Andean Tectonics and Mantle Dynamics as a Pervasive Influence on Amazonian Ecosystem.

The Amazonian landscape evolution is the result of the combined effect of Andean tectonism, climate and the Earth's interior dynamics. To reconstruct the landscape evolution and its influence on paleoenvironmental variations within Amazonia since the Oligocene, we conducted numerical experiments that incorporate different surface and geodynamic processes, reproducing many paleogeographic features as inferred from the sedimentary record. We show that the evolution of the drainage pattern gradually reduced the area of sedimentation derived from the Guiana and Brazilian shields while expanded the Andean derived deposits during the Miocene, affecting the nutrient availability. First order biotic habitats were inferred from these paleogeographical reconstructions, showing an eastward expansion of várzea and terra firme forests and consequent retraction of igapó forests, with a millennial-scale reconfiguration of a mosaic of habitats in the lowlands. We conclude that this dynamism probably guided the observed patterns of speciation in the most biodiverse biome on Earth.

Evidence for mtDNA capture in the jacamar Galbula leucogastra/chalcothorax species-complex and insights on the evolution of white-sand ecosystems in the Amazon basin.

Jacamar species occur throughout Amazonia, with most species occupying forested habitats. One species-complex, Galbula leucogastra/chalcothorax, is associated to white sand ecosystems (WSE). Previous studies of WSE bird species recovered shallow genetic structure in mtDNA coupled with signs of gene flow among WSE patches. Here, we characterize diversification of the G. leucogastra/chalcothorax species-complex with dense sampling across its distribution using mitochondrial and genomic (Ultraconserved Elements, UCEs) DNA sequences. We performed concatenated likelihood and Bayesian analysis, as well as a species-tree analysis using ∗BEAST, to establish the phylogenetic relationships among populations. The mtDNA results recovered at least six geographically-structured lineages, with G. chalcothorax embedded within lineages of G. leucogastra. In contrast, both concatenated and species-tree analyses of UCE data recovered G. chalcothorax as sister to all G. leucogastra lineages. We hypothesize that the mitochondrial genome of one of the G. leucogastra lineage (Madeira) was captured into G. chalcothorax in the past. We discuss how WSE evolution and the coevolution of mtDNA and nuclear genes might have played a role in this apparently rare event.

Molecular systematics of the new world screech-owls (Megascops: Aves, Strigidae): biogeographic and taxonomic implications.

Megascops screech-owls are endemic to the New World and range from southern Canada to the southern cone of South America. The 22 currently recognized Megascops species occupy a wide range of habitats and elevations, from desert to humid montane forest, and from sea level to the Andean tree line. Species and subspecies diagnoses of Megascops are notoriously difficult due to subtle plumage differences among taxa with frequent plumage polymorphism. Using three mitochondrial and three nuclear genes we estimated a phylogeny for all but one Megascops species. Phylogenies were estimated with Maximum Likelihood and Bayesian Inference, and a Bayesian chronogram was reconstructed to assess the spatio-temporal context of Megascops diversification. Megascops was paraphyletic in the recovered tree topologies if the Puerto Rican endemic M. nudipes is included in the genus. However, the remaining taxa are monophyletic and form three major clades: (1) M. choliba, M. koepckeae, M. albogularis, M. clarkii, and M. trichopsis; (2) M. petersoni, M. marshalli, M. hoyi, M. ingens, and M. colombianus; and (3) M. asio, M. kennicottii, M. cooperi, M. barbarus, M. sanctaecatarinae, M. roboratus, M. watsonii, M. atricapilla, M. guatemalae, and M. vermiculatus. Megascops watsonii is paraphyletic with some individuals more closely related to M. atricapilla than to other members in that polytypic species. Also, allopatric populations of some other Megascops species were highly divergent, with levels of genetic differentiation greater than between some recognized species-pairs. Diversification within the genus is hypothesized to have taken place during the last 8 million years, with a likely origin in Central America. The genus later expanded over much of the Americas and then diversified via multiple dispersal events from the Andes into the Neotropical lowlands.

Inferring speciation history in the Andes with reduced-representation sequence data: an example in the bay-backed antpittas (Aves; Grallariidae; Grallaria hypoleuca s. l.).

