Edge effects as the principal cause of area effects on birds in fragmented secondary forest

Bird communities in tropical forests are strongly affected by both patch area and habitat edges. The fact that both effects are intrinsically confounded in space raises questions about how these two widely reported ecological patterns interact, and whether they are independent or simply different spatial manifestations of the same phenomenon. Moreover, do small patches of secondary forest, in landscapes where the most sensitive species have gone locally extinct, exhibit similar patterns to those previously observed in fragmented and continuous primary forests? We addressed these questions by testing edge-related differences in vegetation structure and bird community composition at 31 sites in fragmented and continuous landscapes in the imperilled Atlantic forest of Brazil. Over a two-year period, birds were captured with mist nets to a standardized effort of 680 net-hours at each site (similar to 22 000 net-hours resulting in 3381 captures from 114 species). We found that the bird community in patches of secondary forest was degraded in species composition compared to primary continuous forest, but still exhibited a strong response to edge effects. In fragmented secondary forests, edge and area effects also interacted, such that the magnitude of edge to interior differences on bird community composition declined markedly with patch size. The change in bird species composition between forest interiors and edges was similar to the change in community composition between large and small patches (because species had congruent responses to edge and area), but after controlling for edge effects community composition was no longer affected by patch area. Our results show that although secondary forests hold an impoverished bird community, ecological patterns such as area and edge effects are similar to those reported for primary forests. Our data provide further evidence that edge effects are the main drivers of area effects in fragmented landscapes.

Relief influence on tree species richness in secondary forest fragments of Atlantic Forest, SE, Brazil

(Relief influence on tree species richness in secondary forest fragments of Atlantic Forest, SE, Brazil). The aim of this work was to explore the relationship between tree species richness and morphological characteristics of relief at the Ibiuna Plateau (SE Brazil). We sampled 61 plots of 0.30 ha, systematically established in 20 fragments of secondary forest (2-274 ha) and in three areas within a continuous secondary forest site, Morro Grande Reserve (9,400 ha). At each plot, 100 trees with diameter at breast height > 5 cm were sampled by the point centered quarter method, and total richness and richness per dispersal and succession class were obtained. The relief was characterized by the mean and variance of slope, elevation, aspect and slope location. There was no significant relationship between relief heterogeneity and tree species richness. Relief parameters generally did not affect tree richness, but elevation was particularly important especially in the continuous forest. Despite the limited range of altitudinal variation (150 m), species richness increases with elevation. The highest areas were also those with the largest forest cover and the lowest disturbance degree, which should contribute to the greater richness of those sites. Our results suggest an indirect influence of relief, due to the fact that deforestation is less intense in higher regions, rather than a direct influence of abiotic factors related to the altitudinal gradient.

A comparison of the leaf-litter ant fauna in a secondary atlantic forest with an adjacent pine plantation in southeastern Brazil

We investigated the influence of Pinus afforestation on the structure of leaf-litter ant communities in the southeastern Brazilian Atlantic Forest, studying an old secondary forest and a nearly 30 year-old never managed Pinus elliottii reforested area. A total of 12,826 individual ants distributed among 95 species and 32 genera were obtained from 50 1 m² samples/ habitat. Of these, 60 species were recorded in the pine plantation and 82 in the area of Atlantic forest; almost 50% of the species found in the secondary forest area were also present in the pine plantation. The number of species per sample was significantly higher in the secondary forest than in the pine plantation. Forest-adapted taxa are the most responsible for ant species richness differences between areas, and the pine plantation is richer in species classified as soil or litter omnivorous-dominants. The specialized ant predators registered in the pine plantation, as seven Dacetini, two Basiceros, two Attini and two Discothyrea, belong to widely distributed species. The NMDS (non-metric multidimensional scaling) ordination also suggested strong differences in similarity among samples of the two areas. Furthermore, this analysis indicated higher sample heterogeneity in the secondary forest, with two clusters of species, while in the pine plantation the species belong to a single cluster. We applied the ant mosaic hypothesis to explain the distribution of the leaf-litter fauna and spatial autocorrelation tests among samples. We argue that the results are likely related to differences in quality and distribution of the leaf-litter between the pine plantation and the secondary area.

