Environmental and socio-economic resources at the landscape level - Potentials for sustainable land use in the Georgian Greater Caucasus.

Mountain regions cover one quarter of the Earth's terrestrial surface, and are both valuable and vulnerable areas with complex human-environmental interrelationships. In this coupled system, land-use changes induced by political or socio-economic transformations generate consequences for ecological landscape functions like soil productivity and species richness, and integrative land-use concepts provide the potential of a sustainable land development. In the Kazbegi region in the central Greater Caucasus of Georgia, these transformations further lead to landscape-structure change and population marginalization. Hence, we developed three agricultural land-use scenarios that meet Agenda 2030 Sustainable Development Goals to ensure a sustainable rural land development and the conservation of mountain ecosystems. Our normative scenario approach integrates quantitative and qualitative findings of empirical research in landscape ecology, soil science, vegetation ecology as well as agronomics and socio-economics. According to the examined environmental and socio-economic resources, we defined various scenario logics and normative assumptions that combine optimized livestock production (in dairy cow keeping and cattle fattening) with ecological limitations to maintain the functioning of mountain ecosystems. The rule-based scenarios achieved measurably increased outputs in biomass yields, livestock production and related revenues at the regional scale. Further, GIS generated scenario maps demonstrate the related land-use patterns spatially explicit and in high resolution, and visualize the alternative future from local to the regional scale. In conclusion, scenario development helps to determine region-specific and integrated land-use options to provide a sound base for land users and decision makers. Based on research on multiple landscape functions, this approach can assist sustainable land development in a mountain region.

New insights into island vegetation composition and species diversity-Consistent and conditional responses across contrasting insular habitats at the plot-scale.

Most island-ecology studies focus on the properties of entire island communities, thus neglecting species-environment relationships operating at the habitat-level. Habitat-specific variation in the strength and sign of these relationships will conceal patterns observed on the island scale and may preclude a mechanistic interpretation of patterns and processes. Habitat-specific species-environment relationships may also depend on the descriptor of ecological communities. This paper presents a comprehensive plot-based analysis of local vegetation composition and species diversity (species richness and species evenness) of (i) rocky shore, (ii) semi-natural grassland and (iii) coniferous forest habitats in three Baltic archipelagos in Sweden. To identify differences and consistencies between habitats and descriptors, we assessed the relative contributions of the variable-sets "region", "topography", "soil morphology", "soil fertility", "soil water", "light availability", "distance" and "island configuration" on local vegetation composition, species richness and species evenness. We quantified the impact of "management history" on the descriptors of local grassland communities by a newly introduced grazing history index (GHI). Unlike species diversity, changes in vegetation composition were related to most of the variable-sets. The relative contributions of the variable-sets were mostly habitat-specific and strongly contingent on the descriptor involved. Within each habitat, richness and evenness were only partly affected by the same variable-sets, and if so, their relative contribution varied between diversity proxies. Across all habitats, soil variable-sets showed highly consistent effects on vegetation composition and species diversity and contributed most to the variance explained. GHI was a powerful predictor, explaining high proportions of variation in all three descriptors of grassland species communities. The proportion of unexplained variance was habitat-specific, possibly reflecting a community maturity gradient. Our results reveal that species richness alone is an incomplete representation of local species diversity. Finally, we stress the need of including habitat-based approaches when analyzing complex species-environment relationships on islands.

Restoration of floodplain meadows: Effects on the re-establishment of mosses.

