Effects of forest fragmentation on the vertical stratification of neotropical bats

Vertical stratification is a key component of the biological complexity of rainforests. Understanding community- and species-level responses to disturbance across forest strata is paramount for evidence-based conservation and management. However, even for bats, known to extensively explore multiple layers of the complex three-dimensional forest space, studies are biased towards understory-based surveys and only few assessments of vertical stratification were done in fragmented landscapes. Using both ground and canopy mist-nets, we investigated how the vertical structure of bat assemblages is influenced by forest fragmentation in the experimentally fragmented landscape of the Biological Dynamics of Forest Fragments Project, Central Amazon, Brazil. Over a three year-period, we captured 3077 individuals of 46 species in continuous forest (CF) and in 1, 10 and 100 ha forest fragments. In both CF and forest fragments, the upper forest strata sustained more diverse bat assemblages than the equivalent understory layer, and the midstory layers had significantly higher bat abundance in fragments than in CF. Artibeus lituratus and Rhinophylla pumilio exhibited significant shifts in their vertical stratification patterns between CF and fragments (e.g. R. pumilio was more associated with the upper strata in fragments than in CF). Altogether, our study suggests that fragmentation modulates the vertical stratification of bat assemblages

Grazing improves habitat suitability for many ground foraging birds in Mediterranean wooded grasslands

Wooded grasslands, usually grazed, cover vast areas in Southern Europe and Northern Africa. They host rich resident bird communities and, in winter, receive large numbers of migrants from Central and Northern European woodlands. Many species are partly or entirely dependent on ground foraging, and since in winter food is often the most limiting factor for birds, maintaining suitable ground habitat is crucial. To study how grazing influences suitability of winter ground habitat for birds, we carried out an experiment in a wooded grassland in Southern Iberia, whereby grazing was controlled in 12 purposely fenced two-hectare plots (4 x 15 sheep/ha, 4 x 3 sheep/ha and 4 x no grazing). We quantified ground habitat features, food abundance and intensity of use by ground-foraging birds in each of these 12 plots. In addition, we made focal observations of birds feeding on the ground and compared the habitat of 1m2 foraging patches with those of nearby control patches. We found that virtually all birds prefer to forage in patches with short ground vegetation and high food abundance. Measurements of these parameters in the experimental plots showed that while grazing shortens vegetation it decreases food availability, and thus has opposing effects on important determinants of habitat suitability. Nevertheless, the numbers of birds foraging in the plots indicate that, overall, grazing benefits the assemblage of ground-feeding birds, presumably because for most species the advantages of foraging in less cluttered habitats more than compensate the lower abundance of prey. However, arboreal bird species that make short foraging forays to the ground had lower numbers in grazed plots. Most bird species that forage on the ground benefited from grazing, and although they can forage under a broad range of grazing levels, some showed clear preferences along the gradient of grazing intensity. Such preferences should be taken into consideration by managers. In general, grazing should be maintained at a level sufficient to open up ground vegetation, increasing the area occupied by patches of short vegetation, in which almost all bird species prefer to forage. At moderate levels, grazing is thus a valuable management tool to promote winter bird habitat quality in Mediterranean wooded grasslands, while increasing the economic value of these threatened landscapes

Secondary forest buffers the effects of fragmentation on aerial insectivorous bat species following 30 years of passive forest restoration