Ant community richness and composition across a gradient from Eucalyptus plantations to secondary Atlantic Forest

Secondary forests and exotic tree plantations are expanding across tropical landscapes. However, our current understanding of the value of these human-dominated forest landscapes for invertebrate biodiversity conservation is still very poor. In this paper, we use the leaf-litter ant fauna to assess invertebrate diversity in one commercially managed Eucalyptus plantation (four years old), two abandoned plantations of different regeneration ages (16 and 31 years), and one neighboring secondary Atlantic Forest in Southeastern Brazil. There was a clear gradient in species richness from the secondary forest to the managed Eucalyptus plantation; richness and diversity peaked in secondary forest and in the older regenerating Eucalyptus plantation. Significantly more species were recorded in secondary forest samples than in Eucalyptus plantations, but Eucalyptus plantations had a similar level of richness. Furthermore, a non-metric multidimensional scaling analysis revealed clear differences in species composition between the younger managed Eucalyptus plantation (understory absent) and habitats with sub-developed or developed understory. Eucalyptus plantations were characterized by an assemblage of widespread, generalist species very different from those known to occur in core forest habitats of southeastern Brazil. Our results indicate that while older regenerating Eucalyptus plantations can provide habitat to facilitate the persistence of generalist ant species, it is unlikely to conserve most of the primary forest species, such as specialized predators, Dacetini predators, and nomadic species.

Influence of Post-Clearing Treatment on the Recovery of Herbaceous Plant Communities in Amazonian Secondary Forests

Secondary forests are an increasingly common feature in tropical landscapes worldwide and understanding their regeneration is necessary to design effective restoration strategies. It has previously been shown that the woody species community in secondary forests can follow different successional pathways according to the nature of past human activities in the area, yet little is known about patterns of herbaceous species diversity in secondary forests with different histories of land use. We compared the diversity and abundance of herbaceous plant communities in two types of Central Amazonian secondary forests-those regenerating on pastures created by felling and burning trees and those where trees were felled only. We also tested if plant density and species richness in secondary forests are related to proximity to primary forest. In comparison with primary forest sites, forests regenerating on non-burned habitats had lower herbaceous plant density and species richness than those on burned ones. However, species composition and abundance in non-burned stands were more similar to those of primary forest, whereas several secondary forest specialist species were found in burned stands. In both non-burned and burned forests, distance from the forest edge was not related to herbaceous density and species richness. Overall, our results suggest that the natural regeneration of herbaceous species in secondary tropical forests is dependent on a site`s post-clearing treatment. We recommend evaluating the land history of a site prior to developing and implementing a restoration strategy, as this will influence the biological template on which restoration efforts are overlaid.

Micronutrient dynamics in the forest biomass in succession stages of a seasonal deciduous forest, RS, Brazil

There is a great need of research to assess the behavior of micronutrients in natural forests of southern Brazil. Do to this need, the objective of this work was to study the levels and amounts of micronutrients in forest above ground biomass of the forest, in a comparative way, in two secondary succession stages (SSS) in a Seasonal Deciduous Forest in Rio Grande do Sul, Brazil. The SSS had enjoyed 35 and 55 years of regeneration since the end of agricultural use, respectively for initial secondary forest (ISF) and late secondary forest (LSF). The above-ground biomass was collected and separated into vegetative strata and these in fractions, thereafter chemically analyzed for the levels of B, Fe, Zn, Mn and Cu. Leaf fractions of arboreal, shrubs and herbaceous strata showed the highest levels for most nutrients. Only the levels of iron and manganese were higher in the bark fraction, for both sucession stages. In the LSF, the herbaceous stratum also showed high levels of Fe. The average levels of micronutrients showed differences between the two sucession stages only in relation to Fe and Mn, with higher levels in LSF biomass. The amount of nutrients stored was always higher in LSF, because of the largest biomass and the higher levels of Fe and Mn in the biomass of this SSS. The quantitative order of nutrient storage in biomass was Fe> Mn> Zn> B> Cu.

