Effects of grazing on plant communities and successional processes vary along an aridity gradient at a northern temperate grassland

The intermountain grasslands of North America reach their most northern geographic extent in interior British Columbia’s Cariboo-Chilcotin region. Here, this study examined the long-term effects of livestock grazing exclusion and reductions in grazing severity on plant community characteristics including plant and litter cover, species richness and abundance of leading species of 33 grassland sites across a broad aridity and soil property gradient. Across the aridity gradient, grazing reduced species richness, plant cover, and litter cover. However, the effects of grazing on dominant species varied across the gradient. In more arid grasslands, historical grazing substantially reduced cover of late-seral native bunchgrass Psuedoroegnaria spicata, and repeated measurements indicate that very long time periods are necessary for successional processes associated with recovery of native bunchgrasses. At the cool-wet end of the aridity gradient, successional processes are more rapid but dominated by exotic species Poa pratensis and Tragopogon pratensis. Recent (past 20 years) light grazing and rest-rotation have favored Poa pratensis at the expense of native needlegrasses (Achnatherum spp. and Hesperostipa spp.). We suggest that absence of a dominant large-stature native bunchgrass for mesic grasslands was a key factor in the invasion and dominance of Poa pratensis.

     

Trends and fluctuations in species abundance and richness in Mediterranean annual pastures

Abstract. Floristic data collected from permanent plots during 14 consecutive years are used to model the frequency of the 62 most abundant species in relation to post-ploughing succession, topography and rainfall in annual Mediterranean grasslands located in a Quercus rotundifolia dehesa. The interannual dynamics of species richness are also analysed. From 1980 to 1993, presence/absence data of grassland species were noted in five 20 cm X 20 cm permanent quadrats placed at random in 1980 in 14 permanent plots on a south-facing slope along the topographic gradient. Weekly autumn rainfall data over the 14 years were analysed using a profile attributes index and Hybrid Multidimensional Scaling to arrange the years according to their autumn rainfall pattern. Generalized Linear Models were used to fit the species richness and species frequency according to topographic position, age since the last ploughing episode, total rainfall in the growing season and autumn rainfall pattern using a forward stepwise procedure. The richness model includes all of these variables, and reveals a relatively high goodness-of-fit (71 %). The fact that the meteorological factors play a key role in modelling richness forces us to include them if we wish to use richness as an indicator of the degree of disturbance in these highly fluctuating annual pastures. Models of species dynamics show that although roughly 33 % of the species have a successional behaviour, the majority are more dependent on temporal heterogeneity associated with rainfall or spatial heterogeneity linked to the topographic gradient.     

Measuring the Short‐term Success of Grassland Restoration: The Use of Habitat Affinity Indices in Ecological Restoration

Agricultural intensification threatens grasslands worldwide and the restoration of grasslands from arable lands can at least partially counter this threat. We studied grassland restoration by following early successional changes of arthropod assemblages (spiders, Araneae; true bugs, Heteroptera; orthopterans, Orthoptera; and ground beetles, Carabidae) on 1‐ and 2‐year‐old restorations using arable lands and native grasslands as two ends of the succession timescale. To examine the changes in species composition among the habitat types, we used habitat affinity indices based on fidelity and/or specificity of the species. We found that the number of species did not differ between habitat types, while species composition changed markedly with time. Species richness was thus not adequate to detect favorable changes after grassland restoration. Habitat affinity indices, on the other hand, were useful to detect compositional changes caused by the increasing numbers of species characteristic of target grasslands as early as the second year after restoration. Habitat affinity indices are easy‐to‐use, easy‐to‐interpret measures of restoration success; therefore, we recommend their use as measures complementary to species richness and simple similarity. Our results show that sowing low‐diversity seed mixture followed by mowing and grazing can be particularly successful in grassland restoration in time periods as short as 2 years.     

