The effects of extreme forest fragmentation on the bird community of Singapore Island

Singapore Island suffered one of the highest known deforestation rates in the tropics from the mid-to-late 19th century when over 95% of its native lowland forest was cleared. We compared the current bird community structure and composition among three habitat types, i.e., old (>50 years, 7-935 ha) and young (?50 years, 29-49 ha) naturally regenerating secondary forests and abandoned wooded plantations (27-102 ha) dominated by exotic species. Forest patch area had the strongest influence on the current species richness. The overall bird richness was not greater in most mature forest patches, but 20 species were only found in the old secondary forests and five of these were found in <50% of these patches. The rapid decrease in the number of forest species in plantations was offset by an increase in the number of open habitat species. Comparisons with current bird communities in nearby mainland forest sites (Peninsular Malaysia) suggest that the forest avifauna of Singapore is depauperate. The preservation of larger mature and maturing forests is therefore required for conserving the extant forest avifauna in Singapore. Connecting isolated patches can also be envisioned to facilitate movements of forest birds that have low densities and restricted distribution.     

Birds as agents of seed dispersal in a human-dominated landscape in southern Costa Rica

Birds play vital roles as seed dispersers helping to maintain and restore plant communities. With restoration increasingly key to global conservation, it is important to understand the landscape attributes and bird community characteristics that most influence avian seed dispersal in human-altered landscapes. We examined bird community structure and seed-dispersal patterns in agricultural countryside in Costa Rica that is typical of much of the Neotropics. Contrary to expectations, bird abundance, not richness, best predicted the richness of bird-dispersed seeds. Neither forest patch size or proximity, nor total tree cover, influenced seed dispersal. The richness and abundance of dispersed seeds, however, was strongly correlated with “wetness,” a remotely-sensed metric of vegetation, at several scales. These results suggest that in this human-dominated tropical region: (1) bird abundance, not species richness or size, may drive seed dispersal, and (2) remote-sensing combined with field verification can detect landscape elements that are helpful for maintaining the option of bird-mediated reforestation.     

The responses of understorey birds to forest fragmentation, logging and wildfires: An Amazonian synthesis

We combine mist-net data from 24 disturbance treatments taken from seven studies on the responses of understorey Amazonian birds to selective logging, single and recurrent wildfires, and habitat fragmentation. The different disturbance treatments had distinct effects on avian guild structure, and fire disturbance and the isolation of forest patches resulted in bird communities that were most divergent from those in continuous, undisturbed forest in terms of their species composition. Although low-intensity logging treatments had the least noticeable effects, the composition of understorey birds was still markedly different from the composition in undisturbed forest. This analysis demonstrates the importance of preventing habitat fragmentation and the spread of fires in humid tropical forests, and highlights the need for more research to determine the long-term suitability of large areas of degraded forest for forest birds.     

Long-term effects of fragmentation and fragment properties on bird species richness in Hawaiian forests

Forest fragmentation is a common disturbance affecting biological diversity, yet the impacts of fragmentation on many forest processes remain poorly understood. Forest restoration is likely to be more successful when it proceeds with an understanding of how native and exotic vertebrates utilize forest patches of different size. We used a system of forest fragments isolated by volcanic activity 153 years ago in Hawaii to examine how long-term fragmentation, as well as fragment size and structural features affect the richness of native and exotic bird species. The total number of bird species increased rapidly with forest fragment size, with most of the native species pool found in patches <3 ha. Smaller fragments were dominated by native bird species with several exotic bird species found only in the largest fragments, suggesting that exotic bird species in this landscape show greater area-sensitivity than native species. We used airborne scanning light detection and ranging (LiDAR) to assess whether fragment area was correlated with estimates of fragment vegetation volume as well as measures of tree height. Fragment area was highly correlated with vegetation volume, maximum tree height, and canopy height heterogeneity, and these variables were strong predictors of bird richness, demonstrating that remote sensing can provide key insights into the relationship between fragment structural attributes and biodiversity indicators. Overall, this work demonstrates the value of conserving small remnant mid-elevation forest patches for native birds in Hawaii. This work also provides insight into how newly created forest patches might be used by native and exotic bird species in Hawaii.     

