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.

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.

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.

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.

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.

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.

Diet and food selection by small mammals in an old-growth Atlantic forest of south-eastern Brazil

Here we describe the stomach contents of nine small mammal species (seven rodents and two didelphid marsupials) co-occurring in an old-growth Atlantic forest area. For four terrestrial rodents, we also compared the importance of arthropods in the diet and the selection of arthropod groups by comparing consumption with availability. Small mammals and arthropods were sampled in a 36-ha grid containing 25 sampling stations spaced every 150 m, and 47 stomach contents were analysed. While plant matter was the predominant item in the stomach contents of two rodents (Oligoryzomys nigripes and Rhipidomys mastacalis), four species presented arthropods as the main food item (the rodents Brucepattersonius soricinus and Oxymycterus dasytrichus, and the marsupials Monodelphis n. sp. and Marmosops incanus) and three consumed more plant matter than arthropods, but had significant amounts of both items (the rodents Delomys sublineatus, Euryoryzomys russatus and Thaptomys nigrita). Our results suggest that differences in diet, coupled with differences in habit and microhabitat preferences, are important factors allowing resource partition among species of the diverse group of co-occurring terrestrial small mammals in Atlantic forest areas. Moreover, arthropods were not preyed opportunistically by any of the four terrestrial rodents, since consumption was not proportional to availability. Rather, selection or rejection of arthropod groups seems to be determined by aspects other than availability, such as nutritional value, easiness of capture and handling or palatability.

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.

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.