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.     

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.     

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。     

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.     

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.     

Histological structure of tripartite mushroom bodies in ground beetles (Insecta, Coleoptera: Carabidae)

Contrary to members of the suborder Polyphaga, ground beetles have been found to possess tripartite mushroom bodies, which are poorly developed in members of basal taxa and maximally elaborated in evolutionarily advanced groups. Nevertheless, they do not reach the developmental stage, which has been previously found in particular families of beetles. It has been pointed out that a new formation of the Kenyon cells occurs during at least the first months of adult life, and inactive neuroblasts are found even in one-year-old beetles. It has been suggested that there is a relation between the Kenyon cell number and development of the centers of Kenyon cell new-formation.     

Ground beetles of the Ukraine (Coleoptera, Carabidae)

A review of the ground beetles of the Ukrainian fauna is given. Almost 750 species from 117 genera of Carabidae are known to occur in the Ukraine. Approximately 450 species of ground beetles are registered in the Carpathian region. No less than 300 species of ground beetles are found in the forest zone. Approximately 400 species of Carabidae present in the forest-steppe zone are relatively similar in species composition to those in the forest territories. Some 450 species of Carabidae are inhabitants of the steppe zone. Representatives of many other regions of heterogeneous biotopes such as forest, semi desert, intrazonal, etc. can be found in the steppe areas. The fauna of Carabidae (ca. 100 species) of the lowlands of southern Ukraine (sandy biotopes), situated mostly in the Kherson region, is very peculiar. The fauna of the Crimean mountains contains about 300 species. Conservation measures for the Carabidae are discussed.     

A comparative phylogeographic study reveals discordant evolutionary histories of alpine ground beetles (Coleoptera,Carabidae)

Taiwan, an island with three major mountain ranges, provides an ideal topography to study mountain–island effect on organisms that would be diversified in the isolation areas. Glaciations, however, might drive these organisms to lower elevations, causing gene flow among previously isolated populations. Two hypotheses have been proposed to depict the possible refugia for alpine organisms during glaciations. Nunatak hypothesis suggests that alpine species might have stayed in situ in high mountain areas during glaciations. Massif de refuge, on the other hand, proposes that alpine species might have migrated to lower ice‐free areas. By sampling five sympatric carabid species of Nebria and Leistus, and using two mitochondrial genes and two nuclear genes, we evaluated the mountain–island effect on alpine carabids and tested the two proposed hypotheses with comparative phylogeographic method. Results from the phylogenetic relationships, network analysis, lineage calibration, and genetic structure indicate that the deep divergence among populations in all L. smetanai, N. formosana, and N. niitakana was subjected to long‐term isolation, a phenomenon in agreement with the nunatak hypothesis. However, genetic admixture among populations of N. uenoiana and some populations of L. nokoensis complex suggests that gene flow occurred during glaciations, as a massif de refuge depicts. The speciation event in N. niitakana is estimated to have occurred before 1.89 million years ago (Mya), while differentiation among isolated populations in N. niitakana, N. formosana, L. smetanai, and L. nokoensis complex might have taken place during 0.65–1.65 Mya. While each of the alpine carabids arriving in Taiwan during different glaciation events acquired its evolutionary history, all of them had confronted the existing mountain ranges.     

Restoration ecology meets carabidology: effects of floodplain restitution on ground beetles (Coleoptera,Carabidae)

The effects of different restoration activities on floodplain-inhabiting ground beetles were studied at a lowland river in northwestern Germany. In the 6-km² project area the bank reinforcement was stopped, the dikes that bordered the river on both sides were removed and two oxbows (disconnected parts of the main stream course) were reconnected to the river. We investigated the banks at two sites at the main river, at the two reconnected oxbows and at two unconnected oxbows as reference sites. To analyse the effects of the removal of the dikes we chose three floodplain sites in front and two reference sites behind the new dike. As a consequence of the lack of bank reinforcement and the reconnection of the oxbows, the area of open river banks, a rare bank structure at rivers in northwestern Germany, increased by a factor of 4.6 during the 4 years of investigation. Numerous stenotopic ground beetle species colonized the newly developed habitats at the main river and at the reconnected oxbows quickly, during several weeks or months, and established new populations as indicated by newly hatched individuals. The removal of the dikes should lead to a rewetting of the floodplain area that is incorporated in the flood dynamics, an important aspect of the development of typical floodplain habitats such as wet meadows, reeds and alluvial forests, while reference sites on the other side of the removed dike remained uninfluenced. Four years after the removal of the dikes no species turnover at the sites with a potentially changed water regime was recorded that indicated a shift of the carabid assemblages towards hygrophilous species. One reason could be the low frequency of inundations and their short duration during the investigation period. The flood dynamics could be sustained by the elevation and enlargement of the river bed, and periodic floods are the key factor for a development of floodplain habitats.     

