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.     

Habitat specifity,endemism and the neotropical distribution of Amazonian white‐water floodplain trees

.The Amazon basin is covered by the most species‐rich forests in the world and is considered to house many endemic tree species. Yet, most Amazonian ecosystems lack reliable estimates of their degree of endemism, and causes of tree diversity and endemism are intense matters of debate. We reviewed the spatial distribution of 658 of the most important flood‐tolerant Amazonian white‐water (várzea) tree species across the entire Neotropics by using data from herbaria, floras, inventories and checklists. Our results show that 90% of the várzea tree species are partially or widely distributed across neotropical macro‐regions and biomes. Chi‐square analyses indicated that várzea species richness in non‐várzea macro‐regions was dependent on the flooding gradient and the longitudinal position. Cluster analysis combined with association tests indicated four significant patterns of várzea species distributions depending on species flood‐tolerance (low vs high) and spatial distribution (restricted vs widespread). We predict that the predominance of Andean substrates is the most important factor that determines the distribution of várzea tree species within and beyond the Amazon basin and explains the high floristic similarity to the Orinoco floodplains. Distribution patterns in other extra‐Amazonian macro‐regions are more likely linked to climatic factors, with rainforest climates housing more várzea species than savanna climates. 130 tree species were restricted to South‐American freshwater floodplains, and 68 (> 10%) were endemic to Amazonian várzea. We detected two centers of endemism, one in the western Amazon characterized by low and brief floods, and one in the central Amazon, characterized by high and prolonged floods. Differences in taxonomic composition of endemic centers in the western and central Amazon are the result of different abiotic factors (i.e. flood regimes), as well as the regional species pools from where the species are recruited from. We hypothesize that numerous morphological, physiological and biochemical adaptations permit survival of trees in flooded environments. Furthermore, these adaptations are independently derived across many taxa and result in a highly specialized flora. We attribute higher than expected levels of endemism to the great spatial extent and age of floodplain ecosystems in the Amazon basin, and highlight the role of Amazonian várzea as an potential driver in speciation and diversification processes.     

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.     

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.     

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.     

Forest structure and tree species composition of the understory of two central Amazonian várzea forests of contrasting flood heights

We investigated species composition, distribution, and forest structure of understory trees (≥1 m height, <10 cm diameter at breast height) in two late-successional várzea forests subject to contrasting levels of inundation within the Mamirauá Sustainable Development Reserve, western Brazilian Amazon, and compared it with the overstory flora at the same study sites. In total, 1486 individuals and 116 woody species were recorded on an area totaling 3140 m². Individual densities and tree species richness were considerably higher in the high várzea than in the low várzea, which suggests that the heights and durations of the annual inundations are the main factor limiting species regeneration. In addition, approximately one third of the recorded species with densities ≥8 individuals showed regular or random spatial distribution patterns, which suggests that floodwaters act on dispersal strategies and species establishment.Independent of the forest type, floristic similarity between the understory and the overstory amounted to approximately 35%, and to approximately 10% when compared to other understory inventories in Amazonian várzea. Although the inventoried area of the understory amounted to only 16% of that of the overstory, species richness accounted for approximately 52-56% of that of the overstory. The results indicate that the understory flora of várzea forests is distinct and that it significantly increases local tree species richness. The understory flora of várzea forests therefore should be addressed in floristic inventories that provide the basis for regional and/or basin-wide estimations of tree diversity.     

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.     

Seasonal patterns of spatial variation in understory bird assemblages across a mosaic of flooded and unflooded Amazonian forests

We examined seasonal patterns of spatial variation in understory bird assemblages across a mosaic of upland and floodplain forests in central Amazonia, where variation in flooding patterns and floodwater nutrient load shapes a marked spatial heterogeneity in forest structure and composition. Despite great differences in productivity due to flooding by either nutrient-rich “white waters” (várzea) or nutrient-poor “black waters” (igapó), bird assemblages in the two floodplain forest types were relatively similar, showing lower abundances than adjacent upland forests (terra firme) and sharing a set of species that were absent or scarce elsewhere. Species that breed in pensile nests overhanging water were abundant in floodplain forests, whereas species that feed on the ground were generally scarce. Flooding affected assemblage dynamics in floodplain forests, with some influx of ground-dwelling species such as ant-following birds from adjacent upland during the low-water season, and the occupation by riverine and aquatic species such as kingfishers during floods. Spatial configuration influenced the seasonal pattern of assemblage structuring, with movements from terra firme occurring primarily to adjacent igapó forests. No such influx was detected in várzea forests that were farther from terra firme and isolated by wide river channels. Results support the view that habitat heterogeneity created by flooding strongly contributes to maintain diverse vertebrate assemblages in Amazonia forest landscapes, even in the case of largely sedentary species such as understory forest birds. Including both upland and floodplain forests in Amazonia reserves may thus be essential to preserve bird diversity at the landscape scale.     