In the Andes, humid-forest organisms frequently exhibit pronounced genetic structure and geographic variation in phenotype, often coincident with physical barriers to dispersal. However, phylogenetic relationships of clades have often been difficult to resolve due to short internodes. Consequently, even in taxa with well-defined genetic structure, the temporal and geographic sequences of dispersal and vicariance events that led to this differentiation have remained opaque, hindering efforts to test the association between diversification and earth history and to understand the assembly of species-rich communities on Andean slopes. Here, we use mitochondrial DNA and thousands of short-read sequences generated with genotyping by sequencing (GBS) to examine the geographic history of speciation in a lineage of passerine birds found in the humid forest of the Andes, the 'bay-backed' antpitta complex (Grallaria hypoleuca s. l). Mitochondrial DNA genealogies documented genetic structure among clade but were poorly resolved at nodes relevant for biogeographic inference. By contrast, relationships inferred from GBS loci were highly resolved and suggested a biogeographic history in which the ancestor originated in the northern Andes and dispersed south. Our results are consistent with a scenario of vicariant speciation wherein the range of a widespread ancestor was fragmented as a result of geologic or climatic change, rather than a stepping-stone series of dispersal events across pre-existing barriers. However, our study also highlights challenges of distinguishing dispersal-mediated speciation from static vicariance. Our results further demonstrate the substantial evolutionary timescale over which the diverse biota of the Andes was assembled.

The tempo of trait divergence in geographic isolation: avian speciation across the Marañon Valley of Peru.

Geographic isolation is considered essential to most speciation events, but our understanding of what controls the pace and degree of phenotypic divergence among allopatric populations remains poor. Why do some taxa exhibit phenotypic differentiation across barriers to dispersal, whereas others do not? To test factors controlling phenotypic divergence in allopatry, we employed a comparative phylogeographic approach consisting of replicates of ecologically similar Andean bird species isolated across a major biogeographic barrier, the Marañon Valley of Peru. Our study design leverages variation among codistributed taxa in their degree of plumage, morphometric, and vocal differentiation across the Marañon to examine the tempo of phenotypic evolution. We found that substantial plumage differences between populations required roughly two million years to evolve. In contrast, morphometric trait evolution showed greater idiosyncrasy and stasis. Our results demonstrate that despite a large degree of idiosyncrasy in the relationship between genetic and phenotypic divergence across taxa and environments, comparative studies within regions may reveal predictability in the pace of phenotypic divergence. Our results also suggest that social selection is important for driving differentiation of populations found in similar environments.

Biogeography and spatio-temporal diversification of Selenidera and Andigena toucans (Aves: Ramphastidae).

Andean uplift, Plio-Pleistocene climatic fluctuation, and river dynamics in the Amazon basin have all been implicated in the diversification of the South American avifauna. We reconstructed phylogenetic relationships in the genus Selenidera, which has served as a classic case of putative refugial speciation, and the closely related genus Andigena, to better understand the processes driving their diversification. Using mitochondrial and nuclear DNA sequences, we constructed a phylogeny to estimate the pattern and timing of divergence within and between seven lowland Selenidera toucanets and the five species of Andigena mountain-toucans, which together form a single clade. All phylogenetic analyses supported the monophyly of the montane genus Andigena, but indicated that the genus Selenidera is likely paraphyletic with respect to Andigena. Our time tree analysis is consistent with the orogenic uplift of the northern Andean range having initiated the divergence between Selenidera and Andigena, and that the movement and fragmentation of montane habitats in response to Pleistocene climatic oscillations likely influenced diversification within Andigena. Estimated divergence times for lowland Amazonian Selenidera did not support the Last Glacial Maximum (LGM) refuge hypothesis as an important biogeographic factor for the diversification of lineages studied here. The timing of divergence within Selenidera is consistent with the hypothesis that geographic isolation of areas of endemism generated by Amazonian river dynamics during the Plio-Pleistocene contributed to Selenidera speciation and current species distributions.

