Interactions between plant and insect diversity in the restoration of lowland calcareous grasslands in southern Britain

Abstract. The lowland calcareous grasslands of northwestern Europe are highly prized by ecologists and conservationists as a result of the diversity of their plant and invertebrate communities. Large areas of such grasslands have been lost this century as a result of changes in agricultural land use. Recent changes in agricultural policies, in particular the introduction of agri-environmental incentive schemes, have resulted in an increasing area being managed for the restoration of these communities. This paper reviews the management techniques employed in the restoration of lowland calcareous grasslands and the factors that govern their success. Constraints on the enhancement of the plant diversity of restoration sites include high soil fertility and the presence of undesirable species in the soil seed bank. However, it is thought that the primary constraint is the availability of propagules from which new populations can be established. Similarly, the dispersal mode and ability of insect species is likely to be the major factor limiting the enhancement of insect diversity. Evaluation of the success of restoration management usually involves monitoring changes in the plant community. However, as a result of their short life-cycles and sensitivity to small-scale environmental conditions, insects may respond more rapidly to changes resulting from restoration management and therefore provide better indicators of success. With the exception of a few high-profile butterfly species, the use of insects as indicator taxa has largely been neglected in terrestrial systems. This paper illustrates their potential use with reference to lowland calcareous grasslands in southern Britain.     

An evaluation of the use of translocated blanket bog vegetation for heathland restoration

Abstract. The decision was taken by an opencast coal mining company to translocate on-site blanket bog vegetation, on completion of mining, at a site in Co. Durham, UK, both to preserve it and to use it to enhance recolonization. The vegetation of the treatments was monitored for seven years after site completion and this paper reports on the progress of the translocated material and its effect on recolonization. Translocation of large turves of blanket bog into carefully prepared receptor cells preserved most of the vegetation intact, but resulted in severe decline in the frequency of Sphagnum, while the design of the receptor site as strips of translocated vegetation enclosing strips of spread, stored peat accelerated recolonization of the intervening bare peat by Calluna vulgaris, but not of other target species. This attempt to translocate blanket bog vegetation and at the same time use it to accelerate recolonization was only partly successful. It was concluded that the ecological requirements of species known to be significant for ecosystem function, such as Sphagnum, must be fulfilled if translocation of blanket bog is to be attempted in future.     

Seeds of change: The value of using Rhinanthus minor in grassland restoration

Question: What is the value of using Rhinanthus minor in grassland restoration and can restrictions on its establishment be overcome? Location: England (United Kingdom). Methods: Two experiments were established to determine the efficacy of inoculating R. minor on a suite of four agriculturally improved grasslands and the efficacy of using R. minor in grassland restoration. In Experiment 1, the effect of herbicide gap creation on the establishment and persistence of R. minor in grasslands ranging in productivity was investigated with respect to sward management. In Exp. 2, R. minor was sown at 1000 seeds/m² in conjunction with a standard meadow mix over a randomized plot design into Lolium perenne grassland of moderate productivity. The treatment of scarification was investigated as a treatment to promote R. minor. Results: Gap size had a significant role in the establishment and performance of R. minor, especially the 30 cm diameter gaps (Exp. 1). However, R. minor failed to establish long‐term persistent populations in all of the agriculturally improved grasslands. In Exp. 2, establishment of R. minor was increased by scarification and its presence was associated with a significant increase in Shannon diversity and the number of sown and unsown species. Values of grass above‐ground biomass were significantly lower in plots sown with R. minor, but values of total above‐ground biomass (including R. minor) and forb biomass (not including R. minor) were not affected. Conclusions: The value of introducing R. minor into species‐poor grassland to increase diversity has been demonstrated, but successful establishment was dependent on grassland type. The scope for using R. minor in grassland restoration schemes is therefore conditional, although establishment can be enhanced through disturbance such as sward scarification.     

Changes over three decades in the floristic composition of nutrient-poor grasslands in the Swiss Alps

To investigate recent changes in the floristic composition and nature conservation value of nutrient-poor, semi-natural grasslands of the Swiss Alps, we resurveyed 151 phytosociological relevés in four regions, originally recorded between 1975 and 1985. In the original surveys, the mean number of plant species per plot (25–100 m²) ranged from 47.1 to 58.1 according to region. The flora included a total of 18 species that are protected in Switzerland and a high proportion of habitat specialists of nutrient-poor grasslands (NPG-species). In the second survey, conducted between 2002 and 2004, both mean species number per plot (−3.2 to +11.4) and species evenness (−0.05 to +0.07) were higher in most regions. However, the data revealed clear shifts in community composition, with a higher proportion of nutrient-demanding species (mean nutrient indicator value increased by +0.07 to +0.24 units) and a lower proportion per plot of NPG-species (−3.6 to −11.6%). These changes were greatest in pastures, and in meadows converted to sheep pastures, while the NPG-species were maintained in unfertilized meadows that were managed as ecological compensation areas. To prevent continuing decline in the conservation value of these grasslands, it is important to support low-intensity management, especially mowing, and to prevent further eutrophication.     

