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

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An experimental study of behavioral mechanisms of relationships between red wood ants and ground beetles

Behavioral mechanisms regulating the competitive relations between the red wood ant Formica aquilonia and the ground beetles Carabus regalis, Pterostichus melanarius, P. magus, P. oblongopunctatus, Harpalus smaragdinus, and Amara nitida were studied by artificially initiated collisions between living insects as well as with the use of imitation models. Members of different functional groups within an ant family (aphid-milkers, hunters, and guards) behaved differently towards beetles. Active ants were shown to respond selectively to different features of the possible competitors, such as coloration, the presence of “appendages” (legs, antennae), body symmetry, rate of movement, and scent. Field and laboratory experiments demonstrated the ability of beetles to avoid collisions with active ants. The scent of anthill material attracted ground beetles, which consumed dead ants. The gained individual experience may allow the beetles to use supplementary forage resources in the territories controlled by ants. The flexible tactical patterns facilitate spatial segregation of ground beetles and ants in the same territory and result in a more complete utilization of food resources.     

Forestry alters foraging efficiency and crop contents of aphid-tending red wood ants, Formica aquilonia

Forest management alters species behaviours, distributions and interactions. To evaluate forestry effects on ant foraging performance, we compared the quality and quantity of honeydew harvested by ants among clear-cuts, middle-aged and mature spruce-dominated stands in boreal forests in Sweden. Honeydew quality was examined using honeydew collected by squeezing the gasters of laden Formica aquilonia workers. We used fifteen laden individuals at each study site (four replicates of each stand age) and analysed honeydew chemical composition with gas chromatography-mass spectroscopy. To compare the quantity of honeydew collected by individual ants, we collected and weighed five ants moving up and five ants moving down each of ten trees at the twelve sites (totally 1200 ants). The concentration of trehalose in honeydew was lower in clear-cuts compared with middle aged and mature stands, and similar trends were shown for sucrose, raffinose and melezitose, indicating poorer honeydew quality on clear cuts. Concentrations of the amino acid serine were higher on clear-cuts. The same trend occurred for glutamine, suggesting that increased N-uptake by the trees after clear cutting is reflected in the honeydew of aphids. Ants in mature stands had larger heads and carried proportionally more honeydew and may therefore be more efficient foragers. Human alternation of habitats through clear-cutting thus affects food quality and worker condition in F. aquilonia. This is the first study to show that honeydew quality is affected by anthropogenic disturbances, likely contributing to the reduction in size and abundance of F. aquilonia workers and mounds after clear cutting.     

Molecular markers allow sibling species identification in red wood ants (Formica rufa group)

Red wood ants (Formica rufa group) constitute a group of species that are considered to be among the most promising bioindicators in forest ecosystems. However, because of their morphological similarity and intraspecific variability, morphological species identification can be difficult. Considerable expertise is necessary to discriminate between the sibling species F. lugubris and F. paralugubris, two species that often live in sympatry in the same Alpine forests. New taxonomic tools providing rapid and reliable species identification are needed. We present a simple and reliable molecular technique based on mtDNA (COI gene) and a restriction enzyme for discriminating between F. lugubris and F. paralugubris. We confirm the validity of this method with a Bayesian analysis based on microsatellites. This new molecular tool represents a clear breakthrough for discriminating between F. lugubris and F. paralugubris and is likely to be helpful in large‐scale biomonitoring.     

Population genetics of Formica aquilonia wood ants in Scotland: the effects of long-term forest fragmentation and recent reforestation

Formica aquilonia wood ants are forest specialists which play a key role in the ecology of forests in Europe. Many of the Scottish populations at the edge of the species distribution range occur in highly fragmented landscapes. We used ten microsatellite loci to study the genetic diversity and structure of populations from two contrasting regions (Inverpolly and the Trossachs) to set the Scottish populations in the context of conspecific populations in mainland Europe. Historically, both study regions have experienced extreme habitat loss and fragmentation over several centuries. Inverpolly has remained fragmented whereas large scale reforestation over the last century has greatly increased the forested area in the Trossachs. Despite the long history of fragmentation, genetic diversity in the Scottish populations was greater than in the populations in mainland Europe. Genetic diversity was similar in the two Scottish regions and no evidence of inbreeding was detected. However, the populations in Inverpolly showed more evidence of genetic bottlenecks, possibly due to more frequent stochastic events such as moorland fires. The ant populations in individual forests were genetically distinct and we detected no contemporary gene flow between forests. The most intensively studied forest where non-native conifer plantations now occupy the matrix between the remaining ancient woodland fragments showed evidence that admixture and gene flow between nests was reducing the past differentiation. This may reflect a dynamic response to the reconnection of previously isolated populations in forest fragments by recent reforestation.     

