Chromosomal diversity and evolution in the ground beetle genus Bembidion and related taxa (Coleoptera: Carabidae: Trechitae)

Chromosome numbers and sex chromosome systems of 154 previously unstudied Bembidion species are described. The genus is nearly uniform: males of 176 of 205 species are 2n=22+XY. Karyotypes are presented for 30 species. There is some variation among species in size of Y and size of autosomes. Within most species autosomes are subequal in size, and metacentric or submetacentric. Subterminal secondary constrictions and B chromosomes are reported from several species.The supertribe Trechitae (Zolini + Trechini + Pogonini + Bembidiini) is hypothesized to be primitively male 2n=22+X or 24+X, and the ancestral Bembidion stock 2n=22+XY. Conclusions are based on the most parsimonious hypothesis of ancestral state given an inferred phylogeny of the group, rather than the widespread-is-primitive arguments used previously. Evolution within Bembidion away from the presumably-primitive 2n=22+XY is discussed. Six lineages have lost Y chromosomes; seven have undergone changes in autosome number. It is not known why such changes are so scarce, nor what particular rearrangements led to the observed diversity. Nonetheless, the cytogenetic data can be used to infer a monophyletic origin of groups possessing derived chromosome numbers or sex chromosomes, and to help resolve species limits.     

Broad‐scale geographic patterns in body size and hind wing development of western Palaearctic carabid beetles(Coleoptera: Carabidae)

Research into large‐scale ecological rules has a long tradition but has received increasing attention over the last two decades. Whereas environmental, especially climatic, influences on the geographic distribution of species traits such as body size are well understood in mammals and birds, our knowledge of the determinants and mechanisms which shape spatial patterns in invertebrate traits is still limited. This study analyzes macroecological patterns in two traits of the highly diverse invertebrate taxon of carabid beetles: body size and hind wing development, the latter being directly linked to species’ dispersal abilities. We tested for potential impacts of environmental variables (spatial, areal, topographic and climate‐related) representing both contemporary conditions and historical processes on large‐scale patterns in the two traits. Regression models revealed hump‐shaped relationships with latitude for both traits in the categories 1) all species, 2) widespread and 3) endemic (restricted‐range) species: body size and the proportion of flightless species increased from northern towards southern Europe and then decreased towards North Africa. The shared and independent influence of environmental factors was analyzed by variation partitioning. While contemporary environmental productivity and stability (represented by measures of ambient energy and water energy balance) had strong positive relationships with carabid body size, patterns in hind wing development were most notably influenced by topography (elevation range). Regions with high elevation range and low historical climate variability (since the last ice age), which likely offer long‐term stable habitats (i.e. glacial refugia), coincide with regions with high proportions of flightless species. Thus geographic patterns in carabid traits tend to be formed not only by recent climate but also by dispersal and historical climate and processes (i.e. glaciations and postglacial colonization).     

Non-target effects of repeated methiocarb slug pellet application on carabid beetle (Coleoptera: Carabidae) activity in winter-sown cereals

The effects of repeated annual application of methiocarb-based slug pellets, broadcast on the soil surface and drilled into the seed bed, on carabid beetle activity were investigated over a four year period on a winter-sown cereal field using pitfall traps in barriered plots. Following applications in late autumn all winter-active carabid populations were severely depressed; total carabid activity falling to less than 5% and 10–15% following broadcast and drilled applications, respectively, compared with untreated plots. Spring and summer-active species, not active at the time of application, were largely unaffected by applications and were responsible for a gradual recovery of total activity from early spring onwards. Activity of all affected winter species remained demonstrably depressed on treated areas for the remainder of their seasonal incidence. However, all except one species, Bembidion obtusum, recovered to normal activity levels in the following season prior to reapplication. Recovery patterns are discussed in terms of the known biology of the species involved. Evidence that a minority of summer-active species were also affected by treatments, sometimes positively and sometimes negatively, were attributed to indirect effects possibly involving prey availability and foraging behaviour. The long-term ecological and short-term agronomic implications of methiocarb effects on carabid populations in winter-sown cereals are discussed.     

Population ecology of the rare carabid beetle Carabus variolosus (Coleoptera: Carabidae) in north-west Germany

The declining European ground beetle Carabus variolosus, an indicator of undisturbed woodland brooks and swamps, is listed in the EU Species and Habitats Directive. Little is known about this flightless and endangered beetle, apart from its distribution range and habitat stenotopy. We present the results of a 2-year mark-release-recapture study on two neighbouring populations in Germany, which investigated basic parameters of population ecology that may be influential in the persistence of the beetle. Extremely small population sizes were the key findings. Since population densities corresponded to those of typical Carabus species, the size of the habitats was most probably the limiting factor. The risk of heavy losses during offspring development and hibernation may be alleviated by the considerable proportion of C. variolosus reproducing in more than 1 year. The fact that we did not recapture any marked specimens from the neighbouring population suggests a very limited dispersal power. The results imply a high vulnerability of the species, provide basic methodological information for an implementation of the EC Directive and for further research, and highlight the necessity to enlarge populated habitats.     

