Habitat structure components are effective predictors of trap-nesting Hymenoptera diversity

Habitat heterogeneity can be the major factor affecting species diversity in a community and measuring bee and wasp community habitat preferences in natural systems may provide insights for biodiversity management and conservation. In the present study, we investigate the effects of habitat structure components on solitary bee and wasp species richness and abundance. The research was conducted in an urban forest remnant in southeast Brazil. Our main questions were: (1) is similarity in habitat structure mirrored by similarity in Aculeate assemblage composition? and (2) what are the vegetation features that could be used as predictors of solitary bee and wasp richness and abundance? Aculeate bees and wasps were sampled using trap nests from February to November 2004. Trap nests were placed in sampling units located in 6 ha of secondary mesophytic forest. One hundred and thirty-seven trap nests were occupied by four species of wasps and seven species of bees. Altogether, our sampling units had a mean capture rate (relative to expected richness) of 72% during all the study period. The more similar sampling units were in terms of vegetation structure, the more similar they were in solitary bee and wasp species composition. The variance of tree abundance, shrub height and the abundance of wood logs were good predictors of solitary bee and wasp species richness and abundance in the study area. We demonstrate that even in a small scale it is possible to detect significant influences of habitat features on alpha diversity and that some of them are effective as predictors of trap-nesting Hymenoptera richness and abundance.     

Contrasting Patterns of Species Richness and Composition of Solitary Wasps and Bees (Insecta: Hymenoptera) According to Land‐use

Several species of arthropods inhabiting forest fragments interact with managed areas. The importance of such areas to biodiversity conservation, however, is not well established. Communities of solitary wasps and bees (Insecta: Hymenoptera) play a key role in agroecosystem functioning and they have been used in studies of biodiversity assessment in different land‐use types. We aimed to assess patterns of species richness and composition of solitary wasps and bees over a 1‐yr period in a gradient of decreasing land‐use intensity formed by pastures, alley croppings, young fallows, and old fallows using trap nests. Old fallows had the highest species richness of wasps and bees, harboring all bee species and 86 percent of wasp species occurring in the region, while the remaining land‐uses had similar species richness. Vegetation structure (tree richness) and relative humidity explained most of the variance for the species richness of wasps. For bees, however, there was no influence of environmental factors on the community among land‐use types, indicating better adaptability of this group to environmental variations related to land‐use. The composition of solitary wasp communities (but not those of bees) differed among land‐use types, and the occurrence of rare species in most cases was restricted to old fallow sites. In conclusion, the community of solitary wasps and bees is contingent on land‐use, with solitary wasps more sensitive to anthropized areas. For both groups, less anthropized areas harbor a greater richness and number of rare species while more intensively managed land‐use types harbor higher abundances.     

Spatio-temporal variation in species assemblages in field edges: seasonally distinct responses of solitary bees to local habitat characteristics and landscape conditions

The bees (Hymenoptera: Apiformes) are important pollinators in many ecosystems, but their diversity has declined in Europe during the past century, mainly due to habitat loss. However, some of the habitat requirements of wild bees are met in anthropogenic landscape elements, such as road sides, power-line strips and field edges. Moreover, as the bee species assemblages change throughout the season the habitat requirements of the bee fauna may change accordingly. Understanding such seasonally distinct responses of solitary bees with different phenologies may be of high value for local conservation planning. The purpose of this study was to examine if the habitat quality of field edges for solitary bees change throughout the season, and how this temporal variation relates to local habitat and landscape conditions. By sampling solitary bees in 18 field edges in southeast Norway throughout the season we found that the species richness and abundance of bees was highest in sun exposed field edges, independently of the season. However, we found phenologically distinct responses to the landscape context. Moreover, field edges situated in landscapes with a high proportion of forests and semi-natural landscape elements hosted the most phenologically diverse bee species assemblages. We conclude that in order to fulfil the habitat requirements of bee species assemblages throughout the season, one should conserve and direct habitat restoration schemes towards increasing sun exposure at field edges with a diverse flora and a high proportion of semi-natural areas in the vicinity.     

