Effects of ground fires on understorey bird assemblages in Amazonian forests

Avifaunal responses to understorey fire disturbance and subsequent changes in habitat structure were examined within 20 burnt and unburnt forest plots of 0.25 ha (10×250 m), 10-15 months after an unprecedented understorey fire swept through the Tapajós-Arapiuns Extractive Reserve of central Brazilian Amazonia following the severe 1997-1998 El-Niño dry season. Although these surface fires in the previously undisturbed primary forest were relatively mild, they resulted in dramatic changes in forest structure consistent with those found elsewhere in Amazonia. Bird species negatively affected by these changes tended to be the least common, the most disturbance-sensitive, and habitat specialists. Considering different guilds, ant followers, dead-leaf gleaners, terrestrial gleaners, and arboreal sallying insectivores were the most negatively affected, whereas nectivores and arboreal granivores became more abundant in the burnt forest. The results highlight the severe consequences of even relatively mild surface fires in neotropical forests, and the importance of controlling haphazard frontier expansion for the conservation of susceptible species that are endemic to fire-prone regions.     

Effects of landscape change on clutches of Phyllomedusa tarsius, a neotropical treefrog

Phyllomedusa tarsius is an arboreal frog species that suspends its eggs on leaves above water in order to achieve the high relative humidity necessary for embryo development. I compared the hatching success of P. tarsius in ponds located in pasture, Vismia spp. regrowth, Cecropia spp. regrowth, forest fragments and continuous forest at a site in central Amazônia. Reproductive success of P. tarsius is measured as percentage of clutches producing at least one tadpole and number of tadpoles produced per clutch. Reproductive success was higher in Cecropia regrowth, forest fragments and continuous forest, compared to pasture and Vismia regrowth. Desiccation and flooding were the main causes of clutch loss in pasture and Vismia regrowth, while predation was more frequent in forest and Cecropia regrowth. The levels of successful of P. tarsius clutches observed across all habitat types suggests that a landscape mosaic including pasture, secondary growth and forest fragments may allow for recolonization and genetic exchange to occur in isolated forest patches.     

Do secondary forests act as refuges for old growth forest animals? Recovery of ant diversity in the Atlantic forest of Brazil

The extent to which secondary forests occupying degraded and abandoned lands provide suitable habitat for forest-adapted species is an important conservation issue in times of vanishing old growth forests. We used ants (Hymenoptera: Formicidae), a functionally important and diverse group of invertebrates, to investigate the recovery of soil taxa during secondary forest succession in the Atlantic Forest of Southern Brazil. We compared the resilience of epigeic vs. hypogeic ant assemblages. For this purpose we established 27 sites that encompassed a chronosequence from pastures to old growth forests on two contrasting soil types. Our results are based on a collection of 35 508 individuals in 40 genera.Richness and composition of ant assemblages in secondary forests have recovered slowly and have not approached conditions typical of old growth forests. The distribution of genera along the successional stages was arranged in a nested pattern where ant genera of younger successional stages were a subset of genera present in older stages. Edaphic conditions had no influence on the recovery process. Overall, richness of ants was lower at study sites with water-logged soils than at sites where soils did not exhibit hydromorphic properties. The hypogeic ant assemblage recovered more slowly than the epigeic assemblage.Our results show that secondary forests do not act as refuges for many forest-adapted animals which are currently restricted to discontinuous patches of old growth forest in the highly endangered Atlantic Forest of Brazil. Moreover, estimated recovery times of 50 to several hundred years suggest it would take much longer than previously presumed for complete recolonization.     

