Phyllostomid Bat Assemblage Structure in Amazonian Flooded and Unflooded Forests

In Amazonia, the assemblages of several taxa differ significantly between upland terra firme and white‐water flooded várzea forests, but little is known about the diversity and distribution of bats in these two forest types. We compare the spatio‐temporal patterns of bat assemblage composition and structure in adjacent terra firme and várzea forests in the lower Purus River region of central Brazilian Amazonia. Bats were sampled using mist nets at five sites in each forest type during 40 nights (2400 net‐hours). We captured 1069 bats representing 42 species and Phyllostomidae bats comprised 99.3 percent of all captures. The bat assemblages in várzea and terra firme forests were significantly different, mainly due to a marked dissimilarity in species composition and in the number of captures during high‐water season. In addition, bat assemblages within forest types differed significantly between seasons for both terra firme and várzea. Frugivores dominated the bat assemblages in both forest types. Overall guild structure did not change between várzea and terra firme or between seasons, but frugivore and animalivore abundance increased significantly in várzea forest during the inundation. The difference in assemblage structure observed in the high‐water season is probably caused by the annual várzea flooding, which provides an effective barrier to the persistence of many understory bats. We also hypothesize that some bat species may undertake seasonal movements between forest types in response to fruit abundance, and our results further underline the importance of floodplain habitats for the conservation of species in the Amazon.     

Seasonal flooding decreases fruit-feeding butterfly species dominance and increases spatial turnover in floodplain forests of central Amazonia

The seasonal flood pulse in Amazonia can be considered a primary driver of community structure in floodplain environments. Although this natural periodic disturbance is part of the landscape dynamics, the seasonal inundation presents a considerable challenge to organisms that inhabit floodplain forests. The present study investigated the effect of seasonal flooding on fruit-feeding butterfly assemblages in different forest types and strata in central Amazonia. We sampled fruit-feeding butterflies in the canopy and the understory using baited traps in adjacent upland (unflooded forests—terra firme), white and blackwater floodplain forests (várzea and igapó, respectively) during the low- and high-water seasons. Butterfly abundance decreased in the high-water season, especially of dominant species in várzea, but the number of species was similar between seasons in the three forest types. Species composition differed between strata in all forest types. However, the flood pulse only affected butterfly assemblages in várzea forest. The β-diversity components also differed only in várzea. Species replacement (turnover) dominated the spatial β-diversity in igapó and terra firme in both seasons and várzea in the high-water season. Nonetheless, nestedness was relatively higher in várzea forests during the low-water season, mainly due to the effect of dominant species. These results emphasize the importance of seasonal flooding to structure butterfly assemblages in floodplain forests and reveal the idiosyncrasy of butterfly community responses to flooding in different forest types. Our results also suggest that any major and rapid changes to the hydrological regime could severely affect floodplain communities adapted to this natural seasonal hydrological cycle, threatening the existence of these unique environments.     

Tree species composition and diversity gradients in white-water forests across the Amazon Basin

Aim Attention has increasingly been focused on the floristic variation within forests of the Amazon Basin. Variations in species composition and diversity are poorly understood, especially in Amazonian floodplain forests. We investigated tree species composition, richness and α diversity in the Amazonian white‐water (várzea) forest, looking particularly at: (1) the flood‐level gradient, (2) the successional stage (stand age), and (3) the geographical location of the forests. Location Eastern Amazonia, central Amazonia, equatorial western Amazonia and the southern part of western Amazonia. Methods The data originate from 16 permanent várzea forest plots in the central and western Brazilian Amazon and in the northern Bolivian Amazon. In addition, revised species lists of 28 várzea forest inventories from across the Amazon Basin were used. Most important families and species were determined using importance values. Floristic similarity between plots was calculated to detect similarity variations between forest types and over geographical distances. To check for spatial diversity gradients, α diversity (Fisher) of the plots was correlated with stand age, longitudinal and latitudinal plot location, and flood‐level gradient. Results More than 900 flood‐tolerant tree species were recorded, which indicates that Amazonian várzea forests are the most species‐rich floodplain forests worldwide. The most important plant families recorded also dominate most Neotropical upland forests, and c. 31% of the tree species listed also occur in the uplands. Species distribution and diversity varied: (1) on the flood‐level gradient, with a distinct separation between low‐várzea forests and high‐várzea forests, (2) in relation to natural forest succession, with species‐poor forests in early stages of succession and species‐rich forests in later stages, and (3) as a function of geographical distance between sites, indicating an increasing α diversity from eastern to western Amazonia, and simultaneously from the southern part of western Amazonia to equatorial western Amazonia. Main conclusions The east‐to‐west gradient of increasing species diversity in várzea forests reflects the diversity patterns also described for Amazonian terra firme. Despite the fine‐scale geomorphological heterogeneity of the floodplains, and despite high disturbance of the different forest types by sedimentation and erosion, várzea forests are dominated by a high proportion of generalistic, widely distributed tree species. In contrast to high‐várzea forests, where floristic dissimilarity increases significantly with increasing distance between the sites, low‐várzea forests can exhibit high floristic similarity over large geographical distances. The high várzea may be an important transitional zone for lateral immigration of terra firme species to the floodplains, thus contributing to comparatively high species richness. However, long‐distance dispersal of many low‐várzea trees contributes to comparatively low species richness in highly flooded low várzea.     

