Shading enhances the quality of willow leaves to leaf beetles – but does it matter?

Plant shading is commonly recognised as a factor, which increases susceptibility of plants to attack by herbivorous insects. In this study we experimentally investigated the effect of host plant shading on two willow-feeding leaf beetles, Galerucella lineola feeding upon Salix phylicifolia and Phratora vitellinae feeding upon Salix myrsinifolia . Both beetle species were more abundant on potted willows growing in open habitats than on the same clones placed under the shade of trees. However, in the laboratory the food preference by adults and larval performance showed that the shaded willows are actually better food for both beetle species. On the contrary, when larvae were reared in the field under natural abiotic conditions, we found no difference in larval performance, or if any, even better performance in open habitats. Apparently, higher and more variable daily temperatures in open habitats accelerated the growth of the larvae. When adults were let to emigrate from or immigrate to potted willows, which had been grown in the same conditions but placed either in the open or shady habitats, adults preferred exposed willows. Invertebrate predators were more abundant in open habitats, but we found no differences in leaf beetle mortality by natural enemies between the habitats. Although the larval performance appeared to be approximately equal in the two habitats during the unusually warm study period, we suggest that under suboptimal temperatures the better abiotic conditions of open sites can easily override the better food provided by shaded habitats. The selection of abiotic habitat thus plays a significant role in the adaptive habitat and host plant selection of these beetles within the gradient of shadiness.     

Broad-scale reciprocity in an avian seed dispersal mutualism

Aim Coevolved relationships between individual species of birds and plants rarely occur in seed dispersal mutualisms. This study evaluates whether reciprocal relationships may occur between assemblages of bird and plant species. Location Vancouver Island, British Columbia, Canada (48°50′‐N, 125°22′‐W). Methods The distribution and fruiting phenologies of seven shrub species were compared to seasonal changes in habitat selection and seed dispersal by six fruit‐eating bird species. Results Shrub species inhabiting forest understorey habitat had earlier fruiting phenologies than shrub species inhabiting forest edge habitat along lake and bog margins. Birds showed a parallel pattern in habitat selection, being more abundant in the forest understorey early in the fruiting season, and more abundant in the forest edge later in the season. Rates of seed deposition covaried with avian habitat selection, in such a way that birds directed seed dispersal into habitats preferred by shrubs. Conclusions These results depict a broad‐scale pattern in the abundance of birds and fruits indicative of reciprocal interactions. Seasonal changes in seed dispersal to each habitat appear to reinforce the relationship between shrub habitat affinities and fruiting phenologies. Phenological differences between habitats may also reinforce seasonal changes in avian habitat selection. Therefore, although reciprocal interactions between pairs of bird and plant species are rare, broad‐scale reciprocal relationships may occur between assemblages of bird and plant species.     

Species decline--but why? Explanations of carabid beetle (Coleoptera,Carabidae) declines in Europe

We investigated some of the causes of ground beetle decline using atlas data from Belgium, Denmark and the Netherlands, countries in which natural environments have all but disappeared. We used ordinal regression to identify characteristics that are significantly correlated with the decline of carabid beetle species over the last 50-100 years, using a stepwise selection procedure to select the optimal model according to the Akaike Information Criterion. The results showed that large-bodied carabid populations have declined more than smaller ones, possibly because of their lower reproductive output and lower powers of dispersal. Habitat specialist populations (i.e. species with small niche breadths) have also decreased more than habitat generalist populations. Species with both long- and short-winged individuals have been less prone to decline than those that are exclusively either short-winged or long-winged. Dimorphic species may survive better in highly altered environments because long-winged individuals are good at dispersing between suitable habitats and short-winged individuals are good at surviving and reproducing in these newly colonised habitats. Finally, populations of large carabids associated with coastal, woodland or riparian habitat types were less prone to decline than populations of large carabids associated with various, open or grassland habitat types. The pattern is reversed for carabid species smaller than 8 mm in size. These results are explained in the context of habitat restoration and destruction in these highly modified western European countries.     

