Seedling assemblages and the alternative successional pathways experienced by Atlantic forest fragments

Background: Habitat loss and fragmentation have been argued to drastically alter the composition of tree assemblages inhabiting small forest fragments but the successional trajectory experienced by such edge-affected habitats remains controversial.

Aims: Here we examine whether small fragments (3.4–91.2 ha) support seedling assemblages more similar to those in 10–70-year-old secondary forests than to those in mature forests, in order to infer to what extent fragments move toward early successional systems.

Methods: Using 59 0.1-ha plots distributed in a fragmented landscape of Brazilian Atlantic forest, we evaluated species richness and functional and taxonomic composition of seedling assemblages in 20 small forest fragments, 19 stands of secondary forest and 20 stands of mature forests in the interior of an exceptionally large fragment (ca. 3500 ha).

Results: Small fragments presented the least species-rich seedling assemblages (17.2 ± 5.7 species), followed by secondary (22.5 ± 5.3), and mature forest (28.4 ± 5.3). Small fragments had seedling assemblages with functional and taxonomic composition more similar to those in secondary than in mature forest. Small fragments had a greater relative richness and abundance of pioneer trees (ca. 40% more), vertebrate-dispersed (6–25%), and those bearing medium-sized seeds (30–70%), while large-seeded species and individuals were reduced (>50% decrement) in comparison to seedling assemblages in mature forest.

Conclusions: By comparing seedlings across a wide range of successional habitats we offer evidence that small forest fragments are experiencing an alternative successional pathway towards an early-successional system with reduced plant diversity.     

Understanding and predicting frost‐induced tropical tree mortality patterns

Extreme climatic and weather events are increasing in frequency and intensity across the world causing episodes of widespread tree mortality in many forested ecosystems. However, we have a limited understanding about which local factors influence tree mortality patterns, restricting our ability to predict tree mortality, especially within topographically complex tropical landscapes with a matrix of mature and secondary forests. We investigated the effects of two major local factors, topography and forest successional type, on climate‐induced tropical tree mortality patterns using an observational and modeling approach. The northernmost Neotropical dry forest endured an unprecedented episode of frost‐induced tree mortality after the historic February 2011 cold wave hit northwestern Mexico. In a moderately hilly landscape covering mature and secondary tropical dry forests, we surveyed 454 sites for the presence or absence of frost‐induced tree mortality. In addition, across forty‐eight 1 ha plots equally split into the two forest types, we examined 6,981 woody plants to estimate a frost‐disturbance severity metric using the density of frost‐killed trees. Elevation is the main factor modulating frost effects regardless of forest type. Higher occurrence probabilities of frost‐induced tree mortality at lowland forests can be explained by the strong influence of elevation on temperature distribution since heavier cold air masses move downhill during advective frosts. Holding elevation constant, the probability of frost‐induced tree mortality in mature forests was twice that of secondary forests but severity showed the opposite pattern, suggesting a cautious use of occurrence probabilities of tree mortality to infer severity of climate‐driven disturbances. Extreme frost events, in addition to altering forest successional pathways and ecosystem services, likely maintain and could ultimately shift latitudinal and altitudinal range margins of Neotropical dry forests.     

Canopy bird assemblages are less influenced by habitat age and isolation than understory bird assemblages in Neotropical secondary forest

