Bráulio A. Santos

Multiple successional pathways in human-modified tropical landscapes : New insights from forest succession, forest fragmentation and landscape ecology research

Old‐growth tropical forests are being extensively deforested and fragmented worldwide. Yet forest recovery through succession has led to an expansion of secondary forests in human‐modified tropical landscapes (HMTLs). Secondary forests thus emerge as a potential repository for tropical biodiversity, and also as a source of essential ecosystem functions and services in HMTLs. Such critical roles are controversial, however, as they depend on successional, landscape and socio‐economic dynamics, which can vary widely within and across landscapes and regions. Understanding the main drivers of successional pathways of disturbed tropical forests is critically needed for improving management, conservation, and restoration strategies. Here, we combine emerging knowledge from tropical forest succession, forest fragmentation and landscape ecology research to identify the main driving forces shaping successional pathways at different spatial scales. We also explore causal connections between land‐use dynamics and the level of predictability of successional pathways, and examine potential implications of such connections to determine the importance of secondary forests for biodiversity conservation in HMTLs. We show that secondary succession (SS) in tropical landscapes is a multifactorial phenomenon affected by a myriad of forces operating at multiple spatio‐temporal scales. SS is relatively fast and more predictable in recently modified landscapes and where well‐preserved biodiversity‐rich native forests are still present in the landscape. Yet the increasing variation in landscape spatial configuration and matrix heterogeneity in landscapes with intermediate levels of disturbance increases the uncertainty of successional pathways. In landscapes that have suffered extensive and intensive human disturbances, however, succession can be slow or arrested, with impoverished assemblages and reduced potential to deliver ecosystem functions and services. We conclude that: (i) succession must be examined using more comprehensive explanatory models, providing information about the forces affecting not only the presence but also the persistence of species and ecological groups, particularly of those taxa expected to be extirpated from HMTLs; (ii) SS research should integrate new aspects from forest fragmentation and landscape ecology research to address accurately the potential of secondary forests to serve as biodiversity repositories; and (iii) secondary forest stands, as a dynamic component of HMTLs, must be incorporated as key elements of conservation planning; i.e. secondary forest stands must be actively managed (e.g. using assisted forest restoration) according to conservation goals at broad spatial scales.

Plant β‐diversity in fragmented rain forests: testing floristic homogenization and differentiation hypotheses

Summary Land-use change is the main driver of global biodiversity loss, but its relative impact on species turnover (β-diversity) across multiple spatial scales remains unclear. Plant communities in fragmented rain forests can undergo declines (floristic homogenization) or increases (floristic differentiation) in β-diversity. We tested these alternative hypotheses analysing a large vegetation data base from a hierarchically nested sampling design (450 plots in 45 forest patches in 3 landscapes with different deforestation levels) at Los Tuxtlas rain forest, Mexico. Differences in β-diversity across spatial scales (i.e. among plots, among patches, and among landscapes) were analysed using multiplicative diversity decompositions of Hill numbers. Plant β-diversity among plots within forest patches decreased in landscapes with higher deforestation levels, leading to floristic homogenization within patches. This homogenization process can be explained by the loss of rare and shade-tolerant plant species, and the recruitment and dominance of disturbance-adapted species, and can limit the accumulation of species (γ-diversity) in landscapes with higher deforestation. Nevertheless, the landscape with the highest deforestation level showed the highest floristic differentiation among patches. This landscape showed the greatest isolation distances between patches; a landscape spatial pattern that can limit the interchange of seeds (and species) between patches. Because the study patches are undergoing secondary succession following disturbances (e.g. logging, edge effects), different disturbance regimes and increased distance among patches could lead to higher β-diversity. Synthesis. These findings indicate that patterns of floristic homogenization and differentiation depend on the landscape configuration and on the spatial scale of analysis. At the landscape scale, our results suggest that, in accordance with non-equilibrium dynamics and the landscape-divergence hypothesis, patches located in landscapes with different forest cover and different connectivity can experience contrasting successional pathways due to increasing levels of compositional differentiation between patches. These novel findings add further uncertainties to the maintenance of biodiversity in severely deforested tropical landscapes and have key ecological implications for biodiversity conservation planning.