Methods matter: Different biodiversity survey methodologies identify contrasting biodiversity patterns in a human modified rainforest — A case study with amphibians

Understanding how well tropical forest biodiversity can recover following habitat change is often difficult due to conflicting assessments arising from different studies. One often overlooked potentially confounding factor that may influence assessments of biodiversity response to habitat change, is the possibility that different survey methodologies, targeting the same indicator taxon, may identify different patterns and so lead to different conclusions. Here we investigated whether two different but commonly used survey methodologies used to assess amphibian communities, pitfall trapping and nocturnal transects, indicate the same or different responses of amphibian biodiversity to historic human induced habitat change. We did so in a regenerating rainforest study site located in one of the world’s most biodiverse and important conservation areas: the Manu Biosphere Reserve. We show that the two survey methodologies tested identified contrasting biodiversity patterns in a human modified rainforest. Nocturnal transect surveys indicated biodiversity differences between forest with different human disturbance histories, whereas pitfall trap surveys suggested no differences between forest disturbance types, except for community composition. This pattern was true for species richness, diversity, overall abundance and community evenness and structure. For some fine scale metrics, such as species specific responses and abundances of family groups, both methods detected differences between disturbance types. However, the direction of differences was inconsistent between methods. We highlight that for assessments of rainforest recovery following disturbance, survey methods do matter and that different biodiversity survey methods can identify contrasting patterns in response to different types of historic disturbance. Our results contribute to a growing body of evidence that arboreal species might be more sensitive indicators than terrestrial communities.     

Anthropogenic impact on habitat connectivity: A multidimensional human footprint index evaluated in a highly biodiverse landscape of Mexico

Evaluating the cumulative effects of the human footprint on landscape connectivity is crucial for implementing policies for the appropriate management and conservation of landscapes. We present an adjusted multidimensional spatial human footprint index (SHFI) to analyze the effects of landscape transformation on the remnant habitat connectivity for 40 terrestrial mammal species representative of the Trans-Mexican Volcanic System in Michoacán (TMVS_Mich), in western central Mexico. We adjusted the SHFI by adding fragmentation and habitat loss to its original three components: land use intensity, time of human landscape intervention, and biophysical vulnerability. The adjusted SHFI was applied to four scenarios: one grouping all species and three grouping several species by habitat spatial requirements. Using the SHFI as a dispersal resistance surface and applying a circuit theory based approach, we analyzed the effects of cumulative human impact on habitat connectivity in the different scenarios. For evaluating the relationship between habitat loss and connectivity, we applied graph theory-based equivalent connected area (ECA) index. Results show over 60% of the TMVS_Mich has high SHFI values, considerably lowering current flow for all species. Nevertheless, the effect on connectivity of human impact is higher for species with limited dispersal capacity (100–500 m). Our approach provides a new form of evaluating human impact on habitat connectivity that can be applied to different scales and landscapes. Furthermore, the approach is useful for guiding discussions and implementing future biodiversity conservation initiatives that promote landscape connectivity as an adaptive strategy for climate change.     

Cavity-nesting bee communities in areas with different levels of vegetation disturbance

ABSTRACT

The reasons for the decline of bee diversity and abundance include the destruction and loss of natural habitats. Protected areas are created for biodiversity conservation, but these areas vary strongly in their level of vegetation disturbance. Using trap-nests, we assessed changes in solitary bee abundance, richness, and composition in areas ranging from naturally conserved to degraded. Solitary bees were sampled during an 18-month period in three areas of southeastern Brazil: a preserved area in Rio Preto State Park – PERP; a restored/altered area with exotic plants at the Federal University of the Jequitinhonha and Mucuri Valleys – JK Campus; and a degraded area in Biribiri State Park – PEBi. A total of seven species of bees built 115 nests. In the degraded area, only two nests were built. Abundance of built nests was higher in the preserved area (PERP), but diversity was higher in the restored area (JK Campus). Our results show that the solitary bee population responds positively to habitat complexity (local scale). The presence of a diverse solitary bee fauna in the restored area indicates that altered areas should also be protected as suitable areas for re-colonization of cavity-nesting bees.     

