Leaf shape and size track habitat transitions across forest–grassland boundaries in the grass family (Poaceae)

Grass leaf shape is a strong indicator of their habitat with linear leaves predominating in open areas and ovate leaves distinguishing forest‐associated grasses. This pattern among extant species suggests that ancestral shifts between forest and open habitats may have coincided with changes in leaf shape or size. We tested relationships between habitat, climate, photosynthetic pathway, and leaf shape and size in a phylogenetic framework to evaluate drivers of leaf shape and size variation over the evolutionary history of the family. We also estimated the ancestral habitat of Poaceae and tested whether forest margins served as transitional zones for shifts between forests and grasslands. We found that grass leaf shape is converging toward different shape optima in the forest understory, forest margins, and open habitats. Leaf size also varies with habitat. Grasses have smaller leaves in open and drier areas, and in areas with high solar irradiance. Direct transitions between linear and ovate leaves are rare as are direct shifts between forest and open habitats. The most likely ancestral habitat of the family was the forest understory and forest margins along with an intermediate leaf shape served as important transitional habitat and morphology, respectively, for subsequent shifts across forest–grassland biome boundaries.     

Asymmetric carabid beetle spillover between calcareous grasslands and coniferous forests

Central European calcareous grasslands are considered biodiversity hotspots, but are severely threatened by the change in land-use and by habitat fragmentation. Coniferous forests are typical adjacent habitats to calcareous grasslands, as abandoned calcareous grasslands are often afforested or develop into coniferous forests by succession. To investigate spillover between calcareous grasslands and coniferous forests, a total of 144 pitfall traps for carabid beetles were placed at three different distances (1, 5, 20 m) from the edge in both habitats at eight locations from April to late August. We found that both habitats had a distinct species assemblage and a decrease in spillover with increasing distance from the habitat edge into the adjacent habitat. Calcareous grasslands were more affected by spillover from the adjacent coniferous forests than vice versa because more forest specialists penetrated into calcareous grasslands than grassland specialists penetrated into coniferous forests. We conclude that spillover into small and isolated habitats can severely change species assemblages, which has to be considered in conservation measures. The protection of large sites with small edge-interior ratios can reduce negative effects on species assemblages in endangered calcareous grasslands.     

Responses of ground living arthropods to landscape contrast and context in a forest-grassland mosaic

Landscape context and contrast are major features of transformed landscapes. These concepts are largely described in terms of vegetation and land use, and are rarely used on how other biodiversity responds to these anthropogenic boundaries. South African grassland matrix is naturally dotted with indigenous forest patches which have recently been transformed with plantations of non-native species. We investigate how various arthropod groups (detritivores, predators, ants) respond to juxtaposition of pines, natural forests and grasslands. We assess landscape context effects between natural forests and pines by determining how species that commonly occur in the interiors of these habitats use the adjacent habitat, and how landscape contrast between natural forests and grassland affects these groups proportionately. We sampled arthropods using pitfall traps and active searches in transects running from natural forest interiors across the edge into the matrix interior (grassland or pines). Natural forests had higher predator and detritivore diversity, while grassland had greater ant diversity. Results highlighted the complementarity of natural forests and grassland for arthropod diversity. Higher beta-diversity was recorded across landscape contrast than landscape context. Pine and natural forest associated species overlapped into adjacent habitats indicating that pines are used by certain natural forest species. However, pines are not true natural forest extensions, with only some species being supported. Pines may be connecting naturally isolated arthropod populations, which could have important evolutionary consequences. Only through appreciation of a range of arthropod groups and their response to context and contrast across the whole landscape can we undertake meaningful biodiversity conservation.     