Prospects for biodiversity conservation in the Atlantic Forest: Lessons from aging human-modified landscapes

Recent global assessments have shown the limited coverage of protected areas across tropical biotas, fuelling a growing interest in the potential conservation services provided by anthropogenic landscapes. Here we examine the geographic distribution of biological diversity in the Atlantic Forest of South America, synthesize the most conspicuous forest biodiversity responses to human disturbances, propose further conservation initiatives for this biota, and offer a range of general insights into the prospects of forest species persistence in human-modified tropical forest landscapes worldwide. At the biome scale, the most extensive pre-Columbian habitats across the Atlantic Forest ranged across elevations below 800 masl, which still concentrate most areas within the major centers of species endemism. Unfortunately, up to 88% of the original forest habitat has been lost, mainly across these low to intermediate elevations, whereas protected areas are clearly skewed towards high elevations above 1200 masl. At the landscape scale, most remaining Atlantic Forest cover is embedded within dynamic agro-mosaics including elements such as small forest fragments, early-to-late secondary forest patches and exotic tree mono-cultures. In this sort of aging or long-term modified landscapes, habitat fragmentation appears to effectively drive edge-dominated portions of forest fragments towards an early-successional system, greatly limiting the long-term persistence of forest-obligate and forest-dependent species. However, the extent to which forest habitats approach early-successional systems, thereby threatening the bulk of the Atlantic Forest biodiversity, depends on both past and present landscape configuration. Many elements of human-modified landscapes (e.g. patches of early-secondary forests and tree mono-cultures) may offer excellent conservation opportunities, but they cannot replace the conservation value of protected areas and hitherto unprotected large patches of old-growth forests. Finally, the biodiversity conservation services provided by anthropogenic landscapes across Atlantic Forest and other tropical forest regions can be significantly augmented by coupling biodiversity corridor initiatives with biota-scale attempts to plug existing gaps in the representativeness of protected areas. (C) 2010 Elsevier Ltd. All rights reserved.

Time-lag in biological responses to landscape changes in a highly dynamic Atlantic forest region

Time-lagged responses of biological variables to landscape modifications are widely recognized, but rarely considered in ecological studies. In order to test for the existence of time-lags in the response of trees, small mammals, birds and frogs to changes in fragment area and connectivity, we studied a fragmented and highly dynamic landscape in the Atlantic forest region. We also investigated the biological correlates associated with differential responses among taxonomic groups. Species richness and abundance for four taxonomic groups were measured in 21 secondary forest fragments during the same period (2000-2002), following a standardized protocol. Data analyses were based on power regressions and model selection procedures. The model inputs included present (2000) and past (1962, 1981) fragment areas and connectivity, as well as observed changes in these parameters. Although past landscape structure was particularly relevant for trees, all taxonomic groups (except small mammals) were affected by landscape dynamics, exhibiting a time-lagged response. Furthermore, fragment area was more important for species groups with lower dispersal capacity, while species with higher dispersal ability had stronger responses to connectivity measures. Although these secondary forest fragments still maintain a large fraction of their original biodiversity, the delay in biological response combined with high rates of deforestation and fast forest regeneration imply in a reduction in the average age of the forest. This also indicates that future species losses are likely, especially those that are more strictly-forest dwellers. Conservation actions should be implemented to reduce species extinction, to maintain old-growth forests and to favour the regeneration process. Our results demonstrate that landscape history can strongly affect the present distribution pattern of species in fragmented landscapes, and should be considered in conservation planning. (C) 2009 Elsevier Ltd. All rights reserved.

Using indicators of deforestation and land-use dynamics to support conservation strategies: A case study of central Rondonia, Brazil

In Rondonia State, Brazil, settlement processes have cleared 68,000 km 2 of tropical forests since the 1970s. The intensity of deforestation has differed by region depending on driving factors like roads and economic activities. Different histories of land-use activities and rates of change have resulted in mosaics of forest patches embedded in an agricultural matrix. Yet, most assessments of deforestation and its effects on vegetation, soil and water typically focus on landscape patterns of current conditions, yet historical deforestation dynamics can influence current conditions strongly. Here, we develop and describe the use of four land-use dynamic indicators to capture historical land-use changes of catchments and to measure the rate of deforestation (annual deforestation rate), forest regeneration level (secondary forest mean proportion), time since disturbance (mean time since deforestation) and deforestation profile (deforestation profile curvature). We used the proposed indices to analyze a watershed located in central Rondonia. Landsat TM and ETM+ images were used to produce historical land-use maps of the last 18 years, each even year from 1984 to 2002 for 20 catchments. We found that the land-use dynamics indicators are able to distinguish catchments with different land-use change profiles. Four categories of historical land-use were identified: old and dominant pasture cover on small properties, recent deforestation and dominance of secondary growth, old extensive pastures and large forest remnants and, recent deforestation, pasture and large forest remnants. Knowing historical deforestation processes is important to develop appropriate conservation strategies and define priorities and actions for conserving forests currently under deforestation. (C) 2009 Elsevier B.V. All rights reserved.

