Resource-Mediated Indirect Effects of Grassland Management on Arthropod Diversity

Intensive land use is a driving force for biodiversity decline in many ecosystems. In semi-natural grasslands, land-use activities such as mowing, grazing and fertilization affect the diversity of plants and arthropods, but the combined effects of different drivers and the chain of effects are largely unknown. In this study we used structural equation modelling to analyse how the arthropod communities in managed grasslands respond to land use and whether these responses are mediated through changes in resource diversity or resource quantity (biomass). Plants were considered resources for herbivores which themselves were considered resources for predators. Plant and arthropod (herbivores and predators) communities were sampled on 141 meadows, pastures and mown pastures within three regions in Germany in 2008 and 2009. Increasing land-use intensity generally increased plant biomass and decreased plant diversity, mainly through increasing fertilization. Herbivore diversity decreased together with plant diversity but showed no response to changes in plant biomass. Hence, land-use effects on herbivore diversity were mediated through resource diversity rather than quantity. Land-use effects on predator diversity were mediated by both herbivore diversity (resource diversity) and herbivore quantity (herbivore biomass), but indirect effects through resource quantity were stronger. Our findings highlight the importance of assessing both direct and indirect effects of land-use intensity and mode on different trophic levels. In addition to the overall effects, there were subtle differences between the different regions, pointing to the importance of regional land-use specificities. Our study underlines the commonly observed strong effect of grassland land use on biodiversity. It also highlights that mechanistic approaches help us to understand how different land-use modes affect biodiversity.

Biotic resistance to plant invasion in grassland: Does seed predation increase with resident plant diversity?

Seed predation impacts heavily on plant populations and community composition in grasslands. In particular, generalist seed predators may contribute to biotic resistance, i.e. the ability of resident species in a community to reduce the success of non-indigenous plant invaders. However, little is known of predators' preferences for seeds of indigenous or non-indigenous plant species or how seed predation varies across communities. We hypothesize that seed predation does not differ between indigenous and non-indigenous plant species and that seed predation is positively related to plant species diversity in the resident community. The seed removal of 36 indigenous and non-indigenous grassland species in seven extensively or intensively managed hay meadows across Switzerland covering a species-richness gradient of 18-50 plant species per unit area (c. 2 m(2)) was studied. In mid-summer 2011, c. 24,000 seeds were exposed to predators in Petri dishes filled with sterilized soil, and the proportions of seeds removed were determined after three days' exposure. These proportions varied among species (9.2-62.5%) and hay meadows (17.8-48.6%). Seed removal was not related to seed size. Moreover, it did not differ between indigenous and non-indigenous species, suggesting that mainly generalist seed predators were active. However, seed predation was positively related to plant species richness across a gradient in the range of 18-38 species per unit area, representing common hay meadows in Switzerland. Our results suggest that generalist post-dispersal seed predation contributes to biotic resistance and may act as a filter to plant invasion by reducing the propagule pressure of non-local plant species.

Biotic and Abiotic Properties Mediating Plant Diversity Effects on Soil Microbial Communities in an Experimental Grassland

Plant diversity drives changes in the soil microbial community which may result in alterations in ecosystem functions. However, the governing factors between the composition of soil microbial communities and plant diversity are not well understood. We investigated the impact of plant diversity (plant species richness and functional group richness) and plant functional group identity on soil microbial biomass and soil microbial community structure in experimental grassland ecosystems. Total microbial biomass and community structure were determined by phospholipid fatty acid (PLFA) analysis. The diversity gradient covered 1, 2, 4, 8, 16 and 60 plant species and 1, 2, 3 and 4 plant functional groups (grasses, legumes, small herbs and tall herbs). In May 2007, soil samples were taken from experimental plots and from nearby fields and meadows. Beside soil texture, plant species richness was the main driver of soil microbial biomass. Structural equation modeling revealed that the positive plant diversity effect was mainly mediated by higher leaf area index resulting in higher soil moisture in the top soil layer. The fungal-to-bacterial biomass ratio was positively affected by plant functional group richness and negatively by the presence of legumes. Bacteria were more closely related to abiotic differences caused by plant diversity, while fungi were more affected by plant-derived organic matter inputs. We found diverse plant communities promoted faster transition of soil microbial communities typical for arable land towards grassland communities. Although some mechanisms underlying the plant diversity effect on soil microorganisms could be identified, future studies have to determine plant traits shaping soil microbial community structure. We suspect differences in root traits among different plant communities, such as root turnover rates and chemical composition of root exudates, to structure soil microbial communities.

