No detrimental effects of delayed mowing or uncut grass refuges on plant and bryophyte community structure and phytomass production in low-intensity hay meadows

To maintain European semi-natural grasslands, agri-environment schemes (AES) have been established in many countries but their biodiversity benefits have remained limited. We tested the effects of three new mowing regimes designed to benefit biodiversity in extensively managed meadows across the Swiss lowlands. Our experimental treatments mimicked easily implementable farming practices. We previously showed that invertebrates benefit from delayed mowing and leaving an uncut grass refuge. Here we focus on the effects on plant and bryophyte communities.We compared the standard AES practice (earliest mowing on June 15, no fertilizer input, but no restriction on number of cuts) to three alternative mowing regimes: (i) earliest mowing delayed by one month, (ii) maximum of two cuts per year with at least eight weeks in between, and (iii) leaving an uncut refuge on 10–20% of the meadow area in 12 study areas in the Swiss lowlands. We also tested for the interactive effects of ambient temperature, precipitation, elevation, meadow size, local forest cover, time since AES registration, and phytomass production.After five years of application, we found no difference in the effects of mowing regimes on vascular plant or bryophyte species richness, community composition, phytomass, flowering phenology or average plant height (the latter two indices were derived from the literature). However, cutting frequency and hay nutritional quality (C:N and Ca:P ratios) were lower under delayed mowing. Vascular plant and bryophyte species richness as well as forage quality were negatively related to phytomass, while the latter was positively related to mean summer temperature and negatively to time since AES registration.We conclude that supporting invertebrate biodiversity with alternative mowing regimes has no detrimental effects on the vascular plants and mosses, while the reduced forage quality calls for additional financial compensation of the farmers adopting these agri-environment schemes.     

Restorative mowing on an abandoned semi‐natural meadow: short‐term and predicted long‐term effects

Abstract. When management, in the form of cattle grazing and mowing, ceases the abundance of competitively superior plant species tends to increase in abandoned semi‐natural meadows. Litter accumulation elevates the soil nutrient levels and hinders seedling recruitment. We surveyed changes in plant cover and species composition of a formerly grazed meadow in permanent plots for six years. Some plots were unmown, while others were mown and raked annually in August. The cover of grasses decreased and herb cover remained unchanged regardless of the treatment. Mowing and raking significantly reduced litter accumulation and increased the number of ground layer species. The expected long‐term effects of abandonment and restorative mowing were studied by calculating the transition probabilities for unmown and mown plots and simulating the course of succession as projected by the transition matrices. During a simulation period of 30 yr, abandonment led to (1) a decrease in the cover of small herbs, (2) a slight increase in the cover of tall herbs and (3) a slight decrease in the cover of grasses. In contrast, the cover of small herbs on the mown plots remained unchanged or slightly increased during the course of simulation. These results suggest that mowing late in the season is primarily a management tool for the maintenance of the existing species diversity and composition. However, it may not be an effective restorative tool to induce overall changes in the resident vegetation of abandoned grass‐dominated meadows. Grazing or mowing early in the season may be more effective in this respect. Consequently, mowing early or, alternatively, late in the season may provide management strategies for the maintenance and restoration of species diversity, respectively.     

Evaluating alternative mowing regimes for conservation management of Central European mesic hay meadows: A field experiment

Conservation status of hay meadows highly depends on their management. The main goal of this study was to assess the efficiency of different mowing regimes in maintenance of plant species richness and diversity of mesic hay meadows. The field experiment was carried out on a species rich, mesic hay meadow in Western Hungary. We evaluated the effects of four alternative types of management on the plant community after 7 years of continuous treatment: (1) mowing twice a year, typical traditional management, (2) mowing once a year in May, most practised currently by local farmers, (3) mowing once a year in September, often proposed for conservation management and (4) abandonment of mowing. Traditional mowing resulted in significantly higher number and higher diversity of vascular plant species than other mowing regimes. Mowing twice a year was the only efficient way to control the spread of the invasive Solidago gigantea, and mowing in September was more successful in it than mowing in May. We conclude that the traditional mowing regime is the most suitable to maintain botanical diversity of mesic hay meadows; however, other regimes should also be considered if certain priority species are targeted by conservation.     