Secondary forest habitats are increasingly recognized for their potential to conserve biodiversity in the tropics. However, the development of faunal assemblages in secondary forest systems varies according to habitat quality and species‐specific traits. In this study, we predicted that the recovery of bird assemblages is dependent on secondary forest age and level of isolation, the forest stratum examined, and the species’ traits of feeding guild and body mass. This study was undertaken in secondary forests in central Panama; spanning a chronosequence of 60‐, 90‐, and 120‐year‐old forests, and in neighboring old‐growth forest. To give equal attention to all forest strata, we employed a novel method that paired simultaneous surveys in canopy and understory. This survey method provides a more nuanced picture than ground‐based studies, which are biased toward understory assemblages. Bird reassembly varied according to both habitat age and isolation, although it was challenging to separate these effects, as the older sites were also more isolated than the younger sites. In combination, habitat age and isolation impacted understory birds more than canopy‐dwelling birds. Proportions of dietary guilds did not vary with habitat age, but were significantly different between strata. Body mass distributions were similar across forest ages for small‐bodied birds, but older forest supported more large‐bodied birds, probably due to control of poaching at these sites. Canopy assemblages were characterized by higher species richness, and greater variation in both dietary breadth and body mass, relative to understory assemblages. The results highlight that secondary forests may offer critical refugia for many bird species, particularly specialist canopy‐dwellers. However, understory bird species may be less able to adapt to novel and isolated habitats and should be the focus of conservation efforts encouraging bird colonization of secondary forests.     

Birds,Cattle, and Bracken Ferns: Bird Community Responses to a Neotropical Landscape Shaped by Cattle Grazing Activities

Large areas of tropical moist forests have been converted to cattle pastures, generating complex landscapes where different habitats are represented by small patches with an uneven spatial distribution. Here, we describe how bird communities respond to the different elements present in a livestock landscape that was originally dominated by tropical moist forest. We surveyed six habitats: open pastures, pastures with shrubs, early‐ and middle‐secondary forests, mature forest, and pastures invaded by bracken ferns (Pteridium aquilinum). Bird diversity was high in secondary and mature forests, and low in fern‐invaded sites and open pastures. Fern‐dominated sites had the lowest bird species richness, and trophic guild diversity of all habitats. Habitat structure affected both bird species richness and densities in similar ways. Tree species richness was the habitat attribute that had a bigger positive effect on bird species richness. Bird community structure varied among sampled habitats, separating habitats in two major groups (forests and pastures). Our data indicate that bracken fern‐invaded pastures were the worst habitat condition for avian communities. To increase bird diversity, we recommend to eliminate or manage bracken fern and to increase shrub and tree cover in open pastures to provide food resources and shelter for birds. Finally, we encourage the maintenance of secondary and mature forest remnants as a strategy to conserve resident birds within a landscape dominated by livestock activities.     

Moderate Human Disturbance of Rain Forest Alters Composition of Fruiting Plant and Bird Communities

Forests worldwide are experiencing rapid environmental change due to human activity. We aimed to increase understanding of anthropogenic impacts on community composition and species interactions. In a natural experiment, we asked whether subsistence human land use has altered the community composition of a Neotropical rain forest on the island of Tobago, in the West Indies. We surveyed fruiting plants and birds in three adjacent habitat types that varied in level of disturbance, and used multivariate analyses to determine whether changes in the plant community were associated with differences in avifauna composition. The three forest habitats had similar plant and bird diversities, yet markedly different species compositions and abundances. Primary forest had the most diverse plant community, while disturbed habitats had a more homogeneous plant composition. Primary and disturbed forest had distinct community compositions, with canopy cover and the relative abundance of plant types explaining 83 percent of the variation in bird species assemblages. Seemingly moderate human disturbance has led to substantial changes in the plant and bird assemblages of Tobago's rain forest, outside of a protected reserve. Our study highlights the direct links between human disturbance and the structure of rain forests, underscoring the impact of even moderate activity on community composition.     

Does land-use change affect biodiversity dynamics at a macroecological scale? A case study of birds over the past 20 years in Japan

Because the effects of land-use change on biodiversity have primarily been examined at or below the regional scale, it remains unclear whether such effects scale up to the macroecological scale (i.e. nationwide or continental scale). In Japan, forests have become more mature since the cessation of most forestry efforts in the 1970s. At a nationwide scale, this forest maturation may lead to reductions in the abundance of species that depend on early successional forests (early successional species) and increases in the abundance of species that depend on mature forests (mature forest species). Japan has met its high demand for wood through imports from South-east Asia, resulting in deforestation there. Therefore, the abundance of mature forest species that migrate long distances to overwinter in South-east Asia may decrease. We examined changes in the range sizes of birds in Japan over the past 20 years using the living planet index (LPI). The LPI indicated that the range sizes of early successional species decreased. For mature forest species, the range sizes of long-distance migrants decreased, whereas those of short-distance migrants and residents increased. Our predictions were generally supported. Our results indicate that the effects of land-use change extend to the macroecological scale and that such changes in one country can affect the biodiversity dynamics in other countries. Forest maturation in Japan and concomitant deforestation in South-east Asia have been caused by internationally coupled socioeconomic processes. Therefore, biodiversity conservation at the macroecological scale must consider the role of land use, and such efforts will require both international and socioeconomic perspectives.     