Experimental evidence for the immediate impact of fertilization and irrigation upon the plant and invertebrate communities of mountain grasslands

The response of montane and subalpine hay meadow plant and arthropod communities to the application of liquid manure and aerial irrigation – two novel, rapidly spreading management practices – remains poorly understood, which hampers the formulation of best practice management recommendations for both hay production and biodiversity preservation. In these nutrient-poor mountain grasslands, a moderate management regime could enhance overall conditions for biodiversity. This study experimentally assessed, at the site scale, among low-input montane and subalpine meadows, the short-term effects (1 year) of a moderate intensification (slurry fertilization: 26.7–53.3 kg N·ha−1·year−1; irrigation with sprinklers: 20 mm·week−1; singly or combined together) on plant species richness, vegetation structure, hay production, and arthropod abundance and biomass in the inner European Alps (Valais, SW Switzerland). Results show that (1) montane and subalpine hay meadow ecological communities respond very rapidly to an intensification of management practices; (2) on a short-term basis, a moderate intensification of very low-input hay meadows has positive effects on plant species richness, vegetation structure, hay production, and arthropod abundance and biomass; (3) vegetation structure is likely to be the key factor limiting arthropod abundance and biomass. Our ongoing experiments will in the longer term identify which level of management intensity achieves an optimal balance between biodiversity and hay production.

Promoting Pollinating Insects in Intensive Agricultural Matrices: Field-Scale Experimental Manipulation of Hay-Meadow Mowing Regimes and Its Effects on Bees

Bees are a key component of biodiversity as they ensure a crucial ecosystem service: pollination. This ecosystem service is nowadays threatened, because bees suffer from agricultural intensification. Yet, bees rarely benefit from the measures established to promote biodiversity in farmland, such as agri-environment schemes (AES). We experimentally tested if the spatio-temporal modification of mowing regimes within extensively managed hay meadows, a widespread AES, can promote bees. We applied a randomized block design, replicated 12 times across the Swiss lowlands, that consisted of three different mowing treatments: 1) first cut not before 15 June (conventional regime for meadows within Swiss AES); 2) first cut not before 15 June, as treatment 1 but with 15% of area left uncut serving as a refuge; 3) first cut not before 15 July. Bees were collected with pan traps, twice during the vegetation season (before and after mowing). Wild bee abundance and species richness significantly increased in meadows where uncut refuges were left, in comparison to meadows without refuges: there was both an immediate (within year) and cumulative (from one year to the following) positive effect of the uncut refuge treatment. An immediate positive effect of delayed mowing was also evidenced in both wild bees and honey bees. Conventional AES could easily accommodate such a simple management prescription that promotes farmland biodiversity and is likely to enhance pollination services.

Conservation of threatened habitat types under future climate change – Lessons from plant-distribution models and current extinction trends in southern Germany

A higher risk of future range losses as a result of climate change is expected to be one of the main drivers of extinction trends in vascular plants occurring in habitat types of high conservation value. Nevertheless, the impact of the climate changes of the last 60 years on the current distribution and extinction patterns of plants is still largely unclear. We applied species distribution models to study the impact of environmental variables (climate, soil conditions, land cover, topography), on the current distribution of 18 vascular plant species characteristic of three threatened habitat types in southern Germany: (i) xero-thermophilous vegetation, (ii) mesophilous mountain grasslands (mountain hay meadows and matgrass communities), and (iii) wetland habitats (bogs, fens, and wet meadows). Climate and soil variables were the most important variables affecting plant distributions at a spatial level of 10 × 10 km. Extinction trends in our study area revealed that plant species which occur in wetland habitats faced higher extinction risks than those in xero-thermophilous vegetation, with the risk for species in mesophilous mountain grasslands being intermediary. For three plant species characteristic either of mesophilous mountain grasslands or wetland habitats we showed exemplarily that extinctions from 1950 to the present day have occurred at the edge of the species’ current climatic niche, indicating that climate change has likely been the main driver of extinction. This is largely consistent with current extinction trends reported in other studies. Our study indicates that the analysis of past extinctions is an appropriate means to assess the impact of climate change on species and that vulnerability to climate change is both species- and habitat-specific.