Passive forest restoration can buffer the effects of habitat loss on biodiversity. We acoustically surveyed aerial insectivorous bats in a whole-ecosystem fragmentation experiment in the Brazilian Amazon over a 2-year period, across 33 sites, comprising continuous old-growth forest, remnant fragments, and regenerating secondary forest matrix. We analyzed the activity of 10 species/sonotypes to investigate occupancy across habitat types and responses to fragment size and interior-edge-matrix (IEM) disturbance gradients. Employing a multiscale approach, we investigated guild (edge foragers, forest specialists, flexible forest foragers, and open space specialists) and species-level responses to vegetation structure and forest cover, edge, and patch density across six spatial scales (0.5–3 km). We found species-specific habitat occupancy patterns and nuanced responses to fragment size and the IEM disturbance gradient. For example, Furipterus horrens had lower activity in secondary forest sites and the interior and edge of the smallest fragments (1 and 10 ha) compared to continuous forest, and only two species (Pteronotus spp.) showed no habitat preference and no significant responses across the IEM and fragment size gradients. Only the Molossus sonotype responded negatively to vegetation structure. We uncovered no negative influence of forest cover or edge density at guild or species-level. Our results indicate that reforestation can buffer the negative effects of fragmentation and although these effects can still be detected in some species, generally aerial insectivorous bats appear to be in recovery after 30 years of passive forest restoration. Our findings reinforce the need to protect regenerating forests while conserving vast expanses of old-growth forest

Effects of land‐use change on functional and taxonomic diversity of Neotropical bats

Human land-use changes are particularly extensive in tropical regions, representing one of the greatest threats to terrestrial biodiversity and a key research topic in conservation. However, studies considering the effects of different types of anthropogenic disturbance on the functional dimension of biodiversity in human-modified landscapes are rare. Here, we obtained data through an extensive review of peer-reviewed articles and compared 30 Neotropical bat assemblages in well-preserved primary forest and four different human-disturbed habitats in terms of their functional and taxonomic diversity. We found that disturbed habitats that are structurally less similar to primary forest (pasture, cropland and early-stage secondary forest) were characterized by a lower functional and taxonomic diversity, as well as community level-functional uniqueness. These habitats generally retained fewer species that perform different ecological functions compared to higher-quality landscape matrices, such as agroforestry. According to functional trait composition, different bat ensembles respond differently to landscape change, negatively affecting mainly gleaning insectivorous bats in pasture, narrow-range species in cropland, and heavier animalivorous bats in secondary forest. Although our results highlight the importance of higher-quality matrix habitats to support elevated functional and taxonomic bat diversity, the conservation of bat species that perform different ecological functions in the mosaic of human-modified habitats also depends on the irreplaceable conservation value of well-preserved primary forests. Our study based on a pooled analysis of individual studies provides novel insights into the effects of different human-modified habitats on Neotropical bat assemblages

Functional recovery of Amazonian bat assemblages following secondary forest succession

Regenerating forests occupy large areas in the tropics, mostly as a result of deforestation for livestock and agriculture, followed by land abandonment. Despite the importance of regenerating secondary forests for tropical biodiversity conservation, studies of temporal effects of matrix regeneration on species responses in fragmented landscapes are scarce. Here, we used an Amazonian whole-ecosystem fragmentation experiment to investigate how changes in matrix quality over time through secondary forest regeneration affect bat assemblages from a functional perspective. We found that forest regeneration in the matrix positively affected functional α diversity, as well as species- and community-level functional uniqueness, reflecting an increase of species that perform different ecological functions in secondary forest over time. According to functional trait composition, animalivorous species showed the clearest signs of recovery associated with matrix regeneration. Consequently, between-period differences in functional β-diversity were highest in secondary forest compared to fragments and continuous forest, determined mainly by trait gains. However, ~ 30 years of secondary forest regeneration were not sufficient for the functional recovery of bat assemblages to levels observed in continuous forest. Restoring degraded habitats while protecting primary forest will be an important strategy for safeguarding high functional diversity of bats and their vital contributions to ecosystem functioning in fragmented tropical landscapes

The road to functional recovery : temporal effects of matrix regeneration on Amazonian bats