Habitat quality matters for the distribution of an endangered leaf beetle and its egg parasitoid in a fragmented landscape

Fragmentation, deterioration, and loss of habitat patches threaten the survival of many insect species. Depending on their trophic level, species may be differently affected by these factors. However, studies investigating more than one trophic level on a landscape scale are still rare. In the present study we analyzed the effects of habitat size, isolation, and quality for the occurrence and population density of the endangered leaf beetle Cassida canaliculata Laich. (Coleoptera: Chrysomelidae) and its egg parasitoid, the hymenopteran wasp Foersterella reptans Nees (Hymenoptera: Tetracampidae). C. canaliculata is strictly monophagous on meadow sage (Salvia pratensis), while F. reptans can also parasitize other hosts. Both size and isolation of habitat patches strongly determined the occurrence of the beetle. However, population density increased to a much greater extent with increasing host plant density ( = habitat quality) than with habitat size. The occurrence probability of the egg parasitoid increased with increasing population density of C. canaliculata. In conclusion, although maintaining large, well-connected patches with high host plant density is surely the major conservation goal for the specialized herbivore C. canaliculata, also small patches with high host plant densities can support viable populations and should thus be conserved. The less specialized parasitoid F. reptans is more likely to be found on patches with high beetle density, while patch size and isolation seem to be less important.     

An overview of proximate factors affecting the nesting behavior of solitary wasps and bees (Hymenoptera: Aculeata) in preexisting cavities in wood

Guilds of Aculeate solitary wasps and bees that nest in preexisting cavities in wood are important components of terrestrial ecosystems because they engage in several ecological interactions (e.g. predation and pollination) with other species of plants and animals. Spatial and temporal variations in richness and abundance of solitary wasps and bees can be related to changes in environmental structure and in the diversity of other groups of organisms. The nesting period of these Aculeata is their most critical life cycle stage. Females of solitary wasp and bee species invest relatively more time constructing and provisioning their nests than do females of social species. Differently from species that nest in the soil or construct exposed nests, the main factors affecting the reproductive success of solitary species nesting in preexisting wood holes are still unknown. Our objective is to provide an overview of the role of proximate causes of nesting failure or success among solitary wasps and bees (Aculeata), for designing effective conservation and management strategies for these Hymenoptera.     

Yet Another Empty Forest: Considering the Conservation Value of a Recently Established Tropical Nature Reserve

The primary approach used to conserve tropical biodiversity is in the establishment of protected areas. However, many tropical nature reserves are performing poorly and interventions in the broader landscape may be essential for conserving biodiversity both within reserves and at large. Between October 2010 and 2012, we conducted bird surveys in and around a recently established nature reserve in Xishuangbanna, China. We constructed a checklist of observed species, previously recorded species, and species inferred to have occurred in the area from their distributions and habitat requirements. In addition, we assessed variation in community composition and habitat specificity at a landscape-scale. Despite the fact that the landscape supports a large area of natural forest habitat (~50,000 ha), we estimate that >40% of the bird fauna has been extirpated and abundant evidence suggests hunting is the primary cause. A large proportion (52%) of the bigger birds (>20 cm) were extirpated and for large birds there was a U-shaped relationship between habitat breadth and extirpation probability. Habitat specificity was low and bird communities were dominated by widespread species of limited conservation concern. We question whether extending tropical protected area networks will deliver desired conservation gains, unless much greater effort is channeled into addressing the hunting problem both within existing protected areas and in the broader landscape.     

Patterns and drivers of wild bee community assembly in a Mediterranean IUCN important plant area

Recent reports of pollinator declines have stirred interest in investigating the impacts of habitat exploitation on the conservation of pollinator and plant communities. An important prerequisite to tailor conservation action is to understand the drivers and patterns of species-rich communities, and how they change in space and time during a whole season. To account for this, we surveyed wild bees and flowering plants using standardized transects in 11 natural habitat fragments of an IUCN important plant area along the coast of Israel. We used phylogeny- and taxon-based methods of community structure analyses to study the assembly processes of bee communities, and investigated the effects of several landscape parameters on bee diversity using generalized linear models (GLMs). Our results illustrate that natural habitat sites comprised significantly higher species richness compared to disturbed habitat sites, and show that even the smallest habitat fragments harbored unique bee assemblages, with significant species replacement (turnover) found in both space and time. Our GLMs indicated that flower diversity, and semi-natural habitat within 500 m of habitat fragments were important drivers of bee diversity, but we found no evidence for a species—area relationship among sites. Finally, we document a case of phylogenetic overdispersion despite low species richness, which highlights the importance of accounting for phylogenetic diversity rather than only species richness to reach a more fine-grained understanding of pollinator diversity. This, in turn, is pivotal to developing conservation actions to protect these essential pollinators and their interaction with rare and endemic plant species in this highly threatened ecosystem.     