Vascular plants serve as target species for the evaluation of restoration success as they account for most of the plant species diversity and vegetation cover. Although bryophytes contribute considerably to the species diversity of meadows, they are rarely addressed in restoration projects. This project is a first step toward making recommendations for including mosses in alluvial floodplain restoration projects. The opportunity to assess the diversity and ecological requirements of mosses on floodplain meadows presented itself within the framework of a vegetation monitoring that took place in 2014 on meadows located along the northern Upper Rhine. In this area, large-scale meadow restoration projects have taken place since 1997 in both the functional and fossil floodplains. Other studies have shown that bryophytes are generally present in green hay used in restoration, providing inadvertent bryophyte introduction. We compared bryophyte communities in donor and restored communities and correlated these communities with environmental variables-taking into account that the mosses on the restoration sites possibly developed from green hay. This analysis provided insights as to which species of bryophytes should be included in future restoration projects, what diaspores should be used, and how they should be transferred. Data on bryophyte occurrence were gathered from old meadows, and from restoration sites. We found distinct differences in bryophyte composition (based on frequency) in restored communities in functional flood plains compared to donor communities. Generally, restoration sites are still characterized by a lower species-richness, with a significantly lower occurrence of rare and red listed species and a lower species-heterogeneity. In conclusion, our research establishes what mosses predominate in donor and restored alluvial meadows along the northern Upper River, and what microsite conditions favour particular species. This points the way to deliberate introduction of moss diaspores for more complete alluvial meadow restoration.

Flooding tolerance of four floodplain meadow species depends on age.

Numerous restoration campaigns focused on re-establishing species-rich floodplain meadows of Central Europe, whose species composition is essentially controlled by regular flooding. Climate change predictions expect strong alterations on the discharge regime of Europe's large rivers with little-known consequences on floodplain meadow plants. In this study, we aim to determine the effects of flooding on seedlings of different ages of four typical flood meadow species. To this end, we flooded seedlings of two familial pairs of flood meadow species of wetter and dryer microhabitats for 2 weeks each, starting 2, 4, 6, and 8 weeks after seedling germination, respectively. We show that a 2-week-flooding treatment had a negative effect on performance of seedlings younger than 6 weeks. Summer floods with high floodwater temperatures may have especially detrimental effects on seedlings, which is corroborated by previous findings. As expected, the plants from wet floodplain meadow microhabitats coped better with the flooding treatment than those from dryer microhabitats. In conclusion, our results suggest that restoration measures may perform more successfully if seedlings of restored species are older than the critical age of about 6 weeks before a spring flooding begins. Seasonal flow patterns may influence vegetation dynamics of floodplain meadows and should, therefore, be taken into account when timing future restoration campaigns.

Clonality increases with snow depth in the arctic dwarf shrub Empetrum hermaphroditum.

PREMISE OF THE STUDY: Vegetative reproduction and spread through clonal growth plays an important role in arctic-alpine ecosystems with short cool growing seasons. Local variation in winter snow accumulation leads to discrete habitat types that may provide divergent conditions for sexual and vegetative reproduction. Therefore, we studied variation in clonal structure of a dominant, evergreen, dwarf shrub (Empetrum nigrum s.l. with the two taxa E. nigrum L. s.s. and E. hermaphroditum Hagerup) along a snow cover gradient and compared clonal diversity and spatial genetic structure between habitats. METHODS: We studied 374 individual shoots using 105 polymorphic AFLP markers and analyses based on hierarchical clustering, clonal diversity indices, and small-scale spatial genetic structure with pairwise kinship coefficient. We used two approaches to define a threshold of genotypic distance between two samples that are considered the same clone. Clonality was examined among three habitats (exposed ridges, sheltered depressions, birch forest) differing in snow conditions replicated in four study regions in Norway and Sweden. KEY RESULTS: Clonality of E. hermaphroditum differed between habitats with an increase in clonal diversity with decreasing snow depth. Small-scale spatial genetic structure increased with decreasing clonal diversity and increasing clone size. In three study regions, E. hermaphroditum was the only species, whereas in one region E. nigrum also occurred, largely confined to exposed ridges. CONCLUSIONS: Our results demonstrated that snow cover in conjunction with associated habitat conditions plays an important role for the mode of propagation of the dwarf shrub E. hermaphroditum.

Effects of reduced summer precipitation on productivity and forage quality of floodplain meadows at the Elbe and the Rhine River.