Nestedness of bird assemblages in fragmented Afromontane forest: the effect of plantation forestry in the matrix

Bird species composition (61 species) of Afromontane forest fragments (n = 19) embedded in a natural grassland matrix or a plantation forestry matrix were compared. Forest bird assemblages in a natural grassland matrix displayed a nested non-random species distribution. Species loss was significantly area-dependent and predictable. Assemblages in the plantation-dominated matrix were also nested but were neither area- nor isolation-dependent and these potential processes had equal influence over bird species composition. Forests patches <50 ha in plantation were more species rich than those in a grassland matrix and visa versa for patches >50 ha. The effect of the matrix is revealed here with arboreal plantation cover promoting dispersal across the landscape, favouring abundant generalist bird species in smaller patches than expected in the plantation matrix, and the loss of some specialists and species living at low densities from large (>50 ha) patches. Because of a significant species-area relationship and possible area-dependent extinction, the largest forest patch in the nested series in a grassland matrix protects most bird species (89%). However, in a plantation matrix the absence of a species-area relationship requires the combination of many larger forest patches to protect most bird species. Plantation forests in the matrix significantly alter forest bird assemblage structure and composition. To avoid the effects of insularisation and to maintain the viability of bird communities in Afromontane forests requires conservation of the larger forests in a natural matrix. However, for forests in a plantation matrix, the management of dispersal processes by manipulation of the matrix may be as important to conserving species and communities as is minimising extinctions. Avoiding placing plantations near large forests, increasing the size of small patches, and where afforestation is unavoidable, placing plantations in the vicinity of small rather than large forest patches, are preferable management practices.     

Lowland rainforest avifauna and human disturbance: persistence of primary forest birds in selectively logged forests and mixed-rural habitats of southern Peninsular Malaysia

We compared the composition and structure of primary forest avifauna among primary forests, selectively logged forests and mixed-rural areas (e.g. villages and agricultural areas) of Peninsular Malaysia. We found that forests that were selectively logged at least 30 years ago contained only 73-75% of the 159 species of extant primary forest birds, with an increased proportion of dominant species. We estimated that only 28-32% of the primary forest species utilized the mixed-rural habitat, and that the number of species that bred in the agricultural landscapes might be even lower. The microhabitat of different species most affected their vulnerability to disturbance. Most small, arboreal frugivores and omnivores, and insectivores that fed from tree trunks, showed greater persistence in the mixed-rural habitat than ground dwelling bird species, which were affected most by disturbance. Resource abundance and variables that were closely related to forest disturbance such as the density of large trees, density of dead trees, canopy cover density and shrub volume influenced the distribution of the primary forest birds. Large primary forest reserves and a revision of short-cycle logging regimes (ca. 30 years) are needed if we are to conserve the lowland rainforest avifauna of Peninsular Malaysia and other parts of Southeast Asia.     

The effects of including marine ecological values in terrestrial reserve planning for a forest-nesting seabird

Theoretical advances in systematic reserve design aim to promote the efficient use of limited conservation resources and to increase the likelihood that reserve networks enhance the persistence of valued species and ecosystems. However, these methods have rarely been applied to species that rely on spatially disjunct habitats. We used the marbled murrelet, a seabird that requires old-growth forest in which to nest and high quality marine habitats in which to forage, as a case study to explore methods of incorporating multiple ecological values into single species spatial reserve design. Specifically, we used the cost function in MARXAN to include the ecological value of marine habitats while identifying spatial solutions for terrestrial nesting habitat reserves. Including marine values influenced terrestrial reserve designs most when terrestrial habitat targets were low and little or none of the target was represented in pre-existing protected areas. Our results suggest that including marine values in the planning process will influence marbled murrelet terrestrial reserve designs most where substantial terrestrial nesting habitat still exists, where new reserves are relatively unconstrained by pre-existing reserves, or when conservation resources only allow the protection of a small fraction of available habitat. This paper presents a novel framework for incorporating multiple measures of ecological value in the spatial reserve design process and should be particularly useful for species that rely on multiple habitats during their life cycle.     

Can wood ants distinguish between good and bad food patches on the forest floor?

Wood ants (Formica rufa group) normally feed on secretions of aphids to obtain carbohydrates, and on free-living invertebrates to obtain proteins. The availability of protein resources is usually unpredictable, and the demand for proteins is high during the period when ant larvae are developing. Thus, ants should select high quality food patches, i.e. patches with plenty of prey, when these are available. Foraging on the forest floor is seldom observed, but should be an alternative behaviour during periods of scarce food supply in the trees. To study the hunting behaviour of wood ants on soil invertebrates, ants were offered fly larvae (maggots) in two different quantities (six or two per pah) at two distances from an ant trail. Maggots exposed on the forest floor were found by randomly patrolling ant scouts regardless of bait quality. However, scouts that found the baits with six larvae recruited workers faster and, on average, four times as many workers were recruited to the six-larvae bait than to the bait with only two larvae. This indicates that ants can distinguish between poor and rich patches and that they are able to use this information to recruit more workers. Also, more workers were recruited and more maggots were carried away from patches nearby trails than from those far away. The results indicate that, during the warm season, ants explore and exhibit adaptive ‘foraging behaviour’ on the forest floor in the whole territory, not only close to trails. Consequently, wood ant feeding on soil invertebrates may be a common way for obtaining a large amounts of protein.     

