How to successfully improve the biodiversity of city grasslands?

Urban grasslands (UG) are among the most common types of urban green areas. They are usually species poor, and spontaneous plant migration, which can increase biodiversity, is limited. To improve the range of ecosystem services provided by UG, various seed mixtures are applied during the establishment and restoration of UG. These mixtures vary in content, quality, and cost. High-quality seed mixtures are expensive and are usually only available in small amounts. Meanwhile, alternative methods of seed introduction (e.g., fresh hay application, seeds harvested by brush) have not been well studied in UG restoration, and inexpensive commercial mixtures could have low quality and lead to poor restoration outcomes. Here, we tested the effectiveness of different seed sources to create high-quality UG at two study sites. Based on the results, all seed addition methods increased the species richness of restored grasslands. The outcome of seed addition was satisfactory regardless of differences in residual vegetation species composition and soil properties between the sites. The species richness on plots that received a commercial mixture of flower meadow plants dedicated to pollinators decreased after overwintering. The alternative seed sources (fresh hay and seed incidentally collected during mowing) yielded grassland quality that was comparable to that on plots that received high-quality mixtures with known seed origin (a seminatural meadow mixture and a mixture with the addition of grasses).

From invasive species stand to species-rich grassland: Long-term changes in plant species composition during Solidago invaded site restoration.

Biological invasions degrade ecosystems, negatively affecting human well-being and biodiversity. Restoration of invaded agricultural ecosystems is among specific goals of European Union Biodiversity Strategy. Successful restoration of invaded lands is a long-term process that requires monitoring to assess the effects of interventions. Here, we present the results of a long-term experiment (8 years) on restoration of semi-natural grassland on abandoned arable field overgrown by invasive Solidago species (S. gigantea and S. canadensis). We examined effect of different invaders removal methods (rototilling, turf stripping, herbicide application) and seed application practices (commercial seed mixture, fresh hay) on changes in species composition and taxonomic diversity of restored vegetation. Our results showed a positive effect of grassland restoration on taxonomic diversity and species composition, manifested by a decrease in Solidago cover and an increase in cover and richness of target graminoids and forbs characteristic of grassland. The seed source had a longer lasting and still observable effect on the vegetation composition than the Solidago removal treatments, which ceased to differ significantly in their influence after the first few years. Applying fresh hay as a seed source increased the cover of grassland species such as Arrhenatherum elatius and Poa pratensis. For commercial seed mixture, we observed the high cover of Lolium perenne and Schedonorus pratensis (introduced with seed mixture) at the beginning and the slow decrease along the experiment course. The most striking effect was the fresh hay with herbicide application, which resulted in the lowest Solidago cover and the highest cover of target graminoids. Nonetheless, with years the non-chemical methods, including no treatment, gives comparable to herbicide effectiveness of restoration. Overall, during the experiment, alpha diversity increased, while beta and gamma diversity reached a species maximum in the third year, and then decreased. In conclusion, this study gives guidance to successful restoration of species-rich grasslands on sites invaded by Solidago. It should be emphasised that short-term effect differ considerably from long-term outputs, especially highlighting the importance of seed source, as well as effectiveness of environmentally friendly methods such as regular mowing to control the invader.

Genetic structure of Trifolium pratense populations in a cityscape.

Urban grasslands provide numerous ecosystem services, and their maintenance should be based on naturally regenerating plant populations. However, the urban environment is challenging for preserving viable populations, mostly because of their high fragmentation and small size, which can lead to genetic drift. We examined red clover (Trifolium pratense) in a medium-size city in Central Europe to test the cityscape effect on within- and among-population genetic diversity. We used eight inter-simple sequence repeat markers to examine the genetic structure of 16 populations, each represented by eight individuals. The isolation by resistance was analysed using a least cost patch approach, focusing on gene flow via pollinators. We found great variation among T. pratense populations, with no discernible geographic pattern in genetic diversity. We linked the diversity to the long history of the city and high stochasticity of land use changes that occurred with city development. In particular, we did not find that the Odra River (ca. 100 m wide) was a strong barrier to gene transfer. However, notable isolation was present due to resistance and distance, indicating that the populations are threatened by genetic drift. Therefore, gene movement between populations should be increased by appropriate management of urban green areas. We also found that small urban grassland (UG) patches with small populations can still hold rare alleles which significantly contribute to the overall genetic variation of T. pratense in the city.