Across the tropics, vast deforested areas are undergoing forest regeneration due to land abandonment. Although secondary forest is an expanding type of landscape matrix that has been shown to buffer some of the negative consequences of forest loss and fragmentation on taxonomic diversity, little is known in this regard about the functional dimension of biodiversity. We took advantage of an ecosystem-wide fragmentation experiment to investigate longer term changes in functional diversity of a mega-diverse Amazonian bat assemblage associated with regrowth development in the matrix. We found that matrix regeneration affected several facets of bat functional diversity in secondary forest over time, increasing functional α diversity, species- and community-level functional uniqueness, altering functional trait composition, and resulting in functional β-diversity changes via trait gains. However, approximately 30 years of matrix regeneration were insufficient for functional diversity to recover to the same levels as in continuous forest. Our results suggest that a combination of natural, human-assisted, and active restoration is likely to be the most successful strategy for restoring functional biodiversity of bats in human-modified tropical landscapes, a finding that most likely also applies to many other taxa

Design matters : an evaluation of the impact of small man-made forest clearings on tropical bats using a before-after-control-impact design

In recent years, large clearings (>1000 ha) accounted for gradually smaller amounts of total annual deforestation in the Brazilian Amazon, whereas the proportion of small clearings (<50 ha) nowadays represents more than 80% of annual deforestation. Despite the ubiquity of small clearings in fragmented Amazonian landscapes, most fragmentation research has focused on the effects of large-scale deforestation, leading to a poor understanding of the impacts of smaller barriers on Amazonian vertebrates. We capitalized on the periodical re-isolation of experimental forest fragments at the Biological Dynamics of Forest Fragments Project in the Central Amazon as a before-after-control-impact experiment to investigate the short-term effects of small clearings on bat assemblages. Over the course of three years we sampled six control sites in continuous forest, the interiors and edges of eight forest fragments as well as eight sites in the surrounding matrix. Sampling took place both before and after the experimental manipulation (clearing of a 100 m wide strip of regrowth around each fragment), resulting in ~4000 bat captures. Species were classified as old-growth specialists and habitat generalists according to their habitat affinities and a joint species distribution modeling framework was used to investigate the effect of fragment re-isolation on species occupancy. Following fragment re-isolation, species richness declined in all habitats other than fragment edges and, although responses were idiosyncratic, this decline was more pronounced for forest specialist than for generalist species. Additionally, fragment re-isolation led to a reduction in the similarity between assemblages in modified habitats (fragment interiors, edges and matrix) and continuous forest. Sampling of controls in continuous forest both prior to and after reisolation revealed that much of the variation in bat species occupancy between sampling periods did not arise from fragment re-isolation but rather reflected natural spatiotemporal variability. This emphasizes the need to sample experimental controls both before and after experimental manipulation and suggests caution in the interpretation of results from studies in which the effects of habitat transformations are assessed based solely on data collected using space-for-time substitution approaches

Design matters : an evaluation of the impact of small man-made forest clearings on tropical bats using a before-after-control-impact design

In recent years, large clearings (>1000 ha) accounted for gradually smaller amounts of total annual deforestation in the Brazilian Amazon, whereas the proportion of small clearings (<50 ha) nowadays represents more than 80% of annual deforestation. Despite the ubiquity of small clearings in fragmented Amazonian landscapes, most fragmentation research has focused on the effects of large-scale deforestation, leading to a poor understanding of the impacts of smaller barriers on Amazonian vertebrates. We capitalized on the periodical re-isolation of experimental forest fragments at the Biological Dynamics of Forest Fragments Project in the Central Amazon as a before-after-control-impact experiment to investigate the short-term effects of small clearings on bat assemblages. Over the course of three years we sampled six control sites in continuous forest, the interiors and edges of eight forest fragments as well as eight sites in the surrounding matrix. Sampling took place both before and after the experimental manipulation (clearing of a 100 m wide strip of regrowth around each fragment), resulting in ~4000 bat captures. Species were classified as old-growth specialists and habitat generalists according to their habitat affinities and a joint species distribution modeling framework was used to investigate the effect of fragment re-isolation on species occupancy. Following fragment re-isolation, species richness declined in all habitats other than fragment edges and, although responses were idiosyncratic, this decline was more pronounced for forest specialist than for generalist species. Additionally, fragment re-isolation led to a reduction in the similarity between assemblages in modified habitats (fragment interiors, edges and matrix) and continuous forest. Sampling of controls in continuous forest both prior to and after reisolation revealed that much of the variation in bat species occupancy between sampling periods did not arise from fragment re-isolation but rather reflected natural spatiotemporal variability. This emphasizes the need to sample experimental controls both before and after experimental manipulation and suggests caution in the interpretation of results from studies in which the effects of habitat transformations are assessed based solely on data collected using space-for-time substitution approaches

