Congruence among vascular plants and butterflies in the evaluation of grassland restoration success

Using surrogate taxa in the monitoring of restoration success is strongly advocated but currently under debate. Are butterfly and vascular plant communities good surrogates for each other in monitoring the restoration of calcareous grassland grassland? The research was conducted in a small area in Tuscany (central Italy), which was the focus of a restoration project. We sampled vascular plants using 35 plots (1 m²) and butterflies using 20 transects (100 m in length) belonging to four treatment types: control scrubland, restored ex-arable, restored grassland and reference grassland. A mixed model nested ANOVA showed that reference grassland, 3 years after restoration, had the highest number of vascular plant species and the highest number of butterfly species. The different treatments were found to be distinct from a compositional point of view: pRDA showed that the “treatment” factor, independent of spatial variables, accounted for 22.1 and 21.9% of the total variance in the plant community and butterfly community, respectively (p < 0.01). The butterfly community was significantly correlated with the entire plant community data set and with the phanerophyte community (i.e. shrubs); larval host plants were significantly correlated with the composition of non-migratory butterflies. The results indicated that butterfly and vascular plant communities were congruent, suggesting the use of community composition of different taxa as a valuable tool for the evaluation of management success.     

The seed banks of English lowland calcareous grasslands along a restoration chronosequence

We investigated whether the seed banks of ex-arable lowland calcareous grasslands underwent restoration similar to that of the above-ground restoration, and whether this was influenced by seed-sowing or environmental conditions. We compared 40 sites, where some form of restoration work had been implemented between 2 and 60 years previously, with 40 paired reference sites of good quality calcareous grassland with no history of ploughing or agricultural improvement. We analysed differences between sites and between above- and below-ground vegetation using both a multivariate approach and proportions of selected plant attributes. Seed banks of reference sites were more characteristic of late successional communities, with attributes such as stress tolerance, perenniality and a reliance on fruit as the germinule form more abundant than in restoration sites. In restoration sites, these tended to decrease with restoration site isolation and increase with restoration site age and where soil nutrient conditions were more similar to reference sites (i.e. with relatively low phosphorus and high nitrogen). Seed bank communities of all sites differed considerably from above-ground communities, however, and no overall significant responses to site age, isolation or soil nutrients were detected by multivariate analyses of similarity of species between pairs of sites. Responses to different seeding methods were also barely detectable. While there is some indication from the plant attribute data that the regeneration potential contained in the seed banks of restored sites increasingly resembles that of references sites over time, even seed banks of good quality calcareous grassland are dominated by ruderal species. It is likely, therefore, that permanent seed banks do not facilitate the restoration of ex-arable grasslands.     

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 New Approach for Tracking Vegetation Change after Restoration: A Case Study with Peatlands

Developing objective tools for tracking progress of restored sites is of general concern. Here, we present an innovative approach based on principal response curves (PRC) and species classification according to their preferential habitats to monitor changes in community composition. Following large‐scale restoration of a cut‐over peatland, vegetation was surveyed biannually over 8 years. We evaluated whether the establishing plant communities fell within the range of natural variation. We used both general diversity curves and PRC applied on plant species grouped by preferred habitat to compare restored sites and unrestored sites to a reference ecosystem. After 8 years, diversity and richness differed between the sites, with Forest and Ruderal species more prominent in unrestored sites, and Peatland, Forest, and Wetland species dominant in restored sites. The PRC revealed that the restored site became rapidly dominated by typical peatland plants, the main drivers of temporal changes being Sphagnum rubellum, Pohlia nutans, and Mylia anomala. Some differences remained between the restored and the undisturbed species pools: the former had more herbaceous species associated with wetlands such as Calamagrostis canadensis and Typha latifolia and the latter had more forested species like Kalmia angustifolia throughout the study. PRC revealed to be an efficient tool identifying species driving changes at the community level after restoration. In our case study, examining PRC scores after classifying species according to their preferred habitat allowed to illustrate objectively how restoration promotes target species (associated to peatlands) and how lack of intervention benefits ruderal species.     

