Induced drought strongly affects richness and composition of ground-dwelling ants in the eastern Amazon.

Species loss in tropical regions is forecast to occur under environmental change scenarios of low precipitation. One of the main questions is how drought will affect invertebrates, a key group for ecosystem functioning. We use 1 year of data from a long-term rainwater exclusion experiment in primary Amazonian rainforest to test whether induced water stress and covarying changes in soil moisture, soil respiration, and tree species richness, diversity, size, and total biomass affected species richness and composition (relative abundance) of ground-dwelling ants. Data on ant abundance and environmental variables were collected at two sites (control and experimental) in the Eastern Amazon. Since 2002, drought has been induced in the experimental plot by excluding 50% of normal rainfall. Ant species richness in the experiment plot was reduced and some generalist species responded positively. Ant species richness also increased in the experimental plot with increasing diversity of the plant species of the leaf litter. The relative abundance of ants differed between plots. The experimental plot was characterized by a higher frequency of generalist and other species that appeared to be favored by the reduction in rainfall. Between-plot comparisons suggested loss and changes in ant species composition in tropical forests were affected by increasing dryness. These changes could ultimately lead to cascading effects on ecosystem processes and the services they mediate.

Plant traits controlling growth change in response to a drier climate.

Plant traits are increasingly being used to improve prediction of plant function, including plant demography. However, the capability of plant traits to predict demographic rates remains uncertain, particularly in the context of trees experiencing a changing climate. Here we present data combining 17 plant traits associated with plant structure, metabolism and hydraulic status, with measurements of long-term mean, maximum and relative growth rates for 176 trees from the world's longest running tropical forest drought experiment. We demonstrate that plant traits can predict mean annual tree growth rates with moderate explanatory power. However, only combinations of traits associated more directly with plant functional processes, rather than more commonly employed traits like wood density or leaf mass per area, yield the power to predict growth. Critically, we observe a shift from growth being controlled by traits related to carbon cycling (assimilation and respiration) in well-watered trees, to traits relating to plant hydraulic stress in drought-stressed trees. We also demonstrate that even with a very comprehensive set of plant traits and growth data on large numbers of tropical trees, considerable uncertainty remains in directly interpreting the mechanisms through which traits influence performance in tropical forests.

Small tropical forest trees have a greater capacity to adjust carbon metabolism to long-term drought than large canopy trees.

The response of small understory trees to long-term drought is vital in determining the future composition, carbon stocks and dynamics of tropical forests. Long-term drought is, however, also likely to expose understory trees to increased light availability driven by drought-induced mortality. Relatively little is known about the potential for understory trees to adjust their physiology to both decreasing water and increasing light availability. We analysed data on maximum photosynthetic capacity (Jmax , Vcmax ), leaf respiration (Rleaf ), leaf mass per area (LMA), leaf thickness and leaf nitrogen and phosphorus concentrations from 66 small trees across 12 common genera at the world's longest running tropical rainfall exclusion experiment and compared responses to those from 61 surviving canopy trees. Small trees increased Jmax , Vcmax , Rleaf and LMA (71, 29, 32, 15% respectively) in response to the drought treatment, but leaf thickness and leaf nutrient concentrations did not change. Small trees were significantly more responsive than large canopy trees to the drought treatment, suggesting greater phenotypic plasticity and resilience to prolonged drought, although differences among taxa were observed. Our results highlight that small tropical trees have greater capacity to respond to ecosystem level changes and have the potential to regenerate resilient forests following future droughts.

Amazonia trees have limited capacity to acclimate plant hydraulic properties in response to long-term drought.

The fate of tropical forests under future climate change is dependent on the capacity of their trees to adjust to drier conditions. The capacity of trees to withstand drought is likely to be determined by traits associated with their hydraulic systems. However, data on whether tropical trees can adjust hydraulic traits when experiencing drought remain rare. We measured plant hydraulic traits (e.g. hydraulic conductivity and embolism resistance) and plant hydraulic system status (e.g. leaf water potential, native embolism and safety margin) on >150 trees from 12 genera (36 species) and spanning a stem size range from 14 to 68 cm diameter at breast height at the world's only long-running tropical forest drought experiment. Hydraulic traits showed no adjustment following 15 years of experimentally imposed moisture deficit. This failure to adjust resulted in these drought-stressed trees experiencing significantly lower leaf water potentials, and higher, but variable, levels of native embolism in the branches. This result suggests that hydraulic damage caused by elevated levels of embolism is likely to be one of the key drivers of drought-induced mortality following long-term soil moisture deficit. We demonstrate that some hydraulic traits changed with tree size, however, the direction and magnitude of the change was controlled by taxonomic identity. Our results suggest that Amazonian trees, both small and large, have limited capacity to acclimate their hydraulic systems to future droughts, potentially making them more at risk of drought-induced mortality.