Tree species distribution in<Emphasis Type="Italic">Várzea</Emphasis> forests of Brazilian Amazonia

Amazonian várzea forests are floodplains inundated by nutrient-rich white-water rivers occurring along the Amazon River. They are regularly flooded for up to 210 days per year by water columns of 10–15 m. Topographic variation results in different flooding amplitudes and durations along the flooding gradient, where the different tolerance to flooding of different plant species results in a vegetation zonation. We made a review of literature about the vegetation composition ofvárzea floodplain forests of Brazilian Amazonia along the Amazon River. Twenty-two studies were selected. Basing on the distribution of inventories which are concentrated in three main areas around the three larger cities Belém, Manaus and Tefé, we classified the inventories into three regions: (A) Estuary region with flooding regime influenced by daily inundations linked to the tides; (B) Central Amazonia near Manaus; (C) Western part of Brazilian Amazonia bordering Peru and Colombia, including Tefé and the “Reserva de Desenvolvimento Sustentável Mamirauá”. Summarizing the analyzed species lists, 36 tree species were registered in all sampled regions including the estuary. The regions A +C have 63 species in common, region B+C 143, and A+B 50. In the inventories analyzed here, an increase in species numbers from East to West can be confirmed, but it is difficult to state whether this is not an artefact due to local sampling. Vertical zonation patterns are difficult to discuss due to the lack of comparable data. The inventoried areas are small, and there is an urgent need for comparable floristic inventories throughout the basin. Destruction is spreading rapidly and the traditional use of forests and its resources is changing to a destructive exploitation that already has changed much of the physiognomy and diversity of this unique ecosystem.     

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.     

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.     

Patterns of plant phenology in Amazonian seasonally flooded and unflooded forests

Few studies have successfully monitored community‐wide phenological patterns in seasonally flooded Amazonian várzea forests, where a prolonged annual flood pulse arguably generates the greatest degree of seasonality of any low‐latitude ecosystem on Earth. We monitored the vegetative and reproductive plant phenology of várzea (VZ) floodplain and adjacent terra firme (TF) forests within two contiguous protected areas in western Brazilian Amazonia, using three complementary methods: monthly canopy observations of 1056 individuals (TF: 556, VZ: 500), twice monthly collections from 0.5‐m² litterfall traps within two 100‐ha plots (1 TF, 1 VZ; 96 traps per plot), and monthly ground surveys of residual fruit‐fall along transect‐grids within each 100‐ha plot (12 km per plot). Surveys encompassed the entire annual flood cycle and employed a floating trap design to cope with fluctuating water levels. Phenology patterns were generally similar in both forest types. Leaffall peaked during the aquatic phase in várzea forest and the dry season in terra firme. Flowering typically followed leaffall and leaf flush, extending into the onset of the terrestrial phase and rainy season in várzea and terra firme, respectively. Abiotic seed dispersal modes were relatively more prevalent in várzea than terra firme; the main contrast in fruiting seasonality was more likely a result of differences in community composition and relative abundance of seed dispersal modes than differences within individual genera. We emphasize the difficulty in distinguishing the role of the flood pulse from other seasonal environmental variables without multiannual data or spatially replicated studies across the spectrum of Amazonian forest types.     

Distribution and population structure of four Central Amazonian high-várzea timber species

Amazonian white-water (várzea) floodplains harbor many commercially important timber species which in Brazil are harvested following regulations of the Federal Environmental Agency (IBAMA). Although it is well-known that tree physiology, growth, and species distribution of Amazonian floodplain trees is linked to the heights and durations of the periodical inundations, information about timber stocks and population dynamics is lacking for most tree species. We investigated timber stocks and the population structure of four intensely logged tree species in a western Brazilian várzea forest on an area totaling 7.5 ha. Spatial distribution was investigated in all trees as a function of inundation height and duration and the distance to the river channel, and additionally for saplings (trees <10 cm diameter at breast height––DBH) as a function of the relative photosynthetically active radiation (rPAR). The diameter-class distribution in Hura crepitans and Ocotea cymbarum indicated that populations are subject to density variations that possibly are traced to small-scale flood variability. In all species, saplings concentrated at higher topographic elevations than the mature tree populations, which suggest that the physical ‘escape’ from a flooded environment is an important acclimation to flooding. While Ocotea cymbarum and Guarea guidonia were high-density wood species characterized by random dispersion and a pronounced shade-tolerance, Hura crepitans and Sterculia apetala presented lower wood density, aggregated dispersion, and were more light-demanding. All species presented exploitable stems according to the current harvest regulations, with elevated abundances in comparison to other Amazonian forest types. However, stem densities are below the harvest rates indicating that the harvest regulations are not sustainable. We recommend that the forest management in várzea forests should include specific establishment rates of timber species in dependence of the peculiar site conditions to achieve sustainability.     

