Urbanization reduces gut bacterial microbiome diversity in a specialist ground beetle, Carabus convexus.

Urbanization is rapidly shaping and transforming natural environments, creating networks of modified land types. These urbanization-driven modifications lead to local extinctions of several species, but the surviving ones also face numerous novel selection pressures, including exposure to pollutants, habitat alteration, and shifts in food availability and diversity. Based on the assumption that the environmental pool of microorganisms is reduced in urban habitats due to habitat alteration, biodiversity loss, and pollution, we hypothesized that the diversity of bacterial microbiome in digestive tracts of arthropods would be lower in urban than rural habitats. Investigating the gut bacterial communities of a specialist ground beetle, Carabus convexus, in forested rural versus urban habitats by next generation high-throughput sequencing of the bacterial 16S rRNA gene, we identified 3839 bacterial amplicon sequence variants. The composition of gut bacterial samples did not significantly differ by habitat (rural vs. urban), sex (female vs. male), sampling date (early vs. late spring), or their interaction. The microbiome diversity (evaluated by the Rényi diversity function), however, was higher in rural than urban adults. Our findings demonstrate that urbanization significantly reduced the diversity of the gut bacterial microbiome in C. convexus.

High phytoremediation and translocation potential of an invasive weed species (Amaranthus retroflexus) in Europe in metal-contaminated areas.

We demonstrated the metal accumulation potential of Amaranthus retorflexus, a European weed species, both in moderately and strongly metal-contaminated sites. Metal accumulation in roots, stems, and leaves were studied. We also calculated the bioaccumulation factor (BAF), and translocation factor (TF) values to quantify the metal accumulation, and translocation between plant organs. Our findings indicated that metal accumulation correlated with metal concentration; that is plant organs accumulated higher concentration of metals in the contaminated area than in the control one. We found that the concentrations of Ba, Mn, Sr and Zn were the highest in leaves, and Al, Cr, Cu, Fe and Pb in roots. High BAF value was found for Sr in all studied areas, indicating this metal's high accumulation potential of Amaranthus retorflexus. High TF values were found for Al, Ba, Cu, Fe, Mn, Sr and Zn; these metals were successfully transported to aboveground plant organs. We demonstrated that A. retroflexus, a fast-growing, rapidly spreading weed in Europe, was especially useful for heavy metal phytoremediation and phytoextraction.

Early-stage growth and elemental composition patterns of Brassica napus L. in response to Cd-Zn contamination.

Soil contamination caused by the presence of Cd and the excess amount of Zn is a widespread concern in agricultural areas, posing significant risks to the growth and development of crops. In this paper, the early-stage development and metal (Cd and Zn) accumulation potential of rapeseed (Brassica napus L.) grown under different metal application schemes were assessed by determining radicle and hypocotyl length and the micro- and macro elemental composition of plantlets after 24, 72, and 120 h. The results indicated that the single and co-application of Cd and Zn significantly reduced the radicle and hypocotyl lengths. Accumulation intensity for Cd and Zn was affected by Cd and the combination of Cd and Zn in the solution, respectively. In addition, both metals significantly influenced the tissue Mn and had a minor effect on Cu and Fe concentrations. Both Cd and Zn significantly affected macro element concentrations by decreasing tissue Ca and influencing K and Mg concentrations in a dose- and exposure time-dependent manner. These findings specify the short-term and support the long-term use of rapeseed in remediation processes. However, interactions of metals are crucial in determining the concentration patterns in tissues, which deserves more attention in future investigations.

Heavy metal uptake by plant parts of Populus species: a meta-analysis.

Populus species are well documented for being potentially suitable for phytoremediation purposes regarding their accumulation characteristics. However, published results are contradictory. Based on the data gathered during an extensive literature search, we aimed to assess and revise the metal accumulation potential in the root, stem, and leaf of Populus species growing in contaminated soils, with meta-analysis. We evaluated the influences of pollution level, soil pH, and exposure time on the metal uptake patterns. We found accumulations of Cd, Cr, Cu, Pb, and Zn to be significant in each plant part, while that was only moderate for Ni, and limited for Mn. By calculating the soil pollution index (PI), we observed significantly intensive, PI-independent accumulation for Cd, Cr, Cu, Ni, Pb, and Zn. A decrease in soil pH significantly increased the uptake of Mn and significantly decreased the accumulation of Pb in the stem. Metal uptake was significantly influenced by exposure time as well; Cd concentration was significantly decreased in the stem, while concentrations of Cr in the stem and leaf, and Mn in the stem were significantly increased with time. These aforementioned findings support a well-founded metal-and-growth condition-specific application of poplars in phytoremediation processes, also triggering further in-depth assessments to enhance the efficiency of relevant poplar-based technologies.

