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.

Assessing the impact of deforestation and climate change on the range size and environmental niche of bird species in the Atlantic forests, Brazil

Aim Habitat loss and climate change are two major drivers of biological diversity. Here we quantify how deforestation has already changed, and how future climate scenarios may change, environmental conditions within the highly disturbed Atlantic forests of Brazil. We also examine how environmental conditions have been altered within the range of selected bird species. Location Atlantic forests of south-eastern Brazil. Methods The historical distribution of 21 bird species was estimated using Maxent. After superimposing the present-day forest cover, we examined the environmental niches hypothesized to be occupied by these birds pre- and post-deforestation using environmental niche factor analysis (ENFA). ENFA was also used to compare conditions in the entire Atlantic forest ecosystem pre- and post-deforestation. The relative influence of land use and climate change on environmental conditions was examined using analysis of similarity and principal components analysis. Results Deforestation in the region has resulted in a decrease in suitable habitat of between 78% and 93% for the Atlantic forest birds included here. Further, Atlantic forest birds today experience generally wetter and less seasonal forest environments than they did historically. Models of future environmental conditions within forest remnants suggest generally warmer conditions and lower annual variation in rainfall due to greater precipitation in the driest quarter of the year. We found that deforestation resulted in a greater divergence of environmental conditions within Atlantic forests than that predicted by climate change. Main conclusions The changes in environmental conditions that have occurred with large-scale deforestation suggest that selective regimes may have shifted and, as a consequence, spatial patterns of intra-specific variation in morphology, behaviour and genes have probably been altered. Although the observed shifts in available environmental conditions resulting from deforestation are greater than those predicted by climate change, the latter will result in novel environments that exceed temperatures in any present-day climates and may lead to biotic attrition unless organisms can adapt to these warmer conditions. Conserving intra-specific diversity over the long term will require considering both how changes in the recent past have influenced contemporary populations and the impact of future environmental change.

Rare canopy species in communities within the Atlantic Coastal Forest in Rio de Janeiro State, Brazil

Rare species are one of the principal components of the species richness and diversity encountered in Dense Ombrophilous Tropical Forests. This study sought to analyze the rare canopy species within the Atlantic Coastal Forest in Rio de Janeiro State, Brazil. Six different communities were examined: Dense Ombrophilous alluvial Forest; Dense sub-montane Ombrophilous Forest; Dense Montane Ombrophilous in Serra do Mar and Serra da Mantiqueira. In each area the vegetation was sampled within forty 10 x 25 m plots alternately distributed along a linear transect. All trees with DBH (1.3 m above ground level) a parts per thousand yen5 cm were sampled. The canopy was characterized using the allometric relationship between diameter and height, and included all trees with BDH a parts per thousand yen10 cm and height a parts per thousand yen10 m. A total of 64 families, 206 genera, and 542 species were sampled, of which 297 (54.8%) represented rare species (less than one individual per hectare). The percentage of rare species varied from 34 to 50% in each of the different communities sampled. A majority of these rare trees belonged to the Rosidae, and a smaller proportion to the Dilleniidae. It was concluded that there was no apparent pattern to rarity among families, that rarity was probably derived from a number of processes (such as gap formation), and that a great majority of the rare species sampled were consistently rare. This indicates that the restricted geographic distribution and high degree of endemism of many arboreal taxa justifies the conservation of even small fragments of Atlantic Forest.

Seed development, yield and quality of two palm species growing in different tropical forest types in SE Brazil: implications for ecological restoration

Natural forest remnants have been set as seed production fields to supply seeds of native tree species for tropical forest restoration, but the effect of different forest types on seed production has not been accessed to date for palm species. In this work, we studied seed development, yield, and quality of two palm species in different tropical forest types in SE Brazil. Seed production of palmiteiro (Euterpe edulis) and queen-palm (Syagrus romanzoffiana), which are largely used in restoration efforts due to their importance for vertebrate frugivores, were studied in natural remnants of Atlantic Rainforest, Restinga Forest, Seasonally Dry Forest, and Cerrado Forest. We studied seed development, yield, size, and germination of seed lots produced in some of these forest types, including seeds harvested in 2008, 2009, and both years. Seed yield and quality, as well as seed dry mass in 2009, were higher for palmiteiro seeds produced in the Atlantic Rainforest, while queen-palm seeds produced at the Restinga Forest showed the higher mass and yield, but the lowest physiological potential. Consequently, these natural differences of seed yield and quality have to be taken into account for establishing standards for seed commercialization and analysis, seed pricing, and seedling production in forest nurseries.