Orthoptera as ecological indicators for succession in steppe grassland

Understanding the effects of land-use on threatened ecosystems is of special relevance for nature conservation. The aim of our study was to use Orthoptera as ecological indicators for succession in Central European steppe grasslands.Orthoptera showed a clear response to succession. Each successional stage harboured a unique assemblage. Species richness of habitat specialists was highest in the earliest seral stages. In contrast, density of all species peaked at the intermediate successional stage. Early successional stages are mostly likely to be preferred by specialized Orthoptera because they provide warm suitable oviposition sites (bare ground) and microclimatic conditions. The density peak in the mid-successional stage probably reflects a trade-off between favourable ambient temperatures for optimal development, sufficient food and shelter against predators.Although all successional stages of steppe grassland are relevant for conservation, early and mid-successional stages are the most important. Consequently, conservation management should aim at re-introduction of a traditional, low-intensive land use for abandoned steppe grasslands. As an optimal land use, we recommend traditional rough grazing with sheep and goats, which creates a heterogeneous habitat structure with bare ground, and avoids the accumulation of litter, favouring Orthoptera.     

Restoration of Seminatural Grasslands: What is the Impact on Ants?

The number of species‐rich seminatural grasslands in Northern Europe has decreased significantly due to the abandonment of traditional land use practices. To preserve these habitats, an increasing number of abandoned and overgrown grasslands have been restored by cutting down trees and shrubs and reintroducing grazing. These practices are considered a useful tool to recover the species richness of vascular plants, but their impact on other taxa is hardly known. Here we studied ants as one important group of grassland insects. We investigated (1) the effects of restoration of nongrazed and afforested seminatural grasslands, compared to continuously managed reference sites; and (2) the modulating impacts of habitat characteristics and time elapsed since restoration. We found a total of 27 ant species, 11 of these were characteristic of open habitats and seven characteristic of forests. Neither species richness per site nor the number of open‐habitat species, nor the number of forest species differed between restored and reference sites. Yet, within the restored sites, the total species richness and the number of open‐habitat species was positively related to the time since restoration and the percentage of bare rock. High frequencies of most open‐habitat species were associated with low vegetation, older restored sites, and reference sites. Most forest species showed their highest frequencies in tree‐ and shrub‐dominated habitat. We conclude that restoration efforts have been successful in terms of retrieving species richness. A regular and moderate grazing regime subsequent to the restoration is suggested in order to support a high abundance of open‐habitat species.     

A functional analysis of vegetation dynamics in abandoned and restored limestone grasslands

Question: How do various plant functional groups and types are related to regeneration of limestone grasslands and to temporal changes in cover of trees and shrubs? Which mechanisms are responsible for changes in differently treated grasslands? Location: Southern Poland, 50° 01'N, 19° 50'E. Methods: A 12‐year restoration experiment after the clearing of a 35‐yr‐old secondary pine wood developed on limestone grassland, and a parallel observation of succession in neighbouring abandoned grassland were carried out. Changes in cover of species with different functional traits and established strategies in relation to time and cover of shrub layer were analysed using multiple linear regression. Results: Over 12 years the composition and cover of species changed in the studied grasslands, but significant differences still occurred between the old grassland and the grassland restored in former wood gaps and that developed in former closed wood. Despite the very close proximity of the old grassland, the composition of restored grasslands was still much determined by initial conditions. In these grasslands, the cover of Festuco‐Brometea species was significantly lower than in the old grassland. In all sites, the cover of species with large leaves and with leaves distributed regularly along the stem, and the cover of CS strategists increased. With increasing shrub cover in restored grasslands, the cover of annuals and biennials, tall species, and R and CS strategists decreased. Conclusions: Limited availability of seeds seems the principal reason for the weak regeneration of xerothermic limestone grasslands. Trees and shrubs inhibit the developed of both early successional species and perennials occurring in established communities. They significantly hinder regeneration of grasslands and should be cut every five or six years.     

Dynamics of species richness and composition in a limestone grassland restored after tree cutting

Abstract. After abandonment many limestone grasslands have been overgrown by trees and shrubs; as a result, species-rich communities with many regionally rare and endangered species are vanishing. Some studies suggest that, in cases where grassland is being restored, the species composition and rate of change is strongly dependent on the initial conditions, i.e. the earlier presence of grassland species and the opportunity for colonization of new sites by grassland species. These hypotheses were tested in a five-year restoration experiment after the clearing of a 35-yr-old secondary pine wood developed on abandoned grassland. Tree cutting induced rapid changes in the floristic composition and species cover. The number of grassland species from the class Festuco-Brometea increased significantly in the restored grassland, but their cover was much lower than in the old grassland. Canonical Correspondence Analysis showed significant differences in species composition between the grassland restored in former wood gaps and that developed in former closed wood. In wood gap sites the cover of species from the class Molinio-Arrhenatheretea and tufted perennials was much higher, whereas the cover of Festuco-Brometea species was lower. Significantly more shrubs, woodland species, ruderal and nitrophilous species as well as annual and biennial species occurred in the former closed wood site. It was found that richness and composition of the restored grassland depended strongly on the community composition before tree cutting, as well as on the presence of grassland species in the neighbourhood. Periodical tree cutting enables the maintenance of a temporal-spatial mosaic of scrub-grassland communities in isolated habitats and the preservation of local species diversity.     

