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

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

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

Human activities link fruit bat presence to Ebola virus disease outbreaks

1. A significant link between forest loss and fragmentation and human outbreaks of Ebola virus disease (EVD) has been documented. Deforestation may alter the natural circulation of viruses as well as change the composition, abundance, behaviour and possibly viral exposure of reservoir species. This in turn might increase contact between infected animals and humans. Fruit bats of the family Pteropodidae have been suspected as reservoirs of the Ebola virus. Though there is no solid proof that fruit bats cause human EVD outbreaks, this group of animals have been intermittently infected with the Ebola virus. 2. Our study investigates whether human activities positively affect African fruit bat species’ ranges and whether their ranges are linked to EVD outbreaks, in turn favoured by deforestation. 3. We use species observation records for the 20 fruit bat species found in Africa to determine factors affecting their distribution in two geographical scenarios: 1) the African continent; and 2) inside the predicted Ebola virus range. We do this by employing a hypothetico-deductive approach based on favourability modelling. 4. Our models point to clear associations between human activities and fruit bat distributions that may help scientists understand the anthropogenic settings that could cause the Ebola virus to jump from animals to humans. 5. We show that fruit bat distributions are linked to human activities throughout Africa and particularly within the region where the Ebola virus occurs. More specifically, the areas where human activities favour the presence of five fruit bat species (four of which had recorded seropositive individuals) overlap with EVD outbreak areas, themselves favoured by deforestation. 6. Our work is a useful first step to further investigate the networks and pathways that may lead to an EVD outbreak. The modelling framework we employ here can be employed for other emerging infectious diseases