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.

Relative effects of fragment size and connectivity on bird community in the Atlantic Rain Forest: Implications for conservation

It is known that large fragment sizes and high connectivity levels are key components for maintaining species in fragments; however, their relative effects are poorly understood, especially in tropical areas. In order to test these effects, we built models for explaining understory birds occurrence in a fragmented Atlantic Rain Forest landscape with intermediate habitat cover (3%). Data from over 9000 mist-net hours from 17 fragments differing in size (2-175 ha) and connectivity (considering corridor linkages and distance to nearby fragments) were ranked under a model selection approach. A total 1293 individuals of 62 species were recorded. Species richness, abundance and compositional variation were mainly affected by connectivity indices that consider the capacity of species to use corridors and/or to cross short distances up to 30 m through the matrix. Bird functional groups were differently affected by area and connectivity: while terrestrial insectivores, omnivores and frugivores were affected by both area and connectivity, the other groups (understory insectivores, nectarivores, and others) were affected only by connectivity. In the studied landscape, well connected fragments can sustain an elevated number of species and individuals. Connectivity gives the opportunity for individuals to use multiple fragments, reducing the influence of fragment size. While preserving large fragments is a conservation target worldwide and should continue to be, our results indicated that connectivity between fragments can enhance the area functionally connected and is beneficial to all functional groups and therefore should be a conservation priority. (C) 2008 Elsevier Ltd. All rights reserved.

Can agroforest woodlots work as stepping stones for birds in the Atlantic forest region?

In fragmented landscapes, agroforest woodlots can potentially act as stepping stones facilitating movement between forest fragments. We assessed the influence of agroforest woodlots on bird distribution and diversity in the Atlantic forest region (SE Brazil), and also tested which categories of species can use different types of connection elements, and whether this use is influenced by the distance to large forest patches. We studied two fragmented landscapes, with and without stepping stones linking large fragments, and one forested landscape. Using a point count, a bird survey was undertaken in the fragmented landscapes in five different elements: large remnants (> 400 ha), agroforest woodlots (0.4-1.1 ha), small patches (0.5-7 ha), riparian corridor, and pasture areas (the main matrix). Generalist and open-area species were commonly observed in the agroforest system or other connection elements, whereas only a few forest species were present in these connections. For the latter species, the distance of woodlots to large patches was essential to determine their richness and abundance. Based on our results and data from literature, we suggest that there is an optimal relationship between the permeability of the matrix and the efficiency of stepping stones, which occurs at intermediate degrees of matrix resistance, and is species-dependent. Because the presence of agroforest system favors a higher richness of generalist species, they appeared to be more advantageous for conservation than the monoculture system; for this reason, they should be considered as a management alternative, particularly when the matrix permeability requirement is met.

Importance of estimating matrix quality for modeling species distribution in complex tropical landscapes: a test with Atlantic forest small mammals

Information to guide decision making is especially urgent in human dominated landscapes in the tropics, where urban and agricultural frontiers are still expanding in an unplanned manner. Nevertheless, most studies that have investigated the influence of landscape structure on species distribution have not considered the heterogeneity of altered habitats of the matrix, which is usually high in human dominated landscapes. Using the distribution of small mammals in forest remnants and in the four main altered habitats in an Atlantic forest landscape, we investigated 1) how explanatory power of models describing species distribution in forest remnants varies between landscape structure variables that do or do not incorporate matrix quality and 2) the importance of spatial scale for analyzing the influence of landscape structure. We used standardized sampling in remnants and altered habitats to generate two indices of habitat quality, corresponding to the abundance and to the occurrence of small mammals. For each remnant, we calculated habitat quantity and connectivity in different spatial scales, considering or not the quality of surrounding habitats. The incorporation of matrix quality increased model explanatory power across all spatial scales for half the species that occurred in the matrix, but only when taking into account the distance between habitat patches (connectivity). These connectivity models were also less affected by spatial scale than habitat quantity models. The few consistent responses to the variation in spatial scales indicate that despite their small size, small mammals perceive landscape features at large spatial scales. Matrix quality index corresponding to species occurrence presented a better or similar performance compared to that of species abundance. Results indicate the importance of the matrix for the dynamics of fragmented landscapes and suggest that relatively simple indices can improve our understanding of species distribution, and could be applied in modeling, monitoring and managing complex tropical landscapes.

