Tree Community Change across 700 km of Lowland Amazonian Forest from the Andean Foothills to Brazil

We describe patterns of tree community change along a 700-km transect through terra firme forests of western Amazonia, running from the base of the Andes in Ecuador to the Peru–Brazil border. Our primary question is whether floristic variation at large scales arises from many gradual changes or a few abrupt ones. Data from 54 1-ha tree plots along the transect support the latter model, showing two sharp discontinuities in community structure at the genus level. One is located near the Ecuador–Peru border, where the suite of species that dominates large areas of Ecuadorean forest declines abruptly in importance to the east. This discontinuity is underlain by a subterranean paleoarch and congruent with a change in soil texture. A second discontinuity is associated with the shift from clay to white sand soils near Iquitos. We hypothesize that the first discontinuity is part of an edaphic boundary that runs along the Andean piedmont and causes a transition from tree communities preferring richer, younger soils near the base of the Andes to those preferring poorer, older soils farther east. Because the floristic changes observed at this discontinuity are conserved for large distances to the east and west of it, the discontinuity is potentially key for understanding floristic variation in western Amazonia. The significant floristic turnover at the Ecuador–Peru border suggests that the only large protected area in the region—Ecuador's Yasuní National Park—is not adequate protection for the very diverse tree communities that cover vast areas of northern Peru.     

A comparison of fine-scale distribution patterns of four plant groups in an Amazonian rainforest

We carried out a comparison among the floristic patterns of four different plant groups (palms, trees, melastomes and pteridophytes) in a lowland rainforest site in Peruvian Amazonia. The study site consisted of a mosaic of edaphic patches reflecting the different geological formations that can be found on the surface. We collected the data along a linear transect (500 m long, divided into 20 × 20 m or 5 × 20 m subplots), and recorded of the four plant groups all individuals that exceeded a minimum size limit predefined for each plant group. We also recorded the drainage conditions and soil type classes in each subplot of the transect. The results indicated that different plant groups can produce similar floristic patterns in local spatial scales, and that these patterns reflect similarities in edaphic conditions. All matrix correlations calculated between pairs of the four plant groups were positive and statistically significant. Floristic composition in all plant groups correlated with soil class, and to a somewhat lesser degree with drainage. These results imply that any one of the four plant groups could serve as a rough indicator of more general floristic patterns, and that even the inventory of a limited part of the flora can shed light on the floristic variation found in Amazonian forests.     

Riqueza y Distribución Ecológica de Especies de Pteridofitas en la Zona del Río Yavarí-Mirín, Amazonía Peruana

We studied the ecological distribution of pteridophytes (ferns and fern allies) along eight 8-km transects covering 12.7 ha in Peruvian Amazonia. Subunits of 200 m² of the transects have previously been classified into four different forest types, and here we document and quantify the floristic differences among these forest types. Pteridophytes have been suggested as an indicator group to classify rain forest habitats, but this requires that the ecological preferences of the species are well documented and consistent across geographic regions. Here we analyzed in detail the distribution and diversity patterns of 130 species across the four rain forest types. Relative species abundance and species diversity were similar among some of the forest types and differed among others, but the species composition differed markedly. Our results largely confirmed the earlier interpretation of the edaphic preferences of the pteridophyte species in western Amazonia. This supports the proposition that deterministic processes have an important role in influencing the floristic composition of Amazonian forests.     

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.     

Regional-Scale Drivers of Forest Structure and Function in Northwestern Amazonia

Field studies in Amazonia have found a relationship at continental scales between soil fertility and broad trends in forest structure and function. Little is known at regional scales, however, about how discrete patterns in forest structure or functional attributes map onto underlying edaphic or geological patterns. We collected airborne LiDAR (Light Detection and Ranging) data and VSWIR (Visible to Shortwave Infrared) imaging spectroscopy measurements over 600 km² of northwestern Amazonian lowland forests. We also established 83 inventories of plant species composition and soil properties, distributed between two widespread geological formations. Using these data, we mapped forest structure and canopy reflectance, and compared them to patterns in plant species composition, soils, and underlying geology. We found that variations in soils and species composition explained up to 70% of variation in canopy height, and corresponded to profound changes in forest vertical profiles. We further found that soils and plant species composition explained more than 90% of the variation in canopy reflectance as measured by imaging spectroscopy, indicating edaphic and compositional control of canopy chemical properties. We last found that soils explained between 30% and 70% of the variation in gap frequency in these forests, depending on the height threshold used to define gaps. Our findings indicate that a relatively small number of edaphic and compositional variables, corresponding to underlying geology, may be responsible for variations in canopy structure and chemistry over large expanses of Amazonian forest.     

