Effects of Restoration on Plant Species Richness and Composition in Scandinavian Semi-Natural Grasslands

Plant species richness in rural landscapes of northern Europe has been positively influenced by traditional management for millennia. Owing to abandonment of these practices, the number of species‐rich semi‐natural grasslands has decreased, and remaining habitats suffer from deterioration, fragmentation, and plant species decline. To prevent further extinctions, restoration efforts have increased during the last decades, by reintroducing grazing in former semi‐natural grasslands. To assess the ecological factors that might influence the outcome of such restorations, we made a survey of semi‐natural grasslands in Sweden that have been restored during the last decade. We investigated how plant species richness, species density, species composition, and abundance of 10 species that are indicators of grazing are affected by (1) the size of the restored site, (2) the time between abandonment of grazing and restoration, (3) the time elapsed since restoration, and (4) the abundance of trees and shrubs at the restored site. Only two factors, abundance of trees and shrubs and time since restoration, were positively associated with total species richness and species density per meter square at restored sites. Variation in species composition among restored sites was not related to any of the investigated factors. Species composition was relatively similar among sites, except in mesic/wet grasslands. The investigated factors had small effects on the abundance of the grazing‐indicator species. Only Campanula rotundifolia responded to restoration with increasing abundance and may thus be a suitable indicator of improved habitat quality. In conclusion, positive effects on species richness may appear relatively soon after restoration, but rare, short‐lived species are still absent. Therefore, remnant populations in surrounding areas may be important in fully recreating former species richness and composition.     

Effects of Connectivity and Regional Dynamics on Restoration of Small Mammal Communities in Midwestern Grasslands

Grasslands are among the most imperiled North American ecosystems. State Acres for Wildlife Enhancement (SAFE) is a national conservation program that converts agricultural fields into grasslands mainly to improve habitat for high priority wildlife species. To provide a broader assessment of the contribution of the SAFE program to biodiversity in the Midwest region of North America, we evaluated local and landscape constraints to restoration of small mammal communities. We livetrapped small mammals during three summers (2009–2011) on plots that were recently seeded, seeded 1–4 years prior to sampling, or established references (>10 years old). Restoration trajectories for small mammal communities included a shift over time from dominance by the habitat generalist Peromyscus maniculatus (deer mouse) to communities dominated by grassland Microtus species (prairie voles and meadow voles). Vole abundance during the first year following restoration depended on spatial connectivity provided by linear habitats (roadside ditches and grass waterways) within 300 m of the restored grassland. Patch size and seeding type (cool‐season versus warm‐season grasses) were not predictors of early restoration success. In 2011, voles experienced a severe regional decline consistent with multi‐year population cycles. During the crash, most remaining voles occurred on restored SAFE grasslands, but not on established grasslands. This surprising outcome suggests young restoration plots could function as refuges for voles during population declines in agricultural landscapes.     

Resilience and restoration of tropical and subtropical grasslands,savannas, and grassy woodlands

Despite growing recognition of the conservation values of grassy biomes, our understanding of how to maintain and restore biodiverse tropical grasslands (including savannas and open‐canopy grassy woodlands) remains limited. To incorporate grasslands into large‐scale restoration efforts, we synthesised existing ecological knowledge of tropical grassland resilience and approaches to plant community restoration. Tropical grassland plant communities are resilient to, and often dependent on, the endogenous disturbances with which they evolved – frequent fires and native megafaunal herbivory. In stark contrast, tropical grasslands are extremely vulnerable to human‐caused exogenous disturbances, particularly those that alter soils and destroy belowground biomass (e.g. tillage agriculture, surface mining); tropical grassland restoration after severe soil disturbances is expensive and rarely achieves management targets. Where grasslands have been degraded by altered disturbance regimes (e.g. fire exclusion), exotic plant invasions, or afforestation, restoration efforts can recreate vegetation structure (i.e. historical tree density and herbaceous ground cover), but species‐diverse plant communities, including endemic species, are slow to recover. Complicating plant‐community restoration efforts, many tropical grassland species, particularly those that invest in underground storage organs, are difficult to propagate and re‐establish. To guide restoration decisions, we draw on the old‐growth grassland concept, the novel ecosystem concept, and theory regarding tree cover along resource gradients in savannas to propose a conceptual framework that classifies tropical grasslands into three broad ecosystem states. These states are: (1) old‐growth grasslands (i.e. ancient, biodiverse grassy ecosystems), where management should focus on the maintenance of disturbance regimes; (2) hybrid grasslands, where restoration should emphasise a return towards the old‐growth state; and (3) novel ecosystems, where the magnitude of environmental change (i.e. a shift to an alternative ecosystem state) or the socioecological context preclude a return to historical conditions.     