长白山针阔混交林不同演替阶段的昆虫多样性

昆虫多样性变化对生态系统健康有重要的指示作用, 为研究昆虫群落变化与生境演替之间的关系, 本研究采用网捕、灯诱和诱捕法系统调查了长白山针阔混交林不同演替阶段（次生白桦林、次生针阔混交林、原始阔叶红松林）昆虫群落的组成和多样性, 分析了昆虫在森林演替过程中的规律及与植被群落之间的关系。系统调查共采集昆虫标本8 183头, 隶属于14个目699种, 其中鳞翅目和鞘翅目是主要优势类群。次生针阔混交林昆虫的个体数量最多, 原始阔叶红松林中物种数最多。不同演替阶段昆虫群落的物种数和个体数差异不显著, 但次生针阔混交林、原始阔叶红松林的Fisher’s α指数显著高于次生白桦林。目水平上的昆虫多样性未表现出显著性差异. 昆虫多样性在森林演替过程中和草本植物多样性的变化趋势相同;由于食性和生境选择的不同, 森林演替过程中鳞翅目昆虫多样性逐渐升高, 而鞘翅目多样性逐渐降低。     

Why are leaf‐cutting ants more common in early secondary forests than in old‐growth tropical forests? An evaluation of the palatable forage hypothesis

I evaluated the hypothesis that leaf‐cutting ants are more common in early successional forests than in old‐growth forests because pioneer species, which dominate in early successional habitats, appear more susceptible to leafcutters than shade‐tolerant species, which dominate primary forests (palatable forage hypothesis). The relative importance of pioneer and shade‐tolerant species as plant resources for leaf‐cutting ant colonies was evaluated (1) by literature review of leaf‐cutting ants’ diet, and (2) experimentally, using field assays to determine leafcutter's selectivity. Pioneer species were harvested three times more frequently than shade‐tolerant species and made up the largest component of the diet in all the studies reviewed. The amount harvested was not correlated with the plant species abundance. In addition, leaves from pioneer plants were selected eight times more than leaves from shade‐tolerant species in the field assays. These results support the palatable forage hypothesis. Leafcutters probably select pioneer leaves because of their low level of chemical defenses and high nutrient content. The high availability of pioneer species in early successional forest probably decreases the cost to locate palatable resources. Therefore, early successional habitats support more ant colonies than old‐growth forests. On the other hand, the effective defense mechanisms of mature plant species and the high dispersion of palatable plants could explain the low density of leaf‐cutting ant colonies in old‐growth forests. The palatable forage hypothesis is compared with other hypotheses that explain leaf‐cutting ant density. The preference of foundress queens for forest clearings, the dependence of small colonies on herbs, and the importance of pioneer plant species for mature colonies (palatable forage hypothesis) can be considered complementary, because they focus on different stages of the colony's life history. Consequently, the availability of pioneer plants appears to be one of the most influential factors determining mature leaf‐cutting ant nest densities in Neotropical forests.     

