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.     

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.     

Germinative behaviour of ten tree species in white-water floodplain forests in central Amazonia

Amazonian floodplain forests (known as várzea) are classified into high or low várzea depending on the spatial position on the plains. This topographic feature exposes the terrain over different time periods of inundation, causing a major limiting factor for tree seedling establishment. We hypothesize that, strategically, most of the seeds produced by trees in low várzea forests germinate faster and in synchrony (temporally concentrated germination), and that their seedlings tend to have cotyledons without reserve or foliaceous cotyledons (PEF). By contrast, seeds produced by high-várzea specialist trees exhibit slower and temporally scattered germination, and their seedlings tend to have reserve storage cotyledons (CHR). Generalist species may show no clear pattern or may be related to high-várzea species. To test this hypothesis, diaspores of 10 tree species were collected: five of low-várzea specialist trees, three of high-várzea specialist trees and two of generalist species. Seedling emergence and morphology were monitored daily in a nursery for a period of 150 days of being subjected to non-flooded (sown directly in várzea soil) and flooded conditions (15 days in water before sowing in the same soil). The seedling emergence of low-várzea species showed an increase of 37% in germinability whereas high-várzea and generalist species exhibited a decrease of 38% and 35% of germinability, respectively. Foliaceous cotyledons were preferentially found in seedlings of low-várzea species, and storage cotyledons were more common in those of high-várzea species, indicating how cotyledon morphology may determine the amount and use of resources available to a seedling during the first stages of establishment and growth. Conservation plans aiming for the maintenance of ecosystem services must consider these strategies.     

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.     

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.     

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.     

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.     

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.     

The effects of forestry on carabid beetles (Coleoptera: Carabidae) in boreal forests

As compared to natural forests, managed boreal forests are younger, more homogeneous in terms of tree age and species composition, and consist of smaller fragments. Here we examine the effects of such characteristics caused by forestry on carabid beetles (Coleoptera, Carabidae) in the boreal region. The main results are the following. (1) Fragmentation of forests and the size of a fragment appear not to be crucial for the survival of the majority of forest carabids, as they tend to be distributed over various successional stages, but species requiring old-growth habitats suffer. (2) For carabids there appear to be no or very few edge specialist species, and forest-open land edges appear to be effective barriers for species associated with forest or open habitat. However, generalist species easily cross the edge, and edges of forest fragments may be invaded by species from the surrounding open habitat. (3) Habitat change following clear-cutting dramatically changes the composition of carabid assemblages: species restricted to mature forests disappear and open-habitat species invade, while habitat generalists survive at least in the short term. Carabid diversity can probably best be maintained if forest management mimics natural processes, maintains natural structures and includes the natural composition of vegetation and other structural elements (such as dead wood) within the stands, provided that these forest features can be maintained and recreated through forest management practices. At a larger scale, the whole spectrum of forest types and ages (especially old-growth forests), and different successional processes (especially fire) should be maintained. These require the development and use of innovative logging methods, and the planning, implementation, and assessment of landscape-scale ecological management strategies.     

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.     

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.     

Wood growth patterns of <Emphasis Type="Italic">Macrolobium acaciifolium</Emphasis> (Benth.) Benth. (Fabaceae) in Amazonian black-water and white-water floodplain forests

Macrolobium acaciifolium (Benth.) Benth. (Fabaceae) is a dominant legume tree species occurring at low elevations of nutrient-poor black-water (igapó) and nutrient-rich white-water floodplain forests (várzea) of Amazonia. As a consequence of the annual long-term flooding this species forms distinct annual tree rings allowing dendrochronological analyses. From both floodplain types in Central Amazonia we sampled cores from 20 large canopy trees growing at identical elevations with a flood-height up to 7 m. We determined tree age, wood density (WD) and mean radial increment (MRI) and synchronized ring-width patterns of single trees to construct tree-ring chronologies for every study site. Maximum tree age found in the igapó was more than 500 years, contrary to the várzea with ages not older than 200 years. MRI and WD were significantly lower in the igapó (MRI=1.52±0.38 mm year^?1, WD=0.39±0.05 g cm^?3) than in the várzea (MRI=2.66±0.67 mm year^?1, WD=0.45±0.03 g cm^?3). In both floodplain forests we developed tree-ring chronologies comprising the period 1857–2003 (n=7 trees) in the várzea and 1606–2003 (n=13 trees) in the igapó. The ring-width in both floodplain forests was significantly correlated with the length of the terrestrial phase (vegetation period) derived from the daily recorded water level in the port of Manaus since 1903. In both chronologies we found increased wood growth during El Niño events causing negative precipitation anomalies and a lower water discharge in Amazonian rivers, which leads to an extension of the terrestrial phase. The climate signal of La Niña was not evident in the dendroclimatic proxies.     

