Short-Term Temporal Changes in Tree Live Biomass in a Central Amazonian Forest, Brazil

We monitored seventy-two 1 ha permanent plots spread over 64 km² of terra firme forest at Reserva Ducke (Manaus, Amazonas, Brazil) over 2-yr intervals to assess the effects of a soil and topographic gradient on the rate of change in the aboveground tree live biomass (AGLB). AGLB increased significantly over the 2-yr intervals, exhibiting a mean rate of change of 1.65 Mg/ha/yr (bootstrapped 95% CI: 1.15, 2.79). The rate of change varied according to tree size class; understory and sub-canopy trees exhibited higher rates of change. Over the whole period, the rate of change was not related to soil or topographic features of the plots, but there was evidence that the relationships varied depending on the year of measurement. In the plots monitored between 2001 and 2003 we found a significant relationship between AGLB change and the soil textural gradient, but this relationship was not evident in plots monitored between 2002 and 2004. This suggests that both the temporal variation in the soil–biomass change relationship and the size structure of the forest need to be included in models of biomass change in Amazonia. We also noted that the rate of biomass change is sensitive to the equation used to estimate AGLB. Allometric models that incorporate wood-density data provide higher per plot AGLB estimates, but lower rates of change, suggesting that variations in floristic composition have important implications for carbon cycling in diverse tropical forests.
Abstract in Portuguese is available at http://www.blackwell-synergy.com/loi/btp .     

Species richness,composition, and spatial distribution of vascular epiphytes in Amazonian black-water floodplain forests

This study examines the occurrence of vascular epiphytic species in Central Amazonian black-water floodplain forests (igapó) and considers whether their horizontal and vertical distribution is influenced by the flood pulse, as is the case with tree species (phorophytes). Research was conducted in sixteen forest plots the Jaú National Park. In these, epiphytes on all phorophytes with DBH ≥ 10 cm were identified. We measured flood height using the watermark left by the last high-water period, then estimated the height relative to the ground of every epiphytic individual. We recorded 653 individuals in 37 species, distributed on 109 phorophytes. Igapó floodplain forests have much lower richness and abundance of vascular epiphyte species than do other Amazonian forests. This may reflect the limitation of available sites for colonization (only 24.9% of studied trees were occupied by epiphytes). Holoepiphytes predominated, and the combined presence of a flood-pulse, linked to the nutrient-poor soil poor seems to limit the occurrence of nomadic vines. Horizontal distribution of epiphytes followed the distribution of phorophytes, which in turn followed the flood-level gradient. Also flooding interacted strongly with vertical zonation to determine species richness. As already well-reported for trees, and unlike reports of epiphytes in other floodplains, flooding strongly influenced richness and distribution of vascular epiphytes in the studied igapó forests.     

Pressurized gas transport in Amazonian floodplain trees

Trees of Central Amazonian whitewater floodplain forests are subjected to extended flood periods lasting several months. They must therefore be highly adapted to long-term flooding and to oxygen shortage in the soil and roots which is expected to be the main stress factor in this environment. The improvement of oxygen transport to the roots by pressurized gas transport, known for some tree species of the temperate zone, is one specific adaptation to flooding. To evaluate its significance, gas transport measurements were carried out under experimental conditions with saplings of five Amazonian tree species. Internal aeration of the roots was improved under conditions of pressurized gas transport as shown by polarographical measurements of oxygen exchange between root and rhizosphere. Tracer gas measurements using sulphur hexafluoride (SF₆) showed gas permeability of transport pathways between stem base and roots. Based on the data, we suppose that pressurized gas transport significantly contributes to internal aeration of roots in Amazonian floodplain saplings growing on the higher levels in the flooding gradient, with low water columns, and is an important adaptation for establishment in these temporarily inundated habitats also in the weeks of rising and lowering water levels.     