Environmental correlates of avian diversity in lowland Panama rain forests

Aim The composition of communities is known to be influenced by biogeographical history, but also by local environmental conditions. Yet few studies have evaluated the relative importance of the direct and indirect effects of multiple factors on species diversity in rich Neotropical forests. Our study aims to assess drivers of change in local bird species richness in lowland tropical rain forests. Location Thirty‐two physiographic subregions along the corridor of the Panama Canal, Panama. Methods We mapped the distributions of all forest‐dwelling bird species and quantified the environmental characteristics of all subregions, including mean annual rainfall, topographic complexity, elevational variability, forest age and forest area. Plant species richness, believed to be correlated with structural complexity, was estimated by interpolation through kriging for subregions where data were unavailable. Results The study region has a strong rainfall gradient across a short distance (65 km), which is also accompanied by steep gradients in plant and bird species diversity. Path analysis showed that precipitation strongly affected plant species diversity, which in turn affected avian diversity. Forest age and topography affected bird diversity independently of plant diversity. Forest area and its proportion occurring in the largest two fragments of each subregion (habitat configuration) were also positive correlates of bird species richness. Main conclusions Our results suggest that plant species richness, known to be influenced in part by biogeographical history and geology, also affects bird species assemblages locally. We provide support for the hypothesis that bird species richness increases with structural complexity of the habitat. Our analysis of the distributions of the region's most disturbance‐sensitive bird species showed that subregions with more rainfall, more complex topography and older forests harboured not only richer communities but also more sensitive species; while subregions with the opposite characteristics usually lacked large fractions of the regional forest bird community and hosted only common, widely distributed species. Results also emphasize the importance of preserving forest diversity from habitat loss and fragmentation, and confirm that larger, continuous forest tracts are necessary to maintain the rich avian diversity in the region.     

High Bird Species Diversity in Structurally Heterogeneous Farmland in Western Kenya

Tropical ecosystems are globally important for bird diversity. In many tropical regions, land‐use intensification has caused conversion of natural forests into human‐modified habitats, such as secondary forests and heterogeneous agricultural landscapes. Despite previous research, the distribution of bird communities in these forest‐farmland mosaics is not well understood. To achieve a comprehensive understanding of bird diversity and community turnover in a human‐modified Kenyan landscape, we recorded bird communities at 20 sites covering the complete habitat gradient from forest (near natural forest, secondary forest) to farmland (subsistence farmland, sugarcane plantation) using point counts and distance sampling. Bird density and species richness were on average higher in farmland than in forest habitats. Within forest and farmland, bird density and species richness increased with vegetation structural diversity, i.e., were higher in near natural than in secondary forest and in subsistence farmland than in sugarcane plantations. Bird communities in forest and farmland habitats were very distinct and very few forest specialists occurred in farmland habitats. Moreover, insectivorous bird species declined in farmland habitats whereas carnivores and herbivores increased. Our study confirms that tropical farmlands can hardly accommodate forest specialist species. Contrary to most previous studies, our findings show that structurally rich tropical farmlands hold a surprisingly rich and distinct bird community that is threatened by conversion of subsistence farmland into sugarcane plantations. We conclude that conservation strategies in the tropics must go beyond rain forest protection and should integrate structurally heterogeneous agroecosystems into conservation plans that aim at maintaining the diverse bird communities of tropical forest‐farmland mosaics.     

Temporal variation in the influence of forest succession on caterpillar communities: A long‐term study in a tropical dry forest

Forest succession can influence herbivore communities through changes in host availability, plant quality, microclimate, canopy structure complexity and predator abundance. It is not well known, however, if such influence is constant across years. Caterpillars have been reported to be particularly susceptible to changes in plant community composition across forest succession, as most species are specialists and rely on the presence of their hosts. Nevertheless, in the case of tropical dry forests, plant species have less defined successional boundaries than tropical wet forests, and hence herbivore communities should be able to persist across different successional stages. To test this prediction, caterpillar communities were surveyed during eight consecutive years in a tropical dry forest in four replicated successional stages in Chamela, Jalisco and Mexico. Lepidopteran species richness and diversity were equivalent in mature forests and early successional stages, but a distinctive caterpillar community was found for the recently abandoned pastures. Species composition tended to converge among all four successional stages during the span of eight years. Overall, our results highlight the importance of both primary and secondary forest for the conservation of caterpillar biodiversity at a landscape level. We also highlight the relevance of long‐term studies when assessing the influence of forest succession to account for across year variation in species interactions and climatic factors. Abstract in French is available with online material.     