Taxon specific response of carabids (Coleoptera,Carabidae) and other soil invertebrate taxa on invasive plant Amorpha fruticosa in wetlands

False indigo (Amorpha fruticosa L.) is an invasive exotic plant introduced to Europe in the early eighteenth century. Its spread has been rapid, particularly in disturbed wetland habitats, where it forms dense impermeable monospecific stands and modifies habitat conditions. The impact of A. fruticosa on native plant communities has been well analyzed, however knowledge concerning the possible effects on soil invertebrates and particularly carabid beetles is completely lacking. This study analyzed the impact of an A. fruticosa invasion on carabid beetles and other soil invertebrates. Soil fauna was sampled by pitfall traps at natural habitats, initially colonized by A. fruticosa, and habitats largely invaded by A. fruticosa. In total 2,613 carabid beetles belonging to 50 species and 72,166 soil invertebrates were collected. The invasion of A. fruticosa strongly affected the carabid beetle species composition, which clearly differed between all studied sites. Widespread euritopic carabid beetle species showed positive responses to A. fruticosa invasion, while the activity density of open habitat species strongly declined. Mean individual biomass was significantly higher at invaded sites due to increased incidence of large carabids (genus Carabus Linné, 1758). Carabid beetle activity density and abundance of soil invertebrates were considerably higher at invaded sites than in natural sites. Conversely, the impact of A. fruticosa on carabid beetle species richness and diversity was less pronounced, most likely due to immigration from adjacent habitats. Changes in carabid beetle species composition and abundance of soil invertebrates were most likely due to changes in vegetation structure and microclimate. The results suggest that A. fruticosa invasion considerably affected carabid beetles, an insect group that is only indirectly related to plant composition. Therefore, severe future changes can be expected in invertebrate groups that are closely related to plant composition, since A. fruticosa cannot be completely removed from the habitat and covers relatively large areas.     

Habitat selection of three chrysomelid species associated with Rumex spp.

Summary Habitat selection of three chrysomelid beetles feeding on the Rumex plants was studied in relation to the ecological requirements of the three species and the habitat characteristics arising in their unstable and often unpredictable environment. The different ecological requirements and bionomic characteristics of the three chrysomelids were associated not only with their differing degrees of feeding specialization; they also influenced habitat selection. For example, the small body size and lower food demands of Mantura clavareaui, which is a feeding specialist, allowed it to exploit any habitat, irrespective of habitat size and duration. The other specialist, Gastrophysa atrocyanea, selected larger habitats because of the relatively brief lifespan of its adults after overwintering, its reduced mortality, and greater food demand. On the other hand, Galerucella vittaticollis, which is a multivoltine, general feeder, selected the habitat that lasted longer, rather than one that was larger. This species' life cycle is not synchronized with Rumex phenology, but its adults are long-lived and it has a flexible life cycle. The different patterns in habitat selection among these three chrysomelids prevented serious overlapping of their habitats; thus, the habitat overlapping of 2 or 3 species was 34.62%, that of all 3 species 3.85%. Most of the habitats utilized by Gastrophysa, however, overlapped those of the other two species, because Gastrophysa's habitat utilization range was the narrowest of the three. In Mantura, the degree of overlap with the other two species was lowest because this beetle had the broadest range of habitat utilization. On the other hand, Galerucella adjusted the temporal, rather than the spatial coordinates of its habitat usage when it occupied habitats where the other two specialists were also present. Galerucella, unlike the other two species, is multivoltine. Consequently, it was able to coexist in habitats occupied by the others by exploiting the host plants at times when the other species were numerically few, or even absent. No direct interspecific competition among adults of the three species was observed. Their different patterns of habitat selection created by their different responses to available food allowed them to coexist on Rumex without direct competition.     