Determinants of Plant Community Assembly in a Mosaic of Landscape Units in Central Amazonia: Ecological and Phylogenetic Perspectives

The Amazon harbours one of the richest ecosystems on Earth. Such diversity is likely to be promoted by plant specialization, associated with the occurrence of a mosaic of landscape units. Here, we integrate ecological and phylogenetic data at different spatial scales to assess the importance of habitat specialization in driving compositional and phylogenetic variation across the Amazonian forest. To do so, we evaluated patterns of floristic dissimilarity and phylogenetic turnover, habitat association and phylogenetic structure in three different landscape units occurring in terra firme (Hilly and Terrace) and flooded forests (Igapó). We established two 1-ha tree plots in each of these landscape units at the Caparú Biological Station, SW Colombia, and measured edaphic, topographic and light variables. At large spatial scales, terra firme forests exhibited higher levels of species diversity and phylodiversity than flooded forests. These two types of forests showed conspicuous differences in species and phylogenetic composition, suggesting that environmental sorting due to flood is important, and can go beyond the species level. At a local level, landscape units showed floristic divergence, driven both by geographical distance and by edaphic specialization. In terms of phylogenetic structure, Igapó forests showed phylogenetic clustering, whereas Hilly and Terrace forests showed phylogenetic evenness. Within plots, however, local communities did not show any particular trend. Overall, our findings suggest that flooded forests, characterized by stressful environments, impose limits to species occurrence, whereas terra firme forests, more environmentally heterogeneous, are likely to provide a wider range of ecological conditions and therefore to bear higher diversity. Thus, Amazonia should be considered as a mosaic of landscape units, where the strength of habitat association depends upon their environmental properties.     

Avian Composition Co‐varies with Floristic Composition and Soil Nutrient Concentration in Amazonian Upland Forests

Spatial heterogeneity in the plant species composition of tropical forests is expected to influence animal species abundance and composition because vegetation constitutes the primary habitat feature for forest animals. Floristic variation is tied to variation in soils, so edaphic properties should ultimately influence animal species composition as well. The study of covariation in floristic and faunistic turnover has been hindered by the difficulty of completing coordinated surveys in hyperdiverse tropical communities, but this can be overcome with the use of a few plant taxa that function as surrogates for general floristic turnover. We used avian and plant transect surveys and soil sampling in a western Amazonian upland (terra firme) forest landscape to test whether spatial variation in bird community composition is associated with floristic turnover and corresponding edaphic gradients. Partial Mantel tests and Non‐metric Multidimensional Scaling showed floristic distinctiveness between two forest types closely associated with differences in soil cation concentrations, and differences in both floristic composition and cation concentrations were further linked to compositional differences in avian species, independent of geographic distances among sites. Ten percent of bird species included in Indicator Species Analyses showed significant associations with one of the two forest types. The upland forest types that we sampled, each corresponding to a different geological formation, are intermediate relative to edaphically extreme environments in the region. Models of avian diversification should take into account this environmental heterogeneity, as should conservation planning approaches that aim to represent faunal diversity. Abstract in Spanish is available in the online version of this article.     