Interplay between local and landscape-scale effects on the taxonomic, functional and phylogenetic diversity of aerial insectivorous Neotropical bats

Context Human-modified landscapes are globally ubiquitous. It is critical to understand how habitat loss and fragmentation impact biodiversity from both a local habitat context and landscape-scale perspective to inform land management and conservation strategies. Objectives We used an experimentally fragmented landscape in the Brazilian Amazon to investigate variation in aerial insectivorous bat diversity in response to local habitat and wider landscape characteristics, applying a multiscale approach. Methods We conducted bat acoustic surveys at 33 sites, comprising old secondary forests and fragments of primary forest. Taxonomic, functional and phylogenetic diversity facets were calculated within a Hill numbers framework. We analysed responses to fragment size, interior-edge-matrix gradients, as well as local vegetation structure, continuous forest cover, edge density and patch density across five spatial scales (0.5 km - 3 km) surrounding detector locations. Results Compared with continuous forest, secondary forest matrix around the smallest fragments harboured lower diversity. The overall negative effect of the matrix became less pronounced with increasing fragment size. In contrast, forest edges generally contained higher taxonomic, functional and phylogenetic diversity. We found subtle scale-sensitive associations for FD, responding positively to forest cover (at the 1 km scale) and negatively to edge (1 km scale) and patch density (2.5 km scale). Conclusions Despite a low-contrast matrix of tall secondary forest surrounding fragments after ~30 years of forest recovery, aerial insectivorous bat diversity is not comparable to continuous primary forest. Assemblage functional diversity responds to compositional and configurational landscape characteristics at scales deserving further evaluation at guild and species level

Edge effects and vertical stratification of aerial insectivorous bats across the interface of primary-secondary Amazonian rainforest

Edge effects, abiotic and biotic changes associated with habitat boundaries, are key drivers of community change in fragmented landscapes. Their influence is heavily modulated by matrix composition. With over half of the world’s tropical forests predicted to become forest edge by the end of the century, it is paramount that conservationists gain a better understanding of how tropical biota is impacted by edge gradients. Bats comprise a large fraction of tropical mammalian fauna and are demonstrably sensitive to habitat modification. Yet, knowledge about how bat assemblages are affected by edge effects remains scarce. Capitalizing on a whole-ecosystem manipulation in the Central Amazon, the aims of this study were to i) assess the consequences of edge effects for twelve aerial insectivorous bat species across the interface of primary and secondary forest, and ii) investigate if the activity levels of these species differed between the understory and canopy and if they were modulated by distance from the edge. Acoustic surveys were conducted along four 2-km transects, each traversing equal parts of primary and ca. 30-year-old secondary forest. Five models were used to assess the changes in the relative activity of forest specialists (three species), flexible forest foragers (three species), and edge foragers (six species). Modelling results revealed limited evidence of edge effects, except for forest specialists in the understory. No significant differences in activity were found between the secondary or primary forest but almost all species exhibited pronounced vertical stratification. Previously defined bat guilds appear to hold here as our study highlights that forest bats are more edge-sensitive than edge foraging bats. The absence of pronounced edge effects and the comparable activity levels between primary and old secondary forests indicates that old secondary forest can help ameliorate the consequences of fragmentation on tropical aerial insectivorous bats