Bats in a fragmented landscape: Species composition, diversity and habitat interactions in savannas of Santarém, Central Amazonia, Brazil

Species response to forest fragmentation may strongly vary according to ecological requirements, shape, spatial configuration and connectivity of fragments, and the structure of the surrounding matrix. Bats are natural candidates for studies on forest fragmentation. However, the conclusions of such studies are often contradictory. We present the results of a study on the effects of forest fragmentation on bat communities in Alter do Chão, Santarém area, in Central Amazonia, Brazil, an area composed by forests and fragments surrounded by savanna-like vegetation. We compared the species composition between habitats, tested the effect of size, shape and density of trees of forested areas on the presence species and their relative frequency of captures, and investigated the savannas as an ecological barrier for the dispersion of bats. With an effort of 5678 mistnet-hours, we captured 3740 bats (64 species). Multidimensional Scaling indicated no strong separation between the habitats sampled, however, savanna sites were grouped distinctly of fragments and forest sites. Multiple linear regressions indicated no significant correlation between the number of bat species recorded and the size, shape or tree density in forest sites and fragments. There was a significant correlation between the number of captures and the variables tested, explained by the shape of the sites, but not by size or tree density. In general, sites with bigger shape indices had fewer captures. Ten of the 20 most captured species showed no significant difference between the three habitats. The bat communities in Alter do Chão were not strongly affected by forest fragmentation, and the savannas did not appear to act as an ecological barrier to bats.     

Conservation value of forest fragments to Palaeotropical bats

Forested landscapes in Southeast Asia are becoming increasingly fragmented, making this region a conservation and research priority. Despite its importance, few empirical studies of effects of fragmentation on biodiversity have been undertaken in the region, limiting our ability to inform land-use regimes at a time of increased pressure on forests. We estimated the biodiversity value of forest fragments in peninsular Malaysia by studying fragmentation impacts on insectivorous bat species that vary in dependence of forest. We sampled bats at seven continuous forest sites and 27 forest fragments, and tested the influence of fragment isolation and area on the abundance, species richness, diversity, composition and nestedness of assemblages, and the abundance of the ten most common species. Overall, isolation was a poor predictor of these variables. Conversely, forest area was positively related with abundance and species richness of cavity/foliage-roosting bats, but not for that of cave-roosting or edge/open space foraging species. The smallest of fragments (<150 ha) were more variable in species composition than larger fragments or continuous forest, and larger fragments retained substantial bat diversity, comparable to continuous forest. Some fragments exhibited higher bat abundance and species richness than continuous forest, though declines might occur in the future because of time lags in the manifestation of fragmentation effects. Our findings suggest that fragments >300 ha contribute substantially to landscape-level bat diversity, and that small fragments also have some value. However, large tracts are needed to support rare, forest specialist species and should be the conservation priority in landscape-level planning. Species that roost in tree cavities or foliage may be more vulnerable to habitat fragmentation than those that roost in caves.     

Location of roosts of the lesser long-eared bat Nyctophilus geoffroyi and Gould's wattled bat Chalinolobus gouldii in a fragmented landscape in south-eastern Australia

The location of roosts of two species of vespertilionid bats, the lesser long-eared bat (Nyctophilus geoffroyi) and Gould’s wattled bat (Chalinolobus gouldii), was investigated in a remnant vegetation-farmland mosaic and adjacent floodplain forest in south-eastern Australia. A total of 45 individuals of N. geoffroyi and 27 C. gouldii were fitted with radio transmitters, which resulted in the location of 139 and 89 roosts respectively. Selection of roosting habitat showed both inter- and intra-specific differences. At the landscape level, locations of roosts used by male and female N. geoffroyi differed markedly. Most males roosted in the farmland mosaic within 3 km (mean 1.9±2.9 km) of where they were captured while foraging in remnant vegetation. In contrast, roost sites of females were predominantly in the floodplain forest, 6-12 km from their capture site in farmland (mean for all females, 6.7±2.9 km). All maternity roosts were in the extensive floodplain forest, 4-10 km from foraging areas. Distances moved by this species were greater than previously recorded and further than predicted by flight morphology. Most C. gouldii roosted in the floodplain forest, 4-10 km from their capture site (6.9±1.6 km). Within the floodplain forest, both species roosted in areas that had higher densities of hollow-bearing trees than generally available; dead hollow-bearing trees for N. geoffroyi and large, live trees for C. gouldii. The scale of movements undertaken by these species means that they can obtain resources from multiple landscape elements. Both species used different habitats for roosting and foraging despite the energetic costs of commuting relatively large distances. Conservation of bat populations in rural environments needs to be considered at the landscape scale, with particular attention to identifying landscape elements that provide key resources.     