Restoration of Species‐Rich,Nutrient‐Limited Mountain Grassland by Mowing and Fertilization

Mowing and management to reduce nutrient levels have often been successfully used to restore species‐rich grasslands in various parts of Europe. However, such treatments have failed to restore the species‐rich Central European mountain grasslands dominated by Polygonum bistorta. P. bistorta builds an extensive underground rhizome system that monopolizes available nutrients in these nutrient‐poor grasslands, enabling this species to persist at high densities even in the presence of mowing. Therefore, we tested a restoration approach using a factorial combination of fertilization and mowing, as well as a litter removal treatment. The experiment was run over 5 years and species composition response to these treatments was recorded at two spatial scales. Mowing suppressed flowering and cover of P. bistorta and promoted target grassland species and richness. Fertilization prevented nutrient impoverishment and increased height and dominance of the broad‐leaved grasses with which many species‐rich grassland herbs could coexist. The additive effect of the mowing/fertilization treatments was strong enough to act as a driver of P. bistorta suppression and associated community change. The litter removal treatment, however, had little effect on plant composition. The experiment demonstrates that in nutrient‐limited grasslands, increasing nutrient levels in addition to mowing to manage competition for light can be used to control dominants. This contrasts with restoration of systems where after abandonment increased nutrient levels lead to the establishment of tall dominants that suppress other species by competition for light.     

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.     

Nutrient limitation in species‐rich Calthion grasslands in relation to opportunities for restoration in a peat meadow landscape

Questions: Which nutrient(s) limit(s) vegetation productivity in Calthion grasslands? Is phosphorus release a bottleneck for restoration of species‐rich Calthion grasslands on rewetted dairy meadows? Location: Three species‐rich Calthion grasslands in the Western Peat District in the Netherlands. Methods: We conducted a field fertilization experiment with nitrogen (N), phosphorus (P) and potassium (K) in three existing Calthion grasslands to evaluate the potential for restoration on rewetted dairy meadows. Responses of above‐ground biomass, tissue nutrient concentrations and nutrient ratios were determined after 2 yr of fertilization. Results: Biomass increased with fertilization with N‐only and K‐only but did not react to P‐only additions. Comparisons of tissue nutrient concentrations and nutrient ratios also gave indications of N and K limitation. Conclusions: The strong P release expected after rewetting should not necessarily interfere with restoration of Calthion communities on rewetted dairy meadows. It is concluded that for successful restoration management measures should focus on reducing N and/or K availability. Potassium might be an overlooked bottleneck in the restoration of species‐rich grasslands.     

The early response of subtropical tussock grasslands to restoration treatments

Once widespread, Australia's bluegrass tussock grasslands (dominated by Dichanthium sp.) of the Queensland Central Highlands are now severely endangered. Despite being biodiversity rich and highly valued as low input, nutrient‐dense grazing systems, bluegrass tussock grasslands have suffered extensive clearing and degradation over the last 150 years. Natural recovery of these grasslands is possible but rates of recovery are slow. As such, there is an urgent need to assess practical management strategies to accelerate recovery of these grasslands, with a particular focus on early‐successional stages, when aggressive exotic species are most prevalent. To date, no studies have tested whether commonly used grassland restoration strategies can enhance early‐successional stages and accelerate regeneration in this system. Here, we examine the early short‐term impacts (first two seasons) of two common grassland restoration approaches, with two variations each: direct seeding (single species and low seed diversity) and vegetation clearing (prescribed burning and glyphosate application) across two common starting points: a formerly cropped old field and a historically overgrazed natural grassland. No treatment increased native diversity (Shannon's or richness) in plots but the composition of burned plots in the old field did become more similar to healthy reference sites after two seasons. Burning combined with direct seeding also increased the abundance of the dominant grass, Dichanthium sericeum, toward healthy reference levels within the first two seasons post seeding. This study provides a practical assessment of the short‐term impacts and capacity of common grassland restoration treatments to enhance the recovery of Australia's tussock grassland systems.     