A meta-analysis on the heavy metal uptake in Amaranthus species.

Metals can accumulate in different parts of plant species in high concentrations, which gives the basis for the plant-based technology called phytoremediation. Among annual species, Amaranthus is a well-studied, potential metal accumulator genus; however, some conflicts are found among published results. Thus, we studied the metal (Cd, Cu, Fe, Ni, Pb, and Zn) accumulation potential of Amaranthus plant parts (root, stem, and leaf) by meta-analysis, furthermore, by calculation of bioaccumulation factor (BAF) values. After the extensive literature search and the calculation of relative interaction intensity (RII) values, we found significant accumulation for each metal by Amaranthus individuals growing on contaminated soils compared to plants collected from uncontaminated ones. Differences among plant parts were significant for Cu and Fe, minor for Ni, Pb, and Zn, and negligible for Cd. The BAF values indicated high accumulation in the leaf, moderate in root and stem for Cd, moderate in each plant part for Pb, and very low in each plant part for Fe, Ni, and Zn. We highlight that Amaranthus species are good prospects for metal phytoremediation projects, although, due to specific plant part-metal patterns, special attention should be paid to the harvesting practice.

Discordant spatiotemporal dynamics of functional and phylogenetic diversity of rotiferan communities exposed to aquaculture effluent.

The growth of the human population brought about the global intensification of aquacultural production, and aquaculture became the fastest growing animal husbandry sector. Effluent from aquaculture is an anthropogenic environmental burden, containing organic matter, nutrients and suspended solids that affect water quality especially in the water bodies of high biodiversity and conservation value. Water quality assessment often relies on bioindicators, analysing changes in taxonomic diversity of various freshwater organismal groups. Stepping beyond taxon diversity, we used functional and phylogenetic diversities of rotifers to identify factors affecting their community organization in response to an aquaculture effluent gradient in the largest oxbow lake in the Carpathian Basin, Hungary. Sampling was carried out three times per season at five points along a 3.5 km section of the oxbow lake, including the point of effluent inflow. We used eight traits to evaluate functional diversity: body size, trophi type, feeding mode, protection type, body wall type, corona type, habitat preference and tolerance level. Functional and phylogenetic distances among the 24 species identified indicated trait conservatism. Rotiferan diversity increased with increasing distance from the point of influx in spring and summer. Among the factors affecting community organization in spring and summer, we find examples of environmental filtering, while in autumn the role of biotic interaction is more frequent. Under nutrient-rich conditions in spring and summer, organisms belonging to the same functional group were dominant, whereas under oligotrophic conditions, more diverse but less abundant groups were present. Considering functional and phylogenetic traits allowed us to identify organising forces of rotifer communities in the largest oxbow lake of the Hungarian Lowland.

A meta-analysis-based evaluation of metallic element accumulation in earthworms.

The responses of earthworms to excess soil element concentrations are well studied. However, published information on the metallic element accumulation in individuals is controversial. In this paper, the published data on earthworm As, Cd, Cr, Cu, Ni, Pb, and Zn whole body concentrations were evaluated in individuals collected from contaminated and uncontaminated (control) soils, using meta-analyses. The role of soil pH and exposure time as potential influencing factors on metal accumulation was also assessed. Based on the evaluations, the accumulation of each metallic element was significantly (p < 0.05) more intensive in individuals collected from contaminated soils than in ones from control soils, with minor differences in the order of accumulation intensity among the studied metallic elements. Further, major interspecific differences were indicated in the accumulation, with different species being the most intensive accumulators for individual metallic elements. Among the studied metals, Cu concentration in earthworm bodies increased significantly with increasing soil pH. As for the exposure time-dependent accumulation, Pb concentration was found to decrease significantly with time in whole body tissues of earthworms. These results suggested a high variability in metal- and species-specific accumulation-excretion patterns of earthworms, influenced also by other external factors. Based on the results highlighted in this meta-analysis, accumulation schemes raise the need for further analyses involving other additional variables (e.g., soil type, organic matter content, climatic condition) to get a better understanding of element cycle-earthworm relations.

