Forest Canopy Gap Distributions in the Southern Peruvian Amazon

Canopy gaps express the time-integrated effects of tree failure and mortality as well as regrowth and succession in tropical forests. Quantifying the size and spatial distribution of canopy gaps is requisite to modeling forest functional processes ranging from carbon fluxes to species interactions and biological diversity. Using high-resolution airborne Light Detection and Ranging (LiDAR), we mapped and analyzed 5,877,937 static canopy gaps throughout 125,581 ha of lowland Amazonian forest in Peru. Our LiDAR sampling covered a wide range of forest physiognomies across contrasting geologic and topographic conditions, and on depositional floodplain and erosional terra firme substrates. We used the scaling exponent of the Zeta distribution (λ) as a metric to quantify and compare the negative relationship between canopy gap frequency and size across sites. Despite variable canopy height and forest type, values of λ were highly conservative (λ _mean  = 1.83, s  = 0.09), and little variation was observed regionally among geologic substrates and forest types, or at the landscape level comparing depositional-floodplain and erosional terra firme landscapes. λ-values less than 2.0 indicate that these forests are subjected to large gaps that reset carbon stocks when they occur. Consistency of λ-values strongly suggests similarity in the mechanisms of canopy failure across a diverse array of lowland forests in southwestern Amazonia.     

Holocene fire and occupation in Amazonia: records from two lake districts

While large-scale pre-Columbian human occupation and ecological disturbance have been demonstrated close to major Amazonian waterways, less is known of sites in terra firme settings. Palaeoecological analyses of two lake districts in central and western Amazonia reveal long histories of occupation and land use. At both locations, human activity was centred on one of the lakes, while the others were either lightly used or unused. These analyses indicate that the scale of human impacts in these terra firme settings is localized and probably strongly influenced by the presence of a permanent open-water body. Evidence is found of forest clearance and cultivation of maize and manioc. These data are directly relevant to the resilience of Amazonian conservation, as they do not support the contention that all of Amazonia is a 'built landscape' and therefore a product of past human land use.     

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.     

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.     

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.     

Species turnover in Amazonian frogs: low predictability and large differences among forests

The factors explaining species turnover at different spatial scales have been intensively studied, but most work in Amazonia has mainly focused on plants. For animals, it is not as obvious which environmental variables most affect differences in species composition among sites. We sought to identify what causes anuran turnover in Amazonian terra firme forests, and how the effectiveness of these factors varies among regions and across spatial scales. We sampled frogs in 56 plots along ponds and streams distributed over three terra firme forest areas in Eastern Amazonia. Using multiple regressions on distance matrices, we partitioned the variation in species turnover into components explained by variation in environmental and spatial distances. This was done in parallel for each area separately, and for all areas together, to assess the consistency of results between scales and across areas at the same scale. Each community seemed to respond to a set of factors specific to that area, and the identity of the variables that emerged as significant differed among areas and scales. Both geographical distances and environmental differences had larger explanatory power at the regional scale than at the local scale. The large differences among results from different areas caution against making broad generalizations about species turnover patterns from a single community, as real differences may exist among areas.     

The ancient tropical rainforest tree Symphonia globulifera L. f. (Clusiaceae) was not restricted to postulated Pleistocene refugia in Atlantic Equatorial Africa

Understanding the history of forests and their species'' demographic responses to past disturbances is important for predicting impacts of future environmental changes. Tropical rainforests of the Guineo-Congolian region in Central Africa are believed to have survived the Pleistocene glacial periods in a few major refugia, essentially centred on mountainous regions close to the Atlantic Ocean. We tested this hypothesis by investigating the phylogeographic structure of a widespread, ancient rainforest tree species, Symphonia globulifera L. f. (Clusiaceae), using plastid DNA sequences (chloroplast DNA [cpDNA], psbA-trnH intergenic spacer) and nuclear microsatellites (simple sequence repeats, SSRs). SSRs identified four gene pools located in Benin, West Cameroon, South Cameroon and Gabon, and São Tomé. This structure was also apparent at cpDNA. Approximate Bayesian Computation detected recent bottlenecks approximately dated to the last glacial maximum in Benin, West Cameroon and São Tomé, and an older bottleneck in South Cameroon and Gabon, suggesting a genetic effect of Pleistocene cycles of forest contraction. CpDNA haplotype distribution indicated wide-ranging long-term persistence of S. globulifera both inside and outside of postulated forest refugia. Pollen flow was four times greater than that of seed in South Cameroon and Gabon, which probably enabled rapid population recovery after bottlenecks. Furthermore, our study suggested ecotypic differentiation—coastal or swamp vs terra firme—in S. globulifera. Comparison with other tree phylogeographic studies in Central Africa highlighted the relevance of species-specific responses to environmental change in forest trees.     