Vertebrate responses to fruit production in Amazonian flooded and unflooded forests

We document patterns of fruit and vertebrate abundance within an extensive, virtually undisturbed mosaic of seasonally flooded (várzea and igapó) and unflooded (terra firme) forests of central Amazonia. Using phenological surveys and a standardised series of line-transect censuses we investigate the spatial and temporal patterns of immature and mature fruit availability and how this may affect patterns of habitat use by vertebrates in the landscape. All habitats showed marked peaks in fruiting activity, and vertebrate detection rates varied over time for most species both within and between forest types. Many arboreal and terrestrial vertebrates used both types of flooded forest on a seasonal basis, and fluctuations in the abundance of terrestrial species in várzea forest were correlated with fruit availability. Similarly, the abundance of arboreal seed predators such as buffy saki monkeys (Pithecia albicans) and macaws (Ara spp.) were closely linked with immature fruit availability in terra firme forest. We conclude that highly heterogeneous landscapes consisting of terra firme, várzea and igapó forest appear to play an important role in the dynamics of many vertebrate species in lowland Amazonia, but the extent to which different forest types are used is highly variable in both space and time.     

Do flood pulses structure amphibian communities in floodplain environments?

Beta diversity can provide insights into the processes that regulate communities subjected to frequent disturbances, such as flood pulses, which control biodiversity in floodplains. However, little is known about which processes structure beta diversity of amphibians in floodplains. Here, we tested the influence of flood pulses on the richness, composition, and beta diversity of amphibians in Amazonian floodplain environments. We also evaluated indicator species for each environment. We established linear transects in three environments: low várzea, high várzea, and macrophyte rafts. Species richness decreased and beta diversity increased according to the susceptibility of habitats to flood pulses. Indicator species differed among environments according to forest succession promoted by the flood pulse. The decrease in species richness between high and low várzea is due to non‐random extinctions. The higher rates of species turnover between várzeas and macrophyte rafts are driven by the colonization of species adapted to open areas. Our results highlight that the maintenance of complex environments is needed to protect biodiversity in floodplains.     

Spatial, Temporal, and Economic Constraints to the Commercial Extraction of a Non–timber Forest Product: Copaíba (Copaifera spp.) Oleoresin in Amazonian Reserves

Spatial, Temporal, and Economic Constraints to the Commercial Extraction of a Non?CTimber Forest Product: Copaíba (Copaifera spp.) Oleoresin in Amazonian Reserves. The increasing prevalence of government?C and NGO?Csponsored programs to encourage commercial non?Ctimber forest product (NTFP) extractivism in the humid tropics has highlighted the need for ecological and socioeconomic appraisal of the viability of extractive industries. We adopted a novel integrative approach to examine NTFP resource potential and produced credible landscape?Cscale estimates of the projected value of an economically important Amazonian NTFP, the medicinal oleoresin of Copaifera trees, within two large contiguous extractive reserves in Brazilian Amazonia. We integrated results derived from previous spatial ecology and harvesting studies with socioeconomic and market data, and mapped the distribution of communities within the reserves. We created anisotropic accessibility models that determined the spatial and temporal access to Copaifera trees in permanently unflooded (terra firme) and seasonally flooded (várzea) forests. Just 64.9?% of the total reserve area was accessible, emphasizing the distinction between the actual resource stock and that which is available to extractors. The density of productive tree species was higher in the várzea forests, but per?Ctree productivity was greater in the terra firme forests, resulting in similar estimates of oleoresin yield per unit area (64?C67?ml?ha^?C1) in both forest types. A greater area of the várzea forests was accessible within shorter travel times of ??250 minutes; longer travel times allowed access to increasingly greater volumes of oleoresin from the terra firme forests. The estimated total volume of oleoresin accessible within the two reserves was 38,635 liters for an initial harvest, with projected offtake for a subsequent harvest falling to 8,274 liters. A household that extracted just 2 liters of oleoresin per month could generate 5?% of its mean income; market data suggested that certification could increase the value of the resource fivefold. Our approach is valuable in that it incorporates a range of methodologies and quantitatively accounts for the numerous constraints to the commercial viability of NTFP extraction.     

