Evaluating the legacy of landscape history: extinction debt and species credit in bird and small mammal assemblages in the Brazilian Atlantic Forest

There is now an extensive literature on extinction debt following deforestation. However, the potential for species credit in landscapes that have experienced a change from decreasing to expanding forest cover has received little attention. Both delayed responses should depend on current landscape forest cover and on species life-history traits, such as longevity, as short-lived species are likely to respond faster than long-lived species. We evaluated the effects of historical and present-day local forest cover on two vertebrate groups with different longevities understorey birds and non-flying small mammals - in forest patches at three Atlantic Forest landscapes. Our work investigated how the probability of extinction debt and species credit varies (i) amongst landscapes with different proportions of forest cover and distinct trajectories of forest cover change, and (ii) between taxa with different life spans. Our results suggest that the existence of extinction debt and species credit, as well as the potential for their future payment and/or receipt, is not only related to forest cover trajectory but also to the amount of remaining forest cover at the landscape scale. Moreover, differences in bird and small mammal life spans seem to be insufficient to affect differently their probability of showing time-delayed responses to landscape change. Synthesis and applications. Our work highlights the need for considering not only the trajectory of deforestation/regeneration but also the amount of forest cover at landscape scale when investigating time-delayed responses to landscape change. As many landscapes are experiencing a change from decreasing to expanding forest cover, understanding the association of extinction and immigration processes, as well as their interactions with the landscape dynamic, is a key factor to plan conservation and restoration actions in human-altered landscapes.

THE CONVERSION OF THE ATLANTIC FOREST IN ANTHROPOGENIC LANDSCAPES: LESSONS FOR THE CONSERVATION OF BIOLOGICAL DIVERSITY OF TROPICAL FORESTS

The Brazilian Atlantic forest has been an excellent laboratory for investigations regarding tropical forest ecology and the fragility of tropical ecosystems in face of human disturbances. In this article, we present a synthesis about the spatial distribution of Atlantic forest biodiversity and forest response to human disturbances, as well as the ongoing conservation efforts based on a review of several investigations in this biota. In general, studies have documented an uneven distribution of biodiversity throughout the Atlantic forest region, revealing alarming rates of habitat loss at low altitudes, while protected areas concentrate at higher altitudes. It has been suggested that the remaining forest habitat is moving towards an early-successional systems across human-modified landscapes. Such regressive forest succession increases the threats for several animals and plant groups. Based on these findings, we propose seven guidelines in order to enhance the provision of ecosystem services and the conservation value of human-modified landscapes, reducing the species extinction risk in the Atlantic forest and in other irreplaceable tropical biotas.

Unraveling the drivers of community dissimilarity and species extinction in fragmented landscapes

Communities in fragmented landscapes are often assumed to be structured by species extinction due to habitat loss, which has led to extensive use of the species-area relationship (SAR) in fragmentation studies. However, the use of the SAR presupposes that habitat loss leads species to extinction but does not allow for extinction to be offset by colonization of disturbed-habitat specialists. Moreover, the use of SAR assumes that species richness is a good proxy of community changes in fragmented landscapes. Here, we assessed how communities dwelling in fragmented landscapes are influenced by habitat loss at multiple scales; then we estimated the ability of models ruled by SAR and by species turnover in successfully predicting changes in community composition, and asked whether species richness is indeed an informative community metric. To address these issues, we used a data set consisting of 140 bird species sampled in 65 patches, from six landscapes with different proportions of forest cover in the Atlantic Forest of Brazil. We compared empirical patterns against simulations of over 8 million communities structured by different magnitudes of the power-law SAR and with species-specific rules to assign species to sites. Empirical results showed that, while bird community composition was strongly influenced by habitat loss at the patch and landscape scale, species richness remained largely unaffected. Modeling results revealed that the compositional changes observed in the Atlantic Forest bird metacommunity were only matched by models with either unrealistic magnitudes of the SAR or by models ruled by species turnover, akin to what would be observed along natural gradients. We show that, in the presence of such compositional turnover, species richness is poorly correlated with species extinction, and z values of the SAR strongly underestimate the effects of habitat loss. We suggest that the observed compositional changes are driven by each species reaching its individual extinction threshold: either a threshold of forest cover for species that disappear with habitat loss, or of matrix cover for species that benefit from habitat loss.