Habitat selection and movement ecology of eastern Hermann’s tortoises in a rural Romanian landscape

Understanding wildlife movements and habitat selection are critical to drafting conservation and management plans. We studied a population of eastern Hermann’s tortoise (Testudo hermanni boettgeri) in a traditionally managed rural landscape in Romania, near the northern edge of the species geographic distribution. We used telemetry to radio-track 24 individuals between 2005 and 2008 and performed a Euclidian distance-based habitat selection analysis to investigate habitats preferred by tortoises at both landscapes (second-order order selection) and individual (third-order selection) home range scales. The home range size for tortoises in our study area was 3.79?±?0.62 ha and did not differ by gender or season (pre- and post-nesting seasons). Their movement ecology was characterized by short-distance movements (daily mean?=?31.18?±?1.59 m), apparently unaffected by habitat type. In contrast to other studies, movements of males and females were of similar magnitude. At the landscape (population home range) scale, grasslands and shrubs were preferred, but tortoises also showed affinity to forest edges. At the individual home range scale, tortoises selected grassland and shrub habitats, avoided forests, and used forest edges randomly. Creeks were avoided at both spatial scales. Our results suggest that tortoise home ranges contain well-defined associations of habitats despite a higher selection for grasslands. As such, avoiding land conversion to other uses and maintaining habitat heterogeneity through traditional practices (e.g., manual mowing of grasslands, livestock grazing) are critical for the persistence of tortoise populations.     

Importance of local habitat conditions and past and present habitat connectivity for the species richness of grassland plants and butterflies in power line clearings

Agricultural intensification has caused drastic declines in the area and species richness of semi-natural grasslands across Europe. Novel habitats, such as power line clearings, provide alternative habitats and niches for grassland species, and might therefore mitigate these declines. However, it is not fully understood which environmental factors determine the occurrence of grassland species in the clearings. Identifying the most important drivers for grassland species occurrence would help understand the value of the clearings for grassland conservation and target enhanced management into clearings with most potential as grassland habitat. We studied the effects of local environmental conditions, and past and present connectivity to semi-natural grasslands, on the species richness of grassland plants and butterflies in 43 power line clearings in Finland. The results of generalized linear models and hierarchical partitioning showed that increasing time since clear-cut and amount of clearing residue decreased the species richness of both species groups, while the cover of mesic habitats increased it. However, the two species groups showed also divergent responses. Present-day local environmental conditions appeared to be the sole driver of grassland butterfly species richness, whereas the richness of grassland plants was related both to current conditions and historical connectivity to grasslands in 1870–1880s. This suggests the presence of an extinction debt in the studied grassland plant communities, emphasizing the need for enhanced management to increase suitable grassland habitat in the clearings. This would diminish the potential future losses of grassland plant species in the clearings and create valuable habitat for grassland butterflies as well.     

Invertebrate conservation at the interface between the grassland matrix and natural Afromontane forest fragments

Diversity patterns of amphipods, carabid beetles and ants were investigated in five naturally-fragmented Afromontane forest remnants, and in the surrounding grassland matrix. Forests were architecturally similar. In contrast, grasslands surrounding these forests are subject to great differences in anthropogenic impacts. Consequently, transition from forest to grassland ranged from being abrupt (heavy disturbance) to gradual (little disturbance). Significantly different mean numbers of carabid individuals and species were captured between sites and multivariate analyses showed clear separation in carabid assemblage-structure with level of disturbance. Carabids were, furthermore, significantly more diverse in forests, compared to grasslands. Ants, however, were equally species rich between sites but were significantly more abundant and species rich in grasslands than forests. Amphipods, represented here by a single species, Talistroides africana, was significantly less abundant at highly disturbed sites and significantly more abundant in forests than grasslands. Results support the hypothesis that the dynamics of remnants are influenced by their surrounding landscape. Here, the dynamics of amphipods and carabids (predominantly forest taxa) were influenced by different disturbance regimes in grasslands surrounding these forests. Epigaeic ants, a predominantly grassland taxon here, also showed significant differences in assemblage-composition between sites with varying disturbance. Conserving Afromontane grasslands should be of prime concern because this will include the protection of forest/grassland ecotones and forest remnants.     