Interspecific primate associations in Amazonian flooded and unflooded forests

Stable associations between two or more primate species are a prominent feature of neotropical forest vertebrate communities and many studies have addressed their prevalence, and their costs and benefits. However, little is known about the influence of different habitat types on the frequency, seasonality, and composition of mixed-species groups in Amazonian forest primates. Here we examine the features of interspecific primate groups in a large mosaic of flooded (várzea and igapó) and unflooded (terra firme) forest in central Amazonia. In total, 12 primate species occurred in the study area, nine of which were observed in mixed-species associations. Primates were more than twice as likely to form associations in várzea forest than in terra firme forest. Squirrel monkeys were most frequently found in mixed-species groups in all forest types, most commonly in association with brown capuchins. Another frequent member of interspecific associations was the buffy saki, which often formed mixed-species groups with tamarins or brown capuchins. There was no seasonality in the frequency of associations in terra firme forest whereas associations in várzea forest were twice as frequent during the late-dry and early-wet seasons than in the late-wet and early-dry seasons. Interspecific primate associations were common in all forest types, but the degrees to which different species associate varied between these environments. We suggest that the temporal variation of várzea forest associations is connected with seasonal changes in habitat structure and resource abundance. However, more work is needed to pinpoint the underlying causes of mixed-species associations in all forest types and their strong seasonality in várzea forest.     

Pollen and seed flow patterns of Carapa guianensis Aublet. (Meliaceae) in two types of Amazonian forest

Various factors affect spatial genetic structure in plant populations, including adult density and primary and secondary seed dispersal mechanisms. We evaluated pollen and seed dispersal distances and spatial genetic structure of Carapa guianensis Aublet. (Meliaceae) in occasionally inundated and terra firme forest environments that differed in tree densities and secondary seed dispersal agents. We used parentage analysis to obtain contemporary gene flow estimates and assessed the spatial genetic structure of adults and juveniles. Despite the higher density of adults (diameter at breast height ≥ 25 cm) and spatial aggregation in occasionally inundated forest, the average pollen dispersal distance was similar in both types of forest (195 ± 106 m in terra firme and 175 ± 87 m in occasionally inundated plots). Higher seed flow rates (36.7% of juveniles were from outside the plot) and distances (155 ± 84 m) were found in terra firme compared to the occasionally inundated plot (25.4% and 114 ± 69 m). There was a weak spatial genetic structure in juveniles and in terra firme adults. These results indicate that inundation may not have had a significant role in seed dispersal in the occasionally inundated plot, probably because of the higher levels of seedling mortality.     

Unexpected High Diversity of Galling Insects in the Amazonian Upper Canopy: The Savanna Out There

A relatively large number of studies reassert the strong relationship between galling insect diversity and extreme hydric and thermal status in some habitats, and an overall pattern of a greater number of galling species in the understory of scleromorphic vegetation. We compared galling insect diversity in the forest canopy and its relationship with tree richness among upland terra firme, várzea, and igapó floodplains in Amazonia, Brazil. The soils of these forest types have highly different hydric and nutritional status. Overall, we examined the upper layer of 1,091 tree crowns. Galling species richness and abundance were higher in terra firme forests compared to várzea and igapó forests. GLM-ANCOVA models revealed that the number of tree species sampled in each forest type was determinant in the gall-forming insect diversity. The ratio between galling insect richness and number of tree species sampled (GIR/TSS ratio) was higher in the terra firme forest and in seasonally flooded igapó, while the várzea presented the lowest GIR/TSS ratio. In this study, we recorded unprecedented values of galling species diversity and abundance per sampling point. The GIR/TSS ratio from várzea was approximately 2.5 times higher than the highest value of this ratio ever reported in the literature. Based on this fact, we ascertained that várzea and igapó floodplain forests (with lower GIA and GIR), together with the speciose terra firme galling community emerge as the gall diversity apex landscape among all biogeographic regions already investigated. Contrary to expectation, our results also support the “harsh environment hypothesis”, and unveil the Amazonian upper canopy as similar to Mediterranean vegetation habitats, hygrothermically stressed environments with leaf temperature at lethal limits and high levels of leaf sclerophylly.     

