Secondary Succession and Indigenous Management in Semideciduous Forest Fallows of the Amazon Basin1

To the discussion on secondary succession in tropical forests, we bring data on three under‐addressed issues: understory as well as overstory changes, continuous as opposed to phase changes, and integration of forest succession with indigenous fallow management and plant uses. Changes in vegetation structure and species composition were analyzed in secondary forests following swidden agriculture in a semideciduous forest of Bolivian lowlands. Twenty‐eight fallows, stratified by four successional stages (early = 1–5 yr, intermediate = 6–10 yr, advanced = 12–20 yr, and older = 22–36 yr), and ten stands of mature forests were sampled. The overstory (plants ≥5 cm diameter at breast height [DBH]) was sampled using a 20 × 50 m plot and the understory (plants <5 cm DBH) in three nested 2 × 5 m subplots. Semistructured interviews provided information on fallow management. Canopy height, basal area, and liana density of the overstory increased with secondary forest age. The early stage had the lowest species density and diversity in the overstory, but the highest diversity in the understory. Species composition and abundance differentiated mature forests and early successional stage from other successional stages; however, species showed individualistic responses across the temporal gradient. A total of 123 of 280 species were useful with edible, medicinal, and construction plants being the most abundant for both over‐ and understories. Most of Los Gwarayo preferred mature forests for making new swidden, while fallows were valuable for crops, useful species, and regenerating timber species.     

贵州省普定县不同植被演替阶段的物种组成与群落结构特征

通过对贵州省普定县喀斯特地区不同植被演替阶段群落的调查, 研究了植被演替过程中群落物种组成和群落结构的变化。结果表明, 该地区的植被主要处于5个演替阶段, 即次生乔木林、乔灌过渡林、藤刺灌丛、稀灌草丛以及火烧干扰后的蕨类植物群落。本次调查共记录到植物365种, 隶属89科218属。其中, 蕨类植物31种, 隶属14科23属; 种子植物334种, 隶属75科195属。物种分布较多的科主要有蔷薇科、菊科、禾本科、百合科、忍冬科、唇形科、莎草科、樟科、葡萄科和水龙骨科。随着正向演替的推进, 物种丰富度增加, 群落结构趋于复杂化。藤刺灌丛与乔灌过渡林群落层次不明显, 次生乔木林分层明显。从藤刺灌丛向次生乔木林演替的过程中, 小径级个体所占比例明显降低, 高于1.3 m植物的总密度、乔木密度和藤本密度都先升高后降低, 而灌木密度呈逐渐降低的趋势。对喀斯特地区植被的恢复提出了参考措施。     

Do lianas shape ant communities in an early successional tropical forest?

Almost half of lowland tropical forests are at various stages of regeneration following deforestation or fragmentation. Changes in tree communities along successional gradients have predictable bottom‐up effects on consumers. Liana (woody vine) assemblages also change with succession, but their effects on animal succession remain unexplored. Here we used a large‐scale liana removal experiment across a forest successional chronosequence (7–31 years) to determine the importance of lianas to ant community structure. We conducted 1,088 surveys of ants foraging on and living in trees using tree trunk baiting and hand‐collecting techniques at 34 paired forest plots, half of which had all lianas removed. Ant species composition, β‐diversity, and species richness were not affected by liana removal; however, ant species co‐occurrence (the coexistence of two or more species in a single tree) was more frequent in control plots, where lianas were present, versus removal plots. Forest stand age had a larger effect on ant community structure than the presence of lianas. Mean ant species richness in a forest plot increased by ca. 10% with increasing forest age across the 31‐year chronosequence. Ant surveys from forest >20 years old included more canopy specialists and fewer ground‐nesting ant species versus those from forests <20 years old. Consequently, lianas had a minimal effect on arboreal ant communities in this early successional forest, where rapidly changing tree community structure was more important to ant species richness and composition.     

