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.

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.

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.

Are leaf-litter frogs and lizards affected by edge effects due to forest fragmentation in Brazilian Atlantic forest?

FAPESP Fundacao de Amparo a Pesquisa do Estado de Sao Paulo

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.

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.

Relative effect of litter quality, forest type and their interaction on leaf decomposition in south-east Brazilian forests

Decomposition was studied in a reciprocal litter transplant experiment to examine the effects of forest type, litter quality and their interaction on leaf decomposition in four tropical forests in south-east Brazil. Litterbags were used to measure decomposition of leaves of one tree species from each forest type: Calophyllum brasiliense from restinga forest; Guapira opposita from Atlantic forest; Esenbeckia leiocarpa from semi-deciduous forest; and Copaifera langsdorffii from cerradao. Decomposition rates in rain forests (Atlantic and restinga) were twice as fast as those in seasonal forests (semi-deciduous and cerradao), suggesting that intensity and distribution of precipitation are important predictors of decomposition rates at regional scales. Decomposition rates varied by species, in the following order: E. leiocarpa > C. langsdorffii > G. opposita > C. brasiliense. However, there was no correlation between decomposition rates and chemical litter quality parameters: C:N, C:P, lignin concentration and lignin:N. The interaction between forest type and litter quality was positive mainly because C. langsdorffii decomposed faster than expected in its native forest. This is a potential indication of a decomposer`s adaptation to specific substrates in a tropical forest. These findings suggest that besides climate, interactions between decomposers and plants might play an essential role in decomposition processes and it must be better understood.

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.

Population genetic structure and dispersal in white-lipped peccaries (Tayassu pecari) from the Brazilian Pantanal

In many mammals social organization promotes genetic structuring, which can be influenced by the dispersal pattern of the species. We analyzed the population genetic structure and dispersal of white-lipped peccaries (Tayassu pecan) from the Pantanal, Brazil. We genotyped 100 individuals at 7 microsatellite loci from 2 adjacent locations with no obvious geographic barrier between them. We found a significant but low F(ST) value, and the Bayesian analysis indicated a unique cluster. No significant differences were observed between mean assignment indices of resident males and females from both locations, and the probability of being born at the location sampled of > 30% of the individuals analyzed was lower than average. Mean relatedness between resident female, male, and opposite-sex pairs was not statistically different in both locations. These results suggest a low degree of genetic differentiation between the locations analyzed, and dispersal by both sexes (contrary to the predicted male-biased dispersal of most mammalian species).

Coalescent analysis of mtDNA indicates Pleistocene divergence among three species of howler monkey (Alouatta spp.) and population subdivision within the Atlantic Coastal Forest species, A. guariba

We have used coalescent analysis of mtDNA cytochrome b (cyt b) sequences to estimate times of divergence of three species of Alouatta-A. caraya, A. belzebul, and A. guariba-which are in close geographic proximity. A. caraya is inferred to have diverged from the A. guariba/A. belzebul clade approximately 3.83 million years ago (MYA), with the later pair diverging approximately 1.55 MYA. These dates are much more recent than previous dates based on molecular-clock methods. In addition, analyses of new sequences from the Atlantic Coastal Forest species A. guariba indicate the presence of two distinct haplogroups corresponding to northern and southern populations with both haplogroups occurring in sympatry within Sao Paulo state. The time of divergence of these two haplogroups is estimated to be 1.2 MYA and so follows quite closely after the divergence of A. guariba and A. belzebul. These more recent dates point to the importance of Pleistocene environmental events as important factors in the diversification of A. belzebul and A. guariba. We discuss the diversification of the three Alouatta species in the context of recent models of climatic change and with regard to recent molecular phylogeographic analyses of other animal groups distributed in Brazil.

Variety matters: adaptive genetic diversity and parasite load in two mouse opossums from the Brazilian Atlantic forest

The adaptive potential of a species to a changing environment and in disease defence is primarily based on genetic variation. Immune genes, such as genes of the major histocompatibility complex (MHC), may thereby be of particular importance. In marsupials, however, there is very little knowledge about natural levels and functional importance of MHC polymorphism, despite their key role in the mammalian evolution. In a previous study, we discovered remarkable differences in the MHC class II diversity between two species of mouse opossums (Gracilinanus microtarsus, Marmosops incanus) from the Brazilian Atlantic forest, which is one of the most endangered hotspots for biodiversity conservation. Since the main forces in generating MHC diversity are assumed to be pathogens, we investigated in this study gastrointestinal parasite burden and functional associations between the individual MHC constitution and parasite load. We tested two contrasting scenarios, which might explain differences in MHC diversity between species. We predicted that a species with low MHC diversity would either be under relaxed selection pressure by low parasite diversity (`Evolutionary equilibrium` scenario), or there was a recent loss in MHC diversity leading to a lack of resistance alleles and increased parasite burden (`Unbalanced situation` scenario). In both species it became apparent that the MHC class II is functionally important in defence against gastrointestinal helminths, which was shown here for the first time in marsupials. On the population level, parasite diversity did not markedly differ between the two host species. However, we did observe considerable differences in the individual parasite load (parasite prevalence and infection intensity): while M. incanus revealed low MHC DAB diversity and high parasite load, G. microtarsus showed a tenfold higher population wide MHC DAB diversity and lower parasite burden. These results support the second scenario of an unbalanced situation.