Production and carbon allocation in monocultures and mixed-species plantations of Eucalyptus grandis and Acacia mangium in Brazil

Introducing nitrogen-fixing tree species in fast-growing eucalypt plantations has the potential to improve soil nitrogen availability compared with eucalypt monocultures. Whether or not the changes in soil nutrient status and stand structure will lead to mixtures that out-yield monocultures depends on the balance between positive interactions and the negative effects of interspecific competition, and on their effect on carbon (C) uptake and partitioning. We used a C budget approach to quantify growth, C uptake and C partitioning in monocultures of Eucalyptus grandis (W. Hill ex Maiden) and Acacia mangium (Willd.) (treatments E100 and A100, respectively), and in a mixture at the same stocking density with the two species at a proportion of 1 : 1 (treatment MS). Allometric relationships established over the whole rotation, and measurements of soil CO2 efflux and aboveground litterfall for ages 4-6 years after planting were used to estimate aboveground net primary production (ANPP), total belowground carbon flux (TBCF) and gross primary production (GPP). We tested the hypotheses that (i) species differences for wood production between E. grandis and A. mangium monocultures were partly explained by different C partitioning strategies, and (ii) the observed lower wood production in the mixture compared with eucalypt monoculture was mostly explained by a lower partitioning aboveground. At the end of the rotation, total aboveground biomass was lowest in A100 (10.5 kg DM m(-2)), intermediate in MS (12.2 kg DM m(-2)) and highest in E100 (13.9 kg DM m(-2)). The results did not support our first hypothesis of contrasting C partitioning strategies between E. grandis and A. mangium monocultures: the 21% lower growth (delta B-w) in A100 compared with E100 was almost entirely explained by a 23% lower GPP, with little or no species difference in ratios such as TBCF/GPP, ANPP/TBCF, delta B-w/ANPP and delta B-w/GPP. In contrast, the 28% lower delta B-w in MS than in E100 was explained both by a 15% lower GPP and by a 15% lower fraction of GPP allocated to wood growth, thus partially supporting our second hypothesis: mixing the two species led to shifts in C allocations from above- to belowground, and from growth to litter production, for both species.

A model of road effect using line integrals and a test of the performance of two new road indices using the distribution of small mammals in an Atlantic Forest landscape

Effects of roads on wildlife and its habitat have been measured using metrics, such as the nearest road distance, road density, and effective mesh size. In this work we introduce two new indices: (1) Integral Road Effect (IRE), which measured the sum effects of points in a road at a fixed point in the forest; and (2) Average Value of the Infinitesimal Road Effect (AVIRE), which measured the average of the effects of roads at this point. IRE is formally defined as the line integral of a special function (the infinitesimal road effect) along the curves that model the roads, whereas AVIRE is the quotient of IRE by the length of the roads. Combining tools of ArcGIS software with a numerical algorithm, we calculated these and other road and habitat cover indices in a sample of points in a human-modified landscape in the Brazilian Atlantic Forest, where data on the abundance of two groups of small mammals (forest specialists and habitat generalists) were collected in the field. We then compared through the Akaike Information Criterion (AIC) a set of candidate regression models to explain the variation in small mammal abundance, including models with our two new road indices (AVIRE and IRE) or models with other road effect indices (nearest road distance, mesh size, and road density), and reference models (containing only habitat indices, or only the intercept without the effect of any variable). Compared to other road effect indices, AVIRE showed the best performance to explain abundance of forest specialist species, whereas the nearest road distance obtained the best performance to generalist species. AVIRE and habitat together were included in the best model for both small mammal groups, that is, higher abundance of specialist and generalist small mammals occurred where there is lower average road effect (less AVIRE) and more habitat. Moreover, AVIRE was not significantly correlated with habitat cover of specialists and generalists differing from the other road effect indices, except mesh size, which allows for separating the effect of roads from the effect of habitat on small mammal communities. We suggest that the proposed indices and GIS procedures could also be useful to describe other spatial ecological phenomena, such as edge effect in habitat fragments. (C) 2012 Elsevier B.V. All rights reserved.

