Southern Brazilian Atlantic Forest Research Articles

Phylogeography of Sodreaninae harvestmen (Arachnida: Opiliones: Gonyleptidae): Insights into the biogeography of the southern Brazilian Atlantic Forest

The Brazilian Atlantic Forest has long been considered a global biodiversity hotspot. In the last decade, the phylogeographic patterns of endemic taxa have been unraveling the biogeographic history of the biome. However, highly diverse invertebrate species have still been poorly studied. Sodreana harvestmen (Gonyleptidae) are distributed in most of the humid coastal forests in the southern portion of the Atlantic Forest, a region that has experienced complex topographic evolution and differing climatic conditions since the Early Cretaceous, which likely affected the geographic distribution and diversification of the group. In this study, we investigated the molecular phylogeny and phylogeography of Sodreana to clarify the species relationships and to make inferences about the historical biogeography of the southern Atlantic Forest. We applied coalescent-based phylogenetic analyses using one mitochondrial and three nuclear markers coupled with an ecological niche modeling approach to verify relationships among species, date the main divergence events in the genus, and make inferences concerning possible changes in the geographical distribution and population dynamics from the past. Our results supported the validity of most Sodreana species and suggested that Paleogene-Neogene geomorphologic processes such as the formation of rivers systems, uplift of mountain ranges and related environmental changes have profoundly affected the evolutionary history of Sodreana. The ecological niche models showed that the areas potentially occupied by the species were greatly reduced during Quaternary glacial periods but no recent lineage divergences or genetic bottlenecks were detected, suggesting that climatically stable micro-habitats could have helped maintain populations during drier periods. Our study highlights the importance of humidity-dependent and poor-dispersal taxa in understanding the effects of ancient geological and climate processes on the Atlantic Forest biota.

Consumption of Euterpe edulis fruit by wildlife: implications for conservation and management of the Southern Brazilian Atlantic Forest.

This study aimed to measure the wildlife consumption of Euterpe edulis fruit and use this data to discuss management possibilities. To estimate infructescence fruit volume consumed, collectors were installed in fruit-bearing palms. To characterize consumption from the ground, samples were placed next to fruiting palms. To identify wildlife and their activities, camera traps were installed in infructescences and on the ground. The results suggested that there was a small fruit surplus (1.8 %), and this finding indicated the possibility of a harvest to reduce food for the wildlife. However, recurrent variations in the annual fruit production (21.4 %) were also noted, and suggested that wildlife could tolerate some fruit harvesting. Thus, a harvest could be restricted to fruit volume that exceeds the annual average (94 kg/ha/year). Turdus flavipes, a migratory bird, was the most active species in the dispersal of seeds; this finding indicates the need for broader conservation strategies. Wildlife composition also changed along with the fruiting, and this alteration suggests that dependence on the fruit is variable among different species. Seed germination and seedling mortality were high, results that indicate that local conditions may have a predominant effect on seed volume in natural regeneration density.

Robust volumetric models for supporting the management of secondary forest stands in the Southern Brazilian Atlantic Forest.

The majority of Atlantic Forest fragments in Southern Brazil are second-growth forests dominated by fast-growing species with considerable market-value timber. Nevertheless, volume prediction models are scarce, especially to estimate tree total volume (i.e., stem plus branches). This study approached the issue through the following aims: to fit and select stem and total volume models (generic and species-specific) using data from 288 harvested trees in a management operation, and to fit generic and species-specific bark factors. The power model embedding diameter at breast height (D) and tree stem or total height (H) presented the greatest prediction strength for both stem and total tree volume. Models including only D to predict total tree volume were similar to double-entry models regarding goodness-of-fit. Therefore, they may be useful in the context of subtropical closed-canopy forests, where the difficulty and uncertainty in H measurements are not trivial. Species-specific models fitted for Miconia cinnamomifolia (DC) Naudin. and Hyeronima alchorneoides Allemão surpassed generic models only for the former species. Nevertheless, the prediction improvement should offset the eventual extra efforts implied in the collection of reliable samples of these species. Finally, bark factors stood as a satisfactory tool for inside bark mean volume estimation.