Exocrine glands in the antennae of the carabid beetle,Platynus assimilis (Paykull) 1790 (Coleoptera,Carabidae, Pterostichinae)

The antennae of males and females of the carabid beetle, Platynus, contain numerous type-III gland acini composed of two different gland cells and a canal cell. The findings show that antennal glands are not only found in highly specialized groups, but also in beetles thought to represent ancestral character states. However, in most cases the function of these glands is not well understood.     

Spatial patterns in the distribution of European springtails (Hexapoda: Collembola)

Using a large database on the spatial distribution of European springtails (Collembola) we investigated how range sizes and range distribution across European countries and major islands vary. Irrespective of ecological guild, islands tended to contain more endemic species than mainland countries. Nestedness and species co‐occurrence analysis based on country species lists revealed latitudinal and longitudinal gradients of species occurrences across Europe. Species range sizes were much more coherent and had fewer isolated occurrences than expected from a null model based on random colonization. We did not detect clear postglacial colonization trajectories that shaped the faunal composition across Europe. Our results are consistent with a multiregional postglacial colonization. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 498–506.     

Inventory of the carabid beetle fauna of the Gaoligong Mountains,western Yunnan Province,China: species of the tribe Zabrini (Coleoptera,Carabidae)

A ten-year multidisciplinary, multi-national and multi-institutional biodiversity inventory project in the Gaoligong Shan region of western Yunnan Province, China generated more than 35,000 specimens of the beetle (Coleoptera) family Carabidae. In this report, first of a planned series, we focus on diversity in tribe Zabrini. Our study of just over 1300 specimens of zabrine carabids from the project, all in genus Amara Bonelli, found a total of 13 species, all previously described, to occur in the study area, with none of them strictly endemic. We present a key for identification of adults of these species, as well as nomenclatural data, diagnoses, illustrations of dorsal habitus and male genitalia, and information about geographical, altitudinal and habitat distributions within the study area and overall geographical distribution for each species. Distributions of the species within the study area are compared, and broader geographical range patterns are characterized. We also discuss a possible role of the Gaoligong Shan region as one source area for the present-day fauna of the Himalaya and southern edge of the Qinghai-Xizang (Tibetan) Plateau.     

The effects of forestry on carabid beetles (Coleoptera: Carabidae) in boreal forests

As compared to natural forests, managed boreal forests are younger, more homogeneous in terms of tree age and species composition, and consist of smaller fragments. Here we examine the effects of such characteristics caused by forestry on carabid beetles (Coleoptera, Carabidae) in the boreal region. The main results are the following. (1) Fragmentation of forests and the size of a fragment appear not to be crucial for the survival of the majority of forest carabids, as they tend to be distributed over various successional stages, but species requiring old-growth habitats suffer. (2) For carabids there appear to be no or very few edge specialist species, and forest-open land edges appear to be effective barriers for species associated with forest or open habitat. However, generalist species easily cross the edge, and edges of forest fragments may be invaded by species from the surrounding open habitat. (3) Habitat change following clear-cutting dramatically changes the composition of carabid assemblages: species restricted to mature forests disappear and open-habitat species invade, while habitat generalists survive at least in the short term. Carabid diversity can probably best be maintained if forest management mimics natural processes, maintains natural structures and includes the natural composition of vegetation and other structural elements (such as dead wood) within the stands, provided that these forest features can be maintained and recreated through forest management practices. At a larger scale, the whole spectrum of forest types and ages (especially old-growth forests), and different successional processes (especially fire) should be maintained. These require the development and use of innovative logging methods, and the planning, implementation, and assessment of landscape-scale ecological management strategies.     

Comparison of carabid beetle (Coleoptera: Carabidae) occurrence in rain forest and human-modified sites in south-eastern Madagascar

High species diversity and endemism make Madagascar one of the world's biodiversity hot spots. However, forest cover and, thereby, biodiversity are decreasing rapidly due to human activities. Most of Madagascar is now covered by secondary vegetation, which is species-poor and contains many introduced species. In this study, the effect of human impact through alteration in vegetation on carabid beetles (Coleoptera: Carabidae) was assessed. The study was conducted in montane rain forest in the Parc National de Ranomafana (south-eastern Madagascar), and in a nearby village area with various types of secondary vegetation. Beetles were collected by hand and by pitfall traps from ten sites from both areas. A total of 246 specimens belonging to 54 species were collected. Pitfall traps yielded only six specimens, which clearly shows that pitfall trapping is not a feasible method for catching carabids in such habitats. From the forest in the national park, 38 species were collected and from the village area 28 species. Species compositions and the dominant species were different: 12 species were found from both areas, while 16 species were unique to the village area and 26 species were unique to the forest. Half of the species were singletons and three most abundant species formed 31% of the total number of individuals. We conclude that because forest-dwellers do not occur in the human-modified environment, such species are at great risk of becoming extinct if current rates of forest decline continue in Madagascar.     