The contribution of edaphic heterogeneity to the evolution and diversity of Burseraceae trees in the western Amazon

Environmental heterogeneity in the tropics is thought to lead to specialization in plants and thereby contribute to the diversity of the tropical flora. We examine this idea with data on the habitat specificity of 35 western Amazonian species from the genera Protium, Crepidospermum, and Tetragastris in the monophyletic tribe Protieae (Burseraceae) mapped on a molecular-based phylogeny. We surveyed three edaphic habitats that occur throughout terra firme Amazonia: white-sand, clay, and terrace soils in eight forests across more than 2000 km in the western Amazon. Twenty-six of the 35 species were found to be associated with only one of three soil types, and no species was associated with all three habitats; this pattern of edaphic specialization was consistent across the entire region. Habitat association mapped onto the phylogenetic tree shows association with terrace soils to be the probable ancestral state in the group, with subsequent speciation events onto clay and white-sand soils. The repeated gain of clay association within the clade likely coincides with the emergence of large areas of clay soils in the Miocene deposited during the Andean uplift. Character optimizations revealed that soil association was not phylogenetically clustered for white-sand and clay specialists, suggesting repeated independent evolution of soil specificity is common within the Protieae. This phylogenetic analysis also showed that multiple cases of putative sister taxa with parapatric distributions differ in their edaphic associations, suggesting that edaphic heterogeneity was an important driver of speciation in the Protieae in the Amazon basin.     

THE CONTRIBUTION OF EDAPHIC HETEROGENEIYT TO THE EVOLUTION AND DIVERSITY OF BURSERACEAR TREES IN THE WESTERN AMAZON

Abstract —Environmental heterogeneity in the tropics is thought to lead to specialization in plants and thereby contribute to the diversity of the tropical flora. We examine this idea with data on the habitat specificity of 35 western Amazonian species from the genera Protium, Crepidospermum, and Tetragastris in the monophyletic tribe Protieae (Burseraceae) mapped on a molecular‐based phylogeny. We surveyed three edaphic habitats that occur throughout terra firme Amazonia: white‐sand, clay, and terrace soils in eight forests across more than 2000 km in the western Amazon. Twenty‐six of the 35 species were found to be associated with only one of three soil types, and no species was associated with all three habitats; this pattern of edaphic specialization was consistent across the entire region. Habitat association mapped onto the phylogenetic tree shows association with terrace soils to be the probable ancestral state in the group, with subsequent speciation events onto clay and white‐sand soils. The repeated gain of clay association within the clade likely coincides with the emergence of large areas of clay soils in the Miocene deposited during the Andean uplift. Character optimizations revealed that soil association was not phylogenetically clustered for white‐sand and clay specialists, suggesting repeated independent evolution of soil specificity is common within the Protieae. This phylogenetic analysis also showed that multiple cases of putative sister taxa with parapatric distributions differ in their edaphic associations, suggesting that edaphic heterogeneity was an important driver of speciation in the Protieae in the Amazon basin.     

Edaphic niche differentiation among Polybotrya ferns in western Amazonia: implications for coexistence and speciation

To study the degree of edaphic specialization in Amazonian plants, the distribution patterns of seven species of Polybotrya ferns were studied in 109 sites in a climatically uniform area of northwestern Amazonia (Colombia, Ecuador and northern Peru). The two most abundant species of Polybotrya were found in about two-thirds of the sites with almost 7000 individuals each, the rarest species occurred in just one site with 40 individuals. Each of the seven species appeared to have a unique realised niche, when niche dimensions were defined by gradients in soil texture, soil cation content, and inundation. The species also differed in how broadly or narrowly they were distributed along each gradient. Some species were practically never found in the same sites, whereas others co-occurred with a high frequency, in spite of showing clearly different abundance patterns among sites. A single site only contains a small part of the edaphic variation present in the landscape, and a small proportion of any species' niche space, so broad-scale studies are needed to adequately describe and compare species' niches and to assess to what degree niche differences promote species coexistence. The distribution patterns in Polybotrya are consistent with, but do not prove, that ecological speciation may have been important in the radiation of the genus. If such a pattern is found to be common in other Amazonian plants, this would indicate that each evolutionary lineage has adapted to the available habitats largely independently of the others.     

