Long-term forest-savannah dynamics in the Bolivian Amazon: implications for conservation

The aim of this paper is to evaluate the respective roles of past changes in climate, geomorphology and human activities in shaping the present-day forest-savannah mosaic of the Bolivian Amazon, and consider how this palaeoecological perspective may help inform conservation strategies for the future. To this end, we review a suite of palaeoecological and archaeological data from two distinct forest-savannah environments in lowland Bolivia: Noel Kempff Mercado National Park (NKMNP) on the Precambrian Shield and the 'Llanos de Moxos' in the Beni basin. We show that they contain markedly contrasting legacies of past climatic, geomorphic and anthropogenic influences between the last glacial period and the Spanish Conquest. In NKMNP, increasing precipitation caused evergreen rainforest expansion, at the expense of semi-deciduous dry forest and savannahs, over the last three millennia. In contrast, pre-Hispanic indigenous cultures were instrumental in facilitating recent forest expansion in the Llanos de Moxos by building a vast network of earthworks. Insights from Mid-Holocene palaeodata, together with ecological observations and modelling studies, suggest that there will be progressive replacement of rainforest by dry forest and savannah in NKMNP over the twenty-first century in response to the increased drought predicted by general circulation models. Protection of the latitudinal landscape corridors may be needed to facilitate these future species reassortments. However, devising appropriate conservation strategies for the Llanos de Moxos will be more difficult due to its complex legacy of Palaeo-Indian impact. Without fully understanding the degree to which its current biota has been influenced by past native cultures, the type and intensity of human land use appropriate for this landscape in the future will be difficult to ascertain.     

Impacts of selective logging on inbreeding and gene flow in two Amazonian timber species with contrasting ecological and reproductive characteristics

Selective logging in Brazil allows for the removal of up to 90% of trees above 50 cm diameter of a given timber species, independent of a species’ life history characteristics or how quickly it will recover. The genetic and demographic effects of selective logging on two Amazonian timber species (Dipteryx odorata Leguminosae, Jacaranda copaia Bignoniaceae) with contrasting ecological and reproductive characteristics were assessed in the same forest. Genetic diversity and gene flow were characterized by genotyping adults and seed sampled before and after logging, using hypervariable microsatellite markers. Overall, there were no short‐term genetic impacts on the J. copaia population, with commercial application of current Brazilian forest management regulations. In contrast, for D. Odorata, selective logging showed a range of genetic impacts, with a 10% loss of alleles, and reductions in siring by pollen from trees within the 546‐ha study area (23–11%) and in the number of pollen donors per progeny array (2.8–1.6), illustrating the importance of the surrounding landscape. Asynchrony in flowering between D. odorata trees led to trees with no breeding partners, which could limit the species reproduction and regeneration under current regulations. The results are summarized with other published studies from the same site and the implications for forest management discussed. The different types and levels of impacts associated with each species support the idea that ecological and genetic information by species, ecological guild or reproductive group is essential in helping to derive sustainable logging guidelines for tropical forests.     

Development and characterization of microsatellite markers for the Amazonian blackwing hatchetfish,Carnegiella marthae (Teleostei,Gasteropelecidae)

The blackwing hatchetfish, Carnegiella marthae, is a small characin species distributed in forest streams of the Negro and upper Orinoco River basins in Amazonia. Freshwater hatchetfish are popular in the aquarium trade and represent an economic resource for the riverine people from middle Rio Negro, in Brazil. We isolated and characterized seven microsatellite DNA loci for the blackwing hatchetfish. Number of alleles and heterozygosity per locus in a sample of 30 fish ranged from three to 17 and from 0.19 to 0.87, respectively. These microsatellite loci provide powerful markers for studies on taxonomy, management and phylogeographic history of Amazonian hatchetfish.     

Floral morphology and taxonomic relations among the genera of Amaranthaceae in the New World and the Hawaiian Islands