Distance Decay of Tree Species Similarity in Protected Areas on Terra Firme Forests in Colombian Amazonia

In this study, we investigated the pattern of floristic similarity as a function of geographical distances and environmental variability in well-drained uplands (terra firme) in Colombian Amazonia. The study site comprised three National Natural Parks, Tinigua, Chiribiquete, and Amacayacu, located in different geological units that represent a soil fertility gradient linked to parental materials. Differences in species richness between sites were compared using rarefaction analysis. A clear floristic transition appeared in the east–west direction following a soil fertility gradient along the first PCoA axis. In multiple regression analyses based on distance matrices, both geographical distances and geology explained 64 percent of the total floristic variation. Geographical distances alone accounted for 12 percent of variation in floristic similarities among plots, while geology alone accounted for 1 percent, and the joint effect of both explained 51 percent of the floristic variation. The species richness trend supports the existence of a latitudinal corridor southward of the geographical Equator in the Amazon basin, where tree diversity reaches the maximum expected values. A coupled effect of stochastic dispersal limitation and habitat specialization would certainly appear to be an appropriate explanation for tree species turnover in terra firme forests in Colombian Amazonia, strongly emphasizing that competition and neutrality must be supplementary rather than mutually exclusive processes. This result pinpoints the effect of dispersal on floral mixing as an ongoing active process for structuring tree communities in NW Amazonia, and the size of the reserves as a relevant issue to protect rare species from extinction by chance.
     

Peatland forests are the least diverse tree communities documented in Amazonia,but contribute to high regional beta‐diversity

Western Amazonia is known to harbour some of Earth's most diverse forests, but previous floristic analyses have excluded peatland forests which are extensive in northern Peru and are among the most environmentally extreme ecosystems in the lowland tropics. Understanding patterns of tree species diversity in these ecosystems is important both for quantifying beta‐diversity in this region, and for understanding determinants of diversity more generally in tropical forests. Here we explore patterns of tree diversity and composition in two peatland forest types – palm swamps and peatland pole forests – using 26 forest plots distributed over a large area of northern Peru. We place our results in a regional context by making comparisons with three other major forest types: terra firme forests (29 plots), white‐sand forests (23 plots) and seasonally‐flooded forests (11 plots). Peatland forests had extremely low (within‐plot) alpha‐diversity compared with the other forest types that were sampled. In particular, peatland pole forests had the lowest levels of tree diversity yet recorded in Amazonia (20 species per 500 stems, Fisher's alpha 4.57). However, peatland pole forests and palm swamps were compositionally different from each other as well as from other forest types in the region. Few species appeared to be peatland endemics. Instead, peatland forests were largely characterised by a distinctive combination of generalist species and species previously thought to be specialists of other habitats, especially white‐sand forests. We suggest that the transient nature and extreme environmental conditions of Amazonian peatland ecosystems have shaped their current patterns of tree composition and diversity. Despite their low alpha‐diversity, the unique combination of species found in tree communities in Amazonian peatlands augment regional beta‐diversity. This contribution, alongside their extremely high carbon storage capacity and lack of protection at national level, strengthens their status as a conservation priority.     

Aboveground Roots in Amazonian Floodplain Trees1

Sediment‐rich rivers seasonally flood central Amazonian várzea forests, leading to periodic anoxic conditions in the rhizosphere and requiring morphological and structural adaptations, such as aboveground root systems. We investigated some possible relationships between root types and environmental factors in forest plots covering 3.1 ha of várzea in the Mamirauá Sustainable Development Reserve, Brazil. Digital elevation models of the study sites were obtained; sedimentation and soil texture were investigated to check relationship between position of trees on the flood gradient, soil conditions, and aboveground root systems. Different types of aboveground roots were closely related to flooding duration and habitat dynamics. Species subjected to higher and more prolonged floods tended to produce more aboveground roots than species subjected to lower and shorter inundations. Plank‐buttressing species increased with decreasing flood height and/or flood duration, and with increasing growth height and basal area. Habitats inundated for long periods were dominated by species with low growth heights and low basal areas, which formed stilt roots and aerial roots. Root system and sediment deposition showed a close relationship, plank buttressing being more common in sites subjected to lower sediment rates. In the disturbed sites close to the main river channel colonized by pioneer species, the occurrence of buttresses was lower than in less disturbed climax stages. No clear relationship was found between root systems and sediment grain sizes.     