Disentangling the correlates of species and site contributions to beta diversity in dung beetle assemblages

AbstractAimWe aimed to test whether contributions of individual species (SCBD) and contributions of single sites (LCBD) to overall beta diversity can be predicted by species metrics and species characteristics and also by community metrics and ecological variables, respectively.LocationA mainland‐island landscape in the southern Brazilian Atlantic Forest domain.MethodsDung beetles were sampled along 100 sampling units within four large forest fragments. We partitioned beta diversity into SCBD and LCBD aiming to explore their relationships to species and site characteristics, respectively. We then used a combination of multivariate methods and beta regression analyses to examine patterns in SCBD and LCBD. The occupancy, total abundance, niche position, niche breadth and biological traits of species were used as predictor variables for SCBD values. Community metrics, environmental and spatial variables, and temporal patterns in the beta diversity components of turnover and nestedness were used as predictor variables for LCDB values.ResultsWe found that SCBD was strongly related to various species characteristics, such as occupancy, abundance and niche position, but was not related to biological traits of species and niche breadth. In particular, occupancy and its quadratic term showed a very strong unimodal relationship with SCBD, suggesting that intermediate species in terms of site occupancy contribute most to beta diversity. LCBD was mostly explained by variation in species richness, with a negative relationship being detected. Litter height and large‐scale spatial variables were also important in explaining variation in LCBD.Main conclusionsSCBD and LCBD were highly predictably related to species occupancy and species richness, respectively. Environmental conditions and large‐scale spatial variables also correlated with LCBD values. Understanding the determinants of SCBD and LCBD may thus hold a key to various general ecological, bioassessment and conservation issues. Protecting sites with high LCBD values may be a suitable approach to practical biodiversity conservation.

Tree community dynamics in a forest adjacent to the reservoir of a hydroelectric power plant in the southern Brazilian Atlantic Forest

This study aimed at analyzing the dynamics of tree components in a forest adjacent to the Barra Grande Hydroelectric Power Plant reservoir, in the southern Brazilian Atlantic Forest, located in the municipality of Capão Alto. In 2013, this area was previously inventoried and divided into three sectors according to the distance from the reservoir: lower sector (in contact with the reservoir), intermediate sector (a slope area following the first sector) and upper sector (near the slope top). Fifty plots were allocated in the sectors, where all tree individuals with DBH (diameter at breast height) higher than or equal to 5 cm were inventoried (identified and measured at the circumference). In 2017 the surviving individuals were measured again, the dead ones were registered and the recruits (those with DBH reaching 5 cm) were included into the survey. Dynamics rates in number of individuals and basal area were calculated. The tree community under study has shown species richness stability, though also structural development, which is typical of areas that are facing a self-thinning process. The higher density populations of each sector have shown a similar structural dynamics pattern. There were no differences in the demographic rates among sectors, indicating that, approximately 12 years after the reservoir filling, the forest dynamics has not been influenced by the reservoir but by ecological factors dependent on density, such as competition.

Chusquea kleinii, a new bamboo from the Atlantic forests of Brazil segregated from C. capituliflora (Poaceae: Bambusoideae)

A new species belonging to the woody bamboo genus Chusquea (Poaceae, Bambusoideae) is herein segregated from C. capituliflora and described as C. kleinii, based upon observations of macromorphological characters made on living plants in the field, and the micromorphology of leaf surfaces, especially papillae on the abaxial epidermis. Both taxa are endemic to the southern Brazilian Atlantic forest.

Forest regeneration affects dung beetle assemblages (Coleoptera: Scarabaeinae) in the southern Brazilian Atlantic Forest

The Brazilian Atlantic Forest is one of the most diverse environments, but it is also one of the most threatened areas in terms of loss of biodiversity and ecosystem services. Assessment of changes in the community structure during the recovery of forests can be performed using indicator organisms. Dung beetles perform several ecological functions and show high sensitivity to natural and anthropogenic environmental changes. This study aimed to investigate the effect of regeneration time of Atlantic Forest sites on structure of Scarabaeinae assemblages. We sampled dung beetles using ten baited pitfall traps per site, in six sites grouped into three classes of forest regeneration time (~30, ~60 and >80 years) in the southern Brazilian Atlantic Forest, during January 2015. A total of 520 individuals belonging to 16 species and nine genera of dung beetles were sampled. Rarefied species richness did not differ between sites with different regeneration times. Average species richness and abundance of Scarabaeinae was smaller in areas of shorter recovery time. True alpha diversity was higher in areas with intermediate recovery whereas Shannon diversity showed higher values in areas of shorter recovery. Approximately 29 % of the variation in abundance data of Scarabaeinae was explained by environmental variables, with one-third of this variation explained also by spatial predictors. External factors such as landscape management and farming practices in the surroundings must be taken into consideration in management plans and the management of natural areas for the recovery of biodiversity in the Atlantic Forest. These external factors can considerably affect the structure of communities and lead to scenarios of greater diversity in intermediate regeneration sites due to the heterogeneity of the landscape.