Ecology of pollination in a tropical Venezuelan savanna

Pollination modes ecology of a total of 164 plant species was evaluated according to habitats and plant life forms in the Venezuelan Central Plain. Frequency distribution of nine pollination modes showed that, at the community level bee pollination (38.6%) was dominant. Butterfly (13.9%), fly (12.7%), and wasp (10.8%) pollination were the second most frequent. Moth (6.2%) and wind (10.4%) pollination occurred with similar frequency, and the least common were bird (3.1%), beetle (2.3%) and bat (1.9%) pollination. There was a significant interaction effect indicating that pollination mode was affected by the type of habitat. Bee pollination was the most common pollination mode in all habitats with butterfly, fly and wasp pollination being secondary for forest and forest-savanna transition; and butterfly, wasp, wind and fly pollination being secondary for savanna. Wind, butterfly and fly pollination were found in disturbed areas as secondary pollination modes. Pollination modes were significantly associated and affected by life forms. Bee pollination was dominant in all life forms with wasp, butterfly and fly pollination being the secondary for trees, shrubs, and lianas; and butterfly and wind pollination being the secondary for herbaceous species. The number of pollination modes (richness) among life forms ranged between four and nine for epiphytes and perennial herbs respectively. The highest values of diversity indexes among life forms were found in trees and shrubs. The richness and diversity indices of pollination modes were statistically higher for more structured habitats, forest and forest-savanna transition, than herbaceous habitats, savanna and disturbed areas, which is associated with the highest values of diversity indexes in trees and shrubs. Equitability was higher for forest and disturbed areas than forest-savanna transition and savanna. The results of comparative richness, equitability, diversity indices, and the frequency distribution of pollination modes of 19 samples from tropical and temperate communities indicated that richness of pollination modes may be different between tropical and temperate communities. The proportion of each pollination mode suggests four grouping: (1) rain forests and their strata, (2) grassland savanna, and associated disturbed areas, (3) temperate communities, and (4) the most heterogeneous group, contained mostly neotropical communities, including the four habitats of the Venezuelan Central Plain. The frequency of pollination modes, richness, diversity and equitability of communities, habitats, successional stages, and vegetation strata varies with respect to geography, vegetation structure, and plant species richness.     

Dynamic effects of ground-layer plant communities on beetles in a fragmented farming landscape

Vegetation effects on arthropods are well recognized, but it is unclear how different vegetation attributes might influence arthropod assemblages across mixed-agricultural landscapes. Understanding how plant communities influence arthropods under different habitat and seasonal contexts can identify vegetation management options for arthropod biodiversity. We examined relationships between vegetation structure, plant species richness and plant species composition, and the diversity and composition of beetles in different habitats and time periods. We asked: (1) What is the relative importance of plant species richness, vegetation structure and plant composition in explaining beetle species richness, activity-density and composition? (2) How do plant-beetle relationships vary between different habitats over time? We sampled beetles using pitfall traps and surveyed vegetation in three habitats (woodland, farmland, their edges) during peak crop growth in spring and post-harvest in summer. Plant composition better predicted beetle composition than vegetation structure. Both plant richness and vegetation structure significantly and positively affected beetle activity-density. The influence of all vegetation attributes often varied in strength and direction between habitats and seasons for all trophic groups. The variable nature of plant-beetle relationships suggests that vegetation management could be targeted at specific habitats and time periods to maximize positive outcomes for beetle diversity. In particular, management that promotes plant richness at edges, and promotes herbaceous cover during summer, can support beetle diversity. Conserving ground cover in all habitats may improve activity-density of all beetle trophic groups. The impacts of existing weed control strategies in Australian crop margins on arthropod biodiversity require further study.     

Diversity of coleoptera of the bracket fungus Fomitopsis pinicola in a Norwegian spruce forest

The beetle fauna of 299 sporocarps of the bracket fungus Fomitopsis pinicola in a 200 ha spruce forest in southeastern Norway was investigated in relation to sporocarp, tree and forest variables. The sporocarps contained 36 species of beetles, of which six species are on the Norwegian Red List. Of 12,373 individual beetles collected, 91% were Cis glabratus. Plots of species accumulation curves suggested that there may be more than 60 beetle species present in F. pinicola in the area, but that probably all the specialist Ciidae were found. The major factor influencing beetle diversity turned out to be the level of dead wood at and in the vicinity of the sampling site, with a higher number of species per unit volume of sporocarps in areas with high levels of dead wood. There were also significantly more red-listed species in those areas. Analysing the species occurrence with stepwise logistic regression, we show preferential habitat selection of the six most abundant species of Ciidae. Conservation of beetles associated with bracket fungi using amount of dead wood as a surrogate measurement is discussed.     

Dead trees left in clear-cuts benefit saproxylic Coleoptera adapted to natural disturbances in boreal forest

Forest management alters the pattern of forest dynamics from that in natural conditions in the boreal region. In order to examine how certain forestry measures matching natural dynamics affect forest insects, we compared assemblages of saproxylic Coleoptera on dead, standing birch trunks left behind in eight clear-cut areas with corresponding assemblages in seven mature forests in southern and eastern Finland. We used trunk-window traps for sampling. Distinct beetle assemblages were associated with the different habitats. Median numbers of species or specimens caught did not differ between closed forests and clear-cuts, but individual beetle species occurred unevenly among the habitats. Several beetle species associated with open forest habitat, e.g. burned forests or storm-damage areas, including species regarded as threatened in Finland, were found almost exclusively, in clear-cuts. Correspondingly, a number of beetle species occurring frequently in closed forests were not found in clear-cuts. We conclude that dead trunks left in the clear-cut areas may host not only generalist saproxylic species but also many beetle species specialized to warm, sun-exposed environments, and such species may not be able to survive in closed forests. Management measures matching suppressed natural disturbances are found useful in preserving diversity in managed forests.     