The isotopic ecology of East African mammals

Summary The stable carbon and nitrogen isotope ratios of bone collagen have been used to trace diet and habitat selection of the larger mammals of East Africa. 238 individuals of 43 species from montane forests and grasslands in Kenya and Tanzania have been analyzed. The results show that carbon isotopes discriminate between (1) grazers and browsers in savanna grasslands, (2) forest floor and savanna grassland herbivores and (3) forest floor and forest canopy species. Nitrogen isotopes discriminate between (4) carnivores and herbivores, (5) forest and savanna grassland herbivores, and (6) water-dependent and drought-tolerant herbivores. This technique provides a quantitative approach to assessing long-term habitat and diet selection and the role of resource partitioning in animal community structure.Also Archaeology Program     

Genetic,morphological, and dietary changes associated with novel habitat colonisation in the Canary Island endemic grasshopper Acrostira bellamyi

1. The large flightless grasshopper Acrostira bellamyi Uvarov, endemic to the island of La Gomera (Canary Islands), inhabits two different environments: the xeric euphorb shrubland, as is typical for congeneric Canarian species, and the humid laurel forest, a novel habitat for the genus. 2. We investigate genetic, morphological, and ecological variation among individuals of A. bellamyi from the two habitats. DNA sequence data were used to evaluate whether grasshoppers from the two environments represent distinct lineages. Morphological and trophic analyses were performed to assess phenotypic differentiation between the two different habitats. 3. Population genetic analyses support the hypothesis that the euphorb shrubland is the ancestral habitat for this species. Female laurel forest specimens are larger than those inhabiting the euphorb shrubland, and some external body parts exhibit significant morphometric differences between the two populations. Diet of shrubland individuals is completely different from that of laurel forest individuals. Although in each habitat they consume the most abundant plants, individuals are able to select food plants, which appear to be explained by their nutrient content. 4. Our results suggest that A. bellamyi has colonised laurel forest from shrubland, and that this habitat shift has resulted in genetic, morphological, and ecological changes, perhaps as an adaptation to this new habitat.     

The importance of littoral forest remnants for indigenous bird conservation in southeastern Madagascar

The littoral forests of Madagascar are relatively unexplored ecosystems that are considered seriously threatened by deforestation and habitat fragmentation. We set out to describe the bird communities inhabiting the littoral forest remnants in three different sub-regions of southeastern Madagascar to determine the national importance of these forests for bird conservation. In total, 77 bird species were found inhabiting 14 littoral forest remnants. Of these species, 40 are endemic to Madagascar and a further 21 are endemic to the Indian Ocean sub-region, consisting of Madagascar, the Comoros and the Mascarenes. The matrix habitats (Melaleuca forests, marécage swamp forest, Eucalyptus plantations and Erica grassland) that immediately surround the littoral forests were depauperate of bird species and contained few species that were found within the littoral forests. The geographic location of littoral forest remnants had an important role in determining what bird species occurred within them, with the northern remnants having similar bird communities to nearby humid forest whilst the most southern remnant had a bird community that resembled those of nearby spiny forest habitats. Eleven bird species that have been previously described as being habitat-restricted endemics to either spiny forests or humid forests, were found in littoral forest remnants. These results suggest that these littoral forests may play an important transitional role between the two other major natural habitats (spiny forest and humid forest) of southeastern Madagascar. On this basis we advocate that the littoral forest remnants of southeastern Madagascar should be afforded continuing conservation priority.     

Habitat preferences and distribution characteristics are indicative of species long-term persistence in the Estonian flora

Large-scale changes in regional floras provide direct information about changes in biodiversity through time and enable the evaluation of conservation targets. We compared the distribution ranges in 2004 of Estonian native terrestrial flora with the distribution ranges before 1970, using the Atlas of Estonian Flora. Relative persistence was related to species endemism, commonness, occurrence at its border of the global distribution range, main habitat type, sensitivity to human impact, life-form, conservation category, and Red List category. A literature-based database of the flora of Estonian habitat types was used to evaluate relative persistence of the flora of different habitats. Changes in the flora are largely dependent on human activities. The decrease in mire and grassland habitats and the increase in forests are reflected in the persistences of related species. Flora of mire habitats decreased the most. The fact that an almost ten-fold decrease of grasslands has not resulted in as large a decrease in the ranges of grassland species could serve as evidence of the extinction debt of these habitats. We also found a greater decrease among habitat specialists than habitat generalists and lower average persistence of the species of species-rich habitats. Our data show that current prioritization of species for conservation is in concordance with needs, as reflected in the changes in the range of species. However, conservation has not been entirely successful: the decrease of protected species continues. Our simple method for summarizing large databases was effective for the evaluation of large scale effects of conservation actions.     

