How to survive six months in a flooded soil: Strategies in Chilopoda and Symphyla from Central Amazonian floodplains∗

Annual, long‐term inundation of flood waters for 5–7 months has led to the development of survival strategies in terrestrial invertebrates of Central Amazonian floodplains. Strategies observed are given for terricolous and arboricolous species and include migrants and non‐migrants. Subadult and advanced juvenile stages of the symphylan Ribauliella amazonica Scheller pass inundation in a dormant state inside tree roots in the flooded soil. Reproduction in this univoltine species is restricted to the non‐inundation period. R. amazonica represents the terricolous non‐migrating invertebrate guild with dormant stages in naturally available retreats under water during annual inundation. The undescribed parthenogenetic centipede Lamyctes sp. is found to hatch from flood‐resistant eggs at the beginning of non‐inundation periods. Development is fast, with the first hemi‐edaphic post‐larvae being captured 6–8 weeks after the floods have receded. Their surface structure does not protect the strongly elevated stigmata from lethal inundation. Lamyctes sp. represents the terricolous non‐migrating invertebrate guild, with dormant egg stages under water during annual inundation.     

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.     

Central Amazon Floodplain Forests: Root Adaptations to Prolonged Flooding

The floodplains of Central Amazonia represent a complex system of inundated river valleys and shallow lakes along the Solimões–Amazonas river, which is subjected to an annual flood-pulse lasting up to 10 months. Such flooding reaching an amplitude of about ten meters causes dramatic changes in the bioavailability of nutrients and oxygen levels and poses extreme constraints for plant survival and reproductivity. Tree species of inundation forests in Central Amazonia had to evolve adaptive mechanisms to both desiccation of soils and partial or full submergence. To adapt to flooded conditions, some trees overcome the flood period by dormancy accompanied by defoliation and formation of annual rings in the wood. Other species maintain metabolism and retain the foliage during the flooding, representing another adaptive mechanism to low oxygen availability. This investigation focused on the root physiology and morphology of six species typical of white-water inundation areas (várzea) led to a preliminary classification of adaptive strategies of trees inhabiting forest communities in floodplains of the Amazon basin.     

Climate and geographic distance are more influential than rivers on the beta diversity of passerine birds in Amazonia

Variation in the spatial structure of communities in terms of species composition (beta diversity) is affected by different ecological processes, such as environmental filtering and dispersal limitation. Large rivers are known as barriers for species dispersal (riverine hypothesis) in tropical regions. However, when organisms are not dispersal limited by geographic barriers, other factors, such as climatic conditions and geographic distance per se, may affect species distribution. In order to investigate the relative contribution of major rivers, climate and geographic distance on Passeriformes beta diversity, we divided Amazonia into 549 grid cells (1° of latitude and longitude) and obtained data of species occurrence, climate and geographic position for each cell. Beta diversity was measured using taxonomic, phylogenetic and functional metrics of composition. The influence of climatic variables, geographic distance and rivers on these metrics was tested using regression analyses. Passerine beta diversity is characterized mainly by the change in species taxonomic identity and in phylogenetic lineages across climatic gradients and over geographic distance. However, species with similar traits are found throughout the entire Amazonia. The size of rivers was proportional to their effect on species composition. However, climate and geographic distance are relatively more important than rivers for Amazonian taxonomic and phylogenetic species composition.     

Isolation by distance,not rivers,control the distribution of termite species in the Amazonian rain forest

The spatial distribution of species is affected by dispersal barriers, local environmental conditions and climate. However, the effect of species dispersal and their adaptation to the environment across geographic scales is poorly understood. To investigate the distribution of species from local to broad geographic scales, we sampled termites in 198 transects distributed in 13 sampling grids in the Brazilian Amazonian forest. The sampling grids encompassed an area of 271 500 km² and included the five major biogeographic regions delimited by Amazonian rivers. Environmental data for each transect were obtained from local measurements and remote sensing. Similar to previous studies, termite species composition at the local scale was mostly associated with measures of soil texture and chemistry. In contrast, termite species composition at broad geographic scales was associated with soil nutrients, and the geographic position of the transects. Between 17 and 30% of the variance in termite species composition could be attributed exclusively to the geographic position of the transects, but could not be attributed to measured environmental variables or the presence of major rivers. Isolation by distance may have strong effects on termite species composition due to historic processes and the spatially structured environments along distinct geological formations of Amazonia. However, in contrast to many taxa in Amazonia, there is no evidence that major rivers are important barriers to termite dispersal.     