Evolutionary diversity in tropical tree communities peaks at intermediate precipitation.

Global patterns of species and evolutionary diversity in plants are primarily determined by a temperature gradient, but precipitation gradients may be more important within the tropics, where plant species richness is positively associated with the amount of rainfall. The impact of precipitation on the distribution of evolutionary diversity, however, is largely unexplored. Here we detail how evolutionary diversity varies along precipitation gradients by bringing together a comprehensive database on the composition of angiosperm tree communities across lowland tropical South America (2,025 inventories from wet to arid biomes), and a new, large-scale phylogenetic hypothesis for the genera that occur in these ecosystems. We find a marked reduction in the evolutionary diversity of communities at low precipitation. However, unlike species richness, evolutionary diversity does not continually increase with rainfall. Rather, our results show that the greatest evolutionary diversity is found in intermediate precipitation regimes, and that there is a decline in evolutionary diversity above 1,490 mm of mean annual rainfall. If conservation is to prioritise evolutionary diversity, areas of intermediate precipitation that are found in the South American 'arc of deforestation', but which have been neglected in the design of protected area networks in the tropics, merit increased conservation attention.

Isoprene emission structures tropical tree biogeography and community assembly responses to climate.

The prediction of vegetation responses to climate requires a knowledge of how climate-sensitive plant traits mediate not only the responses of individual plants, but also shifts in the species and functional compositions of whole communities. The emission of isoprene gas - a trait shared by one-third of tree species - is known to protect leaf biochemistry under climatic stress. Here, we test the hypothesis that isoprene emission shapes tree species compositions in tropical forests by enhancing the tolerance of emitting trees to heat and drought. Using forest inventory data, we estimated the proportional abundance of isoprene-emitting trees (pIE) at 103 lowland tropical sites. We also quantified the temporal composition shifts in three tropical forests - two natural and one artificial - subjected to either anomalous warming or drought. Across the landscape, pIE increased with site mean annual temperature, but decreased with dry season length. Through time, pIE strongly increased under high temperatures, and moderately increased following drought. Our analysis shows that isoprene emission is a key plant trait determining species responses to climate. For species adapted to seasonal dry periods, isoprene emission may tradeoff with alternative strategies, such as leaf deciduousness. Community selection for isoprene-emitting species is a potential mechanism for enhanced forest resilience to climatic change.

Drought stress and tree size determine stem CO2 efflux in a tropical forest.

CO2 efflux from stems (CO2_stem ) accounts for a substantial fraction of tropical forest gross primary productivity, but the climate sensitivity of this flux remains poorly understood. We present a study of tropical forest CO2_stem from 215 trees across wet and dry seasons, at the world's longest running tropical forest drought experiment site. We show a 27% increase in wet season CO2_stem in the droughted forest relative to a control forest. This was driven by increasing CO2_stem in trees 10-40 cm diameter. Furthermore, we show that drought increases the proportion of maintenance to growth respiration in trees > 20 cm diameter, including large increases in maintenance respiration in the largest droughted trees, > 40 cm diameter. However, we found no clear taxonomic influence on CO2_stem and were unable to accurately predict how drought sensitivity altered ecosystem scale CO2_stem , due to substantial uncertainty introduced by contrasting methods previously employed to scale CO2_stem fluxes. Our findings indicate that under future scenarios of elevated drought, increases in CO2_stem may augment carbon losses, weakening or potentially reversing the tropical forest carbon sink. However, due to substantial uncertainties in scaling CO2_stem fluxes, stand-scale future estimates of changes in stem CO2 emissions remain highly uncertain.

Stand dynamics modulate water cycling and mortality risk in droughted tropical forest.

Transpiration from the Amazon rainforest generates an essential water source at a global and local scale. However, changes in rainforest function with climate change can disrupt this process, causing significant reductions in precipitation across Amazonia, and potentially at a global scale. We report the only study of forest transpiration following a long-term (>10 year) experimental drought treatment in Amazonian forest. After 15 years of receiving half the normal rainfall, drought-related tree mortality caused total forest transpiration to decrease by 30%. However, the surviving droughted trees maintained or increased transpiration because of reduced competition for water and increased light availability, which is consistent with increased growth rates. Consequently, the amount of water supplied as rainfall reaching the soil and directly recycled as transpiration increased to 100%. This value was 25% greater than for adjacent nondroughted forest. If these drought conditions were accompanied by a modest increase in temperature (e.g., 1.5°C), water demand would exceed supply, making the forest more prone to increased tree mortality.