Aboveground environment type, soil nutrient content and arbuscular mycorrhizal fungi explain establishment success of Centaurea jacea on ex-arable land and in late-successional grasslands

We studied the relative importance of the aboveground and belowground environment for survival and growth of emerged seedlings of Centaurea jacea to better understand the general difficulty of establishing late-successional species at restoration sites on ex-arable land. Potted seedlings growing on soil from six late-successional grasslands and from six ex-arable (restoration) sites were reciprocally exchanged, and survival and relative growth rate of the seedlings monitored. In addition, we assessed aboveground herbivory and colonization of roots by arbuscular myccorhizal fungi of all plants, as well as nutrient availability, and microbial biomass and community composition using PLFA techniques in all twelve soils. Seedling survival was higher in restoration habitat and soil than in grassland habitat and soil, but growth did not differ between the aboveground and belowground environment types. Shoot growth rate was initially correlated with soil nutrient content, and later in the experiment with mycorrhizal colonization levels. Our results indicate that arbuscular mycorhizal fungi may be important for the successful establishment of C. jacea and that abiotic soil factors, like K availability and N:P ratio, can promote mycorrhizal colonization. Hence, the belowground environment should be considered when selecting sites for restoring species-rich grasslands.     

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.     

Spider community responses to grassland restoration: balancing trade‐offs between abundance and diversity

Spiders (Araneae) play key roles in ecosystems, not only as common and abundant generalist predators, but also as major contributors to biodiversity in many areas. In addition, due to their short generation times and high mobility, spiders respond rapidly to small changes in their environment, potentially making them useful indicators for restoration monitoring. However, few studies have focused on spider responses to grassland restoration in the United States. We compared degraded, native, and restored grassland sites to examine how spider communities and habitat respond to arid grassland restoration. We also examined how responses varied with the age of the restoration project. Spider communities in native sites differed from those in restored and degraded sites in several ways: native sites had fewer spiders and a different community composition than degraded and restored sites. However, native and restored sites had more species than degraded sites. Chronosequence data showed trends for lower abundance, higher species richness, and changing community composition as restoration projects mature. Several habitat variables were closely linked to variation in spider communities including cover of invasive annual grasses, litter, and biological soil crusts. Our data suggest that spider and vegetation responses to grassland restoration efforts can be successful in the long term—with resulting communities becoming more similar to native ones—and that spiders are useful indictors of grassland restoration. Our results also suggest that restoration may involve balancing trade‐offs between ecosystem services, with potential losses in predatory control offset by increases in biodiversity with restoration effort.     

Are Ants Useful Indicators of Restoration Success in Temperate Grasslands?

Assessments of restoration are usually made through vegetation community surveys, leaving much of the ecosystem underexamined. Invertebrates, and ants in particular, are good candidates for restoration evaluation because they are sensitive to environmental change and are particularly important in ecosystem functioning. The considerable resources currently employed in restoring calcareous grassland on ex‐arable land mean that it is important to gather as much information as possible on how ecosystems change through restoration. We compared ant communities from 40 ex‐arable sites where some form of restoration work had been implemented between 2 and 60 years previously, with 40 paired reference sites of good quality calcareous grassland with no history of improvement or cultivation. A total of 11 ant species were found, but only two of these were found to be significantly different in abundance between restoration and reference sites: Myrmica sabuleti was more likely to be present in reference sites, whereas Lasius niger was more likely to be found in restoration sites. Myrmica sabuleti abundance was significantly positively correlated with age of restoration sites. The potential number of ant species found in temperate grasslands is small, limiting the information their assemblages can provide about ecosystem change. However, M. sabuleti is a good indicator species for calcareous grassland restoration success and, alongside information from the plant community, could increase the confidence with which restoration success is judged. We found the survey to be quick and simple to carry out and recommend its use.     