Livestock Trampling Reduces the Conservation Value of Beetle Communities on High Quality Exposed Riverine Sediments

Exposed riverine sediments (ERS) are habitat for a large number of rare and threatened specialist species of invertebrates and are of considerable conservation importance. Livestock trampling is believed to be one of the most widespread causes of damage to ERS habitats in the UK, and as such, its effects were the focus of this investigation. Beetle density was measured at two points within 25 distinct patches of habitat along ∼ ∼47 km of the Afon Tywi special site of scientific interest in South Wales, which is known to support an extremely good quality ERS beetle fauna in a UK context. Partial canonical correspondence analyses were used to explore the relationship between beetle assemblage and a range of environmental variables. The percentage of fine (<8 mm) sediments, median sediment size, distance downstream, cattle stocking levels, and counts of sheep faeces were found to best relate to beetle abundance and assemblage structure. Species richness was positively associated with stocking levels, probably because of the addition of species associated with resultant elevated levels of silt and organic matter. The ERS quality score, which is a measure of conservation value based on the rarity of specialist ERS beetles, was negatively associated with measures of trampling damage. It was therefore concluded that livestock trampling reduces the conservation value of beetle communities on high quality ERS and management should restrict trampling in sites of high conservation importance.     

北热带喀斯特森林木本植物花性状及其生境分异

植物花性状的多样化是植物长期进化及自然选择的结果, 不同植物种间花性状的变异与生境存在一定的相关性。北热带喀斯特季节性雨林具有生境异质性强、群落结构复杂、特有成分丰富等特点, 分析该森林植物性状的变化特征及其与生境的关联性, 有助于理解物种共存、协同进化过程和对生境的适应, 可为阐明喀斯特植物的生态适应性、理解生物多样性维持机理提供依据。本研究在弄岗15 ha森林动态监测样地木本植物开花相对集中的时间段进行, 记录并分析了21个物种花性状的变化。根据物种空间分布及其与生境的关联特性, 将21个物种分成偏好谷底、山坡、山顶的3种类型, 分析了这3种类型植物的花性状差异; 另外又根据花性状对21个物种进行聚类分析, 探讨了聚类分组结果和根据偏好生境分组结果的异同。结果表明: 花大小、花色鲜艳度均与物种优势度存在显著的负相关关系, 花小而不鲜艳的物种在群落中更有优势, 表现出更好的适应性; 不同偏好生境的植物花色明度有显著差异, 其他的花性状差异不显著; 聚类分组与根据偏好生境的分组有较好的一致性, 反映出物种的花性状变化响应了生境的变化。综上结果, 我们认为北热带喀斯特季节性雨林植物花性状与物种的优势度及生境条件有密切联系, 在该区域中生境对花性状的影响可能比传粉者更为深刻。     

Tree colonization by the Asian longhorn beetle,Anoplophora glabripennis (Coleoptera: Cerambycidae): effect of habitat and tree suitability

Tree colonization and feeding activity of the invasive wood‐borer Asian longhorn beetle (Anoplophora glabripennis), an Asian pest introduced into North America and Europe, was studied in a newly invaded area in Italy. The hypothesis being tested was that the reproductive success of the insect depend on habitat type and tree suitability. Adult beetles were caged on branches of host and nonhost species, in both urban and forest habitats. Two months later, number and size of feeding patches on plant tissues, eggs laid, and surviving larvae were assessed. Bark concentration of C and N was also measured from the same trees. Results indicated that the mean area of plant tissues consumed by adult feeding was significantly larger on trees growing in forest than in urban habitat, although within the same habitat there were no differences between susceptible and nonsusceptible trees. ALB tree colonization, in terms of number of eggs laid and young larvae survival, was not affected by habitat while it was higher on susceptible trees. Although trees growing in forests had a lower nitrogen concentration, they allowed colonization rates similar to those of trees growing in the urban habitat. Hence, the amount of carbon and nitrogen did not fully explain tree suitability or habitat selection. We suggest compensatory feeding as a potential mechanism that might explain this peculiar situation, as supported by a more intensive feeding activity recorded on trees in the forest. Suitability of different trees may be due to other factors, such as secondary chemical compounds.     