Impact of climatic and hydrological disturbances on blackwater floodplain forests in Central Amazonia

Extreme climatic events and anthropic disturbances affect the hydrological regime of Amazonian rivers and connected floodplain forests. This study aims to investigate the impacts of the Balbina hydroelectric power plant on the floodplain forests of the Uatumã River, in the Central Amazon. For this, tree age and diameter increment from the most abundant tree species of three different topographic levels were obtained and analyzed in the affected area downstream of the Balbina dam (Uatumã River) and compared to an undisturbed site (Abacate River, affluent) considering age structure and mean diameter increments between the topographic levels and the two systems. The occurrence of old trees is much lower at the disturbed site compared to the undisturbed system. Especially at the middle topography of the Uatumã site, we observed tree species with high mean diameter increment indicating a strong disturbance. We suggest that the disturbances may be associated with extreme hydro‐climatic events, such as extreme droughts that occurred during the El Niño years 1925/26 and 1982/83 and that these events may increase vulnerability of igapó floodplains to wildfires. Abstract in Portuguese is available with online material.     

Hardwood floodplain forests in Slovakia: syntaxonomical revision

Hardwood floodplain forests in Slovakia have been traditionally classified within the class Querco-Fagetea, order Fagetalia, alliance Alnion incanae, suballiance Ulmenion. Five main syntaxonomical studies on this topic come from the years 1958–1972 and were published by several authors from different regions. Each author described several new geographically delimited subassociations which were considered to be confined to single lowland region. This resulted in unclear classification scheme of hardwood floodplain forests in Slovakia with numerous floristically poorly differentiated units, especially at the subassociation level. We used 404 relevés ordered to the suballiance Ulmenion in the Slovak national database (http://ibot.sav.sk/cdf/) and analysed the dataset by cluster analysis and detrended correspondence analysis. We distinguished four groups of relevés which represents four subassociations within one association Fraxino pannonicae-Ulmetum: Fraxino pannonicae-Ulmetum caricetosum acutiformis Soó in Aszód 1936 corr. Soó 1964, Fraxino pannonicae-Ulmetum populetosum Jurko 1958 (D?atko 1972), Fraxino pannonicae-Ulmetum carpinetosum (Simon 1957) D?atko 1972 and Fraxino pannonicae-Ulmetum convallarietosum Jurko 1958. Associations Querco pedunculiflorae-Populetum tremuli ?om?ák et Háberová 1979, Arunco-Salicetum capreae Hada? et al. 1969 and Filipendulo-Salicetum capreae Hada? et al. 1969 we propose to exclude from the suballiance Ulmenion.     

Phytoplankton composition and abundance of a central Amazonian floodplain lake

The phytoplankton community of Lake Camaleão, a smallfloodplainlake influenced by a large whitewater river, the Solimões, was monthlyinvestigated for the composition and abundance of itsphytoplankton. The seasonal influence of the floodregime on biomass, species richness and diversity, andits relation with physical and chemical factors(temperature, pH, dissolved oxygen, electricalconductivity, total seston and inorganic nutrients)was analyzed and subjected to principal componentanalysis. Diversity was variable along the seasonalcycle: relatively high values were observed at the endof the dry season supported by high nutrientconcentrations. The phytoplankton was comprised of 262 taxa,with strong dominanceof euglenoids (81%). The three sample stations did not differamong each other, except in the dry season, due todata cluster in relation to theprincipal axis (1 and 2), explaining 63% of thevariation. Biomass accumulation as a function of lakearea reduction contributed to theseresults, indicating that the phytoplankton dynamicswere hydrology-driven.     

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.     