The conservation value of restored landfill sites in the East Midlands, UK for supporting bird communities

There has been a rapid decline of grassland bird species in the UK over the last four decades. In order to stem declines in biodiversity such as this, mitigation in the form of newly created habitat and restoration of degraded habitats is advocated in the UK biodiversity action plan. One potential restored habitat that could support a number of bird species is re-created grassland on restored landfill sites. However, this potential largely remains unexplored. In this study, birds were counted using point sampling on nine restored landfill sites in the East Midlands region of the UK during 2007 and 2008. The effects of restoration were investigated by examining bird species composition, richness, and abundance in relation to habitat and landscape structure on the landfill sites in comparison to paired reference sites of existing wildlife value. Twelve bird species were found in total and species richness and abundance on restored landfill sites was found to be higher than that of reference sites. Restored landfill sites support both common grassland bird species and also UK Red List bird species such as skylark Alauda arvensis, grey partridge Perdix perdix, lapwing Vanellus vanellus, tree sparrow, Passer montanus, and starling Sturnus vulgaris. Size of the site, percentage of bare soil and amount of adjacent hedgerow were found to be the most influential habitat quality factors for the distribution of most bird species. Presence of open habitat and crop land in the surrounding landscape were also found to have an effect on bird species composition. Management of restored landfill sites should be targeted towards UK Red List bird species since such sites could potentially play a significant role in biodiversity action planning.     

Rapid recovery of soil arthropod assemblages after exotic plantation tree removal from hydromorphic soils in a grassland‐timber production mosaic

Many wetland systems are being lost or degraded by human activities such as plantation forestry. Therefore, efforts to restore these wetland systems are important for biodiversity recovery. We assess the recovery of arthropod assemblages that occupy hydromorphic grassland topsoil and leaf litter after the removal of exotic pine trees. We sampled arthropods in three biotopes (natural untransformed hydromorphic grasslands, restored hydromorphic grasslands, and commercial pine plantations) replicated across a large‐scale timber‐grassland mosaic in the KwaZulu‐Natal Midlands, South Africa. In the restored sites, overall species richness, as well as species richness of spiders, ants, and orthopterans, was significantly higher than in plantations, and was as high as in natural, untransformed sites. Additionally, overall assemblage structure along with spider, beetle, ant, and orthopteran assemblages showed no significant differences between restored and natural grasslands. Therefore, pine tree removal enables recovery of these arthropod taxa to levels similar to those in natural hydromorphic grassland. Recovery was rapid, with the assemblages in some restored sites resembling those in untransformed sites after only 6 years, indicating a high level of resilience and recovery in these systems. Contrary to expectations, time since pine removal had a negative effect on arthropod recovery. This was due to the strongly negative effect of alien invasive American bramble (Rubus cuneifolius), which was most common in older restored sites, causing deviation from their restoration trajectory. The potential for arthropod recovery in these hydromorphic grasslands is high, but successful restoration is dependent on ongoing appropriate grassland management, especially removal of bramble.     