Tree Phenology in Adjacent Amazonian Flooded and Unflooded Forests1

Most phenological studies to date have taken place in upland forest above the maximum flood level of nearby streams and rivers. In this paper, we examine the phenological patterns of tree assemblages in a large Amazonian forest landscape, including both upland (terra firme) and seasonally flooded (várzea and igapó) forest. The abundance of vegetative and reproductive phenophases was very seasonal in all forests types. Both types of flooded forest were more deciduous than terra firme, shedding most of their leaves during the inundation period. Pulses of new leaves occurred mainly during the dry season in terra firme, whereas those in the two floodplain forests were largely restricted to the end of the inundation period. Flowering was concentrated in the dry season in all forest types and was strongly correlated with the decrease in rainfall. The two floodplain forests concentrated their fruiting peaks during the inundation period, whereas trees in terra firme tended to bear fruits at the onset of the wet season. The results suggest that the phenological patterns of all forest types are largely predictable and that the regular and prolonged seasonal flood pulse is a major determinant of phenological patterns in várzea and igapó, whereas rainfall and solar irradiance appear to be important in terra firme. The three forest types provide a mosaic of food resources that has important implications for the conservation and maintenance of wide‐ranging frugivore populations in Amazonian forests.     

The role of recruitment and dispersal limitation in tree community assembly in Amazonian forests

Background: Species composition of plant communities is shaped by the interplay between dispersal limitation, environmental filters and stochastic events.

Aims: The aim of this work was to investigate the effects of dispersal limitation and environmental filtering on tree recruitment. To accomplish this, we employed the unified neutral theory of biodiversity and biogeography to examine migration within the metacommunity, defined as a set of interacting local communities linked by the dispersal of multiple potentially interacting species.

Methods: We sampled 12,975 individuals with dbh ≥ 1 cm in 26 1-ha permanent plots, including habitats of terra firme, transitional forests, várzea and campinarana, on the upper Madeira River, Brazilian Amazon.

Results: Campinarana drew individuals from outside the metacommunity species pool at a mean probability of recruitment of 0.06, a much lower probability than terra firme (0.31), transitional (0.21) and várzea forests (0.22). Environmental variables, such as water table depth, soil texture and fertility, were related to differences in community assembly.

Conclusions: Species abundance distribution and diversity patterns of plant assemblages in a large river landscape in the Amazon highlight the importance of environmental heterogeneity that conditions beta-diversity. The high variation in recruitment probabilities from the metacommunity species pool to local communities suggests high habitat variability in the process of maintaining patterns of local diversity.     

Ecological Responses of Frugivorous Bats to Seasonal Fluctuation in Fruit Availability in Amazonian Forests

Lowland Amazon is climatically one of the least seasonal regions on the planet, but little is known about how this is reflected in ecological seasonality. The central objective of this study was to determine whether seasonal fluctuations in the availability of fruit resources in Neotropical forests are sufficiently marked to affect the ecology and physiology of frugivorous bats. Seasonal variations in the overall bat abundance and in the captures, body condition, and reproductive activity of the two most abundant species, Carollia perspicillata and Artibeus planirostris, were studied within a region of central Brazilian Amazonia dominated by a mosaic of nonflooded (terra firme) and seasonally flooded forests (várzea and igapó). Concurrent seasonal changes in fruit availability were measured. The abundance of fruits was markedly seasonal, with far fewer resources available during the low-water season. There was a positive correlation between fruit and bat abundance. Overall, bats did not increase the consumption of arthropods during the period of fruit shortage. In A. planirostris, the body condition declined when fruits were scarcer. In both C. perspicillata and A. planirostris, foraging and reproductive activity were positively correlated with fruit availability. Consequently, the results suggest that resource seasonality is sufficiently marked to affect frugivorous bats and force them to make important eco-physiological adjustments.     