Species richness and composition of epiphytic bryophytes in flooded forests of Caxiuanã National Forest, Eastern Amazon, Brazil.

This study aimed to compare the richness and composition of the epiphytic bryoflora between várzea and igapó forests in Caxiuanã National Forest, Brazilian Amazon. Bryophytes were collected on 502 phorophytes of Virola surinamensis. Average richness per phorophyte and composition between forests and between dry and rainy periods was tested by two-way analysis and by cluster analysis, respectively. In total, 54 species of 13 families were identified. Richness was greater in igapó forest (44 species) compared to várzea forest (38 species). There was no significant difference in the number of species between the studied periods. Cluster analysis showed the bryoflora composition was different between várzea and igapó, but not between dry and rainy periods. Results did not corroborate the hypothesis that várzea forests harbor higher species richness than igapó forests.

Species richness and composition of epiphytic bryophytes in flooded forests of Caxiuanã National Forest, Eastern Amazon, Brazil

ABSTRACT This study aimed to compare the richness and composition of the epiphytic bryoflora between várzea and igapó forests in Caxiuanã National Forest, Brazilian Amazon. Bryophytes were collected on 502 phorophytes of Virola surinamensis. Average richness per phorophyte and composition between forests and between dry and rainy periods was tested by two-way analysis and by cluster analysis, respectively. In total, 54 species of 13 families were identified. Richness was greater in igapó forest (44 species) compared to várzea forest (38 species). There was no significant difference in the number of species between the studied periods. Cluster analysis showed the bryoflora composition was different between várzea and igapó, but not between dry and rainy periods. Results did not corroborate the hypothesis that várzea forests harbor higher species richness than igapó forests.

The effect of exotic grass Urochloa decumbens (Stapf) R.D.Webster (Poaceae) in the reduction of species richness and change of floristic composition of natural regeneration in the Floresta Nacional de Carajás, Brazil.

The introduction of exotic species is considered as one of the major causes of biodiversity loss. The National Forest of Carajás is one of the largest mineral provinces in the world. Mining activities caused changes of the natural habitats, leaving degraded areas after the mineral exploitation. One of the mining areas within FLONA Carajás was used for the extraction of gold. In the process of exploitation, a huge depression was formed by the removal of soil which was mounded up nearby. To prevent soil erosion of these mounds, an exotic grass, Urochloa decumbens (Stapf) R.D.Webster (Poaceae) was planted. The objective of this study was to compare the impact of this non-native grass on species richness and species composition of the natural regeneration in the degraded areas. Four areas were compared, two with and two without presence of U. decumbens. In each area, twenty four 1m²/plots were established. Species richness of the regeneration areas and population sizes were significantly lower in the plots where the exotic grass was present. Our study shows that U. decumbens had a negative effect on species richness and population density, and its presence changed the species composition and distribution of life forms of the natural regeneration.

Seasonality of reproduction of epiphytic bryophytes in flooded forests from the Caxiuanã National Forest, Eastern Amazon.

This work aimed to recognize the reproductive biology of the epiphytic bryoflora of phorophytes of Virola surinamensis (Rol. ex. Rottb.) Warb. in várzea and igapó forests in the Caxiuanã National Forest, to answer the following question: The reproductive period of the bryophyte species is influenced by the environment due the climatic seasonality present in flooded forests, being higher the occurrence of the sexual and asexual reproduction in the rainiest months? The bryophytes were identified and analyzed for the type of reproduction, sexual system and reproductive structures. In total, 502 samples of bryophytes were analyzed, resulting in 54 species, of which 34 were fertile. The comparison of the fertility of the species in different environmental conditions (dry or rainy, and igapó or várzea forest) was assessed using the chi-square test. The fertility of the seven studied species could not be defined by a pattern, considering the forest type and the seasonality. However, two species were associated to the forest type and two further species to the seasonality, showing that, for some bryophyte species, invest in constant fertility may be favoring the maintenance of their populations in tropical forests.

Threshold Responses to Soil Moisture Deficit by Trees and Soil in Tropical Rain Forests: Insights from Field Experiments.

Many tropical rain forest regions are at risk of increased future drought. The net effects of drought on forest ecosystem functioning will be substantial if important ecological thresholds are passed. However, understanding and predicting these effects is challenging using observational studies alone. Field-based rainfall exclusion (canopy throughfall exclusion; TFE) experiments can offer mechanistic insight into the response to extended or severe drought and can be used to help improve model-based simulations, which are currently inadequate. Only eight TFE experiments have been reported for tropical rain forests. We examine them, synthesizing key results and focusing on two processes that have shown threshold behavior in response to drought: (1) tree mortality and (2) the efflux of carbon dioxdie from soil, soil respiration. We show that: (a) where tested using large-scale field experiments, tropical rain forest tree mortality is resistant to long-term soil moisture deficit up to a threshold of 50% of the water that is extractable by vegetation from the soil, but high mortality occurs beyond this value, with evidence from one site of increased autotrophic respiration, and (b) soil respiration reaches its peak value in response to soil moisture at significantly higher soil moisture content for clay-rich soils than for clay-poor soils. This first synthesis of tropical TFE experiments offers the hypothesis that low soil moisture-related thresholds for key stress responses in soil and vegetation may prove to be widely applicable across tropical rain forests despite the diversity of these forests.