Habitat differentiation in the threatened aquatic plant genus Baldellia (L.) Parl. (Alismataceae): Implications for conservation

A comparative analysis of the habitats of Baldellia ranunculoides (subsp. ranunculoides and subsp. repens) and Baldellia alpestris (Alismataceae) was carried out across central and western Europe. Soil samples from 43 populations were analyzed and the composition of accompanying vegetation was analyzed by canonical correspondence and indicator species analysis. Significant differences in the habitat conditions and the accompanying vegetation were found between the three taxa. B. ranunculoides subsp. repens is growing in lowland water bodies on acidic substrates (pH 5–6.3) which are markedly richer in organic matter and poorer in cations, especially in Ca²⁺. B. ranunculoides s. str. is also a lowland taxon, but grows on more mineral and basic substrates (pH 7–9), usually rich in Ca²⁺. The endemic B. alpestris grows in the most distinct habitats, on substrates with low pH and with very low Ca²⁺ concentrations, at much higher altitudes than the other taxa and in very specific plant communities that have almost no affinities with those of the two lowland Baldellia-taxa. The Baldellia-taxa are threatened by eutrophication of their habitats. Moreover, many of the extant populations, in particular those of B. ranunculoides s. str., are very small and will be threatened by environmental stochasticity. Reintroduction and the creation of new populations might reduce the risk of extinction. Information given on the detailed habitat preferences of the Baldellia-taxa could help conservation efforts for these endangered taxa and identify suitable sites for (re-) introductions.     

Habitat Selection and Use of Space by Bald-Faced Sakis (Pithecia irrorata) in Southwestern Amazonia: Lessons from a Multiyear, Multigroup Study

Population density and distribution in tropical forest vertebrates are directly linked to patterns of use of space relative to habitat structure and composition. To examine how forest type may explain the ranging behavior and high variance in group density observed within the geographic range of bald-faced saki monkeys (Pithecia irrorata), we monitored habitat use of 5 neighboring focal groups of this species in southwestern Amazonia over 3 yr. To test whether sakis are unflooded (terra firme) forest specialists, we compared home range (HR) use to the corresponding availability of 4 main forest types and quantified HR size and activity budgets as a function of forest type. HR size varied from 16 to 60 ha, and saki population density at this scale (12.5 ± 6.4 SD individuals/km²) was more closely related to forest type than to group size. Although sakis were not obligate habitat specialists, groups clearly avoided bamboo forest and preferred terra firme forest. Terra firme forests were associated with small HRs, intensive use, high HR overlap, and territorial defense, all of which suggest that saki densities will be higher in areas dominated by terra firme forest where large patches of bamboo (Guadua spp.) are absent. The increased desiccation and subsequent forest fires expected in this region from the combined impacts of climate change and human land use potentially threaten the long-term viability of old-growth terra firme forest specialists such as sakis. Regional-scale conservation efforts should ensure that extensive blocks of terra firme forest are protected in areas that remain relatively free of bamboo.     

Significance of Riparian Forests for the Conservation of Central African Primates

We compared the structure of 12 Central African primate communities, 6 in riparian forests and 6 in adjacent terra firma forests and discussed the implications for primate conservation. The communities in riparian forests included on average 1.5 times more primate species than those in terra firma forests due to the fact that riparian forests shelter 4 specialist species and 6–7 generalist species. The results differ from findings in Amazonia where riparian forests consistently have fewer primate species than terra firma forests accommodate. This may be partly explained by the water level amplitude in Amazonian riparian forests, which deterred the radiation of semiterrestrial species. In Africa, most riparian-specialist primates are terrestrially-adapted and have access to an enlarged food niche. In terms of African primate conservation, we recommend protecting riparian forests and adjacent terra firma forests so that most of the lowland forest diversity is captured. The linear shape of riparian forests (which allows gene flow over long distance) and their persistence in anthropic landscape (because they represent lands of lesser value for agriculture and logging than mainland forests) predispose them to act as biodiversity sanctuaries.     

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

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

A new species of <Emphasis Type="Italic">Pradosia</Emphasis> (Sapotaceae) from Central Amazonia

Pradosia lahoziana is here described as new. It is known from four collections from wet lowland and non-flooded terra firme forests near Manaus in Central Amazonia. Illustrations are provided together with a comparison of the morphological differences with similar species.     