A field assessment of the success of moorland restoration in the rehabilitation of whole plant assemblages

Question: How successful is restoration that is focused on a single dominant plant at enabling the reassembly of the whole vegetation assemblage and what factors affect the relative success of such restoration? Location: Moorlands in England and Scotland, UK. Methods: Vegetation composition was sampled in grassdominated (degraded), restored and long established Calluna vulgaris‐dominated (target) areas within eight moorland restoration sites. Additional soil and biogeographic data were collected. Data were analysed by Canonical Correspondence Analysis, which allowed the impact of moorland management to be examined. Results: All sites showed good restoration success when the dominant managed species (Calluna vulgaris, Molinia caerulea and Nardus stricta) were considered. Restoration success of the remainder of the plant assemblage, disregarding these dominant species, was lower with restored samples at some sites differing little from their respective degraded samples. Moors restored solely by grazing exclusion were more similar to their respective targets than were those restored by more intensive mechanical means. The most important factors in explaining vegetation assemblages were management status (i.e. whether samples represented degraded, restored or target parts of the moor) and latitude. Conclusions: The project demonstrates that, where possible, restoration should be attempted by grazing exclusion alone. Furthermore the importance of applying local restoration targets and of monitoring the whole plant assemblages when assessing the success of moorland restoration is highlighted.     

Grazing with Galloway cattle for floodplain restoration in the Syr Valley, Luxembourg

Three years after a river restoration scheme in the Syr Valley (Luxembourg) we investigated habitat development and habitat use of Galloway cattle deployed in a low-intensity grazing system on a permanent floodplain pasture. Habitats were delimited with a mobile GPS/GIS mapping system and their spatial development was assessed over three consecutive years. During these three years, the patches of the six habitats decreased to 40% of mean initial size, and a rapid net area expansion of wetland habitats (large sedge swamps: +100%, marsh and tall forb grasslands: +43%) was observed. The behavioural patterns and grazing preferences of the cattle were observed directly during the vegetation period in June, August and November. These observations were complemented by a transect analysis of cattle impact indicators in June and November. The cattle grazed the different habitats very selectively, as they preferred the mesophilic, and ruderal grasslands 1.6, and 5.6 times more than expected respectively. During the growing season, the grazing niche breadth declined (3.92 in June to 2.68 in November), and less preferred forage habitats like large sedge swamps were grazed primarily in the autumn. We used bite and step rates to investigate grazing intensity by habitat type. During summer, grazing intensity correlated with forage quality in the different habitat patches, whereas in autumn it was obviously influenced by the effort required to access the desired forage plants in a given habitat. The impact indicators revealed a matter transfer from riparian areas to the valley edge. Here, we give a first insight into habitat development and habitat use of Galloway cattle in a recently restored floodplain area and derive recommendations for the adaptive management of future projects.     

Response of the moss Racomitrium lanuginosum to changes in sheep grazing and snow‐lie due to a snow‐fence

Question: What are the responses of Racomitrium lanuginosum moss to altered snow‐lie and sheep use? Location: ACarexbigelowii‐Racomitrium lanuginosum heath on a Scottish montane plateau affected since 1986 by a fenced ski corridor. Methods: Permanent quadrats were set up along transects 45 m long perpendicular to the snow‐fence. Cover was assessed over a 12‐year period from 1990. Pellet‐group clearance counts provided data on sheep usage between 1990 and 1996. Snow‐lie was mapped in the springs of 1991–1996. Results: The snow‐fence created a gradient in sheep use and altered the duration of snow‐lie. At the start of monitoring Racomitrium cover was lower immediately adjacent to the fence, and after 12 years its cover was significantly reduced within 10 m of the fence. Further away from the fence Racomitrium cover was relatively stable. The loss of Racomitrium was correlated both with increased snow‐lie and heavier sheep usage. Grass cover increased near the fence and was related to sheep use. Dicranum fuscescens responded differently to Racomitrium, increasing significantly near the fence. Conclusions: We found that changes in snow‐lie and grazing pressure quickly brought about vegetation change in this montane ecosystem. Racomitrium was the most sensitive species to the changes in grazing and snow‐lie caused by the fence, having the biggest initial changes. Loss of Racomitrium permitted increases of species more resistant to grazing including Dicranum fuscescens and grasses.     