Abundance, body size and movement patterns of a tropical treefrog in continuous and fragmented forests in the Brazilian Amazon

Phyllomedusa tarsius is a hylid frog that breeds in ponds located in a range of habitats from continuous forests to severely disturbed matrix habitats in Central Amazon. During three reproductive seasons, we followed the movement patterns, measured body size and registered abundance and residency time of this species in five habitats: pasture, Vismia regrowth, Cecropia regrowth, 1 and 10 ha forest fragments, and continuous forest. The frog captures were strongly correlated with rainfall in all disturbed habitats, but not in continuous forest, probably because individuals respond more directly to rainfall patterns in more open habitats. Males in disturbed habitats were smaller than those found in continuous forest, perhaps as a result of differences in habitat quality. Males remained at sites for longer periods in fragments and continuous forest compared to matrix habitats. Here again, the quality and suitability of breeding sites in matrix habitats may be lower than in continuous forest resulting in the need for more movement. We found bigger subpopulations in matrix habitat ponds and a higher percentage of individuals moving among them when compared to continuous forest ponds. Constant movement of individuals among disturbed subpopulations decreases population isolation and increases interbreeding among different subpopulations. No movement between individuals from continuous forest ponds and disturbed habitats was observed. This suggests that the disturbed habitat subpopulations are not acting as sink subpopulations for continuous forest populations. The maintenance of individuals in fragments is more dependent on local conditions for reproduction and on subpopulations in the matrix habitat than on recruitment of individuals from populations in continuous forest.     

The importance of cenotes in conserving bat assemblages in the Yucatan, Mexico

In the Yucatan, cenotes or water sinkholes formed by the dissolution of limestone and surrounded with a characteristic dense layer of heterogeneous vegetation, are the main sources of water for plant and animal assemblages. We investigated their importance to bats by comparing the assemblage structure and diversity between tropical forest and pastureland, with and without cenotes. We set ground level mist nets, sub-canopy nets and harp traps for 96 nights and caught 2819 bats of 26 species and six families. Forest had a greater bat species diversity and abundance than pastureland, as well as rare and threatened species. In forest, sites with and without cenotes had a similar bat assemblage structure and diversity, whereas in pastureland there was a greater species diversity and abundance and number of rare and threatened species at cenotes than sites without cenotes. Chrotopterus auritus and Micronycteris schmidtorum, both threatened in Mexico, and Eptesicus furinalis, were exclusively captured at cenotes. We suggest that it may be the vegetation around cenotes rather than the water bodies per se that attracts many bat species in pastureland. Cenotes are threatened by nitrate contamination from untreated domestic and animal waste and fertilizers and the removal of vegetation to allow livestock access to water. In Yucatan’s agricultural landscapes, protection of cenotes and their characteristic vegetation is key not only for bat conservation but also because they represent potential seed sources for the regeneration of the lowland forest, in which bats play key roles as pollinators and seed dispersers.     

Influence of matrix habitats on the occurrence of insectivorous bird species in Amazonian forest fragments

The influence of matrix habitats on the occurrence of seven understory insectivorous bird species in forest fragments was examined at the Biological Dynamics of Forest Fragments Project, Manaus, Amazonia. Playback techniques were used to detect individuals in the continuous primary forest, fragments of 1, 10 and 100 ha, and in two matrix habitats of secondary forest dominated by Vismia spp. and Cecropia spp. Sampling points in secondary forest areas were distributed at 50, 250 and 500 m from the continuous forest. Using G-tests of frequency distribution, species occurrences were compared in the following ways: (a) continuous forest vs. fragments; (b) continuous forest vs. second growth forest; (c) in second growth at different distances from continuous forest. Species were divided into three categories according to their sensitivity to the fragmentation process. Highly sensitive species (Cyphorhinus arada, Hylophilus ochraceiceps and Thamnomanes ardesiacus) did not occur in small fragments or in matrix habitats. Moderately sensitive species (Formicarius colma and T. caesius) occurred in small fragments and utilized infrequently matrix habitats. Positively affected species (Percnostola rufifrons and Hypocnemis cantator) were frequently detected in small fragments and all matrix habitats. Distances from the continuous forest did not influence the frequency of species occurrences in the secondary forest areas. Species were more frequent in small fragments surrounded by Cecropia spp. than by Vismia spp. Our results support the idea that the maintenance of species in small fragments may depend on their ability to use the matrix, and that increasing the permeability of the matrix may be an option to lessen the effects of forest fragmentation.     