Bird Assemblages in Anthropogenic Habitats: Identifying a Suitability Gradient for Native Species in the Atlantic Forest

Traditional approaches to the study of species persistence in fragmented landscapes generally consider a binary classification of habitat being suitable or unsuitable; however, the range of human‐modified habitats within a region may offer a gradient of habitat suitability (or conservation value) for species. We identified such a gradient by comparing bird assemblages among contrasting land uses (pine plantations of different age, annual crops, clear cuts and cattle pastures) in the Upper Parana Atlantic forest. Bird assemblages and vegetation structure were characterized in an extensive area of 4400 km² in Argentina and Paraguay during the breeding seasons of 2005–2010. Similarity of bird assemblages between anthropogenic habitats and the native forest and the proportion of forest species increased with vegetation vertical structure, while the proportion of open‐area species decreased. As a consequence, mature tree plantations were the most suitable habitats for forest species and were mainly used by frugivores and bark insectivores. In contrast, open habitats were the least suitable habitat for forest species and were used primarily by insectivores. Human‐created habitats that are structurally complex can be used by a subset of forest species, and may improve functional connectivity and mitigate edge effects. The conservation of large tracks of native forests, however, is critical for the long‐term persistence of the entire bird assemblage, especially for native forest dependent species.     

Recovery of amphibian,reptile, bird and mammal diversity during secondary forest succession in the tropics

In tropical regions, many studies have focused on how vegetation and ecosystem processes recover following the abandonment of anthropogenic activities, but less attention has been given to the recovery patterns of vertebrates. Here we conduct a meta‐analysis (n = 147 studies) of amphibian, reptile, bird and mammal recovery during tropical secondary forest succession (i.e. natural regeneration). For each taxonomic group, we compared changes in species richness and compositional similarity during natural secondary succession to reference forests (mature or old growth forest). In addition, we evaluated the response of forest specialists and the change in bird and mammal functional groups during natural secondary succession in the tropical moist forest biome. Overall, species richness of all groups reached levels of the reference forests during natural secondary succession, but this was not the case for species compositional similarity. The delay in recovery of forest specialists may be the reason for the delay in recovery of species compositional similarity. Overall, vertebrate recovery increased with successional stage, but other potential predictors of diversity recovery, such as, the geographical setting (amphibian and reptile species compositional similarity recovered more rapidly on islands), rainfall (mammal species richness and compositional similarity recovered faster in regions of low rainfall), and the landscape context (amphibian, reptile and mammal species compositional similarity recovered faster in regions with more forest patches) influenced vertebrate recovery. These results demonstrate the important role of secondary forests in providing habitat for many vertebrates, but the slow recovery of species compositional similarity, forest specialists and some functional groups (e.g. insectivorous birds) highlighted the challenge of secondary forest persistence, and strongly argues for the continued protection of old growth/mature forest as habitat for forest specialists and as sources for secondary forest sites.     

Altitudinal and latitudinal distribution of east Malagasy forest bird communities

Aim To describe variation in forest bird communities with altitude and latitude. Location Eastern Madagascar. Methods Extraction of data from forest bird inventories conducted in eastern Madagascar. Results There is a strong decline in species richness with altitude, above about 1300 m. Below this altitude, species richness is about constant or declines slightly. Seventy-eight percent of species occurring regularly in forest are absent from at least one of low, mid-altitude or high altitude forest. Of eighty-seven species occurring regularly in forest, only four or possibly five have latitudinally limited distributions, over a latitudinal range of over 1200 km. Three or possibly four are limited to the northern two-thirds, and one appears to be at least much more common in the southern half. Main conclusions Eastern Malagasy rain forest birds show previously unanalysed variation in altitudinal distribution. There is much less latitudinal variation. Species currently considered threatened are concentrated in the lowland and high-altitude zones. This may be at least partly due to lack of survey effort giving the impression that these species are rare, but lowland forests at least are under great human pressure. Bird conservation initiatives would probably have most effect if targeted at lowland east Malagasy rain forest.     