SEASONAL INCREMENT IN TRUNK DIAMETER OF Eucalyptus grandis TREES APPLYING DENDROMETER BANDS

The present work aimed to evaluate the seasonal increment in diameter of Eucalyptus grandis trees for 24 months and its relationship with the climatic variables and fertilization with nitrogen and with sewer mud. The trees were planted in the spacing of 3 x 2 m and fertilized with nitrogen (planting, 6, 12, 18 months) and sewer mud (planting and 8 months). 20 trees were selected by treatment according witch the distribution of basal area and installed dendrometer bands at a 1.3 meter. The results showed a clear effect of the climatic variables on the seasonal increment in diameter of trees, being observed a delay period (lag) of 28 days for the answer of the trees in relation to the climatic variables. Regading to the fertilization effect, it was observed that the increment of trunk diameter was higher in the eucalypt trees with organic in relation to mineral fertilization with nitrogen.

Stand-level patterns of carbon fluxes and partitioning in a Eucalyptus grandis plantation across a gradient of productivity, in Sao Paulo State, Brazil

Wood production represents a large but variable fraction of gross primary production (GPP) in highly productive Eucalyptus plantations. Assessing patterns of carbon (C) partitioning (C flux as a fraction of GPP) between above- and belowground components is essential to understand mechanisms driving the C budget of these plantations. Better knowledge of fluxes and partitioning to woody and non-woody tissues in response to site characteristics and resource availability could provide opportunities to increase forest productivity. Our study aimed at investigating how C allocation varied within one apparently homogeneous 90 ha stand of Eucalyptus grandis (W. Hill ex Maiden) in Southeastern Brazil. We assessed annual above-ground net primary production (ANPP: stem, leaf, and branch production) and total belowground C flux (TBCF: the sum of root production and respiration and mycorrhizal production and respiration), GPP (computed as the sum of ANPP, TBCF and estimated aboveground respiration) on 12 plots representing the gradient of productivity found within the stand. The spatial heterogeneity of topography and associated soil attributes across the stand likely explained this fertility gradient. Component fluxes of GPP and C partitioning were found to vary among plots. Stem NPP ranged from 554 g C m(-2) year(-1) on the plot with lowest GPP to 923 g C m(-2) year(-1) on the plot with highest GPP. Total belowground carbon flux ranged from 497 to 1235 g C m(-2) year(-1) and showed no relationship with ANPP or GPP. Carbon partitioning to stem NPP increased from 0.19 to 0.23, showing a positive trend of increase with GPP (R-2 = 0.29, P = 0.07). Variations in stem wood production across the gradient of productivity observed at our experimental site were a result of the variability in C partitioning to different forest system components.