Species delimitation, phylogeny and evolutionary demography of co-distributed, montane frogs in the southern Brazilian Atlantic Forest

The Brazilian Atlantic Forest (BAF) is recognized as one of the world’s biodiversity hotspots, with even more species per unit of area than the Amazon, however the mechanisms that led to such astonishing diversity are yet to be fully understood. In this study, we investigate the diversification of two co-distributed frog genera associated with montane areas of southern BAF: Melanophryniscus (Bufonidae) and Brachycephalus (Brachycephalidae). Species delimitation methods using mitochondrial and nuclear loci supported the existence of a remarkable number of highly endemic species in each genus, most of which occupy only one or a few adjacent mountaintops. Their timing of diversification was highly congruent, supporting recent speciation events within the past 600 thousand years. Extended Bayesian skyline plots indicate that most populations have remained relatively stable in size across the evolutionary past, with recent growth after 0.15My, suggesting that the drastic changes found in previous studies on lowland frog species were not shared by these montane taxa. These results are consistent with the existence of a montane refugium in southern BAF, allowing species persistence through the climatic shifts experienced along the BAF during the Quaternary.

Crossing the threshold: an amphibian assemblage highly infected with Batrachochytrium dendrobatidis in the southern Brazilian Atlantic forest

ABSTRACTThe fungal infection caused by Batrachochytrium dendrobatidis (Bd) in amphibians is known to be lethal when infection intensity values exceed loads of 10,000 zoospores per individual. We investigated Bd infection intensity in 100 anurans of southern Brazil. Almost half of the individuals were infected and the intensity ranged from four to about 156,000 zoospore genomic equivalents. We found no clinical signs of chytridiomycosis and no evidence of mortality. However, we observed a reduction in the number of infected individuals with loads above 10,000 zoospores. This fact could be considered indirect evidence that individuals with high loads are removed from the population.

Spatial variation of dung beetle assemblages associated with forest structure in remnants of southern Brazilian Atlantic Forest

The Brazilian Atlantic Forest is one of the world's biodiversity hotspots, and is currently highly fragmented and disturbed due to human activities. Variation in environmental conditions in the Atlantic Forest can influence the distribution of species, which may show associations with some environmental features. Dung beetles (Coleoptera: Scarabaeinae) are insects that act in nutrient cycling via organic matter decomposition and have been used for monitoring environmental changes. The aim of this study is to identify associations between the spatial distribution of dung beetle species and Atlantic Forest structure. The spatial distribution of some dung beetle species was associated with structural forest features. The number of species among the sampling sites ranged widely, and few species were found in all remnant areas. Principal coordinates analysis indicated that species composition, abundance and biomass showed a spatially structured distribution, and these results were corroborated by permutational multivariate analysis of variance. The indicator value index and redundancy analysis showed an association of several dung beetle species with some explanatory environmental variables related to Atlantic Forest structure. This work demonstrated the existence of a spatially structured distribution of dung beetles, with significant associations between several species and forest structure in Atlantic Forest remnants from Southern Brazil.

Ancient divergence and recent population expansion in a leaf frog endemic to the southern Brazilian Atlantic forest

The evolutionary history of Neotropical organisms has been often interpreted through broad-scale generalizations. The most accepted model of diversification for the Brazilian Atlantic forest (BAF) rely on putative historical stability of northern areas and massive past habitat replacement of its southern range. Here, we use the leaf frog Phyllomedusa distincta, endemic to the southern BAF, to better understand diversification patterns within this underexplored rainforest region. We used an integrative approach coupling fine-scale sampling and multilocus sequence data, with traditional and statistical phylogeographic (multilocus approximate Bayesian computation) methods to explore alternative hypotheses of diversification. We also employed species paleodistribution modeling to independently verify habitat stability upon a spatially explicit model. Our data support two divergent lineages with coherent geographic distribution that span throughout northern and southern ranges. Demographic estimates suggested the Southern lineage has experienced a recent population expansion, whereas the Northern lineage remained more stable. Hypothesis testing supports a scenario of ancient vicariance with recent population expansion. The paleodistribution model revealed habitat discontinuity during the Last Glacial Maximum (LGM) with one area of putative stability within the range of the Northern lineage. Evidence on genetic structure, demography, and paleodistribution of P. distincta support a historically heterogeneous landscape for the southern BAF, with both areas of forest stability and regions where forest occupation is probably recent. We also associate the southern end of the Cubatao shear zone with a phylogeographic break in the BAF. Taken together, our results argue for the idea of multiple mechanisms generating diversity in this biome and underscore the need of fine-scale data in revealing more detailed pictures.