Testing projected wild bee distributions in agricultural habitats: predictive power depends on species traits and habitat type

Species distribution models (SDM) are increasingly used to understand the factors that regulate variation in biodiversity patterns and to help plan conservation strategies. However, these models are rarely validated with independently collected data and it is unclear whether SDM performance is maintained across distinct habitats and for species with different functional traits. Highly mobile species, such as bees, can be particularly challenging to model. Here, we use independent sets of occurrence data collected systematically in several agricultural habitats to test how the predictive performance of SDMs for wild bee species depends on species traits, habitat type, and sampling technique. We used a species distribution modeling approach parametrized for the Netherlands, with presence records from 1990 to 2010 for 193 Dutch wild bees. For each species, we built a Maxent model based on 13 climate and landscape variables. We tested the predictive performance of the SDMs with independent datasets collected from orchards and arable fields across the Netherlands from 2010 to 2013, using transect surveys or pan traps. Model predictive performance depended on species traits and habitat type. Occurrence of bee species specialized in habitat and diet was better predicted than generalist bees. Predictions of habitat suitability were also more precise for habitats that are temporally more stable (orchards) than for habitats that suffer regular alterations (arable), particularly for small, solitary bees. As a conservation tool, SDMs are best suited to modeling rarer, specialist species than more generalist and will work best in long‐term stable habitats. The variability of complex, short‐term habitats is difficult to capture in such models and historical land use generally has low thematic resolution. To improve SDMs’ usefulness, models require explanatory variables and collection data that include detailed landscape characteristics, for example, variability of crops and flower availability. Additionally, testing SDMs with field surveys should involve multiple collection techniques.     

Effects of habitat simplification on assemblages of cavity nesting bees and wasps in a semiarid neotropical conservation area

Habitat complexity is directly correlated to insect diversity in most natural environments. Structural complexity reflects an increase in vertical stratification and plant diversity and often leads to a greater availability of floral resources and nesting sites. Efficient conservation strategies require understanding of how changes in habitat structure affect insects that provide essential ecosystem services. We analyzed how the diversity and species composition of bees and wasps that nest in pre-existing cavities is affected by habitat complexity. Our study was developed in the semiarid region of northeastern Brazil, in the Ubajara National Park and surrounding area. Four types of habitats within two physiognomies were sampled for two consecutive years. We used 120 trap-nest (9000 cavities) distributed in 40 sample points. Overall, 657 cavities were occupied by 11 species of bees, nine of wasps, and six of cleptoparasitic/parasitoids. Bees and wasp diversity increases with habitat complexity. While species richness was higher in more complex physiognomies, abundance was higher in disturbed areas. Species composition also varied with habitat structure. Habitat simplification has adverse effects on the diversity and composition of assemblages. These effects are stronger in more complex habitats indicating that conservation of humid habitats within semiarid areas is essential to maintain bee and wasp regional diversity.     

Wild bees along an urban gradient: winners and losers

The variability of bee communities along an urban gradient has a potentially strong impact on pollinator-plant interactions. We investigated changes in bee species richness and community composition in the city centre and suburbs of Pozna??, Poland. During 2006?C2008 we recorded 2,495 individuals from 104 species. The most abundant species was Andrena haemmorhoa. Other abundant species were Bombus pascuorum, Andrena vaga, Andrena nitida, Bombus terrestris. Several species appeared to be more abundant in the city centre whereas others showed an opposite tendency. We have found that the urbanized landscape can act as a filter for the bee community since some specific ecological traits facilitate colonization of the city centre. Small-bodied species that start their activity later in the season and are not solitary preferred the city centre. However, bee species diversity and richness remained stable across the urban gradient indicating some advantages of being a city dwelling bee. We suggest that a city can be a very important habitat for a diverse bee fauna and that conservation of a wide range of habitats of different urbanization levels seems to be the most suitable strategy for conservation of bee diversity in cities.     