Effects of urbanization on ground beetle assemblages (Coleoptera, Carabidae) of grassland habitats in western Canada

We pitfall-trapped carabid beetles across an urban-rural gradient in central Alberta, Canada to assess the relative contributions of landscape structure and habitat quality in explaining variation in composition of assemblages. The gradient ranged from the "urban" zone in the city of Edmonton, through a "suburban" zone just outside the city, into a "rural" zone 25–60 km east of the city. We trapped beetles at three replicated sites for each of two habitats (unmanaged grasslands and highly managed graveyards) in each of the three zones. Most of the 24 species (3162 individuals) captured were commonly associated with open habitats. Pterostichus melanarius , an introduced species, made up 80% of the total catch. Carabid assemblages of grasslands and graveyards were distinct, with lower species richness and lower catch rates of native, introduced and brachypterous carabids in graveyards. Graveyard assemblages also had proportionally fewer brachypterous carabid individuals (32 vs 52%) and species (20 vs 36%), suggesting that populations recruit locally from elsewhere after disturbances. Native carabids were least abundant in the urban areas. Brachypterous carabids were similarly abundant across the gradient, but more occurred in the urban zone than elsewhere (32 vs 5–10%). Samples from graveyards were characterized by lower beta diversity (variation among samples) than those from grasslands, but beta diversity was more-or-less equally variable across the gradient within habitats. The amount of unmanaged grassland habitat and the degree of urbanization significantly influence carabid assemblages, and therefore are relevant to management and conservation of urban grassland areas.     

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.     

Carabid beetle diversity and distribution in Boston Harbor Islands national park area (Coleoptera, Carabidae)

As part of an All Taxa Biodiversity Inventory in Boston Harbor Islands national park area, an inventory of carabid beetles on 13 islands was conducted. Intensive sampling on ten of the islands, using an assortment of passive traps and limited hand collecting, resulted in the capture of 6,194 specimens, comprising 128 species. Among these species were seven new state records for Massachusetts (Acupalpus nanellus,Amara aulica,Amara bifrons, Apenes lucidulus, Bradycellus tantillus, Harpalus rubripes and Laemostenus terricola terricola-the last also a new country record; in passing we report also new state records for Harpalus rubripes from New York and Pennsylvania, Amara ovata from Pennsylvania, and the first mainland New York records for Asaphidion curtum). For most islands, there was a clear relationship between species richness and island area. Two islands, however, Calf and Grape, had far more species than their relatively small size would predict. Freshwater marshes on these islands, along with a suite of hygrophilous species, suggested that habitat diversity plays an important role in island species richness. Introduced species (18) comprised 14.0% of the total observed species richness, compared to 5.5% (17 out of 306 species) documented for Rhode Island. We surmise that the higher proportion of introduced species on the islands is, in part, due to a higher proportion of disturbed and open habitats as well as high rates of human traffic. We predict that more active sampling in specialized habitats would bring the total carabid fauna of the Boston Harbor Islands closer to that of Rhode Island or eastern Massachusetts in richness and composition; however, isolation, human disturbance and traffic, and limited habitat diversity all contribute to reducing the species pool on the islands relative to that on the mainland.     

Ski trail effects on a beetle (Coleoptera: Carabidae, Elateridae) community in Vermont

Beetle diversity and species composition were studied using pitfall traps placed along three parallel transects that extended from the center of a downhill ski trail, 100 m into a large, unfragmented spruce-fir forest on Mount Mansfield State Park, VT, USA. Groups of beetles showed associations with three distinct habitats: ski trail openings (Agonum cupripenne, Amara spp., Poecilus lucublandus, Hypnoidus bicolor, and H. abbreviatus), edge (Pterostichus coracinus, Agonum retractum, Sphaeroderus canadensis, and Calathus ingratus), and forest (Pterostichus adstrictus, P. punctatissimus, P. brevicornis, Platynus decentis, Trechas apicalis, T. crassiscapus, and Eanus maculipennis). Consequently, carabid and elaterid species composition in the ski trail, edge, and 5 m into the unfragmented forest showed nearly complete turnover. Most beetles within the ski trail were colonizers from lower elevations; however, one species, H. bicolor, was a tundra species that appeared to have colonized lower elevations along the ski trail opening. Of "forest" beetles, only 2 of 540 individuals (< 0.5%) were collected in the ski trail opening. Thus, ski trails are strong barriers to dispersal for forest beetles, several of which are flightless or dimorphic and primarily short-winged. Consequently, fragmentation of mountain slopes by ski trails, especially those that run perpendicular to the climatic gradient, may put isolated populations at a greater risk of local extirpation.     