Quantitative ecological inventory of terra firme and várzea tropical forest on the Rio Xingu,Brazilian Amazon

Three hectares of Amazonian terra firme forest and an adjacent one-half hectare of várzea forest were quantitatively inventoried at O Deserto, on the Rio Xingu, Pará, Brazil. In the terra firme forest, 1420 individual trees greater than ten cm dbh, in 39 families and 265 species, were inventoried. In the várzea forest, there were 220 individual trees, in 17 families and 40 species.Cenostigma macrophyllum andOrbignya sp. were the most important species in the terra firme forest;Mollia lepidota andLeonia glycycarpa were most important in the várzea forest. Among one-hectare subplots of the total three-hectare terra firme sample, the number of trees ranged from 393 to 460, the number of families was a constant 33, and the number of species ranged from 118 to 162. This variation indicates that one-hectare samples are too small to be used to estimate the species richness of the total forest. The terra firme forest was richer in species and had a greater stature than the várzea forest.     

Primate assemblage structure in Amazonian flooded and unflooded forests

There is considerable variation in primate species richness across neotropical forest sites, and the richest assemblages are found in western Amazonia. Forest type is an important determinant of the patterns of platyrrhine primate diversity, abundance, and biomass. Here we present data on the assemblage structure of primates in adjacent unflooded (terra firme) and seasonally inundated (várzea and igapó) forests in the lower Purús region of central-western Brazilian Amazonia. A line-transect census of 2,026 km in terra firme, 2,309 km in várzea, and 277 km in igapó was conducted. Twelve primate species were recorded from 2,059 primate group sightings. Although terra firme was found to be consistently more species-rich than várzea, the aggregate primate density in terra firme forest was considerably lower than that in the species-poor várzea. Consequently, the total biomass estimate was much higher in várzea compared to either terra firme or igapó forest. Brown capuchin monkeys (Cebus apella) were the most abundant species in terra firme, but were outnumbered by squirrel monkeys (Saimiri cf. ustus) in the várzea. The results suggest that floodplain forest is a crucial complement to terra firme in terms of primate conservation in Amazonian forests.     

The Amazon River as a dispersal barrier to passerine birds: effects of river width, habitat and taxonomy

Aim To evaluate the relative effectiveness of the lower and upper sections, respectively, of the Amazon River as a barrier to bird distribution, and to evaluate ecological and taxonomic factors affecting the efficacy of the river barrier. Location Amazon River of South America between its confluence with the Napo River in the west and its delta in the east. Methods Using published distribution maps for 448 species of passerine birds occurring along the Amazon River, we evaluated whether each was distributed along one bank only (river presumed to be a barrier) or both banks (no barrier) to test the predictions that the river was more effective as a dispersal barrier: (1) along the lower, wider portion of the river than the upper, narrower portion; (2) for species inhabiting forests than open country; (3) for species inhabiting forest understorey than forest canopy; (4) for species restricted to terra firme (never inundated upland forest) than those not restricted to terra firme and (5) for certain taxonomic groups. Results Our analyses demonstrated that the Amazon River was most effective as a dispersal barrier along its lower portion and for species restricted to forests and terra firme. However, the river was not significantly more of a barrier for species inhabiting forest understorey than forest canopy. The river was most significant as a barrier to dispersal for the antbirds (Thamnophilidae) and was less significant as a barrier to species belonging to several large families including woodcreepers (Dendrocolaptidae), ovenbirds (Furnariidae), flycatchers (Tyrannidae), cotingids (Cotingidae), tanagers (Thraupidae), seed‐eating finches (Emberizidae) and blackbirds (Icteridae). Main conclusions The robust widths of Amazonian rivers are widely considered to represent impediments to dispersal and gene flow for many taxa of birds and other animals, and may have represented agents of vicariance in the diversification of species. Our study reaffirms the effectiveness of the lower Amazon River as a current barrier to bird dispersal for forest birds and provides new insights into the effects of habitat and taxonomy on the efficacy of the river barrier. Although supportive of several predictions of the river hypothesis of biological diversification, our study is limited in addressing the historical impact of river barriers as agents of vicariance in the process of diversification.     

