Bats in continuous forest, forest fragments and in an agricultural mosaic habitat-island at Los Tuxtlas, Mexico

In order to document the responses of bats to destruction and fragmentation of their natural habitat and the value of different types of man-made vegetation for bat conservation in the Neotropics, bats were sampled with mist nets to compare species richness and species composition in a tract of continuous forest, forest fragments and a habitat-island consisting of a mosaic of forest and arboreal crops in Los Tuxtlas, southern Mexico. We captured 3835 bats representing 39 species: 76% were captured in continuous forest, 74% in the mosaic habitat and 87% in forest fragments. In the mosaic habitat we captured 43% of the total number of bats, 33% in the forest fragments and 24% in continuous forest. On average the habitats studied had 64% species in common. Evidence of continuous breeding activity was determined for a high number of species at the three habitats (> 70% lactating and 65% with embryos). A few bat species (Carollia brevicauda, Pteronotus parnelli, Sturnira lilium, Artibeus jamaicensis, Dermanura phaeotis, Vampyrodes caraccioli, Glossophaga soricina, Dermanuta toltecus, Cheoroniscus godmani, Platyrrhinus helleri) dominated the sample, but their relative dominance varied among habitats. Recapture of bats provided evidence for inter-habitat movement. The co-occurrence of the three habitats helps conserve a diverse assemblage of bat species in the local landscape.     

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.     

Effects of tropical forest fragmentation on aerial insectivorous bats in a land-bridge island system

Habitat fragmentation causes drastic changes in the biota and it is crucial to understand these modifications to mitigate its consequences. While studies on Neotropical bats have mainly targeted phyllostomid bats, impacts of fragmentation on the equally important aerial insectivores remain largely unexplored. We studied species richness, composition, count abundance and feeding activity of aerial insectivorous bats in a system of land-bridge islands in Panama with acoustic sampling. We predicted negative effects of fragmentation on forest species while bats foraging in open space should remain essentially unaffected. Rarefaction analyses indicated higher species richness for islands than mainland sites. For forest species, multivariate analyses suggested compositional differences between sites due to effects of isolation, area and vegetation structure. Contrary to our expectations, count abundance of forest species was similar across site categories. Feeding activity, however, was curtailed on far islands compared to near islands. As expected, bats hunting in open space did not reveal negative responses to fragmentation. Interestingly, they even displayed higher abundance counts on far and small islands. On the species level, two forest bats responded negatively to size reduction or site isolation, respectively, while a forest bat and a bat hunting in open space were more abundant on islands, irrespectively of island isolation or size. Our findings suggest that small forest remnants are of considerable conservation value as many aerial insectivores intensively use them. Hence high conservation priority should be given to retain or re-establish a high degree of forest integrity and low levels of isolation.     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

Landscape patterns influencing bird assemblages in a fragmented neotropical cloud forest

The cloud forest is one of the rarest and most threatened ecosystems in Mexico, although it contributes highly to the country's biological diversity and provides important ecological services. It is a naturally fragmented ecosystem, but anthropogenic deforestation and fragmentation has been severe. Consequently, it is essential to identify landscape patterns critical for the conservation of cloud forest. In order to understand how landscape patterns affect diversity in this ecosystem, this study explores the consequences of cloud forest fragmentation on bird diversity in eastern Mexico. I analysed the response of bird species richness and abundance as a function of forest fragment size, shape, topographical complexity, altitudinal range, connectivity, and proportion of landscape forested area in a system of 13 cloud forest fragments. Fragment shape was the main characteristic positively related to species richness in the bird community, but a differential response to landscape patterns was also detected. Fragment size was the main characteristic influencing the segment of the bird community depending mostly on forest, that is to say, forest interior and generalist species. In contrast, the extent of forest edge, expressed as fragment shape, produced a positive response of forest border species. Both, forest dependent and border dependent species positively responded to the extent of their suitable habitat. The immediate and most effective ecologically oriented conservation strategy for the region is the conservation of the largest cloud forest fragments.     

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.     

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.     

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.     

Patterns of roost use by bats in a neotropical savanna: implications for conservation

Neotropic bats comprise a highly species-rich group and are considered important regulators of complex ecological processes. Resource partitioning of roosts and food is considered a key element in shaping bat communities. In this paper we examine the roosting patterns of 10 bat species belonging to a neotropical bat community in a Bolivian savanna. Roosts were more common in open woodlands than in forest islands and human settlements. In open woodlands, Tabebuia heptaphylla comprised the largest proportion of roosting trees, whereas in forest islands, Gallesia integrifolia was the most important roosting tree for phyllostomid bats. By comparing active roosts with control holes in Tabebuia heptaphylla, we found that roost site selection by Noctilio albiventris was mainly based on tree diameter and inner width of the cavities. The earlier results are discussed in view of resource partitioning within savanna bat communities and their practical conservation.     