Intra-specific effects of forest loss on the distribution of the forest-dependent northern long-eared bat (Myotis septentrionalis)

Forest fragmentation by the expansion of human activities is acknowledged as an important factor driving declines of forest species worldwide and quantitatively characterizing this trend is essential to understanding the response of populations of forest-dependent species. To examine the impacts of forest cover loss and changes in composition on forest-dependent bats, we examined the effects of these factors on the distribution of the northern long-eared bat (Myotis septentrionalis) in a case study. We surveyed forest fragments in the forest-agricultural landscape of Prince Edward Island, Canada, to assess the occupancy of bats in fragments. Logistic regression and multimodel inference using Akaike’s information criteria were used to identify potentially important predictor variables influencing the distribution of the northern long-eared bat at the fragment and landscape level and quantified their effects. A compositional variable, area of deciduous stands, was found to be the best predictor of the presence of M. septentrionalis. We found the effects of fragmentation were greatest at the fragment level for females and at the landscape level for males. This case study emphasizes the importance of examining intra-specific resource selection in how it affects the response of a forest-dependent species to fragmentation.     

Bird species abundance–occupancy patterns and sensitivity to forest fragmentation: Implications for conservation in the Brazilian Atlantic forest

Developing a predictive theory for species responses to habitat fragmentation is a large, complex challenge in conservation biology, and meeting this challenge likely requires tailoring predictions to specific habitats and taxa. We evaluate the effects of fragmentation on forest birds living in three distinct forest ecosystems found in Brazilian Atlantic forest: seasonal semi-deciduous forest (SF), mixed rain forest (MF), and dense rain forest (DF). We test the hypotheses that (1) bird species most prevalent in SF (relative to other habitat types) will be least vulnerable to population declines in fragmented SF, and (2) species with stronger affiliations with DF or MF will be relatively more sensitive to fragmentation in SF. Using an exploratory statistical technique called “Rank Occupancy–Abundance Profiles (ROAPs),” we compared distribution and abundance of birds among large “continuous” areas of each forest type, then compared abundances in continuous SF forests with patterns of abundance in small fragments of SF, where edge effects could play a marked role in population dynamics. Overall, 39 species showed substantially lower local abundance, occupancy, or both in SF fragments versus continuous SF. As predicted, a higher proportion of bird species associated with DF appeared sensitive to fragmentation in SF; by contrast, species most abundant in SF and MF were similarly abundant in fragmented SF. Our study demonstrates how quantifying distribution and abundance in diverse habitats may enhance managers’ ability to incorporate species-specific responses to human disturbances in their conservation plans, and points out ways that even small reserves may have significant conservation value.     

Dramatic decline in bat species richness in Singapore, with implications for Southeast Asia