Early succession of butterfly and plant communities on set-aside fields

 Hypotheses on secondary succession of butterfly and plant communities were tested using naturally developed 1- to 4-year-old set-aside fields (n = 16), sown fields (n = 8) and old meadows (n = 4) in 1992 in South Germany. Pioneer successional fields (1st and 2nd year of succession, dominated by annuals) and early successional fields (3rd and 4th year of succession where perennials, especially grasses became dominant) had fewer plant species than mid-successional fields (old meadows). In contrast to established hypotheses, mean number of plant species decreased from 1- to 4-year-old set-aside fields. Species richness of butterfly communities did not change during the first four years of succession, but species composition changed greatly. Pioneer successional fields were characterized by (1) specialized butterflies depending on annual pioneer foodplants (e.g. Issoria lathonia), and (2) species preferring the pioneer successions despite their host plants being more abundant on early and mid-successional fields (e.g. Papilio machaon). The variability in butterfly species richness was best explained by flower abundance which was closely correlated with plant species richness. Species whose abundance was correlated with habitat connectivity were significantly smaller than species which correlated with flower abundance. Numbers of caterpillar species were correlated with numbers of adult butterfly species. Life-history features of butterflies changed significantly from pioneer to early and mid-successional fields. We found decreasing body size and migrational ability, decreasing numbers of species hibernating as imago, decreasing numbers of generations and increasing larval stage duration with age of succession, but, contrary to expectation, host plant specialization, numbers of egg-cluster laying species and egg diameter did not change with successional age. Received 18 September 1995 / Accepted: 17 July 1996     

松嫩平原不同生境蝴蝶群落多样性及其保护建议

随着气候变化加剧和人类活动影响,生物多样性变化及其保护逐渐受到广泛关注。蝴蝶作为开花植物的传粉媒介和生态环境监测及评价的关键指示者,其多样性变化能够在一定程度上反映生境状况,因此,有必要清晰认识不同生境中的蝴蝶多样性变化。为明确松嫩平原蝴蝶资源和不同生境的群落多样性差异,采用样线法于2016年5月-2018年8月对松嫩平原的割草草地、湿地、农田、放牧利用草地及恢复草地共五种生境类型进行调查研究。结果发现,调查共记录蝴蝶5108头,隶属于6科21属26种,其中牧女珍眼蝶（Coenonympha amaryllis）和红珠灰蝶（Plebejus argyrognomon）为优势种类,分别占蝴蝶个体总数的25.61%和31.66%,且在五种生境类型中均有分布。不同生境类型中,蝴蝶群落的物种丰富度指数和均匀度指数无明显差异,而恢复草地生境的蝴蝶群落Shannon-Wiener多样性指数较高,优势度指数较低。农田生境中的蝴蝶个体数量较少,且群落组成与其他四种生境之间均具有显著差异。五种生境类型中的蝴蝶数量和多样性均呈现一定的月动态和年动态变化趋势。除湿地和农田外,其余三种生境中蝴蝶物种和个体数量从5月到8月均持续升高。四种生境的蝴蝶物种数量、个体数量（除农田外）在2018年均出现明显下降趋势。物种丰富度指数等指标的月动态和年动态在不同生境类型间存在较大差异。这些结果表明,生境类型和人类活动与蝴蝶多样性变化关系密切,表现为单一生境中蝴蝶多样性较低,复杂生境有利于保护蝴蝶多样性。本研究有助于厘清松嫩平原蝴蝶资源的基础数据,并为该地区蝴蝶多样性保护和利用及评估该区域生态环境提供一定理论支撑。     

Experimental small-scale grassland fragmentation alters competitive interactions among ant species

Different species may respond differently to habitat fragmentation. Theory predicts that abundant generalist species should be less affected by fragmentation than specialist species. In ant communities, the most abundant species is often behaviourally dominant. Thus, habitat fragmentation could alter competitive interactions between the dominant ant species and the other species. We tested this hypothesis in a long-term grassland fragmentation experiment. Fragments of different size (20.25 and 2.25 m²) were isolated by a 5-m wide strip of frequently mown vegetation. Control plots were situated in adjacent undisturbed grassland. Ant density and species composition were assessed 3 and 6 years after initiation of the experimental fragmentation. The effect of the dominant ant species on the resource use of the other species was examined at natural sugar resources (aphids and extrafloral nectaries) and at artificial sugar baits. Lasius paralienus was the most abundant ant species (72% of nests) in the grasslands examined. Species richness and forager density in the other species decreased with increasing density of L. paralienus in fragments but not in control plots. The overall forager density of the other species was positively related to their habitat niche overlap with L. paralienus. The density of foragers of the other species at sugar resources was not affected by L. paralienus forager density. The experimental fragmentation resulted in an increase in natural sugar resources in fragments. This may have reduced the intensity of interspecific competition for sugar resources. Our study shows that the grassland fragmentation altered interactions between the dominant L. paralienus and the other ant species.Electronic Supplementary Material Supplementary material is available for this article at     