Land-use and land-cover change in Atlantic Forest landscapes

The Atlantic Forest is one of the most threatened tropical biomes, with much of the standing forest in small (less than 50 ha), disturbed and isolated patches. The pattern of land-use and land-cover change (LULCC) which has resulted in this critical scenario has not yet been fully investigated. Here, we describe the LULCC in three Atlantic Forest fragmented landscapes (Sao Paulo, Brazil) between 1960-1980s and 1980-2000s. The three studied landscapes differ in the current proportion of forest cover, having 10%, 30% and 50% respectively. Between the 1960s and 1980s. forest cover of two landscapes was reduced while the forest cover in the third landscape increased slightly. The opposite trend was observed between the 1980s and 2000s: forest regeneration was greater than deforestation at the landscapes with 10% and 50% of forest cover and, as a consequence, forest cover increased. By contrast, the percentage of forest cover at the landscape with 30% of forest cover was drastically reduced between the 1980s and 2000s. LULCC deviated from a random trajectory, were not constant through time in two study landscapes and were not constant across space in a given time period. This landscape dynamism in single locations over small temporal scales is a key factor to be considered in models of LULCC to accurately simulate future changes for the Atlantic Forest. In general, forest patches became more isolated when deforestation was greater than forest regeneration and became more connected when forest regeneration was greater than deforestation. As a result of the dynamic experienced by the study landscapes, individual forest patches currently consist of a mosaic of different forest age classes which is likely to impact bio-diversity. Furthermore, landscape dynamics suggests the beginning of a forest transition in some Atlantic Forest regions, what could be of great importance for biodiversity conservation due to the potential effects of young secondary forests in reducing forest isolation and maintaining a significant amount of the original biodiversity. (C) 2012 Elsevier B.V. All rights reserved.

EFEITO DA IDADE E POSIÇÃO DE AMOSTRAGEM NA DENSIDADE E CARACTERÍSTICAS ANATÔMICAS DA MADEIRA DE Eucalyptus grandis

The knowledge of the variations in the wood characteristics produced by eucalyptus trees according to age and sampling positions is essential for its proper use. This study had as objective to evaluate the influence of the age, longitudinal and radial positions on basic density and anatomical characteristics in Eucalyptus grandis wood. The trees were planted in 3x2 m spacing and fertilized with commercial fertilizers in planting, 6th and 12th months. According to basal area distribution, fifteen trees were selected (24, 36 and 72 months of age) - five trees per age. Disks at DBH position (1.3 m) were taken for fiber determination (length, wall thickness, lumen diameter and width) and vessels (tangential diameter, frequency and area occupied) and in other different sampling positions for basic density determination. Wood basic density increased from 0.43 to 0.46 g.cm(-3) as well as the trees age increases with a longitudinal variation model, characterized through a decrease in base-3m (0.42-0.49 -> 0.40-0.46 g.cm(-3)) and an increase to the top of the trunk (0.46 -> 0.54 g.cm(-3)) Fibers and vessels dimensions showed variations related to age and to pit-bark direction. Wood properties behavior and variations indicate that, until this period, the juvenile wood is being formed.

Introducing Acacia mangium trees in Eucalyptus grandis plantations: consequences for soil organic matter stocks and nitrogen mineralization

Background and aims Eucalyptus plantations cover 20 million hectares on highly weathered soils. Large amounts of nitrogen (N) exported during harvesting lead to concerns about their sustainability. Our goal was to assess the potential of introducing A. mangium trees in highly productive Eucalyptus plantations to enhance soil organic matter stocks and N availability. Methods A randomized block design was set up in a Brazilian Ferralsol soil to assess the effects of mono-specific Eucalyptus grandis (100E) and Acacia mangium (100A) stands and mixed plantations (50A:50E)on soil organic matter stocks and net N mineralization. Results A 6-year rotation of mono-specific A. mangium plantations led to carbon (C) and N stocks in the forest floor that were 44% lower and 86% higher than in pure E. grandis stands, respectively. Carbon and N stocks were not significantly different between the three treatments in the 0-15 cm soil layer. Field incubations conducted every 4 weeks for the two last years of the rotation estimated net soil N mineralization in 100A and 100E at 124 and 64 kg ha(-1) yr(-1), respectively. Nitrogen inputs to soil with litterfall were of the same order as net N mineralization. Conclusions Acacia mangium trees largely increased the turnover rate of N in the topsoil. Introducing A. mangium trees might improve mineral N availability in soils where commercial Eucalyptus plantations have been managed for a long time.

