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.     

Influence of temperature in roost selection by Pipistrellus pygmaeus (Chiroptera): relevance for the design of bat boxes

The soprano pipistrelle (Pipistrellus pygmaeus) is a house-dwelling bat, whose maternity colonies are often affected by exclusion from buildings in southern Europe. Bat boxes with a suitable thermal behaviour can provide alternative roosts. However, this is particularly difficult to achieve in hot climates, where boxes may overheat. We studied roost temperatures and roost thermal preferences of this bat in attics and roofs in a Mediterranean region. Roosts reached high temperatures, experienced great thermal fluctuations and had wide spatial thermal ranges. Bats preferred the warmest locations within roosts, indicating that they are a heat tolerant species. However, they avoided temperatures of 40 °C and above, suggesting that this is the upper limit of the species thermally neutral zone. The study of experimental sets of white, grey and black boxes demonstrated that the latter were the ones that most often reached the temperatures observed in roosts in buildings. They were also the ones occupied by larger numbers of bats. These results suggest that, even in hot Mediterranean climates, black boxes are the best option as alternative roosts for evicted colonies. However, on particularly hot days, bats left them, suggesting that black boxes could be improved by creating a wider thermal range.     

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.     

Sex differences in population genetics, home range size and habitat use of the parti-colored bat (Vespertilio murinus, Linnaeus 1758) in Switzerland and their consequences for conservation

When habitats are declining, niche segregation by demographic groups, such as the two sexes, can have a profound impact on the extinction risk of a species as a whole. Thus, differences in the requirements of demographic groups are of importance in conservation. We combined behavioural and genetic data to investigate whether the sexually segregated parti-colored bat (Vespertilio murinus) exhibits sex-specific niche partitioning. We use our data to evaluate implications for conservation of this potentially vulnerable species in Switzerland, the western boundary of its range. Using radio-telemetry, we found sex-specific differences in habitat use. Foraging females strongly relied on lakes while foraging males displayed more flexibility in their habitat use. Moreover, males covered significantly larger foraging areas than females. Sequencing 341 base pairs of the mitochondrial D-loop of 247 individuals revealed sex-specific differences in the genetic structure of colonies, but no such difference was observed for three nuclear micro-satellite markers. We found high mtDNA diversity in two Swiss male colonies and one German female colony, but low mtDNA diversity in two Swiss female colonies. Our genetic data suggest that considerable gene flow occurs via male dispersal and mating. At the same time immigration of females into the existing female colonies in Switzerland is rare compared to the immigration of new males into male colonies. Since we found the sexes in Vespertilio murinus to differ markedly in their ecology, population genetics, and behaviour, we conclude that sex-specific conservation plans are required to protect this species efficiently.     

Roost selection by barbastelle bats (Barbastella barbastellus, Chiroptera: Vespertilionidae) in beech woodlands of central Italy: consequences for conservation

We investigated roost selection by Barbastella barbastellus in a mountainous area of central Italy. Twenty-five bats, mostly lactating females, were radio-tracked to 33 roost trees. Trees in unmanaged woodland were favoured as roost trees; woodland subject to limited logging was used in proportion to availability, and areas where open woodland and pasture occurred were avoided. Selection depended on tree condition (dead beech trees were preferred) and height (roost trees were taller than random ones). Cavity selection was based on cavity type, height and entrance direction: roost cavities were mainly beneath loose bark, at a greater height above ground and facing south more frequently than random cavities. Untouched areas of mature woodland should be preserved to provide roosting conditions for B. barbastellus. In logged areas, harvesting protocols should save dead and mature trees; frequent roost switching and small colony size imply that large numbers of roost trees are needed.     