Diversity and biogeography of vascular epiphytes in Western Amazonia, Yasuní, Ecuador

Aim Although vascular epiphytes are important components of species richness and complexity of Neotropical forests, vascular epiphytes are under‐represented in large scale biogeographical analyses. We studied the diversity, biogeography and floristic relationships of the epiphytic flora of the Yasuní region (Western Amazonia) in a Neotropical context, with special emphasis on the influence of the Andean flora on floristic composition and diversity of surrounding lowland forests. Location Western Amazonian lowland rainforest, Tiputini Biodiversity Station (0°38′ S 76°09′ W, 230 m a.s.l., 650 ha), Yasuní National Park, Ecuador. Methods We compared the vascular epiphyte flora of Yasuní with 16 published Neotropical epiphyte inventories. Secondly, based on a floristic database with records of more than 70,000 specimens of vascular epiphytes from the Neotropics the elevational composition of eight selected inventories was analysed in detail. Results The vascular epiphyte flora of Yasuní is characterized by a very high species richness (313 spp.). A moderate portion of species is endemic to the Upper Napo region (c. 10%). However, this figure is much higher than previous analyses primarily based on woody species suggested. Geographical ranges of these species match with a proposed Pleistocene forest refuge. Compared with Northern and Central Amazonian sites, Western Amazonian epiphyte communities are characterized by a higher portion of montane and submontane species. Species richness of vascular epiphytes at the sites was correlated with the amount of rainfall, which is negatively correlated with the number of dry months. Main conclusion Recent and historic patterns of rainfall are the driving forces behind diversity and floristic composition of vascular epiphytes in Western Amazonia: high annual rainfall in combination with low seasonality provides suitable conditions to harbour high species richness. The proximity to the Andes, the most important centre of speciation for most Neotropical epiphytic taxa, in combination with the climatic setting has allowed a continuous supply of species richness to the region. At least for epiphytes, the borderline between the Andean and Amazonian flora is much hazier than previously thought. Moreover, the comparatively moist climate in Western Amazonia during the Pleistocene has probably led to fewer extinctions and/or more speciation than in more affected surrounding lowlands.     

Predicting and quantifying the structure of tropical dry forests in South Florida and the Neotropics using spaceborne imagery

Aim This research examines environmental theories and remote sensing methods that have been hypothesized to be associated with tropical dry forest structure. Location Tropical dry forests of South Florida and the Neotropics. Methods Field measurements of stand density, basal area and tree height were collected from 22 stands in South Florida and 30 stands in the Neotropics. In South Florida, field measurements were compared to climatic (temperature, precipitation, hurricane disturbance) and edaphic (rockiness, soil depth) variables, spectral indices (NDVI, IRI, MIRI) from Landsat 7 ETM+, and estimates of tree height from the Shuttle Radar Topography Mission (SRTM) and the National Elevation Dataset (NED). Environmental variables associated with tropical dry forest structure in South Florida were compared to tropical dry forest in other Neotropical sites. Results There were significant correlations among temperature and precipitation, and stand density and tree height in South Florida. There were significant correlations between (i) stand density and mean NDVI and standard deviation of NDVI, (ii) MIRI and stand density, basal area and mean tree height, and (iii) estimates of tree height from SRTM with maximum tree height. In the Neotropics, there were no relationships between temperature or precipitation and tropical dry forest structure, however, Neotropical sites that experience hurricane disturbance had significantly shorter tree heights and higher stand densities. Main conclusions It is possible to predict and quantify the forest structure characteristics of tropical dry forests using climatic data, Landsat 7 ETM+ imagery and SRTM data in South Florida. However, results based on climatic data are region‐specific and not necessarily transferable between tropical dry forests at a continental spatial scale. Spectral indices from Landsat 7 ETM+ can be used to quantify forest structure characteristics, but SRTM data are currently not transferable to other regions. Hurricane disturbance has a significant impact on forest structure in the Neotropics.     