Effects of extracts from various parts of invasive Solidago species on the germination and growth of native grassland plant species.

Allelopathy is an important factor influencing whether an invasive plant species can become successfully established in a new range through disrupting the germination and growth of native plant species. Goldenrods (Solidago species) are one of the most widespread invasive taxa in Central Europe of North American origin. Owing to their high environmental impact and wide distribution range, invasive Solidago species should be controlled in Europe, and the areas invaded by them should be restored. Numerous studies have reported the allelopathic effects of Solidago gigantea and Solidago canadensis, but the results are inconsistent regarding differences in the allelopathic effects of particular plant parts and in the sensitivity to Solidago allelopathic effects among native species as well as between the two invasive species themselves. In this study, we aimed to analyse the effect of water extracts from S. canadensis and S. gigantea parts (roots, rhizomes, stems, leaves, and inflorescences) on the germination and initial growth of seedlings of 13 grassland species that typically grow in Central Europe. The tested grassland species differed in susceptibility to Solidago allelopathy, with the most resistant species being Schedonorus pratensis, Lolium perenne, Trifolium pratense, Daucus carota and Leucanthemum vulgare. The inhibitory effect of 10% water extracts from leaves and flowers were stronger than those from rhizomes, roots, and stems without leaves, regardless of the Solidago species. Our study results imply that reducing the allelopathic effect of Solidago during habitat restoration requires removal of the aboveground parts, including fallen leaves. The allelopathic effects of roots and rhizomes seem to be of secondary importance.

Spatial patterns of vascular plant species richness in Poland - a data set.

Recognition of species richness spatial patterns is important for nature conservation and theoretical studies. Inventorying species richness, especially at a larger spatial extent is challenging, thus different data sources are joined and harmonized to obtain a comprehensive data set. Here we present a new data set showing vascular plant species richness in Poland based on a grid of 10 × 10 km squares. The data set was created using data from two sources: the Atlas of Distribution of Vascular Plants in Poland and the Polish Vegetation Database. Using this data set, we analysed 2,160 species with taxonomical nomenclature according to the Euro + Med PlantBase checklist in 3,283 squares covering the entire territory of Poland (ca. 312,000 km2). The species were divided into groups according to their status and frequency of distribution, and the statistics for each square were obtained. For purposes of analysis, sampling bias was assessed. The data set promotes theoretical analysis on species richness and reinforces the planning of nature conservations.

The effect of grassland type and proximity to the city center on urban soil and vegetation coverage.

Urban soils with associated vegetation are important components of urban ecosystems, providing multiple regulating and supporting ecosystem services. This study aimed to analyze the differences in the soil chemistry and vegetation of urban grasslands considering urbanization gradient and urban grassland type (UGT). We hypothesized that the chemical properties of soil, such as metal content, as well as vegetation traits, differ according to grassland type (lawns, grasslands in parks, grasslands on river embankments, and roadsides) and the location of grassland patches (city center versus peripheries). Our samples included 94 UGT patches which each patch represented by four square sampling plots sized 1 m2. The results showed high differentiation of measured traits unrelated to UGT and location. The exception was K content, with a relatively high concentration in lawns, and some metals (Cd, Cu, Pb, Zn), with higher concentrations in the city center than in the peripheries. We found two grassland patches located in the city center where the concentrations of Pb, Zn, and Cu exceeded the level authorized by Polish standards. In the case of vegetation traits, the variability was not structured considering the UGT and location of the patches, except for bare soil cover, which was higher in lawns in the city center compared to embankments in the peripheries. We observed correlations between vegetation traits and soil chemical properties. The vascular plant species richness decreased when N, P, and C content, along with an increase in grass cover and a decrease in herbs.

Drivers of systematic bias in alien plant species distribution data.