Literature searches and recent surveys of the bat fauna of Singapore indicate that of the 24 species of Microchiroptera and six species of Megachiroptera documented for this small equatorial island just 15 and 5, respectively, are still present. These recorded declines in chiropteran species richness almost certainly understate the true losses as extensive land transformation/habitat loss (>95%) and biota loss occurred early in Singapore’s colonial history before comprehensive surveys of bats were made. In an effort to reconstruct the pre-settlement bat fauna, we inferred an upper bound of pre-settlement species richness using a checklist from a well-known bat assemblage in neighbouring Peninsular Malaysia, and a lower bound based on species common to Peninsular Malaysia, Borneo and Sumatra. The Singapore records were compared with these two species list predictions. Based on this analysis, we infer that between 60 and 72 species would have inhabited Singapore before 1819. We also estimate that between 33% (based on the confirmed inventory) and 72% of the species (based on the upper-bound estimate of species richness) are now locally extinct. For Microchiroptera the data suggest that the documented local extinction rate of 38% may project to between 69% and 75%. Forest-dependant bats are particularly affected and comprise a much lower proportion of the bat fauna than is seen in intact forest in Peninsular Malaysia. All hipposiderids and 40% of the documented rhinolophid taxa have been lost and almost half (6) of the surviving microchiropteran species are locally endangered. Projected local extinction rates for Megachiroptera raise the documented value (17%) to about 60%, with most of the survivors being widespread species known to forage in cultivated or secondary forest habitats or to commute long distances between fragmented resources. The dramatic decline in Singapore bat species richness and a concomitant change in chiropteran guild and trophic structure (Microchiroptera vs. Megachiroptera) reflect patterns of diversity change seen elsewhere in the region in response to loss of forest habitat. In Singapore, the decline in diversity (species richness and abundances) for both mega- and microbats may also relate to urbanisation and deliberate or accidental destruction of bats and their roost sites in a land that has one of the highest human population densities on the planet. Although these losses (actual and inferred) represent a microcosm of mainly local extinctions, a wider geographical extrapolation over the 21st century indicates that heavy deforestation in progress in Southeast Asia might be expected to lead to extinction of many bat taxa, with upper-bound estimates of regional species losses exceeding 40% and global extirpation anticipated for at least 23% of Southeast Asia’s bat fauna by 2100.     

Phyllostomid bat diversity in a variegated coffee landscape

We examined bat diversity at two different spatial scales: habitat and matrix, in the Quindío coffee region in Colombia. Habitats were: forest, shaded coffee and associated coffee; and matrices were: associated coffee (M1) and shaded coffee (M2). Three sampling sites from each type of habitat were located at each matrix. The forest areas of the Quindío region are severely fragmented and less structurally complex than coffee patches. The shaded coffee habitat had patches that were larger and more complex. In spite of limited patch size and lower complexity, the forest remnants were those with greatest species richness and demonstrated clear similarities even between the two matrices. This was not observed in coffee plantations, neither in associated coffee nor shaded coffee. On the landscape scale, M2 showed lower β diversity and greater edge density (ED) than M1. This fact explains that greater connectivity between different habitats exists in M2 than in M1. Our results suggest that production and conservation are compatible, as maintenance of forest remnants in a mosaic structure by landowners of the vegetation is sufficient to conserve phyllostomid bats at landscape level.     

The importance of gallery forests in the tropical lowlands of Costa Rica for understorey forest birds

Rivers and streams lined by narrow forest strips are common in the lowland countryside of south-western Costa Rica. We studied the importance of these gallery forests for understorey birds, especially forest species. Using mist-nets, we captured 1110 birds belonging to 90 species between June and September 2007 at 16 sites spread equally over four habitat types: forest interior, forest margin, gallery forest connected to closed forest, and isolated gallery forest. Though isolated gallery forests had the greatest number of expected species in total, they supported the lowest number of forest-specialist species, lower even than connected gallery forests. Granivorous birds showed an increase from forest to isolated gallery forest, all other feeding guilds however showed no change. The studied habitats differed from each other in their faunal composition. Assemblages of three species groups categorised by their habitat preferences (forest specialists, forest generalists, non-forest species) showed nested distribution patterns across the four habitats. There was no significant difference in the proportion of birds with brood patches or of recaptures across the studied habitat types, leading to the assumption that birds not only use gallery forests for movement and foraging but also for reproduction. Though of limited conservation value for most forest understorey birds, at least for a small fraction of forest species gallery forests constitute an important secondary habitat. More significantly, they can form corridors or stepping stones that allow movements within the matrix of human-dominated habitats, and represent an important landscape component benefiting total bird species richness in the Costa Rican countryside.     