Edge-Related Loss of Tree Phylogenetic Diversity in the Severely Fragmented Brazilian Atlantic Forest

Deforestation and forest fragmentation are known major causes of nonrandom extinction, but there is no information about their impact on the phylogenetic diversity of the remaining species assemblages. Using a large vegetation dataset from an old hyper-fragmented landscape in the Brazilian Atlantic rainforest we assess whether the local extirpation of tree species and functional impoverishment of tree assemblages reduce the phylogenetic diversity of the remaining tree assemblages. We detected a significant loss of tree phylogenetic diversity in forest edges, but not in core areas of small (<80 ha) forest fragments. This was attributed to a reduction of 11% in the average phylogenetic distance between any two randomly chosen individuals from forest edges; an increase of 17% in the average phylogenetic distance to closest non-conspecific relative for each individual in forest edges; and to the potential manifestation of late edge effects in the core areas of small forest remnants. We found no evidence supporting fragmentation-induced phylogenetic clustering or evenness. This could be explained by the low phylogenetic conservatism of key life-history traits corresponding to vulnerable species. Edge effects must be reduced to effectively protect tree phylogenetic diversity in the severely fragmented Brazilian Atlantic forest.

Chronic anthropogenic disturbance drives the biological impoverishment of the Brazilian Caatinga vegetation

Summary In addition to acute transformations of ecosystems caused by deforestation, old-growth forests world-wide are being increasingly altered by low-intensity but chronic human disturbance. Overgrazing and the continuous extraction of forest products are important drivers of chronic disturbance, which can lead to the gradual local extinction of species and the alteration of vegetation structure. We tested this hypothesis in the Brazilian Caatinga vegetation, one of the most species-rich and populated semi-arid regions of the world. Using a multimodel averaging approach, we examined the impact of five recognized indicators of chronic disturbance (i.e. proximity to urban centre, houses, roads, density of people and livestock) on the diversity, abundance and evenness of 30 woody plant communities. We separately tested the response of seedlings, saplings and adults to identify the ontogenetic stages that are most susceptible to chronic disturbance. We recorded over 11 000 individuals belonging to 51 plant species. As expected, most indicators of chronic disturbance were negatively related to species diversity and stem abundance, with a variable effect on community evenness. The density of people and density of livestock were the main factors driving changes in plant communities, with a stronger negative impact on seedling and sapling diversities. Species composition also varied significantly with disturbance indicators, irrespective of ontogeny. Our results show the potential negative impact that chronic disturbance can have on Caatinga plant assemblages and highlight the fact that disturbance resulting from an extractivism-based and subsistence economy are probably driving old-growth forest stands towards shrub-dominated secondary stands. Synthesis and applications. These findings indicate that chronic disturbance should not continue to be neglected and we argue for: (i) research and rural programmes able to support better practices in terms of land use and sustainable exploitation of forest resources, (ii) improved governance and law enforcement to shift extractivism towards sustainable standards, and (iii) expanding the coverage and effective implementation of strictly protected areas.

Conserving Tropical Tree Diversity and Forest Structure: The Value of Small Rainforest Patches in Moderately-Managed Landscapes

Rainforests are undergoing severe deforestation and fragmentation worldwide. A huge amount of small forest patches are being created, but their value in conserving biodiversity and forest structure is still controversial. Here, we demonstrate that in a species-rich and moderately-managed Mexican tropical landscape small rainforest patches (<100 ha) can be highly valuable for the conservation of tree diversity and forest structure. These patches showed diverse communities of native plants, including endangered species, and a new record for the country. Although the number of logged trees increased in smaller patches, patch size was a poor indicator of basal area, stem density, number of species, genera and families, and community evenness. Cumulative species-area curves indicated that all patches had a similar contribution to the regional species diversity. This idea also was supported by the fact that patches strongly differed in floristic composition (high β-diversity), independently of patch size. Thus, in agreement with the land-sharing approach, our findings support that small forest patches in moderately-managed landscapes should be included in conservation initiatives to maintain landscape heterogeneity, species diversity, and ecosystem services.

