Scale‐dependence of vegetation‐environment relationships in semi‐natural grasslands

Questions: Which environmental and management factors determine plant species composition in semi‐natural grasslands within a local study area? Are vegetation and explanatory factors scale‐dependent? Location: Semi‐natural grasslands in Lærdal, Sognog Fjordane County, western Norway. Methods: We recorded plant species composition and explanatory variables in six grassland sites using a hierarchically nested sampling design with three levels: plots randomly placed within blocks selected within sites. We evaluated vegetation‐environment relationships at all three levels by means of DCA ordination and split‐plot GLM analyses. Results: The most important complex gradient determining variation in grassland species composition showed a broad‐scale relationship with management. Soil moisture conditions were related to vegetation variation on block scale, whereas element concentrations in the soil were significantly related to variation in species composition on all spatial scales. Our results show that vegetation‐environment relationships are dependent on the scale of observation. We suggest that scale‐related (and therefore methodological) issues may explain the wide range of vegetation‐environment relationships reported in the literature, for semi‐natural grassland in particular but also for other ecosystems. Conclusions: Interpretation of the variation in species composition of semi‐natural grasslands requires consideration of the spatial scales on which important environmental variables vary.     

Population dynamics of the perennial Plantago media in semi‐natural grasslands

Abstract. In this study we investigated population dynamics of a perennial forb, Plantago media, in semi‐natural grasslands in southeastern Sweden. Plantago media is a rather common inhabitant of grasslands in this region, but it has been shown to experience dispersal limitations both among managed grassland sites and at potential dispersal routes along road verges. The demography of P. media was studied in 22 permanent plots at two sites over five years. A seed sowing experiment was also performed at each site. The life‐cycle of P. media includes seedlings, juveniles, small vegetative rosettes, large vegetative rosettes and flowering rosettes. The population growth rates (λ‐values) were negative for both populations, with one exception: the first year transition matrix at one site. The projected life span for individuals surviving from seedlings to flowering rosettes varied between 19.6 yr and 227.8 yr. Elasticity analysis showed that remaining in the large rosette stages, both vegetative and flowering, was the most important factor influencing population growth rate. LTRE analysis gave similar results, indicating that spatial and temporal variation have similar effects on the population growth rate. The expected time to extinction in populations with an initial size of between 100 and 1000 individuals varied between 60 and 200 yr. The seed sowing experiment showed that seedling emergence was enhanced by both seed addition and disturbance, suggesting that recruitment is limited by seed and microsite availability. Recruitment was not found to have much impact on population growth rate. However, long‐term population maintenance may depend on occasional small‐scale disturbances that enhance pulses of recruitment. Fragmented landscape is likely to effect plant populations including rare and endangered species as well as more common species which have limited dispersal mechanisms.     

Trait but not species convergence during plant community assembly in restored semi‐natural grasslands

Community assembly or succession was traditionally thought of as being deterministic and directional, leading to a clearly defined climax state. The alternative view, however, keeps gaining attention. This view states that community assembly is influenced by historical processes, where differences in the sequence and timing of species arrival result in distinct communities. Here we tested the hypothesis that both views are valid, but at a different level, with increasing dissimilarity in species composition among sites with increasing age (divergence), caused by historical processes (priority effects), and with increasing similarity in mean trait composition (convergence) among sites, indicating a directional development at the niche level. We surveyed a chronosequence of restored semi‐natural grassland patches on former pine plantations over seven restoration age classes, covering 22 grasslands. Pairwise multivariate distances were calculated between the different grassland patches based on species abundance on the one hand, and on mean community trait values for 28 plant life history traits on the other. Trait composition showed a clear decrease in multivariate distance with increasing restoration age, indicating trait convergence through time. At the species level, we found no evidence of convergence through time, with even a trend towards divergence. Furthermore, spatial variation and environmental heterogeneity were found to remain constant through time. These results confirm our hypothesis. At the trait level, limited niches occur, only filled by species having the appropriate traits, resulting in a clear deterministic model of assembly. Species identity, on the contrary, has no role in this niche filling. The first appropriate species to reach a restoration site will be most likely the ones that get established, resulting in divergence of the species composition among restored grasslands.     