Temporal and spatial diversification of Pteroglossus araçaris (AVES Ramphastidae) in the neotropics: Constant rate of diversification does not support an increase in radiation during the Pleistocene

We use the small-bodied toucan genus Pteroglossus to test hypotheses about diversification in the lowlandNeotropics. We sequenced three mitochondrial genes and one nuclear intron from all Pteroglossus speciesand used these data to reconstruct phylogenetic trees based on maximum parsimony, maximumlikelihood, and Bayesian analyses. These phylogenetic trees were used to make inferences regarding boththe pattern and timing of diversification for the group. We used the uplift of the Talamanca highlands ofCosta Rica and western Panama as a geologic calibration for estimating divergence times on the Pteroglossustree and compared these results with a standard molecular clock calibration. Then, we used likelihoodmethods to model the rate of diversification. Based on our analyses, the onset of the Pteroglossus radiationpredates the Pleistocene, which has been predicted to have played a pivotal role in diversification in theAmazon rainforest biota. We found a constant rate of diversification in Pteroglossus evolutionary history,and thus no support that events during the Pleistocene caused an increase in diversification. We compareour data to other avian phylogenies to better understand major biogeographic events in the Neotropics.These comparisons support recurring forest connections between the Amazonian and Atlantic forests,and the splitting of cis/trans Andean species after the final uplift of the Andes. At the subspecies level, thereis evidence for reciprocal monophyly and groups are often separated by major rivers, demonstrating theimportant role of rivers in causing or maintaining divergence. Because some of the results presented hereconflict with current taxonomy of Pteroglossus, new taxonomic arrangements are suggested.

Evolutionary history of Ramphastos toucans: molecular phylogenetics, temporal diversification, and biogeography.

The toucan genus Ramphastos (Piciformes: Ramphastidae) has been a model in the formulation of Neotropical paleobiogeographic hypotheses. Weckstein (2005) reported on the phylogenetic history of this genus based on three mitochondrial genes, but some relationships were weakly supported and one of the subspecies of R. vitellinus (citreolaemus) was unsampled. This study expands on Weckstein (2005) by adding more DNA sequence data (including a nuclear marker) and more samples, including R. v. citreolaemus. Maximum parsimony, maximum likelihood, and Bayesian methods recovered similar trees, with nodes showing high support. A monophyletic R. vitellinus complex was strongly supported as the sister-group to R. brevis. The results also confirmed that the southeastern and northern populations of R. vitellinus ariel are paraphyletic. R. v. citreolaemus is sister to the Amazonian subspecies of the vitellinus complex. Using three protein-coding genes (COI, cytochrome-b and ND2) and interval-calibrated nodes under a Bayesian relaxed-clock framework, we infer that ramphastid genera originated in the middle Miocene to early Pliocene, Ramphastos species originated between late Miocene and early Pleistocene, and intra-specific divergences took place throughout the Pleistocene. Parsimony-based reconstruction of ancestral areas indicated that evolution of the four trans-Andean Ramphastos taxa (R. v. citreolaemus, R. a. swainsonii, R. brevis and R. sulfuratus) was associated with four independent dispersals from the cis-Andean region. The last pulse of Andean uplift may have been important for the evolution of R. sulfuratus, whereas the origin of the other trans-Andean Ramphastos taxa is consistent with vicariance due to drying events in the lowland forests north of the Andes. Estimated rates of molecular evolution were higher than the "standard" bird rate of 2% substitutions/site/million years for two of the three genes analyzed (cytochrome-b and ND2).

Amphibian DNA shows marked genetic structure and tracks pleistocene climate change in northeastern Brazil.

The glacial refugia paradigm has been broadly applied to patterns of species dynamics and population diversification. However, recent geological studies have demonstrated striking Pleistocene climate changes in currently semiarid northeastern Brazil at time intervals much more frequent than the climatic oscillations associated with glacial and interglacial periods. These geomorphic data documented recurrent pulses of wet regimes in the past 210,000 years that correlate with climate anomalies affecting multiple continents. While analyzing DNA sequences of two mitochondrial genes (cytochrome b and NADH-dehydrogenase subunit 2) and one nuclear marker (cellular-myelocytomatosis proto-oncogene) in the forest-associated frogs Proceratophrys boiei and Ischnocnema gr. ramagii, we found evidence of biological responses consistent with these pluvial maxima events. Sampled areas included old, naturally isolated forest enclaves within the semiarid Caatinga, as well as recent man-made fragments of humid coastal Atlantic forest. Results show that mtDNA lineages in enclave populations are monophyletic or nearly so, whereas nonenclave populations are polyphyletic and more diverse. The studied taxa show evidence of demographic expansions at times that match phases of pluvial maxima inferred from geological data. Divergence times between several populations fall within comparatively drier intervals suggested by geomorphology. Mitochondrial and nuclear data show local populations to be genetically structured, with some high levels of differentiation that suggest the need of further taxonomic work.