Quality of remnant indigenous grassland linkages for adult butterflies (Lepidoptera) in an afforested African landscape

Retention of interconnected, remnant grassland linkages is proposed here to reduce the adverse effects of alien pine afforestation in Afromontane grasslands. Adult butterflies were sampled at 38 grassland sites, representing increasing levels of disturbance both within the afforested area and outside it. Butterfly species richness and abundance in the lesser disturbed grassland remnants within the afforested area were similar to those of the surrounding natural grasslands. In contrast, butterfly species richness, but not necessarily abundance, decreased significantly in the highly disturbed sites, both in the grassland linkages and outside. Although some highly disturbed sites were relatively rich in species, most were visited by geographically widespread and vagile species. In contrast, wide, relatively undisturbed grassland linkages, as well as grasslands outside, were important for localised, sedentary and local endemic butterfly species. Nectar plants, especially the alien Verbena bonariensis, were the most significant variable explaining local butterfly distribution. In addition, tall grasses, hills, topographical landmarks, thermoregulatory sites, shelter and water features were also vital for particular species. It did not matter how deep the grassland linkages were situated inside the afforested area, as long as they were made up of good habitat. Retention of wide, quality grassland linkages are a way forward to maximise biodiversity alongside agroforestry.     

Avian Community Structure Along a Mountain Big Sagebrush Successional Gradient

Abstract: We compared vegetative structure and bird communities among 4 successional states in central Oregon representing a continuum from 1) postburn grassland, 2) mountain big sagebrush—Idaho fescue (Artemisia tridentatA—Festuca idahoensis) shrub—steppe, 3) sagebrush—steppe—juniper (Juniperus occidentalis), to 4) old-growth western juniper. Species richness, evenness, and diversity of bird communities were highest in old-growth and mid-successional juniper (22.9 species/transect and 23.6 species/transect, respectively) but lowest in the grasslands (17.6 species/transect). Bird species diversity was positively correlated with physiognomic cover diversity (r = 0.74, P = 0.001). Density of breeding birds was greatest in old-growth juniper (6.6 birds/ha) and lowest in postburn grasslands (3.6 birds/ha) but similar in shrub—steppe and sagebrush—steppe—juniper (6.0 birds/ha and 5.5 birds/ha, respectively). Old-growth juniper had the highest total densities of both tree and cavity nesters. Mountain chickadees (Parus gambeli), Cassin's finches (Carpodacus cassinii), chipping sparrows (Spizella passerina), brown-headed cowbirds (Molothrus ater), mountain bluebirds (Sialia currucoides), dark-eyed juncos (Junco hyemalis), Empidonax flycatchers, ash-throated flycatchers (Myiarchus cinerascens), and northern flickers (Colaptes auratus) were more abundant in cover types dominated by junipers. Vesper sparrows (Pooecetes gramineus), western meadowlarks (Sturnella neglecta), green-tailed towhees (Pipilo chlorurus), and horned larks (Eremophila alpestris) were associated with grassland communities. Brewer's sparrows (Spizella breweri), sage sparrows (Amphispiza belli), sage thrashers (Oreoscoptes montanus), and horned larks (Eremophila alpestris) were most abundant in sagebrush cover types. Management strategies should restore or maintain the desired proportions of the different successional states to maintain populations of grassland and sagebrush birds while providing habitat for tree and cavity nesting species.     

Species and functional trait re-assembly of ground beetle communities in restored grasslands

Ecosystem restoration provides unique opportunities to study community dynamics under succession and can reveal how consumer communities re-assemble and respond to successional changes. Studying community dynamics from both taxonomic and functional trait perspectives also may provide more robust assessments of restoration progress or success and allow cross-system comparisons. We studied ground beetle (Coleoptera: Carabidae) communities for three years in a restored grassland chronosequence with sites from 0 to 28 years old. We measured traditional community metrics (abundance, richness, Shannon diversity) and functional trait metrics based on species’ body length, wing morphology, activity time, phenology, and diet. Communities had high species richness and abundance in early successional stages, but these declined in later stages to low levels comparable to an adjacent grassland remnant. Species composition also shifted with time, converging with the remnant. Although functional richness, like species richness, declined as succession progressed, functional divergence quickly increased and was maintained over time, suggesting niche differentiation in established communities. Young sites were typified by small, macropterous, phytophagous species, while older sites contained larger species more likely to be flightless and carnivorous. Prescribed burns also affected traits, decreasing prevalence of larger species. This study demonstrates that functionally diverse consumer communities can self-assemble under restoration practices. In a relatively short amount of time both morphological and trophic level diversity are established. However, prescribed fire intended to control non-desirable plants may also shape beetle community functional composition, and restoration managers should consider if plant community benefits of fire outweigh potential declines in consumer function.     