Vegetation development after a large scale restoration of species-rich grasslands in a Central European floodplain

Large scale restoration using local high-diversity seed mixture combined with turf transfer was applied on ex-arable land in the Morava River floodplain in the western Slovakia in the years 1999–2012. The post-restoration vegetation development was recorded during 12 years after the restoration using floristic records per restored polygons with cover estimation in simple 3-degree scale. Temporal changes in species composition were evaluated by gradient analysis and number of characteristic grassland and ruderal species on restored sites was analysed by general linear models. Species composition changed gradually towards the species composition typical for species-rich floodplain grasslands, but the trajectory was not straightforward and several irregularities were observed. They were probably induced by extreme weather events (drought, floods). The decrease in ruderal species and increase in the number of typical floodplain grassland species were observed, when floodplain grassland species permanently outcompeted ruderal species since 8th year after the restoration. However the development in large scale was slower, than expected from previous small-scale experiments, it is evident, that combination of local seed mixture sowing with a turf transfer is a feasible method for the restoration of species-rich floodplain grasslands from arable land.     

Sowing different mixtures in dry acidic grassland produced priority effects of varying strength

Several longer-term assembly studies on ex-arable land have found that species that arrive first at a disturbed site can play a key role in the further development of the community and that this priority effect influences aboveground productivity, species diversity and stability of the grassland communities that develop. Restoration of nutrient poor, species rich grasslands is often limited by seed dispersal as well as the accessibility of suitable microsites for establishment. Sowing species (i.e. creating priority effects for further assembly) may help overcome such dispersal barriers, but the potential of using priority effects for restoration has not been tested in this type of dry grassland. We tested the hypothesis that sowing two different seed mixtures used for dry acidic grassland restoration onto a sandy substrate (which formed an equivalent to a primary succession) would create priority effects, and that these priority effects would be sustained over a number of years. We followed community assembly and measured aboveground productivity for four years after sowing. We found that priority effects caused by sowing of differently diverse mixtures did also occur in dry acidic grassland habitat, but that how persistent they were over time depended on the response variable considered. Priority effects on species number were not as strong as found in previous ex-arable land studies, whereas priority effects for aboveground productivity were still visible after 4 years. In addition, functional composition of the community still reflected the composition of the seed mixtures 4 years later. Our results suggest that priority effects can occur in nutrient-poor dry acidic grassland but in contrast to more nutrient-rich sites the breadth of responses affected may not be as wide.     

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.     

Strengths and potential pitfalls of hay transfer for ecological restoration revealed by RAD‐seq analysis in floodplain Arabis species

Achieving high intraspecific genetic diversity is a critical goal in ecological restoration as it increases the adaptive potential and long‐term resilience of populations. Thus, we investigated genetic diversity within and between pristine sites in a fossil floodplain and compared it to sites restored by hay transfer between 1997 and 2014. RAD‐seq genotyping revealed that the stenoecious floodplain species Arabis nemorensis is co‐occurring with individuals that, based on ploidy, ITS‐sequencing and morphology, probably belong to the close relative Arabis sagittata, which has a documented preference for dry calcareous grasslands but has not been reported in floodplain meadows. We show that hay transfer maintains genetic diversity for both species. Additionally, in A. sagittata, transfer from multiple genetically isolated pristine sites resulted in restored sites with increased diversity and admixed local genotypes. In A. nemorensis, transfer did not create novel admixture dynamics because genetic diversity between pristine sites was less differentiated. Thus, the effects of hay transfer on genetic diversity also depend on the genetic make‐up of the donor communities of each species, especially when local material is mixed. Our results demonstrate the efficiency of hay transfer for habitat restoration and emphasize the importance of prerestoration characterization of microgeographic patterns of intraspecific diversity of the community to guarantee that restoration practices reach their goal, that is maximize the adaptive potential of the entire restored plant community. Overlooking these patterns may alter the balance between species in the community. Additionally, our comparison of summary statistics obtained from de novo‐ and reference‐based RAD‐seq pipelines shows that the genomic impact of restoration can be reliably monitored in species lacking prior genomic knowledge.     