Model of succession in degraded areas based on carabid beetles (Coleoptera, Carabidae)

Degraded areas constitute challenging tasks with respect to sustainable management of natural resources. Maintaining or even establishing certain successional stages seems to be particularly important. This paper presents a model of the succession in five different types of degraded areas in Poland based on changes in the carabid fauna. Mean Individual Biomass of Carabidae (MIB) was used as a numerical measure for the stage of succession. The run of succession differed clearly among the different types of degraded areas. Initial conditions (origin of soil and origin of vegetation) and landscape related aspects seem to be important with respect to these differences. As characteristic phases, a 'delay phase', an 'increase phase' and a 'stagnation phase' were identified. In general, the runs of succession could be described by four different parameters: (1) 'Initial degradation level', (2) 'delay', (3) 'increase rate' and (4) 'recovery level'. Applying the analytic solution of the logistic equation, characteristic values for the parameters were identified for each of the five area types. The model is of practical use, because it provides a possibility to compare the values of the parameters elaborated in different areas, to give hints for intervention and to provide prognoses about future succession in the areas. Furthermore, it is possible to transfer the model to other indicators of succession.     

Taxonomic review of the Pterostichini and Loxandrini fauna of New Caledonia (Coleoptera, Carabidae)

The generic-level taxa included in Pterostichini and Loxandrini from New Caledonia are reviewed and a key to genera and species provided. Two new genera are described, Paniestichus and Abacophrastus, with the following new species: Paniestichus subsolianus, Abacophrastus millei, Abacophrastus chapes, Abacophrastus carnifex, Abacophrastus hobbit, Abacophrastus megalops, Abacophrastus reflexus and Abacophrastus bellorum. Abacoleptus curtus new species, is described. Notonomus irideus and Notonomus savesi are moved to Prosopogmus. Four new species of Prosopogmus are described: Prosopogmus koghisensis, Prosopogmus lescheni, Prosopogmus fortis and Prosopogmus aoupiniensis. Homalosoma griseolum is moved to Sphodrosomus. Cerabilia is newly recorded from New Caledonia and the genus is moved from Platynini to Loxandrini and therefore is the first report of Loxandrini from New Caledonia. An apparent adventive from Australia, Darodilia, is newly reported from New Caledonia.     

Hybridization and speciation in the carabid beetles of the subgenusOhomopterus (Coleoptera, Carabidae, genusCarabus)

Natural hybridization among wingless carabid beetles of the subgenusOhomopterus (Carabidae, genusCarabus) is reviewed, and its significance in the evolution of this subgenus discussed. Natural hybridization occurs between parapatric species of similar size. Two case studies of natural hybridization suggest that natural hybridization could have affected the evolution of this subgenus in different ways. When there is a large difference in genital morphology between hybridizing species, interspecific copulation often results in genital injuries that causes mortality of copulating individuals, and hence reduces the fitness of hybridizing individuals greatly. In such a case, hybridization may be effective in maintaining the parapatric distribution of the two species, and in the long term, may promote reinforcement selection for traits which are effective in prezygotic reproductive isolation. When the morphological difference in genitalia is not so large as to cause genital injury, a hybrid population may be established at the intermediate zone between two parental species, provided that the immigration rates of the two species into the intermediate zone are small. Thus, natural hybridization may have contributed to both divergence and reticulate evolution in this subgenus.     

Poleward range expansion without a southern contraction in the ground beetle Agonum viridicupreum (Coleoptera, Carabidae)

We investigated the extent of poleward shifts in the distribution range of Agonum viridicupreum due to climate change in the western Palaearctic. Species' records were obtained from extensive literature sources as well as from collections, and consistent amateur entomologists' recordings. Within the general geographic range of the species, we analyzed in detail two parts of both, the northern and southern distribution range boundaries: (1 and 2) north-western Germany (leading or high-latitude edge), (3) Israel and (4) southern Italy (rear or low-latitude edge). Temporal changes in the occurrence data of the species indicated a northward shift of the leading edge of a minimum of 100 km within the last 50 to 100 years. In contrast, according to the data gathered, the rear edge has not changed during the last decades. Further studies are needed in order to fully understand the underlying mechanisms of the different behaviour of leading and rear range edges of Agonum viridicupreum in the current context of global change. Despite our incomplete understanding, chronosequences of the occurrence of the given species have the potential to optimize climate niche modelling to predict trends in the distribution range in the future.