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.     

The Finnish forest site type approach: ordination and classification studies of mesic forest sites in southern Finland

A.K. Cajander's forest site type classification system is based on definition of plant communities typical to certain climatical and edaphical site conditions, but the structure and composition of the tree stands in Finland are considered sensitive to random variation and are therefore not used as primary classification criteria. The system has often received criticism, usually that the effects of the tree stand and successional stage of the stand have been underestimated. Most of the present-day forest stands in Finland represent young successional stages and are subjected to intensive management. This should result in an additional difficulty in the application of the forest site types in the field.The present study is based on three independent data sets representing forests on mineral soil in southern part of Finland. TWINSPAN classification, DCA ordination and canonical correspondence analysis (CCA) techniques were applied in successive stages of the data analysis. It was found that the definition of the intermediately fertile, mesic site types was clearly confused by the effects of the tree species and age of the stand. The analyses also revealed that the succession pathways on mesic forest sites are largely determined by the tree species composition. In stands dominated by Pinus sylvestris, the succession follows the competitive hierarchy model, whereas in stands dominated by Picea abies, severe shading of the tree canopy governs the development of understorey vegetation.Abbreviations CCA Canonical correspondence Analysis - DCA Detrended correspondence Analysis - TWINSPAN Two-way indicator species analysis     

Removal of cattle accelerates tropical dry forest succession in Northwestern Mexico

Domestic livestock influence patterns of secondary succession across forest ecosystems. However, the effects of cattle on the regeneration of tropical dry forests (TDF) in Mexico are poorly understood, largely because it is difficult to locate forests that are not grazed by cattle or other livestock. We describe changes in forest composition and structure along a successional chronosequence of TDF stands with and without cattle (chronic grazing or exclusion from grazing for ~ 8 year). Forest stands were grouped into five successional stages, ranging from recently abandoned to mature forest, for a total of 2.7 ha of the sampled area. The absence of cattle increased woody plant (tree and shrub) density and species richness, particularly in mid-successional and mature forest stands. Species diversity and evenness were generally greater in sites where cattle were removed and cattle grazing in early successional stands reduced establishment and/or recruitment of new individuals and species. Removal of cattle from forest stands undergoing succession appears to facilitate a progressive and non-linear change of forest structure and compositional attributes associated with rapid recovery, while cattle browsing acts as a chronic disturbance factor that compromises the resilience and structural and functional integrity of the TDF in northwestern Mexico. These results are important for the conservation, management, and restoration of Neotropical dry forests.     

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.     

Forest Succession in Tropical Hardwood Hammocks of the Florida Keys: Effects of Direct Mortality from Hurricane Andrew1

A tree species replacement sequence for dry broadleaved forests (tropical hardwood hammocks) in the upper Florida Keys was inferred from species abundances in stands abandoned from agriculture or other anthropogenic acitivities at different times in the past. Stands were sampled soon after Hurricane Andrew, with live and hurricane‐killed trees recorded separately; thus it was also possible to assess the immediate effect of Hurricane Andrew on stand successional status. We used weighted averaging regression to calculate successional age optima and tolerances for all species, based on the species composition of the pre‐hurricane stands. Then we used weighted averaging calibration to calculate and compare inferred successional ages for stands based on (1) the species composition of the pre‐hurricane stands and (2) the hurricane‐killed species assemblages. Species characteristic of the earliest stages of post‐agricultural stand development remains a significant component of the forest for many years, but are gradually replaced by taxa not present, even as seedlings, during the first few decades. This compositional sequence of a century or more is characterized by the replacement of deciduous by evergreen species, which is hypothesized to be driven by increasing moisture storage capacity in the young organic soils. Mortality from Hurricane Andrew was concentrated among early‐successional species, thus tending to amplify the long‐term trend in species composition.     

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.     

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.