The relationship between British ground beetles (Coleoptera, Carabidae) and land cover

Aims and methods Ground beetle and satellite‐derived land cover data from 1687 United Kingdom 10 km national grid squares were used to assess the relationship between species pool and cover data in Great Britain using fuzzy classification and constrained ordination. Results Ground beetle species pools classified into nine groups which were related to land cover variables using constrained ordination. There was a strong relationship between upland land cover and three ground beetle groups. Deciduous woodland, coastal and tilled land were associated with three other groups. Three further groups did not appear to be strongly associated with any particular cover, but differed in geographical position. Conclusion The distribution of species pools derived from the British national recording scheme at the 10 km scale was strongly related to satellite‐derived land cover data. There appears to be considerable potential for the use of a synthesis of land cover and ground beetle data in the monitoring of environmental change over a large, countrywide, area.     

Response of the foraging behaviour of red wood ants (Formica rufa group) to exclusion from trees

Abstract 1 Because of the large numbers within a colony and their aggressive nature, red wood ants (Formica rufa group) have a potential to greatly influence the cold‐temperate forest ecosystem. Wood ants are omnivorous and hunt in trees as well as on the forest floor. 2 A field experiment in a mixed forest in central Sweden was carried out to examine (i) the foraging behaviour of wood ants on the forest floor and (ii) the impact of increased numbers of wood ants on the soil fauna. The foraging behaviour of wood ants was manipulated by excluding the ants from their food resources in the tree canopy, with the intention to increase ant activity on the forest floor. To estimate this activity, the number of trees with foraging ants, the numbers of ants going to and from their nests and the prey carried by home‐running wood ants were determined during the summer period. Pitfall traps were placed in the soil to determine effects on mobile soil invertebrates. 3 When excluded from local trees, wood ants searched other trees further away from the nests rather than searching more intensively for prey on the forest floor. By contrast to the initial hypothesis, more soil‐living prey were caught by ants in the control plots than in the plots where the local trees were not accessible to the wood ants. The proportion of soil‐living to tree‐living prey tended to be greater in the control plots. 4 In the treated plots (no access to the trees), wood ants had a negative effect on the activity of Linyphiidae spiders. There was little effect of wood ants on other soil invertebrates. 5 This study suggests that the role of wood ants as top predators in the forest soil food‐web in central Sweden is limited.     

Temperature and rate of seed consumption by ground beetles (Carabidae)

Predation is a key source of seed mortality in many weed species and thus is a part of natural control. In the field, the intensity of seed predation by invertebrates varies during the course of a year. One source of this variation is fluctuations in ambient temperature. Here, the effect of temperature on seed consumption is investigated for the first time, using two abundant carabid seed predators, Pseudoophonus rufipes and Harpalus affinis (Coleoptera: Carabidae), and dandelion (Taraxacum officinale) as a model system. Field collected individuals were sexed, kept at one of six constant temperatures between 10 and 28 °C and provided with a surplus of seed. Seed consumption was recorded over a period of 4 days. Averaged over all the temperatures, the smaller H. affinis consumed 12.2 seeds day^?1 and larger P. rufipes 29 seeds day^?1. On average, females consumed more seeds than males. Seed consumption by both species increased with temperature. In H. affinis the increase was linear and different for males and females. In P. rufipes the consumption was similar in both sexes but curvilinear because there was no further increase in consumption above 20 °C. Assuming a linear relationship between temperature and consumption at up to 20 °C we calculated the temperature at which seed consumption ceased (?0.1 to 0.3 °C in H. affinis and 6.3–6.9 °C in P. rufipes) and the increment in seed consumption per 1 °C increase in temperature above this threshold (0.4–1.0 and 1.5–4.2 seeds individual^?1 day^?1, respectively) for the two species. Thus, it is possible to calculate the average daily consumption of each species over a range of temperatures up to 20 °C.     