The effect of the surrounding landscape matrix on mangrove bird community assembly in north Australia

Mangroves are highly threatened ecosystems yet their community ecology is poorly understood. We examined the ecological determinants of bird community assemblage in floristically depauperate mangroves. Birds were surveyed using line transect methods. Large mangrove patches supported fewer species than smaller patches. Patches did not comprise nested species subsets and the bird species richness of several small patches combined was greater than a single large area. The number of mangrove dependent species in a patch was area-dependent suggesting these species may be resource limited, although there was no species density compensation. There was a clear effect of the surrounding habitat, with matrix species accounting for ∼45% of bird species in a patch. Patches surrounded by tropical savanna were relatively species-poor, while regardless of size, patches including monsoon rainforest were relatively species rich. Null model analysis of non-random assemblage structure (nestedness and species co-occurrence) revealed no deterministic structure to the overall mangrove species assemblage. These analyses described a random pattern of bird distribution and with no evidence of density compensation this suggests that competition is a weak structuring force of mangrove bird assemblages. The lack of nestedness and the random co-occurrence of species are consistent with the matrix-dependence of bird community composition. Conservation plans should treat mangrove patches as part of a habitat mosaic and incorporate many smaller mangrove patches rather than just big ones. Consideration of the nature, extent and diversity of the surrounding matrices is vital in managing and conserving mangrove bird communities.     

Dissecting habitat loss and fragmentation effects following logging in boreal forest: Conservation perspectives from landscape simulations

Habitat loss and fragmentation are recognized as major threats to biodiversity. Their respective effects, however, are sometimes not well distinguished, even though habitat loss is recognized as the most important source of variation affecting species abundance and richness at the landscape scale. As ‘habitat’ is a species-specific concept (based on species perception of its environment), habitat loss and fragmentation studies should be conducted on a species-specific basis. We here assessed the influence of habitat loss and fragmentation in the context of a boreal forest considering forest clearcutting as an anthropogenic disturbance inducing mature forest loss and fragmentation that has a potential impact on wildlife. Using 16 simulated patterns of mature forest loss and fragmentation and three natural landscapes as replicates, we assessed the respective influence of forest loss and fragmentation on the abundance of 10 bird species common in the boreal forest of eastern Canada. Species–habitat relationships were modeled through habitat use models that were utilized to predict abundance of the 10 species within each combination of loss and fragmentation patterns (3 landscapes × 16 patterns). We used three-way ANOVAs to assess the effects of mature forest loss, fragmentation and replicates (random effect) on species abundance. Our results indicated that: (1) variation in species abundance mostly depended on mature forest loss, followed by static landscape attributes other than cutovers (e.g. streams, lakes, roads) and finally by fragmentation and (2) responses to mature forest loss and fragmentation differed among species, not necessary in relation to the successional status but in relation to their perception of their environment. Decreasing detrimental effects of mature forest loss through conservation of large continuous patches of forest may be suitable to maintain abundances of mature forest bird species. Our results highlight that studies aiming to quantify effects of habitat loss and fragmentation on wildlife should be conducted on a species-specific basis and use several landscape replicates to avoid potentially biased results.     

Landscape and patch-level factors influence bird communities in an urbanized tropical island

As human population continues to increase and intensification of human land use escalates, it is important to address the role of urban forest patches in supporting bird communities. We related bird species richness and community assemblage to landscape- and patch- level factors in 40 forest patches in the densely populated metropolitan area of San Juan, Puerto Rico. In total, 54 bird species were observed including 26 resident, 10 endemic, 12 migratory, and 6 introduced species. Patch size, level of urbanization in the surrounding matrix, and vertical heterogeneity of forest patches were the most important variables in explaining species richness. Patch size had the highest predictive power in explaining species richness for all groups except migrants, which were best predicted by patch-level factors (vegetation heterogeneity). The degree of matrix urbanization was correlated negatively with endemic species richness, but positively with introduced species. Endemic species were particularly sensitive to landscape factors (patch size, matrix urbanization, and canopy texture). Introduced species richness was not dependent on any patch-level factors. Eight species were relatively unaffected by urbanization, whereas two species were only found in large forest patches, and two species were sensitive to urban development. Seven species demonstrated a preference for small patches. Recommendations for land managers and conservation agencies to maintain a high bird species richness and diversity include: (1) preserve both large and small forest patches, (2) limit urban development near forest patches, (3) manage forest patch structure to maintain vertical heterogeneity, and (4) maintain forest patches with different vegetation types.     