Phyllostomid Bat Assemblage Structure in Amazonian Flooded and Unflooded Forests

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

Disentangling stand and environmental correlates of aboveground biomass in Amazonian forests

Tropical forests contain an important proportion of the carbon stored in terrestrial vegetation, but estimated aboveground biomass (AGB) in tropical forests varies two‐fold, with little consensus on the relative importance of climate, soil and forest structure in explaining spatial patterns. Here, we present analyses from a plot network designed to examine differences among contrasting forest habitats (terra firme, seasonally flooded, and white‐sand forests) that span the gradient of climate and soil conditions of the Amazon basin. We installed 0.5‐ha plots in 74 sites representing the three lowland forest habitats in both Loreto, Peru and French Guiana, and we integrated data describing climate, soil physical and chemical characteristics and stand variables, including local measures of wood specific gravity (WSG). We use a hierarchical model to separate the contributions of stand variables from climate and soil variables in explaining spatial variation in AGB. AGB differed among both habitats and regions, varying from 78 Mg ha^?1 in white‐sand forest in Peru to 605 Mg ha^?1 in terra firme clay forest of French Guiana. Stand variables including tree size and basal area, and to a lesser extent WSG, were strong predictors of spatial variation in AGB. In contrast, soil and climate variables explained little overall variation in AGB, though they did co‐vary to a limited extent with stand parameters that explained AGB. Our results suggest that positive feedbacks in forest structure and turnover control AGB in Amazonian forests, with richer soils (Peruvian terra firme and all seasonally flooded habitats) supporting smaller trees with lower wood density and moderate soils (French Guianan terra firme) supporting many larger trees with high wood density. The weak direct relationships we observed between soil and climate variables and AGB suggest that the most appropriate approaches to landscape scale modeling of AGB in the Amazon would be based on remote sensing methods to map stand structure.     

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.     

The impact of global climate change on tropical forest biodiversity in Amazonia

Aim To model long‐term trends in plant species distributions in response to predicted changes in global climate. Location Amazonia. Methods The impacts of expected global climate change on the potential and realized distributions of a representative sample of 69 individual Angiosperm species in Amazonia were simulated from 1990 to 2095. The climate trend followed the HADCM2GSa1 scenario, which assumes an annual 1% increase of atmospheric CO₂ content with effects mitigated by sulphate forcing. Potential distributions of species in one‐degree grid cells were modelled using a suitability index and rectilinear envelope based on bioclimate variables. Realized distributions were additionally limited by spatial contiguity with, and proximity to, known record sites. A size‐structured population model was simulated for each cell in the realized distributions to allow for lags in response to climate change, but dispersal was not included. Results In the resulting simulations, 43% of all species became non‐viable by 2095 because their potential distributions had changed drastically, but there was little change in the realized distributions of most species, owing to delays in population responses. Widely distributed species with high tolerance to environmental variation exhibited the least response to climate change, and species with narrow ranges and short generation times the greatest. Climate changed most in north‐east Amazonia while the best remaining conditions for lowland moist forest species were in western Amazonia. Main conclusions To maintain the greatest resilience of Amazonian biodiversity to climate change as modelled by HADCM2GSa1, highest priority should be given to strengthening and extending protected areas in western Amazonia that encompass lowland and montane forests.     

There's no place like home: seedling mortality contributes to the habitat specialisation of tree species across Amazonia

Understanding the mechanisms generating species distributions remains a challenge, especially in hyperdiverse tropical forests. We evaluated the role of rainfall variation, soil gradients and herbivory on seedling mortality, and how variation in seedling performance along these gradients contributes to habitat specialisation. In a 4‐year experiment, replicated at the two extremes of the Amazon basin, we reciprocally transplanted 4638 tree seedlings of 41 habitat‐specialist species from seven phylogenetic lineages among the three most important forest habitats of lowland Amazonia. Rainfall variation, flooding and soil gradients strongly influenced seedling mortality, whereas herbivory had negligible impact. Seedling mortality varied strongly among habitats, consistent with predictions for habitat specialists in most lineages. This suggests that seedling performance is a primary determinant of the habitat associations of adult trees across Amazonia. It further suggests that tree diversity, currently mostly harboured in terra firme forests, may be strongly impacted by the predicted climate changes in Amazonia.     

Low Primate Diversity and Abundance in Northern Amazonia and its Implications for Conservation

Large areas of the Rio Negro basin in Amazonia are covered by continuous tracts of tropical forest, but have few primate species. This is anomalous considering the general relationship between area and number of species. One possibility is that much of the forest is unsuitable habitat for most primates and the area of suitable habitat is much less than the forested area. This has important consequences for the design of reserves and predictions of the consequences of climate change, which tend to be based on broad categories based on satellite images, and not on information of species distributions within those broad categories. The study was conducted through diurnal and nocturnal line‐transect surveys in the Biodiversity Research Program 25‐km² permanent grid in Viruá National Park, which has vegetation associations typical of much of northern Amazonia. The highest primate diversity and abundances occurred in tall terra firme forests (58%), whereas inundated forests and scrublands, which cover 42 percent of the survey grid and 90.8 percent of the Viruá National Park, have virtually no primates. This suggests that parks and reserves in northwestern Amazonia will have to be very large to maintain viable populations of most primates and their ecological interactions, and that very broad habitat categories are not sufficient to make predictions about actual and future suitability of areas for primate conservation.     

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.