Large‐Scale Restoration of Dry Grasslands on Ex‐Arable Land Using a Regional Seed Mixture: Establishment of Target Species

Large‐scale (circa 500 ha) restoration of species‐rich dry grasslands was conducted using a high‐diversity regional seed mixture in the White Carpathians Protected Landscape Area and Biosphere Reserve, Czech Republic, Central Europe. After sowing, the restored grasslands were regularly mown. Vegetation was analyzed at sites restored 1–12 years ago and compared with that of ancient, extremely species‐rich grasslands nearby. Nearly all (98%) sown target species successfully established and nearly half of unsown target species established spontaneously, partly dependent on distance to the ancient grasslands. Early mowing in the first half of June appeared to support species diversity and broad‐leaved forbs at the expense of competitive grasses. Using a regional seed mixture appeared to be an effective way of restoring dry grasslands and is especially recommended in the proximity of still existing ancient grasslands where spontaneous establishment of unsown target species may reinforce the success of restoration more easily.     

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.     

Near‐natural methods promote restoration of species‐rich grassland vegetation—revisiting a road verge trial after 9 years

The present loss of species‐rich grasslands makes it vital to restore these valuable habitat types, including novel habitat variants such as road verges. Due to the lack of knowledge on long‐term outcomes of restoration initiatives, well‐designed studies comparing different restoration methods are needed. In this study, we examined fine‐scale vegetation recovery patterns over 9 years in a field experiment with several near‐natural restoration methods (adding local seed mixtures, transferring hay from local grasslands using hard or light raking, and natural regeneration) in a road verge. We compared this to standard revegetation (hydroseeding species‐poor commercial seed mixtures). We found major temporal changes in vegetation restored by local seed or hay transfer, before it gradually became more similar to the donor grasslands and seed mixtures, which served as references for the experiment. Natural (spontaneous) regeneration with seed dispersal from surroundings gave similar results, whereas areas revegetated using standard methods became more dissimilar to the reference sites during the study period. The main variation in species composition reflected the contrast between local donor grasslands and seed mixtures and the species‐poor early successional grasslands. We conclude that near‐natural methods (hay transfer and seeding) successfully restored species‐rich grassland, including road verges. This study underlines the importance of comparing several treatments over a sufficiently long period to assess their success in restoring species‐rich grassland.     

Epiphytic lichens on Juniperus communis – an unexplored component of biodiversity in threatened alvar grassland

Semi‐natural grasslands, among them thin‐soil calcareous grasslands (alvars), have great conservation value but have become increasingly rare in Europe. The main threat to alvar grasslands is the encroachment by juniper Juniperus communis and therefore it is usually removed during the restoration practice. Juniper can also be a host plant for many epiphytic lichens, but its role as a phorophyte is poorly known. We studied epiphytic lichen diversity on 126 junipers in 17 sites in western Estonia and found 140 lichenized taxa including several rare and red‐listed species. Using indirect and direct multivariate analyses (DCA, pCCA) and general linear models we revealed that both habitat and phorophyte properties affect lichen assemblies on juniper. Lichen species richness per site showed a unimodal relationship with compound factors of site productivity and juniper characteristics (stem circumference and juniper width). Lichen species richness per phorophyte was increasing with its size and with the proportion of dead branches, and was twice higher in plate alvars than in ryhk alvars. Also, the species composition in plate alvars differed from ryhk alvars by having 42 characteristic lichen species in plate alvars vs three indicators of ryhk alvars. The composition of lichens was significantly influenced by encroachment of alvars, e.g. by high juniper cover and shrub layer height, as well as by the proportion of dead branches and stem circumference of juniper. We conclude that the epiphytic lichen assemblies on junipers are threatened by grassland encroachment similarly to ground layer lichen assemblies. We suggest that some old and scencent junipers should be preserved during the restoration of alvar grasslands.     