SEASONAL MICROHABITAT SELECTION BY TERRESTRIAL RODENTS IN AN OLD-GROWTH ATLANTIC FOREST

We tested the hypothesis that microhabitat variables, abundance of terrestrial rodents, and microhabitat selection patterns of terrestrial rodents vary between the cool-dry and warm-wet season in the Atlantic forest of Brazil. We selected variables associated with ecological factors potentially important to terrestrial rodents (physical structure of litter and woody debris, and arthropod availability) and established 25 small, independent sampling units covering 36 ha of a homogenous, mature Atlantic forest patch. Litter humidity and height, amount of small woody debris, arthropod availability, and terrestrial rodent abundance increased, whereas the quantity of large woody debris decreased in the warm-wet season. Greater spatial segregation among terrestrial rodents also was observed in this season, especially between morphologically similar species. The distribution of 3 of the 4 most common terrestrial rodents was influenced by microhabitat variables in at least I of the seasons, and these species also differed in their pattern of microhabitat selection between seasons. In general, the amount of small woody debris and litter humidity were more important for the microscale distribution of terrestrial rodents in the cool-dry season, whereas in the mild warm-wet season species distributions were associated with food availability or were not clearly influenced by the measured variables. The patterns of microhabitat selection by 3 common terrestrial rodents, which were associated with features that characterize old-growth forest, may be responsible for their vulnerability to forest fragmentation.

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.

Sex ratio and morphological characteristics of Rufous Gnateaters, Conopophaga lineata (Aves, Passeriformes) in Atlantic forest fragments

Unequal sex ratios lead to the loss of genetic variability, decreasing the viability of populations in the long term. Anthropogenic activities often disturb the natural habitats and can cause alterations in sex ratio and morphological characteristics of several species. Forest fragmentation is a major conservation concern, so that understanding its effects in natural populations is essential. In this study, we evaluated the sex ratio and the morphological characteristics of Rufous Gnateaters (Conopophaga lineata (Wied, 1831)) in small and large forest fragments in Minas Gerais, Brazil. Birds (n = 89) were sexed by plumage characteristics and molecular markers. The molecular analysis showed that plumage is not a totally reliable method for sexing Rufous Gnateaters. We observed that sex ratio did not differ between large and small forest fragments, but birds in small fragments had larger wings and tarsus. Wing and tarsus changes may affect the movement ability of individuals within and among forest fragments. In conclusion, Rufous Gnateaters have been able to survive in both small and large Atlantic rain forest fragments without altering their sex ratio, but morphological changes can be prejudicial to their long term survival.

The genetic effects of Late Quaternary climatic changes over a tropical latitudinal gradient: diversification of an Atlantic Forest passerine

The increase in biodiversity from high to low latitudes is a widely recognized biogeographical pattern. According to the latitudinal gradient hypothesis (LGH), this pattern was shaped by differential effects of Late Quaternary climatic changes across a latitudinal gradient. Here, we evaluate the effects of climatic changes across a tropical latitudinal gradient and its implications to diversification of an Atlantic Forest (AF) endemic passerine. We studied the intraspecific diversification and historical demography of Sclerurus scansor, based on mitochondrial (ND2, ND3 and cytb) and nuclear (FIB7) gene sequences. Phylogenetic analyses recovered three well-supported clades associated with distinct latitudinal zones. Coalescent-based methods were applied to estimate divergence times and changes in effective population sizes. Estimates of divergence times indicate that intraspecific diversification took place during Middle-Late Pleistocene. Distinct demographic scenarios were identified, with the southern lineage exhibiting a clear signature of demographic expansion, while the central one remained more stable. The northern lineage, contrasting with LGH predictions, exhibited a clear sign of a recent bottleneck. Our results suggest that different AF regions reacted distinctly, even in opposite ways, under the same climatic period, producing simultaneously favourable scenarios for isolation and contact among populations.