Vegetation re‐establishment on land previously subject to control of Pteridium aquilinum by herbicide

Abstract. Vegetation development after conventional bracken control with herbicide in the uplands is often slow and rarely results in the development of the target community. Two experiments were established on contrasting sites to investigate cost‐effective methods for the re‐establishment of vegetation dominated by Calluna vulgaris (heather), which is the usual target community seen as the desired replacement for Pteridium aquilinum in the uplands by many land managers. At one site, where grazing intensity was low and shelter was sufficient to prevent rapid Pteridium litter loss, the necessary treatments to establish Calluna were to disturb the litter and add a source of Calluna seeds. A difference between continued Pteridium control or a relaxation of control was not observed over the time‐scale of the experiment. At the second site, where grazing intensity was high and shelter minimal, vegetation dominated by Rumex acetosella had developed on a thin but dense litter layer. To enhance Calluna regeneration it was necessary to prevent sheep grazing, disturb the remaining litter layer and add a source of Calluna propagules. A nurse crop hindered Calluna establishment due to its vigorous growth. At neither site was an initial application of fertilizer an aid to Calluna establishment. It is possible to devise methods for the re‐establishment of Calluna at sites where previous attempts at Pteridium control alone have not had the desired effect on vegetation development. However, the precise choice of methods used is dependent on the grazing management at the site. High levels of livestock grazing prevented Calluna establishment, but some trampling may aid litter breakdown. The ability to adjust stocking rates and access to the site by machinery are critical factors to take into account in producing an effective strategy for long‐term Pteridium control and its replacement by target vegetation.     

A multi-site assessment of the effectiveness of Pteridium aquilinum control in Great Britain

Questions: The objectives of this study were to assess the effect of Pteridium aquilinum (bracken) control treatments, at the national scale, and the impact of restoration practices, at the local scale, on P. aquilinum performance. Hypotheses: 1. Geographical location (locally between and within sites) affects the control of P. aquilinum through time. 2. Are the P. aquilinum control treatments successful at all sites, and if so which ones? 3. The treatments applied at the individual site level to restore vegetation influences the performance of P. aquilinum through time. Location: Four geographically distinct acid grassland and heathland sites infested with P. aquilinum across Great Britain. Methods: Six main-plot, bracken control treatments were applied to all sites with site-specific vegetation restoration treatments. Response variables (P. aquilinum cover, frond length and density) were monitored twice yearly, in June and August between 1993 and 2003. Results: Between- and within-site spatial variation was found, although impact is perhaps less than suggested from shorter-term data. Despite local variation all sites responded similarly to bracken control treatments; asulam treatment resulted in a rapid reduction in frond performance followed by a continued recovery taking approximately ten years to return to untreated values. Cutting treatments tended to have a slower impact at the start but an increasing one over time, especially cutting twice per year. Restoration treatments had a limited impact; the only significant effect in August was grass seeding on frond length at Sourhope. In June only, the plots where sheep were fenced out showed a significant reduction in P. aquilinum cover at Peak. Conclusions: Long-term control of Pteridium aquilinum at all sites and on all measures was best achieved using a continuous cutting treatment, preferably twice per year.     

Blue Data Books - an encouraging new instrument for restoration and conservation

Abstract. In order to stress successes in restoration and conservation of species, regionally, nationally or globally, a new instrument has been developed, the ‘Blue Lists’, ‘registers of those Red List species that show a durable overall stabilization or increase of abundance in the region (nation, world) considered’. ‘Blue Data Books’ include additional information on the ecology, conservation, and promotion of the species. For describing the overall change of abundance of every individual species six categories are defined. Furthermore, the effects of nature conservation techniques (NCT) on a species are evaluated using six additional categories. In a test area of 3 431 km² in northern Switzerland, information was compiled on the change in abundance of the 708 Red List species of higher plants (spermatophyta) over the last 10-15 yr. Overall, 33% of these species showed a stabilization or even increase in abundance; these are Blue List species; ca.20% showed a decline and for almost 50% the change in abundance is not known. NCTs have been successfully applied to more than 50 % of all the species, at least locally. For about a further 30%, the required NCTs are known but have not yet been tried out. If these techniques were applied on a large scale, the decline of very many species in the test area could be stopped.     