No general edge effects for invertebrates at Afromontane forest/grassland ecotones

The Afromontane region of South Africa is characterised by numerous small, remnant forests in a grassland matrix. The edges, or ecotones between forests and grasslands are usually sharp (typically just over a few metres) and are mainly maintained by both natural and, more recently, anthropogenic fires. We investigated epigaeic amphipod, carabid and ant distribution patterns across Afromontane forest/grassland ecotones and found little evidence to support the biological edge effect. Five of the fifty-two sampled species however, did increase significantly in abundance at the ecotone. Among these was a very distinct edge species, the amphipod Talistroides africana. Overall, carabids were more abundant and species rich in forests while for ants it was in the grasslands. Ants and carabids were both more abundant and species rich in spring and summer than in autumn and winter. More interestingly, the abundance and species richness patterns across the ecotone did not change with the passing of the seasons. We argue that a conservation strategy for the Afromontane forest patches must also incorporate the surrounding grassland. The grassland habitat is often perceived as less valuable than forest and, as a consequence, is subject to many anthropogenic disturbances such as fragmentation, cattle grazing and afforestation. Protecting grasslands around forest patches not only conserves the rich ant diversity, but also conserves the biota in the forests and at the edges, and would therefore be more meaningful in terms of the overall conservation of Afromontane biodiversity.     

The influence of biogeographic history on the functional and phylogenetic diversity of passerine birds in savannas and forests of the Brazilian Amazon

Passeriformes is the largest and most diverse avian order in the world and comprises the Passeri and Tyranni suborders. These suborders constitute a monophyletic group, but differ in their ecology and history of occupation of South America. We investigated the influence of biogeographic history on functional and phylogenetic diversities of Passeri and Tyranni in forest and savanna habitats in the Brazilian Amazon. We compiled species composition data for 34 Passeriformes assemblages, 12 in savannas and 22 in forests. We calculated the functional (Rao's quadratic entropy, FD_Q) and phylogenetic diversities (mean pairwise distance, MPD, and mean nearest taxon distance, MNTD), and the functional beta diversity to investigate the potential role of biogeographic history in shaping ecological traits and species lineages of both suborders. The functional diversity of Passeri was higher than for Tyranni in both habitats. The MPD for Tyranni was higher than for Passeri in forests; however, there was no difference between the suborders in savannas. In savannas, Passeri presented higher MNTD than Tyranni, while in forest areas, Tyranni assemblages showed higher MNTD than Passeri. We found a high functional turnover (~75%) between Passeri and Tyranni in both habitats. The high functional diversity of Passeri in both habitats is due to the high diversity of ecological traits exhibited by species of this group, which enables the exploitation of a wide variety of resources and foraging strategies. The higher Tyranni MPD and MNTD in forests is likely due to Tyranni being older settlers in this habitat, resulting in the emergence and persistence of more lineages. The higher Passeri MNTD in savannas can be explained by the existence of a larger number of different Passeri lineages adapted to this severe habitat. The high functional turnover between the suborders in both habitats suggests an ecological strategy to avoid niche overlap.     

新疆吉木萨尔县蝴蝶群落多样性

蝴蝶作为指示生物, 被广泛地应用于生物多样性监测及环境质量评估。探究新疆吉木萨尔县蝴蝶群落多样性, 可为当地蝴蝶多样性的保护及环境监测提供基础数据。本研究采用样线法在新疆吉木萨尔县选取山前荒漠、农田、山地草原、山地森林、亚高山草甸5种不同的生境类型, 对蝴蝶种类和群落多样性进行调查。共记录蝴蝶4,401号, 隶属于7科26属38种。其中蛱蝶科有9属12种, 为优势科; 粉蝶科的个体数最多, 占比55.01%; 绢蝶科、凤蝶科和弄蝶科的种类数和个体数最少, 均为单科种, 是该地区的稀有类群。对不同生境蝴蝶群落多样性和相似度分析比较的结果显示: 5种生境中多样性指数从高到低依次为亚高山草甸、山地森林、山地草原、农田及山前荒漠, 其中山地森林和亚高山草甸的相似性系数较高, 达到0.77, 山前荒漠和山地草原的相似性系数最低, 为0.37。蝴蝶物种数及多样性指数随海拔的增加呈上升趋势。蝴蝶群落随月份发生变化, 蝴蝶种类和数量在5月发生、7月达到峰值。蝶类个体数在3年内呈下降趋势。研究结果表明, 蝴蝶物种的组成和多样性与生境类型具有密切联系, 保护生态环境, 维持该地区植物群落的多样性、降低人为干扰程度是保护蝶类多样性的关键。     

