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.

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.

Behavioral suites mediate group-level foraging dynamics in communities of tropical stingless bees

Competition for floral resources is a key force shaping pollinator communities, particularly among social bees. The ability of social bees to recruit nestmates for group foraging is hypothesized to be a major factor in their ability to dominate rich resources such as mass-flowering trees. We tested the role of group foraging in attaining dominance by stingless bees, eusocial tropical pollinators that exhibit high diversity in foraging strategies. We provide the first experimental evidence that meliponine group foraging strategies, large colony sizes and aggressive behavior form a suite of traits that enable colonies to improve dominance of rich resources. Using a diverse assemblage of Brazilian stingless bee species and an array of artificial ""flowers"" that provided a sucrose reward, we compared species` dominance and visitation under unrestricted foraging conditions and with experimental removal of group-foraging species. Dominance does not vary with individual body size, but rather with foraging group size. Species that recruit larger numbers of nestmates (Scaptotrigona aff. depilis, Trigona hyalinata, Trigona spinipes) dominated both numerically (high local abundance) and behaviorally (controlling feeders). Removal of group-foraging species increased feeding opportunities for solitary foragers (Frieseomelitta varia, Melipona quadrifasciata and Nannotrigona testaceicornis). Trigona hyalinata always dominated under unrestricted conditions. When this species was removed, T. spinipes or S. aff. depilis controlled feeders and limited visitation by solitary-foraging species. Because bee foraging patterns determine plant pollination success, understanding the forces that shape these patterns is crucial to ensuring pollination of both crops and natural areas in the face of current pollinator declines.

Effects of roads, topography, and land use on forest cover dynamics in the Brazilian Atlantic Forest

Roads and topography can determine patterns of land use and distribution of forest cover, particularly in tropical regions. We evaluated how road density, land use, and topography affected forest fragmentation, deforestation and forest regrowth in a Brazilian Atlantic Forest region near the city of Sao Paulo. We mapped roads and land use/land cover for three years (1962, 1981 and 2000) from historical aerial photographs, and summarized the distribution of roads, land use/land cover and topography within a grid of 94 non-overlapping 100 ha squares. We used generalized least squares regression models for data analysis. Our models showed that forest fragmentation and deforestation depended on topography, land use and road density, whereas forest regrowth depended primarily on land use. However, the relationships between these variables and forest dynamics changed in the two studied periods; land use and slope were the strongest predictors from 1962 to 1981, and past (1962) road density and land use were the strongest predictors for the following period (1981-2000). Roads had the strongest relationship with deforestation and forest fragmentation when the expansions of agriculture and buildings were limited to already deforested areas, and when there was a rapid expansion of development, under influence of Sao Paulo city. Furthermore, the past(1962)road network was more important than the recent road network (1981) when explaining forest dynamics between 1981 and 2000, suggesting a long-term effect of roads. Roads are permanent scars on the landscape and facilitate deforestation and forest fragmentation due to increased accessibility and land valorization, which control land-use and land-cover dynamics. Topography directly affected deforestation, agriculture and road expansion, mainly between 1962 and 1981. Forest are thus in peril where there are more roads, and long-term conservation strategies should consider ways to mitigate roads as permanent landscape features and drivers facilitators of deforestation and forest fragmentation. (C) 2009 Elsevier B.V. All rights reserved.

Habitat fragmentation reduces genetic diversity and connectivity among toad populations in the Brazilian Atlantic Coastal Forest

Tropical rainforests are becoming increasingly fragmented and understanding the genetic consequences of fragmentation is crucial for conservation of their flora and fauna. We examined populations of the toad Rhinella ornata, a species endemic to Atlantic Coastal Forest in Brazil, and compared genetic diversity among small and medium forest fragments that were either isolated or connected to large forest areas by corridors. Genetic differentiation, as measured by F(ST), was not related to geographic distance among study sites and the size of the fragments did not significantly alter patterns of genetic connectivity. However, population genetic diversity was positively related to fragment size, thus haplotype diversity was lowest in the smallest fragments, likely due to decreases in population sizes. Spatial analyses of genetic discontinuities among groups of populations showed a higher proportion of barriers to gene flow among small and medium fragments than between populations in continuous forest. Our results underscore that even species with relatively high dispersal capacities may, over time, suffer the negative genetic effects of fragmentation, possibly leading to reduced fitness of population and cases of localized extinction. (C) 2008 Elsevier Ltd. All rights reserved.

Decomposition in tropical forests: a pan-tropical study of the effects of litter type, litter placement and mesofaunal exclusion across a precipitation gradient

1. Litter decomposition recycles nutrients and causes large fluxes of carbon dioxide into the atmosphere. It is typically assumed that climate, litter quality and decomposer communities determine litter decay rates, yet few comparative studies have examined their relative contributions in tropical forests. 2. We used a short-term litterbag experiment to quantify the effects of litter quality, placement and mesofaunal exclusion on decomposition in 23 tropical forests in 14 countries. Annual precipitation varied among sites (760-5797 mm). At each site, two standard substrates (Raphia farinifera and Laurus nobilis) were decomposed in fine- and coarse-mesh litterbags both above and below ground for approximately 1 year. 3. Decomposition was rapid, with >95% mass loss within a year at most sites. Litter quality, placement and mesofaunal exclusion all independently affected decomposition, but the magnitude depended upon site. Both the average decomposition rate at each site and the ratio of above- to below-ground decay increased linearly with annual precipitation, explaining 60-65% of among-site variation. Excluding mesofauna had the largest impact on decomposition, reducing decomposition rates by half on average, but the magnitude of decrease was largely independent of climate. This suggests that the decomposer community might play an important role in explaining patterns of decomposition among sites. Which litter type decomposed fastest varied by site, but was not related to climate. 4. Synthesis. A key goal of ecology is to identify general patterns across ecological communities, as well as relevant site-specific details to understand local dynamics. Our pan-tropical study shows that certain aspects of decomposition, including average decomposition rates and the ratio of above- to below-ground decomposition are highly correlated with a simple climatic index: mean annual precipitation. However, we found no relationship between precipitation and effects of mesofaunal exclusion or litter type, suggesting that site-specific details may also be required to understand how these factors affect decomposition at local scales.

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.

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

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.

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.

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.

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.

On the first terrestrial ostracod of the Superfamily Cytheroidea (Crustacea, Ostracoda): description of Intrepidocythere ibipora n. gen. n. sp from forest leaf litter in Sao Paulo State, Brazil

Of the three superfamilies of Ostracoda present in fresh water, only the Cytheroidea had thus far no records in terrestrial environments. Here, we report on a new genus and species, Intrepidocythere ibipora n. gen. n. sp., of the ostracod superfamily Cytheroidea, from forest leaf litter in Sao Paulo State, Brazil. Judging from morphological similarities, this new genus is believed to be closely related to the genus Elpidium. Possible pathways that led to the colonisation of terrestrial habitats are discussed, and an overview is given on the distribution of the known terrestrial ostracod lineages. The present findings strengthen the idea that terrestrial ostracods are more common than previously thought, at least in tropical areas.

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.

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.