Passive restoration contributes to bird conservation in Brazilian Pampa grasslands

The global decline of biodiversity makes it important to find affordable ways to conserve and restore habitats. Restoration is useful for conserving native grasslands, with passive restoration defined as either natural colonization or unassisted recovery. Grasslands in southeastern South America have been transformed into croplands and impacted by other human activities. We describe the first assessment of passive restoration as a management tool to conserve birds in the Pampa grasslands of Brazil. We compared bird species richness using coverage‐based rarefaction and extrapolation, applying PERMANOVA for composition, and the abundance of bird communities between sites undergoing passive restoration (PR) and sites with native grasslands (NG). We employed fitted generalized linear mixed models (GLMM) to quantify relationships between bird occurrence and vegetation structure and cover. We recorded 61 species of birds during our study (45 in PR and 46 in NG) and 762 individuals (333 in PR and 429 in NG). Of these species, 15 were restricted to PR and 16 to NG. Grassland specialists and threatened species were found in both PR and NG, and only vegetation height differed between PR and NG. We detected eight species of conservation concern, including three recorded only in PR, three only in NG, and two in both PR and NG. The absence of marked differences in species richness and composition of bird communities between passive‐restoration and native grasslands in our study suggests that grasslands in the process of passive restoration can provide habitat for many species of grassland birds and that passive restoration is an appropriate management tool for biodiversity conservation in Brazilian grasslands.     

Problems, Approaches, and Results in Restoration of Dutch Calcareous Grassland During the Last 30 Years

This paper is based on research of the restoration of species‐rich calcareous grasslands in The Netherlands, over the last 30 years. Chalk grassland is a semi‐natural vegetation with a high density of species at a small scale. This type of vegetation was once widespread in Western Europe as common grazing land, mainly for flocks of sheep for which the main function was dung production. In some regions of Central Europe, these grasslands were also used for hay production. The dung was used to maintain arable field production at a reasonable level. In the chalk district in the southernmost part of The Netherlands some 25 sites of this vegetation, varying in area from 0.05–4.5 ha, are still present. Chalk grassland completely lost its significance for modern agricultural production after the wide application of artificial fertilizer following World War II. This grassland has a high conservation value both for plants and animal species, of which a large number of species are exclusively restricted to this biotope. When conservation activities started at a large scale in the early 1960s, three different types of restoration activities could be distinguished: (1) restoration of fertilized sites; (2) restoration of abandoned grasslands; and (3) recreation of chalk grassland on former arable fields. The main aim of the restoration attempt is to create and/or improve sustainable conditions for both plant and animal species characteristic of the chalk grassland ecosystem. In the process of restoration, several phases of different activities can be distinguished: (1) pre‐restoration phase, during which information of the land use history is collected and, based on these data, clear restoration goals are established; (2) initial restoration phase, during which effects of former, non‐conservational land use has to be undone in order to stimulate germination and establishment of target species originating from soil seed bank and species pool; (3) consolidation phase, including the introduction and continuation of a regular management system for sustainable conservation; and (4) long‐term conservation strategy, including measures to prevent disturbance from the outside and genetic erosion and extinction of locally endangered plant populations.     

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.     

Local grassland restoration affects insect communities

1. It is hypothesised that ecological restoration in grasslands can induce an alternative stable state shift in vegetation. The change in vegetation influences insect community assemblages and allows for greater functional redundancy in pollination and refuge for native insect species. 2. Insect community assemblages at eight coastal California grassland sites were evaluated. Half of these sites had undergone restoration through active revegetation of native grassland flora and half were non‐restored. Insects were collected from Lupinus bicolor (Fabaceae) within 2 × 2‐m² plots in spring 2017. Lupinus bicolor is a common native species that is used in California restoration projects, and home and state landscaping projects. 3. Ordination demonstrated that insect community assemblages were different between restored and non‐restored sites. These differences were seen in insect functional groups as well as taxa‐specific differences and were found to be driven by environmental characteristics such as non‐native forb cover. 4. Functional redundancy of herbivores decreased at restored sites, while pollinators became more redundant compared with non‐restored sites. The assemblages of the common species found at restoration sites contained more native insects than those found at non‐restored sites, including species such as Bombus vosnesenskii. 5. Local grassland restoration has the potential to induce an alternative stable state change and affect insect community assemblages. Additionally, it was found that grassland restoration can be a potential conservation tool to provide refugia for bumblebees (Bombus), but additional studies are required to fully understand its broader applicability.     