A mowing experiment to evaluate the influence of management on the activity of host ants of <Emphasis Type="Italic">Maculinea</Emphasis> butterflies

Maculinea butterflies obligatory parasitize certain species of Myrmica ants. Thus, the presence of the host ant species is a limiting factor for the survival of a Maculinea population. Here, we analyse the influence of vegetation structure and ground temperature on ant diversity and abundance on Maculinea habitats, with the final aim of identifying the environmental variables determining patterns of variation in species composition in order to recommend a mowing regime that will promote our three target species: Maculinea teleius, M. nausithous and M. alcon. Experimental plots with different mowing regimes were established at eight sites in South-Eastern Germany, a region which still contains a number of relatively large, stable populations of these threatened butterfly species. Among the seven different ant species recorded, four belong to the genus Myrmica (M. scabrinodis, M. rubra, M. ruginodis and M. vandeli). Among these, M. scabrinodis results most abundant at all sites. In a CCA analysis of environmental variables recorded at the studied plots, ant species diversity appears largely determined by litter cover, mean temperature, and mean grass cover. Mowing once a year, in the second half of September, after the larvae have left their host plants, enhances the abundance of Myrmica ants in the meadows, and would be the best management compromise for all three species.     

Habitat selection of the globally threatened Aquatic Warbler Acrocephalus paludicola at the western margin of its breeding range and implications for management

The globally threatened Aquatic Warbler Acrocephalus paludicola is an umbrella species for fen mires and is at risk of extinction in its westernmost breeding population due to severe habitat loss. We used boosted regression trees to model Aquatic Warbler habitat selection in order to make recommendations for effective management of the last remnant habitats. Habitat data were collected in the years 2004–2006 in all remaining breeding sites in Pomerania (eastern Germany and western Poland) as well as in recently abandoned sites. Models were validated using data from similar Aquatic Warbler habitats in Lithuania. The probability of occurrence of Aquatic Warblers in late May/early June was positively associated with low isolation from other occupied sites, less eutrophic conditions, a high proportion of area mown early in the preceding year, high availability of vegetation 60–70 cm high, high prey abundance and high habitat heterogeneity. Early summer land management is needed in the more productive sites to prevent habitat deterioration by succession to higher and denser vegetation. As this also poses a serious threat to broods, management that creates a mosaic of early and late used patches is recommended to preserve and restore productive Aquatic Warbler sites. In less productive sites, winter mowing can maintain suitable habitat conditions. Aquatic Warbler‐friendly land use supports a variety of other threatened plant and animal species typical of fens and sedge meadows and can meet the economic interests of local land users.     

Direct and indirect effects of land‐use intensification on ant communities in temperate grasslands

Land‐use intensification is a major driver of local species extinction and homogenization. Temperate grasslands, managed at low intensities over centuries harbored a high species diversity, which is increasingly threatened by the management intensification over the last decades. This includes key taxa like ants. However, the underlying mechanisms leading to a decrease in ant abundance and species richness as well as changes in functional community composition are not well understood. We sampled ants on 110 grassland plots in three regions in Germany. The sampled grasslands are used as meadows or pastures, being mown, grazed or fertilized at different intensities. We analyzed the effect of the different aspects of land use on ant species richness, functional trait spaces, and community composition by using a multimodel inference approach and structural equation models. Overall, we found 31 ant species belonging to 8 genera, mostly open habitat specialists. Ant species richness, functional trait space of communities, and abundance of nests decreased with increasing land‐use intensity. The land‐use practice most harmful to ants was mowing, followed by heavy grazing by cattle. Fertilization did not strongly affect ant species richness. Grazing by sheep increased the ant species richness. The effect of mowing differed between species and was strongly negative for Formica species while Myrmica and common Lasius species were less affected. Rare species occurred mainly in plots managed at low intensity. Our results show that mowing less often or later in the season would retain a higher ant species richness—similarly to most other grassland taxa. The transformation from (sheep) pastures to intensively managed meadows and especially mowing directly affects ants via the destruction of nests and indirectly via loss of grassland heterogeneity (reduced plant species richness) and increased soil moisture by shading of fast‐growing plant species.     