Zur Langzeitökologie des Lärchen-Arvengürtels in den südlichen Walliser Alpen

Palaeoecological studies in the "Alpe d'Essertse" area have provided much information about Vegetation changes and timberline fluctuations during the Holocene In this study we repeated previous biostratigraphic investigations using plant macrofossils to improve their temporal and taxonomie resolution and to test their reliability. By analyzing 0.5-cm layers of a lake sediment we reached a temporal resolution of 44 years, and we were able to reconstruct Vegetation changes in the surrounding area at species level. The sedimentary record analyzed extends from the Late-Glacial to the late Holocene Alpine grasslands (12'000-11'000 cal. BP) were afforested by Larix decidua, Juniperus nana, and Pinus cembra (11'000-9'600 cal. B.P). Stable subalpine larch-stone pine-forests (9'600^4'900 cal. BP) were followed by shrublands and meadows as a consequence of the climatically and anthropogenically induced destruction of forest Vegetation (4'900-2'600 cal. BP). Changes in the abundance of P. cembra and L. decidua needles as well as changes of the other taxa were consistent with those found in previous studies from the same lake. Our results demonstrate that plant-macrofossil records can be reproduced spatially and temporally on separate cores with independent 14C chronologies.

Middle to Late Holocene vegetation history of the Upper Engadine (Swiss Alps): the role of man and fire

o reconstruct the vegetation and fire history of the Upper Engadine, two continuous sediment cores from Lej da Champfèr and Lej da San Murezzan (Upper Engadine Valley, southeastern Switzerland) were analysed for pollen, plant macrofossils, charcoal and kerogen. The chronologies of the cores are based on 38 radiocarbon dates. Pollen and macrofossil data suggest a rapid afforestation with Betula, Pinus sylvestris, Pinus cembra, and Larix decidua after the retreat of the glaciers from the lake catchments 11,000 cal years ago. This vegetation type persisted until ca. 7300 cal b.p. (5350 b.c.) when Picea replaced Pinus cembra. Pollen indicative of human impact suggests that in this high-mountain region of the central Alps strong anthropogenic activities began during the Early Bronze Age (3900 cal b.p., 1950 b.c.). Local human settlements led to vegetational changes, promoting the expansion of Larix decidua and Alnus viridis. In the case of Larix, continuing land use and especially grazing after fire led to the formation of Larix meadows. The expansion of Alnus viridis was directly induced by fire, as evidenced by time-series analysis. Subsequently, the process of forest conversion into open landscapes continued for millennia and reached its maximum at the end of the Middle Ages at around 500 cal b.p. (a.d. 1450).

Treeline Fluctuations Recorded for 12,500 Years by Soil Profiles, Pollen, and Plant Macrofossils in the Central Swiss Alps

Past treelines can rarely be recorded by pollen percentages alone, but pollen concentration, pollen influx, and plant macrofossils (including stomata of conifers) are more reliable indicators. In addition, ancient forest soils above today's treeline may trace the maximum upper expansion of the forest since the last glaciation. Charcoal in such soil profiles may be radiocarbon dated. Our example from the Central Swiss Alps at the Alpe d'Essertse consists of a plant-macrofossil diagram and pollen diagrams of the pond Gouille Rion at 2343 m a.s.l. and a sequence of soil profiles from 1780 m to 2600 m a.s.l. The area around the pond was forested with LariJc decidua and Pinus cembra between 9500 and 3600 BP. After 4700 BP the forest became more open and Juniperus nana and Alnus viridis expanded (together with Picea abies in the subalpine forest). Between 1700 and 900 BP the Juniperus nana and Alnus viridis scrubs declined while meadows and pastures took over, so that the pond Gouille Rion was definitively above timber­ line. The highest Holocene treeline was at 2400 to 2450 m a.s.l. (i.e. 50 to 100 m higher than the uppermost single specimen of Pinus cembra today) between 9000 and 4700 BP, but it is not yet dated in more detail. The highest charcoal of Pinus cembra at 2380 m a.s.l. has a radiocarbon date of 6010 ± 70 BP. Around 6900 BP a strong climatic deterioration caused an opening of timberline forest. First indicators of anthropogenic influence occurred at 4700 BP, when the forest limit started to move down. The lowering of timberline after 4700 BP was probably due to combined effects of human and climatic impact.

Lichen species richness is highest in non-intensively used grasslands promoting suitable microhabitats and low vascular plant competition

