Landscape history and soil properties affect grassland decline and plant species richness in rural landscapes

Past semi-natural grassland extent is thought to have a major influence on contemporary species richness in rural landscapes. The loss of grasslands over the last 300 years was reconstructed for 12 rural landscapes in Sweden, ranging from open modern agricultural landscapes to more forested landscapes. Old maps and aerial photographs from 1950s and today were used to reconstruct landscape patterns in four time-steps to investigate how present plant species richness relates to past grassland extent and decline in old and new grassland habitats. The relative importance of soil properties on the timing of grassland decline was assessed. Plant species occurrence was recorded in managed and abandoned grassland habitats in each landscape.Past and present grassland distribution was a major factor in determining plant species patterns found in grasslands today. All landscapes had an average of 80% grassland 300 years ago. Since then grassland has declined by 90% across all landscapes. Proportion of clay soils influenced the timing of grassland decline, where grasslands in landscapes dominated by clay soils were conversed to crop-fields more than 100 years ago. Grasslands on coarser soils declined later, primarily to forest. Landscapes with more than 10% semi-natural grassland left today had 50% higher species richness in all grasslands, including both abandoned and new grassland. Time since major grassland decline also seems to have an effect on the landscapes’ species richness. The results show that plant species patterns in grasslands at local scales are determined by broader landscape processes which may have occurred many centuries ago.     

Responses of butterfly and moth species to restored cattle grazing in semi-natural grasslands

The effects of restorative grazing on the abundance of butterfly and moth species were studied in mesic semi-natural grasslands of SW Finland differing in management history: (1) old continuously grazed, (2) restored (with ca 5 years of reinitiated grazing), and (3) abandoned former pastures.Generalized linear modelling of species abundances and indicator species analysis produced qualitatively similar results. Only three species (Polyommatus icarus, Lycaena hippothoe and Camptogramma bilineatum) were most abundant in old pastures, whereas 12 species (Polyommatus semiargus, Polyommatus amandus, Brenthis ino, Aphantopus hyperantus, Scopula immorata, Idaea serpentata, Scotopteryx chenopodiata, Epirrhoe alternata, Cybosia mesomella, Polypogon tentacularius, Hypena proboscidalis and Cryptocala chardinyi) were most abundant in abandoned pastures. None of the old-pasture species had become more abundant in restored pastures. Three species, Epirrhoe hastulata, Xanthorhoe montanata and Chiasmia clathrata, occurred equally abundantly in abandoned and in restored pastures indicating a slow progress of restoration.Species associated with old pastures differed from species associated with abandoned pastures in their recent distributional changes in Finland. The species of old pastures showed decreasing trends, whereas those of abandoned pastures showed mainly increasing trends in their distribution. In five out of 11 species, the preferred successional stage differed markedly between this study and previous studies conducted in Central Europe.We conclude that (1) ca 5 years of restorative grazing in mesic grasslands has been insufficient for the colonisation of old-pasture species in the restored sites, (2) different management intensities are needed regionally for the maintenance of grassland insect diversity and (3) application of the knowledge on successional preferences of different species in conservation management, even in climatically similar regions, should be made with caution.     

Plant species richness and composition in managed grasslands: The relative importance of field management and environmental factors

Decline of grassland diversity throughout Europe within the last decades is threatening biological diversity and is a major conservation problem. There is an urgent need to determine the underlying factors that control vascular plant species richness and composition in managed grasslands. In this study, 117 grasslands were sampled using standardised methods. Explanatory variables were recorded for each grassland site, reflecting the local field management, site-specific environmental conditions and large-scale spatial trends. Using variation partitioning methods, we determined the pure and shared effects of these three sets of explanatory variables on the plant species richness and composition in grasslands. Most of the explained variation in plant species richness was related to the joint effect of local field management and environmental variables. However, the applied variation partitioning approach revealed that the pure effect of spatial variables contributed relatively little to explaining variation in both the plant species richness and species composition. The largest fractions of explained variation in plant species composition were accounted for by the pure effects of environmental and local field management variables. Moreover, the results revealed that the main mechanisms by which these sets of explanatory variables affect plant species vary according to the type of management regime under study. From our findings we could conclude that particularly a reduction of nitrogen fertilisation on meadows and grazing at a low stocking rate on pastures can help to conserve biodiversity.     