Among the main challenges in modelling biological invasion is a lack of valid data on the absence of invasive species. Absence data are important for assessing the reliability of models, but multiple surveys at a location are needed. In practice, omission errors are more frequent than commission errors. We therefore quantified how eliminating potentially biased areas from invasive species distribution models (iSDMs) affected the models' performance, and we assessed how the distribution of biased areas correlated with environmental factors. We hypothesized that for neophytes, the distribution of biased areas corresponds to specific land relief and/or particular landscape and land use, but not the density of roads and urbanized areas. The data on neophytes were obtained from a distribution atlas covering approximately 31,000 km2 in Central Europe overlaid with a 2 × 2 km square grid. One hundred fifty-three species were used for modelling neophyte richness, and negative residuals from the model were assumed to indicate biased squares. Twenty invasive species were used as an independent dataset for testing the effect of excluding the biased squares on iSDM performance. The exclusion of biased squares increased the iSDM performance from an area under the curve value of 0.73 to 0.78. The best results were obtained by excluding 30 % of the squares from the original dataset. The presence of damp sites explained the distribution of biased squares; the density of roads and urbanized areas had no impact. The applied method allows distinguishing biased, plausibly undersampled squares in a species distribution atlas, the exclusion of which significantly improves iSDM performance. The results suggest that the commonly observed low sampling effort in areas distant from communication routes and urbanized areas was not crucial in modelling invasive species distribution, which can be related to smaller neophyte richness in remote areas resulting from low propagule pressure.

The impact of restoration methods for Solidago-invaded land on soil invertebrates.

The belowground community structure of soil biota depends on plant composition and may be affected by invasive plant species. We hypothesized that the type of land restoration method applied affects the abundance and composition of soil invertebrates. Our field experiment centred on Solidago species control using different seed mixtures and methods of seed introduction (sowing mixtures: grasses, grasses with legumes, seeds from a seminatural meadow, and application of fresh hay) and different frequencies of mowing (one, two, or three times per year). Soil invertebrates were identified to the taxa, using light microscopes. Richness and diversity indices were calculated, and a redundancy analysis was conducted. Generally, mowing intensity negatively influenced soil organisms, although increased mowing frequency positively affected the abundance of some taxa (Symphyla, Hemiptera). Mowing twice per year decreased the abundance of soil invertebrates, but not their diversity. Soil invertebrate taxa had the greatest abundance in the plots sown with a seed mixture containing grasses with legumes. Among the restoration methods studied, mowing once a year and introducing grasses with legumes represented the least harmful strategy with regard to soil invertebrate abundance. Further studies are needed to investigate the dynamics of soil mesofauna exposed to long-term mowing and changes in vegetation characteristics.

Towards a better understanding of linear species distribution in river valleys: The abundance of river corridor plants is linked to soil exchangeable potassium and pH.

The phenomenon that some vascular plant species grow mainly or exclusively in the large river valleys of Central Europe constitutes a long-standing distribution puzzle. In our study, we focused on determining which physicochemical properties of soil affect the distribution of river corridor plant (RCP) species. Research that included RCP coverage, the physicochemical properties of soil as well as several topographic and spatial variables were conducted in 10 transects perpendicular to the San River (SE Poland). The sampled plots were located both in close proximity to the river, where the greatest number of RCP populations are concentrated, and along anthropogenic linear landscape elements which have proven to be important for RCP occurrence in areas located away from the riverbed. Spearman rank correlation matrices were constructed to observe the general relationships between particular variables and Boosted Regression Tree models were used for explaining RCP coverage. Our study indicated that in the case of plots located in proximity to the river, the expected coverage of RCP species was highest in plots characterized by a higher soil exchangeable potassium (K) content, as well as in those located closer to the San River and at lower altitudes. In plots situated along anthropogenic linear landscape elements, the expected coverage of RCPs was highest in plots with a high soil exchangeable K content and in those with a high soil pH. The analyses clearly showed that the physicochemical properties of soils indeed affect RCP species occurrence and they require soils with a high exchangeable K content and a high pH. Since these two soil parameters are closely related to soil fertility, and additionally K enhances root development, we suppose that both factors strengthen the competitiveness of RCPs in floodplain ecosystems.

Seeds harvested during mowing from semi-natural grasslands as an ad hoc but effective solution for grassland restoration.

Species-rich grasslands formed by local ecotypes of native species provide numerous ecosystem services both in rural areas as well as urban grasslands. Nonetheless, their area is still too small, making grasslands one of the most frequently restored habitats. Successful restoration requires high-quality seed material, which is expensive and often not easy to acquire. In this study, we tested the potential of seeds accidentally collected during the mowing of a semi-natural grassland for grassland restoration. We tested seed purity, species composition, and germination capability in both laboratory and field conditions. Ninety percent of the collected material consisted of pure seeds of numerous species. Their germination capability was relatively low but still sufficient for successful grassland restoration under a typical seed density/mass per unit area seeding ratio. The germination capacity was the highest in the first two weeks after sowing and increased with overwintering seed storage. The results suggested that the seeds could be successfully used for species-rich grassland restoration. In terms of advantages, the seed mixture had a low cost and contained native species seeds representing local ecotypes. In terms of disadvantages, there was a relatively low amount of seed material and an inability to plan the time of seed harvesting. Thus, the use of the accidentally collected seeds can be considered an effective but rather ad hoc solution.