Phylogenetic impoverishment of plant communities following chronic human disturbances in the Brazilian Caatinga.

Chronic disturbances, such as selective logging, firewood extraction and extensive grazing, may lead to the taxonomic and phylogenetic impoverishment of remaining old-growth forest communities worldwide; however, the empirical evidence on this topic is limited. We tested this hypothesis in the Caatinga vegetation--a seasonally dry tropical forest restricted to northeast Brazil. We sampled 11,653 individuals (adults, saplings, and seedlings) from 51 species in 29 plots distributed along a gradient of chronic disturbance. The gradient was assessed using a chronic disturbance index (CDI) based on five recognized indicators of chronic disturbances: proximity to urban center, houses and roads and the density of both people and livestock. We used linear models to test if mean effective number of lineages, mean phylogenetic distance and phylogenetic dispersion decreased with CDI and if such relationships differed among ontogenetic stages. As expected, the mean effective number of lineages and the mean phylogenetic distance were negatively related to CDI, and such diversity losses occurred irrespective of ontogeny. Yet the increase in phylogenetic clustering in more disturbed plots was only evident in seedlings and saplings, mostly because clades with more descendent taxa than expected by chance (e.g., Euphorbiaceae) thrived in more disturbed plots. This novel study indicates that chronic human disturbances are promoting the phylogenetic impoverishment of the irreplaceable woody flora of the Brazilian Caatinga forest. The highest impoverishment was observed in seedlings and saplings, indicating that if current chronic disturbances remain, they will result in increasingly poorer phylogenetically forests. This loss of evolutionary history will potentially limit the capacity of this ecosystem to respond to human disturbances (i.e., lower ecological resilience) and particularly their ability to adapt to rapid climatic changes in the region.

Phylogenetic Impoverishment of Amazonian Tree Communities in an Experimentally Fragmented Forest Landscape

Amazonian rainforests sustain some of the richest tree communities on Earth, but their ecological and evolutionary responses to human threats remain poorly known. We used one of the largest experimental datasets currently available on tree dynamics in fragmented tropical forests and a recent phylogeny of angiosperms to test whether tree communities have lost phylogenetic diversity since their isolation about two decades previously. Our findings revealed an overall trend toward phylogenetic impoverishment across the experimentally fragmented landscape, irrespective of whether tree communities were in 1-ha, 10-ha, or 100-ha forest fragments, near forest edges, or in continuous forest. The magnitude of the phylogenetic diversity loss was low (<2% relative to before-fragmentation values) but widespread throughout the study landscape, occurring in 32 of 40 1-ha plots. Consistent with this loss in phylogenetic diversity, we observed a significant decrease of 50% in phylogenetic dispersion since forest isolation, irrespective of plot location. Analyses based on tree genera that have significantly increased (28 genera) or declined (31 genera) in abundance and basal area in the landscape revealed that increasing genera are more phylogenetically related than decreasing ones. Also, the loss of phylogenetic diversity was greater in tree communities where increasing genera proliferated and decreasing genera reduced their importance values, suggesting that this taxonomic replacement is partially underlying the phylogenetic impoverishment at the landscape scale. This finding has clear implications for the current debate about the role human-modified landscapes play in sustaining biodiversity persistence and key ecosystem services, such as carbon storage. Although the generalization of our findings to other fragmented tropical forests is uncertain, it could negatively affect ecosystem productivity and stability and have broader impacts on coevolved organisms.