Urban Individuals of Three Rove Beetle Species Are Not More Exploratory or Risk-Taking Than Rural Conspecifics.

Urbanization is creating changes in environmental and habitat conditions, as well as creating disturbance and threats to urban-associated species. Some traits, such as high exploratory and risk-taking behavior, are beneficial to allow colonization of urban habitats and coping with urbanization-derived pressures. In this study the exploratory and risk-taking behavior of rural and urban individuals of three forest-associated rove beetle species were tested during their main reproductive period by five frequently used behavioral measures. Individuals of all studied species were similarly ranked by all behavioral measures, indicating that the studied rove beetles responded consistently in the different contexts. However, the behavior of beetles was consistent over time for all/most studied species only by using two measures of exploratory behavior. These provide evidence for the existence of the exploratory dimension of personality in rove beetles. We found a higher exploratory behavior in males than females in Ocypus nitens which can be explained by the active searching of males for mating partners. There were no urbanization-related differences in the exploratory behavior of individuals, suggesting that behavioral changes (being more exploratory) may not yield additional fitness benefits in these rove beetle species with good dispersal capacity.

Land use and soil characteristics affect soil organisms differently from above-ground assemblages.

BACKGROUND: Land-use is a major driver of changes in biodiversity worldwide, but studies have overwhelmingly focused on above-ground taxa: the effects on soil biodiversity are less well known, despite the importance of soil organisms in ecosystem functioning. We modelled data from a global biodiversity database to compare how the abundance of soil-dwelling and above-ground organisms responded to land use and soil properties. RESULTS: We found that land use affects overall abundance differently in soil and above-ground assemblages. The abundance of soil organisms was markedly lower in cropland and plantation habitats than in primary vegetation and pasture. Soil properties influenced the abundance of soil biota in ways that differed among land uses, suggesting they shape both abundance and its response to land use. CONCLUSIONS: Our results caution against assuming models or indicators derived from above-ground data can apply to soil assemblages and highlight the potential value of incorporating soil properties into biodiversity models.

Are There Personality Differences between Rural vs. Urban-Living Individuals of a Specialist Ground Beetle, Carabus convexus?

The world-wide, rapid urbanization is leading to substantial changes in environmental and habitat conditions. These changes, as well as disturbances accompanying urbanization have considerable effects at various levels of the biological organization on wildlife. Understanding behavioral responses to such changes is essential for identifying which organisms may successfully adapt to the altered conditions. In this study, individuals of a forest specialist ground beetle, Carabus convexus, from rural and urban forest patches were tested for their exploratory and risk-taking behavior. Beetles responded consistently in the different contexts; furthermore, by behaving consistently over time, demonstrated that they had personalities. Agglomerative cluster analysis identified two groups of behavioral traits: the exploratory and the risk-taking dimension of personality. Urban females were significantly more exploratory than urban males which can be an adaptation to find high quality food needed to mature eggs in urban habitats, as well as to select favorable microsites for oviposition. Moreover, urban females and males showed more risk-taking behavior than rural females. Urban beetles with more risk-taking behavior may be better able to cope with frequent urbanization-driven disturbance events.

Differences in Life History Traits in Rural vs. Urban Populations of a Specialist Ground Beetle, Carabus convexus.

Urbanization is increasing worldwide and causes substantial changes in environmental parameters, generating various kinds of stress on arthropods, with several harmful consequences. We examined a forest specialist ground beetle, Carabus convexus, in forested habitats to evaluate the changes in four important life history traits between rural and urban populations. Analyzing beetles from the overwintered cohort in their first breeding season, we found no significant differences in body length or body mass between the rural and urban individuals. Body condition, judged by fat reserve scores, was similarly poor in both habitats, indicating that beetles were not able to accumulate substantial fat reserves at either habitat. Females with ripe eggs in their ovaries were first captured at the same time in both areas. The number of ripe eggs, however, was significantly higher in females of the low-density urban population (6.13 eggs/female) than in those of the high-density rural population (4.14 eggs/female), indicating density-dependent fecundity. Altered environmental and habitat conditions by urbanization, however, seemed to cause high mortality during egg hatching and/or larval development, preventing the growth of the urban population to the level of rural one.