Annual burning of semi‐natural grasslands for conservation: winners and losers among plant species

Species‐rich semi‐natural grasslands in Europe are a main target for conservation efforts, and alternative methods to the traditional management of mowing or grazing would be welcome due to the difficulties in maintaining traditional management practices. One such method proposed is burning of grassland vegetation during late winter or spring. To evaluate the effects of annual spring burning vs annual mowing on semi‐natural grassland vegetation, we compared the frequency of species in eleven field experiments in southern Sweden after ca 14 years. Out of the 88 species analyzed, five were more frequent in burnt plots compared with mowed plots (Vicia cracca, Cirsium arvense, Urtica dioica, Galium verum, Convallaria majalis). In contrast, 37 species were significantly less frequent in burnt plots compared with mowed ones, those with the largest differences being Ranunculus acris, Briza media, Veronica chamaedrys, Festuca ovina, Plantago lanceolata and Anthoxanthum odoratum. Tall‐grown species and those with preferences for N‐rich soils increased in frequency under an annual spring‐burn regime, compared with annual mowing, as did species producing larger amounts of nectar. Hence, although vegetation composition becomes more trivial with annual spring burns, there might be long‐term benefits for nectar‐feeding insects.     

Rediscovery of Celes akitanus (Orthoptera: Acrididae) from semi‐natural grasslands in Japan

Celes akitanus was rediscovered in semi‐natural grasslands in Japan. This species is one of the most endangered insects in Japan; most local populations were thought to be extinct. Reliable records of this rare species had been obtained only from Yamagata, Niigata, Tokyo and Nagano Prefectures. We compared the morphology of the holotype of Oedipoda akitana (=C. akitanus) and newly collected specimens and found that they matched with each other. The species was redescribed based on the holotype and new specimens. A molecular phylogenetic analysis involving other acridid species indicated that our samples were practically identical to C. akitanus from China. We conclude that the specimens we collected are C. akitanus, which has been rediscovered in Japan after about 30 years. Because the discovered populations were restricted to narrow and scattered areas, conservation of these populations should be a high priority. Population monitoring and habitat assessment are required. Because C. akitanus and many other endangered species are found in the grassland areas, these habitats should be protected and maintained.     

Size and heterogeneity rather than landscape context determine plant species richness in semi‐natural grasslands

Question: Is plant diversity in fragmented semi‐natural grasslands related to present and historical landscape context? Location: Southern Sweden. Methods: Plant diversity was described at 30 semi‐natural grassland sites in terms of total and specialist plant species richness at the site and species density at different scales (0.5–10 m²). These measures are commonly used to assess conservation value of semi‐natural grasslands. Landscape context was measured as contemporary connectivity to other semi‐natural grasslands, historical connectivity 50 years ago, amount of linear elements potentially suitable for dispersal (road verges, power line clearings), and amount of forest (inverse of the openness of the landscape). Results: The diversity measures were generally correlated with each other, implying that species richness in a subset of the grassland can predict the total richness. Plant species density at three scales (0.5 m², 10 m² and total) was related to the landscape context using an information theoretic approach. Results showed that total species richness increased with increased size of grasslands, contrary to earlier diversity studies in semi‐natural grasslands. Larger grasslands were more heterogeneous than smaller grasslands, and this is a likely reason for the species‐area relationship. Heterogeneity was also of high importance at the smaller scales (0.5 m², 10 m²). With increased amount of forest, total species richness increased but species density on 10 m² decreased. There was no influence of connectivity in either the contemporary or the historical landscape, contrary to previous studies. Conclusions: Grassland size and heterogeneity are of greater importance for plant diversity in semi‐natural grassland, than grassland connectivity in the landscape.