Establishment and persistence of target species in newly created calcareous grasslands on former arable fields

The effects of different restoration measures and management variants on the vegetation development of newly created calcareous grasslands were studied in southern Germany from 1993 to 2002. In 1993, fresh seed-containing hay from a nature reserve with ancient calcareous grasslands was transferred onto ex-arable fields with and without topsoil removal. Nine years after start of the restoration, the standing crop was lower and the cover of bare soil was higher on topsoil-removal sites than on sites without soil removal. Topsoil removal had a positive effect on the proportion of target species (class Festuco-Brometea), because the number and cover of productive meadow species (class Molinio-Arrhenatheretea) were reduced. Persistence of hay-transfer species and the number of newly colonizing target species were highest on topsoil-removal sites. On plots with and without soil removal, species richness and the number of target species increased quickly after hay transfer and were always higher on hay-transfer plots than on plots that had not received hay in 1993. In 2002, differences induced by hay transfer were still much more pronounced than differences between management regimes. Management by mowing, however, led to higher species richness, a greater number of target species and a lower number of ruderals in comparison to no management on restoration fields without soil removal. A detrended correspondence analysis (DCA) indicated that vegetation composition of the hay-transfer plots of the restoration fields still differed from the vegetation of ancient grasslands in the nature reserve. Vegetation of an ex-arable field in the nature reserve (last ploughed in 1959) showed an intermediate successional stage. In general our results indicate that the transfer of autochthonous hay is an efficient method for the restoration of species-rich vegetation, which allows not only quick establishment but also long-term persistence of target species.     

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.     

A six-year experimental restoration of biodiversity by shrub-clearing and grazing in calcareous grasslands of the French Prealps

The conservation of dry calcareous grasslands in the French Prealps strongly depends on the maintenance of low-intensity farming systems supported by agri-environmental schemes. An experimental assessment of the effect of current agro-pastoral management on the biodiversity of plant communities was conducted during a six-year permanent plot survey in four sites with contrasting habitat conditions (mesic to xeric). Analyses of species changes showed: (i) a strong increase in species richness and open grassland species frequencies four years after shrub-clearing, and (ii) a noticeable recovery of rare annuals and perennial species of conservation interest establishing in gaps created by grazing. At the community level, the restoration effect was evaluated by a between-year Correspondence Analysis, explaining 10.9% of the total floristic variability versus 29.5% for the site effect (between-site CA). Species ordination by between-year CA showed similar trajectories of vegetation changes during restoration despite different habitat conditions and grazing regimes between sites. The successful restoration of prealpine calcareous grasslands was explained by the availability of seed sources during the study in adjacent grazed or mown grasslands. Thus, restoration assessment should focus on dispersal possibilities and functional roles of species rather than species richness only. Finally, the spatial (i.e. the area of patches that need to be restored) and temporal (i.e. the frequency of shrub-clearing) implications for the large-scale conservation of prealpine calcareous grasslands by current agro-pastoral management are discussed.     

Post-Soviet recovery of grassland vegetation on abandoned fields in the forest steppe zone of Western Siberia

Following the collapse of the Soviet Union in 1991 around 45 million hectares of arable land became abandoned across Russia. Our study focused on the recovery potential and conservation value of grassland vegetation on ex-arable land in the Tyumen region of the Western Siberian grain belt. We compared ex-arable grasslands of different successional stages with ancient grasslands as reference for the final stage of succession along a climatic gradient from the pre-taiga to the forest steppe zone. Plant community composition and species richness of ex-arable land clearly developed towards reference sites over time, but even after 24 years of abandonment, the grassland vegetation had not totally recovered. The γ-diversity of vascular plants was slightly higher on ex-arable land than in ancient grasslands but the mean α-diversity was still moderately lower. A significant proportion of the vegetation of ex-arable land still consisted of ruderal and mesic grassland species and the number and cover of meadow-steppe species was significantly lower than in ancient grasslands. Grazing and time since abandonment positively affected the reestablishment of target grassland species, whereas it was negatively affected by the cover of grasses. In contrast to ex-arable land, the conservation value of arable land is only modest. Therefore, future intensification of land use is most likely less harmful if directed to existing arable land. Re-cultivation of ex-arable land and grassland improvement operations such as seeding of competitive grass species are major threats for the biodiversity of secondary grasslands on ex-arable land in the forest steppe zone of Western Siberia.     