Biodiversity differences between managed and unmanaged forests: meta-analysis of species richness in Europe.

Past and present pressures on forest resources have led to a drastic decrease in the surface area of unmanaged forests in Europe. Changes in forest structure, composition, and dynamics inevitably lead to changes in the biodiversity of forest-dwelling species. The possible biodiversity gains and losses due to forest management (i.e., anthropogenic pressures related to direct forest resource use), however, have never been assessed at a pan-European scale. We used meta-analysis to review 49 published papers containing 120 individual comparisons of species richness between unmanaged and managed forests throughout Europe. We explored the response of different taxonomic groups and the variability of their response with respect to time since abandonment and intensity of forest management. Species richness was slightly higher in unmanaged than in managed forests. Species dependent on forest cover continuity, deadwood, and large trees (bryophytes, lichens, fungi, saproxylic beetles) and carabids were negatively affected by forest management. In contrast, vascular plant species were favored. The response for birds was heterogeneous and probably depended more on factors such as landscape patterns. The global difference in species richness between unmanaged and managed forests increased with time since abandonment and indicated a gradual recovery of biodiversity. Clearcut forests in which the composition of tree species changed had the strongest effect on species richness, but the effects of different types of management on taxa could not be assessed in a robust way because of low numbers of replications in the management-intensity classes. Our results show that some taxa are more affected by forestry than others, but there is a need for research into poorly studied species groups in Europe and in particular locations. Our meta-analysis supports the need for a coordinated European research network to study and monitor the biodiversity of different taxa in managed and unmanaged forests.

Predation Pressure on Sentinel Insect Prey Along a Riverside Urbanization Gradient in Hungary.

Urbanization is one of the most important global trends which causes habitat reduction and alteration which are, in turn, the main reasons for the welldocumented reduction in structural and functional diversity in urbanized environments. In contrast, effects on ecological mechanisms are less known. Predation is one of the most important ecological functions because of its communitystructuring effects. We studied six forest habitats along a riverside urbanization gradient in Szeged, a major city in southern Hungary, crossed by the river Tisza, to describe how extreme events (e.g., floods) as primary selective pressure act on adaptation in riparian habitats. We found a generally decreasing predation pressure from rural to urban habitats as predicted by the increasing disturbance hypothesis (higher predator abundances in rural than in urban habitats). The only predators that reacted differently to urbanization were ground active arthropods, where results conformed to the prediction of the intermediate disturbance hypothesis (higher abundance in moderately disturbed suburban habitats). We did not find any evidence that communities exposed to extreme flood events were preadapted to the effects of urbanization. The probable reason is that changes accompanied by urbanization are much faster than natural landscape change, so the communities cannot adapt to them.

Environmental filtering is the main assembly rule of ground beetles in the forest and its edge but not in the adjacent grassland.

In a fragmented landscape, transitional zones between neighboring habitats are common, and our understanding of community organizational forces across such habitats is important. Edge studies are numerous, but the majority of them utilize information on species richness and abundance. Abundance and taxonomic diversity, however, provide little information on the functioning and phylogeny of the co-existing species. Combining the evaluation of their functional and phylogenetic relationships, we aimed to assess whether ground beetle assemblages are deterministically or stochastically structured along grassland-forest gradients. Our results showed different community assembly rules on opposite sides of the forest edge. In the grassland, co-occurring species were functionally and phylogenetically not different from the random null model, indicating a random assembly process. Contrary to this, at the forest edge and the interior, co-occurring species showed functional and phylogenetic clustering, thus environmental filtering was the likely process structuring carabid assemblages. Community assembly in the grassland was considerably affected by asymmetrical species flows (spillover) across the forest edge: more forest species penetrated into the grassland than open-habitat and generalist species entered into the forest. This asymmetrical species flow underlines the importance of the filter function of forest edges. As unfavorable, human-induced changes to the structure, composition and characteristics of forest edges may alter their filter function, edges have to be specifically considered during conservation management.

Pollution intensity-dependent metal accumulation in ground beetles: a meta-analysis.