Ecological association of habitats and bird species during the breeding season in southeastern New South Wales

Bird communities in southeastern New South Wales, as determined by classification and ordination techniques, were seldom discrete. The most cohesive assemblage was associated with wet sclerophyll and rainforest habitats. These moist forests also supported the richest avifaunas and had high population densities. Although most birds occurred throughout the region, most species were uncommon. Of the 116 species recorded, 51 (44%) were found on fewer than 10% of the 143 plots censused. Only 19 species occurred on more than half the plots, with six of these recorded on more than 90% of the plots. Other than a small group of birds primarily restricted to moist forest, only the most common and widely distributed species regularly occurred together. Species were grouped with habitats on an elevational gradient from coastal forests to the tablelands. Within this elevational range, species were associated with major structural features of the vegetation. In terms of bird assemblages, open habitats were distinguished from those with dense vegetation. Plots which had been recently logged were separated from mature forest. The results indicate that plans for wildlife management should consider both the assemblages of species which are associated with major forest habitats and the requirements of individual species.     

Game Vertebrate Densities in Hunted and Nonhunted Forest Sites in Manu National Park,Peru

Manu National Park of southern Peru is one of the most renowned protected areas in the world, yet large-bodied vertebrate surveys conducted to date have been restricted to Cocha Cashu Biological Station, a research station covering <0.06 percent of the 1.7 Mha park. Manu Park is occupied by >460 settled Matsigenka Amerindians, 300–400 isolated Matsigenka, and several, little-known groups of isolated hunter–gatherers, yet the impact of these native Amazonians on game vertebrate populations within the park remains poorly understood. On the basis of 1495 km of standardized line-transect censuses, we present density and biomass estimates for 23 mammal, bird, and reptile species for seven lowland and upland forest sites in Manu Park, including Cocha Cashu. We compare these estimates between hunted and nonhunted sites within Manu Park, and with other Neotropical forest sites. Manu Park safeguards some of the most species-rich and highest biomass assemblages of arboreal and terrestrial mammals ever recorded in Neotropical forests, most likely because of its direct Andean influence and high levels of soil fertility. Relative to Barro Colorado Island, seed predators and arboreal folivores in Manu are rare, and generalist frugivores specializing on mature fruit pulp are abundant. The impact of such a qualitative shift in the vertebrate community on the dynamics of plant regeneration, and therefore, on our understanding of tropical plant ecology, must be profound. Despite a number of external threats, Manu Park continues to serve as a baseline against which other Neotropical forests can be gauged. Abstract in Spanish is available at http://www.blackwell-synergy.com/loi/btp .     

Postdispersal Seed Fate of Tropical Montane Trees in an Agricultural Landscape, Southern Costa Rica

Variation in postdispersal seed fate is an important factor driving patterns of forest regeneration. Because most previous studies have not tracked final seed fate and have commonly equated seed removal with predation without considering the possibility of secondary dispersal, little is known about individual seed mortality factors in successional and degraded habitats. This study tracked the postdispersal fate of large-seeded tropical montane trees in abandoned pasture, young secondary forests, and small forest fragments during two consecutive years in an agricultural landscape in southern Costa Rica. The incidence of secondary dispersal by animals, scatterhoarding in particular, and the effects of seed burial on germination were measured. Overall, seeds survived through germination more often in secondary forests with high levels of mortality occurring in abandoned pastures and forest fragments. The majority of seed mortality was caused by rodent predation in forest fragments, insects and fungal pathogens in secondary forests, and a combination of desiccation, insects, and fungal pathogens in pastures. Seeds were frequently secondarily dispersed in larger forest fragments, whereas they were only rarely moved in pastures and secondary forests. Burial tended to improve germination in pastures and was important for an often scatterhoarded species, Otoba novogranatensis, in all habitats. The results of this study suggest that: (1) seed mortality factors differ in response to the type of habitat degradation; (2) large-seeded species have high potential for survival when dispersed to young secondary forests; and (3) seed removal is not a reliable proxy for seed predation, particularly in less degraded forest fragments.     