Geographical patterns of terrestrial herbs: a new component in planning the conservation of the Brazilian Atlantic Forest

The Brazilian Atlantic Forest (sensu lato) is constituted by Dense Ombrophilous Forest, Mixed Ombrophilous Forest and Seasonal Semideciduous Forest, in which tree species are distributed in gradients, whichever latitude, longitude, or altitude is considered. We investigated whether herb species living on the floor of these forests would have the same geographic distribution pattern. If tree and herb-layer species have coincident distribution patterns, the same conservation units can conserve both species sets; otherwise, specific conservation actions should be taken. We built a presence/absence matrix of 803 terrestrial native species in 80 surveys across the southern and southeastern Brazilian Atlantic Forest. We performed UPGMA, CA and RDA multivariate analyses in order to investigate the distribution patterns. The high degree of agreement among the analyses led us to assume that the geographic distribution of the herb-layer flora occurs as gradients among three main floristic groups: Tropical Ombrophilous forests (mostly lowland forests), Subtropical Ombrophilous forests (mostly montane and upper-montane forests in lower latitudes and sub-montane and lowland forests in higher latitudes, i.e., where temperature is lower), and Seasonal Semideciduous forests (with a marked dry season). Based on different patterns and processes between herbs and trees, and in order to encompass the highest flora diversity, we highlight the importance of creating more reserves in each one of the three main floristic groups of herb-layer species in the Brazilian Atlantic Forest.

Scale-dependence of processes structuring dung beetle metacommunities using functional diversity and community deconstruction approaches.

Community structure is driven by mechanisms linked to environmental, spatial and temporal processes, which have been successfully addressed using metacommunity framework. The relative importance of processes shaping community structure can be identified using several different approaches. Two approaches that are increasingly being used are functional diversity and community deconstruction. Functional diversity is measured using various indices that incorporate distinct community attributes. Community deconstruction is a way to disentangle species responses to ecological processes by grouping species with similar traits. We used these two approaches to determine whether they are improvements over traditional measures (e.g., species composition, abundance, biomass) for identification of the main processes driving dung beetle (Scarabaeinae) community structure in a fragmented mainland-island landscape in southern Brazilian Atlantic Forest. We sampled five sites in each of four large forest areas, two on the mainland and two on the island. Sampling was performed in 2012 and 2013. We collected abundance and biomass data from 100 sampling points distributed over 20 sampling sites. We studied environmental, spatial and temporal effects on dung beetle community across three spatial scales, i.e., between sites, between areas and mainland-island. The γ-diversity based on species abundance was mainly attributed to β-diversity as a consequence of the increase in mean α- and β-diversity between areas. Variation partitioning on abundance, biomass and functional diversity showed scale-dependence of processes structuring dung beetle metacommunities. We identified two major groups of responses among 17 functional groups. In general, environmental filters were important at both local and regional scales. Spatial factors were important at the intermediate scale. Our study supports the notion of scale-dependence of environmental, spatial and temporal processes in the distribution and functional organization of Scarabaeinae beetles. We conclude that functional diversity may be used as a complementary approach to traditional measures, and that community deconstruction allows sufficient disentangling of responses of different trait-based groups.