Concordant Phylogeographies of 2 Malaria Vectors Attest to Common Spatial and Demographic Histories

The phylogeography of South American lineages is a topic of heated debate. Although a single process is unlikely to describe entire ecosystems, related species, which incur similar habitat limitations, can inform the history for a subsection of assemblages. We compared the phylogeographic patterns of the cytochrome oxidase I marker from Anopheles triannulatus (N = 72) and previous results for A. darlingi (N = 126) in a broad portion of their South American distributions. Both species share similar population subdivisions, with aggregations northeast of the Amazon River, in southern coastal Brazil and 2 regions in central Brazil. The average (ST) between these groups was 0.39 for A. triannulatus. Populations northeast of the Amazon and in southeastern Brazil are generally reciprocally monophyletic to the remaining groups. Based on these initial analyses, we constructed the a priori hypothesis that the Amazon and regions of high declivity pose geographic barriers to dispersal in these taxa. Mantel tests confirmed that these areas block gene flow for more than 1000 km for both species. The efficacy of these impediments was tested using landscape genetics, which could not reject our a priori hypothesis but did reject simpler scenarios. Results form summary statistics and phylogenetics suggest that both lineages originated in central Amazonia (south of the Amazon River) during the late Pleistocene (579 000 years ago) and that they followed the same paths of expansion into their contemporary distributions. These results may have implications for other species sharing similar ecological limitations but probably are not applicable as a general paradigm of Neotropical biogeography.

Forest-obligate Sabethes mosquitoes suggest palaeoecological perturbations

The origin of tropical forest diversity has been hotly debated for decades. Although specific mechanisms vary, many such explanations propose some vicariance in the distribution of species during glacial cycles and several have been supported by genetic evidence in Neotropical taxa. However, no consensus exists with regard to the extent or time frame of the vicariance events. Here, we analyse the cytochrome oxidase II mitochondrial gene of 250 Sabethes albiprivus B mosquitoes sampled from western Sao Paulo in Brazil. There was very low population structuring among collection sites (Phi(ST) = 0.03, P = 0.04). Historic demographic analyses and the contemporary geographic distribution of genetic diversity suggest that the populations sampled are not at demographic equilibrium. Three distinct mitochondrial clades were observed in the samples, one of which differed significantly in its geographic distribution relative to the other two within a small sampling area (similar to 70 x 35 km). This fact, supported by the inability of maximum likelihood analyses to achieve adequate fits to simple models for the population demography of the species, suggests a more complex history, possibly involving disjunct forest refugia. This hypothesis is supported by a genetic signal of recent population growth, which is expected if population sizes of this forest-obligate insect increased during the forest expansions that followed glacial periods. Although a time frame cannot be reliably inferred for the vicariance event leading to the three genetic clades, molecular clock estimates place this at similar to 1 Myr before present.

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.

Are corridors, fragment size and forest structure important for the conservation of leaf-litter lizards in a fragmented landscape?

To investigate the implications of forest fragmentation for conservation of leaf-litter lizards the importance of fragment size, corridors and forest structure was examined in 20 forest fragments and six localities within a continuous forest in the Atlantic Plateau of Sao Paulo state, Brazil. The fragments were 2-276 ha in area and had different degrees of connectivity depending on the presence or absence of corridors. Two species of lizards were dominant, Ecpleopus gaudichaudii and Enyalius perditus. Variation in forest structure among sites was important only in explaining the abundance of E. perditus. Regardless of variation in forest structure, lizard species composition, total lizard abundance, number of species and abundance of E. perditus were sensitive to fragmentation per se but not to fragment size or corridor linkage. The inhospitable matrix surrounding fragments is probably what determines the presence and abundance of E. perditus and the high er lizard richness in continuous forests. These conditions may have prevented lizard species from recolonizing the forest fragments. Our results emphasize that the conservation of this leaf-litter fauna depends on the maintenance of large tracts of continuous forests and not on the size of fragments or on the presence of forest connections. Strategies for conservation of leaf-litter lizards in such highly fragmented Atlantic Forest landscapes should consider the enlargement of landscape connectivity between fragments and continuous forest, allowing the latter areas to act as a source of individuals for fragments.

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.