BACKGROUND: Floodplain meadows along rivers are semi-natural habitats and depend on regular land use. When used non-intensively, they offer suitable habitats for many plant species including rare ones. Floodplains are hydrologically dynamic ecosystems with both periods of flooding and of dry conditions. In German floodplains, dry periods may increase due to reduced summer precipitation as projected by climate change scenarios. Against this background, the question arises, how the forage quantity and quality of these meadows might change in future. METHODS: We report results of two field trials that investigated effects of experimentally reduced summer precipitation on hay quantity and quality of floodplain meadows at the Rhine River (2011-2012) and at two Elbe tributaries (2009-2011). We measured annual yield, the amount of hay biomass, and contents of crude protein, crude fibre, energy, fructan, nitrogen, phosphorus, and potassium. RESULTS: The annual yield decreased under precipitation reduction at the Rhine River. This was due to reduced productivity in the second cut hay at the Rhine River in which, interestingly, the contents of nitrogen and crude protein increased. The first cut at the Rhine River was unaffected by the treatments. At the Elbe tributaries, the annual yield and the hay quantity and quality of both cuts were only marginally affected by the treatments. CONCLUSION: We conclude that the yield of floodplain meadows may become less reliable in future since the annual yield decreased under precipitation reduction at the Rhine River. However, the first and agriculturally more important cut was almost unaffected by the precipitation reduction, which is probably due to sufficient soil moisture from winter/spring. As long as future water levels of the rivers will not decrease during spring, at least the use of the hay from the first cut of floodplain meadows appears reliable under climate change.

Invasibility of a nutrient-poor pasture through resident and non-resident herbs is controlled by litter, gap size and propagule pressure.

Since inference concerning the relative effects of propagule pressure, biotic interactions, site conditions and species traits on the invasibility of plant communities is limited, we carried out a field experiment to study the role of these factors for absolute and relative seedling emergence in three resident and three non-resident confamilial herb species on a nutrient-poor temperate pasture. We set up a factorial field experiment with two levels each of the factors litter cover (0 and 400 g m(-2)), gap size (0.01 and 0.1 m(2)) and propagule pressure (5 and 50 seeds) and documented soil temperature, soil water content and relative light availability. Recruitment was recorded in spring and autumn 2010 and in spring 2011 to cover initial seedling emergence, establishment after summer drought and final establishment after the first winter. Litter alleviated temperature and moisture conditions and had positive effects on proportional and absolute seedling emergence during all phases of recruitment. Large gaps presented competition-free space with high light availability but showed higher temperature amplitudes and lower soil moisture. Proportional and absolute seedling recruitment was significantly higher in large than in small gaps. In contrast, propagule pressure facilitated absolute seedling emergence but had no effects on proportional emergence or the chance for successful colonisation. Despite significantly higher initial seedling emergence of resident than non-resident species, seed mass and other species-specific traits may be better predictors for idiosyncratic variation in seedling establishment than status. Our data support the fluctuating resource hypothesis and demonstrate that the reserve effect of seeds may facilitate seedling emergence. The direct comparison of propagule pressure with other environmental factors showed that propagule pressure affects absolute seedling abundance, which may be crucial for species that depend on other individuals for sexual reproduction. However, propagule batch size did not significantly affect the chance for successful colonisation of disturbed plots.

Potential effects of direct transfer payments on farmland habitat diversity in a marginal European landscape.