Central Amazonian floodplain forests: Tree adaptations in a pulsing system

Amazonian floodplain forests are characterized by an annual flood pulse with changes of the water table that exceed 10 meters. Seedlings and adult trees are waterlogged or submerged for continuous periods lasting up to seven months per year. The monomodal flood pulse of the rivers causes drastic changes in the bioavailability of nutrients, oxygen levels, and concentrations of phytotoxins. The aquatic phase occurs during a period in which temperature and light conditions are optimal for plant growth and development, implying the need for adaptations. Not only do trees persist in a dormant state, they grow vigorously during most of the year, including the aquatic period. The regularity of flooding may have enhanced the evolution of specific traits, which partially are well known from floodplain trees in other tropical and in temperate regions. Different kinds of adaptations are found at the level of structural, physiological, and phenological traits. Combinations of adaptations regarding seed germination, seedling development, and traits of roots, shoots, and leaves result in a variety of growth strategies among trees. These lead to specific species distributions and zonations along the flooding gradient and within Amazonian floodplain systems (nutrient-rich white-water várzea and nutrient-poor black-water igapó).     

Extremely long-distance seed dispersal by an overfished Amazonian frugivore

Throughout Amazonia, overfishing has decimated populations of fruit-eating fishes, especially the large-bodied characid, Colossoma macropomum. During lengthy annual floods, frugivorous fishes enter vast Amazonian floodplains, consume massive quantities of fallen fruits and egest viable seeds. Many tree and liana species are clearly specialized for icthyochory, and seed dispersal by fish may be crucial for the maintenance of Amazonian wetland forests. Unlike frugivorous mammals and birds, little is known about seed dispersal effectiveness of fishes. Extensive mobility of frugivorous fish could result in extremely effective, multi-directional, long-distance seed dispersal. Over three annual flood seasons, we tracked fine-scale movement patterns and habitat use of wild Colossoma, and seed retention in the digestive tracts of captive individuals. Our mechanistic model predicts that Colossoma disperses seeds extremely long distances to favourable habitats. Modelled mean dispersal distances of 337-552 m and maximum of 5495 m are among the longest ever reported. At least 5 per cent of seeds are predicted to disperse 1700-2110 m, farther than dispersal by almost all other frugivores reported in the literature. Additionally, seed dispersal distances increased with fish size, but overfishing has biased Colossoma populations to smaller individuals. Thus, overexploitation probably disrupts an ancient coevolutionary relationship between Colossoma and Amazonian plants.     

The interplay of dispersal limitation,rivers, and historical events shapes the genetic structure of an Amazonian frog

Disentangling the impact of landscape features such as rivers and historical events on dispersal is a challenging but necessary task to gain a comprehensive picture of the evolution of diverse biota such as that found in Amazonia. Adenomera andreae, a small, territorial, terrestrial frog species of the Amazonian forest represents a good model for such studies. We combined cytochrome b sequences with 12 microsatellites to investigate the genetic structure at two contrasted spatial scales in French Guiana: along a ～6‐km transect, to evaluate dispersal ability, and between paired bank populations along a ～65‐km stretch of the Approuague river, to test the effect of rivers as barriers to dispersal. We observed significant spatial genetic structure between individuals at a remarkably small geographical scale, and conclude that the species has a restricted dispersal ability that is probably tied to its life‐history traits. Mitochondrial and microsatellite data also indicate a high level of differentiation among populations on opposite banks of the river, and, in some cases, among populations on the same riverbank. These results suggest that the observed population structure in A. andreae is the result of restricted dispersal abilities combined with the action of rivers and Quaternary population isolation. Given that Amazonia hosts a great portion of anurans, as well as other small vertebrates, that display life‐history traits comparable with A. andreae, we argue that our analyses provide new insights into the complex interactions among evolutionary processes shaping Amazonian biodiversity. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 356–373.     