Survival of organisms in polluted habitats is a key factor regarding their long-term population persistence. To avoid harmful physiological effects of pollutants' accumulation in organisms, decontamination and excretion could be effective mechanisms. Among invertebrates, ground beetles are reliable indicators of environmental pollution. Published results, however, are inconsistent, as some studies showed effective decontamination and excretion of pollutants, while others demonstrated severe toxic symptoms due to extreme accumulation. Using ground beetles as model organisms, we tested our pollution intensity-dependent disposal hypothesis for five pollutants (Cd, Cu, Mn, Pb, and Zn) among four soil pollution intensity levels (low, moderate, high, and extreme) by categorical meta-analysis on published data. According to our hypothesis, decontamination and excretion of pollutants in ground beetles are effective in lowly or moderately polluted habitats, while disposal is ineffective in highly or extremely polluted ones, contributing to intense accumulation of pollutants in ground beetles. In accordance with the hypothesis, we found that in an extremely polluted habitat, accumulation of Cd and Pb in ground beetles was significantly higher than in lowly polluted ones. These findings may suggest the entomoremediation potential of ground beetles in an extremely polluted environment.

Conversion from environmental filtering to randomness as assembly rule of ground beetle assemblages along an urbanization gradient.

Urbanization fragments, isolates or eliminates natural habitats, and changes the structure and composition of assemblages living in the remaining natural fragments. Knowing assembly rules is necessary to support and/or maintain biodiversity in urban habitats. We hypothesized that forest communities in rural sites are organized by environmental filtering, but this may be changed by urbanization, and in the suburban and urban forest fragments replaced by randomly organized assemblages, influenced by the colonization of species from the surrounding matrix. Evaluating simultaneously the functional and phylogenetic relationships of co-existing species, we showed that at the rural sites, co-existing ground beetle species were functionally and phylogenetically more similar than expected by chance, indicating that environmental filtering was the likely process structuring these communities. Contrary to this, in urban and suburban sites, the co-occurring species were functionally and phylogenetically not different from the null model, indicating randomly structured assemblages. According to our findings, changes in environmental and habitat characteristics accompanied by urbanization lead to assemblages of randomly colonized species from the surrounding matrix, threatening proper ecosystem functioning. To reassemble stochastically assembled species of urban and suburban fragments to structured, properly functioning communities, appropriate management strategies are needed which simultaneously consider recreational, economic and conservation criteria.

Edge responses are different in edges under natural versus anthropogenic influence: a meta-analysis using ground beetles.

Most edges are anthropogenic in origin, but are distinguishable by their maintaining processes (natural vs. continued anthropogenic interventions: forestry, agriculture, urbanization). We hypothesized that the dissimilar edge histories will be reflected in the diversity and assemblage composition of inhabitants. Testing this "history-based edge effect" hypothesis, we evaluated published information on a common insect group, ground beetles (Coleoptera: Carabidae) in forest edges. A meta-analysis showed that the diversity-enhancing properties of edges significantly differed according to their history. Forest edges maintained by natural processes had significantly higher species richness than their interiors, while edges with continued anthropogenic influence did not. The filter function of edges was also essentially different depending on their history. For forest specialist species, edges maintained by natural processes were penetrable, allowing these species to move right through the edges, while edges still under anthropogenic interventions were impenetrable, preventing the dispersal of forest specialists out of the forest. For species inhabiting the surrounding matrix (open-habitat and generalist species), edges created by forestry activities were penetrable, and such species also invaded the forest interior. However, natural forest edges constituted a barrier and prevented the invasion of matrix species into the forest interior. Preserving and protecting all edges maintained by natural processes, and preventing anthropogenic changes to their structure, composition, and characteristics are key factors to sustain biodiversity in forests. Moreover, the increasing presence of anthropogenic edges in a landscape is to be avoided, as they contribute to the loss of biodiversity. Simultaneously, edges under continued anthropogenic disturbance should be restored by increasing habitat heterogeneity.

Heavy metal uptake by plant parts of willow species: A meta-analysis.