Diversity of plant assemblages in isolated depressional wetlands from Central-Western Europe

A closed depression is defined as a geomorphologic element where a sediment depository is encircled by hillslopes. Despite the fact that closed depressions are often the only stagnant water points in many European landscapes, few ecological researches on their plant assemblages have been done. The main goal of this study was to give first results of the environmental factors responsible for the vegetation composition, richness and rarity in the closed depressions of the Lorraine biogeographical district (Belgium, France and grand-duché de Luxembourg). We surveyed for plant presence 85 forest and 77 grassland closed depressions. For each site, wetland area, local environmental factors and regional connectivities registered. For each species, the Ellenberg values were compiled. To investigate the main source of variation in species composition and in species richness (including richness in rare species), Non-metric Multidimensional Scaling analyses and Generalized Linear Models were respectively used. Species pools in forest (forest and preforest species) and in grassland (bog plants, pioneers, helophytes) were quite different. In both landscapes, a gradient from plants typical of basic high-productive soils to plants typical of acid low-productive soils reflects a shared successional gradient. The accumulation of organic matter allowed the establishment of Sphagnum spp., which slowly acidified the soil and thus acted as ecosystem engineers for the arrival of bog plants. Moreover, the species composition was additionally driven by the plant light tolerance in forests and by the plant water requirements in grasslands. Mechanisms of species accumulation (increase in species richness) were different in forests and in grasslands: respectively related to the plant light tolerance and to the wetland area. At the regional level, the averaged soil productivity was negatively related to the richness in rare species. Indeed, perennial highly-competitive plants such as Glyceria spp., Iris pseudacorus or Urtica dioica impeded the establishment of smaller and rarer species. At the habitat level, isolated closed depressions (due to either low connectivity or low grazing pressure) have more habitat rare species, giving evidences of dispersal limitation in plant assemblages of closed depressions.     

Landscape composition and habitat area affects butterfly species richness in semi-natural grasslands

During the last 50 years, the distribution and abundance of many European butterfly species associated with semi-natural grasslands have declined. This may be the result of deteriorating habitat quality, but habitat loss, resulting in decreasing area and increasing isolation of remaining habitat, is also predicted to result in reduced species richness. To investigate the effects of habitat loss on species richness, we surveyed butterflies in semi-natural grasslands of similar quality and structure, but situated in landscapes of different habitat composition. Using spatially explicit habitat data, we selected one large (6–10 ha) and one small (0.5–2 ha) grassland site (pasture) in each of 24 non-overlapping 28.2 km² landscapes belonging to three categories differing in the proportion of the area that consisted of semi-natural grasslands. After controlling for local habitat quality, species richness was higher in grassland sites situated in landscapes consisting of a high proportion of grasslands. Species richness was also higher in larger grassland sites, and this effect was more pronounced for sedentary than for mobile species. However, the number of species for a given area did not differ between large and small grasslands. There was also a significant relationship between butterfly species richness and habitat quality in the form of vegetation height and abundance of flowers. In contrast, butterfly density was not related to landscape composition or grassland size. When species respond differently to habitat area or landscape composition this leads to effects on community structure, and nestedness analysis showed that depauperate communities were subsets of richer ones. Both grassland area and landscape composition may have contributed to this pattern, implying that small habitat fragments and landscapes with low proportions of habitat are both likely to mainly contain common generalist species. Based on these results, conservation efforts should aim at preserving landscapes with high proportions of the focal habitat.     