How too much care kills species: Grassland reserves,agri-environmental schemes and extinction of <Emphasis Type="Italic">Colias myrmidone</Emphasis> (Lepidoptera: Pieridae) from its former stronghold

We describe a rapid extirpation of a strong population of globally threatened butterfly, Colias myrmidone, from the White Carpathians, Czech Republic. The area is renowned for a high plant and insect diversity associated with savannah-like Carpathian meadows, traditionally managed by mowing once a year and subsequent autumn grazing. Although many meadows fell victim to land consolidation schemes in the 1970s and 1980s, over 2000 hectares have been preserved in a dense network of reserves. The reserve management practised since the mid-1990s has consisted of uniform machine mowing of entire areas and this was further accelerated after accession of the Czech Republic to the EU, when agri-environmental schemes (AES) required two cuts per year. This starkly conflicted with the life history of C. myrmidone, because both spring and autumn cuts interfere with larval development. Despite intensive surveys, only five and then two C. myrmidone individuals were seen in the area in 2005 and 2006, respectively, rendering the species effectively extinct. Conservation authorities failed by neglecting early warning signals issued by amateur entomologists, and by ignoring the basic tenets of conservation of insect diversity, such as the need to maintain heterogeneity of resources. Unless AES provide for this crucial aspect of habitat suitability for invertebrates, they may turn from a great hope into a massive failure.     

Effects of vegetation management by mowing on ground-dwelling arthropods

Species-rich grasslands are rare in the Netherlands and need consistent vegetation management to retain their characteristic biodiversity. Roadside verges are important refuges for grassland plants since the mowing management no longer aims at traffic safety only but also strives for botanical diversity. Although arthropods are highly abundant in roadside verges, the effect of different mowing practices on this group is largely unknown. During 4 years, we studied ground beetles, weevils, ants and ground-dwelling spiders with pitfall traps in experimental plots in roadside verges with five different mowing treatments: (i) no management, (ii) and (iii) mowing once a year with and without hay removal, (iv) and (v) mowing twice a year with and without hay removal. This was done in a plant productivity gradient; the experiment was repeated in low-, medium- and high-productive verges. In the low-productive site, the effect of management on the arthropods only existed in a higher abundance in plots mown twice per year with hay removal. In the medium- and high-productive sites, mowing twice a year with hay removal resulted not only in highest abundances but also in highest arthropod species richness. Mowing twice without hay removal and mowing once with removal showed intermediate values, while mowing once per year without removal and particularly the absence of management resulted in low diversity and low abundance. To promote ground-dwelling arthropods in medium-to-high-productive grassland verges, we recommend a management of mowing twice a year with the removal of hay. It is reasoned that some form of rotational management, aiming at leaving some vegetation refuges intact after mowing events, may further promote arthropod survival. However, caution should be taken that these refuges are not too large, as overall suitability for ground-dwelling arthropod decreases rapidly in such patches. Out of several studied vegetation characteristics, the number of flowering plant species (medium-productive verge) and total flower abundance (high-productive verge) appeared to represent suitable, and easily monitored, proxies that significantly mirror arthropod diversity.     

Rotational fallows as overwintering habitat for grassland arthropods: the case of spiders in fen meadows

Regular mowing of grassland is often necessary for plant conservation, but uncut vegetation is needed by many arthropods for overwintering. This may lead to conflicting management strategies for plant and arthropod conservation. Rotational fallows are a possible solution. They provide a spatio-temporal mosaic of mown and unmown areas that may combine benefits to both plants and arthropods. We tested if rotational fallows enhance spider overwintering in fen meadows. Rotational fallows consisted of three adjoining strips 10 m wide and 35–50 m long. Each year, one of these strips was left unmown (fallow) in an alternating manner so that each strip was mown two out of three years. Spiders were sampled during spring with emergence traps in nine pairs of currently unmown fallow strips and completely mown reference plots. Fallows significantly enhanced orb-weavers (Araneidae), sac spiders (Clubionidae) and ground spiders (Gnaphosidae). However, only 4.7% of the total variation in community composition was attributable to fallows. Community variation was larger between landscapes (34.5%) and sites (38.2%). Also β diversity was much higher between landscapes (45 species) and sites (22 species) than between fallows and mown reference plots (10 species). We conclude that the first priority for spider conservation is to preserve as many fen meadows in different landscapes as possible. Locally, rotational fallows enhance overwintering of the above-mentioned spider families, which are sensitive to mowing in other grassland types as well. Thus, rotational fallows would probably foster spider conservation in a wide range of situations. However, stronger effects can be expected from larger and/or older fallow areas.     