Lichens are very sensitive to habitat changes and their species richness is likely to decline under intensive land use. Currently, a comprehensive study analyzing lichen species richness in relation to land-use types, extending over different regions and including information on habitat variables, is missing for temperate grasslands. In three German regions we studied lichen species richness in 490 plots of 16 m2 representing different land-use types, livestock types, and habitat variables. Due to the absence of low-intensity pastures and substrates such as woody plants, deadwood and stones, there were no lichens in the 78 plots in Schorfheide-Chorin. In the two other regions, the richness of lichen species was 45 % higher in pastures than in meadows, and 77 % higher than in mown pastures, respectively. Among the pastures, the richness of all lichen species was on average 10 times higher in sheep-grazed pastures than in the ones grazed by cattle or horses. On average, the richness of all lichen species increased by 3.3 species per additional microhabitat. Furthermore, the richness of corticolous lichens increased by 1.2 species with 10 % higher cover of woody plants, lignicolous lichen species richness increased by 4.8 species with 1 % higher cover of deadwood, and saxicolous lichen species richness increased by 1.0 species with 1 % higher cover of stones. Our findings highlight the importance of low-intensity land use for lichen conservation. In particular, the degradation of grasslands rich in microhabitats and the destruction of lichen substrates by intensification, and conversion of unfertilized pastures formerly grazed at low intensity to meadows should be avoided to maintain lichen diversity.

Reconstruction of Holocene vegetation dynamics at Lac de Bretaye, a high-mountain lake in the Swiss Alps

A deeper understanding of past vegetation dynamics is required to better assess future vegetation responses to global warming in the Alps. Lake sediments from Lac de Bretaye, a small subalpine lake in the Northern Swiss Alps (1780 m a.s.l.), were analysed to reconstruct past vegetation dynamics for the entire Holocene, using pollen, macrofossil and charcoal analyses as main proxies. The results show that timberline reached the lake’s catchment area at around 10,300 cal. BP, supporting the hypothesis of a delayed postglacial afforestation in the Northern Alps. At the same time, thermophilous trees such as Ulmus, Tilia and Acer established in the lowlands and expanded to the altitude of the lake, forming distinctive boreo-nemoral forests with Betula, Pinus cembra and Larix decidua. From about 5000 to 3500 cal. BP, thermophilous trees declined because of increasing human land use, mainly driven by the mass expansion of Picea abies and severe anthropogenic fire activity. From the Bronze Age onwards (c. 4200–2800 cal. BP), grazing indicators and high values for charcoal concentration and influx attest an intensifying human impact, fostering the expansion of Alnus viridis and Picea abies. Hence, biodiversity in alpine meadows increased, whereas forest diversity declined, as can be seen in other regional records. We argue that the anticipated climate change and decreasing human impact in the Alps today will not only lead to an upward movement of timberline with consequent loss of area for grasslands, but also to a disruption of Picea abies forests, which may allow the re-expansion of thermophilous tree species.

Dispersal and microsite limitation in an abandoned calcareous grassland of the southern prealps

Dispersal limitation is often involved when the species composition of a dry abandoned grassland shows a slow response to resumed regular mowing. A seed-addition experiment, using 32 species which do not belong to the local species pool, was performed on Monte San Giorgio (southern Switzerland) to test whether the low recruitment success was due to dispersal limitation or due to unfavourable microsite conditions. In October 1997, 20 species were individually sown in six 3 × 4 m blocks of a 2 × 2 factorial “partial” split-plot design with treatments of abandonment vs. mowing and undisturbed vs. root-removed soil, this last being applied in small naturally-degradable pots. Moreover, 12 species were sown only in the treatments on undisturbed soil. Seedlings of sown and spontaneously germinating seeds were observed on 16 occasions over one 12-month period. Seeds of 31 out of the 32 species germinated. Twenty-four species showed germination rates higher than 5% and different seasonal germination patterns. Established vegetation, especially the tussocks ofMolinia arundinacea, reduced the quality of microsites for germination. Whereas a few species germinated better under the litter ofMolinia arundinacea, many more germinated better under the more variable microsite conditions of a mown grassland. Only a few seedlings of 25 species out of the 31 germinated species survived until October 1998. Seedling survival was negatively affected by litter, unfavourable weather conditions (frost and dry periods followed by heavy rains) and herbivory (slugs and grasshoppers). Tussocks ofMolinia arundinacea, however, tended to protect seedlings. The poor establishment success of “new” species observed in abandoned meadows on Monte San Giorgio after resumed mowing is due to dispersal and microsite limitations.

Present-day vegetation and the Holocene and recent development of Egelsee-Moor, Salzburg province, Austria