Ground beetles (Coleoptera: Carabidae) as bioindicators

One of the primary goals of research on bioindicators is to identifyspecies or other taxonomic units that would reliably indicate disturbances inthe environment, and reflect the responses of other species or the overallbiodiversity. However, there is no perfect bioindicator and selecting the mostsuitable one depends to a great extent on the goal of the survey. In this paperwe examine the suitability of carabids as bioindicators. Carabids are frequentlyused to indicate habitat alteration. They have been used in grasslands andboreal forests where species number and/or abundances have been noted to changealong a habitat disturbance gradient. A common trend is that large, poorlydispersing specialist species decrease with increased disturbance while smallgeneralist species with good dispersal ability increase. Some species are notaffected by moderate disturbance. There is, however, not enough research todetermine how suitable carabids are for biodiversity studies, or how well theyrepresent the response of other species. We conclude that carabids are usefulbioindicators, but as crucial understanding of their relationship with otherspecies is incomplete, they should be used with caution.     

The thoracic morphology of cave-dwelling and free-living ground beetles from China (Coleoptera,Carabidae, Trechinae)

External and internal thoracic structures of two carabid species (Trechini) were examined and documented with different techniques. The study has a main focus on the eyeless cave-dwelling specialist Sinaphaenops wangorum, but detailed information is also provided for a species occurring in cave entrances. The phylogenetic background of the structural features of the thoracic skeletomuscular system was addressed. The thoracic morphology of the examined species was compared to conditions observed in previously studied carabids and non-related subterranean leiodids (Staphylinoidea) in order to identify cave adaptations. Main thoracic character complexes linked with cavernicolous habits in Trechini are elongation of the pro- and mesothorax and the legs, and a complete and irreversible reduction of the flight apparatus. The lost flight capacity is linked with a far reaching modification of skeletal elements of the metathorax including a strongly shortened and simplified metanotum, a shortened metaventrite, and completely reduced wings and sclerites of the wing base. The elongate prothorax together with the long and slender head and elongated legs distinctly increases the activity range in the subterranean lightless environment, which likely facilitates foraging of the carnivorous beetles. Some of the observed features like wing loss and elongation of the anterior thorax and legs are also found in some cave-dwelling Leiodidae (Leptodirini), whereas some other subterranean members of the staphylinoid family have a compact body and legs of normal length. In contrast to the predaceous Trechini, Leptodirini are scavengers.     

Phylogenetic diversification patterns and divergence times in ground beetles (Coleoptera: Carabidae: Harpalinae)

Background  

Harpalinae is a species rich clade of carabid beetles with many unusual morphological forms and ecological interactions. How this diversity evolved has been difficult to reconstruct, perhaps because harpalines underwent a rapid burst of diversification early in their evolutionary history. Here we investigate the tempo of evolution in harpalines using molecular divergence dating techniques and explore the rates of lineage accumulation in harpalines and their sister group.     

Dynamics of abundance of ground beetles (Coleoptera,Carabidae) in fenced areas in the Cis-Altai plain (Western Siberia)

The density of ground beetle populations was estimated by means of fenced pitfall traps in 2004–2008, in the environs of Biisk (Altai Territory) and Karasuk (Novosibirsk Province). In all, 64 species of ground beetles were revealed. The capture rates were usually higher in open than in fenced plots. No correlation was observed between the time of trap exposure and the captures. By the end of the 3rd–6th week of exposure, the fraction of juvenile and immature adults increased. The beetles captured within a fenced area 1 m² usually fell into the traps in 2–3 days. The maximal density of carabids in the fenced areas was estimated at 55–60 ind./m².     