Comparative range use by three Atlantic Forest understorey bird species in relation to forest fragmentation

In this paper, we report on range use patterns of birds in relation to tropical forest fragmentation. Between 2003 and 2005, three understorey passerine species were radio-tracked in five locations of a fragmented and in two locations of a contiguous forest landscape on the Atlantic Plateau of Sao Paulo in south-eastern Brazil. Standardized ten-day home ranges of 55 individuals were used to determine influences of landscape pattern, season, species, sex and age. In addition, total observed home ranges of 76 individuals were reported as minimum measures of spatial requirements of the species. Further, seasonal home ranges of recaptured individuals were compared to examine site fidelity. Chiroxiphia caudata, but not Pyriglena leucoptera or Sclerurus scansor, used home ranges more than twice as large in the fragmented versus contiguous forest. Home range sizes of C. caudata differed in relation to sex, age, breeding status and season. Seasonal home ranges greatly overlapped in both C. caudata and in S. scansor. Our results suggest that one response by some forest bird species to habitat fragmentation entails enlarging their home ranges to include several habitat fragments, whereas more habitat-sensitive species remain restricted to larger forest patches.

Fragmentation drives tropical forest fragments to early successional states: A modelling study for Brazilian Atlantic forests

Land use leads to massive habitat destruction and fragmentation in tropical forests. Despite its global dimensions the effects of fragmentation on ecosystem dynamics are not well understood due to the complexity of the problem. We present a simulation analysis performed by the individual-based model FORMIND. The model was applied to the Brazilian Atlantic Forest, one of the world`s biodiversity hot spots, at the Plateau of Sao Paulo. This study investigates the long-term effects of fragmentation processes on structure and dynamics of different sized remnant tropical forest fragments (1-100 ha) at community and plant functional type (PFT) level. We disentangle the interplay of single effects of different key fragmentation processes (edge mortality, increased mortality of large trees, local seed loss and external seed rain) using simulation experiments in a full factorial design. Our analysis reveals that particularly small forest fragments below 25 ha suffer substantial structural changes, biomass and biodiversity loss in the long term. At community level biomass is reduced up to 60%. Two thirds of the mid- and late-successional species groups, especially shade-tolerant (late successional climax) species groups are prone of extinction in small fragments. The shade-tolerant species groups were most strongly affected; its tree number was reduced more than 60% mainly by increased edge mortality. This process proved to be the most powerful of those investigated, explaining alone more than 80% of the changes observed for this group. External seed rain was able to compensate approximately 30% of the observed fragmentation effects for shade-tolerant species. Our results suggest that tropical forest fragments will suffer strong structural changes in the long term, leading to tree species impoverishment. They may reach a new equilibrium with a substantially reduced subset of the initial species pool, and are driven towards an earlier successional state. The natural regeneration potential of a landscape scattered with forest fragments appears to be limited, as external seed rain is not able to fully compensate for the observed fragmentation-induced changes. Our findings suggest basic recommendations for the management of fragmented tropical forest landscapes. (C) 2011 Elsevier B.V. All rights reserved.

Movements of neotropical understory passerines affected by anthropogenic forest edges in the Brazilian Atlantic rainforest

Edge effects are suggested to have great impact on the persistence of species in fragmented landscapes. We tested edge avoidance by forest understory passerines in the Brazilian Atlantic Rainforest and also compared their mobility and movement patterns in contiguous and fragmented landscapes to assess whether movements would increase in the fragmented landscape. Between 2003 and 2005, 96 Chiroxiphia caudata, 38 Pyriglena leucoptera and 27 Sclerurus scansor were radio-tracked. The most strictly forest species C. caudata and S scansor avoided forest edges while P leucoptera showed affinities for the edge Both sensitive species showed larger mean step length and maximal observed daily distance in the fragmented forest versus the unfragmented forest. P. leucoptera did not show any significant difference. There were no significant differences in proportional daily home range use for any of the three species. Our results suggested that fragmentation and the consequent increase in edge areas do influence movement behavior of sensitive forest understory birds that avoided the use of edges and increased the speed and distance they covered daily. For the most restricted forest species, it would be advisable to protect larger patches of forest instead of many small or medium fragments connected by narrow corridors. However, by comparing our data with that obtained earlier, we concluded that movement behavior of resident birds differs from that of dispersing birds and might not allow to infer functional connectivity or landscape-scale sensitivity to fragmentation; a fact that should be taken into consideration when suggesting conservation strategies. (c) 2008 Elsevier Ltd. All rights reserved.