Influence of abandonment on syrphid assemblages in mountainous meadows

Grasslands are often managed with different intensities in the European Alps. Studies have shown that a medium management can benefit plant and animal biodiversity in these ecosystems. However, in recent decades abandonment of extensively managed meadows is an ongoing global challenge. Syrphids (hoverflies) have been recognized as a threatened group due to environmental drivers, and these managed grasslands provide preferred habitats for them. Yet, at what extent syrphids respond to grassland management strategies is little known. We investigated whether abandonment of managed mountain meadows (mown once a year, no fertilizer use) affects syrphid abundance, richness and species assemblages. Moreover, the influence of plant richness, flower frequency and surrounding landscape on syrphids were assessed. Four managed meadows were compared with four abandoned meadows in a mountainous region in Austria. Adult syrphids were sampled over two consecutive years (June and August 2015–2016) by sweep netting using line transects and observation plots. Syrphid abundance was significantly higher in managed than abandoned meadows and increased with increasing plant richness and flower frequency across management types. Management and sampling time also significantly affected syrphid species assemblages. Syrphid richness and abundance were not influenced by surrounding landscape. We conclude that both abandoned and managed meadows provide important habitat types for syrphids. Therefore, efforts should be made to maintain both managed and abandoned meadows in order to protect syrphid species within mountainous landscapes.     

Vegetation changes in semi-natural meadows with unchanged management in southern Sweden, 1965–1990

Vegetation records from 1963–66 and 1990 were compared in eight semi-permanent plots from continuously managed, semi-natural hay-meadows from five sites in southern Sweden to find out whether changes in plant communities could be detected in these grasslands over a period of ca. 25 years in spite of unchanged management. It was found that species richness, α-diversity and species frequency distribution remained fairly constant over the period, while there was considerable species turnover. However, it was also found that the total cover of vascular plants increased as well as the proportion of graminoids, both in cover and in species number, with a corresponding decrease of forbs. The number of species favoured by mowing also decreased as well as the number of typical grassland species. The detected changes are interpreted partly as effects of increased loads of atmospheric nitrogen deposition, and partly related to small changes in management intensity (changed frequency of mowing and/or aftermath grazing). It is possible that the significant land use changes in adjacent areas resulting in large landscape changes (increase of forests and woodlands) might have negatively influenced the interchange of grassland species, leading to an increase in the number of non-typical grassland species in the meadows.     

Use of sheep grazing in the restoration of semi‐natural meadows in northern Finland

Abstract. The biodiversity of species‐rich semi‐natural meadows is declining across Europe due to ceased management. In this study we aimed to find out how successfully the local species richness of an overgrown semi‐natural mesic meadow could be restored by sheep grazing after a long period of abandonment. The cover of vascular plant species in grazed plots and ungrazed exclosures was studied for five years and the responses of different functional plant groups were followed (herbs vs grasses, tall vs short species, species differing in flowering time, species representing different Grime's CSR strategies and species indicative of rich vs poor soil). Grazing increased species number by nearly 30%. On grazed plots the litter cover practically disappeared, favouring small herbs such as Rhinanthus minor, Ranunculus acris, Trifolium pratense and the grass Agrostis capillaris. Grazing decreased the cover of the late flowering tall herb Epilobium angustifolium but had no effect on the abundance of the early flowering tall herbs Anthriscus sylvestris or Geranium sylvaticum. We suggest that to succeed in restoration it is useful to determine the responses of different functional plant groups to grazing. Grassland managers need this information to optimize the methods and timing of management used in restoration. Additional management practices, such as mowing, may be needed in mesic meadows to decrease the dominance of tall species. The availability of propagules seemed to restrict further increase of species richness in our study area.     