Biology outside the classroom: the SNAB visit/issue report

Pitfall trap sampling of Carabid beetle species on roundabouts in Bracknell, Berkshire, was used to assess the biodiversity of this taxon by its use as an indicator. The aim of the study was to discover the role of traffic islands in the provision of refugia for invertebrate fauna in fragmented urban habitats. Sampling was performed on 15 roundabouts where a total of 24 species were recorded during four trapping periods over a total of 10 days. The resulting asymptotic curves indicated that the total number of species present on all but two of the sites had been represented in the samples taken. There was found to be a positive correlation between the area of the islands and the number of ground beetle species and between the log of the area and the number of individuals found. The log/log relationship between the area of the islands and the number of species was significant. The total abundance of the beetles present was also positively and significantly correlated with roundabout area when both variables were logged. The number of habitats on the islands was positively correlated with the total abundance and species richness. The relationship between species, area and habitat was also positively and significantly correlated. In conclusion, it is obvious that roundabouts of large area with higher numbers of habitat types are greater in Carabid beetle diversity than small, sparsely vegetated roundabouts. Thus, roundabouts can promote the maintenance of biodiversity in fragmented urban habitats.We present this study as an example of a simple method that could be used or easily adapted for educational use, and suggest how some of the perceived problems in its use may be overcome. We discuss how such a study could be useful in illustrating concepts such as biodiversity and species richness, and some of the factors that influence it, as well as demonstrating the level of biodiversity that can be found in urban environments.     

Evaluation of Swedish woodland key habitats using red-listed bryophytes and lichens

All Swedish forest land is at present being surveyed with the aim of mapping woodland key habitats which have an estimated number of ca. 70 000. An untested requirement of these habitats is that they should contain red-listed species. In order to investigate if they fulfill their aim, an inventory of nationally red-listed bryophytes and lichens was performed in ca. 120 randomly selected woodland key habitats distributed throughout the country. The species were recorded in line transects, covering the whole surface of the habitats. The mean number of red-listed species per habitat was 0.7 for bryophytes and 1.4 for lichens with 11 species at the richest site and 71% of the sites having at least one species. Nemoral forests and coniferous forests were the most species-rich habitat types. The habitats of northern Sweden were more species rich than the southern ones. There were weak correlations between presence of bryophytes and lichens. The study shows that a majority of the woodland key habitats contain red-listed bryophyte and lichen species. Further studies are needed in order to record more organism groups and to compare the woodland key habitats with the surrounding matrix forests.     

Grouse moor management: habitat and conservation implications for invertebrates in southern Scotland

The distribution of ground beetle, rove beetle, spider and plant bug species assemblages, and the occurrence of other beetle species, was investigated in a five-year survey of a grouse moor in the Scottish borders, some of which was managed by burning, cutting and herbicide application. Multivariate analyses were used to assess the relationship between assemblage distribution and a number of environmental variables. The management of dry Calluna moor had a positive effect on the habitat diversity for ground beetles and plant bugs but had little effect on rove beetles and spiders. A total of 39 nationally rare and scarce species (38 beetle, 1 plant bug) was recorded in the survey. The most important habitat present was on streamside sites, especially sediment. A number of these rarer species were restricted to sites managed by burning and cutting but other species were only found on unmanaged wet Calluna moor. Molinia-dominated moor was generally of poorer quality than Calluna sites, with fewer rare and scarce species and lower site rarity values based on the ground beetle species recorded. The highest median site rarity scores were for dry, open, managed Calluna sites. In order to maximise both habitat diversity and the incidence of rare and scarce species on grouse moor, a mosaic of both managed and undisturbed patches differing in soil characteristics, plant composition and vegetation structure appears to be required.     