The challenge of maintaining Atlantic forest biodiversity: A multi-taxa conservation assessment of specialist and generalist species in an agro-forestry mosaic in southern Bahia

Recent developments have highlighted the importance of forest amount at large spatial scales and of matrix quality for ecological processes in remnants. These developments, in turn, suggest the potential for reducing biodiversity loss through the maintenance of a high percentage of forest combined with sensitive management of anthropogenic areas. We conducted a multi-taxa survey to evaluate the potential for biodiversity maintenance in an Atlantic forest landscape that presented a favorable context from a theoretical perspective (high proportion of mature forest partly surrounded by structurally complex matrices). We sampled ferns, butterflies, frogs, lizards, bats, small mammals and birds in interiors and edges of large and small mature forest remnants and two matrices (second-growth forests and shade cacao plantations), as well as trees in interiors of small and large remnants. By considering richness, abundance and composition of forest specialists and generalists, we investigated the biodiversity value of matrix habitats (comparing them with interiors of large remnants for all groups except tree), and evaluated area (for all groups) and edge effects (for all groups except trees) in mature forest remnants. our results suggest that in landscapes comprising high amounts of mature forest and low contrasting matrices: (1) shade cacao plantations and second-growth forests harbor an appreciable number of forest specialists; (2) most forest specialist assemblages are not affected by area or edge effects, while most generalist assemblages proliferate at edges of small remnants. Nevertheless, differences in tree assemblages, especially among smaller trees, Suggest that observed patterns are unlikely to be stable over time. (C) 2009 Elsevier Ltd. All rights reserved.

STRUCTURE OF THE SHRUB-ARBOREAL COMPONENT OF AN ATLANTIC FOREST FRAGMENT ON A HILLOCK IN THE CENTRAL LOWLAND OF RIO DE JANEIRO, BRAZIL

The present study describes and evaluates the horizontal and vertical structures of a lowland forest fragment on a hillock in the municipality of Silva Jardim, Rio de Janeiro State, Brazil (22 degrees 31`56 `` S and 42 degrees 20`46 `` W). Twenty plots (10x2m) totaling 0.5ha were laid out following the slope grade using DBH >= 5cm as the inclusion criterion. A total of 734 individuals were encountered, yielding a total density of 1468 ind./ha and a total basal area of 10783m(2). The richness values (129 species/41 families), Shannon-Wiener diversity (4.22) and equitability (0.87) indices indicated an accentuated floristic heterogeneity and low ecological dominance. Lauraceae, Myrtaceae, Fabaceae and Euphorbiaceae showed the greatest species richness, corroborating other studies that indicated these species as the most representative of Atlantic Forest areas in southeastern Brazil. The species with the greatest importance values (VI) were Aparisthmium cordatum, Guapira opposita, Lacistema pubescens, Xylopia sericea, Tapirira guianensis and Piptocarpha macropoda. The high diversity observed was influenced by earlier anthropogenic actions and by the current successional stage. The forest fragment studied demonstrated closer floristic similarity to areas inventoried in a close-by biological reserve than to fragments dispersed throughout the coastal plain. Similarities in soil type, degree of soil saturation and use-history of forest resources all support these relationships. The fragmented physiognomy of the central lowland in this region and the use-history of the landscape make these small remnant forest areas important in terms of establishing strategies for landscape restoration and species conservation.

Landscape and farm scale management to enhance biodiversity conservation in the cocoa producing region of southern Bahia, Brazil

In southern Bahia, Brazil, large land areas are used for the production of cocoa (Theobroma cacao), which is predominantly grown under the shade of native trees in an agroforestry system locally known as cabruca. As a dominant forest-like landscape element of the cocoa region, the cabrucas play an important role in the conservation of the region`s biodiversity. The purpose of this review is to provide the scientific basis for an action plan to reconcile cocoa production and biodiversity conservation in southern Bahia. The available research collectively highlights the diversity of responses of different species and biological groups to both the habitat quality of the cabrucas themselves and to the general characteristics of the landscape, such as the relative extent and spatial configuration of different vegetation types within the landscape mosaic. We identify factors that influence directly or indirectly the occurrence of native species in the cabrucas and the wider landscape of the cocoa region and develop recommendations for their conservation management. We show that the current scientific knowledge already provides a good basis for a biodiversity friendly management of the cocoa region of southern Bahia, although more work is needed to refine some management recommendations, especially on shade canopy composition and density, and verify their economic viability. The implementation of our recommendations should be accompanied by appropriate biological and socioeconomic monitoring and the findings should inform a broad program of adaptive management of the cabrucas and the wider cocoa landscape.

