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.

The impact of fragmentation and density regulation on forest succession in the Atlantic rain forest

The Atlantic Rain Forest, an important biodiversity hot spot, has faced severe habitat loss since the last century which has resulted in a highly fragmented landscape with a large number of small forest patches (<100 ha). For conservation planning it is essential to understand how current and future forest regeneration depends on ecological processes, fragment size and the connection to the regional seed pool. We have investigated the following questions by applying the forest growth simulation model FORMIND to the situation of the Atlantic Forest in the state of Sao Paulo, SE Brazil: (1) which set of parameters describing the local regeneration and level of density regulation can reproduce the biomass distribution and stem density of an old growth forest in a reserve? (2) Which additional processes apart from those describing the dynamics of an old growth forest, drive forest succession of small isolated fragments? (3) Which role does external seed input play during succession? Therefore, more than 300 tree species have been classified into nine plant functional types (PFTs), which are characterized by maximum potential height and shade tolerance. We differentiate between two seed dispersal modes: (i) local dispersal, i.e. all seedlings originated from fertile trees within the simulated area and (ii) external seed rain. Local seed dispersal has been parameterized following the pattern oriented approach, using biomass estimates of old growth forest. We have found that moderate density regulation is essential to achieve coexistence for a broad range of regeneration parameters. Considering the expected uncertainty and variability in the regeneration processes it is important that the forest dynamics are robust to variations in the regeneration parameters. Furthermore, edge effects such as increased mortality at the border and external seed rain have been necessary to reproduce the patterns for small isolated fragments. Overall, simulated biomass is much lower in the fragments compared to the continuous forest, whereas shade tolerant species are affected most strongly by fragmentation. Our simulations can supplement empirical studies by extrapolating local knowledge on edge effects of fragments to larger temporal and spatial scales. In particular our results show the importance of external seed rain and therefore highlight the importance of structural connectivity between regenerating fragments and mature forest stands. (C) 2009 Elsevier B.V. All rights reserved.

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.

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.

Composição da fauna de formigas (Hymenoptera: Formicidae) de serapilheira em florestas semidecídua e de Eucalyptus spp., na região sudeste do Brasil

Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)

Gap size measurement: the proposal of a new field method

In the tropical Atlantic Forest, 42 canopy gaps had their areas estimated using four different field methods of measurement: Runkle, Brokaw and Green [Runkle, J.R., 1981. Gap formation in some old-growth forests of the eastern United States. Ecology 62, 1041-1051; Brokaw, N.V.L., 1982. The definition of treefall gap and its effect on measures of forest dynamics. Biotropica 14, 158-160; Green, P.T., 1996. Canopy Gaps in rain forest on Christmas Island, Indian Ocean: size distribution and methods of measurement. J. Trop. Ecol. 12, 427-434] and a new method proposed in this work. It was found that within the same gap delimitation, average gap size varied from 56.0 up to 88.3 m(3) while total sum of gap area varied from 2351.3 to 3707.9 m(3) Differences among all methods and between pairs of method proved to be statistically significant. As a consequence, gap size-class distribution was also different between methods. When one method is held as a standard, deviation on average values of gap size ranged between 11.8 and 59.7% as deviations on single gap size can reach 172.8%. Implications on forest dynamics were expressed by the forest turnover rate that was 24% faster or 15% slower depending on the method adopted for gap measurement. Based on my results and on methods' evaluation, the use of a new method is proposed here for future research involving the measure of gap size in forest ecosystems. Finally, it is concluded that forest comparisons disregarding the influence of different methods of gap measurement should be reconsidered. (c) 2005 Elsevier B.V. All rights reserved.

Jaguar (Panthera onca) food habits in Atlantic rain forest of southeastern Brazil

Between January and December 1996, the food habits of a relict population of jaguars were studied in 220 km(2) Linhares Forest Preserve, which comprises much of the remaining old-growth Atlantic Forest of Espirito Santo, Brazil. Fecal analysis indicated opportunistic feeding on 24 prey species (N = 101 scats). Mammals represented 87 percent of the total items, followed by reptiles (9.8%) and birds (2.8%). Considering prey weight, 23.4 percent of the items weighed 1-3 kg, 40.5 percent were 3-10 kg, and 27.7 percent weighed more than 10 kg. Analysis of relative prey frequency and biomass indicated that the diet was concentrated in two prey types: long-nosed armadillo and white-lipped peccary. Literature data suggest that forest jaguars rely on the same mammal prey over their entire geographic range.