Relating bat species presence to simple habitat measures in a central Appalachian forest

We actively sampled the bat community at 63 sites using detection and non-detection metrics on the Fernow Experimental Forest (FEF) in the central Appalachians of West Virginia using Anabat acoustical equipment May-June 2001-2003 to relate species presence to simple habitat measures such as proximity to riparian areas, forest canopy cover, forest canopy gap width, and forest canopy height. We acoustically detected eight species on the FEF, including the endangered Myotis sodalis. The presence of Lasiurus cinereus, M. lucifugus, M. sodalis, and Pipistrellus subflavus was associated more with riparian areas than upland areas. Both univariate comparisons and multiple logistic regression modeling showed that the probability that clutter-adapted foraging species such as M. septentrionalis and M. sodalis would be detected was greater as forest canopy cover increased or forest canopy gap size decreased, whereas the opposite was true for open-adapted foraging species such as Eptesicus fuscus and L. cinereus. The overall proportion of unidentifiable bat echolocation sequences to those identified to species was related to upland sites with increasing forest canopy cover indicating some sampling bias between cluttered and uncluttered habitats. However, given sufficient sample points, bat community surveys using acoustical detection show the ability to quickly develop generalized habitat associations for rugged areas such as the central Appalachians where traditional mist-net survey efforts often are logistically difficult and are lacking in scope. Moreover, these acoustical surveys also could lend themselves to species-specific predictive mapping of foraging habitat as well as allowing researchers to formulate testable hypotheses about detailed bat habitat relationships to be definitively tested with radio-telemetry techniques.     

Bat diversity in montane rainforest and shaded coffee under different management regimes in southeastern Chiapas, Mexico

Differences in alpha and beta bat diversity among montane rainforest and five shaded coffee plantations under different management regimes, as well as some environmental factors and vegetation parameters influencing bat richness, were evaluated for the first time in southeastern Chiapas, Mexico. In each site, bats were captured every 2 months from March 2004 to July 2005, with six mist-nets, during two nights, using the capture-recapture method. We captured 2970 individuals of 43 bat species. Montane rainforest had the greatest alpha diversity (H′ = 2.681; n = 37), whereas alpha diversity was similar among coffee plantations (H′ = 2.229-2.364; n = 23-26). The number of frugivorous and nectarivorous species was similar among the sites; the greatest exchange in species composition (beta diversity) occurred for insectivorous bats, which reduce their number in coffee plantations as pesticides are incorporated. Bat richness species was significantly related to the number of vegetation strata, height, and cover of trees. We suggest that coffee plantations could act as corridors, facilitating connection among different elements of the landscape in the Sierra Madre de Chiapas for some frugivorous and nectarivorous bats.     

Scattered trees in rural landscapes: foraging habitat for insectivorous bats in south-eastern Australia

Scattered trees and small patches of vegetation among farmland are typical of rural landscapes throughout the world, often comprising a significant proportion of remaining habitats for native fauna. Insectivorous bats can use such isolated resources owing to the high mobility of most species, but little is known of the relationship between bats and tree density, or of the impact of incremental loss of trees in the landscape. Bats were surveyed at 30 sites in south-eastern Australia, in five habitat categories representing a range of tree densities from remnant woodland blocks (>35 trees/ha) to sparsely scattered trees (<1 tree/ha), and open paddocks devoid of trees. Sampling was undertaken by using harp traps and ultrasonic bat detectors. The abundance of potential arthropod prey was assessed by using light traps. Eleven species of insectivorous bats were recorded and bats were widespread in all habitat categories; all species were recorded around scattered trees. Overall activity, as revealed by detectors, did not differ significantly between the four treed categories, but in open paddocks there was a lower level of activity and a different community composition. However, a regression model revealed a significant quadratic relationship between activity of bats and tree density, with highest activity at 20-30 trees/ha. All species were recorded in open paddocks, but for eight of the ten species this represented <7% of their total activity recorded across all habitat categories. For six species, significant positive correlations were found between activity levels and the abundance of potential arthropod prey. In contrast to bat detectors, trapping results showed a significant difference in overall abundance of bats between wooded habitats, with decreased abundance as tree density declined. This study highlights the value of scattered trees as foraging habitat for bats, and emphasizes that, in rural land mosaics, such small and isolated habitat components have value for the conservation of biodiversity, and require greater recognition and protection.     