Swarming of bats at underground sites in Britain—implications for conservation

We investigated the extent of visitation of underground sites during late summer and autumn (‘swarming’) by bats. Bats were captured at eight cave, mine and tunnel sites in southern England over 6 years (1995-2000). A total of 3077 bats representing 11 of Britain's 16 species was caught. Myotis bats predominated in autumn. At some sites these are rarely seen during winter hibernation counts. Myotis nattereri and M. daubentonii were most common. However, species composition changed seasonally. At one site, M. brandtii dominated early in the season and at all sites peak activity of M. daubentonii was earlier than in M. nattereri. Relatively high numbers of internationally vulnerable species such as M. bechsteinii and Barbastella barbastellus were recorded. There was a strong male bias in captures of swarming species. Swarming probably has an important reproductive function, most males are reproductively active at this time and such sites may be important for outbreeding. Due to this and the large number of individuals and species concerned, the conservation of ‘swarming sites’ is vitally important. Clearly swarming sites merit special consideration in conservation management strategies for cave-dwelling bats.     

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.     

Divergence in the soil and rhizosphere microbial communities of monoculture and silvopastoral traditional C. dodecandra agroforestry systems in Yucatan,Mexico

Numerous works have reported the impact on soil properties and microbial communities of intensive soil management, but very little is known about the impact caused by traditional agroforestry systems carried out by smallholders. To investigate whether the different smallholder's management between a monoculture plantation of Cordia dodecandra trees and a Silvopastoral system has led to a divergence in these ecosystems, soil properties, as well as soil and Cordia dodecandra rhizosphere microbial communities were analysed by MiSeq amplicon sequencing. The main findings were (i) Large variation in the soil properties of the Silvopastoral system suggests that it has a greater heterogeneity; (ii) Organic carbon, organic matter, carbonates, nitrogen, inorganic phosphorous and calcium, was significantly higher in the soil of the Silvopastoral system. (iii) The relative abundance of the major prokaryotic orders in soil and rhizospheres displayed small differences between the two agroforestry systems, and diversity indexes were slightly higher in the Silvopastoral system. (iv) The fungal orders Hypocreales and Pleosporales were more abundant in the Silvopastoral system than in the monoculture. Other evaluated parameters showed only minor or no difference. Thus, the main conclusion is that these soils have diverged in some properties and fungal orders, but not in their prokaryotic communities. To our knowledge, this is the first report on the divergence in soil properties and microbiota of these two extended smallholder agroforestry systems and therefore can serve as reference for future works.     

Predictive modelling and ground validation of the spatial distribution of the New Zealand long-tailed bat (Chalinolobus tuberculatus)

The use of predictive models is continually increasing, but few models are subsequently field-checked and evaluated. This study evaluates the statistical strength and usefulness for conservation purposes of a predictive habitat use model developed for Chalinolobus tuberculatus, a threatened microchiropteran bat species found in the temperate rainforests of New Zealand. The relationship between various environmental variables and the presence/absence of the species was investigated using generalised linear modelling. The model developed was coupled with GIS data to develop maps of predicted occurrence within the West Coast region of New Zealand’s South Island. It was found that distance to forest boundary, slope, presence of Nothofagus, general land cover, variability in mean annual solar radiation, and mean ambient winter minimum temperature were significantly associated with the occurrence of the species. Evaluation of the statistical strength of the distribution model with independent data of species’ occurrence collected at 152 sites found that the C. tuberculatus model showed a moderate ability to predict both species presence and absence (τ(b) coefficient = 0.37). The field detection rate (0.45) using this model was significantly higher than that of historical surveys (0.12). The value of the species habitat model and the need to evaluate its utility in the development of conservation strategies is discussed.     

Undesignated sites in Mexico qualifying as wetlands of international importance

Those sites designated under the Ramsar Convention on Wetlands of International Importance especially as Waterfowl Habitat constitute the most important wetland conservation network at a global level, with more than 1000 sites of international importance. With only seven designated sites, waterfowl and wetlands in Mexico are evidently under-represented in the list. We identify 34 currently undesignated sites in Mexico that qualify as wetlands of international importance, based on waterfowl count data from 1991-1997, using the Ramsar Convention criteria based on waterfowl. Using a complementarity approach implemented by linear integer programming, the sites were prioritised into two categories on the basis of their importance for designation. Twelve sites were categorised as Priority 1 (higher), and 22 sites as Priority 2 (lower). The Priority 1 set has held a waterfowl count average of 1.2 million individuals, and between 1% (ruddy duck, Oxyura j. jamaicensis) and 51% (black brant, Branta bernicla nigricans) of the populations of 10 species of waterfowl, and includes sites from eight biogeographic regions.     