How deforestation pattern in the Amazon influences vertebrate richness and community composition

The effects of habitat configuration on species persistence are predicted to be most apparent when remaining habitat cover is below 30%. We tested this prediction by comparing vertebrate communities in 21 landscapes located in the southern Amazonia, including 7 control landscapes (similar to 100% of forest cover) and 14 fragmented landscapes (4 x 4 km). The fragmented landscapes retained similar proportions of forest (similar to 25%), but had contrasting configurations, resulting from two different deforestation patterns: the "fish-bone pattern" common in small properties, and the large-property pattern generally used by large ranchers. Vertebrates were surveyed in all landscapes in February-July 2009 with interviews (n = 150). We found a significant difference in reported species richness among the fish-bone, large-property, and control areas (mean = 29.3, 38.8 and 43.5 respectively). Control areas and large-properties tended to have a higher number of specialist species (mean = 13.7, and 11.7, respectively), when compared with the fish-bone pattern (5.1). Vertebrate community composition in the control and large-properties was more similar to one another than to those of the fish-bone landscapes. The number of fragments was the main factor affecting the persistence of species, being negatively associated with specialist species richness. Species richness was also positively related with the size of the largest fragment structurally connected to the studied landscapes (i.e., a regional scale effect). Our results demonstrated that the large-property pattern, which results in less fragmented landscapes, can maintain a more diverse community of large vertebrates, including top predators, which are considered fundamental for maintaining ecosystem integrity. These results support the hypothesis that landscape configuration contributes to the persistence and/or extirpation of species.

Decisions on Temporal Sampling Protocol Influence the Detection of Ecological Patterns

Within-site variability in species detectability is a problem common to many biodiversity assessments and can strongly bias the results. Such variability can be caused by many factors, including simple counting inaccuracies, which can be solved by increasing sample size, or by temporal changes in species behavior, meaning that the way the temporal sampling protocol is designed is also very important. Here we use the example of mist-netted tropical birds to determine how design decisions in the temporal sampling protocol can alter the data collected and how these changes might affect the detection of ecological patterns, such as the species-area relationship (SAR). Using data from almost 3400 birds captured from 21,000 net-hours at 31 sites in the Brazilian Atlantic Forest, we found that the magnitude of ecological trends remained fairly stable, but the probability of detecting statistically significant ecological patterns varied depending on sampling effort, time of day and season in which sampling was conducted. For example, more species were detected in the wet season, but the SAR was strongest in the dry season. We found that the temporal distribution of sampling effort was more important than its total amount, discovering that similar ecological results could have been obtained with one-third of the total effort, as long as each site had been equally sampled over 2 yr. We discuss that projects with the same sampling effort and spatial design, but with different temporal sampling protocol are likely to report different ecological patterns, which may ultimately lead to inappropriate conservation strategies.

The importance of landscape structure for seed dispersal in rain forest fragments

Questions What are the main features of the seed rain in a fragmented Atlantic forest landscape? Can seed rain species attributes (life form, dispersal mode, successional status) relate to the spatial arrangement (size and number of fragments, edge density and presence of corridor) of forest fragments in the landscape? How does the rain forest landscape structure affect the seed rain? Location Atlantic rainforest, Sao Paulo State, Southeastern Brazil. Methods Seed rain samples were collected monthly throughout 1yr, counted, identified and classified according to species dispersal mode, successional status and life form. Seed rain composition was compared with woody species near the seed traps. Relationships between seed rain composition and landscape spatial arrangement (fragment area, presence of corridor, number of fragments in the surroundings, proximity of fragments, and edge density) were tested using canonical correspondence analysis (CCA). Results We collected 20142 seeds belonging to 115 taxa, most of them early successional and anemochorous trees. In general, the seed rain had a species composition distinct from that of the nearby forest tree community. Small isolated fragments contained more seeds, mainly of anemochorous, epiphytic and early-successional species; large fragments showed higher association with zoochorous and late-successional species compared to small fragments. The CCA significantly distinguished the species dispersal mode according to fragment size and isolation, anemochorous species being associated to small and isolated fragments, and zoochorous species to larger areas and fragment aggregation. Nevertheless, a gradient driven by proximity (PROX) and edge density (ED) segregated lianas (in the positive extremity), early successional and epiphyte species (in the negative end); large fragments were positively associated to PROX and ED. Conclusions The results highlight the importance of the size and spatial arrangement of forest patches to promote habitat connectivity and improve the flux of animal-dispersed seeds. Landscape structure controls seed fluxes and affects plant dispersal capacity, potentially influencing the composition and structure of forest fragments. The seed rain composition may be used to assess the effects of landscape spatial structure on plant assemblages, and provide relevant information for biodiversity conservation.

