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An implicit assumption of speciation biology is that population differentiation is an important stage of evolutionary diversification, but its significance as a rate-limiting control on phylogenetic speciation dynamics remains largely untested. If population differentiation within a species is related to its speciation rate over evolutionary time, the causes of differentiation could also be driving dynamics of organismal diversity across time and space. Alternatively, geographic variants might be short-lived entities with rates of formation that are unlinked to speciation rates, in which case the causes of differentiation would have only ephemeral impacts. By pairing population genetics datasets from173 NewWorld bird species (>17,000 individuals) with phylogenetic estimates of speciation rate, we show that the population differentiation rates within species are positively correlated with their speciation rates over long timescales. Although population differentiation rate explains relatively little of the variation in speciation rate among lineages, the positive relationship between differentiation rate and speciation rate is robust to species-delimitation schemes and to alternative measures of both rates. Population differentiation occurs at least three times faster than speciation, which suggests that most populations are ephemeral. Speciation and population differentiation rates are more tightly linked in tropical species than in temperate species, consistent with a history of more stable diversification dynamics through time in the Tropics. Overall, our results suggest that the processes responsible for population differentiation are tied to those that underlie broad-scale patterns of diversity.

High tropical species diversity is often attributed to evolutionary dynamics over long timescales. It is possible, however, that latitudinal variation in diversification begins when divergence occurs within species. Phylogeographic data capture this initial stage of diversification in which populations become geographically isolated and begin to differentiate genetically. There is limited understanding of the broader implications of intraspecific diversification because comparative analyses have focused on species inhabiting and evolving in restricted regions and environments. Here, we scale comparative phylogeography up to the hemisphere level and examine whether the processes driving latitudinal differences in species diversity are also evident within species. We collected genetic data for 210 New World bird species distributed across a broad latitudinal gradient and estimated a suite of metrics characterizing phylogeographic history. We found that lower latitude species had, on average, greater phylogeographic diversity than higher latitude species and that intraspecific diversity showed evidence of greater persistence in the tropics. Factors associated with species ecologies, life histories, and habitats explained little of the variation in phylogeographic structure across the latitudinal gradient. Our results suggest that the latitudinal gradient in species richness originates, at least partly, from population-level processes within species and are consistent with hypotheses implicating age and environmental stability in the formation of diversity gradients. Comparative phylogeographic analyses scaled up to large geographic regions and hundreds of species can show connections between population-level processes and broad-scale species-richness patterns.

We studied the phylogeography and plumage variation of the Russet-crowned Warbler (Myiothlypis coronata), from Venezuela to Bolivia, with focus on populations from Ecuador and northern Peru. We analyzed sequences of mitochondrial and nuclear genes, geographic distributions, as well as photographs of specimens deposited at museum collections. Phylogenetic analyses identified three major lineages formed by populations from: Venezuela and Colombia (M. c. regulus), Ecuador and northern Peru (M. elata, M. castaneiceps, M. orientalis, M. c. chapmani), and central Peru and Bolivia (M. c. coronata). We found further population structure within M. c. regulus and M. c. coronata, and population structure and complexity of plumage variation within the Ecuador-northern Peru lineage. Time-calibrated trees estimated that most intraspecific variation originated during the Pleistocene; however, this pattern may not be attributed to an increase in diversification rate during that period. We discuss these results in the context of the importance of geographic-ecological barriers in promoting lineage diversification along the Andes and put forward a preliminary taxonomic proposal for major lineages identified in this study.

Aim: To evaluate the role of historical processes in the evolution of Sclerurus leaftossers by integrating phylogenetic and phylogeographical approaches. Location: Humid forests of the Neotropical region. Methods: We reconstructed the evolutionary history of Sclerurus based on DNA sequences representing all species and 20 of the 26 recognized subspecies using one autosomal nuclear locus and three protein-coding mitochondrial gene sequences. Phylogenetic relationships were inferred using Bayesian and maximum-likelihood methods. We used Bayesian coalescent-based approaches to evaluate demographic changes through time, and to estimate the timing of diversification events. Based on these results, we examined the temporal accumulation of divergence events using lineage-through-time plots. Results: The monophyly of all Sclerurus species was strongly supported except for Sclerurus mexicanus, which was paraphyletic in relation to Sclerurus rufigularis, and for the sister pair Sclerurus scansor—Sclerurus albigularis, which were not reciprocally monophyletic in the nuclear tree. We found remarkably deep phylogeographical structure within all Sclerurus species, and overall this structure was congruent with currently recognized subspecies and Neotropical areas of endemism. Diversification within Sclerurus has occurred at a relatively constant rate since the Middle Miocene. Main conclusions: Our results strongly support the relevance of physiographical (e.g. Nicaragua Depression, Isthmus of Panama, Andean Cordillera, great rivers of Amazonia) and ecological barriers (open vegetation corridor) and ecological gradients (elevational zonation) to the diversification of Neotropical forest-dwelling organisms. Despite the high congruence among the spatial patterns identified, the variance in divergence times suggests multiple speciation events occurring independently across the same barrier, and a role for dispersal. The phylogenetic patterns and cryptic diversity uncovered in this study demonstrate that the current taxonomy of Sclerurus underestimates the number of species.