How do management and restoration needs of mountain grasslands depend on moisture regime? Experimental study from north‐western Slovakia (Western Carpathians)

Question: How does species composition change in traditionally managed meadows after mowing has ceased, and in abandoned meadows after re‐introduction of mowing? Are there differences in the dynamics of dry and moderately wet meadows? Location: Zázrivá‐Ple?ivá (19°11′N, 49°16′E), north‐western Slovakia, western Carpathians. Methods: Pairs of experimental plots (mown and unmown) were established to replicate each combination of dry/wet and traditionally managed/abandoned meadows. Changes in species composition were studied over 5 years. The data on changes in species composition was analysed by constrained and unconstrained ordinations, and visualized using Principal Response Curves. Results: Species composition of newly abandoned wet grasslands was changing towards the corresponding long‐abandoned plots even in the first year of abandonment. Similarly, newly established restoration mowing in abandoned dry grasslands rapidly shifted the stand species composition towards that of traditionally managed ones. Nevertheless, 4 year after reintroduction of mowing, the species composition of the restored plots was still far from the target composition. The effect of mowing in abandoned wet grasslands and abandonment in dry grasslands was much less pronounced and slower. Conclusions: Moisture regime is a very important factor determining the management needs of various grassland types. Wet grasslands are much more sensitive to abandonment, with a rapid degradation rate and limited possibilities for restoration, which can be extremely slow. Even in the dry grasslands, that quickly responded to restoration mowing, restoration is a long‐term process.     

Mowing may bring about vegetation change,but its effect is strongly modified by hydrological factors

We assessed the effect of long-term (11 years) mowing with a hand scythe on the distributions of plant species and species turnover within meadow communities dominated by Carex acutiformis and C. acuta in a small lowland river valley subjected to annual flooding. We hypothesized that mowing would trigger the process of species exchange toward multispecies communities according to the abiotic environmental gradients, as has been reported for traditionally used wet meadows. We found that mowing had a much greater impact on the increase in plant species abundance in wetter, subjected to deeper and longer flooding, zone of the valley than in its drier part. The treatment and hydrology had no substantial effect on the sedges and cryptogams while the graminoids and the forbs were the least stable components of the plant communities. We found that annual management was conducive to the appearance of numerous seedlings of Alnus glutinosa in the part of wetland situated close to the valley edge. Surface flooding was found to be a driving force modifying the impact of mowing on sedge meadows. This “hydrological resetting impulse” created a specific floristic reset of the community, bringing it to the more simplified forms. Mowing every 4–5 years at the beginning of August is advisable to protect sedge meadows distributed in flooded small lowland river valleys.     

Environmental and historical effects on lichen diversity in managed and unmanaged wooded meadows

Question: Which environmental and historical variables affect epiphytic lichen diversity in managed and unmanaged wooded meadows? Location: The island of Gotland located in the Baltic Sea east of the Swedish mainland. Methods: We examined total epiphytic lichen diversity (crustose, foliose and fruticose species) on 1148 trees in eight grazed, eight traditionally managed (mowing, hay gathering and pollarding) and seven unmanaged wooded meadows. In addition to management, data on site location, habitat structure, history and adjacent habitat were analysed. Results: Lichen species richness increased with wooded meadow area and was greater within managed sites than in unmanaged meadows. Historic crown cover (ca. 1930) also influenced present‐day lichen richness. Geographic location, distance to sea, wooded meadow area and average tree circumference were important determinants of lichen species composition. Tree circumference was the strongest overall predictor of the number of species on individual trees. However, tree circumference interacted with management regime, whereby cessation of management appeared to reduce species richness most on large trees. Traditionally managed sites, on average, supported the greatest richness of red‐listed species. Conclusions: Management regime, wooded meadow area and canopy cover were the main drivers of lichen species richness, while geographic location, wooded meadow area, distance to sea and average tree circumference were important determinants of species composition.     

Effectiveness of mowing for the flora diversity preservation: a case study of steppe-like grassland in Croatia (NATURA 2000 site)

Intensive human activities have resulted in a critical reduction of grasslands in Croatia, one of the richest Europian countries in terms of biodiversity. Sub-Pannonic steppic grasslands are very rare, only few occur in the continental part of Croatia and most of them are protected as NATURA 2000 sites. We studied vascular flora of the Sub-Pannonic stepic grassland with the aim to examine the changes in plant communities 15 years from the mowing application. Results showed that hand mowing once per year at the end of vegetation season increased the species diversity as well as the number of medium and low growth taxa. The critically endangered, nearly threatened and vulnerable species remained preserved. The occurrence of trees, shrubs and invasive species as a consequence of succession and anthropogenic influences could be important threat for steppe-like grassland flora. From a conservation point of view infrequent mowing regime could be an effective management tool, although it needs to be adapted to regional and local circumstances for maintaining high biodiversity of steppe-like grasslands and valuable plant species.     