Secondary Succession and Indigenous Management in Semideciduous Forest Fallows of the Amazon Basin1

To the discussion on secondary succession in tropical forests, we bring data on three under‐addressed issues: understory as well as overstory changes, continuous as opposed to phase changes, and integration of forest succession with indigenous fallow management and plant uses. Changes in vegetation structure and species composition were analyzed in secondary forests following swidden agriculture in a semideciduous forest of Bolivian lowlands. Twenty‐eight fallows, stratified by four successional stages (early = 1–5 yr, intermediate = 6–10 yr, advanced = 12–20 yr, and older = 22–36 yr), and ten stands of mature forests were sampled. The overstory (plants ≥5 cm diameter at breast height [DBH]) was sampled using a 20 × 50 m plot and the understory (plants <5 cm DBH) in three nested 2 × 5 m subplots. Semistructured interviews provided information on fallow management. Canopy height, basal area, and liana density of the overstory increased with secondary forest age. The early stage had the lowest species density and diversity in the overstory, but the highest diversity in the understory. Species composition and abundance differentiated mature forests and early successional stage from other successional stages; however, species showed individualistic responses across the temporal gradient. A total of 123 of 280 species were useful with edible, medicinal, and construction plants being the most abundant for both over‐ and understories. Most of Los Gwarayo preferred mature forests for making new swidden, while fallows were valuable for crops, useful species, and regenerating timber species.     

Ecological Stoichiometry along a Gradient of Forest Succession: Bait Preferences of Litter Ants

We examined bait preferences of litter ants along a successional gradient of forests in the Atlantic Forest of Brazil. The ants preferred protein-based baits in secondary forests, yet preferred carbohydrate-based baits in old-growth forests. This suggests a shift in stoichiometric balances between secondary and old-growth forest which might have an impact on the functional interactions in these ecosystems. Therefore, the ecological services offered by secondary habitats may be quite different from those offered by old-growth forests.     

Forest patch size and isolation as drivers of bird species richness in Maputaland,Mozambique

Aim To examine the response of forest‐dependent and generalist bird assemblages and feeding guilds to patch characteristics of forest fragments in the context of island biogeography and metapopulation theories in an area that supports high levels of species diversity and endemism. Location Dune, riverine, sand and swamp forest fragments in southern Mozambique’s Maputaland. Methods We recorded 20 forest‐dependent and 69 generalist bird species at 220 survey points in 30 forest fragments that ranged in size from 5 to 7432 ha and that were 100 to 5100 m apart. We used linear regressions to relate the number of species to forest fragment size, isolation, perimeter, fractal dimension, shape and core area for frugivores, insectivores, granivores, nectarivores, carnivores and omnivores separately for forest‐dependent and generalist bird assemblages. We tested for modality in occupancy frequency distributions of species in forest fragments and used the binary matrix temperature calculator to determine whether the assemblages had a nested structure separately for forest‐dependent and generalist bird species. Results The number of forest‐dependent and generalist bird species categorized into the feeding guilds varied independently from forest patch parameter values. Occupancy models showed a random distribution for forest‐dependent birds, and generalists had a unimodal distribution. Both the forest‐dependent and generalist bird assemblages had a non‐nested structure. The presence of eight rare forest‐dependent and four endemic bird species in 22 of the 30 fragments contributed to the non‐nested structure of these bird assemblages. Main conclusions Fragmented forests did not induce the expected responses of the forest bird assemblages in that the number of species was not explained by patch characteristics, nor could we find evidence for metapopulation dynamics. Non‐nestedness may be caused by the few rare and endemic species occupying forest fragments with a wide range of patch characteristics. Knowledge of the response of species to forest fragmentation can benefit conservation planning catering for rare and endemic species. Our study suggests that all forest fragments in the region may have to be included in a conservation network to complement existing landscape‐based plans to protect the region.