Flying shells: historical dispersal of marine snails across Central America

The geological rise of the Central American Isthmus separated the Pacific and the Atlantic oceans about 3 Ma, creating a formidable barrier to dispersal for marine species. However, similar to Simpson's proposal that terrestrial species can 'win sweepstakes routes'-whereby highly improbable dispersal events result in colonization across geographical barriers-marine species may also breach land barriers given enough time. To test this hypothesis, we asked whether intertidal marine snails have crossed Central America to successfully establish in new ocean basins. We used a mitochondrial DNA genetic comparison of sister snails (Cerithideopsis spp.) separated by the rise of the Isthmus. Genetic variation in these snails revealed evidence of at least two successful dispersal events between the Pacific and the Atlantic after the final closure of the Isthmus. A combination of ancestral area analyses and molecular dating techniques indicated that dispersal from the Pacific to the Atlantic occurred about 750 000 years ago and that dispersal in the opposite direction occurred about 72 000 years ago. The geographical distribution of haplotypes and published field evidence further suggest that migratory shorebirds transported the snails across Central America at the Isthmus of Tehuantepec in southern Mexico. Migratory birds could disperse other intertidal invertebrates this way, suggesting the Central American Isthmus may not be as impassable for marine species as previously assumed.     

Pre-Columbian Floristic Legacies in Modern Homegardens of Central Amazonia

Historical ecologists have demonstrated legacy effects in apparently wild landscapes in Europe, North America, Mesoamerica, Amazonia, Africa and Oceania. People live and farm in archaeological sites today in many parts of the world, but nobody has looked for the legacies of past human occupations in the most dynamic areas in these sites: homegardens. Here we show that the useful flora of modern homegardens is partially a legacy of pre-Columbian occupations in Central Amazonia: the more complex the archaeological context, the more variable the floristic composition of useful native plants in homegardens cultivated there today. Species diversity was 10% higher in homegardens situated in multi-occupational archaeological contexts compared with homegardens situated in single-occupational ones. Species heterogeneity (β-diversity) among archaeological contexts was similar for the whole set of species, but markedly different when only native Amazonian species were included, suggesting the influence of pre-conquest indigenous occupations on current homegarden species composition. Our findings show that the legacy of pre-Columbian occupations is visible in the most dynamic of all agroecosystems, adding another dimension to the human footprint in the Amazonian landscape.     

Amazonian Amphibian Diversity Is Primarily Derived from Late Miocene Andean Lineages

The Neotropics contains half of remaining rainforests and Earth's largest reservoir of amphibian biodiversity. However, determinants of Neotropical biodiversity (i.e., vicariance, dispersals, extinctions, and radiations) earlier than the Quaternary are largely unstudied. Using a novel method of ancestral area reconstruction and relaxed Bayesian clock analyses, we reconstructed the biogeography of the poison frog clade (Dendrobatidae). We rejected an Amazonian center-of-origin in favor of a complex connectivity model expanding over the Neotropics. We inferred 14 dispersals into and 18 out of Amazonia to adjacent regions; the Andes were the major source of dispersals into Amazonia. We found three episodes of lineage dispersal with two interleaved periods of vicariant events between South and Central America. During the late Miocene, Amazonian, and Central American-Chocoan lineages significantly increased their diversity compared to the Andean and Guianan-Venezuelan-Brazilian Shield counterparts. Significant percentage of dendrobatid diversity in Amazonia and Chocó resulted from repeated immigrations, with radiations at <10.0 million years ago (MYA), rather than in situ diversification. In contrast, the Andes, Venezuelan Highlands, and Guiana Shield have undergone extended in situ diversification at near constant rate since the Oligocene. The effects of Miocene paleogeographic events on Neotropical diversification dynamics provided the framework under which Quaternary patterns of endemism evolved.     