Previous studies on phytoremediation reported contradictory or inconsistent results on the Cd, Pb, and Zn accumulation in and among plant parts of willow (Salix) species. We hypothesized that metals could accumulate in all plant organs in different concentrations and the metal accumulation in tissues would be increased with exposure time. Furthermore, we analysed the effect of soil pH on metal accumulation, and the correlation between metals. We evaluated published information on Cd, Pb, and Zn accumulation in root, stem, twig, and leaf of willow species using meta-analysis. Results showed that all parts of willow species accumulated significantly more Cd, Pb, and Zn in contaminated soils than in uncontaminated soils. However, the metal accumulation was significantly different among plant parts. We concluded that willow species were proven to be prosperous accumulators of Cd (twigs and leaves), Pb (roots and twigs) and Zn (twigs). We found that Cd accumulation rate in stems is higher in soils with lower pH. Significant positive correlation was found between the accumulations of Cd and Zn in stems. Accumulation rates of Cd (both in leaves and twigs) and Zn (in twigs) were increased significantly with exposure time and the accumulation was successful for at least 3 years.

Species Richness Responses to Structural or Compositional Habitat Diversity between and within Grassland Patches: A Multi-Taxon Approach.

Habitat diversity (spatial heterogeneity within and between habitat patches in a landscape, HD) is often invoked as a driver of species diversity at small spatial scales. However, the effect of HD on species richness (SR) of multiple taxa is not well understood. We quantified HD and SR in a wet-dry gradient of open grassland habitats in Hortobágy National Park (E-Hungary) and tested the effect of compositional and structural factors of HD on SR of flowering plants, orthopterans, true bugs, spiders, ground beetles and birds. Our dataset on 434 grassland species (170 plants, 264 animals) showed that the wet-dry gradient (compositional HD at the between-patch scale) was primarily related to SR in orthopterans, ground-dwelling arthropods, and all animals combined. The patchiness, or plant association richness, of the vegetation (compositional HD at the within-patch scale) was related to SR of vegetation-dwelling arthropods, whereas vegetation height (structural HD at the within-patch scale) was related to SR of ground-dwelling arthropods and birds. Patch area was related to SR only in birds, whereas management (grazing, mowing, none) was related to SR of plants and true bugs. All relationships between HD and SR were positive, indicating increasing SR with increasing HD. However, total SR was not related to HD because different taxa showed similar positive responses to different HD variables. Our findings, therefore, show that even though HD positively influences SR in a wide range of grassland taxa, each taxon responds to different compositional or structural measures of HD, resulting in the lack of a consistent relationship between HD and SR when taxon responses are pooled. The idiosyncratic responses shown here exemplify the difficulties in detecting general HD-SR relationships over multiple taxa. Our results also suggest that management and restoration aimed specifically to sustain or increase the diversity of habitats are required to conserve biodiversity in complex landscapes.

Supporting biodiversity by prescribed burning in grasslands - A multi-taxa approach.

There are contrasting opinions on the use of prescribed burning management in European grasslands. On the one hand, prescribed burning can be effectively used for the management of open landscapes, controlling dominant species, reducing accumulated litter or decreasing wildfire risk. On the other hand burning can have a detrimental impact on grassland biodiversity by supporting competitor grasses and by threatening several rare and endangered species, especially arthropods. We studied the effects of prescribed burning in alkaline grasslands of high conservation interest. Our aim was to test whether dormant-season prescribed burning can be an alternative conservation measure in these grasslands. We selected six sites in East-Hungary: in three sites, a prescribed fire was applied in November 2011, while three sites remained unburnt. We studied the effects of burning on soil characteristics, plant biomass and on the composition of vegetation and arthropod assemblages (isopods, spiders, ground beetles and rove beetles). Soil pH, organic matter, potassium and phosphorous did not change, but soluble salt content increased significantly in the burnt sites. Prescribed burning had several positive effects from the nature conservation viewpoint. Shannon diversity and the number of flowering shoots were higher, and the cover of the dominant grass Festuca pseudovina was lower in the burnt sites. Graminoid biomass was lower, while total, green and forb biomass were higher in the burnt plots compared to the control. The key finding of our study was that prescribed burning did not decrease the abundance and diversity of arthropod taxa. Species-level analyses showed that out of the most abundant invertebrate species, 10 were not affected, 1 was negatively and 1 was positively affected by burning. Moreover, our results suggest that prescribed burning leaving unburnt patches can be a viable management tool in open landscapes, because it supports plant diversity and does not threaten arthropods.