Success of active restoration in grasslands: a case study of birds in southern Brazil

Grasslands in southeastern South America have been extensively converted to various land uses such as agriculture, threatening regional biodiversity. Active restoration has been viewed as a management alternative for recovery of degraded areas worldwide, although most studies are conducted in forests and none has evaluated the effect of active restoration of grasslands in southeastern South America. From 2015 through 2017 we monitored a federally owned tract of grassland from the beginning of the active‐restoration process. We compared the bird community in this active‐restoration area (AR) with a reference area (NG) in Pampa grasslands in southern Brazil. We sampled birds by point counts and surveyed vegetation structure in plots. Over the 3 years of active restoration, bird species richness and abundance were higher in AR (30 species, 171 individuals) than NG (22 species, 154 individuals). The species composition also differed between the two habitats. Grassland bird species were present in both AR and NG. The vegetation structure differed between AR and NG in five attributes: height, short and tall grasses, herbs, and shrubs. Since it has been found that active restoration is useful in promoting species diversity, we encourage studies of the use of long‐term restoration efforts. Our study, even on a local scale, showed a rapid recovery of the bird community in the active‐restoration compared to native grassland, and suggests the potential for recovery of the degraded grasslands of the Brazilian Pampa biome.     

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.     

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.     

Recovery of Native Plant Community Characteristics on a Chronosequence of Restored Prairies Seeded into Pastures in West‐Central Iowa

Restored grasslands comprise an ever‐increasing proportion of grasslands in North America and elsewhere. However, floristic studies of restored grasslands indicate that our ability to restore plant communities is limited. Our goal was to assess the effectiveness of restoration seeding for recovery of key plant community components on former exotic, cool‐season pastures using a chronosequence of six restoration sites and three nearby remnant tallgrass prairie sites in West‐Central Iowa. We assessed trends in Simpson's diversity and evenness, richness and abundance of selected native and exotic plant guilds, and mean coefficient of conservatism (mean C). Simpson's diversity and evenness and perennial invasive species abundance all declined with restoration site age. As a group, restoration sites had greater richness of native C₃ species with late phenology, but lower richness and abundance of species with early phenology relative to remnant sites. Total native richness, total native abundance (cover), mean C, and abundance of late phenology C₃ plants were similar between restoration and remnant sites. Observed declines in diversity and evenness with restoration age reflect increases in C₄ grass abundance rather than absolute decreases in the abundance of perennial C₃ species. In contrast to other studies, restoration seeding appears to have led to successful establishment of tallgrass prairie species that were likely to be included in seeding mixtures. While several floristic measures indicate convergence of restoration and remnant sites, biodiversity may be further enhanced by including early phenology species in seeding mixes in proportion to their abundance on remnant prairies.     

Long‐term experiment manipulating community assembly results in favorable restoration outcomes for invaded prairies

Invasive species are a common problem in restoration projects. Manipulating soil fertility and species arrival order has the potential to lower their abundance and achieve higher abundances of seeded native species. In a 7‐year experiment in Missouri, United States, we tested how nutrient addition and the timing of arrival of the invasive legume Lespedeza cuneata and seeded native prairie grass and forb species influenced overall community composition. Treatments that involved early arrival of seeded forb and grass species and late arrival of L. cuneata were most successful at creating community structure that fulfilled our restoration goals, displaying high abundance of seeded native forb species, low abundances of L. cuneata, and non‐native species. There were few treatment interactions, with the exception that timing seeded native forbs and timing of L. cuneata arrival interactively influenced the abundance of seeded native forbs. This suggests that the individual treatments are supporting the restoration goals, such as creating a community with low abundance of L. cuneate or high abundance of native seeded species, without restricting each other. This study demonstrates the importance of priority effects in disturbed habitats prone to invasion, the lasting effects of initial seeding on long‐term community composition, and the potential for fertilization to positively benefit restoration of degraded grasslands.