Nutrient transport within and between habitats through seed dispersal processes by woolly monkeys in north‐western Amazonia

The contribution of vertebrate animals to nutrient cycling has proven to be important in various ecosystems. However, the role of large bodied primates in nutrient transport in neotropical forests is not well documented. Here, we assess the role of a population of woolly monkeys (Lagothrix lagothricha lugens) as vectors of nutrient movement through seed dispersal. We estimated total seed biomass transported by the population within and between two habitats (terra firme and flooded forests) at Tinigua Park, Colombia, and quantified potassium (K), phosphorus (P) and nitrogen (N) content in seeds of 20 plant species from both forests. Overall, the population transported an estimated minimum of 11.5 (±1.2 SD) g of potassium, 13.2 (±0.7) g of phosphorus and 34.3 (±0.1) g nitrogen, within 22.4 (±2.0) kg of seeds ha^?1 y^?1. Approximately 84% of all nutrients were deposited in the terra firme forest mostly through recycling processes, and also through translocation from the flooded forest. This type of translocation represents an important and high‐quality route of transport since abiotic mechanisms do not usually move nutrients upwards, and since chemical tests show that seeds from flooded forests have comparatively higher nutrient contents. The overall contribution to nutrient movement by the population of woolly monkeys is significant because of the large amount of biomass transported, and the high phosphorus content of seeds. As a result, the phosphorus input generated by these monkeys is of the same order of magnitude as other abiotic mechanisms of nutrient transport such as atmospheric deposition and some weathering processes. Our results suggest that via seed dispersal processes, woolly monkey populations can contribute to nutrient movement in tropical forests, and may act as important nutrient input vectors in terra firme forests. Am. J. Primatol. 72:992–1003, 2010. © 2010 Wiley‐Liss, Inc.     

Combining phylogeography and landscape genetics of Xenopipo atronitens (Aves: Pipridae), a white sand campina specialist,to understand Pleistocene landscape evolution in Amazonia

Open vegetation (campinas and campinaranas) associated with white sand patches occurs in the form of islands in a forested matrix throughout the Amazon basin. Bird species restricted to these habitats have patchy distributions, although connectivity may have been influenced by past glacial cycles as a result of the substitution of forest by savanna. Because these landscape changes are a matter of debate in the history of Amazonia, we studied the diversification of Xenopipo atronitens, a white sand specialist, aiming to infer the effects of past climate changes. The split of Xenopipo atronitens from its sister species, Xenopipo uniformis, may be related to Tepuis erosion and retreat of escarpments during the Miocene, or to a dispersal event. Compared with birds from terra firme forest, X. atronitens has low genetic structure. Low levels of unidirectional gene flow were found from the Guyana Shield to adjacent areas. Demographic expansion starting approximately 25 kyr BP was detected for some populations and is probably related to the Last Glacial Maximum and subsequent climate improvement. Landscape genetic analyses indicate that the forested (terra firme) matrix acts as a barrier for the dispersal of X. atronitens. The results of the present study indicate that glacial cycles have deeply influenced Amazonian biogeographical history, demonstrating a complex interaction between forest and nonforest habitats during the Pleistocene. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 60–76.     

A comparison of the phytosociology and dynamics of three floodplain (Várzea) forests of known ages, Rio Jurua, western Brazilian Amazon

CAMPBELL, D. G., STONE, J. L. & ROSAS Jr, A., 1992. A comparison of the phytosociology and dynamics of three floodplain (Várzea) forests of known ages, Rio Juruá, western Brazilian Amazon. An unusual historic accident has resulted in three adjacent inundated (vórzea) forests of known ages (50 years old, 14–50 years old, and 14 years old), on the margin of the Rio Juruá, near the village of Rodrigues Alves in western Acre, Brazilian Amazon. One-hectare quantitative inventories (500 × 20 m) were conducted on each of the three forests. Physical characteristics of the plots, including soil particle size, soil cation levels, level of inundation and rate of soil deposition were measured. Plot 1, steeply sloping and ecotonal between várzea and terra firme, was infrequently flooded, with a mean high water mark of 0.0. Plot 2, on slightly undulating terrain, had a mean high water mark of 1.16 m. Plot 3 was level and deeply-flooded, with a mean high water mark of 4.00 m. All trees ≥ 10 cm diameter at breast height (dbh) were mapped, measured and identified to species. Plots 1, 2 and 3 contained 523, 420 and 777 trees, 106, 73 and 20 species, Shannon diversity indices of 3.60, 3.51 and 1.82, and stand basal areas of 25.46 m², 27.01 m² and 25.72 m², respectively. Alchomea triplinervia was the superior competitor on species-rich plot 1; Annona sp. strongly dominated in the limiting conditions of plot 3. On plot 2, no single species was able to achieve dominance. For the 25 species with N≥ 10, habitat specialization as a function of the level of annual inundation was demonstrated, and for five of these species that occurred on plot 1, further refinement of niche as a function of gradient was demonstrated. One-year mortality in plots 1, 2 and 3, was 1.99%, 1.56% and 1.70%, respectively. In Plot 3, 77% of the mortalities were trees that had died standing, vs 40% and 43% in plots 1 and 2.     