Enhancement of late successional plants on ex-arable land by soil inoculations

Restoration of species-rich grasslands on ex-arable land can help the conservation of biodiversity but faces three big challenges: absence of target plant propagules, high residual soil fertility and restoration of soil communities. Seed additions and top soil removal can solve some of these constraints, but restoring beneficial biotic soil conditions remains a challenge. Here we test the hypotheses that inoculation of soil from late secondary succession grasslands in arable receptor soil enhances performance of late successional plants, especially after top soil removal but pending on the added dose. To test this we grew mixtures of late successional plants in arable top (organic) soil or in underlying mineral soil mixed with donor soil in small or large proportions. Donor soils were collected from different grasslands that had been under restoration for 5 to 41 years, or from semi-natural grassland that has not been used intensively. Donor soil addition, especially when collected from older restoration sites, increased plant community biomass without altering its evenness. In contrast, addition of soil from semi-natural grassland promoted plant community evenness, and hence its diversity, but reduced community biomass. Effects of donor soil additions were stronger in mineral than in organic soil and larger with bigger proportions added. The variation in plant community composition was explained best by the abundances of nematodes, ergosterol concentration and soil pH. We show that in controlled conditions inoculation of soil from secondary succession grassland into ex-arable land can strongly promote target plant species, and that the role of soil biota in promoting target plant species is greatest when added after top soil removal. Together our results point out that transplantation of later secondary succession soil can promote grassland restoration on ex-arable land.     

Flower resource and land management drives hoverfly communities and bee abundance in seminatural and agricultural grasslands

Pollination is a key ecosystem service, and appropriate management, particularly in agricultural systems, is essential to maintain a diversity of pollinator guilds. However, management recommendations frequently focus on maintaining plant communities, with the assumption that associated invertebrate populations will be sustained. We tested whether plant community, flower resources, and soil moisture would influence hoverfly (Syrphidae) abundance and species richness in floristically‐rich seminatural and floristically impoverished agricultural grassland communities in Wales (U.K.) and compared these to two Hymenoptera genera, Bombus, and Lasioglossum. Interactions between environmental variables were tested using generalized linear modeling, and hoverfly community composition examined using canonical correspondence analysis. There was no difference in hoverfly abundance, species richness, or bee abundance, between grassland types. There was a positive association between hoverfly abundance, species richness, and flower abundance in unimproved grasslands. However, this was not evident in agriculturally improved grassland, possibly reflecting intrinsically low flower resource in these habitats, or the presence of plant species with low or relatively inaccessible nectar resources. There was no association between soil moisture content and hoverfly abundance or species richness. Hoverfly community composition was influenced by agricultural improvement and the amount of flower resource. Hoverfly species with semiaquatic larvae were associated with both seminatural and agricultural wet grasslands, possibly because of localized larval habitat. Despite the absence of differences in hoverfly abundance and species richness, distinct hoverfly communities are associated with marshy grasslands, agriculturally improved marshy grasslands, and unimproved dry grasslands, but not with improved dry grasslands. Grassland plant community cannot be used as a proxy for pollinator community. Management of grasslands should aim to maximize the pollinator feeding resource, as well as maintain plant communities. Retaining waterlogged ground may enhance the number of hoverflies with semiaquatic larvae.     

Rapid genetic restoration of a keystone species exhibiting delayed demographic response

Genetic founder effects are often expected when animals colonize restored habitat in fragmented landscapes, but empirical data on genetic responses to restoration are limited. We examined the genetic response of banner‐tailed kangaroo rats (Dipodomys spectabilis) to landscape‐scale grassland restoration in the Chihuahuan Desert of New Mexico, USA. Dipodomys spectabilis is a grassland specialist and keystone species. At sites treated with herbicide to remove shrubs, colonization by D. spectabilis is slow and populations persist at low density for ≥10 years (≥6 generations). Persistence at low density and low gene flow may cause strong founder effects. We compared genetic structure of D. spectabilis populations between treated sites and remnant grasslands, and we examined how the genetic response to restoration depended on treatment age, area, and connectivity to source populations. Allelic richness and heterozygosity were similar between treated sites and remnant grasslands. Allelic richness at treated sites was greatest early in the restoration trajectory, and genetic divergence did not differ between recently colonized and established populations. These results indicated that founder effects during colonization of treated sites were weak or absent. Moreover, our results suggested founder effects were not mitigated by treatment area or connectivity. Dispersal is negatively density‐dependent in D. spectabilis, and we hypothesize that high gene flow may occur early in the restoration trajectory when density is low. Our study shows genetic diversity can be recovered more rapidly than demographic components of populations after habitat restoration and that founder effects are not inevitable for animals colonizing restored habitat in fragmented landscapes.