Carnivore mammals in a fragmented landscape in northeast of Sao Paulo State, Brazil

Sao Paulo is the most developed state in Brazil and little of its native vegetation remains. In Luiz Antonio and Santa Rita do Passa Quatro municipalities, only small fragments of cerrado (Brazilian savanna) physiognomies (cerrado, cerrado sensu stricto) and of semideciduous forest have been left, surrounded by eucalyptus silviculture and sugar-cane agriculture. However, that vegetation mosaic still shelters large mammals, including several carnivore species. To detect the carnivores present in such a mosaic area (50,000 ha), and to find out how they use the landscape, we recorded them through 21 camera traps and 21 track plots, during 18 months. Species richness, diversity and relative frequency were evaluated according to the habitat. Ten species were recorded, some of them locally threatened to extinction (Puma concolor, Leopardus pardalis, Chrysocyon brachyurus). Species diversity did not significantly differ among fragments, and although most species preferred one or another habitat, the carnivore community as a whole explored all the study area regardless of the vegetation cover;eucalyptus plantations were as used by the carnivores as the native fragments. Therefore, it seems possible to maintain such animals in agricultural landscapes, where some large native fragments are left and the matrix is permeable to native fauna.

Dynamics of Dissolved Forms of Carbon and Inorganic Nitrogen in Small Watersheds of the Coastal Atlantic Forest in Southeast Brazil

Based on the fact that streamwater quality reflects landscape conditions, the objectives of this study were: to investigate nitrogen (N), carbon (C), and major ion concentrations in six streams crossing minimally disturbed Atlantic Forest areas, with similar geomorphological characteristics; to determine N and C fluxes in one of these pristine streams (Indaia); and assess the impact of human activity on the biogeochemistry of two other streams in the same region, crossing urbanized areas. The distribution pattern of carbon and inorganic nitrogen dissolved forms, as well as the major ion and biogenic gas concentrations in the streamwater, was similar in pristine streams, indicating that the C and N dynamics were determined by influence of some factors, such as climate, atmospheric deposition, geology, soil type, and land covering, which were analogous in the forested watersheds. The urban streams were significantly different from the pristine streams, showing low dissolved oxygen concentrations, high respiration rates, and high concentrations of carbon dioxide, dissolved inorganic nitrogen, dissolved inorganic carbon, and major ion. These differences were attributed to anthropogenic impact on water quality, especially domestic sewage discharge. Additionally, in the Indaia stream, it was possible to observe the importance of rainfall over temporal dynamics of dissolved carbon forms, and also, the obtained specific flux of dissolved inorganic nitrogen was relatively elevated (approximately 11 kg ha(-1) year(-1)). These results reveal the influence of human activity over the biogeochemistry of coastal streams and also indicate the importance N export of Atlantic Forest to the ocean.

Influence of multi-scale landscape structure on the occurrence of carnivorous mammals in a human-modified savanna, Brazil

So Paulo is the most developed state in Brazil and contains few fragments of native ecosystems, generally surrounded by intensive agriculture lands. Despite this, some areas still shelter large native animals. We aimed at understanding how medium and large carnivores use a mosaic landscape of forest/savanna and agroecosystems, and how the species respond to different landscape parameters (percentage of landcover and edge density), in a multi-scale perspective. The response variables were: species richness, carnivore frequency and frequency for the three most recorded species (Puma concolor, Chrysocyon brachyurus and Leopardus pardalis). We compared 11 competing models using Akaike`s information criterion (AIC) and assessed model support using weight of AIC. Concurrent models were combinations of landcover types (native vegetation, ""cerrado"" formations, ""cerrado"" and eucalypt plantation), landscape feature (percentage of landcover and edge density) and spatial scale. Herein, spatial scale refers to the radius around a sampling point defining a circular landscape. The scales analyzed were 250 (fine), 1,000 (medium) and 2,000 m (coarse). The shape of curves for response variables (linear, exponential and power) was also assessed. Our results indicate that species with high mobility, P. concolor and C. brachyurus, were best explained by edge density of the native vegetation at a coarse scale (2,000 m). The relationship between P. concolor and C. brachyurus frequency had a negative power-shaped response to explanatory variables. This general trend was also observed for species richness and carnivore frequency. Species richness and P. concolor frequency were also well explained by a second concurrent model: edge density of cerrado at the fine (250 m) scale. A different response was recorded for L. pardalis, as the frequency was best explained for the amount of cerrado at the fine (250 m) scale. The curve of response was linearly positive. The contrasting results (P. concolor and C. brachyurus vs L. pardalis) may be due to the much higher mobility of the two first species, in comparison with the third. Still, L. pardalis requires habitat with higher quality when compared with other two species. This study highlights the importance of considering multiple spatial scales when evaluating species responses to different habitats. An important and new finding was the prevalence of edge density over the habitat extension to explain overall carnivore distribution, a key information for planning and management of protected areas.