Edaphic and Floristic Variation within a 1‐ha Plot of Lowland Amazonian Rain Forest1

Several studies in lowland tropical rain forests have documented effects of local‐scale topographic variation on plant species distribution and abundance patterns. Few studies have compared the distribution patterns of more than one plant group, however, and even fewer have related these to measured physical and chemical soil characteristics. Here, we document such soil characteristics within a square 1‐ha plot in Amazonian Ecuador, and compare them to the distribution patterns of terrestrial pteridophytes, angiosperm ground herbs, and palms. Substantial variation in soil properties was found within the plot. The three plant groups showed highly correlated floristic patterns within the 1‐ha plot even after the effect of geographical distances had been taken into account. Mantel tests yielded significant correlations between edaphic patterns, as measured by distances in various soil and topographic characteristics, and floristic patterns. For all three plant groups, differences in elevation within the plot were highly correlated with floristic distances, and for terrestrial pteridophytes and palms, distances in soil calcium content and sand content were also important. Our results resembled those obtained at wider spatial scales with the same plant groups, which indicates that soil factors may play an important role for distribution and beta diversity of plants, even at the local scale.     

Evidence for ecological divergence across a mosaic of soil types in an Amazonian tropical tree: Protium subserratum (Burseraceae)

Soil heterogeneity is an important driver of divergent natural selection in plants. Neotropical forests have the highest tree diversity on earth, and frequently, soil specialist congeners are distributed parapatrically. While the role of edaphic heterogeneity in the origin and maintenance of tropical tree diversity is unknown, it has been posited that natural selection across the patchwork of soils in the Amazon rainforest is important in driving and maintaining tree diversity. We examined genetic and morphological differentiation among populations of the tropical tree Protium subserratum growing parapatrically on the mosaic of white‐sand, brown‐sand and clay soils found throughout western Amazonia. Nuclear microsatellites and leaf morphology were used to (i) quantify the extent of phenotypic and genetic divergence across habitat types, (ii) assess the importance of natural selection vs. drift in population divergence, (iii) determine the extent of hybridization and introgression across habitat types, (iv) estimate migration rates among populations. We found significant morphological variation correlated with soil type. Higher levels of genetic differentiation and lower migration rates were observed between adjacent populations found on different soil types than between geographically distant populations on the same soil type. P_ST‐F_ST comparisons indicate a role for natural selection in population divergence among soil types. A small number of hybrids were detected suggesting that gene flow among soil specialist populations may occur at low frequencies. Our results suggest that edaphic specialization has occurred multiple times in P. subserratum and that divergent natural selection across edaphic boundaries may be a general mechanism promoting and maintaining Amazonian tree diversity.     

Distribution and Diversity of Pteridophytes and Melastomataceae along Edaphic Gradients in Yasuní National Park, Ecuadorian Amazonia1

We documented the floristic composition of pteridophytes (ferns and fern allies) and Melastomataceae in Yasuní National Park, Amazonian Ecuador. Our main questions were: (1) Are the density of individuals, species richness, and/or species diversity (measured with Shannon's H′) of the two plant groups related to edaphic differences? and (2) How many of the pteridophyte and Melastomataceae species are non–randomly distributed in relation to a soil base content gradient within terra firme (non–inundated forest). To answer these questions, we sampled 27 line transects of 500 × 5 m distributed in an area of ca 20 × 25 km. The study area included a permanent 50 ha plot established to monitor forest dynamics; thus, our results also provide information on landscape–scale floristic variability to which results from within the plot can be compared. A total of 45,608 individuals and 140 species of pteridophytes, and 4893 individuals and 89 species of the Melastomataceae, were counted in the transects. Both with pteridophytes and with Melastomataceae, a clear negative correlation was found between the amount of extractable bases in the soil and the number of plant individuals encountered in a transect. With Melastomataceae, species richness and species diversity also were negatively correlated with soil base content, but with pteridophytes they were not. More than 50 percent of the common species of both pteridophytes and Melastomataceae were nonrandomly distributed along the soil cation content gradient within terra firme. We conclude that while the species richness patterns observed in one plant group are not indicative of similar patterns in other plant groups, it seems likely that a substantial (but unknown) proportion of species belonging to other plant groups will be found to show distribution patterns that reflect edaphic preferences within terra firme forests.     