Sapling communities in Amazonian white-water forests

Aim Structure and floristic composition of forest regeneration (trees between 1 and 10 cm diameter at breast height (d.b.h), ≥ 1 m growth height) was described in three forest types of Amazonian white‐water forests (várzea), in order to analyse whether floristic composition of saplings is related to the successional stage of the forests, whether it differs in comparision with the mature flora, and if there exists a zonation of sapling species along the gradients of flooding and irradiation. Location Mamirauá Sustainable Development Reserve, Western Brazilian Amazon. Methods The investigated forests were of the low and the high várzea type, on an annual average flooded around 4 and 1 months, respectively. The two low‐várzea forests belonged to the secondary and to the late‐successional stage, the high‐várzea forest also to the late‐successional stage. A total of 24 circular sample plots covering 1885 m² were installed. They were nested within three rectangular 1 ha permanent sample plots where individuals ≥ 10 cm d.b.h. were formerly inventoried. Average inundation and radiation [relative photosynthetically active radiation (rPAR)] at the forest floor was recorded in all inventoried plots. The Mean spatial Distribution Center (MDC; Ebdon, 1998 ) for all sapling species was calculated. Sapling species were grouped into associations with respect to their light‐demand and their location along the flood‐level gradient. Results Average flood height and average rPAR at the forest floor was highest in the secondary stage of the low várzea and lowest in the high‐várzea forest. Overall average density and basal area of the saplings averaged 2250 individuals and 2.3 m² ha^?1 in the secondary stage, 2330 individuals and 2.6 m² ha^?1 in the late‐successional stage of the low várzea and 5000 individuals and 4.8 m² ha^?1 in the high várzea. In all forest types, species richness of saplings was lower than species richness of trees ≥ 10 cm d.b.h., and amounted to 25 species in the secondary stage, to 35 species in the late‐successional stage of the low várzea and to 88 species in the high‐várzea forest. The amount of ‘immigrants’ ( Bazzaz, 1991 ) at the sapling level increased with proceeding forest succession and amounted to 24 and 29% in the low várzea and to 31% in the high várzea. Sapling species distribution, species richness and individual density was linked to both, the gradient of flooding and the gradient of irradiation. Species richness and individual density was highest in the high várzea and decreased with increasing influence of flooding. The high várzea was predominated by pronounced shade‐tolerant sapling species, whereas in the low várzea occured both, light demanding and shade‐tolerant species groups. Main conclusions Influence of flooding seems to be the main factor triggering species composition and structure of sapling communities in várzea forests. Proceeding forest succession reduces the impact of flooding because of the biogenical induced silting up of the forested sites. However, forest succession also alters forest architecture of the overstory and such the light conditions at the sapling level. Therefore, radiation is an important factor influencing species composition of sapling communities in várzea forests.     

Flooding and soil composition determine beta diversity of lowland forests in Northern South America

Beta diversity may be determined by dispersal limitation, environment, and phylogeographic history. Our objective was to advance the understanding of plant species turnover in rain forests in northern South America and determine which factors are affecting species beta diversity. We evaluated the relative effect of environmental variables (i.e., soil, climate, fragmentation, and flooding frequency) and dispersal limitation (i.e., geographical distance and resistance distance due mountain barriers) on tree beta diversity in 32 1‐ha lowland forest plots. We found that tree species turnover was better explained by environmental distance than by geographical distance. Although soil conditions and flooding regime were good predictors of tree species composition, almost half of the variance remained unexplained. In our study system, the eastern Andean ridge had no significant effect on plant beta diversity, probably because of its young age in relation to the phylogeny. Our results provide support for the importance of environmental factors and suggest a more restricted role of dispersal limitation. Therefore, we advise that conservation strategies of lowland trees should consider specific forest types (e.g., seasonally flooded vs. terra firme, as well as piedmont vs. central Amazonian forests).