Plant species richness in the Chaco Serrano Woodland from central Argentina: Ecological traits and habitat fragmentation effects

The Chaco Serrano Woodland from central Argentina has been dramatically reduced during the past 30 years, and is currently confined to several isolates of different size. In this study, we evaluated the effects of forest size, isolation and edge formation on plant species richness. Furthermore, we tested whether plants species with particular ecological traits were differentially affected by habitat fragmentation. Habitat area showed the highest explanatory value for plant species richness in stepwise multiple regressions. The effect of area was most pronounced for rare species, suggesting that large forests are necessary to preserve species with low local or regional abundance. Differences between edge and interior of Chaco Serrano were more pronounced for native and shrub species richness. The analysis of individual species cover revealed that native and biotically pollinated plants were less abundant in woodland edges. Our results showed that forest transformation into smaller remnants has lead to an impoverishment of plant communities, with particular subsets of species defined by ecological traits (rarity, origin and pollination mode) being more susceptible.     

Changes in Collembola richness and diversity along a gradient of land-use intensity: A pan European study

Changes in Collembola richness and diversity along a land-use intensity gradient were studied in eight European countries (Portugal, Spain, France, Switzerland, Hungary, UK, Ireland and Finland). In each country a set of six 1 km² land-use units (LUUs) were selected forming a gradient ranging from natural forest to agricultural dominated landscapes, passing through mixed-use ones. In addition to data on Collembola, detailed information regarding landscape diversity and structure was collected for each LUU. A total of 47,774 individuals were identified from 281 species. Collembola reacted not only to changes in the diversity of the landscape, but also to the composition of that diversity and the area occupied by each land-use type at each LUU. Although species richness patterns were not concordant among the different countries, the total number of species per LUU (landscape richness) was generally higher in natural forests and mixed-used landscapes, and lower in agricultural dominated landscapes. Moreover, high richness and diversity of Collembola at each LUU were associated with a diverse landscape structure, both in terms of number of patches and patch richness. Despite this comparable species richness between mixed-use landscapes and those dominated by natural forests, average species richness on forested areas (local richness) decreased along the gradient, showing that forest patches on mixed-use landscapes support a lower richness than in landscapes dominated by forest. This aspect is important when addressing the role of native forests in structuring biodiversity in disturbed and fragmented landscapes. Although a diverse landscape can support a high biodiversity, the results suggest that intensive fragmentation should be avoided with the risk of collapsing local species richness with the consequent result for regional biodiversity.     

Soil phosphorus and tree cover modify the effects of livestock grazing on plant species richness in Australian grassy woodland

The effects of grazing on the richness of understorey plant communities are predicted to vary along gradients of resources and tree cover. In temperate Australia livestock management has involved phosphorus addition and tree removal but little research has examined how the effects of grazing on plant species richness may vary with these management regimes. Patterns of understorey plant species richness were examined in 519, 0.09 ha quadrats in grazed pastures and remnant grassy forests and woodlands in southern Australia. Sheep grazing was the primary land use and sites varied widely in grazing frequency and density, tree cover and phosphorus fertiliser history. Using an information theoretic approach the available data provides strong evidence that the effect of grazing on total species richness varies according to available phosphorus and tree cover. Intermittent grazing and no grazing were associated with high total and native plant richness, but only at low phosphorus concentrations. Phosphorus was strongly negatively correlated with richness, particularly at low grazing frequency. Total species richness was positively correlated with tree cover except under frequent grazing at high stocking rates, suggesting that heavy grazing eliminates spatial and temporal heterogeneity imposed by trees. Native plant species richness was negatively correlated with a history of cultivation, positively correlated with tree cover and varied according to landscape position and geological substrate. Frequent high density grazing, particularly when associated with clearing, cultivation and fertiliser addition, was associated with the persistence of very few native plant species. In contrast, the richness of exotic plant species was relatively invariant and performance of the best model was low. While several studies have highlighted the importance of the grazed and cleared matrix for the conservation of native plant species, this benefit may be limited in landscapes where intensive grazing management systems dominate. Strong evidence for interactions between grazing, phosphorus and tree cover suggest that failure to consider other land use practices associated with grazing management systems could lead to erroneous conclusions regarding vegetation responses to livestock grazing.     