Hybridization in headwater regions,and the role of rivers as drivers of speciation in Amazonian birds

Many understory birds and other groups form genetically differentiated subspecies or closely related species on opposite sides of major rivers of Amazonia, but are proposed to come into geographic contact in headwater regions where narrower river widths may present less of a dispersal barrier. Whether such forms hybridize in headwater regions is generally unknown, but has important implications to our understanding of the role of rivers as drivers of speciation. We used a dataset of several thousand single nucleotide polymorphisms to show that seven taxon pairs that differentiate across a major Amazonian river come into geographic contact and hybridize in headwater regions. All taxon pairs possessed hybrids with low numbers of loci in which alleles were inherited from both parental species, suggesting they are backcrossed with parentals, and indicating gene flow between parental populations. Ongoing gene flow challenges rivers as the sole cause of in situ speciation, but is compatible with the view that the wide river courses in the heart of Amazonia may have driven interfluvial divergence during episodes of wet forest retraction away from headwater regions. Taxa as old as 4 Ma in our Amazonian dataset continue to hybridize at contact zones, suggesting reproductive isolation evolves at a slow pace.     

Do flood pulses structure amphibian communities in floodplain environments?

Beta diversity can provide insights into the processes that regulate communities subjected to frequent disturbances, such as flood pulses, which control biodiversity in floodplains. However, little is known about which processes structure beta diversity of amphibians in floodplains. Here, we tested the influence of flood pulses on the richness, composition, and beta diversity of amphibians in Amazonian floodplain environments. We also evaluated indicator species for each environment. We established linear transects in three environments: low várzea, high várzea, and macrophyte rafts. Species richness decreased and beta diversity increased according to the susceptibility of habitats to flood pulses. Indicator species differed among environments according to forest succession promoted by the flood pulse. The decrease in species richness between high and low várzea is due to non‐random extinctions. The higher rates of species turnover between várzeas and macrophyte rafts are driven by the colonization of species adapted to open areas. Our results highlight that the maintenance of complex environments is needed to protect biodiversity in floodplains.     

Tracing an invasion: landbridges, refugia, and the phylogeography of the Neotropical rattlesnake (Serpentes: Viperidae: Crotalus durissus)

Abstract Pleistocene fragmentation of the Amazonian rainforest has been hypothesized to be a major cause of Neotropical speciation and diversity. However, the role and even the reality of Pleistocene forest refugia have attracted much scepticism. In Amazonia, previous phylogeographical studies have focused mostly on organisms found in the forests themselves, and generally found speciation events to have predated the Pleistocene. However, molecular studies of open-formation taxa found both north and south of the Amazonian forests, probably because of vicariance resulting from expansion of the rainforests, may provide novel insights into the age of continuous forest cover across the Amazon basin. Here, we analyse three mitochondrial genes to infer the phylogeography of one such trans-Amazonian vicariant, the Neotropical rattlesnake (Crotalus durissus), which occupies primarily seasonal formations from Mexico to Argentina, but avoids the rainforests of Central and tropical South America. The phylogeographical pattern is consistent with gradual dispersal along the Central American Isthmus, followed by more rapid dispersal into and across South America after the uplift of the Isthmus of Panama. Low sequence divergence between populations from north and south of the Amazon rainforest is consistent with mid-Pleistocene divergence, approximately 1.1 million years ago (Ma). This suggests that the Amazonian rainforests must have become fragmented or at least shrunk considerably during that period, lending support to the Pleistocene refugia theory as an important cause of distribution patterns, if not necessarily speciation, in Amazonian forest organisms. These results highlight the potential of nonforest species to contribute to an understanding of the history of the Amazonian rainforests themselves.     

