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1.Environmental changes may not always result in rapid changes in species distributions, abundances or diversity. In order to estimate the effects of, for example, land-use changes caused by agri-environment schemes (AES) on biodiversity and ecosystem services, information on the time-lag between the application of the scheme and the responses of organisms is essential.2.We examined the effects of time since transition (TST) to organic farming on plant species richness and butterfly species richness and abundance. Surveys were conducted in cereal fields and adjacent field margins on 60 farms, 20 conventional and 40 organic, in two regions in Sweden. The organic farms were transferred from conventional management between 1 and 25 years before the survey took place. The farms were selected along a gradient of landscape complexity, indicated by the proportion of arable land, so that farms with similar TST were represented in all landscape types. Organism responses were assessed using model averaging.3.Plant and butterfly species richness was c.20% higher on organic farms and butterfly abundance was about 60% higher, compared with conventional farms. Time since transition affected butterfly abundance gradually over the 25-year period, resulting in a 100% increase. In contrast, no TST effect on plant or butterfly species richness was found, indicating that the main effect took place immediately after the transition to organic farming.4.Increasing landscape complexity had a positive effect on butterfly species richness, but not on butterfly abundance or plant species richness. There was no indication that the speed of response to organic farming was affected by landscape complexity.5.Synthesis and applications. The effect of organic farming on diversity was rapid for plant and butterfly species richness, whereas butterfly abundance increased gradually with time since transition. If time-lags in responses to AESs turn out to be common, long-term effects would need to be included in management recommendations and policy to capture the full potential of such schemes.

There is a lack of quantitative syntheses of fragmentation effects across species and biogeographic regions, especially with respect to species life-history traits. We used data from 24 independent studies of butterflies and moths from a wide range of habitats and landscapes in Europe and North America to test whether traits associated with dispersal capacity, niche breadth and reproductive rate modify the effect of habitat fragmentation on species richness. Overall, species richness increased with habitat patch area and connectivity. Life-history traits improved the explanatory power of the statistical models considerably and modified the butterfly species-area relationship. Species with low mobility, a narrow feeding niche and low reproduction were most strongly affected by habitat loss. This demonstrates the importance of considering life-history traits in fragmentation studies and implies that both species richness and composition change in a predictable manner with habitat loss and fragmentation.

Habitat loss poses a major threat to biodiversity, and species-specific extinction risks are inextricably linked to life-history characteristics. This relationship is still poorly documented for many functionally important taxa, and at larger continental scales. With data from five replicated field studies from three countries, we examined how species richness of wild bees varies with habitat patch size. We hypothesized that the form of this relationship is affected by body size, degree of host plant specialization and sociality. Across all species, we found a positive species-area slope (z = 0.19), and species traits modified this relationship. Large-bodied generalists had a lower z value than small generalists. Contrary to predictions, small specialists had similar or slightly lower z value compared with large specialists, and small generalists also tended to be more strongly affected by habitat loss as compared with small specialists. Social bees were negatively affected by habitat loss (z = 0.11) irrespective of body size. We conclude that habitat loss leads to clear shifts in the species composition of wild bee communities.

The population dynamics of organisms living in short-lived habitats will largely depend on the turnover of habitat patches. It has been suggested that epiphytes, whose host plants can be regarded as habitat patches, often form such patch-tracking populations. However, very little is known about the long-term fate of epiphyte individuals and populations. We estimated life span and assessed environmental factors influencing changes in vitality, fertility, abundance, and distribution of the epiphytic lichen species Lobaria pulmonaria on two spatial scales, individual trees and forest patches, over a period of approximately 10 years in 66 old-growth forest fragments. The lichen had gone extinct from 7 of the 66 sites (13.0%) where it was found 10 years earlier, even though the sites remained unchanged. The risk of local population extinction increased with decreasing population size. In contrast to the decrease in the number of occupied trees and sites, the mean area of the lichen per tree increased by 43.0%. The number of trees with fertile ramets of L. pulmonaria increased from 7 (approximately 1%) to 61 (approximately 10%) trees, and the number of forest fragments with fertile ramets increased from 4 to 23 fragments. The mean annual rate of L. pulmonaria extinction at the tree level was estimated to be 2.52%, translating into an expected lifetime of 39.7 years. This disappearance rate is higher than estimated mortality rates for potential host trees. The risk of extinction at the tree level was significantly positively related to tree circumference and differed between tree species. The probability of presence of fertile ramets increased significantly with local population size. Our results show a long expected lifetime of Lobaria pulmonaria ramets on individual trees and a recent increase in vitality, probably due to decreasing air pollution. The population is, however, declining slowly even though remaining stands are left uncut, which we interpret as an extinction debt.

Intensification or abandonment of agricultural land use has led to a severe decline of semi-natural habitats across Europe. This can cause immediate loss of species but also time-delayed extinctions, known as the extinction debt. In a pan-European study of 147 fragmented grassland remnants, we found differences in the extinction debt of species from different trophic levels. Present-day species richness of long-lived vascular plant specialists was better explained by past than current landscape patterns, indicating an extinction debt. In contrast, short-lived butterfly specialists showed no evidence for an extinction debt at a time scale of c. 40 years. Our results indicate that management strategies maintaining the status quo of fragmented habitats are insufficient, as time-delayed extinctions and associated co-extinctions will lead to further biodiversity loss in the future.

Local extinction of species can occur with a substantial delay following habitat loss or degradation. Accumulating evidence suggests that such extinction debts pose a significant but often unrecognized challenge for biodiversity conservation across a wide range of taxa and ecosystems. Species with long generation times and populations near their extinction threshold are most likely to have an extinction debt. However, as long as a species that is predicted to become extinct still persists, there is time for conservation measures such as habitat restoration and landscape management. Standardized long-term monitoring, more high-quality empirical studies on different taxa and ecosystems and further development of analytical methods will help to better quantify extinction debt and protect biodiversity.

During the last 50 years, the distribution and abundance of many European butterfly species associated with semi-natural grasslands have declined. This may be the result of deteriorating habitat quality, but habitat loss, resulting in decreasing area and increasing isolation of remaining habitat, is also predicted to result in reduced species richness. To investigate the effects of habitat loss on species richness, we surveyed butterflies in semi-natural grasslands of similar quality and structure, but situated in landscapes of different habitat composition. Using spatially explicit habitat data, we selected one large (6-10 ha) and one small (0.5-2 ha) grassland site (pasture) in each of 24 non-overlapping 28.2 km(2) landscapes belonging to three categories differing in the proportion of the area that consisted of semi-natural grasslands. After controlling for local habitat quality, species richness was higher in grassland sites situated in landscapes consisting of a high proportion of grasslands. Species richness was also higher in larger grassland sites, and this effect was more pronounced for sedentary than for mobile species. However, the number of species for a given area did not differ between large and small grasslands. There was also a significant relationship between butterfly species richness and habitat quality in the form of vegetation height and abundance of flowers. In contrast, butterfly density was not related to landscape composition or grassland size. When species respond differently to habitat area or landscape composition this leads to effects on community structure, and nestedness analysis showed that depauperate communities were subsets of richer ones. Both grassland area and landscape composition may have contributed to this pattern, implying that small habitat fragments and landscapes with low proportions of habitat are both likely to mainly contain common generalist species. Based on these results, conservation efforts should aim at preserving landscapes with high proportions of the focal habitat.