Effects of habitat area, isolation, and landscape diversity on plant species richness of calcareous grasslands

Calcareous grasslands harbour a high biodiversity, but are highly fragmented and endangered in central Europe. We tested the relative importance of habitat area, habitat isolation, and landscape diversity for species richness of vascular plants. Plants were recorded on 31 calcareous grasslands in the vicinity of the city of Göttingen (Germany) and were divided into habitat specialist and generalist species. We expected that habitat specialists were more affected by area and isolation, and habitat generalists more by landscape diversity. In multiple regression analysis, the species richness of habitat specialists (n = 66 species) and habitat generalists (n = 242) increased with habitat area, while habitat isolation or landscape diversity did not have significant effects. Contrary to predictions, habitat specialists were not more affected by reduced habitat area than generalists. This may have been caused by delayed extinction of long-living plant specialists in small grasslands. Additionally, non-specialists may profit more from high habitat heterogeneity in large grasslands compared to habitat specialists. Although habitat isolation and landscape diversity revealed no significant effect on local plant diversity, only an average of 54% of habitat specialists of the total species pool were found within one study site. In conclusion, habitat area was important for plant species conservation, but regional variation between habitats contributed also an important 46% of total species richness.     

Movement Behaviour of the Carabid Beetle Pterostichus melanarius in Crops and at a Habitat Interface Explains Patterns of Population Redistribution in the Field

Animals may respond to habitat quality and habitat edges and these responses may affect their distribution between habitats. We studied the movement behaviour of a ground-dwelling generalist predator, the carabid beetle Pterostichus melanarius (Illiger). We performed a mark-recapture experiment in two adjacent habitats; a large plot with oilseed radish (Raphanus sativus) and a plot with rye (Secale cereale). We used model selection to identify a minimal model representing the mark-recapture data, and determine whether habitat-specific motility and boundary behaviour affected population redistribution. We determined movement characteristics of P. melanarius in laboratory arenas with the same plant species using video recording. Both the field and arena results showed preference behaviour of P. melanarius at the habitat interface. In the field, significantly more beetles moved from rye to oilseed radish than from radish to rye. In the arena, habitat entry was more frequent into oilseed radish than into rye. In the field, movement was best described by a Fokker-Planck diffusion model that contained preference behaviour at the interface and did not account for habitat specific motility. Likewise, motility calculated from movement data using the Patlak model was not different between habitats in the arena studies. Motility (m² d⁻¹) calculated from behavioural data resulted in estimates that were similar to those determined in the field. Thus individual behaviour explained population redistribution in the field qualitatively as well as quantitatively. The findings provide a basis for evaluating movement within and across habitats in complex agricultural landscapes with multiple habitats and habitat interfaces.     

Multilevel phenotypic selection on morphological characters in a metapopulation of Silene tatarica

This study partitions selection in a natural metapopulation of a riparian plant species, Silene tatarica, into individual- and patch-level components by using contextual analysis, in which a patch refers to a spatially distinct stand of individual plants. We estimated selection gradients for two morphological characters (plant height and number of stems), their respective patch means, and plant density with respect to reproductive success in a two-year study. The approach was also extended to partition selection separately within habitats with varying degrees of exposure to river disturbances and herbivory. The selection differentials and gradients for plant height were positive at both individual and patch levels, with selection forces highest in the closed habitat with low exposure to disturbance. This pattern suggests that local groups with taller than average plants are more visible to pollinators than to groups that are shorter than average plants; and, within patches, individuals with short stature are visited less often than taller ones. Selection on the number of stems was in opposition at individual and patch levels. At the individual level the character was selected toward higher values, whereas selection at the patch-level favored smaller mean number of stems. The strength of the latter component was associated with the intensity of herbivory in different habitats, suggesting that the patch-level selection against a large number of stems might be due to high attractiveness of such patches to the main herbivore, reindeer. Consequently, direction and strength of selection in spatially structured populations may depend significantly on fitness effects arising at the group level.     

Selection against ruderals in Danish grasslands over an eight-year period

In order to investigate what community selection forces are operating on Grime's CSR strategies in different semi-natural grassland habitats that were not subjected to successional processes, a time-series analysis of 2946 Danish grassland plots from an eight-year period with pin-point plant cover data was conducted. Grime's CSR strategies were used for grouping plant species into functional types, which were then treated as dependent variables in a time-series analysis. Across four grassland habitat types, there was significant selection against ruderal plant species during the eight-year period. Furthermore, the monitoring data indicated that the decreasing cover of ruderals may, in part be, due to reduced grazing pressure of large herbivores. As a consequence of the findings in this study and others, it may be suggested that increased disturbance, e.g. by large herbivores, is a suitable management strategies for grassland habitats.