Habitat selection of endangered and endemic large flying-foxes in Subic Bay, Philippines

Large flying-foxes in insular Southeast Asia are the most threatened of the Old World fruit bats due to high levels of deforestation and hunting and effectively little local conservation commitment. The forest at Subic Bay, Philippines, supports a rare, large colony of vulnerable Philippine giant fruit bats (Pteropus vampyrus lanensis) and endangered and endemic golden-crowned flying-foxes (Acerodon jubatus). These large flying-foxes are optimal for conservation focus, because in addition to being keystone, flagship, and umbrella species, the bats are important to Subic Bay’s economy and its indigenous cultures. Habitat selection information streamlines management’s efforts to protect and conserve these popular but threatened animals. We used radio telemetry to describe the bats’ nighttime use of habitat on two ecological scales: vegetation and microhabitat. The fruit bats used the entire 14,000 ha study area, including all of Subic Bay Watershed Reserve, as well as neighboring forests just outside the protected area boundaries. Their recorded foraging locations ranged between 0.4 and 12 km from the roost. We compared the bats’ use to the availability of vegetative habitat types, riparian areas, and bat trees. The fruit bats’ locations showed a preference for undisturbed forest types and selection against disturbed and agricultural areas. Bat locations also showed selection for particular fruiting/flowering bat trees. The bats showed strong preference for riparian areas; locations were in riparian areas over four times more than expected. From these results we recommend that management focus flying-fox conservation efforts on undisturbed forest and riparian areas.     

Habitat structure and proximity to forest edge affect the abundance and distribution of forest-dependent birds in tropical coastal forests of southeastern Madagascar

Despite the fact that Madagascar is classified a biological `hotspot' due to having both high levels of species endemism and high forest loss, there has been no published research on how Madagascan bird species respond to the creation of a forest edge or to degradation of their habitat. In this study, we examined how forest bird communities and different foraging guilds were affected by patch habitat quality and landscape context (forest core, forest edge and matrix habitat) in the threatened littoral forests of coastal southeastern Madagascar. We quantified habitat use and community composition of birds by conducting 20 point counts in each landscape contextual element in October and November 2002. We found that littoral forest core habitats had significantly (p<0.01) more bird species than forest edge and matrix habitats. Thirty-one (68%) forest dependent species were found to be edge-sensitive. Forest edge sites had fewer species, and a higher representation of common species than forest interior sites. Twenty-nine species were found in the matrix habitat, and the majority of matrix-tolerant forest species had their greatest abundance within littoral forest edge habitats. Guild composition also changed with landscape context. Unlike other tropical studies with which we are familiar, we found that frugivorous species were edge-sensitive while sallying insectivores were edge-preferring. The majority of canopy insectivores (n=15, 88%), including all six endemic vanga species, were edge-sensitive. When habitat quality was assessed, the distributions of nine edge-sensitive species were significantly (p<0.01) affected by changes in habitat complexity and vegetation vertical structure in core or edge point counts. Therefore, we believe that changes in vegetation structure at the edge of littoral forest remnants may be a key indicator of mechanisms involved in edge sensitivity of forest dependent species in these forests. Our findings indicate that habitat fragmentation and degradation affect Madagascan bird communities and that these processes threaten many species. With continued deforestation and habitat degradation in Madagascar, we predict the further decline of many bird species.     

Mixed-species bird flocks from Brazilian Atlantic forest: the effects of forest fragmentation and seasonality on their size, richness and stability

The effects of forest fragmentation on species richness, size and stability of bird mixed-species flocks were studied during rainy and dry seasons at nine Atlantic Forest fragments in Minas Gerais state, southeastern Brazil. Two distinct kinds of mixed-species flocks were identified at the study area: heterogeneous flock and understory flocks. The first type of association was observed in all forest fragments and was composed mainly of canopy and midstory species, but may include a few understory species. Understory flocks were composed mainly of understory species and were detected in only three forest fragments. Forest fragment area and season significantly affected species richness, size and stability of heterogeneous flocks. Moreover, species richness in heterogeneous flocks was correlated with total species pool in forest fragments. Species richness and size of understory flocks were different among the three forest fragments, but season influenced only flock size. Understory flock stability differed neither among forest fragments nor between seasons. The red-crowned ant-tanager Habia rubica was the nuclear and leader species of understory flocks, and determined the occurrence of this association in forest fragments. Our results show that forest fragmentation can affect species’ interactions in foraging associations, and that conservation plans for the region should preserve nuclear species of mixed-species flocks and their preferred habitats.     