High Bee and Wasp Diversity in a Heterogeneous Tropical Farming System Compared to Protected Forest

It is a globally important challenge to meet increasing demands for resources and, at the same time, protect biodiversity and ecosystem services. Farming is usually regarded as a major threat to biodiversity due to its expansion into natural areas. We compared biodiversity of bees and wasps between heterogeneous small-scale farming areas and protected forest in northern coastal Belize, Central America. Malaise traps operated for three months during the transition from wet to dry season. Farming areas consisted of a mosaic of mixed crop types, open habitat, secondary forest, and agroforestry. Mean species richness per site (alpha diversity), as well as spatial and temporal community variation (beta diversity) of bees and wasps were equal or higher in farming areas compared to protected forest. The higher species richness and community variation in farmland was due to additional species that did not occur in the forest, whereas most species trapped in forest were also found in farming areas. The overall regional species richness (gamma diversity) increased by 70% with the inclusion of farming areas. Our results suggest that small-scale farming systems adjacent to protected forest may not only conserve, but even favour, biodiversity of some taxonomic groups. We can, however, not exclude possible declines of bee and wasp diversity in more intensified farmland or in landscapes completely covered by heterogeneous farming systems.     

Agricultural buffer zone thresholds to safeguard functional bee diversity: Insights from a community modeling approach

Wild bee species are important pollinators in agricultural landscapes. However, population decline was reported over the last decades and is still ongoing. While agricultural intensification is a major driver of the rapid loss of pollinating species, transition zones between arable fields and forest or grassland patches, i.e., agricultural buffer zones, are frequently mentioned as suitable mitigation measures to support wild bee populations and other pollinator species. Despite the reported general positive effect, it remains unclear which amount of buffer zones is needed to ensure a sustainable and permanent impact for enhancing bee diversity and abundance. To address this question at a pollinator community level, we implemented a process‐based, spatially explicit simulation model of functional bee diversity dynamics in an agricultural landscape. More specifically, we introduced a variable amount of agricultural buffer zones (ABZs) at the transition of arable to grassland, or arable to forest patches to analyze the impact on bee functional diversity and functional richness. We focused our study on solitary bees in a typical agricultural area in the Northeast of Germany. Our results showed positive effects with at least 25% of virtually implemented agricultural buffer zones. However, higher amounts of ABZs of at least 75% should be considered to ensure a sufficient increase in Shannon diversity and decrease in quasi‐extinction risks. These high amounts of ABZs represent effective conservation measures to safeguard the stability of pollination services provided by solitary bee species. As the model structure can be easily adapted to other mobile species in agricultural landscapes, our community approach offers the chance to compare the effectiveness of conservation measures also for other pollinator communities in future.     

Fruit set of highland coffee increases with the diversity of pollinating bees

The worldwide decline of pollinators may negatively affect the fruit set of wild and cultivated plants. Here, we show that fruit set of the self-fertilizing highland coffee (Coffea arabica) is highly variable and related to bee pollination. In a comparison of 24 agroforestry systems in Indonesia, the fruit set of coffee could be predicted by the number of flower-visiting bee species, and it ranged from ca. 60% (three species) to 90% (20 species). Diversity, not abundance, explained variation in fruit set, so the collective role of a species-rich bee community was important for pollination success. Additional experiments showed that single flower visits from rare solitary species led to higher fruit set than with abundant social species. Pollinator diversity was affected by two habitat parameters indicating guild-specific nesting requirements: the diversity of social bees decreased with forest distance, whereas the diversity of solitary bees increased with light intensity of the agroforestry systems. These results give empirical evidence for a positive relationship between ecosystem functions such as pollination and biodiversity. Conservation of rainforest adjacent to adequately managed agroforestry systems could improve the yields of farmers.     

Kitam Bird Sanctuary,the only low elevation protected area of Sikkim: A conservation hotspot for butterflies in the Eastern Himalaya

The lowland areas of the Himalayan region are subjected to immense anthropogenic pressure because of least representation in the protected area network. Kitam Bird Sanctuary is the only representative protected area that occurs below 1000 m in Sikkim state of India (a part of globally significant biodiversity hotspot of Himalayas) and serves as the refuge for various species of flora and fauna. Here we studied butterfly diversity and community composition in Kitam Bird Sanctuary (a small protected area of 6 km² geographical area) following point count method spread across predetermined transects. Altogether 1674 butterflies belonging to 111 species and six families were recorded after completion of 240 point counts. Among these, 18 species are federally protected under the Wildlife (Protection) Act (1972) of India. Most of the butterflies were forest specialist in terms of habitat preference, whereas based on host plant specificity, the butterfly community was mostly dominated by generalist feeder (Oligophagous II and Polyphagous). Butterfly community parameters showed a strong correlation with habitat variables. While Kitam Bird Sanctuary is primarily designated for conservation of lowland birds, the high diversity of butterflies both in terms of taxonomic richness and trait composition suggests that the sanctuary harbors an ideal habitat for butterflies of the tropical region and invites conservation attention.     