Are the vegetation structure and composition of the shrubby Caatinga free from edge influence

A influencia de borda, ou efeito de borda, leva a diversas alteracoes biologicas em paisagens fragmentadas. Embora muito estudada em ecossistemas florestais, a influencia de borda permanece desconhecida para a Caatinga. Baseada nas condicoes fisicas e biologicas da Caatinga arbustiva, nossa predicao a priori foi que este tipo de vegetacao nao sofre influencia de borda em termos de estrutura e composicao de especies. Nos amostramos arbustos, cactos e arvores em 20 parcelas de 200 m2 em bordas antigas (> 60 anos de idade) e interiores de um fragmento de 690 ha, parcialmente isolado e circundado por Opuntia ficus-indica. Altura das plantas, diâmetro ao nivel solo, densidade de individuos e riqueza e diversidade de especies foram estatisticamente iguais entre os habitats de borda e interior. A magnitude da influencia de borda variou de -0,027 a 0,027, indicando baixa importância ecologica da criacao de bordas para as variaveis analisadas. Adicionalmente, a similaridade floristica entre os habitats alcancou 90% e uma analise de componentes principais mostrou que a composicao de especies varia de forma similar na borda e no interior. O conjunto dos resultados indica que a vegetacao do fragmento estudado nao e ecologicamente afetada pela criacao de bordas e sugere que alteracoes nas condicoes fisicas e na disponibilidade de recursos apos a criacao das bordas sao insuficientes para eliminar plantas ja estabelecidas ou alterar o recrutamento e a sobrevivencia de novos individuos.

Community-Level Patterns of Insect Herbivory in a Fragmented Atlantic Forest Landscape

ABSTRACT Insect herbivores largely affect plant population structure, community organization, and ecosystem functioning, but little is known on how insect herbivory is altered in human-modified landscapes. Here we assessed 3,566 woody seedlings inhabiting 20 Atlantic forest fragments (3-91 ha) in northeast Brazil to examine the extent to which standing levels of herbivory on woody seedlings correlated with forest fragment metrics (fragment area and distance to forest edge) and resource availability (pioneer plants). Overall, 78% of all seedlings and 36% of the 23,003 recorded leaves experienced injuries caused by folivorous insects, the bulk of them promoted by chewing insects (85.9% of damaged leaves). This insect guild removed 9.2 ±1.9 cm2 of foliar tissue per leaf, which represented 10.2 ±1.8% of the standing leaf area. Contrary to our expectations, frequency and magnitude of foliar damage by insects were statistically uncorrelated to either basic forest fragment metrics (fragment area, edge proximity) or resource abundance (percentage of pioneer seedlings). Our findings indicate that insect herbivory is a pervasive ecological process in fragmented landscapes. However, rather than being a function of simple fragment metrics or resource availability, its variation seems to be caused by a range of drivers, such as dispersal ability of folivorous insects and vulnerability of their parasitoids and predators to human disturbance.

Does forest fragmentation cause an increase in forest temperature

Forest fragmentation is considered by many to be a primary cause of the current biodiversity crisis. The underlying mechanisms are poorly known, but a potentially important one is associated with altered thermal conditions within the remaining forest patches, especially at forest edges. Yet, large uncertainty remains about the effect of fragmentation on forest temperature, as it is unclear whether temperature decreases from forest edge to forest interior, and whether this local gradient scales up to an effect of fragmentation (landscape attribute) on temperature. We calculated the effect size (correlation coefficient) of distance from forest edge on air temperature, and tested for differences among forest types surrounded by different matrices using meta-analysis techniques. We found a negative edge-interior temperature gradient, but correlation coefficients were highly variable, and significant only for temperate and tropical forests surrounded by a highly contrasting open matrix. Nevertheless, it is unclear if these local-scale changes in temperature can be scaled up to an effect of fragmentation on temperature. Although it may be valid when considering “fragmentation” as forest loss only, the landscape-scale inference is not so clear when we consider the second aspect of fragmentation, where a given amount of forest is divided into a large number of small patches (fragmentation per se). Therefore, care is needed when assuming that fragmentation changes forest temperature, as thermal changes at forest edges depend on forest type and matrix composition, and it is still uncertain if this local gradient can be scaled up to the landscape.