Effects of reflective groundcovers on ground beetles (Coleoptera: Carabidae) in red raspberry (Rubus idaeus) cropping systems

Reflective groundcovers are management tools used in temperate regions where light can be a limiting factor to crop productivity. Although their effects on crops have been extensively studied, little is known about their ecological impacts on ground‐dwelling organisms. Theoretically, reflective groundcovers add structural complexity to the system and thus have the potential to create refuges for ground‐dwelling invertebrates. At the same time, groundcovers can create abiotic conditions underneath them that could potentially cause declines in the abundance and richness of ground‐dwelling invertebrates. During the summers of 2006 and 2007, white, woven, reflective polymer groundcovers were placed in two red raspberry (Rubus idaeus L.) cropping systems (organic and conventional) in south‐eastern New Brunswick to assess their effects on ground beetle richness, activity density and overall assemblage structure. A total of 8723 ground beetles belonging to 23 species were collected using pitfall traps. In contrast with the combined effects of cropping systems and sampling years, the use of a groundcover accounted for a small proportion of the variance in trap catches and caused little change in ground beetle richness, activity density and overall assemblage structure. Based on these results, we suggest that reflective groundcovers constitute promising management tools to increase light environment quality in row crops without being detrimental to ground beetle species that may contribute to pest suppression.     

大步甲的分子系统与进化模式

大步甲主要分布于北半球,全世界大约有1000余种。因大步甲后翅退化,不能飞行,其移动能力和扩散范围都受到很大程度的限制,因此,容易产生地理隔离和遗传分化,从而成为研究物种分化和生物多样化的好材料。近年来,通过对大步甲分子系统的详细研究,不仅很大程度上解明了大步甲各分类群间的系统发育关系,而且对大步甲的系统演化过程和形态进化模式也有了较深的理解,获得了很多重要的见解。在此做一个简单的综述。     

Trophic position and seasonal changes in the diet of the red wood ant Formica aquilonia as indicated by stable isotope analysis

1. Red wood ants are among the most numerous generalist predators and strongly affect the composition of arthropod communities in forest ecosystems. However, their trophic position remains poorly understood. Stable isotope analysis was applied to study the trophic position of Formica aquilonia and reveal seasonal changes in its trophic links with both myrmecophilous aphids and other invertebrates in a mixed forest of western Siberia. 2. The δ¹⁵N values of F. aquilonia exceeded those of herbivores and aphids by approximately 3.5‰. Despite obligate trophobiotic relationships with aphids, F. aquilonia occupied the trophic position of first‐order predator. The higher content of ¹³C in the worker ants, compared with members of grazing food chains, was explained by their consumption of ¹³C‐enriched aphid honeydew. 3. Myrmecophilous tree‐dwelling aphids were enriched in ¹³C and ¹⁵N relative to grass‐inhabiting species, and the honeydew of tree‐dwelling aphids had higher δ¹³C values than those of the honeydew of grass‐inhabiting aphids. 4. The decrease in δ¹³C values of the worker ants from spring and summer to autumn apparently reflected the transition from the collection of tree sap and feeding on the aphid honeydew from trees with high ¹³C content in the spring and early summer to a more diverse liquid diet in late summer, which included ¹³C‐depleted honeydew of aphids from herbs. 5. The prevalence of the ¹⁵N‐depleted aphid honeydew in the ants' diet in the second half of the summer is discussed as one possible explanation for the seasonal decline in δ¹⁵N values of the worker ants.     

Winter quarters of wetland ground beetles (Coleoptera,Carabidae) in South Scandinavia

Sites at various distances from eutrophic and mesotrophic lakes and fens on the island Öland in southern Sweden were systematically surveyed in summer (May–August) and autumn (late September–October) to elucidate the hibernation sites of wetland carabid beetles. Thirty-five of 47 wetland species were found in their winter quarters. In areas ≤50 m from the lakes and fens, the observed as well as the estimated number of hibernating wetland species was higher in mesic to dry sites than in wet or moist open sites. Very few overwintering beetles were detected far (≥120 m) from the water. There was no significant difference in the frequencies of macropterous and dimorphic (brachypterous) species and individuals hibernating close to, and more distantly from, water. Individuals of small species hibernated closer to water than larger ones. At least in Scandinavia, there is no convincing evidence that wetland species hibernate far away from water, migrating by flight. A considerable number of the species that were collected in their winter quarters are rare and red- listed in at least one of the Scandinavian countries. Winter quarters situated in mesic and dry meadows and woodland close to lakes and fens had the highest number of red-listed species. Therefore, it is of fundamental importance to protect such sites. Threats to the integrity of the current habitats are logging and various transformations of meadows, e.g. digging and removal of large objects such as stones. Cessation of grazing by cattle may also have a negative effect.     