Composition and seasonal flux of the soil seed bank of species-rich Themeda triandra grasslands in relation to burning history

Abstract. Most species-rich grasslands dominated by Themeda triandra in southeastern Australia have been ungrazed and frequently burnt for decades. The seedling emergence technique was used to determine the size and taxonomic composition of the soil seed bank of five grasslands that had different fire histories (i.e. burnt at 1 yr, 3 yr and > 10 yr intervals) and this was compared to the standing vegetation at each site. A nested sampling design (subplot, plot, site) was used to determine the effect of spatial scale on the patterns observed in both the vegetation and the seed bank. Temporal variation in the seed bank was assessed by repeated soil sampling over a two year period. 61 native and 30 exotic species were recorded in the vegetation. Richness varied more between sites than within sites. Sites were therefore internally homogeneous for species richness. However, no correlation between burning frequency and richness was found. DCA ordination separated the sites into distinct groups, but sites with similar fire history did not necessarily group closely. 60 taxa germinated from the soil seed bank, comprising 32 native and 28 exotic species; 11 species, mostly therophytes, were restricted to the seed bank. The richness of the seed bank was significantly lower than the vegetation at all spatial scales. No correlation between seed bank richness and fire history was found. The seed bank of species-rich grasslands is dominated by a limited number of widespread, highly clumped, annual, native and exotic monocots. Most native hemicryptophytes, and perennials in general, were represented in the soil by a transient seed bank. Only 12 % of study species, all therophytes, were considered to form large, persistent seed banks, the size of which was greater in unburnt grasslands at all times of the year. The distinct floristic patterns observed in the vegetation are less clearly represented in the seed bank. The seed bank represents a floristically distinct (and less variable) component of the vegetation when compared to the standing flora. The size of the long-term soil seed bank suggests that it has little functional importance for many native species and probably contributes little to seedling regeneration processes following disturbance. Altering established fire regimes is likely to only change the composition and small-scale richness of the existing site vegetation and will not re-integrate species previously lost from the vegetation due to past management. It is suggested that the maintenance of vegetation remnants and processes that encompass a range of long-term burning histories will be necessary if the flora is to be conserved in situ. Restoration of degraded grasslands cannot rely on the soil seed bank but rather, will be dependent on the reintroduction of propagules.     

Effects of topsoil removal,seed transfer with plant material and moderate grazing on restoration of riparian fen grasslands

Question: How do moderate grazing, topsoil removal and hay transfer affect species diversity and abundance on a eutrophic fen grassland site? Location: Northern Germany. Method: A three-factorial field experiment with the factors grazing, topsoil removal and hay transfer of diaspore-rich material was established in 2001. Soil nutrients and seed bank were analysed at the beginning of the experiment, species composition and vegetation development was monitored for four years (2002–2005). Results: Topsoil removal had a significant effect on the abundance of different plant species groups: resident vegetation of agricultural grasslands was suppressed, while clonal reed species were facilitated in recolonising the area. The establishment of regionally rare and endangered species of nutrient-poor fens and wet meadows introduced with hay was achieved mainly on plots with topsoil removal, with the exception of Rhinan-thus angustifolius, which also established on plots with intact topsoil. Effects of grazing after four years of experiments were of minor influence on species composition. Conclusion: The establishment of target plant species of nutrient-poor fens is most successful when both an adequate number of viable diaspores and suitable sites for germination and establishment are available. In our experiment this was achieved by the combination of topsoil removal and hay transfer. We recommend this combination, together with continuous management (grazing/cutting), for further restoration in fen grasslands.     

Hydrology of Dutch Cirsio‐Molinietum meadows: Prospects for restoration

Abstract. Fen meadows (Cirsio dissecti‐Molinietum) are seriously threatened by desiccation, acidification and eutro‐phication. In The Netherlands several projects were launched to restore damaged fen meadows. This review describes how successes and failures of these restoration projects depend on hydrological systems. Six hydrological systems have been distinguished, which all provide the site conditions required by this community. Nowadays, the best developed fen meadows are found in the higher Pleistocene landscape of The Netherlands, where they depend on base‐rich groundwater discharging from local or large groundwater systems. Fen meadows of the lower Holocene landscape usually occur in man‐made surface water systems. Almost all stands have been severely deteriorated. Restoration of fen meadows in the Pleistocene landscape is promising when the hydrology is only slightly disturbed or when hydrological measures are taken in combination with sod cutting. Restoration prospects of fen meadows in the Holocene landscape are low. Until now a complete regeneration of Cirsio‐Molinietum meadows has not been realized. Restoration measures failed to restore high pH values in the top soil. It is hypothesized that viable seeds of many target species lack in the soil seed bank. In addition, the dispersal capacities of these species seem to be limited.     

