indicate the age of the adults; some females lived for at least 2 years and probably three, with a distinct regression of their ovaries between each breeding season.
3. Mandible wear affected their ability to feed and thus build up fat reserves or develop eggs. Individuals with extreme mandible wear may die of starvation.     

Studies on predation of Prostephanus truncatus (Horn) (Col., Bostrichidae) and Sitophilus zeamais Mots. (Col., Curculionidae) at different densities on maize by Teretriosoma nigrescens Lewis (Col., Histeridae)

Laboratory experiments using whole cobs were conducted to examine the effect of varying densities of the larger grain borer, Prostephanus truncatus and the maize weevil, Sitophilus zeamais on rate of population increase by the histerid beetle, Teretriosoma nigrescens , a predator primarily of P. truncatus . Densities of all species of insects were determined at the end of the experiment, and an electrophoretic analysis of gut content was conducted on larval and adult T. nigrescens sampled during the experiments. Results indicated that T. nigrescens has a strong preference for P. truncatus and densities of T. nigrescens were associated only with densities of P. truncatus . The maize weevil played little role as an alternative prey or in interfering with T. nigrescens reproduction.     

Satiation time and appetite revival of Coccinella septempunctata L. (Col., Coccinellidae), a predator of Lipaphis erysimi Kalt. (Hom., Aphididae)

After a 24 h period of starvation the Coccinella septempunctata grub satiated in 85 min after eating 31.57 ± 7.12 prey (aphids). After 6, 12, 18, 24, 30 and 36 h of prey deprivation, the grub consumed 14.31 ± 5.67, 22.05 ± 5.39, 27.10 ± 6.49, 31.57 + 7.12, 33.76 + 6.52, and 36.98 ± 8.56 prey, respectively. The voluntary intake (appetite) shows a sigmoid relationship with the time since the last meal, with an upper limit of 44.30 prey irrespective of the prey deprivation period. As maximum appetite was attained after 24–36 h of deprivation, it is recommended that C. septempunctata grubs should be starved for 24 h prior to release in the biological control programme of Lipaphis erysimi.     

Food preferences and the value of animal food for the carabid beetle Amara similata (Gyll.) (Col., Carabidae)

Abstract:  Several studies have shown that the mainly granivorous carabid beetles, e.g. Amara spp., include animal food in their diet to a considerable extent. We therefore hypothesized that the performance of these beetles would be enhanced by dietary mixing including both seeds and animal food. In order to test this, we conducted laboratory feeding experiments with adults and larvae of Amara similata . Both adults and larvae were subjected to different diet treatments including: seeds, houseflies, grasshoppers, earthworms, slugs and snails in pure and mixed diets. Larval survival, development time, pupal and teneral weights were used as indicators of food quality for the larvae. For the adult beetles, mass change was used as an indicator of food quality. We found seeds to be high-quality food, while all pure animal diets were of low quality for both adults and larvae. Animal foods added to the seed diet had both positive and negative effects. A mixed diet of all foods enhanced the mass gain of adults compared with the seed diet, but reduced larval performance dramatically. Earthworms and grasshoppers added to seeds increased the pupal and teneral weights, while reduced larval survival. Thus, A. similata is omnivorous with a mainly granivorous feeding habit. It may gain benefits on some fitness parameters and incur costs on others from preying or scavenging on animal food. Therefore, the overall fitness consequences of a mixed seed-animal diet are uncertain.     

Entomopathogenic fungi in predatory beetles (Col.: Carabidae andStaphylinidae) from agricultural fields

Prevalence of entomopathogenic fungi was studied in overwintering ground beetles (Col.: Carabidae) and rove beetles (Col.: Staphylinidae) collected from fields of lucerne, white cabbage and white cabbage undersown with white clover. In general infection levels in adult ground beetles and rove beetles were low (Carabidae: max. 7.6%,Staphylinidae: max. 7.0%). In comparison, prevalence of entomopathogenic fungi in carabid larvae was high (19–50%). At one study site an epizootic ofBeauveria bassiana was observed, infecting 67% of the staphylinidAnotylus rugosus and 37% of the staphylinidGyrohypnus angustatus. Beauveria bassiana was the predominant fungus isolated from ground beetles and rove beetles from all studied sites. Other fungal species included the hyphomycetesMetarhizium anisopliae, Paecilomyces farinosus andVerticillium lecanii as well asZoophthora radicans andZoophthora philonthi (Zygomycetes: Entomophthorales). Two individuals ofAnotylus rugosus were found to have a dual infection ofZoophthora philonthi andBeauveria bassiana.