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.     

Reduced-impact logging and temporal activity of understorey bats in lowland Amazonia

Because global timber demands continue to threaten tropical rain forests, identification of sustainable-use forest management protocols that meet human needs while preserving biodiversity is critical. Reduced-impact logging (RIL) protocols are increasingly common in the tropics and may be a viable option for sustainable forest use; however, few studies have documented faunal responses to RIL. Moreover, evaluations of abundance or diversity may not be sufficient for a comprehensive understanding of faunal responses to human activities, especially in the short-term. We assessed the effects of RIL on the temporal activity patterns of abundant bats in lowland Amazonian rain forest. More specifically, we characterized temporal patterns of activity, overlap of temporal activity, and dispersion of activity modes for seven abundant bat species and for four common bat guilds in RIL forest and in undisturbed forest in Tapajós National Forest, Pará, Brazil. Temporal activity of aerial insectivores, nectarivores, and gleaning animalivores did not change in response to RIL. In contrast, three of five species of frugivores and frugivores as a group changed their patterns of temporal activity in response to RIL. RIL had a greater effect on temporal activity of frugivores that foraged in the understorey than on frugivores that foraged in multiple forest strata. Overlap of temporal activity of frugivores was greater than expected by chance in undisturbed forest, but was random in RIL forest. Changes in activity may be a response to a combination of increased predation risk and reduced distances of early evening flights between day roosts and feeding areas in RIL forest compared to undisturbed forest.     

Landscape-moderated importance of hedges in conserving farmland bird diversity of organic vs. conventional croplands and grasslands

Hedges provide important nesting, feeding and sheltering sites for birds in agricultural areas, while organic farming also enhances farmland birds. However, it is little known how the interaction of these local variables (amount of hedges and organic management) with landscape scale variables affects birds. We selected paired conventional and organic winter wheat fields and meadows in each of 10 landscapes in Germany. Birds were surveyed in the fields and in the adjoining hedges. More bird species occurred in organic than in conventional fields regardless of land-use type (wheat fields and meadows). However, hedge length had a much stronger effect on bird richness than organic farmland management. We found an interaction of landscape complexity and hedge length in that hedge length enhanced bird richness only in case of simple landscapes (<17% semi-natural areas within a 500 m radius around the centre of bird survey plots). In more complex landscapes the local effect of hedge length levelled off because bird richness was high even without local hedges. Therefore, adding hedges or introducing organic farming practices should be primarily promoted in simple landscapes, where it really makes a difference for biodiversity.     

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.     

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.     

The distribution of cave-dwelling bats and conservation status of underground habitats in Northwestern Turkey

The distribution and abundance of cave-dwelling bats were investigated in the Thrace region of northwest Turkey. Data were collected in two periods, January-March and April-May 2001, from 32 underground sites, 25 of which had not been surveyed previously. Approximately 76,000 bats representing 13 species were recorded. The most abundant species were Miniopterus schreibersii, Rhinolophus euryale, Myotis myotis/blythii, Myotis capaccinii, and Rhinolophus ferrumequinum. The roosts were evaluated for their conservation importance. The most important sites in Turkish Thrace are Dupnisa and Koyunbaba. The Dupnisa Cave serves as a hibernaculum to approximately 28,000 bats representing five species. The Koyunbaba Cave is a nursery roost to approximately 23,000 bats of six species. Presently, none of the caves in the region has adequate protection and some bat populations are under serious threat.     

Uncovering the fruit bat bushmeat commodity chain and the true extent of fruit bat hunting in Ghana, West Africa

Harvesting, consumption and trade of bushmeat are important causes of both biodiversity loss and potential zoonotic disease emergence. In order to identify possible ways to mitigate these threats, it is essential to improve our understanding of the mechanisms by which bushmeat gets from the site of capture to the consumer’s table. In this paper we highlight the previously unrecognized scale of hunting of the African straw-colored fruit bat, Eidolon helvum, a species which is important in both ecological and public health contexts, and describe the commodity chain in southern Ghana for its trade. Based on interviews with 551 Ghanaians, including bat hunters, vendors and consumers, we estimate that a minimum of 128,000 E. helvum bats are sold each year through a commodity chain stretching up to 400 km and involving multiple vendors. Unlike the general bushmeat trade in Ghana, where animals are sold in both specialized bushmeat markets and in restaurants, E. helvum is sold primarily in marketplaces; many bats are also kept by hunters for personal consumption. The offtake estimated in this paper raises serious conservation concerns, while the commodity chain identified in this study may offer possible points for management intervention. The separation of the E. helvum commodity chain from that of other bushmeat highlights the need for species-specific research in this area, particularly for bats, whose status as bushmeat is largely unknown.     