Twenty-three genera of Amaranthaceae occur in the New World. Two endemic genera occur in the Hawaiian Islands. Among the genera of the subfamily Amaranthoideae, Celosia, Cyathula and Achyranthes have their main distributions in the Old World; the two last-named genera are represented in the Americas only by widespread weeds. All the New World genera of the subfamily Gomphrenoideae are mainly or entirely restricted to this region. Characters of androecium and gynoecium are fundamental in the recognition of genera within the family. Androecia of different genera may be structurally and phylogenetically more similar than would appear from a cursory examination. It is suggested that the type of staminal tubes found in Pseudogomphrena and Froelichia can be derived from that in Alternanthera and Froelichiella by reduction of filament length and a fusion of pseudostaminodia with the filaments. The staminal tube in Gomphrena could result from a further decrease in distance between pseudostaminodia of the Pseudogomphrena type, and a deeper forking of the pseudostaminodia; each so-called apical filament lobe in Gomphrena would then be homologous with half a pseudostaminodium in Pseudogomphrena. Much of the variation in the androecia of these and other genera, as well as within genera such as Pfaffia, can be explained as the combined results of coalescence and splitting-up tendencies. Splitting up of staminal tubes may not necessarily take place along the borders of phylogenetically original filaments and pseudostaminodia. The Amaranthus-type of pollen is found in the majority of genera of the subfamily Amaranthoideae, but also in the Chenopodiaceae. A group of genera within the subfamily Gomphrenoideae also has pollen very similar to, or identical with, this type. Most genera of the subfamily Gomphrenoideae have pollen of the Gomphrena-type. Pseudoplantago has unilocular (at anthesis) anthers, a characteristic of the subfamily Gomphrenoideae, but floral structure as well as pollen morphology connect the genus to a group of genera within the Amaranthoideae, subtribe Achyranthinae. The combination of subcuboidal shape and opercula with radially arranged hooked protuberances, makes the pollen of Pseudoplantago unique among the angiosperms studied so far. Floral morphology and palynological characteristics indicate a close relationship between Pfaffia and Alternanthera. Both genera, as currently accepted, are relatively homogeneous from pollen morphological points of view. There are no correlations between pollen morphology and the variation in the androecium in Pfaffia, nor would pollen structure support recognition of Hebanthe as a distinct genus. Woehleria and Irenella may be derived from, or be of the same origin as, Dicraurus and Iresine. All four genera are placed in the subfamily Gomphrenoideae because of the bisporangiate anthers, but their pollen structure is very close to, or identical with, that of the Amaranthus-type. Pseudogomphrena combines characteristics of Gomphrena and Pfaffia.     

Characterization of 10 microsatellite markers for the understorey Amazonian herb Heliconia acuminata

We characterized 10 microsatellite loci for the plant Heliconia acuminata from the Biological Dynamics of Forest Fragments Project (Manaus, Brazil). Markers were screened in 61 individuals from one population and were found to be polymorphic with an average of eight alleles per locus. We found moderate to high levels of polymorphic information content, and observed and expected heterozygosities. All 10 markers are suitable for spatial genetic structure and parentage analyses and will be used for understanding H. acuminata dynamics across a fragmented landscape.     

Characterization of microsatellite loci for the Amazonian rummy‐nose tetra,Hemigrammus bleheri (Teleostei,Characidae)

The rummy‐nose tetra, Hemigrammus bleheri, is a small fish found in forest streams of the Rio Negro floodplain, central Amazonia. This species is popular in aquaria and used as an ornamental fishery resource by the riverine people from middle Rio Negro. Here we describe eight microsatellite DNA loci for rummy‐nose tetras. Number of alleles and heterozygosity per locus in a sample of 30 fish ranged from five to 22 and from 0.52 to 0.96, respectively. These highly variable genetic markers provide important tools for investigating population history and identifying conservation units in rummy‐nose tetras.     

Warming,drought, and disturbances lead to shifts in functional composition: A millennial-scale analysis for Amazonian and Andean sites

Tropical forests are changing in composition and productivity, probably in response to changes in climate and disturbances. The responses to these multiple environmental drivers, and the mechanisms underlying the changes, remain largely unknown. Here, we use a functional trait approach on timescales of 10,000 years to assess how climate and disturbances influence the community-mean adult height, leaf area, seed mass, and wood density for eight lowland and highland forest landscapes. To do so, we combine data of eight fossil pollen records with functional traits and proxies for climate (temperature, precipitation, and El Niño frequency) and disturbances (fire and general disturbances). We found that temperature and disturbances were the most important drivers of changes in functional composition. Increased water availability (high precipitation and low El Niño frequency) generally led to more acquisitive trait composition (large leaves and soft wood). In lowland forests, warmer climates decreased community-mean height probably because of increased water stress, whereas in highland forests warmer climates increased height probably because of upslope migration of taller species. Disturbance increased the abundance of acquisitive, disturbance-adapted taxa with small seeds for quick colonization of disturbed sites, large leaves for light capture, and soft wood to attain fast height growth. Fire had weak effects on lowland forests but led to more stress-adapted taxa that are tall with fast life cycles and small seeds that can quickly colonize burned sites. Site-specific analyses were largely in line with cross-site analyses, except for varying site-level effects of El Niño frequency and fire activity, possibly because regional patterns in El Niño are not a good predictor of local changes, and charcoal abundances do not reflect fire intensity or severity. With future global changes, tropical Amazonian and Andean forests may transition toward shorter, drought- and disturbance-adapted forests in the lowlands but taller forests in the highlands.     