The impact of logging on the abundance, species richness and community composition of butterfly guilds in Borneo

Abstract:  The impact of logging on the relative abundance, species richness and community composition of four butterfly guilds (generalists, herb specialists, liana specialists and tree specialists) was assessed in differentially disturbed logging treatments located within the province of Central Kalimantan, Indonesian Borneo. Logging affected the overall butterfly composition by increasing the dominance of liana specialists at the expense of tree specialists and increasing the species richness of herb specialists and, to a lesser extent, generalists. There was no significant difference, however, in the species richness of liana specialists or tree specialists across logging treatments. The species composition of all guilds differed significantly among logging treatments with the greatest difference between unlogged forest and road sites with older and recently logged forest intermediate between these extremes. Our results indicate that logging has a disparate effect on abundance and species richness within each butterfly guild. We suggest that shifts in the dominance of butterfly guilds are due to large-scale logging-induced changes in vegetation, particularly in relation to liana abundance. The higher butterfly species richness in logged areas on the other hand is probably because of butterfly species associated with open areas entering the logged forest from roads or other large open areas.     

Phyllostomid bat assemblages in different successional stages of tropical rain forest in Chiapas,Mexico

Due to their role in seed dispersal, changes in the community of phyllostomid bats have direct consequences on ecological succession. The objective of this work was to document changes in the structure of bat assemblages among secondary successional stages of tropical rain forest in Chiapas, Mexico. Bats were mist-netted at ground level during 18 months in 10 sites belonging to 3 successional stages: four sites represented early succession (2–8 years of abandonment), four intermediate succession (10–20 years of abandonment), and two late succession (mature old-growth forest).We captured 1,179 phyllostomids comprising 29 species. Phyllostomid species richness was 17 (58% of all species) in the early stage, 18 (62%) in the intermediate stage and 24 (83%) in the late stage. The late successional mature forest possessed nine species that were exclusively found there, whereas early and intermediate successional stages contained only one exclusive species. Sturnira lilium, Artibeus lituratus, Carollia perpicillata, Artibeus jamaicensis and Glossophaga soricina represented 88% of all captured phyllostomid bats. Frugivores made up more than 90% of the species captured in early and intermediate successional stages and 84% in late successional forest. The Bray–Curtis index of dissimilarity showed a replacement of species through successional stages with the largest dissimilarity between early and late stages, followed by intermediate and late, and the lowest dissimilarity between early and intermediate stages. The number of gleaning insectivore species increased during succession. The carnivorous guild was exclusively found in the late stage (three species). We conclude that the late successional mature forest was the main reservoir for the gleaning insectivore and carnivore guilds; however, early and intermediate successional stages possessed a great diversity of species including many frugivores.     

Stability in ecosystem functioning across a climatic threshold and contrasting forest regimes

Classical ecological theory predicts that changes in the availability of essential resources such as nitrogen should lead to changes in plant community composition due to differences in species-specific nutrient requirements. What remains unknown, however, is the extent to which climate change will alter the relationship between plant communities and the nitrogen cycle. During intervals of climate change, do changes in nitrogen cycling lead to vegetation change or do changes in community composition alter the nitrogen dynamics? We used long-term ecological data to determine the role of nitrogen availability in changes of forest species composition under a rapidly changing climate during the early Holocene (16k to 8k cal. yrs. BP). A statistical computational analysis of ecological data spanning 8,000 years showed that secondary succession from a coniferous to deciduous forest occurred independently of changes in the nitrogen cycle. As oak replaced pine under a warming climate, nitrogen cycling rates increased. Interestingly, the mechanism by which the species interacted with nitrogen remained stable across this threshold change in climate and in the dominant tree species. This suggests that changes in tree population density over successional time scales are not driven by nitrogen availability. Thus, current models of forest succession that incorporate the effects of available nitrogen may be over-estimating tree population responses to changes in this resource, which may result in biased predictions of future forest dynamics under climate warming.     

Soil phosphorus and disturbance influence liana communities in a subtropical montane forest

Questions: What are the effects of soil, topography, treefall gaps, tree species composition, and tree density on liana species composition and total liana abundance? Location: A 6‐ha permanent plot in a subtropical montane forest in northwest Argentina. Methods: Multiple regressions were used to quantify associations of liana species composition and total liana abundance with edaphic, disturbance and tree community variables. Gradients in liana and tree species composition were quantified using principal components analysis (PCA). Results: Liana species composition was correlated most strongly with soil phosphorus concentration (R²=0.55). Total liana aanased with phosphorus and the density of recent treefall gaps (R²=0.60). Conclusions: In our study area, liana composition and abundance are most strongly correlated with features of the physical environment, rather than host tree characteristics. Our results support the hypothesis that recent increases in liana abundance in mature tropical forests may be related to increased rates of gap formation.     