Alternative models of vertebrate speciation in Amazonia: an overview

The main hypotheses proposed to explain barrier formation separating populations and causing the differentiation of vertebrate species in Amazonia are based on different (mostly historical) factors, as follows. (1) Changes in the distribution of land and sea or in the landscape due to tectonic movements or sea-level fluctuations (Paleogeography hypothesis). (2) The barrier effect of Amazonian rivers (River hypothesis). (3) A combination of the barrier effect of broad rivers and vegetational changes in Northern and Southern Amazonia (River-refuge hypothesis). (4) The isolation of forest blocks near areas of surface relief in the periphery of Amazonia during dry climatic periods of the Tertiary and Quaternary (Refuge theory). (5) Competitive species interactions and local species isolations in peripheral regions of Amazonia due to invasion and counterinvasion during cold/warm periods of the Pleistocene (Disturbance-vicariance hypothesis). (6) Parapatric speciation across steep environmental gradients without separation of the representative populations (Gradient hypothesis). Several of these hypotheses are probably relevant to a different degree for the speciation processes in different faunal groups or during different geological periods. The paleogeography hypothesis refers mainly to faunal differentiation during the Tertiary and in combination with the Refuge hypothesis; Milankovitch cycles leading to global climatic-vegetational changes affected the biomes of the world not only during the Pleistocene but also during the Tertiary and earlier geological periods. New geoscientific evidence for the effect of dry climatic periods in Amazonia supports the predictions of the Refuge theory.     

Patterns of plant phenology in Amazonian seasonally flooded and unflooded forests

Few studies have successfully monitored community‐wide phenological patterns in seasonally flooded Amazonian várzea forests, where a prolonged annual flood pulse arguably generates the greatest degree of seasonality of any low‐latitude ecosystem on Earth. We monitored the vegetative and reproductive plant phenology of várzea (VZ) floodplain and adjacent terra firme (TF) forests within two contiguous protected areas in western Brazilian Amazonia, using three complementary methods: monthly canopy observations of 1056 individuals (TF: 556, VZ: 500), twice monthly collections from 0.5‐m² litterfall traps within two 100‐ha plots (1 TF, 1 VZ; 96 traps per plot), and monthly ground surveys of residual fruit‐fall along transect‐grids within each 100‐ha plot (12 km per plot). Surveys encompassed the entire annual flood cycle and employed a floating trap design to cope with fluctuating water levels. Phenology patterns were generally similar in both forest types. Leaffall peaked during the aquatic phase in várzea forest and the dry season in terra firme. Flowering typically followed leaffall and leaf flush, extending into the onset of the terrestrial phase and rainy season in várzea and terra firme, respectively. Abiotic seed dispersal modes were relatively more prevalent in várzea than terra firme; the main contrast in fruiting seasonality was more likely a result of differences in community composition and relative abundance of seed dispersal modes than differences within individual genera. We emphasize the difficulty in distinguishing the role of the flood pulse from other seasonal environmental variables without multiannual data or spatially replicated studies across the spectrum of Amazonian forest types.     

Crop Diversity on Anthropogenic Dark Earths in Central Amazonia

A recent archaeological survey demonstrates that one of the most durable of all forms of pre-Columbian landscape transformation, Amazonian Dark Earths (ADE; soils formed by pre-Columbian settlement), are widespread along the course of the Madeira River, Central Amazonia, Brazil. We hypothesize that processes of crop cultivation and management by human populations today in landscapes that were intensively transformed during the pre-Columbian period will diverge from those in environments where human agency has not left such a heavy footprint. In order to test this hypothesis, we compare bitter manioc fields, homegardens and secondary forests on ADE with those on non-anthropogenic soils along the lower and middle Madeira River. We demonstrate that crop species and landrace populations diverge on anthropogenic and non-anthropogenic soils as a result of the interaction between human selection and management, soil physical and chemical properties, and plant responses over time. Hence, crop species selection and abundance and therefore agrobiodiversity is contingent on anthropogenic soils in Central Amazonia.     

The legacy of human use in Amazonian palm communities along environmental and accessibility gradients

Aim

Palms are iconic and dominant elements of neotropical forests. In the Amazon region, palms have been used and managed by humans for food, material, medicine and other purposes for millennia. It is, however, debated to what extent the structure of modern palm communities reflects long-term human modification. Here, we investigate the complex interplay of ecological and societal factors that influence the distributions of both human-used and non-used palms in western Amazonia.

Location

Amazonia.

Time period

Present.

Major taxa studied

Palms (Arecaceae).

Methods

We used Bayesian hierarchical joint species distribution models to predict the distributions and environmental niche dimensions of 78 western Amazonian species, and to explore their relationships with their diversity of human uses and with specific uses (food, construction and medicine). The models were parameterized with a comprehensive set of field- and satellite-derived environmental predictors.