Macroecological drivers of vascular plant species composition in semi-natural grasslands: A regional study from Lower Silesia (Poland).

In agricultural landscapes of Europe, species-rich semi-natural grasslands provide numerous ecosystem services, such as forage production, ground and surface water regulation, and carbon sequestration. In the face of the declining area of grasslands, information about the macroecological drivers of grasslands diversity is an important element of studies on their ecology and management. Here, we tested the effect of factors representing environmental gradients, landscape structure, human pressure, habitat continuity in time, and spatial structure on species composition of vascular plants in semi-natural grasslands. The analysis was performed using a multivariate approach for 689 vegetation plots distributed across ~20,000 km2 in the Lower Silesia region (Poland, Central Europe). We found that species composition was significantly influenced by factors representing all studied groups. The most influential factors were temperature, long-term habitat continuity, and topography. Moreover, we captured the correlation between habitat conditions and habitat continuity. Old grasslands (existing since the 1940s) usually occurred in wet habitats, and medium grasslands (existing since the 1980s) existed in highlands in most cases. We highlighted the potential of freely available environmental databases, as well as historical topographic maps, in the exploration of patterns of species composition at a large spatial extent. The result show also the usefulness of vegetation databases in recognition of grassland complexes contributing the regional biodiversity.

Drivers of Solidago species invasion in Central Europe-Case study in the landscape of the Carpathian Mountains and their foreground.

AIM: The invasion process is a complex, context-dependent phenomenon; nevertheless, it can be described using the PAB framework. This framework encompasses the joint effect of propagule pressure (P), abiotic characteristics of the environment (A), and biotic characteristics of both the invader and recipient vegetation (B). We analyzed the effectiveness of proxies of PAB factors to explain the spatial pattern of Solidago canadensis and S. gigantea invasion using invasive species distribution models. LOCATION: Carpathian Mountains and their foreground, Central Europe. METHODS: The data on species presence or absence were from an atlas of neophyte distribution based on a 2 × 2 km grid, covering approximately 31,200 km2 (7,752 grid cells). Proxies of PAB factors, along with data on historical distribution of invaders, were used as explanatory variables in Boosted Regression Trees models to explain the distribution of invasive Solidago. The areas with potentially lower sampling effort were excluded from analysis based on a target species approach. RESULTS: Proxies of the PAB factors helped to explain the distribution of both S. canadensis and S. gigantea. Distributions of both species were limited climatically because a mountain climate is not conducive to their growth; however, the S. canadensis distribution pattern was correlated with proxies of human pressure, whereas S. gigantea distribution was connected with environmental characteristics. The varied responses of species with regard to distance from their historical distribution sites indicated differences in their invasion drivers. MAIN CONCLUSIONS: Proxies of PAB are helpful in the choice of explanatory variables as well as the ecological interpretation of species distribution models. The results underline that human activity can cause variation in the invasion of ecologically similar species.

Connectivity assessment and prioritization of urban grasslands as a helpful tool for effective management of urban ecosystem services.

Urban grasslands are usually managed as short-cut lawns and have limited biodiversity. Urban grasslands with low-intensity management are species rich and can perform numerous ecosystem services, but they are not accepted by citizens everywhere. Further, increasing and/or maintaining a relatively high level of plant species richness in an urban environment is limited by restricted plant dispersal. In this study, we examined the connectivity of urban grasslands and prioritized the grassland patches with regard to their role in connectivity in an urban landscape. We used high-resolution data from a land use system to map grassland patches in Wroclaw city, Silesia, southwest Poland, Central Europe, and applied a graph theory approach to assess their connectivity and prioritization. We next constructed a model for several dispersal distance thresholds (2, 20, 44, 100, and 1000 m), reflecting plants with differing dispersal potential. Our results revealed low connectivity of urban grassland patches, especially for plants with low dispersal ability (2-20 m). The priority of patches was correlated with their area for all dispersal distance thresholds. Most of the large patches important to overall connectivity were located in urban peripheries, while in the city center, connectivity was more restricted and grassland area per capita was the lowest. The presence of a river created a corridor, allowing plants to migrate along watercourse, but it also created a barrier dividing the system. The results suggest that increasing the plant species richness in urban grasslands in the city center requires seed addition.