First report on new evidence for the occurrence of Podocarpus and possible human presence at the mouth of the Amazon during the Late-glacial

Palynological studies on late Quaternary lake sediments from the region of the Amazon estuary, 100 km north-east of Belém, Pará State, Brazil, enable reconstruction of lowland Amazonian rain forest during the Late-glacial and Holocene periods. Late-glacial forests included populations of Podocarpus which suggests a distinct climatic cooling. Ilex was abundant in the early Holocene. Records of the mangrove taxon, Rhizophora, indicate rapid Atlantic sea-level rise in the beginning of the Holocene. High charcoal representation may reflect the first arrival of Amerindians in the Amazon coastal area, probably about 10 800 B.P.     

Phylogeography of the piranha genera Serrasalmus and Pygocentrus: implications for the diversification of the Neotropical ichthyofauna

The phylogenetic relationships within the piranhas were assessed using mitochondrial sequences with the aim of testing several hypotheses proposed to explain the origin of Neotropical diversity (palaeogeography, hydrogeology and museum hypotheses). Sequences of the ribosomal 16S gene (510 bp) and control region (980 bp) were obtained from 15 localities throughout the main South American rivers for 21 of the 28 extant piranha species. The results indicate that the genus Serrasalmus is monophyletic and comprises three major clades. The phylogeographical analyses of these clades allowed the identification of five vicariant events, extensive dispersal and four lineage duplications suggesting the occurrence of sympatric speciation. Biogeographical patterns are consistent with the prediction made by the museum hypothesis that lineages from the Precambrian shields are older than those from the lowlands of the Amazon. The vicariant events inferred here match the distribution of the palaeoarches and several postdispersal speciation events are identified, thereby matching the predictions of the palaeogeography and hydrogeology hypotheses, respectively. Molecular clock calibration of the control region sequences indicates that the main lineages differentiated from their most recent common ancestor at 9 million years ago in the proto Amazon-Orinoco and the present rate of diversification is the highest reported to date for large carnivorous Characiformes. The present results emphasize that an interaction among geology, sea-level changes, and hydrography created opportunities for cladogenesis in the piranhas at different temporal and geographical scales.     

Teleconnection between tree growth in the Amazonian floodplains and the El Niño–Southern Oscillation effect

There is a limited knowledge about the El Niño–Southern Oscillation (ENSO) effects on the Amazon basin, the world's largest tropical rain forest and a major factor in the global carbon cycle. Seasonal precipitation in the Andean watershed annually causes a several month‐long inundation of the floodplains along the Amazon River that induces the formation of annual rings in trees of the flooded forests. Radial growth of trees is mainly restricted to the nonflooded period and thus the ring width corresponds to its duration. This allows the construction of a tree‐ring chronology of the long‐living hardwood species Piranhea trifoliata Baill. (Euphorbiaceae). El Niño causes anomalously low precipitation in the catchment that results in a significantly lower water discharge of the Amazon River and consequently in an extension of the vegetation period. In those years tree rings are significantly wider. Thus the tree‐ring record can be considered as a robust indicator reflecting the mean climate conditions of the whole Western Amazon basin. We present a more than 200‐year long chronology, which is the first ENSO‐sensitive dendroclimatic proxy of the Amazon basin and permits the dating of preinstrumental El Niño events. Time series analyses of our data indicate that during the last two centuries the severity of El Niño increased significantly.     