Plant species occurrences in a rural hemiboreal landscape: effects of remnant habitats, site history, topography and soil

Changes in land use during the last century have caused fragmentation and a reduction in area of many species-rich habitats in the hemiboreal region. We examined abundances of plant species and their occurrence in different habitats in south-east Sweden. We found 361 plant species in 146 sample sites, which represented 14 different types of habitat. Most species were rare and occurred only in a few habitats. Almost half of all species (49%) were found in one or two habitats. Of these, 99 species occurred in one habitat only. The habitats with largest number of restricted species, i.e. habitat specialists, were dry to mesic semi-natural grasslands and remnant habitats such as road verges and mid field islets. The occurrence of 52 species was analysed with respect to topography, top- and subsoil and land use history. Few of the 52 species were affected by aspect or type of topsoil. Subsoil affected nearly half of the species and habitats with a convex landform influenced occurrence of > 90% of the species. Seventeen species were positively associated with a long continuity of grassland management, whereas two species were associated with lack of management. Open grasslands that are encroached by trees and shrubs show a decline in species number. Deciduous forests, especially wet deciduous forests, have a potential for restoring moist to mesic grassland habitats. Small remnant habitats are important for many of the species restricted to semi-natural grasslands. These habitats may function as "rescue sites" for the species, which in turn may promote dispersal and increase likelihood of restoration success. Therefore, remnant habitats are important for maintaining and restoring species richness in rural landscapes.     

Comparison of butterfly communities and abundances between marginal grasslands and conservation lands in the eastern Great Plains

The alteration and fragmentation of native tallgrass prairie in the Midwestern United States has created a need to identify other land types with the ability to support grassland butterfly species. This study examines butterfly usage of marginal grasslands, which consist of semi-natural grasslands existing within in a larger agricultural matrix, compared to grasslands managed for conservation of prairie species. Using generalized linear mixed models we analyzed how land purpose (marginal vs. conservation grasslands) affected butterfly abundance. We found grassland butterfly species to be significantly more common on conservation grasslands, whereas generalist species were significantly more common on marginal grasslands. Results of ordination analyses indicated that while many species used both types of habitats, butterfly species assemblages were distinct between habitat types and that edge to interior ratio and the floristic quality index of sites were important habitat characteristics driving this distinction. Within conservation grasslands we examined the relationship between butterfly abundance and the planting diversity used in restoring each site. We found higher diversity restorations hosted more individuals of butterflies considered habitat generalists, as well as species considered to be of conservation concern.     

Are Montane Forests Demographic Sinks for Bornean White‐bearded Gibbons Hylobates albibarbis?

I estimated habitat-specific population densities for a population of Bornean white-bearded gibbons Hylobates albibarbis inhabiting seven distinct forest types at Gunung Palung National Park, West Kalimantan, Indonesia. Population densities in montane forests (0.44 individuals/km²) were almost ten times lower than those in the next best habitat (upland granite forest; 4.2 individuals/km²) and far lower than those in lowland forest types. Demographic data on 33 gibbon groups living across the seven forest types showed that reproduction was substantially depressed in montane forests compared to high-quality lowland habitats. A simple model suggests that montane forests are demographic sinks for gibbons at Gunung Palung. Follow-up data from observations of montane groups 5 yr after the initial observation period support this result. As high-quality lowland forests (source habitat for gibbons) are being disproportionately lost in and around Gunung Palung National Park due to illegal logging and conversion to oil palm plantations, an increasing percentage of the remaining forest in the park comprises sink habitat for gibbons. This result has disquieting implications for the long-term viability of gibbon populations at Gunung Palung. In addition, as montane forests are generally low-quality habitat for most rainforest vertebrates, and since lowland forests are being lost at alarming rates across the tropics, source-sink population dynamics similar to those I describe here may characterize populations of other tropical vertebrate species.     