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Résumé La présence de champignons entomopathogènes a été recherchée chez des carabes (Col.: Carabidae) et des staphylins (Col.: Staphylinidae) récoltés dans des champs de luzerne, de chou et d'une culture mélangée chou-trèfle blanc. Chez les adultes, le taux d'infection dans les deux groupes d'insectes est en général très faible (Carabidae: max. 7,6%, Staphylinidae: max. 7%). Chez les larves de carabes par contre, le taux d'infection par les champignons est élevé (19–50%). Sur l'un des sites de l'étude, une épizootie àBeauveria bassiana a été observée, l'infection portant sur 67% des individus récoltés appartenant à l'espèceAnotylus rugosus et 37% desGyrohypnus angustatus. B. bassiana est le champignon prédominant isolé à partir des coléoptères de tous les sites étudiés. D'autres espèces fongiques ont été relevées: ce sont les hyphomycètesMetarhizium anisopliae, Paecilomyces farinosus etVerticillium lecanii ainsi queZoophthora radicans etZoophthora philonthi. Deux individus d'A. rugosus ont été retrouvés infectés à la fois parErynia etB. bassiana.

     

Adult feeding affects fecundity of the predator,Calosoma sycophanta (Col.: Carabidae)

Calosoma sycophanta L. adults were fed either gypsy moth (Lymantria dispar L.) larvae or split grapes for set periods of time while their reproduction was monitored. Few female beetles reproduced unless fed gypsy moth larvae during the first week after they ended hibernation. Even females initially fed grapes that were later fed larvae had reduced reproduction. The implications these results have for relationships between beetle and gypsy moth populations are discussed.     

Microsatellites and allozymes as the genetic memory of habitat fragmentation and defragmentation in populations of the ground beetle Carabus auronitens (Col., Carabidae)

Aim This paper has three aims: (1) to reconstruct the population history of a flightless silvicolous (forest) ground beetle in a region where strong anthropogenic activity has altered the connectivity of the landscape; (2) to estimate the effects of both contemporary and historical landscape structure on the genetics of populations; and (3) to investigate the reasons for clinal variation in one gene locus found in an earlier study carried out in the same geographical location. Location Münster (Westphalia), north‐west Germany. Methods We investigated 26 populations of the ground beetle Carabus auronitens Fabricius, 1792 by analysing seven polymorphic microsatellite loci and an allozyme locus. Samples of at least 16 individuals per site were studied. These were obtained from dry pitfall traps placed at 23 sites and from three additional (refuge) populations. We used regression and correlation analyses to investigate the effects of both historical and contemporary landscape structure on the allele frequency distributions of the investigated loci. Spatial autocorrelation analysis was used to study possible clinal variations, and admixture rates were calculated in order to assess the genetic influence of populations from possible refuges. Possible reasons for the development of the cline were examined using simulation models. Results The allele frequency distributions at the investigated loci could not be explained by selection or isolation by distance. We found clinal variation in 50% of the investigated loci and our simulations indicated that this was unlikely to have developed by chance. Admixture rates show significant influences of the investigated refuge populations on the populations under study. Main conclusions The findings strongly suggest that the clinal variation is secondary. It results from recolonization of the area by C. auronitens from multiple refuges after anthropogenic landscape changes caused forest fragmentation and led to species isolation.     

Prolonged development time of the bark beetle predator Thanasimus formicarius (Col.: Cleridae) in relation to its prey species Tomicus piniperda (L.) and Ips typographus (L.) (Col.: Scolytidae)

1 The generation time of the bark beetle predator Thanasimus formicarius (L.) (Col.: Cleridae) was found to be predominantly two years both in the field and in rearing experiments conducted with two of its main prey species, the pine shoot beetle Tomicus piniperda (L.) and the spruce bark beetle Ips typographus (L.) (Col.: Scolytidae). 2 Emergence of T. formicarius adults in the first summer was only observed in one of the two rearing experiments, and these individuals represented only 6% of that generation. 3 All individuals not emerging as adults in the first summer remained as larvae in their pupal chambers until the second summer. Pupae were found starting around mid-June, and adults (in pupal chambers) were found from late July through to the end of August. 4 Newly emerged adults had to feed in order to survive hibernation. 5 The existence of T. formicarius races, specialized on certain bark beetle species and with phenologies matching their hosts, could not be demonstrated. After hibernation there was no difference in feeding activity, timing of egg-laying or proportion of egg-laying females between the T. formicarius adults reared as larvae on T. piniperda (flight period in April) and those reared as larvae on I. typographus (main flight period generally starting in late May or early June) when exposed to a temperature and day-length typical of the early spring conditions prevailing during the flight period of T. piniperda. 6 T. formicarius was parasitized by Enclisis vindex (Tschek) (Hym.: Ichneumonidae) in the pupal chamber. 7 The importance of these findings for the population dynamics of bark beetles is discussed.