Anthropogenic determinants of primate and carnivore local extinctions in a fragmented forest landscape of southern Amazonia

Habitat fragmentation has been shown to influence the abundance, movements and persistence of many species. Here, we examine the effects of forest patch and landscape metrics, and levels of forest disturbance on the patterns of local extinction of five primate and 14 carnivore species within 129 forest patches in a highly fragmented forest landscape of southern Brazilian Amazonia. Classic habitat area effects were the strongest predictors of species persistence, explaining between 42% and 55% of the overall variation in primate and carnivore species richness. Logistic regression models showed that anthropogenic disturbance, including surface wildfires, timber extraction and hunting pressure, had detrimental effects on the persistence of some species over and above those of fragment size. Different species ranged in their responses from highly sensitive to highly tolerant to forest fragmentation. Patterns of local extinction documented here were by no means chance events, and the nestedness of the overall species-by-site matrix was highly nonrandom in terms of the sets of species extirpated from the most to the least species-rich forest patches.     

Tolerance of frugivorous birds to habitat disturbance in a tropical cloud forest

In view of the continued decline in tropical forest cover around the globe, forest restoration has become a key tool in tropical rainforest conservation. One of the main - and least expensive - restoration strategies is natural forest regeneration. By aiding forest seed influx both into disturbed and undisturbed habitats, frugivorous birds facilitate forest regeneration. This study focuses on the tolerance of a frugivorous bird community to anthropogenic habitat disturbance within the broader context of natural forest regeneration with conservation purposes. It was carried out in the tropical cloud forest of Costa Rica’s Talamanca Mountains. Bird community response and tolerance to habitat disturbance was assessed by comparing bird presence and densities along a disturbance gradient, ranging from open pastures to closed mature forests. Birds were censused along nine transects applying the variable width line transect procedure. Forty relevant frugivorous bird species were observed during 102 h of survey time. Densities were calculated for 33 species; nine species responded negatively to increasing level of disturbance and nine others positively. Results indicate that large frugivores are generally moderately tolerant to intermediate, but intolerant to severe habitat disturbance, and that tolerance is often higher for medium and small frugivores. It appears that moderately disturbed habitats in tropical cloud forests are highly suitable for restoration through natural regeneration aided by frugivorous birds. Due to a lack of large forest seed dispersers, severely disturbed habitats appear less suitable.     

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.     

Composition and abundance of birds in Andean alder (Alnus acuminata) patches with past and present harvest in Bolivia

We compared species richness and abundance of birds between five patches under selective Alnus exploitation and five patches that have not been harvested for at least 10 years prior to our study, during the early dry season (April-July 2001), in Cotapata National Park. Using “point counts” we recorded birds and their distribution in two (<1.5 m and >1.5 m) forest layers. Simultaneously we evaluated the floristic structure (size [dbh] distribution, basal area, tree density, tree height, and vegetation cover) and composition (diversity) on three transects placed within each Alnus patch. Both bird diversity and vegetation cover were significantly higher in not presently used patches but only for the higher layer of the forest, whereas plant diversity was higher in presently used patches. Lack of differences between the two types of Alnus patches in any of the vegetation parameters measured in the lower layer was coupled with an indistinct avifauna. Small changes in habitat characteristics following a perturbation like selective logging have the potential to affect richness and abundance of birds, at least within the habitats directly affected by the perturbation.     

Six years of habitat modification in a tropical rainforest margin of Indonesia do not affect bird diversity but endemic forest species

Studies on temporal changes of tropical bird communities in response to habitat modification are rare. We quantified changes in bird assemblages at the rainforest margin of Lore Lindu National Park, Central Sulawesi, over an interval of 6 years. Standardized bird counts were conducted in the years 2001/2002 and 2008 at 15 census points representing natural forest, secondary forest, agroforest and openland sites. Although overall species richness remained nearly identical, different species groups were affected unequally by habitat modification within the forest margin landscape. The mostly endemic forest species declined in abundance (72.0% of forest species) and were detected at fewer census points in 2008 (56.0%). In contrast, 81.8% of the solely widespread openland birds became more abundant and 63.6% of the species were recorded at a larger number of census points. Hence, recent human activities in the forest margin ecotone negatively affected species of high conservation value. Species richness turned out to be a poor indicator of habitat change, and our results underline the importance of considering species identities. Biotic homogenization as result of habitat conversion is a global phenomenon. In our study, the winners were widespread openland species, while the losers were endemic forest birds. In conclusion, our study shows that 6 years of land-use change had negative impacts on bird community structure and endangered species, but not on overall bird species richness.     