Herbivory on Handroanthus ochraceus (Bignoniaceae) along a successional gradient in a tropical dry forest

This study determined the temporal patterns of herbivory on Handroanthus ochraceus (Cham.) Mattos (Bignoniaceae) along a successional gradient in a seasonally dry tropical forest (SDTF) in southeastern Brazil. We assessed the diversity of free-feeding herbivore insects (sap-suckers and leaf-chewers), leaf herbivory rates, leaf nitrogen content, phenolic compounds, and spider abundance through the rainy season in three different successional stages (early, intermediate, and late). Sampling was conducted in December, at the beginning of the rainy season (with fully expanded young leaves), February (mid-aged leaves), and April, at the end of rainy season (old leaves). Fifteen reproductive trees of H. ochraceus were sampled per successional stage in each month of sampling. Herbivore diversity was highest in the early stage of succession, but herbivory rates were highest in the intermediate and late stages. This result was probably related to differences in herbivore community composition and leaf quality across successional stages. The highest herbivore abundance was found in April in the early successional stage. In addition, we found low levels of herbivory in the intermediate and late successional stages in the second half of the rainy season. For each successional stage, leaf nitrogen content decreased through the rainy season, whereas the concentration of phenolic compounds increased. For the intermediate and late successional stages, the temporal changes that took place as the rainy season progressed corroborated the following hypotheses postulated for SDTFs: (1) both the abundance of chewing insects and herbivory rates decreased, (2) the abundance of natural enemies (i.e., spiders) increased, and (3) leaf quality decreased. These results suggest that the described herbivory patterns are robust for advanced successional stages (intermediate and late) of the SDTFs, but may not apply to early successional stages of these forests.     

Nitrogen Uptake by Trees and Mycorrhizal Fungi in a Successional Northern Temperate Forest: Insights from Multiple Isotopic Methods

Forest succession may cause changes in nitrogen (N) availability, vegetation and fungal community composition that affect N uptake by trees and their mycorrhizal symbionts. Understanding how these changes affect the functioning of the mycorrhizal symbiosis is of interest to ecosystem ecology because of the fundamental roles mycorrhizae play in providing nutrition to trees and structuring forest ecosystems. We investigated changes in tree and mycorrhizal fungal community composition, the availability and uptake of N by trees and mycorrhizal fungi in a forest undergoing a successional transition (age-related loss of early successional tree taxa). In this system, 82–96% of mycorrhizal hyphae were ectomycorrhizal (EM). As biomass production of arbuscular mycorrhizal (AM) trees increased, AM hyphae comprised a significantly greater proportion of total fungal hyphae, and the EM contribution to the N requirement of EM-associated tree taxa declined from greater than 75% to less than 60%. Increasing N availability was associated with lower EM hyphal foraging and ¹⁵N tracer uptake, yet the EM-associated later-successional species Quercus rubra was nonetheless a stronger competitor for ¹⁵N than AM-associated Acer rubrum, likely due to the more extensive nature of the persistent EM hyphal network. These results indicate that successional increases in N availability and co-dominance by AM-associated trees have increased the importance of AM fungi in the mycorrhizal community, while down-regulating EM N acquisition and transfer processes. This work advances understanding of linkages between tree and fungal community composition, and indicates that successional changes in N availability may affect competition between tree taxa with divergent resource acquisition strategies.     

Structural changes of vegetation and its association with microclimate in a successional gradient of low thorn forest in northeastern Mexico

Understanding how microclimate and vegetation are associated during secondary succession is of primary importance for plant conservation in the face of the increasing land cover modification. However, these patterns are still unstudied for many plant communities. This study aimed to evaluate the structure (species richness, Shannon's diversity index, Simpson´s dominance index, abundance of each species, average height of species, species cover (%), species composition, and indicator values) of a low thorn forest fragment and to analyze its relation with microclimate along a successional gradient. Four stages of succession were delimited by the analysis of Landsat images, in the state of Tamaulipas, northeast Mexico. Statistical models incorporated species richness, diversity indices, abundance, height, and cover, as variables for searching differences between stages, or to evaluate microclimate associations. A total of 70 species, 54 genera, and 27 families were determined. Height of tree layer was the most important variable for discrimination of the successional stages. Conserved areas differed floristically from other stages, associated mainly with the lowest values of wind speed originated by tree layer characteristics. A significant association between species and microclimate was found, being wind speed and relative humidity the most important variables. Some species, due to their high importance values and their patterns of association with microclimate, may be considered as key taxa for low thorn forest, which is a threatened semitropical community in northeast Mexico. Conserved and late successional areas account for climatic regulation of this plant community, and the importance of these forest patches may be considered when establishing biodiversity protection areas.