Frugivory in cotingas of the Atlantic Forest of southeast Brazil

In this paper we present a list of plant species whose fruits are eaten by eight cotinga species (Carpornis cucullatus, C. melanocephalus, Laniisoma elegans, Lipaugus lanioides, Oxyruncus cristatus, Phibalura flavirostris, Procnias nudicollis, and Pyroderus scutatus) at Parque Estadual Intervales (PEI), an Atlantic Forest reserve in southeast Brazil. Besides providing a list of fruits eaten by cotingas, our goal is to allow insights into fruit use by these birds whose ecology is poorly known. From 1990 to 2001 four sites located within PEI were regularly visited. These sites form an altitudinal gradient (70-800 m a.s.l.) and also a gradient of forest disturbance (second-growth to old-growth vegetation). Feeding records were made along several trails and dirt roads that crossed the study sites. Cotingas ate a variety of fruits (99 species in 34 families) that ranged from small (< 5 mm diameter) to large (up to 37 mm) and included some non-ornithochorous ones. With the exception of P. nudicollis, cotingas exploited not only fruits typical of the forest interior but also fruits of secondary vegetation, suggesting that concerning fruit exploitation early successional vegetation does not represent a barrier for the survival of these cotingas.

Can overharvesting of a non-timber-forest-product change the regeneration dynamics of a tropical rainforest? The case study of Euterpe edulis

The exploitation of non-timber forest products is often considered a low-impact activity in tropical forests. However, assessments of the impacts of such activity are mostly focused on the harvested species and not on the plant community, thus limiting our understanding for establishing forest management recommendations. We investigated the consequences of Euterpe edulis palm heart harvesting on the seed rain in the Brazilian Atlantic rainforest. We compared the density of E. edulis individuals, as well as the density of E. edulis seeds, and the density, richness and functional composition of seed rain of the whole plant community, before and after palm heart harvesting in a 10 ha permanent plot. This assessment was carried out in preserved (typical old-growth Atlantic rainforest) and in disturbed (more open habitat dominated by the native bamboo Guadua tagoara) forest patches. Palm harvesting reduced the E. edulis population from 202.16 to 25.67 ind/ha and its seed rain density from 0.362 to 0.3 seeds/m2 and from 2.395 to 0.15 seeds/m2 in preserved and disturbed forest patches, respectively. Seed density of light-dependent climbers, pioneer trees, bamboo and animal-dispersed seeds increased after palm harvesting, especially in the disturbed forest patches, where palm harvesting was more intense and may have changed the light regime of the understory. On the other hand, species richness of the plant community declined by half. We observed a remarkable decline in the number of animal-dispersed species, especially for those with large seeds, suggesting that the activity of seed dispersers, including many species attracted by E. edulis fruits, was reduced. Therefore, harvesting of E. edulis palm heart may change the regeneration dynamics of the Atlantic rainforest, both due to shifts in forest structure, mediated by the removal of individuals from the forest canopy, and in community functioning, mediated by the interference on the activity of seed dispersers. © 2013 Elsevier B.V. All rights reserved.

Composição da avifauna e frugivoria por ave em um mosaico sucessional na Mata Atlântica

Pós-graduação em Ciências Biológicas (Zoologia) - IBRC

Seasonal variation in the fate of seeds under contrasting logging regimes

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Helminth Fauna of Two Species of Physalaemus (Anura: Leiuperidae) from an Undisturbed Fragment of the Atlantic Rainforest, Southeastern Brazil

Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)

Relações entre tamanho populacional, uso do habitat, dieta e predação de ninhos por Nasua nasua (Carnivora, Procyonidae) em remanescentes florestais

Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES)

Seedling fate across different habitats: the effects of herbivory and soil fertility

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Habitat structure and food resources for wildlife across successional stages in a tropical forest

Tropical forests are experiencing an increase in the proportion of secondary forests as a result of the balance between the widespread harvesting of old-growth forests and the regeneration in abandoned areas. The impacts of such a process on biodiversity are poorly known and intensely debated. Recent reviews and multi-taxa studies indicate that species replacement in wildlife assemblages is a consistent pattern, sometimes stronger than changes in diversity, with a replacement from habitat generalists to old-growth specialists being commonly observed during tropical forest regeneration. However, the ecological drivers of such compositional changes are rarely investigated, despite its importance in assessing the conservation value of secondary forests, and to support and guide management techniques for restoration. By sampling 28 sites in a continuous Atlantic forest area in Southeastern Brazil, we assessed how important aspects of habitat structure and food resources for wildlife change across successional stages, and point out hypotheses on the implications of these changes for wildlife recovery. Old-growth areas presented a more complex structure at ground level (deeper leaf litter, and higher woody debris volume) and higher fruit availability from an understorey palm, whereas vegetation connectivity, ground-dwelling arthropod biomass, and total fruit availability were higher in earlier successional stages. From these results we hypothetize that generalist species adapted to fast population growth in resource-rich environments should proliferate and dominate earlier successional stages, while species with higher competitive ability in resource-limited environments, or those that depend on resources such as palm fruits, on higher complexity at the ground level, or on open space for flying, should dominate older-growth forests. Since the identification of the drivers of wildlife recovery is crucial for restoration strategies, it is important that future work test and further develop the proposed hypotheses. We also found structural and functional differences between old-growth forests and secondary forests with more than 80 years of regeneration, suggesting that restoration strategies may be crucial to recover structural and functional aspects expected to be important for wildlife in much altered ecosystems, such as the Brazilian Atlantic forest. (C) 2012 Elsevier B.V. All rights reserved.