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

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

The importance of 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.     

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

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

Severe declines of ant-following birds in African rainforest fragments are facilitated by a subtle change in army ant communities

Tropical rainforests are disappearing at a rapid rate. Although several studies have revealed the detrimental effects of forest fragmentation on tropical birds, the ecological mechanisms facilitating the decline of populations have been poorly documented. In the tropical rainforests of Africa and America, ant-following birds track the massive swarm raids of army ants and prey on animals which are flushed by the ants. We analyzed the persistence of five species of ant-following birds along a habitat fragmentation gradient in western Kenya and tested if bird populations are limited by the abundance of army ant colonies in forests. Abundances of four of five ant-followers declined by 52–100% in forest fragments 113 ha. Multi-model Bayesian inference suggests that the decline of the three most specialized ant-followers is facilitated by a decrease in the abundance of the army ant Dorylus wilverthi in small forest fragments. Our data suggests that a second, fragmentation-tolerant army ant species, Dorylus molestus, does not functionally compensate for the decline of D. wilverthi because, first, of a higher affinity of birds to D. wilverthi raids (found for one species). Second, because the daytime activity of D. molestus is dependent on high humidity conditions, a pattern which was not found for D. wilverthi. Consequently, specialized ant-followers in small fragments, where D. wilverthi is missing, probably suffer from food scarcity due to a cease of army ant foraging in the dry season. Our results suggest that a subtle alteration of army ant communities caused by habitat fragmentation may have large ecological consequences.     

Conservation-oriented forestry and early successional saproxylic beetles: Responses of functional groups to manipulated dead wood substrates

New conservation-oriented forestry aims to maintain intact populations of forest organisms by improving the conservation value of managed forests and providing protected areas. We tested the conservation value of treatments of dead wood for assemblages of early successional saproxylic beetles. In nine areas in northern Sweden, we selected one clear-cut, one mature managed forest and one reserve. In 2001-2002, we placed three blocks of spruce logs, each containing control, burned and shaded logs and a high stump (“snag”) at each site. Saproxylic beetles emerging from the dead wood were collected using emergence traps and beetles flying close to it were collected using flight-intercept traps. After one year of exposure, assemblage composition was examined, with respect to nutritionally-defined functional groups, red-listed species and fire-favoured species. Experimental snags were most complementary to control logs, supporting different assemblages of cambium consumers and fungivores and supporting more red-listed individuals. Burned logs supported depauperate assemblages, particularly with respect to cambium consumers, while shading of logs affected assemblages of fungivores, but only on clear-cuts. Despite containing less dead wood, managed forests provided valuable habitat, supporting similar assemblages of saproxylic beetles to reserves. Most functional groups were less abundant on clear-cuts than in older forests, but fire-favoured species were more common on clear-cuts, suggesting that clear-cuts may support assemblages of species associated with natural disturbances, if suitable substrates are available. Utilization of logs by saproxylic beetles changes over time, so long-term monitoring of our experimental logs will determine their lifetime conservation value.     