     

岷江冷杉林皆伐后次生群落结构和物种多样性的演替动态

为阐明岷江冷杉林皆伐后次生群落的演替过程,采用空间代替时间的方法,在川西米亚罗林区海拔3100—3600 m的阴坡选择岷江冷杉林皆伐后次生演替10、20、30、40和50a阶段的次生群落作为研究对象,对其群落结构和物种多样性的动态进行了研究。不同演替阶段的树木均呈显著聚集分布。按群落中优势种的重要值将该演替序列划分为3个类型:悬钩子-蔷薇灌丛、白桦阔叶林和桦木-岷江冷杉针阔混交林。随次生演替,乔木和灌木种的物种丰富度趋于增加,而草本种的物种丰富度趋于减少;乔木和灌木种的Shannon-Wiener多样性指数趋于增大,而草本的Shannon-Wiener多样性指数趋于减小;乔木、灌木和草本层的Pielou均匀度指数均趋于增大;乔木层的Simpson优势度指数趋于减小,灌木和草本层的Simpson优势度指数在演替0—40年阶段趋于增大,而在演替50a阶段趋于减小。在该演替序列中,乔木、灌木和草本层的物种组成均呈耐荫种替代非耐荫种的趋势。     

Earthworms, stand age, and species composition interact to influence particulate organic matter chemistry during forest succession

The landscapes colonized by invasive earthworms in the eastern U.S. are often patchworks of forest stands in various stages of successional development. We established six field sites in tulip poplar dominated forests in the Smithsonian Environmental Research Center in Edgewater, MD, that span mid (50–70 years-three plots) and late (120–150 years-three plots) successional stages where younger sites had greater earthworm density and biomass than older sites and were dominated by non-native lumbricid species. In particular Lumbricus rubellus, a litter-feeding species, was abundant in mid successional forests. Here, we separated particulate organic matter (POM) from the bulk soil by a combination of size and density fractionation and found that patterns in soil POM chemistry were similar to those found previously during litter decay: in younger forests with high abundance of earthworms, organic carbon normalized cutin- and suberin-derived substituted fatty acid (SFA) concentration was lower and lignin-derived phenols greater than in older forests where earthworms were less abundant. The chemistry of the dominant litter from mid versus late successional tree species did not fully explain the differences in POM chemistry between age classes. Instead, the differences in leaf body versus petiole and leaf versus root chemistry were the dominant drivers of POM chemistry in mid versus late successional stands, although aspects of stand age and tree species also impacted POM chemistry. Our results indicate that preferential ingestion of leaf body tissue by earthworms and the subsequent shifts in sources of plant biopolymers in soil influenced POM chemistry in mid successional forests. These results indicate that invasive earthworm activity in North American forests contributes to a shift in the aromatic and aliphatic composition of POM and thus potentially influences carbon stabilization in soil.     

Impact of a hurricane on the herpetofaunal assemblages of a successional chronosequence in a tropical dry forest

Land‐use change is the main cause of deforestation and degradation of tropical forest in Mexico. Frequently, these lands are abandoned leading to a mosaic of natural vegetation in secondary succession. Further degradation of the natural vegetation in these lands could be exacerbated by stochastic catastrophic events such as hurricanes. Information on the impact of human disturbance parallel to natural disturbance has not yet been evaluated for faunal assemblages in tropical dry forests. To evaluate the response of herpetofaunal assemblages to the interaction of human and natural disturbances, we used information of pre‐ and post‐hurricane herpetofaunal assemblages inhabiting different successional stages (pasture, early forest, young forest, intermediate forest, and old growth forest) of dry forest. Herpetofaunal assemblages were surveyed in all successional stages two years before and two years after the hurricane Jova that hit the Pacific Coast of Mexico on October 2011. We registered 4093 individuals of 61 species. Overall, there were only slight effects of successional stage, hurricane Jova or the interaction between them on abundance, observed species richness and diversity of the herpetofauna. However, we found marked changes in estimated richness and composition of frogs, lizards, and snakes among successional stages in response to hurricane Jova. Modifications in vegetation structure as result of hurricane pass promoted particular changes in each successional stage and taxonomic group (anurans, lizards, and snakes). Secondary forests at different stages of succession may attenuate the negative effects of an intense, short‐duration, and low‐frequency natural disturbance such as hurricane Jova on successional herpetofaunal trajectories and species turnover.     

