D. Johan Kotze

Carabid beetle assemblages (Coleoptera, Carabidae) across urban-rural gradients: an international comparison

We studied communities of carabid beetles in residual forest patchesalong urban-suburban-rural gradients in three cities (Helsinki, Finland; Sofia,Bulgaria and Edmonton, Canada) to examine their responses to urbanisation. OnlyFinnish carabids showed a marked division of community structure along thegradient. In Bulgaria and Canada, carabids did not separate into distincturban,suburban and rural communities. Our results provide some support for thepredictions that species richness will decrease, that opportunistic specieswillgain dominance, and that small-sized species will become more numerous underdisturbance such as that provided by urbanisation. The rather weak and variedresponse of carabids to this disturbance suggests that local factors and theirinteraction are of primary importance for community composition. Occurrence ofreasonably similar carabid communities across the gradient at each of the threelevels of urbanisation suggests that habitat changes commonly associated withurbanisation have not affected the ecological integrity of carabid assemblagesin residual urban forest patches.

Using the ecosystem services approach for better planning and conservation of urban green spaces: a Finland case study

Ecosystem services are vital for humans in urban regions. However, urban development poses a great risk for the ability of ecosystems to provide these services. In this paper we first address the most important ecosystem services in functional urban regions in Finland. Well accessible and good quality recreational ecosystem services, for example, provided by urban nature, are an important part of a high-quality living environment and important for public health. Vegetation of urban regions can have a role in carbon dioxide sequestration and thus in climate change mitigation. For instance, estimates of carbon sinks can be compared to total CO2 emissions of an urban region, and the municipality can aim at both increasing carbon sinks and decreasing CO2 emissions with proper land-use planning. Large and contiguous core nature areas, smaller green areas and ecological connections between them are the essence of regional ecological networks and are essential for maintaining interconnected habitats for species and thus biological diversity. Thus, both local and regional level ecological networks are vital for maintaining ecosystem services in urban regions. The impacts of climate change coupled with land-use and land cover change will bring serious challenges for maintaining ecosystem services in urban areas. Although not yet widely used in planning practices, the ecosystem services approach can provide an opportunity for land-use planning to develop ecologically sustainable urban regions. Currently, information on ecosystem services of urban regions is lacking and there is a need to improve the knowledge base for land-use planning.

Forty years of carabid beetle research in Europe – from taxonomy, biology, ecology and population studies to bioindication, habitat assessment and conservation

Abstract ‘Carabidologists do it all’ (Niemelä 1996a) is a phrase with which most European carabidologists are familiar. Indeed, during the last half a century, professional and amateur entomologists have contributed enormously to our understanding of the basic biology of carabid beetles. The success of the field is in no small part due to regular European Carabidologists’ Meetings, which started in 1969 in Wijster, the Netherlands, with the 14th meeting again held in the Netherlands in 2009, celebrating the 40th anniversary of the first meeting and 50 years of long-term research in the Dwingelderveld. This paper offers a subjective summary of some of the major developments in carabidology since the 1960s. Taxonomy of the family Carabidae is now reasonably established, and the application of modern taxonomic tools has brought up several surprises like elsewhere in the animal kingdom. Progress has been made on the ultimate and proximate factors of seasonality and timing of reproduction, which only exceptionally show non-seasonality. Triggers can be linked to evolutionary events and plausibly explained by the “taxon cycle” theory. Fairly little is still known about certain feeding preferences, including granivory and ants, as well as unique life history strategies, such as ectoparasitism and predation on higher taxa. The study of carabids has been instrumental in developing metapopulation theory (even if it was termed differently). Dispersal is one of the areas intensively studied, and results show an intricate interaction between walking and flying as the major mechanisms. The ecological study of carabids is still hampered by some unresolved questions about sampling and data evaluation. It is recognised that knowledge is uneven, especially concerning larvae and species in tropical areas. By their abundance and wide distribution, carabid beetles can be useful in population studies, bioindication, conservation biology and landscape ecology. Indeed, 40 years of carabidological research have provided so much data and insights, that among insects - and arguably most other terrestrial organisms - carabid beetles are one of the most worthwhile model groups for biological studies.

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.

Carabid beetle and spider assemblages along a forested urban-rural gradient in southern Finland

To investigate the effects of urbanization on carabid beetles (Carabidae) and ground dwelling spiders (Araneae) a study was completed along a 20 km urban–rural forest gradient in the Helsinki–Espoo area of southern Finland. To study changes in assemblage structure, abundance and species richness, these taxa were collected in the year 2000 using pitfall traps, which had been placed in four forest sites within each of the urban, suburban and rural zones. We expected to find changes in the abundances and species richnesses in the two taxa across the urban–rural gradient, but did not find any. Our second and third hypotheses, stating that generalist species and small-bodied species should gain dominance along the gradient from rural to urban sites, were partly supported as carabid specialists were more characteristic of suburban and rural environments whereas generalists were more likely to be collected from rural areas compared to suburban or urban sites. Furthermore, medium to large-sized carabid individuals were more likely to be collected in the rural sites compared to urban forests. We found no evidence for significant changes in spider abundance or species richness across the urban–rural gradient in relation to body size or habitat specialization. We suggest that urbanization does not have significant effects on the total abundances and species richnesses in these two taxa. However, individual species responded differently to urbanization, and there were significant differences in the specialization and body sizes of carabids across the gradient.