The Interplay between Environmental Filtering and Spatial Processes in Structuring Communities: The Case of Neotropical Snake Communities

Both habitat filters and spatial processes can influence community structure. Space alone affects species immigration from the regional species pool, whereas habitat filters affect species distribution and inter-specific interactions. This study aimed to understand how the interplay between environmental and geographical processes influenced the structure of Neotropical snake communities in different habitat types. We selected six studies that sampled snakes in forests, four conducted in savannas and two in grasslands (the latter two are grouped in a non-forest category). We used the net relatedness and nearest taxon indices to assess phylogenetic structure within forest and non-forest areas. We also used the phylogenetic fuzzy-weighting algorithm to characterize phylogenetic structure across communities and the relation of phylogenetic composition patterns to habitat type, structure, and latitude. Finally, we tested for morphological trait convergence and phylogenetic niche conservatism using four forest and four non-forest areas for which morphological data were available. Community phylogenetic composition changed across forest and non-forest areas suggesting that environmental filtering influences community structure. Species traits were affected by habitat type, indicating convergence at the metacommunity level. Tail length, robustness, and number of ventral scales maximized community convergence among forest and non-forest areas. The observed patterns suggested environmental filtering, indicating that less vertically structured habitats represent a strong filter. Despite the fact that phylogenetic structure was not detected individually for each community, we observed a trend towards communities composed by more closely related species in higher latitudes and more overdispersed compositions in lower latitudes. Such pattern suggests that the limited distribution of major snake lineages constrained species distributions. Structure indices for each community were also related to habitat type, showing that communities from non-forest areas tend to be more clustered. Our study showed that both environmental filtering and spatial gradients play important roles in shaping the composition of Neotropical snake communities.     

The effects of reproductive specialization on energy costs and fitness genetic variances in cyclical and obligate parthenogenetic aphids

Organisms with coexisting sexual and asexual populations are ideal models for studying the consequences of either reproductive mode on the quantitative genetic architecture of life-history traits. In the aphid Rhopalosiphum padi, lineages differing in their sex investment coexist but all share a common parthenogenetic phase. Here, we studied multiple genotypes of R. padi specialized either for sexual and asexual reproduction and compared their genetic variation in fitness during the parthenogenetic phase. Specifically, we estimated maintenance costs as standard metabolic rate (SMR), together with fitness (measured as the intrinsic rate of increase and the net reproductive rate). We found that genetic variation (in terms of broad-sense heritability) in fitness was higher in asexual genotypes compared with sexual genotypes. Also, we found that asexual genotypes exhibited several positive genetic correlations indicating that body mass, whole-animal SMR, and apterous individuals production are contributing to fitness. Hence, it appears that in asexual genotypes, energy is fully allocated to maximize the production of parthenogenetic individuals, the simplest possible form of aphid repertoire of life-histories strategies.     

Forest eternal? Endemic butterflies of the Bamenda highlands,Cameroon, avoid close‐canopy forest

The Gulf of Guinea Highlands, a centre of endemism and high conservation importance, represent the only large mountain system in West and Central Africa. We studied habitat use of three common endemic butterflies Colias electo manengoubensis, Bicyclus anisops and Mylothris jacksoni knutsoni, using time‐standardized surveys in four distinct habitats: close‐canopy forest, scrub and forest edges, bracken and grasslands. All three species avoided close‐canopy forests and bracken; the Colias preferring grassland, whereas Bicyclus and Mylothris scrub and forest edges. Ordination analyses of surrounding habitats indicated that all three taxa required heterogeneous landscape mosaics. We argue that the life history traits of taxa with limited geographic distribution may reflect past habitat conditions within their ranges, and that these habitat preferences can indicate the continuous existence of mosaic of forest and nonforest habitats in the West African mountains. Such a landscape was probably maintained by climatic fluctuation and large herbivores, further modified by human impact. This conclusion is consistent with the palaeoenvironmental record and with the requirements of Afromontane endemics from other groups. Recent conservation activities focus on patches of continuous forests, but the mosaic landscapes are no less threatened by intensive agriculture, and should be included to protected areas.     

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.