Conditional allelopathic potential of temperate lianas

Allelopathy is often treated as an innate characteristic of a species rather than a phenotypically plastic trait that can vary with environmental conditions. Lianas are a highly competitive, phenotypically plastic life form that typically occur in both shaded and unshaded environments. As such, we hypothesized that temperate lianas may conditionally change allocation to allelopathic chemicals in response to light availability though the expected direction of change is unclear. Shading may reduce resource availability and therefore reduce allocation to allelochemicals, induce allelopathy as a competitive mechanism, or may not be related to allelopathy. To test the conditionality of allelopathy, sun and shade leaves of five common liana species (Toxicodendron radicans, Parthenocissus quinquefolia, Celastrus orbiculatus, Lonicera japonica, and Vitis vulpina) were collected from a young deciduous forest in New Jersey, USA, and tested with laboratory bioassays to detect allelopathic potential. All liana species showed allelopathic potential, and three species exhibited induction of increased allelopathic potential in shaded environments. The two species that were not shade induced are late successional lianas that persist for long periods in forest canopies. In contrast, the inducible lianas were early successional species that typically decline with canopy closure. This research indicates that lianas have the potential to be allelopathic and allelopathic potential conditionally responds to shading only for species that would normally be excluded from the forest canopy. As early successional lianas are present throughout forest regeneration in a range of light environments, allelopathic plasticity may increase their success by differentially allocating resources based on environmental conditions.     

Determinants of litter decomposition rates in a tropical forest: functional traits,phylogeny and ecological succession

Plant litter decomposition is one of the most important processes in terrestrial ecosystems, as it is a key factor in nutrient cycling. Decomposition rates depend on environmental factors, but also plant traits, as these determine the character of detritus. We measured litter decomposition rate for 57 common tree species displaying a variety of functional traits within four sites in primary and four sites in secondary tropical forest in Madang Province, Papua New Guinea. The phylogenetic relationships between these trees were also estimated using molecular data. The leaves collected from different tree species were dried for two days, placed into detritus bags and exposed to ambient conditions for two months. Nitrogen, carbon and ash content were assessed as quantitative traits and used together with a phylogenetic variance– covariance matrix as predictors of decomposition rate. The analysis of the tree species composition from 96 quadrats located along a successional gradient of swidden agriculture enabled us to determine successional preferences for individual species. Nitrogen content was the only functional trait measured to be significantly positively correlated with decomposition rate. Controlling for plant phylogeny did not influence our conclusions, but including phylogeny demonstrated that the mainly early successional family Euphorbiaceae is characterized by a particularly high decomposition rate. The acquisitive traits (high nitrogen content and low wood density) correlated with rapid decomposition were characteristic for early successional species. Decomposition rate thus decreased from early successional to primary forest species. However, the decomposition of leaves from the same species was significantly faster in primary than in secondary forest stands, very probably because the high humidity of primary forest environments keeps the decomposing material wetter.     

Temporal variation in the influence of forest succession on caterpillar communities: A long‐term study in a tropical dry forest

Forest succession can influence herbivore communities through changes in host availability, plant quality, microclimate, canopy structure complexity and predator abundance. It is not well known, however, if such influence is constant across years. Caterpillars have been reported to be particularly susceptible to changes in plant community composition across forest succession, as most species are specialists and rely on the presence of their hosts. Nevertheless, in the case of tropical dry forests, plant species have less defined successional boundaries than tropical wet forests, and hence herbivore communities should be able to persist across different successional stages. To test this prediction, caterpillar communities were surveyed during eight consecutive years in a tropical dry forest in four replicated successional stages in Chamela, Jalisco and Mexico. Lepidopteran species richness and diversity were equivalent in mature forests and early successional stages, but a distinctive caterpillar community was found for the recently abandoned pastures. Species composition tended to converge among all four successional stages during the span of eight years. Overall, our results highlight the importance of both primary and secondary forest for the conservation of caterpillar biodiversity at a landscape level. We also highlight the relevance of long‐term studies when assessing the influence of forest succession to account for across year variation in species interactions and climatic factors. Abstract in French is available with online material.     