Sustaining recovered grasslands is not likely without proper management: vegetation changes after cessation of mowing

Grasslands recovered by sowing low diversity seed mixtures of local provenance are usually managed by mowing. Besides restoration success only a few studies have focused on the direct effects of post-restoration mowing on recovered grassland vegetation. In this study we followed vegetation changes in 13 successfully recovered grasslands in 5 × 5-m-sized sites with continuous and ceased mowing at Hortobágy National Park, East-Hungary. We asked the following questions: (i) What are the effects of cessation of mowing on the vegetation structure and diversity of recovered grasslands? (ii) What are the effects of cessation of mowing on the abundance of sown grasses, target and undesirable species? (iii) Is yearly mowing an appropriate management tool for the maintenance of recovered grasslands? Our results showed that the cessation of mowing caused litter accumulation, while diversity, total vegetation cover and the cover of sown grasses decreased compared to the mown sites. The cover of undesirable perennial species was significantly higher in unmown sites than in mown ones. The species composition of mown sites remained more similar to near-natural grasslands than the unmown ones. Our results suggest that without regular post-restoration mowing the favourable status of recovered grasslands can rapidly decline due to litter accumulation and by the expansion of undesirable species, even in the short-run. We also stress that while yearly mowing is enough to maintain grasslands recovered by low-diversity seed sowing, it cannot be considered to be enough to recover target vegetation composition.     

Invertebrate herbivory along a gradient of plant species diversity in extensively managed grasslands

Increasing plant diversity has long been hypothesized to negatively affect levels of invertebrate herbivory due to a lower number of specialist insect herbivores in more diverse sites, but studies of natural systems have been rare. We used a planned comparison to study herbivory in a set of 19 semi-natural montane grasslands managed as hay meadows. Herbivory was measured in transects through the plant communities, and in individuals of Plantago lanceolata and Trifolium pratense that were transplanted into each meadow. In addition, plant community biomass and arthropod abundances were determined in the grasslands. Before the first mowing in June, mean herbivory levels correlated negatively with plant species richness, as predicted by theory, but they were also significantly affected by plant community biomass and plant community composition. After mowing, herbivory levels were only significantly related to plant community composition. Damage levels in the transplants were lower than herbivory levels in the established plant communities. Most insect herbivores were generalists and not specialists. The number of insect herbivores and spiders were positively correlated and tended to increase with increasing plant species richness. Herbivory levels were correlated negatively with spider abundances. We conclude that while the predicted negative relationship between plant species richness and insect herbivory can be found in grasslands, the underlying mechanism involves generalist rather than specialist herbivores. Our data also suggest a role of natural enemies in generalist herbivore activities.Electronic Supplementary Material Supplementary material is available to authorised users in the online version of this article at .     

Vegetation development in sown field margins and on adjacent ditch banks

The creation of temporal and newly sown field margins for 6 years is a common agri-environment scheme (AES) in the Netherlands. Conservation profits resulting from AES vary over different areas and need further studying. We examined plant species richness in such field margins and adjacent ditch banks in the province of Zeeland, where these linear elements do not experience plant biomass removal after mowing as management strategy. First, during 2 years, we inventoried field margins sown with a wildflower mixture and related the species composition and richness to the age of the margins. In a second assessment, we studied plant species richness on ditch banks protected from arable fields by these margins. Major clusters in a principal component analysis (PCA) on species composition in the field margins showed a succession from sown and ruderal annual species (year 1), to sown perennial species (year 2) and ending with a dominance by tussock forming grass species and Urtica dioica (year 5–6). Total plant species richness decreased with increasing age of the margins, and this was caused by the combination of a decline in sown species and a stable number of not-sown species. The presence of field margins during several years did not result in an increase in plant species richness on adjacent ditch banks. In both the field margins and on the ditch banks, mowing management is not followed by the removal of the cuttings. For plant conservation, the results of these field margins are disappointing, probably due to the lack of a proper management. Therefore, we recommend implementing a hay-making and opening management, to increase plant richness and to reduce noxious weeds in the margins and on the ditch banks.