Regrowth forests on abandoned agricultural land: A review of their habitat values for recovering forest fauna

Human land use has modified the structure and function of terrestrial landscapes throughout much of the world, with cropping and livestock grazing the major drivers of landscape change. In many tropical, sub-tropical, temperate and Mediterranean regions, regrowth forests regenerate naturally on abandoned agricultural land if human disturbance declines. With the exception of some tropical forest literature, the broader ecological and conservation literature has largely ignored the potential of regrowth forests to facilitate passive landscape restoration and the recovery of fauna communities in fragmented agricultural landscapes. This paper addresses this deficiency by reviewing the available global evidence of fauna recovery in regrowth forest from 68 papers, identifying the main gaps in current knowledge, and providing directions for further research. The majority of reviewed studies focus on regrowth in tropical regions, which often contain large areas of mature forest. Species’ utilisation of regrowth forest is highly variable and is particularly influenced by land-use history, an important determinant of the structural and compositional characteristics of regrowth forests. While site-scale (<1 ha) forest structure and floristic diversity were frequently studied, only 11 studies considered the spatial configuration and context of habitat patches and just two studies explicitly considered landscape structure. Based on this review, six key research questions are posed to direct future research on this important issue. We conclude that a broader perspective of the role of regrowth forest in the landscape is required if we are to realise the potential benefits of regrowth forest for passive landscape restoration and fauna conservation and recovery.     

The effects of forest patch size and matrix type on changes in carabid beetle assemblages in an urbanized landscape

In this study we compared ground beetles (Carabidae) from a range of different forest fragments along an urbanization gradient in Brussels, Belgium. We address the following questions: (i) How does the degree of urbanization in the surrounding habitat affect forest beetles, and does it interact with the effects of patch size and distance to forest edge? (ii) Do these factors have a different effect at the level of individual species, habitat affinity groups or total community? During 2002 we sampled 13 forest plots in 10 forest patches, ranging in size from 5.27 to 4383 ha. The beetles were captured using transects of pitfall traps from the edge to a distance of 100 m into each woodland and identified to species level. Effects of urbanization, forest size and forest edge were evaluated on total species number, abundance and habitat affinity groups and ten abundant, widespread model carabid species. Overall, the effects of urbanization, forest size and edge effects slightly influenced total species richness and abundance but appeared to have a major effect on ground beetle assemblages through species specific responses. More urbanized sites had significantly fewer forest specialists and more generalist species. Large forest fragments were favoured by forest specialist species while generalist species and species frequently associated with forest (forest generalists) dominated the smaller forests. Forest edges mainly harboured generalist species while forest specialist species were more frequent into the forests if the forest patches were large enough, otherwise they disappeared due to the destruction or impoverishment of their habitat. Our results show the importance of differentiating between habitat affinity, especially habitat generalists versus specialists, the latter having a higher value in nature conservation, and merely the quantity of species represented in human-dominated areas.     

Sampling for ants in different-aged spruce forests: A comparison of methods

It is necessary to use special sampling method for studying of each animal group. However each method has its specificity and describes the studied community a bit differently. Three common methods for sampling of ants were compared: soil excavation, pitfall traps and baits. Ants were sampled in 25 patches in a large spruce forest in the Czech Republic during 2005 and 2006. Each patch represented one of five age classes (0–2, 3–5, 8–12, 26–41 and 85–105 years old), and the patches were located in five sites such that the five patches in each site formed a chronosequence. The percentage of variability in ants as explained by other sampling techniques was as follows: 60.0% of the variability in bait data was explained by pitfall trap data, while only 19.7% of the variability in pitfall trap data was explained by excavation data. Species and numbers of ants on baits were strongly affected by bait type: Myrmica species preferred honey whereas Formicinae preferred tuna bait. Assembly composition in pitfall traps varied significantly between study sites. The position of sampling site in landscape (location in the forest regardless of the immediate habitat type where the trap was located) hence strongly affected ants. Proportions of ants in particular methods were equal in the majority of cases. For excavation samples, ant numbers and species were more influenced by the immediate habitat than the wider habitat, but the opposite was true for samples from pitfall traps.     

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.     

Acari (mite) assemblages under plantations of bluegum, Eucalyptus globulus, in southwestern Australia

Farm forestry, in particular the planting of exotic bluegums Eucalyptus globulus, is a strategy increasingly used in southwestern Australia to mitigate the damaging effects of land clearing and to improve agricultural productivity. At the same time, such agroforestry systems have the potential to at least partially impede the biodiversity decline associated with habitat destruction and agriculture. The soil/litter habitat, an important repository of biodiversity in terrestrial systems, is one of the habitats most affected by clearing and conventional agricultural practices. Within this habitat, free-living mites are one of the most diverse and abundant animal groups contributing significantly to key ecosystem functions. This study compares the soil and litter mite assemblages of 7-year-old E. globulus plantations with those of adjacent native forest and pasture sites. The assemblages associated with E. globulus were substantially more diverse than those of the pasture soils but well below that of the native forest. Particularly low densities of oribatid mites were observed in the plantation sites. We suggest that the young plantation age, the exotic nature of the plantation species, and the homogeneous, mono-specific litter all contributed to limit the potential for these plantations to enhance mite biodiversity.     