Impact of a drier Early-Mid-Holocene climate upon Amazonian forests

This paper uses a palaeoecological approach to examine the impact of drier climatic conditions of the Early-Mid-Holocene (ca 8000-4000 years ago) upon Amazonia's forests and their fire regimes. Palaeovegetation (pollen data) and palaeofire (charcoal) records are synthesized from 20 sites within the present tropical forest biome, and the underlying causes of any emergent patterns or changes are explored by reference to independent palaeoclimate data and present-day patterns of precipitation, forest cover and fire activity across Amazonia. During the Early-Mid-Holocene, Andean cloud forest taxa were replaced by lowland tree taxa as the cloud base rose while lowland ecotonal areas, which are presently covered by evergreen rainforest, were instead dominated by savannahs and/or semi-deciduous dry forests. Elsewhere in the Amazon Basin there is considerable spatial and temporal variation in patterns of vegetation disturbance and fire, which probably reflects the complex heterogeneous patterns in precipitation and seasonality across the basin, and the interactions between climate change, drought- and fire susceptibility of the forests, and Palaeo-Indian land use. Our analysis shows that the forest biome in most parts of Amazonia appears to have been remarkably resilient to climatic conditions significantly drier than those of today, despite widespread evidence of forest burning. Only in ecotonal areas is there evidence of biome replacement in the Holocene. From this palaeoecological perspective, we argue against the Amazon forest 'dieback' scenario simulated for the future.     

Tree and stand level variables influencing diversity of lichens on temperate broad-leaved trees in boreo-nemoral floodplain forests

Tree and stand level variables affecting the species richness, cover and composition of epiphytic lichens on temperate broad-leaved trees (Fraxinus excelsior, Quercus robur, Tilia cordata, Ulmus glabra, and U. laevis) were analysed in floodplain forest stands in Estonia. The effect of tree species, substrate characteristics, and stand and regional variables were tested by partial canonical correspondence analysis (pCCA) and by general linear mixed models (GLMM). The most pronounced factors affecting the species richness, cover and composition of epiphytic lichens are acidity of tree bark, bryophyte cover and circumference of tree stems. Stand level characteristics have less effects on the species richness of epiphytic lichens, however, lichen cover and composition was influenced by stand age and light availability. The boreo-nemoral floodplain forests represent valuable habitats for epiphytic lichens. As substrate-related factors influence the species diversity of lichens on temperate broad-leaved trees differently, it is important to consider the effect of each tree species in biodiversity and conservation studies of lichens. Nomenclature Randlane et al. (2007) for lichens; Leht (2007) for vascular plants.     

Dietary–morphological relationships in a fish assemblage of the Bolivian Amazonian floodplain

Morphological correlates of diet were examined in 48 species of freshwater fishes from floodplain lakes in the central part of the Mamoré River (Bolivian Amazon). The species were classified, according to the percentage occurrence of seven food items, into eight broad trophic categories: mud feeders, algivores, herbivores, terrestrial invertivores and omnivores, carnivores, zooplanktivores, aquatic invertivores and piscivores. There were significant relationships between the diet and morphology of the fishes even when the effect of taxonomical relatedness between species was eliminated. Relative gut length was the main morphological variable used to order species on a carnivore to mud feeder gradient. Standard length and head and mouth size were the morphological variables most closely associated with prey size. Mud feeder, algivore and piscivore species appeared as the most dietary and morphologically specialized. These results support both the hypotheses that species morphology influences the diet and that morphological similarity is conserved even in comparison with taxonomically unrelated species.     

Three fishes in one: cryptic species in an Amazonian floodplain forest specialist

Accurately describing biodiversity in tropical regions such as Amazonia is difficult because of insufficient morphological inventories and the lack of studies on the distribution of genetic diversity. Aquatic organisms from Amazonian flooded forests are generally expected to move laterally along the forests during the annual inundation cycle, a behaviour that should promote admixture of populations and reduce within‐drainage speciation. We used an unprecedented fine‐scale sampling effort and multiple DNA markers to quantify region‐wide population differentiation in an Amazonian floodplain forest specialist, the black‐wing hatchet fish Carnegiella marthae ( Myers, 1927 ). Our study revealed three previously unsuspected and ancient cryptic species of black‐wing hatchet fish in the Rio Negro floodplain (RNF), in central Amazonia. Two species produce occasional first‐generation hybrids. The third and rarer species, although found in extreme sympatry with another species, appears to be reproductively isolated, and also differs in external morphology and dentition. Our findings have important implications for guiding conservation management because C. marthae is harvested commercially in the RNF ornamental fishery. They also suggest that the diversity of Amazonian ichthyofauna is vastly underestimated, including that found in landscapes lacking contemporary barriers to account for population divergence and speciation. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 391–403.     