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.     

Comparison of breeding bird and vegetation communities in primary and secondary forests of Great Smoky Mountains National Park

We compared breeding bird communities and vegetation characteristics at paired point locations in primary (undisturbed) and mature secondary forest (70-100 years old) sites in Great Smoky Mountains National Park, USA to understand how sites logged prior to creation of the park compare to undisturbed sites following 70 years of protection from human disturbance. We found that bird and vegetation communities are currently similar, but retain some differences in species composition. Rank abundance curves for primary and secondary forest bird communities showed very similar patterns of species dominance. Species composition was also similar on the two sites which shared 24 of the 25 most frequently recorded species. Nonetheless, comparisons of density estimates derived from distance sampling showed three bird species were more abundant on primary forest sites and that one bird species was significantly more abundant on secondary forest sites. Notably, comparisons based on raw counts (unadjusted for potential differences in detectability) produced somewhat different results. Analyses of vegetation samples for the paired sites also showed relative similarity, but with some differences between primary and secondary forests. Primary forest sites had more large trees (trees greater than 50 cm diameter at breast height) and late successional species. Primary forest sites had a denser tall shrub layer while secondary forest sites had a denser canopy layer. Nonetheless, tree species richness, basal area of live trees and number of standing snags did not differ between primary and secondary forest sites. Results indicate that breeding bird communities on sites within the park that were logged commercially 70 years ago are currently quite similar to bird communities on sites with no history of human disturbance. Similarities between the bird communities on previously disturbed and undisturbed sites in Great Smoky Mountains National Park may exceed those on more fragmented landscapes because large patches of primary forest, adjacent to commercially logged sites, remained in the park when it was established in 1935. These patches of primary forest may have served as source areas for commercially logged sites.     

Simulating anthropogenic impacts to bird communities in tropical rain forests

We used an aggregated modelling approach to simulate the impacts of anthropogenic disturbances on the long-term dynamics of faunal diversity in tropical rain forests. We restricted our study to bird communities even though the approach is more general. We developed a model called BioDiv which simulated the establishment of hypothetical bird species in a forest. Our model was based on the results of a simple matrix model which calculated the spatio-temporal dynamics of a tropical rain forest in Malaysia. We analysed the establishment of bird species in a secondary forest succession and the impacts of 60 different logging scenarios on the diversity of the bird community. Of the three logging parameters (cycle length, method, intensity), logging intensity had the most servere impact on the bird community. In the worst case the number of bird species was reduced to 23% of the species richness found in a primary forest.     

Regional scale effects of human density and forest disturbance on large-bodied vertebrates throughout the Yucatán Peninsula, Mexico

The occupancy probability of 35 large-bodied bird and mammal species was examined in relation to patch- and landscape-scale habitat and disturbance variables in 147 forest patches distributed throughout the Mexican Yucatán Peninsula. Occupancy was assessed on the basis of interviews with local informants. The most important predictors of vertebrate species richness, composition, and patch occupancy were human population density and the extent and quality of forest cover. Most forest species responded positively to forest extent, while felids in particular were sensitive to human population density. However, the effects of human density on patch occupancy operated at extremely local scales. Effects were stronger at a smaller grain size, offering optimistic prospects for conservation strategies that incorporate human population effects. Three arboreal frugivores (Ateles geoffroyi, Alouatta pigra, and Ramphastos sulfuratus) were strongly associated with total basal area of trees bearing fleshy fruits. The degree of hunting pressure was not related to human population density, and affected the occupancy probability of three game species, two of which (Mazama spp., Crax rubra) are among the most preferred prey across the Yucatán Peninsula. Levels of patch occupancy across this region varied considerably among species, and were best explained by body size and degree of forest habitat specificity, large-bodied species and habitat specialists being the most vulnerable. This study provides a quantitative assessment of the conservation potential of large vertebrates in Mesoamerica and identifies disturbance-sensitive species. This can inform regional-scale conservation planning at a time when low deforestation in parts of the Yucatán Peninsula still provides a narrow window of conservation opportunity given the rapid human population growth.     