Passive Restoration in Biodiversity Hotspots: Consequences for an Atlantic Rainforest Lizard Taxocene

Long-term conservation in biodiversity hotspots depends on the recovery of communities in secondary forest fragments. In most cases, however, recovery strategies for these areas are based only on passive restoration. It is therefore necessary to determine the efficiency of such strategies. In this study, we assess the efficiency of passive restoration on a 567-ha 28-yr-old fragment of Atlantic Rainforest in Northeastern Brazil. We measured richness, composition, abundance and biomass of a lizard taxocene and also vegetation structure and availability of several microhabitat descriptors in 18 plots of this secondary forest. We then compared them with measures in 29 plots from two neighboring reference sites. Species richness, abundance, biomass and microhabitat descriptors availability inside the secondary fragment did not differ from reference sites. However, composition and vegetation structure showed small differences. Some forest specialist lizards, which should be a focus of conservation efforts in fragmented landscapes of the Atlantic Rainforest, were not found in the fragment and data indicate that this was not due to sampling or a lack of suitable habitat or microhabitat. In the presence of preserved source sites, passive restoration may be a cheap and effective way to recover lizard taxocenes of the Atlantic Rainforest. Some of the species may need to be re-introduced to accelerate the full recovery of original composition of lizard taxocenes in secondary Atlantic Rainforests.

Allometric models for estimating the phytomass of a secondary Atlantic Forest area of southeastern Brazil

The purpose of this study was to develop and validate equations to estimate the aboveground phytomass of a 30 years old plot of Atlantic Forest. In two plots of 100 m², a total of 82 trees were cut down at ground level. For each tree, height and diameter were measured. Leaves and woody material were separated in order to determine their fresh weights in field conditions. Samples of each fraction were oven dried at 80 °C to constant weight to determine their dry weight. Tree data were divided into two random samples. One sample was used for the development of the regression equations, and the other for validation. The models were developed using single linear regression analysis, where the dependent variable was the dry mass, and the independent variables were height (h), diameter (d) and d²h. The validation was carried out using Pearson correlation coefficient, paired t-Student test and standard error of estimation. The best equations to estimate aboveground phytomass were: lnDW = -3.068+2.522lnd (r² = 0.91; s y/x = 0.67) and lnDW = -3.676+0.951ln d²h (r² = 0.94; s y/x = 0.56).

Habitat suitability of Anopheles vector species and association with human malaria in the Atlantic Forest in south-eastern Brazil

Every year, autochthonous cases of Plasmodium vivax malaria occur in low-endemicity areas of Vale do Ribeira in the south-eastern part of the Atlantic Forest, state of São Paulo, where Anopheles cruzii and Anopheles bellator are considered the primary vectors. However, other species in the subgenus Nyssorhynchus of Anopheles (e.g., Anopheles marajoara) are abundant and may participate in the dynamics of malarial transmission in that region. The objectives of the present study were to assess the spatial distribution of An. cruzii, An. bellator and An. marajoara and to associate the presence of these species with malaria cases in the municipalities of the Vale do Ribeira. Potential habitat suitability modelling was applied to determine both the spatial distribution of An. cruzii, An. bellator and An. marajoara and to establish the density of each species. Poisson regression was utilized to associate malaria cases with estimated vector densities. As a result, An. cruzii was correlated with the forested slopes of the Serra do Mar, An. bellator with the coastal plain and An. marajoara with the deforested areas. Moreover, both An. marajoara and An. cruzii were positively associated with malaria cases. Considering that An. marajoara was demonstrated to be a primary vector of human Plasmodium in the rural areas of the state of Amapá, more attention should be given to the species in the deforested areas of the Atlantic Forest, where it might be a secondary vector.