Adaptations of fish species to oxygen depletion in a central Amazonian floodplain lake

Pronounced seasonal and daily oxygen concentration changes are characteristic for Amazonian floodplain lakes. Studies on the fish fauna of the Lago Camaleão, Solimões River, Amazonas, Brazil, showed several fish species which are able to survive prolonged periods of heavy hypoxia. Twenty species belonging to eight families were observed in the laboratory in order to determine their respiratory adaptations to hypoxic conditions and oxygen concentrations at which the fish present respiratory adaptations. Finally, the fish species were distributed throughout the habitats of Lake Camaleão according to their adaptation responses. Ten fish species used the surface water for aquatic surface respiration, four species used atmospheric oxygen for aerial respiration, four species used oxygen supplied by the exudation of the roots of floating macrophytes and two exhibited a high tolerance to hypoxic conditions, and well-developed physiological biochemical mechanisms. The fish fauna is well adapted to low oxygen concentrations. The large variety of morpho-anatomical adaptations associated with biochemical and physiological mechanisms to tolerate hypoxic and anoxic conditions enable the 20 fish species to exploit several habitats of Lago Camaleão, such as floating aquatic macrophyte meadows, open water and near the shoreline.     

High-quality seed dispersal by fruit-eating fishes in Amazonian floodplain habitats

Seed dispersal is a critical stage in the life history of plants. It determines the initial pattern of juvenile distribution, and can influence community dynamics and the evolutionary trajectories of individual species. Vertebrate frugivores are the primary vector of seed dispersal in tropical forests; however, most studies of seed dispersal focus on birds, bats and monkeys. Nevertheless, South America harbors at least 200 species of frugivorous fishes, which move into temporarily flooded habitats during lengthy flood seasons and consume fruits that fall into the water; and yet, we know remarkably little about the quality of seed dispersal they effect. We investigated the seed dispersal activities of two species of large-bodied, commercially important fishes (Colossoma macropomum and Piaractus brachypomus, Characidae) over 3 years in Pacaya-Samiria National Reserve (Peru). We assessed the diet of these fishes during the flood season, conducted germination trials with seeds collected from digestive tracts, and quantified fruit availability. In the laboratory, we fed fruits to captive Colossoma, quantified the proportion of seeds defecated by adult and juvenile fish, and used these seeds in additional germination experiments. Our results indicate that Colossoma and Piaractus disperse large quantities of seeds from up to 35% of the trees and lianas that fruit during the flood season. Additionally, these seeds can germinate after floodwaters recede. Overexploitation has reduced the abundance of our focal fish species, as well as changed the age structure of populations. Moreover, older fish are more effective seed dispersers than smaller, juvenile fish. Overfishing, therefore, likely selects for the poorest seed dispersers, thus disrupting an ancient interaction between seeds and their dispersal agents. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Geological control of floristic composition in Amazonian forests