Do changes in carbon allocation account for the growth response to potassium and sodium applications in tropical Eucalyptus plantations?

Understanding the underlying mechanisms that account for the impact of potassium (K) fertilization and its replacement by sodium (Na) on tree growth is key to improving the management of forest plantations that are expanding over weathered tropical soils with low amounts of exchangeable bases. A complete randomized block design was planted with Eucalyptus grandis (W. Hill ex Maiden) to quantify growth, carbon uptake and carbon partitioning using a carbon budget approach. A combination of approaches including the establishment of allometric relationships over the whole rotation and measurements of soil CO2 efflux and aboveground litterfall at the end of the rotation were used to estimate aboveground net production (ANPP), total belowground carbon flux and gross primary production (GPP). The stable carbon isotope (delta C-13) of stem wood alpha-cellulose produced every year was used as a proxy for stomatal limitation of photosynthesis. Potassium fertilization increased GPP and decreased the fraction of carbon allocated belowground. Aboveground net production was strongly enhanced, and because leaf lifespan increased, leaf biomass was enhanced without any change in leaf production, and wood production (P-W) was dramatically increased. Sodium application decreased the fraction of carbon allocated belowground in a similar way, and enhanced GPP, ANPP and P-W, but to a lesser extent compared with K fertilization. Neither K nor Na affected delta C-13 of stem wood alpha-cellulose, suggesting that water-use efficiency was the same among the treatments and that the inferred increase in leaf photosynthesis was not only related to a higher stomatal conductance. We concluded that the response to K fertilization and Na addition on P-W resulted from drastic changes in carbon allocation.

Associations of Forest Cover, Fragment Area, and Connectivity with Neotropical Understory Bird Species Richness and Abundance

Theoretical and empirical studies demonstrate that the total amount of forest and the size and connectivity of fragments have nonlinear effects on species survival. We tested how habitat amount and configuration affect understory bird species richness and abundance. We used mist nets (almost 34,000 net hours) to sample birds in 53 Atlantic Forest fragments in southeastern Brazil. Fragments were distributed among 3 10,800-ha landscapes. The remaining forest in these landscapes was below (10% forest cover), similar to (30%), and above (50%) the theoretical fragmentation threshold (approximately 30%) below which the effects of fragmentation should be intensified. Species-richness estimates were significantly higher (F = 3715, p = 0.00) where 50% of the forest remained, which suggests a species occurrence threshold of 30-50% forest, which is higher than usually occurs (<30%). Relations between forest cover and species richness differed depending on species sensitivity to forest conversion and fragmentation. For less sensitive species, species richness decreased as forest cover increased, whereas for highly sensitive species the opposite occurred. For sensitive species, species richness and the amount of forest cover were positively related, particularly when forest cover was 30-50%. Fragment size and connectivity were related to species richness and abundance in all landscapes, not just below the 30% threshold. Where 10% of the forest remained, fragment size was more related to species richness and abundance than connectivity. However, the relation between connectivity and species richness and abundance was stronger where 30% of the landscape was forested. Where 50% of the landscape was forested, fragment size and connectivity were both related to species richness and abundance. Our results demonstrated a rapid loss of species at relatively high levels of forest cover (30-50%). Highly sensitive species were 3-4 times more common above the 30-50% threshold than below it; however, our results do not support a unique fragmentation threshold.