Typology of life cycles of ground beetles (Coleoptera,Carabidae) in Western Palaearctic

An original classification of the life cycles of ground beetles from Western Palaearctic is proposed. The classification is based on a combination of five criteria: duration, number of generations per season, phenology of reproduction, stability, and repeatability of reproduction. According to the individual lifespan, the cycles are subdivided into annual and biennial ones. The annual life cycles may be uni-and bivoltine, whereas biennial ones are always univoltine. By the time of reproduction, winter-spring, spring, spring-summer, early summer, summer, late summer, summer-autumnal, autumnal, autumn-winter, winter, and aseasonal species are distinguished. The biennial and bivoltine cycles may be of both facultative and obligate nature. Species living only one season and having a continuous reproductive period are designated as semelparous, while those breeding during two or more years or having several distinct periods of reproduction in one season, as iteroparous. By now, 30 variants of life cycles in Carabidae from western Palaearctic have been established. Repeated similarly directed modifications of the life cycle may produce essentially different seasonal rhythms in some individuals. In this case, two subpopulation groups usually appear within the population. Under the most unfavorable conditions, these groups become practically isolated and hibernate at different ontogenetic stages. The individual development in each of these groups takes two years with the same seasonal rhythm. Among the types considered, only obligate-bivoltine life cycles are always polyvariant, but annual univoltine and obligate-biennial ones are always univariant. The facultative-bivoltine and biennial life cycles may be realized as uni-and polyvariant ones, depending on the environmental conditions.     

Characterization of microsatellite loci in red wood ants Formica (s. str.) spp. and the related genus Polyergus

Ants are interesting subjects for studies of evolution of altruism. We developed 13 microsatellite loci in a red wood ant Formica (s. str.) yessensis from random amplified polymorphic DNA fragments to study genetic structure within populations and colonies. Five loci bore two to five alleles in both F. (s. str.) yessensis and F. (s. str.) truncorum and two were also polymorphic in a related species, Polyergus samurai. Results suggest that the loci will be useful in evolutionary studies on Formica and Polyergus species.     

Diel activity patterns of South Scandinavian high mountain ground beetles (Coleoptera, Carabidae)

Fourteen species of carabids were studied throughout the snowfree season in outdoor arenas above the timberline at Finse, South Norway (60°36'N, 7°30'E). Ten species were nocturnal. Onset of nocturnal activity occurred after termination of twilight with peak activity before midnight, but in June, with less than 1 h of darkness, activity was extended into evening twilight, the peak being around midnight. Linear regressions between ground surface temperature and nocturnal activity predicted zero activity in the range −2.0 to0.8°C, mean −0.5°C, indicating that activity is possible on most nights throughout the snowfree season. Two diurnal species had activity threshold temperatures at 4 and 8°C, respectively, the former with a peak at 16°C and an upper threshold at 28°C.
The proportion of nocturnal species in the study area was significantly greater than in lowland open fields. Large and common species were mainly nocturnal, while small and rare species were mainly diurnal. It is suggested that nocturnalism in the South Scandinavian high mountains is a response to avoid bird predation and desiccation.     

The impact of two insecticides on predatory ground beetles (Carabidae) in newly-sown grass

Field scale experiments carried out over three years showed that chlorpyrifos reduced the numbers of several carabid beetle species caught in pitfall traps. Fonofos seed treatment was also shown to affect selected species. Consideration of the biology of individuals of groups of species provided explanations of why some species such as Nebria brevicollis, Notiophilus spp., Bembidion spp., and Trechus quadristriatus were severely affected by one or other of the chemicals, whereas other species (e.g. Amara spp., Harpalus spp.) were able to survive or recolonise pastures shortly after chemical application. Measurements of elytral and pronotal widths and counts of mature eggs in N. brevicollis, which was trapped in reduced numbers over a long period, were made and provided some information about the mode of action of these chemicals.     

An unexpected clade of South American ground beetles (Coleoptera,Carabidae, Bembidion)

Phylogenetic relationships of the Antiperyphanes Complex of the genus Bembidion are inferred using DNA sequences from seven genes (two nuclear ribosomal, four nuclear protein coding, and one mitochondrial protein coding). Redefined subgenera within the complex are each well-supported as monophyletic. Most striking was the discovery that a small set of morphologically and ecologically heterogeneous species formed a clade, here called subgenus Nothonepha. This unexpected result was corroborated by the discovery of deep pits in the lateral body wall (in the mesepisternum) of all Nothonepha, a trait unique within Bembidion. These pits are filled with a waxy substance in ethanol-preserved specimens. In one newly discovered species (Bembidion tetrapholeon sp. n., described here), these pits are so deep that their projections into the body cavity from the two sides touch each other internally. These structures in Bembidion (Nothonepha) are compared to very similar mesepisternal pits which have convergently evolved in two other groups of carabid beetles. The function of these thoracic pits is unknown. Most members of subgenus Nothonepha have in addition similar but smaller pits in the abdomen. A revised classification is proposed for the Antiperyphanes Complex.