Environment versus dispersal in the assembly of western Amazonian palm communities

Aim It is a central issue in ecology and biogeography to understand what governs community assembly and the maintenance of biodiversity in tropical rain forest ecosystems. A key question is the relative importance of environmental species sorting (niche assembly) and dispersal limitation (dispersal assembly), which we investigate using a large dataset from diverse palm communities. Location Lowland rain forest, western Amazon River Basin, Peru. Methods We inventoried palm communities, registering all palm individuals and recording environmental conditions in 149 transects of 5 m × 500 m. We used ordination, Mantel tests and indicator species analysis (ISA) to assess compositional patterns, species responses to geographical location and environmental factors. Mantel tests were used to assess the relative importance of geographical distance (as a proxy for dispersal limitation) and environmental differences as possible drivers of dissimilarity in palm species composition. We repeated the Mantel tests for subsets of species that differ in traits of likely importance for habitat specialization and dispersal (height and range size). Results We found a strong relationship between compositional dissimilarity and environmental distance and a weaker but also significant relationship between compositional dissimilarity and geographical distance. Consistent with expectations, relationships with environmental and geographical distance were stronger for understorey species than for canopy species. Geographical distance had a higher correlation with compositional dissimilarity for small‐ranged species compared with large‐ranged species, whereas the opposite was true for environmental distance. The main environmental correlates were inundation and soil nutrient levels. Main conclusions The assembly of palm communities in the western Amazon appears to be driven primarily by species sorting according to hydrology and soil, but with dispersal limitation also playing an important role. The importance of environmental characteristics and geographical distance varies depending on plant height and geographical range size in agreement with functional predictions, increasing our confidence in the inferred assembly mechanisms.     

Implications of collection patterns of botanical specimens on their usefulness for conservation planning: an example of two neotropical plant families (Moraceae and Myristicaceae) in Peru

We evaluated the usefulness of herbarium collection databases for assessing patterns of species diversity and distribution based on a dataset from the flowering plant families Moraceae and Myristicaceae from the Peruvian Amazon. For Moraceae, a total of 3523 collections were used representing 134 species. The Myristicaceae were represented by 2113 collections of 46 species. We evaluated the distribution of collections based on 252 grid cells (0.5° size) covering all lowland rainforest in the Peruvian Amazon. We found that collections were concentrated in a few cells and that species diversity clearly increases in relation to collection density. Moraceae were collected in only 45% and Myristicaceae in only 31% of the 252 grid cells. Fifty percent of the collections came from just six and three cells, respectively. Most species were represented by only a small number of collections and collected only in a few grid cells, meaning a few widespread common species tend to dominate the collection records. Not surprisingly, most collections were made close to towns and transport routes. We evaluated the usefulness of rarefaction curves and diversity estimators for comparing diversity between regions. These techniques seem to be of little use for botanical collections due to violations of underlying assumptions. Problems such as accuracy of geographic and taxonomic data and strong bias in the spatial representation of the whole dataset are important to consider when basing conservation analysis, planning, and decision-making on seemingly large databases of biodiversity collections and are discussed in detail.     

A 7000‐year history of changing plant trait composition in an Amazonian landscape; the role of humans and climate

Tropical forests are shifting in species and trait composition, but the main underlying causes remain unclear because of the short temporal scales of most studies. Here, we develop a novel approach by linking functional trait data with 7000 years of forest dynamics from a fossil pollen record of Lake Sauce in the Peruvian Amazon. We evaluate how climate and human disturbances affect community trait composition. We found weak relationships between environmental conditions and traits at the taxon level, but strong effects for community‐mean traits. Overall, community‐mean traits were more responsive to human disturbances than to climate change; human‐induced erosion increased the dominance of dense‐wooded, non‐zoochorous species with compound leaves, and human‐induced fire increased the dominance of tall, zoochorous taxa with large seeds and simple leaves. This information can help to enhance our understanding of forest responses to past environmental changes, and improve predictions of future changes in tropical forest composition.     

Late Quaternary vegetation and climate change in the Panama Basin: Palynological evidence from marine cores ODP 677B and TR 163-38