Population structure of relict Metasequoiaglyptostroboides and its habitat fragmentation and degradation in south-central China

The relict dawn redwood Metasequoia glyptostroboides Hu et Cheng is endemic to the region bordering Hubei and Hunan provinces and Chongqing municipality in south-central China. It is critically endangered. We investigated its population size and age structure, and provided a comparison to the study of Chu and Cooper (1950), documenting the changes of the past 60 years. Our study included all the known wild individuals of the species as well as analyses of the floristic diversity of their habitats. In the last 41 years, habitat changes have effectively ended recruitment of M. glyptostroboides and have reduced species richness in direct response to human disturbance, as shown on several indices. The remaining specimens ranged from roughly 41 to some 265 years for an average near 95 years, with heights of 12–51 m averaging 27 m. The detrimental activities of human residents include cultivation of profitable plants in the understory, selective cutting, harvesting of wood for fuel, and thoroughgoing collection of seeds for sale on the active market. Under present environmental conditions and land use, the dawn redwood will not maintain its natural range in south-central China. Our work detailing the plant populations in the habitats of this living fossil will be useful in establishing priorities for its recovery and conservation.     

Wild bee species increase tomato production and respond differently to surrounding land use in Northern California

Pollination provided by bees enhances the production of many crops. However, the contribution of wild bees remains unmeasured for many crops, and the effects of anthropogenic change on many bee species are unstudied. We experimentally investigated how pollination by wild bees affects tomato production in northern California. We found that wild bees substantially increase the production of field-grown tomato, a crop generally considered self-pollinating. Surveys of the bee community on 14 organic fields that varied in proximity to natural habitat showed that the primary bee visitors, Anthophora urbana Cresson and Bombus vosnesenskii Radoszkowski, were affected differently by land management practices. B. vosnesenskii was found primarily on farms proximate to natural habitats, but neither proximity to natural habitat nor tomato floral abundance, temperature, or year explained variation in the visitation rates of A. urbana. Natural habitat appears to increase B. vosnesenskii populations and should be preserved near farms. Additional research is needed to determine how to maintain A. urbana. Species-specific differences in dependency on natural habitats underscore the importance of considering the natural histories of individual bee species when projecting population trends of pollinators and designing management plans for pollination services. Thus, to maintain an entire bee community, multiple approaches, including maintaining natural habitat, should be implemented.     

Conservation management of calcareous grasslands. Changes in plant species composition and response of functional traits during 25 years

Five management treatments have been applied to a calcareous grassland since 1974 to study their efficiency in maintaining high diversity of grassland species. The treatments were: continuing sheep grazing, mowing, mulching, burning and leaving fallow (unmanaged). After 25 years of management, the treatments mowing and mulching resulted in a species composition similar to that from grazing by sheep. Thus, mowing and mulching can be useful management alternatives to grazing. In contrast, both burning and leaving fallow considerably changed the species composition. The changes were similar in both treatments and neither can be recommended for management purposes. We also evaluated eight traits with assumed sensitivity to the treatments (life form, life cycle, growth form, runners, lateral spread, fecundity, seed mass, germination season). In 1999, all traits except life cycle were found to respond to the different treatments. Trait response is discussed in detail.     

Large-scale changes in the abundance of common higher plant species across Britain between 1978, 1990 and 1998 as a consequence of human activity: Tests of hypothesised changes in trait representation

Presence of higher plant species was recorded in 1455 permanently marked quadrats located across Britain in 1978, 1990 and 1998 in a stratified, random sample of 259 1 km squares. Significant increases and decreases in frequency of each species were summarised as changes in the representation of simple traits, each of which had an established relationship with varying levels of fertility or disturbance. By testing the null hypothesis that the trait values represented among increasing or decreasing species were a random draw from the 1978 species pool, we determined the consistency of botanical change with changes in land-use in different vegetation types and landscape locations across Britain.Overall, 63% of significant changes in species occupancy were decreases and 37% increases. Trait changes were largely consistent with the impact of increased nutrient availability across vegetation types associated with inherently low fertility, such as infertile grassland, heath, bog and moorland. Linear habitats in lowland Britain saw trait changes consistent with secondary succession. Although trait changes were highly consistent with eutrophication in upland vegetation, the identity of the changing species left open the possibility that increased N deposition, sheep grazing intensity and local improvement could all have played a part.Recent changes in common plant species across Britain suggest that objectives for large-scale restoration and maintenance of plant communities must address four problems: (a) the increasing scarcity of stress-tolerant species in lowland species pools, (b) exploitation and maintenance of species populations on habitat fragments and linear features in the lowlands, (c) the spread and persistence of generalist species in upland Britain, (d) systemic nutrient enrichment in both lowland and upland environments.     