Results

Our results suggest that a combination of ecological and anthropogenic factors drive the present-day distributions of Amazonian palms. The modelled ecological niches of the species revealed use-related species-sorting along soil, climatic, accessibility and drainage gradients. We found peaks in the proportions of useful palms and their diversity of uses in fertile soils, close to rivers, and on floodplains. These are habitats favourable for human settlement, although they harbour naturally restricted palm species pools. We also found a negative correlation between predicted palm species richness and number of human uses across western Amazonia.

Main conclusions

Soil characteristics, accessibility, and species pool size all contribute to defining palm–human relationships. At the basin scale, the signature of human use on palm communities was predicted to be stronger in the species-poor south-west than in central-western Amazonia. Overall, we conclude that environmental conditions have influenced modern Amazonian palm distributions both directly and indirectly, by regulating human settlement patterns and natural resource use over extended time periods.     

The relative role of rivers,environmental heterogeneity and species traits in driving compositional changes in southeastern Amazonian bird assemblages

Amazonian rivers have been proposed to act as geographic barriers to species dispersal, either driving allopatric speciation or defining current distribution limits. The strength of the barrier varies according to the species’ ecological characteristics and the river's physical properties. Environmental heterogeneity may also drive compositional changes but has not been well assessed in Amazonia. Aiming to understand the contributions of riverine barriers and environmental heterogeneity in shaping compositional changes in Amazonian forest bird assemblages, we focus on the Tapajós River. We investigate how spatial variation in species composition is related to physical barriers (Tapajós and Jamanxim rivers), species’ ecological characteristics (distinct guilds), and environmental heterogeneity (canopy reflectance, soils, and elevation). We sampled birds through point-counts and mist nets on both banks of the Tapajós and Jamanxim rivers. To test for relationships between bird composition and environmental data, we used Mantel and partial Mantel tests, NMDS, and ANOVA + Tukey HSD. The Mantel tests showed that the clearest compositional changes occurred across the Tapajós River, which seems to act unequally as a significant barrier to the bird guilds. The Jamanxim River was not associated with differences in bird communities. Our results reinforce that the Tapajós River is a biogeographical boundary for birds, while environmental heterogeneity influences compositional variation within interfluves. We discuss the combined influence of geographical barriers, environmental heterogeneity, and ecological characteristics of species in shaping species distributions and community composition and the complexity of extrapolating the patterns found for birds to other Amazonian organisms. Abstract in Portuguese is available with online material.     

A Phylogenetic Lineage of Closely Related Trypanosomes (Trypanosomatidae,Kinetoplastida) of Anurans and Sand Flies (Psychodidae,Diptera) Sharing the Same Ecotopes in Brazilian Amazonia1

ABSTRACT. Analysis of the phylogenetic relationships among trypanosomes from vertebrates and invertebrates disclosed a new lineage of trypanosomes circulating among anurans and sand flies that share the same ecotopes in Brazilian Amazonia. This assemblage of closely related trypanosomes was determined by comparing whole SSU rDNA sequences of anuran trypanosomes from the Brazilian biomes of Amazonia, the Pantanal, and the Atlantic Forest and from Europe, North America, and Africa, and from trypanosomes of sand flies from Amazonia. Phylogenetic trees based on maximum likelihood and parsimony corroborated the positioning of all new anuran trypanosomes in the aquatic clade but did not support the monophyly of anuran trypanosomes. However, all analyses always supported four major clades (An01‐04) of anuran trypanosomes. Clade An04 is composed of trypanosomes from exotic anurans. Isolates in clades An01 and An02 were from Brazilian frogs and toads captured in the three biomes studied, Amazonia, the Pantanal and the Atlantic Forest. Clade An01 contains mostly isolates from Hylidae whereas clade An02 comprises mostly isolates from Bufonidae; and clade An03 contains trypanosomes from sand flies and anurans of Bufonidae, Leptodactylidae, and Leiuperidae exclusively from Amazonia. To our knowledge, this is the first study describing morphological and growth features, and molecular phylogenetic affiliation of trypanosomes from anurans and phlebotomines, incriminating these flies as invertebrate hosts and probably also as important vectors of Amazonian terrestrial anuran trypanosomes.