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.     

Wood growth of Tabebuia barbata (E. Mey.) Sandwith (Bignoniaceae) and Vatairea guianensis Aubl. (Fabaceae) in Central Amazonian black-water (igapó) and white-water (várzea) floodplain forests

Tree growth is a fundamental indicator for conservation plans of Amazonian floodplain forests. In this study we use dendrochronology to analyze wood growth patterns of Tabebuia barbata and Vatairea guianensis, two tree species occurring in nutrient-rich white-water (várzea, Mamirauá Sustainable Development Reserve, MSDR) and nutrient-poor black-water (igapó, Amanã Sustainable Development Reserve, ASDR) floodplain forests of Central Amazonia. From 20 trees per species and floodplain system (total of 80 trees) growing under a similar flooding regime with a mean inundation height of about 4 m we measured diameter at breast height (dbh). We sampled two cores per tree with an increment corer at the height of dbh to determine wood density (WD), tree age and mean radial increment (MRI) rates. The wood samples were macroscopically analyzed. Both tree species show distinct annual tree rings characterized by marginal parenchyma tissues. MRI was measured by a digital measuring device and WD was determined by the ratio dry mass/fresh volume. MRI of both tree species was significantly higher in the várzea than in the igapó, which can be traced back to the contrasting nutrient status. WD showed no difference comparing both floodplain forest types. Tree ages of a species for the same diameter are more than twofold higher in the igapó than in the várzea. To insure a sustainable harvest, felling cycles in these forests should be adjusted according to rates of growth.     

A classification of major natural habitats of Amazonian white-water river floodplains (várzeas)

Most countries sharing the Amazon basin have signed the Ramsar Convention on Wetlands of International Importance but still lack complete wetland inventories, classification systems, and management plans. Amazonian wetlands vary considerably with respect to hydrology, water and soil fertility, vegetation cover, diversity in plant and animal species and primary and secondary productivity. Here, we propose a classification system of major natural habitats of Amazonian white-water river floodplains (várzeas) based on hydrological, water and soil chemistry and biological parameters. The Amazonian várzea is one of the largest Amazonian wetlands. It is exceptionally rich in plant and animal species and plays important roles in landscape history, evolution, hydrology and biogeochemical cycles of the Amazon basin. Most of Amazonia??s rural population lives in or along the várzea, emphasizing the economic importance of its natural resources. Our classification indicates five major systems, which are subdivided into 10 main habitats and up to 40 functional (vegetation) units of which the most important mesohabitats are described. We understand this classification as a dynamic system, as it is open to the inclusion of future research attempts and habitats without affecting the entire classification system. Our classification may be used for scientific purposes, such as comparative studies on biomass, productivity, biogeochemical cycles and biodiversity. Also, because the classification builds on habitat types and/or vegetation and functional units already distinguished by the local population it may be especially useful in guiding intelligent use of várzea habitat for specific management activities, such as agriculture, animal husbandry, forestry, fisheries, and conservation.     

Tree community structure reflects niche segregation of three parapatric squirrel monkey species (<Emphasis Type="Italic">Saimiri</Emphasis> spp.)

Integration between ecology and biogeography provides insights into how niche specialization affects the geographical distribution of species. Given that rivers are not effective barriers to dispersal in three parapatric species of squirrel monkeys (Saimiri vanzolinii, S. cassiquiarensis and S. macrodon) inhabiting floodplain forests of Central Amazonia, we tested whether forest structure and tree diversity may explain species differences in niche specialization and spatial segregation. We sampled 6617 trees of 326 species in three habitats (high várzea, low várzea and chavascal) used by three Saimiri species, and estimated tree species richness in each of them. For each tree, we measured variables known to influence habitat use in primates, such as crown area and presence of lianas, epiphytes and hemi-epiphytes. We used ANOVA to compare these variables and performed multivariate analyses (NMDS, ANOSIM and SIMPER) to evaluate dissimilarities in forest structure among each habitat inhabited by the three Saimiri species. We identified differences in the tree species richness, crown area and presence of lianas, epiphytes and hemi-epiphytes between the three habitats for all Saimiri species. NMDS demonstrated that areas of high and low várzeas occupied by S. vanzolinii were clearly separated from the other species. We also found that different plant species contributed to dissimilarity among Saimiri ranges. Our findings support the hypothesis that tree community structure may promote niche specialization and spatial segregation among primates. We discuss how these patterns could have been favored by historical changes in forest flood patterns, the evolutionary history of Saimiri spp., and past competition.