Effect of forest fragmentation on fruit and seed predation of the tropical dry forest tree Ceiba aesculifolia

Forests fragmentation reduces the density of natural plant populations forming patches of the remaining individuals. One of the biotic interactions that can be affected by forest fragmentation and is poorly studied is seed predation. We determined the effects of forest fragmentation on seed and fruit predation in Ceiba aesculifolia by comparing trees in continuous forest with trees in fragmented forest. We compared the following variables: (a) frequency of fruit predation by Collie’s squirrel (Sciurus colliaei) in each habitat; (b) frequency of the cotton-staining bug seed predator (Dysdercus, Orden Hemiptera) in each habitat; (c) the effect of seed predation on germination frequency and time; and (d) the effect of different life stages of Dysdercus on seed viability. In continuous habitat, 100% of the trees presented fruits with squirrel predation while only 34% of trees in fragmented habitats presented fruit predation. In continuous forest 27% of the trees contained fruits with the seed predator Dysdercus, while only 2% of the trees in fragmented forest presented Dysdercus. The initial weight of damaged seeds was greater than seeds that were not damaged indicating that seed predators select heavier seeds to feed upon. Frequency of seed germination was affected by different life stages; pre-adults decreased germination significantly more than nymphs and adults. Seed predation significantly increased the time it took for germination to occur. Our study shows that forest fragmentation significantly affects predation patterns of squirrels and cotton-staining bugs. Reduction of natural seed predators in forest fragments may have long-term consequences on forest structure and diversity.     

Does the matrix matter? A forest primate in a complex agricultural landscape

Many threatened primates now exist in fragmented forest habitats. The survival of these populations may depend on their ability to utilise agricultural or other matrix habitats between forest fragments, but this is poorly known. Here, we systematically investigate an arboreal primate’s use of a heterogeneous matrix in a fragmented forest landscape: the Angola black-and-white colobus (Colobus angolensis palliatus) in southern Kenya. We used a novel technique, based on semi-structured interviews with local informants, to address the difficulty of sampling relatively rare but important events, such as dispersal between fragments. We found that colobus frequently travelled and foraged in indigenous matrix vegetation (such as mangrove, wooded shrubland and shrubland) up to 4 km from the nearest forest fragments. Agricultural habitats, such as perennial plantation (coconut, mango and cashew nut) were also used by colobus for travelling and foraging (in remnant indigenous trees). The probability of sighting colobus in the matrix increased with the proportion of both tall (>6 m) vegetation cover and food tree cover, but declined with distance from forest habitat. Our findings suggest that certain matrix habitats are important for C. a. palliatus, and that future primate conservation initiatives might benefit from adopting a ‘landscape-level’ approach to habitat management, particularly in fragmented forest systems.     

Drastic erosion in functional attributes of tree assemblages in Atlantic forest fragments of northeastern Brazil

The long-term dynamics of plant communities remain poorly understood in isolated tropical forest fragments. Here we test the hypothesis that tropical tree assemblages in both small forest fragments and along forest edges of very large fragments are functionally much more similar to stands of secondary growth (5-65-yr old) than to core primary forest patches. The study was carried out in a severely fragmented landscape of the Brazilian Atlantic forest. Nine functional attributes of tree assemblages were quantified by sampling all trees (DBH ? 10 cm) within 75 plots of 0.1 ha distributed in four forest habitats: small forest fragments (3.4-79.6 ha), forest edges, second-growth patches, and primary forest interior areas within a large forest fragment (3500 ha). These habitats were markedly different in terms of tree species richness, and in the proportion of pioneer, large-seeded, and emergent species. Age of second-growth stands explained between 31.4% and 88.2% of the variation in the functional attributes of tree assemblages in this habitat. As expected, most traits associated with forest edges and small forest fragments fell within the range shown by early (<25-yr old) and intermediate-aged secondary forest stands (25-45-yr old). In contrast to habitat type, tree assemblage attributes were not affected by vegetation type, soil type and the spatial location of plots. An ordination analysis documented a striking floristic drift in edge-affected habitats. Our results suggest that conservation policy guidelines will fail to protect aging, hyper-fragmented landscapes from drastic impoverishment if the remaining forest patches are heavily dominated by edge habitat.