Sensitivity to fragmentation and spatial distribution of birds in forest fragments of northern paraná

In this study we investigated the relationship between the sensitivity to forest fragmentation of bird species and their habitat spatial distribution. We hypothesized that a homogenous spatial distribution is associated to habitat-generalist species, which would tend to be less sensitive to forest fragmentation; in contrast, a heterogeneous spatial distribution would drive to an increase of sensitivity to forest fragmentation. We concentrated on analyses of the suboscine birds, which are known to be closely associated to microhabitats. Field data was obtained in the Parque Estadual Mata dos Godoy (23°27′ S, 51°15′ W), southern Brazilian Atlantic forest. This reserve has high Biotic Integrity levels, with a relatively high number of recorded suboscines. Forty three species of suboscines were recorded, of which 21 species were considered to have high sensitivity and 22 to have low sensitivity to forest fragmentation. There was no association between levels of sensitivity to forest fragmentation and spatial distribution of suboscines. We only found significant correlation with the group of the subcanopy suboscines at the northern portion of Parque Estadual Mata dos Godoy. Therefore, we found a weak relationship between spatial distribution (habitat specialization) and sensitivity to forest fragmentation. The fact that habitat-specialist birds tend to live at the northern portion of Godoy State Park, which has the first type of terrain in a landscape to be converted into agriculture, indicate that that region has special importance for conservation.

Phylobetadiversity among forest types in the Brazilian Atlantic Forest complex.

Phylobetadiversity is defined as the phylogenetic resemblance between communities or biomes. Analyzing phylobetadiversity patterns among different vegetation physiognomies within a single biome is crucial to understand the historical affinities between them. Based on the widely accepted idea that different forest physiognomies within the Southern Brazilian Atlantic Forest constitute different facies of a single biome, we hypothesize that more recent phylogenetic nodes should drive phylobetadiversity gradients between the different forest types within the Atlantic Forest, as the phylogenetic divergence among those forest types is biogeographically recent. We compiled information from 206 checklists describing the occurrence of shrub/tree species across three different forest physiognomies within the Southern Brazilian Atlantic Forest (Dense, Mixed and Seasonal forests). We analyzed intra-site phylogenetic structure (phylogenetic diversity, net relatedness index and nearest taxon index) and phylobetadiversity between plots located at different forest types, using five different methods differing in sensitivity to either basal or terminal nodes (phylogenetic fuzzy weighting, COMDIST, COMDISTNT, UniFrac and Rao’s H). Mixed forests showed higher phylogenetic diversity and overdispersion than the other forest types. Furthermore, all forest types differed from each other in relation phylobetadiversity patterns, particularly when phylobetadiversity methods more sensitive to terminal nodes were employed. Mixed forests tended to show higher phylogenetic differentiation to Dense and Seasonal forests than these latter from each other. The higher phylogenetic diversity and phylobetadiversity levels found in Mixed forests when compared to the others likely result from the biogeographical origin of several taxa occurring in these forests. On one hand, Mixed forests shelter several temperate taxa, like the conifers Araucaria and Podocarpus. On the other hand, tropical groups, like Myrtaceae, are also very representative of this forest type. We point out to the need of more attention to Mixed forests as a conservation target within the Brazilian Atlantic Forest given their high phylogenetic uniqueness.

Seasonal changes in dominant bacterial taxa from acidic peatlands of the Atlantic Rain Forest

The acidic peatlands of southern Brazil are essential for maintenance of the Atlantic Rain Forest, one of the 25 hot-spots of biodiversity in the world. While these ecosystems are closely linked to conservation issues, their microbial community ecology and composition remain unknown. In this work, histosol samples were collected from three acidic peatland regions during dry and rainy seasons and their chemical and microbial characteristics were evaluated. Culturing and culture-independent approaches based on SSU rRNA gene pyrosequencing were used to survey the bacterial community and to identify environmental factors affecting the biodiversity and microbial metabolic potential of the Brazilian peatlands. All acidic peatlands were dominated by the Acidobacteria phylum (56-22%) followed by Proteobacteria (28-12%). The OTU richness of these phyla and the abundance of their Gp1, Gp2, Gp3, Gp13, Rhodospirillales and Caulobacteriales members varied according to the period of collection and significantly correlated with the rainy season. However, despite changes in acidobacterial and proteobacterial communities, rainfall did not affect the microbial metabolic potential of the southern Brazilian Atlantic Rain Forest peatlands, as judged by the metabolic capabilities of the microbial community.