The ground beetles (Coleoptera: Carabidae) of exposed riverine sediments in Scotland and northern England

A survey of invertebrates on exposed riverine sediments (ERS) in four catchments of rivers in Scotland and northern England was carried out in 1996 and 1997 using pitfall traps. The resulting 179 lists of ground beetle species were used in ordination and classification analyses to determine the different types of habitats and the environmental factors influencing species assemblage distribution. Types of ERS habitat differed between highland and lowland catchments; there were also different numbers of habitat types in each catchment. The distribution of ERS within catchments was related to geology, position in catchment, sediment composition and the amount of vegetation. Evidence of the positive nature of river engineering was identified and the role of river management is discussed. The numbers of nationally rare and scarce ground beetle species recorded from the ERS habitats in each catchment are reported and the factors affecting their distribution discussed.     

Proventricular trituration in adult carabid beetles (Coleoptera: Carabidae)

Adult carabids that ingest particles of food have a muscular proventriculus lined internally with an intima that bears sclerotized projections. In experiments where six species of carabid were allowed to feed upon Drosophila, the average size of cuticular remains found in post-proventricular regions of the gut was generally significantly smaller than those in the crop. This reduction in size of cuticular particles is likely to be due to proventricular trituration rather than prolonged exposure to digestive secretions. There is no evidence that material is triturated by the proventriculus and returned to the crop, the main seat of digestion in carabids, so proventricular trituration is unlikely to enhance digestion. By crushing large, potentially abrasive, cuticular particles, the proventriculus may serve to protect midgut epithelial tissue.     

Testing abundance-range size relationships in European carabid beetles (Coleoptera, Carabidae)

Four of the eight hypotheses proposed in the literature for explaining the relationship between abundance and range size (the sampling artifact, phylogenetic non-independence, range position and resource breadth hypotheses) were tested by using atlas data for carabid beetles (Coleoptera, Carabidae) from Belgium, Denmark and the Netherlands. A positive relationship between abundance and partial range size was found in all three countries, and the variation in abundance was lower for widespread species. Analysis of the data did not support three of the proposed hypotheses, but did support the resource breadth hypothesis (species having broader environmental tolerances and being able to use a wider range or resources will have higher local densities and be more widely distributed than more specialised species). Examination of species' characteristics revealed that widespread species are generally large bodied, generalists (species with wide niche breadths occurring in a variety of habitat types) and are little influenced by human-altered landscapes, while species with restricted distributions are smaller bodied, specialists (species with small niche breadths occurring in only one or two habitat types), and favour natural habitat. Landscape alteration may be an important factor influencing carabid abundance and range size in these three countries with a long history of human-induced environmental changes.     

Nocturnal windborne migration of ground beetles,particularly Pseudoophonus griseus (Coleoptera: Carabidae), in China

1 A network of light‐traps, an aerial net carried by kytoon (balloon) and two entomological radars were used to investigate whether ground beetles migrate nocturnally through China. The network‐wide, simultaneous sudden increase in light trap catches, and after subsequent decrease, indicated a seasonal long‐distance night migration of ground beetles, with Pseudoophonus griseus (Panzer) predominant, in August. 2 Aerial net trapping indicated that carabids were able to ascend to altitudes of at least 200 m and become windborne. Radar observations indicated that the migratory beetles formed high‐density layer concentrations at approximately 200–300 m. 3 These concentrations were coincident with the top of the temperature inversion and a wind speed maximum, which suggested that the carabids tended to select warm, fast moving air for their long‐distance migration. 4 The ground beetles orientated and displaced towards the downwind direction in southerly winds. Their air speed decreased as the tailwind increased and, thus, migrating beetles appeared to be conserving energy. 5 The mean ± SD displacement speed (ground speed) and air speed were 6.85 ± 1.73 m/s (n= 172) and 4.45 ± 1.54 m/s (n= 172), respectively. The duration of flight, estimated from the variation in area density derived from radar data, was approximately 9–10 h, indicating that the beetles might migrate hundreds of kilometres in a single flight.