Untangling associations between chironomid taxa in Neotropical streams using local and landscape filters

1. Analyses of species association have major implications for selecting indicators for freshwater biomonitoring and conservation, because they allow for the elimination of redundant information and focus on taxa that can be easily handled and identified. These analyses are particularly relevant in the debate about using speciose groups (such as the Chironomidae) as indicators in the tropics, because they require difficult and time-consuming analysis, and their responses to environmental gradients, including anthropogenic stressors, are poorly known. 2. Our objective was to show whether chironomid assemblages in Neotropical streams include clear associations of taxa and, if so, how well these associations could be explained by a set of models containing information from different spatial scales. For this, we formulated a priori models that allowed for the influence of local, landscape and spatial factors on chironomid taxon associations (CTA). These models represented biological hypotheses capable of explaining associations between chironomid taxa. For instance, CTA could be best explained by local variables (e.g. pH, conductivity and water temperature) or by processes acting at wider landscape scales (e.g. percentage of forest cover). 3. Biological data were taken from 61 streams in Southeastern Brazil, 47 of which were in well-preserved regions, and 14 of which drained areas severely affected by anthropogenic activities. We adopted a model selection procedure using Akaike`s information criterion to determine the most parsimonious models for explaining CTA. 4. Applying Kendall`s coefficient of concordance, seven genera (Tanytarsus/Caladomyia, Ablabesmyia, Parametriocnemus, Pentaneura, Nanocladius, Polypedilum and Rheotanytarsus) were identified as associated taxa. The best-supported model explained 42.6% of the total variance in the abundance of associated taxa. This model combined local and landscape environmental filters and spatial variables (which were derived from eigenfunction analysis). However, the model with local filters and spatial variables also had a good chance of being selected as the best model. 5. Standardised partial regression coefficients of local and landscape filters, including spatial variables, derived from model averaging allowed an estimation of which variables were best correlated with the abundance of associated taxa. In general, the abundance of the associated genera tended to be lower in streams characterised by a high percentage of forest cover (landscape scale), lower proportion of muddy substrata and high values of pH and conductivity (local scale). 6. Overall, our main result adds to the increasing number of studies that have indicated the importance of local and landscape variables, as well as the spatial relationships among sampling sites, for explaining aquatic insect community patterns in streams. Furthermore, our findings open new possibilities for the elimination of redundant data in the assessment of anthropogenic impacts on tropical streams.

Conservation in tropical landscape mosaics: the case of the cacao landscape of southern Bahia, Brazil

A recent debate has contrasted two conservation strategies in agricultural landscapes; either ""land sparing`` farm development combining intensive production practices with forest set-asides, or ""wildlife-friendly`` farming with greater on-farm habitat value but lower yields. We argue that in established mosaic landscapes including old cacao production regions where natural forest has already been reduced to relatively small fragments, a combination of both strategies is needed to conserve biodiversity. After reviewing the evidence for the insufficiency of either strategy alone if applied to such landscapes, the paper focuses on the cacao production landscape of southern Bahia, Brazil, once the world`s second largest cacao producer. Here, small remaining areas of Atlantic Forest are embedded in a matrix dominated by traditional cacao agroforests, resulting in a landscape mosaic that has proven favorable to the conservation of the region`s high biodiversity. We show that current land use dynamics and public policies pose threats but also offer opportunities to conservation and describe a three- pronged landscape conservation strategy, consisting of (i) expansion of the protected areas system, (ii) promotion of productive yet biodiversity-friendly cacao farming practices, and (iii) assistance to land users to implement legally mandated on-farm reserves and voluntary private reserves. We discuss recent experiences concerning the implementation of this strategy, discuss likely future scenarios, and reflect on the applicability of the Bahian experience to biodiversity rich cacao production regions elsewhere in the tropics.