Response of plant functional traits during the restoration of calcareous grasslands from forest stands

In this survey, we studied the response of plant functional traits to calcareous grassland restoration in the Calestienne region, Southern Belgium (restoration protocol: forest clear-cutting followed by grazing at all sites). We considered traits related to dispersal, establishment, and persistence that integrate the main challenges of plants to re-establish and survive in restored areas. Functional traits were compiled from databases and compared among (i) pre-restoration and young restoration forests; (ii) restoration areas of different ages; and (iii) old restorations and reference grasslands. The following questions were addressed: (i) What is the early response (2–4 years) in terms of plant functional trait following one restorative clear-cut event? (ii) What plants functional trait responses occur from restorative management (i.e., sheep and goat grazing)? (iii) Which differences still persist between the oldest restored parcels (10–15 years), and the historical reference grasslands? Forest clear-cuts induced several changes among functional traits, including decreased mean seed mass and certain vegetative traits (i.e., decreased phanerophytes, branching species; and increased short lifespan species i.e., annuals and biennials). During restorative management, clonal, epizoochorous and autumn germinating species were favored. Despite numerous other changes during this phase, many differences remained compared to reference grasslands. In particular, geophytes, mycorrhizal and evergreen species abundance were not approaching reference grassland values. The observed pattern helped to draw inferences on the possible mechanisms operating under vegetation recovery following restorative forest clear-cut and subsequent management were identified and described in this study. Results indicated grazing was an important factor, which increased epizoochorous species, and autumn germinating taxa that filled niches in vegetation opened by summer grazing animals. Finally, differences between old restoration and reference grasslands emphasized that management should focus on reduction in soil fertility, and geophyte rhizomatous grasses. Long-term monitoring is vital to assess if management plans are effective in the complete restoration of species functional trait assemblages.     

Soil Organism and Plant Introductions in Restoration of Species-Rich Grassland Communities

Soil organisms can strongly affect competitive interactions and successional replacements of grassland plant species. However, introduction of whole soil communities as management strategy in grassland restoration has received little experimental testing. In a 5-year field experiment at a topsoil-removed ex-arable site ( receptor site ), we tested effects of (1) spreading hay and soil, independently or combined, and (2) transplanting intact turfs on plant and soil nematode community development. Material for the treatments was obtained from later successional, species-rich grassland ( donor site ). Spreading hay affected plant community composition, whereas spreading soil did not have additional effects. Plant species composition of transplanted turfs became less similar to that in the donor site. Moreover, most plants did not expand into the receiving plots. Soil spreading and turf transplantation did not affect soil nematode community composition. Unfavorable soil conditions (e.g., low organic matter content and seasonal fluctuations in water level) at the receptor site may have limited plant and nematode survival in the turfs and may have precluded successful establishment outside the turfs. We conclude that introduction of later successional soil organisms into a topsoil-removed soil did not facilitate the establishment of later successional plants, probably because of the "mismatch" in abiotic soil conditions between the donor and the receptor site. Further research should focus on the required conditions for establishment of soil organisms at restoration sites in order to make use of their contribution to grassland restoration. We propose that introduction of organisms from "intermediate" stages will be more effective as management strategy than introduction of organisms from "target" stages.     

Fruit‐feeding Butterfly Communities are Influenced by Restoration Age in Tropical Forests

Currently, a large‐scale restoration project aims to restore around 15 million hectares of Atlantic Forest in Brazil. This will increase forest cover and connectivity among remnant sites as well as restore environmental services. Currently, studies on recovery of fauna in restored areas of the Atlantic Forest are practically nonexistent. To address this knowledge vacuum, our study compares diversity patterns of fruit‐feeding butterflies in three forest areas with different restoration ages (11, 22, and 54 years), and uses a native forest area as reference. Results showed butterfly communities in maturing restored areas becoming more similar to the ones found in the native forest, with an increase in the proportional abundance of forest species, and a decrease of edge and grassland species. Moreover, we found a higher diversity among sites at the intermediate restoration age, with a community composed of both grassland and forest species. Butterfly species composition differed significantly among sites, showing interesting patterns of potential species replacement over time. Our results indicate that, although restored sites were located in a fragmented landscape, they provide suitable habitats for recolonization by fruit‐feeding butterfly assemblages. Hence, restored areas can be considered important habitat for forest animal species, increasing local biodiversity and, possibly, restoring some of the ecosystem services provided by them.