Alternative pathways of liana communities in the forests of northwestern Argentina

Liana dynamics in secondary and mature forests are well known in tropical areas dominated by native tree species. Outside the tropics and in secondary forests invaded by exotic species, knowledge is scarce. In this study, we compare liana communities between secondary and mature forests dominated by native species in a subtropical montane area of Sierra de San Javier, Tucuman, Argentina. Additionally, we evaluate changes of liana communities in secondary forests with increasing densities of Ligustrum lucidum and Morus alba, two of the most invasive exotic trees of the area. We surveyed liana species richness and density in three 30-year secondary patches, four 60-year secondary patches, and four mature patches dominated by native tree species, to analyze changes in liana communities with forest age. Within each patch, we sampled 10–25 20 × 20 m quadrats. Additionally, we surveyed liana density and species richness in secondary forest patches with different densities of L. lucidum and M. alba. In native-dominated forests, liana species richness increased and showed a tendency of increasing basal area from 30-year secondary forests to mature forests. Liana density was highly variable, and most of the species were shared between native-dominated secondary and mature forests. Liana density and species richness decreased with L. lucidum density, whereas in secondary forests highly dominated by M. alba, lianas increased in density. Overall, lianas followed different pathways influenced by native forest succession and exotic tree invasions.     

Changes in two Minnesota forests during 14 years following catastrophic windthrow

Abstract. We measured tree damage and mortality following a catastrophic windthrow in permanent plots in an oak forest and a pine forest in central Minnesota. We monitored changes in forest structure and composition over the next 14 years. Prior to the storm, the oak forest was dominated by Quercus ellipsoidalis, and the pine forest by Pinus strobus. The immediate impacts of the storm were to differentially damage and kill large, early‐successional hardwoods and pines. Subsequent recovery was characterized by the growth of late‐successional hardwoods. In both forests the disturbance acted to accelerate succession. Ordination of tree species composition confirmed the trend of accelerated succession, and suggested a convergence of composition between the two forests.     

Contrasting genetic diversity and differentiation of populations of two successional stages in a Neotropical pioneer tree (Eremanthus erythropappus, Asteraceae)

Eremanthus erythropappus, commonly known as "candeia", is an abundant pioneer tree species, forming dense populations known as "candeial", but it is also found in forests at middle stages of succession. Trees from forests are bigger and occur in lower density than in the "candeial". The objectives of the present study were to investigate if the decrease in population density during successional process is accompanied by 1) changes in within-population genetic diversity, and 2) differentiation of populations. Eight populations, four of early successional stage ("candeial") and four of middle successional stages (forest), were analyzed with RAPD markers. The genetic diversity found was high compared to other tree species analyzed with RAPD markers. AMOVA revealed that most of the genetic variations of E. erythropappus were found within populations (85.7%), suggesting that this species is predominantly outcrossing. The relatively low differentiation among the populations can be attributed to small distances among the populations analyzed (0.2 to 10.8 km). No indication that populations from middle successional habitats show lower genetic variation than populations from early successional stages was found. The percentage of polymorphic fragments (82.8 and 84.8%) and the Shannon indexes (0.442 and 0.455) were similar in "candeial" and forest, respectively. These results suggest that if an increase in selection intensity occurred during succession, it did not result in a decrease in genetic diversity or that the selection effect was balanced by other factors, such as gene flow. Higher significant differentiation among E. erythropappus populations from "candeial" in relation to that among populations from forest was also not detected.     

On the dynamics,floristic subdivision and geographical distribution of várzea forests in Central Amazonia

Abstract. Within different stands of the white-water inundation forest (várzea forest) in the Central Amazon region, composition, abundance, frequency and basal area of tree species were recorded. Determinations of age and radial growth rates were conducted using dendrochronological methods. Results show significant differences in age, history and species composition between stands as well as different growth strategies among dominant species. Assignment of tree species to growth strategies by means of anatomical and morphological features together with quantitative aspects of vegetational analysis permit the further differentiation of successional stages of várzea forests. General features of successional stages were quantitatively described and compared with forest types from outside the várzea. Many tree species of the várzea forests are widespread in South America, and not limited to floodplains. Their occurrence on sites with distinct dry seasons suggests that they are not specifically adapted to flooding but are tolerant to seasonality in general.