Kerteszia subgenus of Anopheles associated with the Brazilian Atlantic rainforest: current knowledge and future challenges

Background: The Atlantic rainforest ecosystem, where bromeliads are abundant, provides an excellent environment for Kerteszia species, because these anophelines use the axils of those plants as larval habitat. Anopheles (K.) cruzii and Anopheles (K.) bellator are considered the primary vectors of malaria in the Atlantic forest. Although the incidence of malaria has declined in some areas of the Atlantic forest, autochthonous cases are still registered every year, with Anopheles cruzii being considered to be a primary vector of both human and simian Plasmodium. Methods: Recent publications that addressed ecological aspects that are important for understanding the involvement of Kerteszia species in the epidemiology of malaria in the Atlantic rainforest in the Neotropical Region were analysed. Conclusion: The current state of knowledge about Kerteszia species in relation to the Atlantic rainforest ecosystem was discussed. Emphasis was placed on ecological characteristics related to epidemiological aspects of this group of mosquitoes. The main objective was to investigate biological aspects of the species that should be given priority in future studies

Phylogeography of the common vampire bat (Desmodus rotundus): Marked population structure, Neotropical Pleistocene vicariance and incongruence between nuclear and mtDNA markers

Background: The common vampire bat Desmodus rotundus is an excellent model organism for studying ecological vicariance in the Neotropics due to its broad geographic range and its preference for forested areas as roosting sites. With the objective of testing for Pleistocene ecological vicariance, we sequenced a mitocondrial DNA (mtDNA) marker and two nuclear markers (RAG2 and DRB) to try to understand how Pleistocene glaciations affected the distribution of intraspecific lineages in this bat. Results: Five reciprocally monophyletic clades were evident in the mitochondrial gene tree, and in most cases with high bootstrap support: Central America (CA), Amazon and Cerrado (AMC), Pantanal (PAN), Northern Atlantic Forest (NAF) and Southern Atlantic Forest (SAF). The Atlantic forest clades formed a monophyletic clade with high bootstrap support, creating an east/west division for this species in South America. On the one hand, all coalescent and non-coalescent estimates point to a Pleistocene time of divergence between the clades. On the other hand, the nuclear markers showed extensive sharing of haplotypes between distant localities, a result compatible with male-biased gene flow. In order to test if the disparity between the mitochondrial and nuclear markers was due to the difference in mutation rate and effective size, we performed a coalescent simulation to examine the feasibility that, given the time of separation between the observed lineages, even with a gene flow rate close to zero, there would not be reciprocal monophyly for a neutral nuclear marker. We used the observed values of theta and an estimated mutation rate for the nuclear marker gene to perform 1000 iterations of the simulation. The results of this simulation were inconclusive: the number of iterations with and without reciprocal monophyly of one or more clades are similar. Conclusions: We therefore conclude that the pattern exhibited by the common vampire bat, with marked geographical structure for a mitochondrial marker and no phylogeographic structure for nuclear markers is compatible with a historical scenario of complete isolation of refuge-like populations during the Pleistocene. The results on demographic history on this species is compatible with the Carnaval-Moritz model of Pleistocene vicariance, with demographic expansions in the southern Atlantic forest.

Nitrogen dynamics during ecosystem development in tropical forest restoration

We considered whether ecological restoration using high diversity of native tree species serves to restore nitrogen dynamics in the Brazilian Atlantic Forest. We measured delta(15)N and N content in green foliage and soil; vegetation N:P ratio; and soil N mineralization in a preserved natural forest and restored forests of ages 21 and 52 years. Green foliage delta(15)N values, N content, N:P ratio, inorganic N and net mineralization and nitrification rates were all higher, the older the forest. Our findings indicate that the recuperation of N cycling has not been achieved yet in the restored forests even after 52 years, but show that they are following a trajectory of development that is characterized by their N cycling intensity becoming similar to a natural mature forest of the same original forest formation. This study demonstrated that some young restored forests are more limited by N compared to mature natural forests. We document that the recuperation of N cycling in tropical forests can be achieved through ecological restoration actions. (C) 2011 Elsevier B.V. All rights reserved.