Species diversity patterns in managed Scots pine stands in ancient forest sites.

Continuity in forest habitats is crucial for species diversity and richness. Ancient Scots pine forests are usually under forest management, which disturbs vegetation and causes differentiation in terms of tree stand age. To date, vegetation variability in ancient Scots pine forests has not been examined based on tree stand age classes. In the present study the continuity of a large Scots pine forest complex was investigated, and a system of sampling plots established in five tree stand age classes: initiation stands (4-10 years), young stands (20-35 years), middle-aged stands (45-60 years), pre-mature stands (70-85 years) and mature stands (95-110 years). Species composition, including vascular plants, bryophytes and lichens, on soil, tree trunks, and coarse woody debris, was analyzed. Based on existing classifications systems, forest species and ancient forest species groups were distinguished. In the studied ancient Scots pine forests the species pool and richness were relatively low, and the vegetation consisted mostly of generalist species. Cryptogams, which can grow on diverse substrates, were the most abundant species. Moreover, most species could tolerate both forest and non-forest conditions. Age class forests provided different environmental niches for species. Initiation stands were optimal for terrestrial light-demanding species, and in terms of species composition, initiation stands were most specific. Young stands were most preferred by species on coarse woody debris, and at this stage of stand maturation epiphytic species re-appeared. The oldest stands were not rich in forest specialists, i.e. species of closed forest and ancient forest species. Cryptogams of closed forests inhabited different substrates, and they were not associated only with the oldest stands. The low number of forest specialists in the oldest stands may be a general feature of acidophilus pine forests. However, it may also be a result of the lack of species sources in the vicinity of maturing pine stands. In managed forests a frequent diversity pattern is an increase in a species pool and richness after clear-cut logging. In the present study we obtained higher species pools in initiation and young stands, but richness was similar in all tree stand age classes. This resulted from taking into account species of different substrates (terrestrial, epixylous and epiphytic species) which changed their participation in the vegetation of subsequent stages of tree stand development.

Effect of anthropogenic factors, landscape structure, land relief, soil and climate on risk of alien plant invasion at regional scale.

We compared the effectiveness of explanatory variables representing different environmental spheres on the risk of alien plant invasion. Using boosted regression trees (BRT), we assessed the effect of anthropogenic factors, soil variables, land relief, climate and landscape structure on neophyte richness (NR) (alien plant species introduced after the 15th century). Data on NR were derived from a 2 × 2 km grid covering a total area of 31,200 km2 of the Carpathian massif and its foreground, Central Europe. Each of the examined environmental spheres explained NR, but their explanatory ability varied more than two-folds. Climatic variables explained the highest fraction of deviation, followed by anthropogenic factors, soil type, land relief and landscape structure. The global model, which incorporated crucial variables from all studied environmental spheres, had the best explanatory ability. However, the explained deviation was far smaller than the sum of the deviations explained by the single-sphere models. The global model showed that the deviation that could be explained by variables representing particular spheres, overlapped. The variables representing landscape structure were not included in the global model as they were found to be redundant. Finally, the climatic variables explained a smaller fraction of the deviation than the anthropogenic factors. The partial dependency plots allowed the assessment of the course of dependencies between NR and particular explanatory variables after eliminating the average effect of all other variables. The relationships were usually curvilinear and revealed some values of environmental variables beyond which NR changed considerably.

Effect of petrol fuel contamination on the growth of mature Scots pine, Pinus sylvestris L., trees.

The radial increment and crown status of mature Scots pine trees growing in polluted and unpolluted sites were compared. In 1996, as a result of some malfunction, unleaded petrol penetrated into the soil next to a plantation. Detailed geological and hydrological studies revealed the route of the spread of contamination and extent of the pollution. The trees growing in polluted sites revealed strong depletion of radial growth starting immediately after pollution. Such depletion lasted 2-3 years before the ring widths stabilised at a low level. After a few years the radial increment increased, and now do not differ from the increment of trees in the unpolluted sites.