Forest structure of artificial islands in the Tucuruí dam reservoir in northern Brazil: a test core-area model / Estrutura da floresta em ilhas artificiais no reservatório da usina hidrelétrica de Tucuruí, Brasil: um teste do modelo de área nuclear

Construction of hydroelectric dams in tropical regions has been contributing significantly to forest fragmentation. Alterations at edges of forest fragments impact plant communities that suffer increases in tree damage and dead, and decreases in seedling recruitment. This study aimed to test the core-area model in a fragmented landscape caused by construction of a hydroelectric power plant in the Brazilian Amazon. We studied variations in forest structure between the margin and interiors of 17 islands of 8-100 hectares in the Tucuruí dam reservoir, in two plots (30 and >100m from the margin) per island. Mean tree density, basal area, seedling density and forest cover did not significantly differ between marginal and interior island plots. Also, no significant differences were found in liana density, dead tree or damage for margin and interior plots. The peculiar topographic conditions associated with the matrix habitat and shapes of the island seem to extend edge effects to the islands' centers independently of the island size, giving the interior similar physical microclimatic conditions as at the edges. We propose a protocol for assessing the ecological impacts of edge effects in fragments of natural habitat surrounded by induced (artificial) edges. The protocol involves three steps: (1) identification of focal taxa of particular conservation or management interest, (2) measurement of an "edge function" that describes the response of these taxa to induced edges, and (3) use of a "Core-Area Model" to extrapolate edge function parameters to existing or novel situations.

A construção de usinas hidrelétricas em regiões tropicais tem contribuído significativamente para a fragmentação da floresta. As alterações nas bordas de fragmentos florestais causam profundos impactos na comunidade de plantas, tais como, o aumento em de árvores mortas ou danificadas e a diminuição do recrutamento de plântulas. Este estudo tem como objetivo testar o modelo de área nuclear (core-area model) em uma paisagem de floresta fragmentada resultante da formação do lago da Usina Hidrelétrica de Tucucui na Amazônia. Foram medidas as variações na estrutura em 17 ilhas, com tamanhos variando de 8 a 100 hectares. Em cada ilha foram colocadas duas parcelas, a 1ª parcela a 30 metros da margem e a 2ª parcela a mais de 100 metros da margem da ilha. A densidade e a área basal de árvores, a densidade da regeneração natural e a cobertura do dossel da floresta não foram significativamente diferentes entre as parcelas da borda e do interior das ilhas. Também não foram encontradas diferenças significativas na densidade de lianas, no número de troncos mortos ou árvores danificadas entre as parcelas da borda e do interior das ilhas. A grande declividade do terreno das ilhas, a matriz circundante (água) e a forma irregular da ilhas podem ser os fatores responsáveis me provocar a extensão do efeito de borda em todas as ilhas analisadas, independente do tamanho e grau de isolamento. Nós propomos um novo protocolo para avaliar os impactos ecológicos dos efeitos de borda em fragmentos de habitat criados artificialmente. Este protocolo envolve três etapas: (1) identificação dos táxons biológicos de particular interesse de conservação ou de gestão (2) medição das das variáveis responsaveis pela resposta dos táxons induzidas pelo efeito de borda e (3) Usar o modelo de área nuclear (core area model) na extrapolação das variáveis medidas em situacões novas ou existentes.

The cost-effectiveness of biodiversity surveys in tropical forests.

The identification of high-performance indicator taxa that combine practical feasibility and ecological value requires an understanding of the costs and benefits of surveying different taxa. We present a generic and novel framework for identifying such taxa, and illustrate our approach using a large-scale assessment of 14 different higher taxa across three forest types in the Brazilian Amazon, estimating both the standardized survey cost and the ecological and biodiversity indicator value for each taxon. Survey costs varied by three orders of magnitude, and dung beetles and birds were identified as especially suitable for evaluating and monitoring the ecological consequences of habitat change in our study region. However, an exclusive focus on such taxa occurs at the expense of understanding patterns of diversity in other groups. To improve the cost-effectiveness of biodiversity research we encourage a combination of clearer research goals and the use of an objective evidence-based approach to selecting study taxa.