Seed bank classification and its importance for the restoration of species-rich flood-meadows

Abstract. This paper focuses on the persistence of seeds in the seed bank of a formerly fertilized flood-meadow into which seeds from an original Alopecurus pratensis-Sanguisorba officinalis community have been introduced. The longevity of seeds in the seed bank of the original community is also estimated. The established vegetation was compared with the seed bank, as divided into two layers (0 - 5 cm and 5–10 cm); this allowed a classification of species into three groups, with (1) transient, (2) short-term persistent and (3) long-term persistent seed bank. The majority of the species of the unfertilized flood-meadow community have a transient or short-term persistent seed bank with seeds showing a large variance in shape i.e. flattened or elongate. Because of this, the characteristic flood-meadow species will disappear soon after the beginning of fertilizer application and will not re-establish from the seed bank, once the fertilizer application is ceased. The formerly fertilized flood-meadow contains many ruderal and arable weed species, the seeds of which tend to be compact or round. Significantly more seeds were found in the seed bank of the formerly fertilized flood-meadow under cattle-grazing than under sheep-grazing and no grazing.     

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.     

The use of spatial concepts as a basis for designing a viable-habitat network: Conserving redstart (Phoenicurus phoenicurus) populations in Sherwood Forest, England

This paper presents a method for the design of a habitat network in a fragmented landscape. It employs metapopulation dynamics and connectivity to develop spatial rules which, together with the requirements of a target species, can be used to guide the creation of a network to improve population persistence. Using these guidelines, two conservation strategies of patch enlargement and corridor creation are developed. The spatial models used to produce the habitat network are described. The resulting maps show the network consisting of the source patches that could be occupied by stable populations and a number of patches, which could support viable local populations, situated in the proximity of the sources or linked to them. The approach could be used to inform decision-making in nature conservation and landscape planning.     

Mechanism and stoichiometry of the redox reaction between iron(III) and caffeic acid

The stoichiometry of the redox reaction of caffeic acid with iron(III) was determined at pH 2.5. A linear increase in the yield of iron(II) was found with increasing iron(III) concentration until reached constant values when iron(III)/caffeic acid molar ratios were higher than 9. The reaction proceeds through two steps each having different rates, and involving intermediates with different redox activities. A mechanism of the redox reaction consistent with our results is proposed.     

Genetic diversity of the endangered Chinese endemic herb Saruma henryi Oliv. (Aristolochiaceae) and its implications for conservation

Saruma henryi Oliv., the only representative of the monotypic genus Saruma Oliv. (Aristolochiaceae), is an endangered perennial herb endemic to China. It is a phylogenetically, ecologically, and medicinally important species. In the present study, inter-simple sequence repeat (ISSR) markers were employed to investigate the genetic diversity and differentiation of 14 populations. A total of 16 selected primers yielded 175 bright and discernible bands, with an average of 10.94 per primer. POPGENE analysis showed that the genetic diversity was quite low at the population level (h = 0.0447–0.1243; I = 0.0642–0.1853; PPB = 10.29–36.57%), but pretty high at the species level (h = 0.2603; I = 0.3857; PPB = 73.71%). The hierarchical analysis of molecular variance (AMOVA) revealed a high level of genetic differentiation among populations (67.18% of total variance components, P < 0.001), in line with the gene differentiation coefficient (G _ST = 0.6903) and the limited among-population gene flow (N _m = 0.2243). Both Principal Coordinates Analysis (PCoA) and UPGMA cluster analysis supported the grouping of all 14 populations into three geographic groups, among which there occurred a moderate level of genetic differentiation (33.18% of total variance components, P < 0.001) as shown by AMOVA analysis. In addition, Mantel test revealed a significant correlation between genetic and geographic distances among populations (r = 0.7792, P = 0.001), indicating the role of geographic isolation in shaping its present population genetic structure. The present levels and patterns of genetic diversity of S. henryi were assumed to result largely from its breeding system, geographic isolation, clonal growth, its unique biological traits and evolutionary history. The high genetic differentiation among populations implies that the conservation efforts should aim to preserve all the extant populations of this endangered herb.