Temporal Diversity Patterns and Phenology in Fruit-feeding Butterflies in the Atlantic Forest

The Atlantic Forest deserves special attention due to its high level of species endemism and degree of threat. As in other tropical biomes, there is little information about the ecology of the organisms that occur there. The objectives of this study were to verify how fruit-feeding butterflies are distributed through time, and the relation with meteorological conditions. Species richness and Shannon index were partitioned additively at the monthly level, and beta diversity, used as a hierarchical measure of temporal species turnover, was calculated among months, trimesters, and semesters. Circular analysis was used to verify how butterflies are distributed along seasons and its relation with meteorological conditions. We sampled 6488 individuals of 73 species. Temporal diversity of butterflies was more grouped than expected by chance among the months of each trimester. Circular analyses revealed that diversity is concentrated in hot months (September-March), with the subfamily Brassolinae strongly concentrated in February-March. Average temperature was correlated with total abundance of butterflies, abundance of Biblidinae, Brassolinae and Morphinae, and richness of Satyrinae. The present results show that 3mo of sampling between September and March is enough to produce a nonbiased sample of the local assemblage of butterflies, containing at least 70 percent of the richness and 25 percent of abundance. The influence of temperature on sampling is probably related to butterfly physiology. Moreover, temperature affects resource availability for larvae and adults, which is higher in hot months. The difference in seasonality patterns among subfamilies is probably a consequence of different evolutionary pressures through time.

Forest structure in a mosaic of rainforest sites: The effect of fragmentation and recovery after clear cut

A variety of human-induced disturbances such as forest fragmentation and recovery after deforestation for pasture or agricultural activities have resulted in a complex landscape mosaic in the Una region of northeastern Brazil. Using a set of vegetation descriptors, we investigated the main structural changes observed in forest categories that comprise the major components of the regional landscape and searched for potential key descriptors that could be used to discriminate among different forest categories. We assessed the forest structure of five habitat categories defined as (I) interiors and (2) edges of large fragments of old-growth forest (>1000 ha), (3) interiors and (4) edges of small forest fragments (<100 ha), and (5) early secondary forests. Forest descriptors used here were: frequency of herbaceous lianas and woody climbers, number of standing dead trees, number of fallen trunks, litter depth, number of pioneer plants (early secondary and shade-intolerant species), vertical foliage stratification profile and distribution Of trees in different diameter classes. Edges and interiors of forest fragments were significantly different only in the number of standing dead trees. Secondary forests and edges of fragments showed differences in litter depth, fallen trunks and number of pioneer trees, and secondary forests were significantly different from fragment interiors in the number of standing dead trees and the number of pioneer trees. Horizontal and vertical structure evaluated via ordination analysis showed that fragment interiors, compared to secondary forests, were characterized by a greater number of medium (25-35 cm) and large (35-50 cm) trees and smaller numbers of thin trees (5-10 cm). There was great heterogeneity at the edges of small and large fragments, as these sites were distributed along almost the entire gradient. Most interiors of large and small fragments presented higher values of foliage densities at higher strata ( 15-20 m and at 20-25 m height), and lower densities at 1-5 m. All secondary forests and some fragment edge sites showed an opposite tendency. A discriminant function highlighted differences among forest categories, with transects of large fragment interiors and secondary forests representing two extremes along a disturbance gradient determined by foliage structure (densities at 15-20 m and 20-25 m), with the edges of both large and small fragments and the interiors of small fragments scattered across the gradient. The major underlying processes determining patterns of forest disturbance in the study region are discussed, highlighting the importance of forest fragments, independently of its size, as forests recovery after clear cut show a greatly distinct structure, with profound implications on fauna movements. (C) 2009 Elsevier BY. All rights reserved.

Successional stage and geographic features determine floristic similarity among Atlantic Forest remnants, Sao Paulo State, Brazil

Analysis of floristic similarity relationships between plant communities can detect patterns of species occurrence and also explain conditioning factors. Searching for such patterns, floristic similarity relationships among Atlantic Forest sites situated at Ibiuna Plateau, Sao Paulo state, Brazil, were analyzed by multivariate techniques. Twenty one forest fragments and six sites within a continuous Forest Reserve were included in the analyses. Floristic composition and structure of the tree community (minimum dbh 5 cm) were assessed using the point centered quarter method. Two methods were used for multivariate analysis: Detrended Correspondence Analysis (DCA) and Two-Way Indicator Species Analysis (TWINSPAN). Similarity relationships among the study areas were based on the successional stage of the community and also on spatial proximity. The more similar the successional stage of the communities, the higher the floristic similarity between them, especially if the communities are geographically close. A floristic gradient from north to south was observed, suggesting a transition between biomes, since northern indicator species are mostly heliophytes, occurring also in cerrado vegetation and seasonal semideciduous forest, while southern indicator species are mostly typical ombrophilous and climax species from typical dense evergreen Atlantic Forest.