Using digital soil maps to infer edaphic affinities of plant species in Amazonia: Problems and prospects

Amazonia combines semi‐continental size with difficult access, so both current ranges of species and their ability to cope with environmental change have to be inferred from sparse field data. Although efficient techniques for modeling species distributions on the basis of a small number of species occurrences exist, their success depends on the availability of relevant environmental data layers. Soil data are important in this context, because soil properties have been found to determine plant occurrence patterns in Amazonian lowlands at all spatial scales. Here we evaluate the potential for this purpose of three digital soil maps that are freely available online: SOTERLAC, HWSD, and SoilGrids. We first tested how well they reflect local soil cation concentration as documented with 1,500 widely distributed soil samples. We found that measured soil cation concentration differed by up to two orders of magnitude between sites mapped into the same soil class. The best map‐based predictor of local soil cation concentration was obtained with a regression model combining soil classes from HWSD with cation exchange capacity (CEC) from SoilGrids. Next, we evaluated to what degree the known edaphic affinities of thirteen plant species (as documented with field data from 1,200 of the soil sample sites) can be inferred from the soil maps. The species segregated clearly along the soil cation concentration gradient in the field, but only partially along the model‐estimated cation concentration gradient, and hardly at all along the mapped CEC gradient. The main problems reducing the predictive ability of the soil maps were insufficient spatial resolution and/or georeferencing errors combined with thematic inaccuracy and absence of the most relevant edaphic variables. Addressing these problems would provide better models of the edaphic environment for ecological studies in Amazonia.     

Primate population densities in three nutrient-poor amazonian terra firme forests of south-eastern Colombia

We censused primate populations at three non-hunted 'terra firme' forests of south-eastern Colombian Amazonia. The aggregate biomass densities of diurnal primates at all sites were amongst the lowest recorded for any non-hunted forest in western Amazonia and elsewhere in the Neotropics. Densities of red howler monkeys were low, as is typical in Amazonian terra firme forests far removed from white-water rivers, and densities of woolly monkeys were 1.5-3.5 times lower than those estimated for this species in central-western Brazilian Amazonia. Densities of small to mid-sized primates except for brown capuchins (Cebus apella) and white-faced capuchins (Cebus albifrons) were similar to those of other oligotrophic Amazonian forest sites. Our results are in agreement with other studies showing that terra firme forests of lowland Amazonia typically sustain a low biomass density of primates and other mid-sized to large vertebrates. Large reserves are therefore required to assure the viability of primate populations in oligotrophic systems. Given the escalating negative impacts of human habitat disturbance and hunting in Colombian Amazonia, we urge that a baseline sampling protocol to quantify the abundance and distribution of the harvest-sensitive vertebrate fauna be established within protected areas and the large indigenous reserves so that conservation efforts can be defined and implemented.     

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.
     

Mapping gradual landscape-scale floristic changes in Amazonian primary rain forests by combining ordination and remote sensing

Aim We present a new method to economically map gradual changes in plant species composition in lowland rain forests using field data and satellite images. Such a method will be a useful tool in planning the sustainable use and conservation of Amazonian rain forests. Location The study covered an area of c. 700 km² of primary rain forest in Amazonian Ecuador. Methods We field inventoried the species composition of pteridophytes and Melastomataceae in 340 inventory plots (5 m × 50 m), described the prevailing topography and analysed soil cation concentration and texture. We used non‐metric multidimensional scaling (NMDS) to summarize the floristic variation among the inventory plots in three ordination dimensions. The scores of the three ordination axes were predicted to non‐visited places using a Landsat TM (thematic mapper) satellite image and the k nearest neighbours (knn) estimation method. To avoid extrapolation, we excluded from the analysis those pixel windows whose spectral values were not represented in the areas covered by field sampling. The accuracy of the predictions was evaluated by cross‐validation and by comparing the predictions based on spectrally nearest neighbours to the predictions based on random neighbours. Results The floristic gradients presented by NMDS ordination were interpretable in terms of topography, drainage and soil cation content. Thirteen percent of the cloud‐free pixels were excluded from the knn analysis to avoid extrapolation. The estimates of the floristic ordination scores based on spectrally nearest neighbours were always more accurate than estimates based on random neighbours. Main conclusions The presented method needs a relatively small input of work and resources, is mechanistic and produces maps that give relevant information on floristic variation over forest areas that are traditionally considered essentially homogeneous. Therefore, the method appears to have a great potential for use in mapping large areas of Amazonian rain forests.