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.     

Matrix effects on bird assemblages in fragmented Afromontane forests in South Africa

We examine the effects of matrix type on forest bird species richness in historically fragmented forests of the KwaZulu-Natal midlands, South Africa. Bird species-area relationships for forest fragments within natural grassland were compared with those surrounded by plantation forestry (Pinus spp.). While fragments in grasslands displayed a species-area effect, no such effect (slope ∼0) was detected for fragments in plantations. The critical fragment size to avoid an island effect on species richness was 302 ha for fragments in grasslands, and contained 51 of the 61 forest bird species encountered. Small forest fragments (<50 ha) within grasslands were less species-rich than those surrounded by plantation, while the reverse was true for forests larger than 50 ha. Bird density was significantly lower in larger forest patches within plantation when compared to those in grassland. However, a slight decline in species density with increasing species richness in a patch (i.e. weak density compensation) characterised only those bird assemblages surrounded by a grassland matrix, suggesting these are approaching species saturation. Commercial plantations may increase the likelihood of colonisation of, and immigration from, small forest fragments by birds, reducing the incidence of area-dependent extinction in small fragments, but may also result in lower species richness in larger fragments. The latter is likely a consequence of the promotion by plantations of a wider distribution of generalist species in, and the loss of some specialised or rare species from, large patches. Management options are to avoid planting plantations near large forest patches and to increase the size of small patches where possible. Where afforestation is unavoidable, placing plantations in the vicinity of small forest patches rather than large forest patches may be preferable.     

Landscape-level effects on avifauna within tropical agriculture in the Western Ghats: Insights for management and conservation

A critical handicap to tropical biodiversity conservation efforts in agroecosystems is the unknowns regarding the influence of landscape-scale factors on the persistence of species. To address these uncertainties, we explored two essential landscape-scale questions, within India’s biologically-rich Western Ghats, examining two nearby human-dominated landscapes that dramatically differed in their pattern of land cover. First, how does the proximity of intact forest patches affect bird community composition within agricultural landscapes? Second, can simple remote sensing-derived measures (brightness, wetness, and NDVI) be used to estimate native bird species composition within those landscapes? In both landscapes, as distance to intact forest decreased, the similarity in bird community composition between agricultural areas and intact forest increased. This suggests that the retention of tropical forest bird communities within human-dominated landscapes critically depends on the maintenance of nearby intact forest. In an answer to the second question, the remote sensing measures correlated with forest-affiliated avian species richness in only one of the two landscapes, reflecting an ecological difference between the two in the response of forest bird species to local agricultural conditions. In the landscape where a correlation was found, there was high variation in vegetative structure, which strongly impacted both the remote sensing measures and forest bird species richness. In the other landscape, forest species richness strongly correlated with changes in tree species composition in the agriculture, a factor that could not be detected by the remote sensing metrics. In order to successfully conserve biodiversity in tropical agricultural landscapes, our findings show that it is essential to conserve intact forest within those landscapes and to understand the effect of local agricultural practices on species.     

Bird conservation in fragmented Mediterranean forests of Spain: effects of geographical location, habitat and landscape degradation

The effects of habitat fragmentation on forest bird assemblages were analysed in 214 holm oak (Quercus ilex) remnants spread across the northern and southern plateaux of central Spain. Bird richness was highly dependent on fragment area for all species regardless of isolation, and barely affected by habitat traits. Geographical location was associated with high differences in richness of bird assemblages, which included 17 species exclusive to northern remnants and one exclusive to southern remnants. This supports the hypothesis that habitat suitability deteriorates sharply from north to south for forest birds in Spain. The species-area relationships of bird assemblages sampled in fragmented forests along a broad continental gradient (from Norway to southern Spain) showed that true forest birds only nest in woodlands >100 ha in southern Spain, whereas the full complement of forest species occurs in much smaller fragments in central-western Europe. Loss of species that are particularly sensitive to habitat fragmentation accounts for these differences between dry Spanish and mesic European woodlands. These results are explained by the low habitat suitability of Spanish woodlands, associated with the restrictive conditions for plant regeneration in the Mediterranean climate and long-standing human usage. There is, therefore, a particular need to develop management strategies that conserve birds, and probably other forest organisms, in Mediterranean regions by preventing habitat deterioration and decreases in fragment size, and by conserving all woods >100 ha.