The use of highlands by the Lowland Tapir (Tapirus terrestris) in the southern Brazilian Atlantic Forest

The Lowland Tapir (Tapirus terrestris), out of the four living tapir species, has the broadest distribution. Although it naturally occurs in a variety of habitats in the Neotropics, including rainforest lowlands, gallery forests, dry chaco forests, and open grassy habitats it is currently classified as vulnerable to extinction. The Lowland Tapir is usually found near water, but there have been a few occurrence records in highland habitats. Here we report on seven records of the Lowland Tapir using highland habitats in four localities in the state of Paraná, southern Brazil. Records included tracks and feces in cloud forests (1,150-1,800 m a.s.l.) and tracks in grasslands (“campos de altitude”; 1,515-1,760 m a.s.l.). Six records were obtained between spring and summer, and one during winter. Although the use of high altitude locations by the Lowland Tapir offers an additional area for population maintenance, the occurrence of the species in these areas could have been driven by increased anthropic pressures in the Atlantic Forest lowlands.Key words: Campos de altitude, cloud forest, conservation, Serra do Mar.

Prokaryotic communities of acidic peatlands from the southern Brazilian Atlantic Forest

The acidic peatlands of southern Brazil are ecosystems essential for the maintenance of the Atlantic Forest, one of the 25 hot-spots of biodiversity in the world. In this work, we investigated the composition of prokaryotic communities in four histosols of three acidic peatland regions by constructing small-subunit (SSU) rRNA gene libraries and sequencing. SSU rRNA gene sequence analysis showed the prevalence of Acidobacteria (38.8%) and Proteobacteria (27.4%) of the Bacteria domain and Miscellaneous (58%) and Terrestrial (24%) groups of Crenarchaeota of the Archaea domain. As observed in other ecosystems, archaeal communities showed lower richness than bacterial communities. We also found a limited number of Euryarchaeota and of known methanotrophic bacteria in the clone libraries.

Mata Atlântica enchytraeids (Paraná, Brazil): A new genus, Xetadrilus gen. nov., with three new species, and four new species of Guaranidrilus Černosvitov (Enchytraeidae, Oligochaeta)

Seven new species of terrestrial Enchytraeidae (Oligochaeta) are described from soils of the southern Brazilian Atlantic rain forest. They were found in the framework of the German-Brazilian project SOLOBIOMA, which studied rain forest recovery. Specimens were investigated in vivo and as stained whole mounts. Three species belong to a new genus, named Xetadrilus. Xetadrilus is similar to Guaranidrilus Č ernosvitov, 1937 and Tupidrilus Righi, 1974; it differs in the absence of lateral chaetae from segment VIII on and in peculiar structures of the prostomium: ganglia, inner papillae, and a frontal epithelial recess. Further peculiarities are small body size, absence of oesophageal appendages, and a variable pattern of pharyngeal glands. The three species are named X. maacki, X. aphanus, and X. fabryi. Three further nominal species are transferred to Xetadrilus: Marionina pituca Righi, 1974, Marionina righiana Xie & Rota, 2001, and Stercutus ugandensis Bell, 1954. The other four new species belong to Guaranidrilus, the species-richest genus at the sampling sites. They are named G. andreolii, G. marquesi, G. cingulatus, and G. hoeferi. All species have elongate transverse epidermal gland cells, three pairs of post-pharyngeal ganglia, and a dorsal vesicle in the nephridial postseptale. These traits may be included in the genus diagnosis. In all species described here, most of the species-specific taxonomic traits are found in non-reproductive structures, which means that specimens of all age groups can be identified to the species level, juveniles included.

Lysimeter soil water balance evaluation for an experiment developed in the Southern Brazilian Atlantic Forest region

AbstractThis study aims at monitoring the behaviour of the rainfall, runoff, drainage, soil water storage, and evapotranspiration variables involved in the water balance measured by lysimeter data. The evaluation of the water balance considered different time scales, where the components were monitored daily and in 10‐day accumulated period intervals. The results demonstrated that in wet periods the soil water content was greater at a depth of 10 cm, whereas in the dry periods a greater concentration was observed at 70 cm depth. At the depth of 30 cm, the lowest values of soil water content were observed for both wet and dry periods. The results, obtained through the use of tensiometers and time domain reflectometry installed internally and externally to the lysimeter, were very close, which was more noticeable during the periods of lower water loss by the soil. The water balance, calculated from the lysimeter data, demonstrated that 70% of the total rainfall was lost by the process of evapotranspiration. The drainage accounted for 27·5% of the precipitated water, highlighting the fact that this component should not be disregarded in the water balance calculation. Copyright © 2011 John Wiley & Sons, Ltd.