AIM: Conservation and land-use planning require accurate maps of patterns in species composition and an understanding of the factors that control them. Substantial doubt exists, however, about the existence and determinants of large-area floristic divisions in Amazonia. Here we ask whether Amazonian forests are partitioned into broad-scale floristic units on the basis of geological formations and their edaphic properties. LOCATION: Western and central Amazonia. METHODS: We used Landsat imagery and Shuttle Radar Topography Mission (SRTM) digital elevation data to identify a possible floristic and geological discontinuity of over 300 km in northern Peru. We then used plant inventories and soil sampling to document changes in species composition and soil properties across this boundary. Data were obtained from 138 sites distributed along more than 450 km of road and river. On the basis of our findings, we used broad-scale Landsat and SRTM mosaics to identify similar patterns across western and central Amazonia. RESULTS: The discontinuity identified in Landsat and SRTM data corresponded to a 15-fold change in soil cation concentrations and an almost total change in plant species composition. This discontinuity appears to be caused by the widespread removal of cation-poor surface sediments by river incision to expose cation-rich sediments beneath. Examination of broad-scale Landsat and SRTM mosaics indicated that equivalent processes have generated a north-south discontinuity of over 1500 km in western Brazil. Due to similarities with our study area, we suggest that this discontinuity represents a chemical and ecological limit between western and central Amazonia. MAIN CONCLUSIONS: Our findings suggest that Amazonian forests are partitioned into large-area units on the basis of geological formations and their edaphic properties. The evolution of these units through geological time may provide a general mechanism for biotic diversification in Amazonia. These compositional units, moreover, may correspond to broad-scale functional units. The existence of large-area compositional and functional units would suggest that protected-area, carbon sequestration, and other land-use strategies in Amazonia be implemented on a region-by-region basis. The methods described here can be used to map these patterns, and thus enable effective conservation and management of Amazonian forests.     

Seasonal movements of river dolphins (Inia geoffrensis) in a protected Amazonian floodplain

Deliberate killing for use as bait in a regional catfish (Calophysus macropterus) fishery is the primary threat affecting the survival of the Amazon river dolphin, or boto (Inia geoffrensis). Establishing and improving freshwater protected areas has been suggested as a possible course of action to protect the species. However, the ecology of the boto is poorly understood and more information is needed on the species’ habitat use and movement patterns to ensure that spatial protection initiatives meet conservation needs. In this study, mark‐recapture/resight data collected in and near the Mamirauá Sustainable Development Reserve, a várzea in the Brazilian Amazon, was used to examine transition probabilities of botos between various habitat types. Our findings suggest that it is imperative that spatial protection initiatives focused on the boto take into account the importance of várzea habitat and bay systems for individuals in early life stages. Moreover, because botos frequently occupy the main rivers, particularly during the low water period, protecting areas of the main river waters adjacent to várzeas could aid in the protection of hunted populations.     

The role of fungi in periodically inundated Amazonian forests

In a comparative survey the terms (terra-firme) flood plain, igapó, várzea with and without black water influence are reviewed and the importance of ectotrophic mycorrhiza for their definition emphasized. Várzea alta with fluvisol base is briefly characterized and a first list of Basidiomycetes collected in it is provided. These are all non-mycorrhizal litter fungi.     

Lowland tapirs facilitate seed dispersal in degraded Amazonian forests

The forests of southeastern Amazonia are highly threatened by disturbances such as fragmentation, understory fires, and extreme climatic events. Large‐bodied frugivores such as the lowland tapir (Tapirus terrestris) have the potential to offset this process, supporting natural forest regeneration by dispersing a variety of seeds over long distances to disturbed forests. However, we know little about their effectiveness as seed dispersers in degraded forest landscapes. Here, we investigate the seed dispersal function of lowland tapirs in Amazonian forests subject to a range of human (fire and fragmentation) and natural (extreme droughts and windstorms) disturbances, using a combination of field observations, camera traps, and light detection and ranging (LiDAR) data. Tapirs travel and defecate more often in degraded forests, dispersing much more seeds in these areas [9,822 seeds per ha/year (CI_95% = 9,106; 11,838)] than in undisturbed forests [2,950 seeds per ha/year (CI_95% = 2,961; 3,771)]. By effectively dispersing seeds across disturbed forests, tapirs may contribute to natural forest regeneration—the cheapest and usually the most feasible way to achieve large‐scale restoration of tropical forests. Through the dispersal of large‐seeded species that eventually become large trees, such frugivores also contribute indirectly to maintaining forest carbon stocks. These functions may be critical in helping tropical countries to achieve their goals to maintain and restore biodiversity and its ecosystem services. Ultimately, preserving these animals along with their habitats may help in the process of natural recovery of degraded forests throughout the tropics. Abstract in Portuguese is available with online material.