A simple and distinctive microbiota associated with honey bees and bumble bees

Specialized relationships with bacteria often allow animals to exploit a new diet by providing a novel set of metabolic capabilities. Bees are a monophyletic group of Hymenoptera that transitioned to a completely herbivorous diet from the carnivorous diet of their wasp ancestors. Recent culture-independent studies suggest that a set of distinctive bacterial species inhabits the gut of the honey bee, Apis mellifera. Here we survey the gut microbiotae of diverse bee and wasp species to test whether acquisition of these bacteria was associated with the transition to herbivory in bees generally. We found that most bee species lack phylotypes that are the same or similar to those typical of A. mellifera, rejecting the hypothesis that this dietary transition was symbiont-dependent. The most common bacteria in solitary bee species are a widespread phylotype of Burkholderia and the pervasive insect associate, Wolbachia. In contrast, several social representatives of corbiculate bees do possess distinctive bacterial phylotypes. Samples of A. mellifera harboured the same microbiota as in previous surveys, and closely related bacterial phylotypes were identified in two Asian honey bees (Apis andreniformis and Apis dorsata) and several bumble bee (Bombus) species. Potentially, the sociality of Apis and Bombus species facilitates symbiont transmission and thus is key to the maintenance of a more consistent gut microbiota. Phylogenetic analyses provide a more refined taxonomic placement of the A. mellifera symbionts.     

Identifying potential conservation areas in the Cuitzeo Lake basin,Mexico by multitemporal analysis of landscape connectivity

Lake Cuitzeo basin is an important ecological area subjected to strong human pressure on forest covers that are key elements for the long-term support of biodiversity. We studied landscape connectivity changes for the years 1975, 1996, 2000, 2003 and 2008 to identify potential conservation areas in the basin. We modeled potential distributions of the Mexican bobcat (Lynx rufus escuinapae) and the ringtail (Bassariscus astutus) – two terrestrial mammal focal species with contrasting dispersal capacities – and we determined their habitat availability and suitability in the basin. We then identified their optimal habitat patches and produced landscape cumulative resistance maps, estimated least-cost paths (graph theory approach), and elaborated current flow maps (circuit theory approach). For evaluation of landscape connectivity, we applied an integral index of connectivity (IIC) to each study period, and determined individual habitat patch contribution to the overall landscape connectivity. The IIC index had very low values associated with reduced availability of focal species habitat. However, our study showed the conservation importance of the surface of optimal habitat patch areas. The combined application of a graph-based approach and current flow mapping were useful, and complementary both in terms of estimating potential dispersal corridors and identifying high probability dispersal areas. This indicated that landscape connectivity analysis is a useful tool for identification of potential conservation areas and for local landscape planning.     

Winter bird communities in the northern Negev: species dispersal patterns, habitat use and implications for habitat conservation

Bird habitat conservation may require different management strategies for different seasonal bird assemblages. We studied habitat use by winter birds in forest and scrubland habitat patches in the northern Negev, Israel. Our goal was to assess whether differences in responses to landscape and habitat structure between breeding and non-breeding seasons require changes in future conservation plans that have been suggested for the Negev breeding bird community. We evaluated habitat and area effects on bird abundance and distribution and tested whether species habitat use during winter involves niche shifts. Compared with breeding birds, a larger proportion of winter bird species occupied both scrubland and forest. As in summer, forest bird species responded to habitat structure, whereas scrubland species were associated with both habitat structure and area. Resident birds disperse into habitats in which they were not present during summer. Consequently, for several species, the correlation between bird densities and environmental factors showed a better fit at the landscape rather than at the habitat scale. In addition, rather than niche shift, birds actually extended their niche breadth. Nest site selection may constrain bird distribution into a realized niche, smaller than their fundamental niche. Despite the scale differences in habitat use, the similar species diversity patterns between seasons suggest that both winter and summer birds would benefit from conservation of scrub patches larger than 50 ha, and enrichment of foliage layers within the planted forests.