The influence of field shape, area and surrounding landscape on plant species richness in grazed ex-fields

Over the past 100 years species-rich semi-natural grasslands have decreased dramatically in Western Europe, where former arable fields (ex-fields) are used instead as pasture. The disappearance of semi-natural grasslands has caused a threat to the biodiversity in agricultural landscapes. Many typical grassland plants are dispersal limited, thus grazed ex-fields can be used to investigate if species spontaneously colonise these new grassland habitats. We examined the relationship between surrounding landscape, field area, shape, distance between edge and centre, and plant species diversity in ex-fields that had been grazed for 15-18 years. The results showed that there were 35% more plant species in fields surrounded by commercial forestry production compared to those surrounded by open agricultural landscape. Area and shape did not influence species richness, although there was increasing number of species in the centre with decreasing distance from the edge. Twenty-five percent of the species where typical grassland species, and ex-fields surrounded by forest had 91% more grassland species compared to those in the open landscape. It is possible to increase grassland plant occurrences by grazing ex-fields surrounded by forest or other grassland remnant habitats, particularly in landscapes where grazed semi-natural grasslands are scarce.     

Old cultural traditions, in addition to land use and topography, are shaping plant diversity of grasslands in the Alps

Socio-economically motivated land use changes are a major threat for species diversity of grasslands throughout the world. Here, we comprehensively explore how plant species diversity of grasslands in the species-rich cultural landscape of the Swiss Alps depends on recent land use changes, and, neglected in previous studies, on old cultural traditions. We studied diversity in 216 grassland parcels at three altitudinal levels in 12 villages of three cultural traditions (Romanic, Germanic, and Walser). In valleys of Romanic villages more different parcel types tended to occur than in those of Germanic and Walser villages, suggesting that socio-economic differences among cultural traditions still play a role in shaping landscape diversity. Moreover, at the village level, higher man-made landscape diversity was associated with higher plant species richness. All observed changes in land use reduced the farmers’ workload. Plant species richness was lower in fertilized than in unfertilized parcels and in abandoned compared with used parcels. Grazing slightly reduced species richness compared with mowing among unfertilized parcels, while in fertilized parcels it had a positive influence. The highest species diversity was found in mown unfertilized subalpine grasslands. Nevertheless, moderate grazing of former meadows can be a valuable alternative to abandonment. We conclude that the ongoing changes in land use reduce plant species richness within parcels and at the landscape level. To preserve plant species diversity at the landscape level a high diversity of land use types has to be maintained.     

Responses of birds of different biogeographic origins and habitat requirements to agricultural land abandonment in northern Spain

Agricultural land-use changes in Europe have taken two opposing directions: towards agricultural intensification or land abandonment. While in the Mediterranean region land abandonment is a main cause of avian diversity decline, in northern Europe species diversity often increases with successional age. We examined the hypothesis that the biogeographic origin of the avifauna determines whether abandonment brings conservation benefits or detriment by studying the bird community of agricultural land in northern Spain, at the boundary of the Mediterranean and Eurosiberian regions. Using a successional gradient, we examined landscape-scale effects of agricultural abandonment on birds during the breeding and non-breeding seasons.The trend in avian diversity with successional stage differed little between Mediterranean and Eurosiberian species in winter. In the spring, however, there was an increase in diversity with stage in abandonment for Eurosiberian birds but not for Mediterranean species. Analysis of individual species showed a preference among Eurosiberian birds for more wooded habitats whereas Mediterranean birds preferred open areas and shrubland.The introduction of agricultural policies to geo-political units that do not coincide with eco-regions cannot be assumed to bring uniform conservation benefits. In the Mediterranean region, agricultural mosaics of low intensity cultivation maintain the highest diversity of priority bird species. Agricultural land abandonment should not be assumed to benefit conservation.     

Species composition of arbuscular mycorrhizal fungi in two mountain meadows with differing management types and levels of plant biodiversity

Species composition of arbuscular mycorrhizal fungi (AMF) was analysed in two differently managed mountain grasslands in Thuringia (Germany). Arbuscular mycorrhizal fungi were studied in the roots of 18 dominant plant species from a total of 56 (32%). Additionally, spores of AMF were isolated from soil samples. Arbuscular mycorrhizal fungi species composition was analysed based on 96 sequences of the internal transcribed spacer of the nuclear ribosomal DNA, 72 originated from mycorrhizal roots, and 24 originated from AMF spores. Phylogenetic analyses revealed a total of 19 AMF species representing all genera of the Glomeromycota except Scutellospora and Pacispora. Despite a different farming intensity, resulting in remarkable differences concerning their plant species diversity (27 against 43 plant species), the diversity of AMF was found to be similar with 11 species on the intensively farmed meadow and ten species on the extensively farmed one. Nevertheless, species composition between both sites was clearly different. It thus seems likely that the AMF species composition, but not necessarily the species number, is related to above ground plant biodiversity in the system under study.     