This paper describes the present-day vegetation, stratigraphy and developmental history of the mire of Egelsee-Moor (Salzburg, Austria; 45°45′N, 13°8.5′E, 700 m a.s.l., 15 ha in area) since the early Late Glacial on the basis of 4 transects with 14 trial borings across the peatland. We present a vegetation map of the mire, a longitudinal section through the peat body based on six cores showing the peat types, overview macrofossil diagrams of six cores showing the local mire development and two pollen diagrams covering the Late Glacial and Holocene. The chronology of the diagrams depends on biostratigraphic dating for the Late Glacial and early Holocene and radiocarbon dating for the remaining Holocene. The northern part of the mire originated through terrestrialisation of nutrient-rich, mostly inundated fen and the southern part through paludification of wet soils. The very small lake of today was a reservoir until recently for providing water-power for timber rafting (‘Holztrift’). The mire vegetation today is a complex of forested parts (mainly planted Pinus sylvestris and Thuja occidentalis, but also spontaneous Picea abies, Betula pubescens and Frangula alnus), reed-lands (Phragmites) and litter meadows (Molinietum, Schoenetum, etc.). The central part has hummock-hollow complexes with regionally rare species of transitional mires (Drosera anglica, D. intermedia, Lycopodiella inundata, Scorpidium scorpioides, Sphagnum platyphyllum, S. subnitens). The results indicate that some of the mid-Holocene sediments may have been removed by the timber-rafting practices, and that water extraction from the hydrological catchment since 1967 has resulted in a partial shift of transitional mire to ombrotrophic bog. The latter potentially endangers the regionally rare species and was used as an argument to stop further water extraction.

Modeling postglacial vegetation dynamics of temperate forests on the Olympic Peninsula (WA, USA) with special regard to snowpack

Past and future forest composition and distribution in temperate mountain ranges is strongly influenced by temperature and snowpack. We used LANDCLIM, a spatially explicit, dynamic vegetation model, to simulate forest dynamics for the last 16,000 years and compared the simulation results to pollen and macrofossil records at five sites on the Olympic Peninsula (Washington, USA). To address the hydrological effects of climate-driven variations in snowpack on simulated forest dynamics, we added a simple snow accumulation-and-melt module to the vegetation model and compared simulations with and without the module. LANDCLIM produced realistic present-day species composition with respect to elevation and precipitation gradients. Over the last 16,000 years, simulations driven by transient climate data from an atmosphere-ocean general circulation model (AOGCM) and by a chironomid-based temperature reconstruction captured Late-glacial to Late Holocene transitions in forest communities. Overall, the reconstruction-driven vegetation simulations matched observed vegetation changes better than the AOGCM-driven simulations. This study also indicates that forest composition is very sensitive to snowpack-mediated changes in soil moisture. Simulations without the snow module showed a strong effect of snowpack on key bioclimatic variables and species composition at higher elevations. A projected upward shift of the snow line and a decrease in snowpack might lead to drastic changes in mountain forests composition and even a shift to dry meadows due to insufficient moisture availability in shallow alpine soils.

Uppermost Limit, Extent, and Fluctuations of the Timberline and Treeline Ecocline in the Swiss Central Alps during the past 11,500 Years

Pollen and macrofossils were analyzed at two sites above today's treeline (or tree limit) in the Swiss Central Alps (Gouillé Loéré, 2503 m a.s.l., and Lengi Egga, 2557 m a.s.l.) to test two contrasting hypotheses about the natural formation of timberline (the upper limit of closed forest) in the Alps. Our results revealed that Pinus cembra--Larix decidua forests near timberline were rather closed between 9000 and 2500 B.C. (9600-4000 ¹⁴C yr BP), when timberline fluctuations occurred within a belt 100-150 m above today's tree limit. The treeline ecocline above timberline was characterized by the mixed occurrence of tree, shrub, dwarf-shrub, and herbaceous species, but it did not encompass more than 100-150 altitudinal meters. The uppermost limit reached by timberline and treeline during the Holocene was ca. 2420 and 2530 m, respectively, i.e., about 120 to 180 m higher than today. Between 3500 and 2500 B.C. (4700-4000 ¹⁴C yr BP) timberline progressively sank by about 300 m, while treeline was lowered only ca. 100 m. This change led to an enlargement of the treeline-ecocline belt (by ca. 300 m) after 2500 B.C. (4000 ¹⁴C yr BP). Above the treeline ecocline, natural meadows dominated by dwarf shrubs (e.g., Salix herbacea) and herbaceous species (e.g., Helianthemum, Taraxacum, Potentialla, Leontodon t., Cerastium alpinum t., Cirsium spinosissimum, Silene exscapa t., and Saxifraga stellaris) have been present since at least 11,000 cal yr ago. In these meadows tree and tall shrub species (>0.5 m) never played a major role. These results support the conventional hypothesis of a narrow ecocline with rather sharp upper timberline and treeline boundaries and imply that today's treeless alpine communities in the Alps are close to a natural stage. Pollen (percentages and influx), stomata, and charcoal data may be useful for determining whether or not a site was treeless. Nevertheless, a reliable and detailed record of past local vegetation near and above timberline is best achieved through the inclusion of macrofossil analysis.