The Late Quaternary paleoenvironmental history from Pacific slopes of the western Andes is reconstructed by pollen analysis of 32 samples from two marine sediment cores from the Panama Basin, eastern equatorial Pacific: core ODP 677B (83°44.2200′ W, 1°12.1440′ N, 3473 m water depth) is 185 cm long and spans the last 39,410 years, core TR 163-38 (81.583° W, 1.337° N, 2200 m water depth) is 103 cm long and covers the last 17,380 years. Six ecological groups were established: mangrove, brackish and fresh water swamps, terra firma lowland forests, broad range taxa, Andean forests, and open vegetation. A good correspondence was found between the changes of these ecological groups in the two cores. The records evidence the continuous presence of all vegetation types during the last 39,410 years and specially the uninterrupted occurrence of tropical rain forest. They record a development from: (1) a cold and humid phase (39,410-28,120 yr cal BP) with moderately high sea levels, (2) the coldest and driest phase in the record (28,120-14,500 yr cal BP) accompanied by the lowest sea levels, (3) a transitional phase when sea level rose and humid conditions dominated, (4) a stage (11,300-5600 yr cal BP) of the highest sea levels and moisture conditions including a drier period ∼7000 yr BP, to (5) a final period (5600 yr cal BP-Present) when sea level reached its present height, humidity persisted, and indicators of disturbance expanded. Peaks in pollen and spore concentration, associated with high river discharge periods, indicate periods of higher precipitation around 33,500, 28,000 and 12,000-9000 yr cal BP. Although main vegetation responses seem to reflect rainfall and moisture variations, a good correspondence was found between δ¹⁸O values and percentages of Andean and lowland pollen, suggesting that vegetation also responded to tempearture changes.     

A set of microsatellite DNA markers for the one‐lined pencilfish Nannostomus unifasciatus,an Amazonian flooded forest fish

The one‐lined pencilfish Nannostomus unifasciatus is a small fish from the flooded forests of the Amazon basin. Pencilfish are popular in aquaria and are used as ornamental fishery resource by riverine communities from central Amazonia. Here we report a set of nine microsatellite loci for N. unifasciatus. Number of alleles and heterozygosity per locus in a sample of 30 pencilfish ranged from 4 to 19 and from 0.39 to 0.86, respectively. These markers will prove useful to investigate population genetic structure, identify conservation units, and to conduct phylogeographical reconstructions in pencilfish from the flooded forests of Amazonia.     

Spatial organization and population density of the fish community of the litter banks within a central Amazonian blackwater stream

Results of a study of the fish community of the leaf litter banks in the lower reaches of the Taruma-Mirim, a small central Amazonian stream is presented. The Taruma-Mirim is an acidic, blackwater, forest stream which flows through igapo forest and enters the Rio Negro close to Manaus. The commonly held view that these blackwaters hold an impoverished ichthyofauna is challenged. The studied litter banks hold about 20 species of fish, all of which were found to live within distinct subregions of the habitat. Average fish density was about 100 individuals m⁻². Absolute population sizes were found to be remarkably small; within a 200 m² area the most abundant species had a population size of about 10⁴ and the least abundant <10² individuals. It is argued that high species richness linked to specialized habitat requirements and small population size indicates considerable population stability. The factors leading to the evolution of such species richness are discussed.     

Vegetation cover of Brazil in the last 21 ka: New insights into the Amazonian refugia and Pleistocenic arc hypotheses

Aim

The two main hypotheses about the Neotropical palaeovegetation, namely that of Amazonian refugia by Haffer and of the Pleistocene arc by Prado and Gibbs, are still constantly debated. We offer new insights on this debate using ecological niche modelling with combined climate–soil predictors to test both hypotheses, reconstruct the palaeovegetation of the Last Glacial Maximum (LGM; 21 ka) and Mid‐Holocene (Mid‐H; 6 ka) and indicate the configuration of refugia areas.

Location

Brazil.

Time period

Last 21 ka.

Major taxa studied

Biomes.

Methods

We modelled the environmental space of the 10 most representative biomes with the RandomForest classifier, using climate predictors from three atmospheric general circulation models (CCSM4, MPI‐ESM‐P and MIROC‐ESM) and soil predictors, the same for the different situations. Based on the consensus among the models, we reconstructed the palaeovegetation cover for LGM and Mid‐H and used fossil pollen sites to validate the reconstructions in a direct comparison.

Results

The climate in the past was cooler and wetter throughout most of the territory. The Amazon basin region was the most affected by climate change in the last 21 ka, with equatorial rain forest retracting to refugia areas, while the tropical rain forest (with climatic preferences similar to the Atlantic forest) expanded in the basin. In southern Brazil, the mixed forest (Araucaria forest) shifted to lower latitudes, while the grasslands expanded. In most biomes, the greatest changes occurred in the ecotonal zones, supported by pollen fossils.

Main conclusions

With regard to Haffer's hypothesis, the forests of the Amazonian lowlands retreated to refugia areas, while the colder and wetter climate of the basin created a favourable niche for another type of forest, instead of savanna. The advance of dry vegetation was restricted to ecotonal conditions, preventing the formation of a continuous Pleistocene arc, predicted by Prado and Gibbs's hypothesis.