Elevated CO2 and plant species diversity interact to slow root decomposition

Changes in plant species diversity can result in synergistic increases in decomposition rates, while elevated atmospheric CO₂ can slow the decomposition rates; yet it remains unclear how diversity and changes in atmospheric CO₂ may interact to alter root decomposition. To investigate how elevated CO₂ interacts with changes in root-litter diversity to alter decomposition rates, we conducted a 120-day laboratory incubation. Roots from three species (Trifolium repens, Lespedeza cuneata, and Festuca pratense) grown under ambient or elevated CO₂ were incubated individually or in combination in soils that were exposed to ambient or elevated CO₂ for five years. Our experiment resulted in two main findings: (1) Roots from T. repens and L. cuneata, both nitrogen (N) fixers, grown under elevated CO₂ treatments had significantly slower decomposition rates than similar roots grown under ambient CO₂ treatments; but the decomposition rate of F. pratense roots (a non-N-fixing species) was similar regardless of CO₂ treatment. (2) Roots of the three species grown under ambient CO₂ and decomposed in combination with each other had faster decomposition rates than when they were decomposed as single species. However, roots of the three species grown under elevated CO₂ had similar decomposition rates when they were incubated alone or in combination with other species. These data suggest that if elevated CO₂ reduces the root decomposition rate of even a few species in the community, it may slow root decomposition of the entire plant community.     

Changes in local species richness of farmland birds in relation to land-use changes and landscape structure

It has been suggested that an increase in the area of low-intensity land-use on arable land (e.g. set-aside fields and short-rotation coppice), and high or increased farmland habitat heterogeneity, may halt or reverse the observed population decline of farmland birds. We tested these hypotheses by undertaking farmland bird censuses during two contrasting periods of agricultural policies and land-use (i.e. 1994 and 2004) in a farmland region covering a gradient of forest- to farmland-dominated landscapes in Sweden. Local species richness (i.e. at 3 hectare sites) declined significantly between 1994 and 2004. Local species richness was positively related to habitat heterogeneity in both years of study whereas temporal change in species richness was not. Local change in species richness was positively associated with a change in the proportion of non-rotational set aside and short-rotation coppice (i.e. low-intensity land-use forms), but also to changes in the amount of spring-sown crops. However, the effect of low-intensity land-use was significantly dependent on the amount of forest in the surrounding landscape. An increase in low-intensity land-use was linked to an increase (or less marked decrease) in species richness at sites located in open farmland surroundings but to a decrease in richness at sites located in forest surroundings. This interaction between amount of forest and low-intensity land-use could be interpreted as a “rare habitat effect”, where an increase in a farmland habitat only positively affects biodiversity when it was originally uncommon (i.e. open farmland areas). Our results suggest that conservation measures of farmland biodiversity have to be put in a landscape context.     

Concentration of trace and major elements in natural grasslands of Bosnia and Herzegovina in relation to soil properties and plant species

Deficient trace elements concentration in soils, forages, and animals have been reported in several areas of Balkan region. Main challenge in overcoming low productivity of forage and animal production in this region is the lack of data on the nutritional status of the pastures and soils. This study examined the nutrient and pseudo total concentration of trace elements in soil and herbage plants, and the potential deficiency or excess with regards to crop and livestock production. Soil and plant samples from 100 sampling points were collected in the mountainous grasslands of Manja?a (between longitudes 16°56′ and 17°8′ east; and latitudes 44°33′ and 44°42′ north) and Vla?i? (between longitudes 17°14′ and 17°29′ east; and latitudes 44°25′ and 44°37′ north). Soil samples were analysed for soil texture, pseudo total concentration (5?ml HNO₃) of trace elements (TE), pH, soil organic carbon (SOC), while plant samples were analysed for TE. The soil pH varied from strongly acidic to moderately alkaline. The concentration of SOC varied from 0.5% to 12.3%. Most of the soil samples were finer-textured soils high in silt content. The average concentrations of Na, P, Zn, Se, Cu, Co, and B were low in both soil and herbage plants. Plant K, Ca, Mg, Mo, and Mn concentrations were sufficiently high to meet the requirements of grazing animals, while Fe concentrations was elevated in certain areas. High levels of Mo were found in both soil and plants. The results suggest that imbalances observed in natural pastures of Manja?a and Vla?i? area, caused by low soil TE concentration and other soil and plant properties, could contribute to poor animal nutrition.