Farmland habitat diversity in marginal European landscapes changed significantly in the past decades. Further changes toward homogenization are expected, particularly in the course of European agricultural policy. Based on three alternative transfer payment schemes, we modeled spatially explicit potential effects on the farmland habitat diversity in a marginal European landscape. We defined (1) a scenario with direct transfer payments coupled to production, (2) a scenario with direct transfer payments decoupled from production, and (3) a scenario phasing out all direct transfer payments. We characterized habitat diversity with three indices: habitat richness, evenness, and rarity. The habitat pattern in 1995 served as reference for comparison. All scenarios predicted a general trend of homogenization of the farmland habitat pattern, yet to a differing extent. Transfer payments coupled to production (Scenario 1) favored the abandonment of agricultural production, particularly in low-productive areas and arable land use in more productive areas. Habitat richness and habitat evenness had intermediate values in this scenario. Decoupling transfer payments from production (Scenario 2) supported grassland as most profitable farming system. This led to a grassland-dominated landscape with low values of all habitat diversity indices. Phasing out transfer payments (Scenario 3) resulted in complete abandonment or afforestation of agricultural land and extremely low values in all habitat diversity indices. Scenario results indicate that transfer payments may prevent cessation of agricultural production, but may not counteract homogenization in marginal landscapes. Conserving high farmland habitat diversity in such landscapes may require support schemes, e.g., Pillar Two of EU Common Agricultural Policy.

Assessing the spatial distribution of grassland age in a marginal European landscape.

Grassland age is increasingly recognised to be an indicator for present-day biodiversity, e.g. plant species richness, and is also important for other landscape functions. We developed a methodological approach to systematically assess the spatial distribution of grassland age in marginal European landscapes. This approach - applied to the Lahn-Dill Highlands (1270 km(2)), a marginal landscape in Hesse, Germany - comprises three steps: (1) in a two-stage stratification process, we pre-stratified the study area according to recent land-cover patterns and their changes between 1955 and 1995 (stratification I) and classified grassland types by combining data on soil moisture, base-richness, and elevation (stratification II). From 50 grassland types, we randomly selected 1000 representative grassland patches. (2) We determined the age of these patches by means of aerial photograph interpretation of a chronosequence dating back to 1953 and classified each patch with respect to the age classes young (<18 years), mid-aged (18-47 years), and old (>47 years). (3) Based on this information, we calculated grassland type-specific probabilities for grassland patches to belong to the respective age classes. These probabilities were projected to districts by direct extrapolation. An exemplary validation of extrapolation results for two test areas was performed. The results revealed that 49% of the investigated patches were old grassland. The remaining patches were mid-aged (36%) or young grassland (15%). The extrapolation results indicated accordingly a predominance of old grassland at the district scale. Occurrences of mid-aged grassland were concentrated in districts with a pronounced land-cover change, whereas young grassland is apparently evenly distributed across the study area. Validation results suggest that our approach is suitable for a realistic estimation of grassland age in marginal European landscapes. The method may be applied in landscape models of various disciplines that rely on large-scale information on grassland age.

Linking socio-economic factors, environment and land cover in the German Highlands, 1945-1999.

Land-cover changes are caused by human activities and natural ecological processes. Thus, our study uses an interdisciplinary approach to research land-cover changes. We present a method to (i) link socio-economic/environmental factors and land-cover changes, (ii) identify indicators of land-cover changes, and (iii) distinguish between socio-economic and environmental indicators associated with local types of overall land-cover changes. The study was conducted in the Lahn-Dill Highlands, Germany, a typical marginal rural landscape. In this region, we investigated land-cover changes occurring over the period 1945-1999. Land-cover data were derived from multi-temporal aerial photographs. Types of overall land-cover changes characterising the districts within the study area were differentiated. With the help of redundancy analysis (RDA), we analysed the relationships between land-cover changes and widely available socio-economic/environmental factors. The results reveal that both individual land-cover changes at patch level and types of overall land-cover changes characterising districts are correlated with socio-economic and environmental factors. Whereas the stable environmental factors are drivers of land-cover changes in our rural study area, socio-economic factors introduced into the analysis mostly result from land-cover changes. We identified correlative socio-economic indicators that cannot explain land-cover changes, but that in combination with the environmental factors can be used to greatly facilitate the reconstruction of past land-cover changes and thus lead to a better knowledge of land-cover history. Based on the types of overall land-cover changes, the results of the study can be adopted for the study of land-cover changes in other regions.