Conservation importance of limestone karst outcrops for Palaeotropical bats in a fragmented landscape

Limestone karst landscapes are important for biodiversity yet are increasingly threatened by development activities such as mining. Furthermore, karsts are often scattered and isolated by agriculture, and are rarely considered in landscape planning because of a paucity of biodiversity data. We determined the conservation significance of an isolated limestone karst outcrop for insectivorous bats by quantifying the influence of this roosting resource on local assemblage structure across a fragmented landscape in peninsular Malaysia. Using a combination of rank abundance, gradient and randomisation analyses, we demonstrate that bat assemblages at nine forest sites are structured following a spatial gradient of increasing distance from a karst roosting resource. The assemblage at our karst site was dominated by a superabundance of three cave-roosting species, two of which were also found to dominate assemblages up to 11 km away. Cave-roosting bats exhibited a significant decay in abundance related to the distance from karst, with sites closest to karst also characterised by a rarity of tree cavity/foliage-roosting species that were otherwise common. Gradient analysis revealed that differences in assemblage composition were largely associated with the distance from the karst and, to a lesser extent, forest isolation and area. Our findings suggest that isolated karst outcrops can serve as important population reservoirs for cave-roosting bats, which subsidise diversity levels in forest fragments that might otherwise be expected to decline over time. While conservation efforts need to focus on maintaining large areas of connected forest, landscape management needs to ensure protection of karsts as point resources for cave-roosting bats.     

The role of forest structure, fragment size and corridors in maintaining small mammal abundance and diversity in an Atlantic forest landscape

Using the abundance and distribution of small mammals at 26 sites in an Atlantic forest landscape, we investigated how species abundance and alpha and beta diversity are affected by fragment size and the presence of corridors. To account for the variability in forest structure among fragments, we described and minimized the influence of foliage density and stratification on small mammal data. Sites were distributed among three categories of fragment size and in continuous forest. For small and medium-sized categories, we considered isolated fragments and fragments connected by corridors to larger remnants. Small mammal abundance and alpha and beta diversity were regressed against site scores from the first axis of a Principal Component Analysis on forest structure variables. Residuals were used in analyses of variance to compare fragment size and connectivity categories. Forest structure influenced total abundance and abundance of some species individually, but not the diversity of small mammal communities. Total abundance and alpha diversity were lower in small and medium-sized fragments than in large fragments and continuous forest, and in isolated compared to connected fragments. Three species were less common, but none was more abundant in smaller fragments. At least one species was more abundant in connected compared to isolated fragments. Beta diversity showed an opposite relationship to fragment size and corridors, increasing in small and isolated fragments. Results highlight the importance of secondary forest for the conservation of tropical fauna, the hyper-dynamism of small isolated fragments and the potential of corridors to buffer habitat fragmentation effects in tropical landscapes.     

Influence of forest fragmentation on an endangered large-bodied lemur in northwestern Madagascar

Madagascar’s diverse and mostly endemic fauna and flora suffer from recent landscape changes that are primarily caused by high levels of human activities. The loss and fragmentation of forest habitats are well known consequences of human activities. In this study, we investigate the effects of forest fragmentation on presence, abundance and genetic diversity in a larger-bodied lemur species, Lepilemur edwardsi, in northwestern Madagascar. In addition, we characterized the genetic differentiation among populations and demographic changes. We found L. edwardsi at only 13 (76.5%) of 17 visited sites, 11 of which were situated in the Ankarafantsika National Park (ANP). We captured between two and 17 individuals per site. We sequenced the mtDNA d-loop of all samples and genotyped 14 microsatellite loci in two exemplary populations for demographic analyses. A negative influence on forest fragmentation could be detected, since the fragments had a lower genetic diversity than sites in the ANP. Genetic differentiation between populations ranged from low to high but was almost always significant. A typical pattern of isolation-by-distance could not be detected and the data could rather be interpreted as results of random genetic drift. The data furthermore revealed signals of a demographic collapse of about two orders of magnitude in the two exemplary sites. This decline probably started during the last few hundred years of intensified human disturbances and population growth. Given the results of this study, urgent conservation actions are needed and should concentrate on an effective protection of the few remaining populations in order to ensure the long-term survival of L. edwardsi.     

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.     

The importance of forest area and configuration relative to local habitat factors for conserving forest mammals: A case study of koalas in Queensland, Australia

The loss and fragmentation of forest habitats by human land use are recognised as important factors influencing the decline of forest-dependent fauna. Mammal species that are dependent upon forest habitats are particularly sensitive to habitat loss and fragmentation because they have highly specific habitat requirements, and in many cases have limited ability to move through and utilise the land use matrix. We addressed this problem using a case study of the koala (Phascolarctos cinereus) surveyed in a fragmented rural-urban landscape in southeast Queensland, Australia. We applied a logistic modelling and hierarchical partitioning analysis to determine the importance of forest area and its configuration relative to site (local) and patch-level habitat variables. After taking into account spatial autocorrelation and the year of survey, we found koala occurrence increased with the area of all forest habitats, habitat patch size and the proportion of primary Eucalyptus tree species; and decreased with mean nearest neighbour distance between forest patches, the density of forest patches, and the density of sealed roads. The difference between the effect of habitat area and configuration was not as strong as theory predicts, with the configuration of remnant forest becoming increasingly important as the area of forest habitat declines. We conclude that the area of forest, its configuration across the landscape, as well as the land use matrix, are important determinants of koala occurrence, and that habitat configuration should not be overlooked in the conservation of forest-dependent mammals, such as the koala. We highlight the implications of these findings for koala conservation.