Forest structure and live aboveground biomass variation along an elevational gradient of tropical Atlantic moist forest (Brazil)

Live aboveground biomass (AGB) is an important source of uncertainty in the carbon balance from the tropical regions in part due scarcity of reliable estimates of live AGB and its variation across landscapes and forest types. Studies of forest structure and biomass stocks of Neotropical forests are biased toward Amazonian and Central American sites. In particular, standardized estimates of aboveground biomass stocks for the Brazilian Atlantic forest are rarely available. Notwithstanding the role of environmental variables that control the distribution and abundance of biomass in tropical lowland forests has been the subject of considerable research, the effect of short, steep elevational gradients on tropical forest structure and carbon dynamics is not well known. In order to evaluate forest structure and live AGB variation along an elevational gradient (0-1100 m a.s.l.) of coastal Atlantic Forest in SE Brazil, we carried out a standard census of woody stems >= 4.8 cm dbh in 13 1-ha permanent plots established on four different sites in 2006-2007. Live AGB ranged from 166.3 Mg ha(-1) (bootstrapped 95% CI: 1444,187.0) to 283.2 Mg ha(-1) (bootstrapped 95% CI: 253.0,325.2) and increased with elevation. We found that local-scale topographic variation associated with elevation influences the distribution of trees >50 cm dbh and total live AGB. Across all elevations, we found more stems (64-75%) with limited crown illumination but the largest proportion of the live AGB (68-85%) was stored in stems with highly illuminated or fully exposed crowns. Topography, disturbance and associated changes in light and nutrient supply probably control biomass distribution along this short but representative elevational gradient. Our findings also showed that intact Atlantic forest sites stored substantial amounts of carbon aboveground. The live tree AGB of the stands was found to be lower than Central Amazonian forests, but within the range of Neotropical forests, in particular when compared to Central American forests. Our comparative data suggests that differences in live tree AGB among Neotropical forests are probably related to the heterogeneous distribution of large and medium-sized diameter trees within forests and how the live biomass is partitioned among those size classes, in accordance with general trends found by previous studies. In addition, the elevational variation in live AGB stocks suggests a large spatial variability over coastal Atlantic forests in Brazil, clearly indicating that it is important to consider regional differences in biomass stocks for evaluating the role of this threatened tropical biome in the global carbon cycle. (C) 2010 Elsevier B.V. All rights reserved.

The genus Rhagomys (Thomas 1917) (Rodentia, Cricetidae, Sigmodontinae) in South America: morphological considerations, geographic distribution and zoogeographic comments

American Museum of Natural History

Large-scale ecological restoration of high-diversity tropical forests in SE Brazil

The complex interactions among endangered ecosystems, landowners` interests, and different models of land tenure and use, constitute an important series of challenges for those seeking to maintain and restore biodiversity and augment the flow of ecosystem services. Over the past 10 years, we have developed a data-based approach to address these challenges and to achieve medium and large-scale ecological restoration of riparian areas on private lands in the state of Sao Paulo, southeastern Brazil. Given varying motivations for ecological restoration, the location of riparian areas within landholdings, environmental zoning of different riparian areas, and best-practice restoration methods were developed for each situation. A total of 32 ongoing projects, covering 527,982 ha, were evaluated in large sugarcane farms and small mixed farms, and six different restoration techniques have been developed to help upscale the effort. Small mixed farms had higher portions of land requiring protection as riparian areas (13.3%), and lower forest cover of riparian areas (18.3%), than large sugarcane farms (10.0% and 36.9%, respectively for riparian areas and forest cover values). In both types of farms, forest fragments required some degree of restoration. Historical anthropogenic degradation has compromised forest ecosystem structure and functioning, despite their high-diversity of native tree and shrub species. Notably, land use patterns in riparian areas differed markedly. Large sugarcane farms had higher portions of riparian areas occupied by highly mechanized agriculture, abandoned fields, and anthropogenic wet fields created by siltation in water courses. In contrast, in small mixed crop farms, low or non-mechanized agriculture and pasturelands were predominant. Despite these differences, plantations of native tree species covering the entire area was by far the main restoration method needed both by large sugarcane farms (76.0%) and small mixed farms (92.4%), in view of the low resilience of target sites, reduced forest cover, and high fragmentation, all of which limit the potential for autogenic restoration. We propose that plantations should be carried out with a high-diversity of native species in order to create biologically viable restored forests, and to assist long-term biodiversity persistence at the landscape scale. Finally, we propose strategies to integrate the political, socio-economic and methodological aspects needed to upscale restoration efforts in tropical forest regions throughout Latin America and elsewhere. (C) 2010 Elsevier BA/. All rights reserved.