Trophic level modulates carabid beetle responses to habitat and landscape structure : a pan-European study

1. Anthropogenic pressures have produced heterogeneous landscapes expected to influence diversity differently across trophic levels and spatial scales.

Urban snow indicates pollution originating from road traffic

Traffic is a major source of pollutants in cities. In this well-replicated study we analysed a broad array of contaminants in snowpacks along roads of different traffic intensities. The majority of pollutants showed a similar pattern with respect to traffic intensity: pH and conductivity as well as concentrations of PAHs, total suspended solids, phosphorus and most heavy metals were higher next to high intensity roads compared to low intensity roads. These pollutant levels also decreased considerably up to 5 m distance from the roads. Furthermore, apart from nitrogen, these variables increased in concentration from control sites in urban forest patches to road bank sites next to roads of low, intermediate and high traffic intensities. The deposition pattern of various traffic-derived pollutants--whether gaseous or particle-bound--was the same. Such information can be useful for the purposes of managing pollutants in urban areas.

Effects of fragmentation and trampling on carabid beetle assemblages in urban woodlands in Helsinki, Finland

We studied the effects of fragmentation (edge effects and patch size) and trampling (path cover) on carabid beetle assemblages in urban woodland patches in Helsinki, Finland. We expected that (1) open habitat and generalist species would benefit and forest species would suffer from increased woodland fragmentation, and (2) most carabid species would respond negatively to increased levels of trampling. A total of 2088 carabid individuals representing 37 species were collected. A cluster analysis distinguished sites in the interior of large woodland patches, with low or moderate path cover, from the other sites. The other sites did not cluster meaningfully, suggesting increased variation in the carabid fauna with increasing human impact. All species and ecological species-groups decreased with increasing distance from the edge toward the woodland interior and with increasing patch size. This pattern is in accordance with our expectation for open habitat and generalist species but opposite to what we expected for forest species. The reason for these surprising results may be that (1) the species we collected are not true forest interior species, (2) urban woodland edges are optimal habitats for many forest carabids, or (3) edges are actually sub-optimal, and high catches simply reflect increased activity of beetles moving away from the edge. Trampling did not have an overall negative effect on carabids as hypothesized. Species associated with moist forest habitat responded as predicted: they decreased in abundance with increasing path cover. Furthermore, open habitat species decreased with increasing path cover but more straightforward than we had predicted. Model elaboration, by dropping the highly trampled sites from the analyses, suggested that our data of high trampling may be too scarce: the results without these sites were more in accordance with our predictions than with the full dataset.

Effects of habitat edges and trampling on the distribution of ground beetles (Coleoptera, Carabidae) in urban forests

Urban forest patches are generally small with highly contrasting edges bordering non-forest habitat, landscape features that increase with urbanisation. These forest patches are also subject to high human foot traffic resulting in trampling and other user disturbances that affect their quality as habitat for invertebrates. We studied the effects of these factors on carabid beetles in urban forests in the cities of Helsinki (Finland) and Edmonton (Canada). In both cities, the structure of carabid assemblages was affected by trampling intensity and distance to the forest edge. Moderate intensity of trail use was associated with increased beetle captures, especially in Edmonton. The effects on individual species were apparent in Edmonton but harder to demonstrate in Helsinki where forest specialist species may have been largely extirpated already. We suggest that these differences result because there has been a long history of fragmentation of urban forest patches in Helsinki, species loss from such patches is gradual, and understorey vegetation structure constrains human foot traffic in forest patches more in Edmonton than it does in Helsinki.

Quantifying the effects of trampling and habitat edges on forest understory vegetation – A field experiment

We investigated the effects of human trampling on boreal forest understory vegetation on, and off paths from suburban forest edges towards the interiors and on the likelihood of trampling-aided dispersal into the forests for three years by carrying out a trampling experiment. We showed that the vegetation was highly sensitive to trampling. Even low levels of trampling considerably decreased covers of the most abundant species on the paths. Cover decreased between 10 and 30% on paths which had been trampled 35 times, and at least by 50% on those trampled 70-270 times. On-path vegetation cover decreased similarly at forest edges and in the interiors. However, some open habitat plant species that occurred outside the forest patches and at forest edges dispersed into the forests, possibly through the action of trampling. A higher cover percentage of an open habitat species at the forest edge line increased its probability to disperse into the forest interior. The vegetation community on, next to, and away from lightly trampled paths remained the same throughout the trampling experiment. For heavily trampled paths, the community changed drastically on the paths, but stayed relatively similar next to and away from the paths. As boreal vegetation is highly sensitive to the effects of trampling, overall ease of access throughout the forest floor should be restricted to avoid the excessive creation of spontaneous paths. To minimize the effects of trampling, recreational use could be guided to the maintained path network in heavily used areas.