Relative efficiency of extensive grazing vs. wild ungulates management for dung beetle conservation in a heterogeneous landscape from Southern Europe (Scarabaeinae, Aphodiinae, Geotrupinae)

The goal of this paper is to evaluate the relative efficiency of two strategies that favour dung beetles conservation (Scarabaeinae, Aphodiinae, Geotrupinae) in Southern Europe: extensive grazing and wild ungulate management. We conducted a study in the French Cévennes national park/UNESCO World Biosphere Reserve where dung beetles are distributed in a dichotomous sheep (grasslands, shrublands) and deer (clearings, shrublands, forests) droppings dominating abroad landscape. Natural sheep droppings and deer lumps dung beetle assemblages have been sampled two consecutive years during spring, summer and autumn in five representative habitats. This sampling of natural assemblages allowed for the estimation of (i) the density of trophic resource in the habitats, (ii) the regional dung beetle fauna and the distribution of species among the habitats and (iii) the density of beetles in natural droppings and the evenness of assemblages. High diversity, high species density in droppings and high evenness were observed in grazed shrubland, whereas fewer species were observed in deer lumps. Our results clearly showed that, while wild ungulates manure is not enough to ensure the conservation of the regional dung beetle species pool, the spatial habitat heterogeneity of grazed shrubland allows the local coexistence of numerous species. Consequently, one may expect that the conservation of European dung beetle fauna, especially Scarabaeinae and several long lifetime species, could be enhanced by extensive grazing.     

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.     

Herpetofauna diversity and microenvironment correlates across a pasture-edge-interior ecotone in tropical rainforest fragments in the Los Tuxtlas Biosphere Reserve of Veracruz, Mexico

We evaluated the relationship between amphibian and reptile diversity and microhabitat dynamics along pasture-edge-interior ecotones in a tropical rainforest in Veracruz, Mexico. To evaluate the main correlation patterns among microhabitat variables and species composition and richness, 14 ecotones were each divided into three habitats (pasture, forest edge and forest interior) with three transects per habitat, and sampled four times between June 2003 and May 2004 using equal day and night efforts. We measured 12 environmental variables describing the microclimate, vegetation structure, topography and distance to forest edge and streams.After sampling 126 transects (672 man-hours effort) we recorded 1256 amphibians belonging to 21 species (pasture: 12, edge: 14, and interior: 13 species), and 623 reptiles belonging to 33 species (pasture: 11, edge: 25, and interior: 22 species). There was a difference in species composition between pasture and both forest edge and interior habitats. A high correlation between distance to forest edge and temperature, understorey density, canopy cover, leaf litter cover, and leaf litter depth was found. There was also a strong relationship between the composition of amphibian and reptile ensembles and the measured environmental variables. The most important variables related to amphibian and reptile ensembles were canopy cover, understorey density, leaf litter cover and temperature.Based on amphibian and reptile affinity for the habitats along the ecotone, species were classified into five ensembles (generalist, pasture, forest, forest edge and forest interior species). We detected six species that could indicate good habitat quality of forest interior and their disappearance may be an indication of habitat degradation within a fragment, or that a fragment is not large enough to exclude edge effects. Different responses to spatial and environmental gradients and different degrees of tolerance to microclimatic changes indicated that each ensemble requires a different conservation strategy. We propose to maintain in the Los Tuxtlas Biosphere Reserve the forest remnants in the lowlands that have gentler slopes and a deep cover of leaf litter, a dense understorey, and high relative humidity and low temperature, to buffer the effects of edge related environmental changes and the invasion of species from the matrix.