Canopy arthropod communities on Mexican oaks at sites with different disturbance regimes

We examined the effects of forest disturbance and fragmentation on diversity and composition of canopy arthropod fauna in six oak species in central Mexico. Canopy fogging of 54 trees during rainy and dry seasons was conducted on three temperate forests with different levels of disturbance and habitat fragmentation. In total, we found 831 morphospecies belonging to 20 arthropod orders. Diversity of arthropods was higher in the rainy season. Acari, Hymenoptera and Diptera orders dominated during the dry season, and Acari and Collembola during the rainy season. Oak species distributed in sites with lower disturbance had significantly higher arthropod diversity than those distributed in disturbed and fragmented sites. Mean density of arthropod fauna ranged from 47.0 individuals m⁻² in the dry season to 375.5 m⁻² in the rainy season. Forest disturbance and fragmentation reduced the number of arthropod morphospecies and modified the relative abundance of arthropods at order level.     

Gap-crossing by gliding marsupials: thresholds for use of isolated woodland patches in an agricultural landscape

Single trees and small patches of trees in farmland are conspicuous components of agricultural landscapes around the world, but their value for the conservation of biodiversity is not well known. In this study, arboreal mammals were censused by using hair-sampling tubes in small patches of woodland (single trees to patches <1.0 ha) in cleared farmland adjacent to a linear network of woodland known to support resident populations of arboreal mammals. Ninety-one small isolates were stratified by size (single trees or small patches) and distance from the linear network to test the capacity of animals to cross habitat ‘gaps’. The genus Petaurus (small gliding marsupials), the most commonly detected taxon, was recorded in 31% of hair-tubes (98 of 316). It occurred in 21% of sites in isolated trees and patches, and in all linear strips. A logistic regression model demonstrated that Petaurus sp. was most likely to occur in isolates in close proximity to linear strips and other patches of woodland. Ninety-five per cent of sites at which this taxon occurred were within 75 m of the linear network. This threshold corresponds with the maximum distance that animals can glide in a single movement between trees. The size of isolates did not influence utilisation rates. Such isolates are smaller than a single home range and were probably used to supplement home ranges centred on the linear network, by providing additional foraging habitat and den sites. Protection and restoration of isolated trees and small woodland clumps in cleared landscapes contributes to mammal conservation and this study provides quantitative data that can assist landscape design and habitat restoration in rural environments.     

Bird responses to broad-leaved forest patch area in a plantation landscape across seasons

Although plantation forests have widely replaced native forests worldwide, few studies have examined the effects of plantation forestry on organisms at the landscape level. In this study, we examined the effects of broad-leaved forest patch area (1.4–312 ha) on bird assemblages in a conifer plantation-dominated landscape during wintering and breeding seasons. We also surveyed birds in the plantation matrix and treated the patch area of the matrix as 0. We examined whether the detection rates of species changed suddenly across patch area (threshold) using segmented regression. We found that species richness increased with patch area in both seasons. Responses of detection rates to patch area varied and were categorized as follows: insensitive, linear increase and decrease, quadratic increase and decrease, concave and convex. Thresholds indicating that the detection rate suddenly increased above a certain patch area were found for two species only in the winter. Species responses varied more in the winter than in the breeding season. Certain ecological traits of species were consistently associated with the responses to patch area across seasons (migratory strategy, nesting substrate and foraging trait), while body weight was only associated in the winter. Because species richness and detection rates of many species were high in large patches, large broad-leaved forests (especially >40 ha) should be retained and restored with high priority. Additionally, bird responses to patch area in the winter should not be ignored because many species required large patches during the winter season.