AIM: To examine the effect of geographical barriers and habitat dynamics related to climatic oscillations on the phylogeography of a widespread passerine of Neotropical cloud forests, the spotted barbtail (Premnoplex brunnescens). LOCATION: Neotropical humid forests of montane areas in lower Central America and South America. METHODS: We sequenced two mitochondrial genes and one nuclear intron from specimens collected across the distribution of P. brunnescens. Phylogenetic relationships were inferred using Bayesian and maximum‐likelihood methods. Groups with maximum differentiation were estimated with spatial analysis of molecular variance (SAMOVA). We estimated timing of differentiation and relationships among groups with a species‐tree approach and historical demography with extended Bayesian skyline plots. RESULTS: Six highly differentiated clades of P. brunnescens are distributed in lower Central America, Sierra Nevada de Santa Marta, northern Venezuelan mountains, the Northern Andes, central Peru, and southern Peru and Bolivia. Within the Northern Andes clade, six phylogroups were identified associated with different slopes and isolated cordilleras. Most clades occupy opposite sides of low‐lying valleys and ridgelines, but little differentiation was observed across several putative barriers. Population divergence occurred in the late Miocene and Pliocene, perhaps in association with Andean uplift. Historical fluctuations in population sizes suggest that populations tracked the spatial dynamics of montane forests associated with glacial cycles. MAIN CONCLUSIONS: Extensive genetic differentiation in mitochondrial and nuclear DNA exists among populations of P. brunnescens. Such marked divergence was probably promoted by the rugged topography and dynamic ecological history of the Neotropical mountains. Our study sheds light on mechanisms promoting population differentiation in the montane Neotropics.

Ecological speciation can proceed despite genetic interchange when selection counteracts the homogenizing effects of migration. We tested predictions of this divergence-with-gene-flow model in Coeligena helianthea and C. bonapartei, 2 parapatric Andean hummingbirds with marked plumage divergence. We sequenced putatively neutral markers (mitochondrial DNA [mtDNA] and nuclear ultraconserved elements [UCEs]) to examine genetic structure and gene flow, and a candidate gene (MC1R) to assess its role underlying divergence in coloration. We also tested the prediction of Gloger's rule that darker forms occur in more humid environments, and examined morphological variation to assess adaptive mechanisms potentially promoting divergence. Genetic differentiation between species was low in both ND2 and UCEs. Coalescent estimates of migration were consistent with divergence with gene flow, but we cannot reject incomplete lineage sorting reflecting recent speciation as an explanation for patterns of genetic variation. MC1R variation was unrelated to phenotypic differences. Species did not differ in macroclimatic niches but were distinct in morphology. Although we reject adaptation to variation in macroclimatic conditions as a cause of divergence, speciation may have occurred in the face of gene flow driven by other ecological pressures or by sexual selection. Marked phenotypic divergence with no neutral genetic differentiation is remarkable for Neotropical birds, and makes C. helianthea and C. bonapartei an appropriate system in which to search for the genetic basis of species differences employing genomics.

We describe a new species of wren in the genus Thryophilus (Troglodytidae) based on analysis of morphological, vocal, and genetic variation. Individuals of the new species are readily separated in the field or the museum from those of any other wren species, including its closest relatives T. rufalbus and T. nicefori, by a combination of traits including, but not limited to, plumage coloration of the upperparts, the pattern of barring on the wings and tail, overall smaller body size, a richer repertoire of syllable types, shorter trills, and distinctive terminal syllables. The new species is allopatrically distributed in relation to its congeners, being restricted to the dry Cauca River Canyon, a narrow inter-Andean valley enclosed by the Nechí Refuge rainforests and the northern sectors of the Western and Central Andes of Colombia. Individuals or pairs have been found only in remnant patches of dry forest and scrub at 250– 850 m elevation. This newly discovered species is uncommon and threatened because of ongoing transformation of natural habitats in the Cauca River Canyon, and especially because of the planned construction of a major dam in the region; immediate conservation actions are thus imperative.

Diversification of Neotropical birds is not directly linked to the Andean uplift, the major landscape change of the Neogene period; instead, most diversification is post-Neogene and species diversity is dependent on how long lineages have persisted in the landscape and how easily they disperse. Landscape not decisive in neotropical speciation The idea that landscape change drives diversification is firmly embedded in the biogeographical literature. It has been difficult to test this against alternative possibilities, including a model in which diversification is driven by evolutionary persistence and geographic structuring of populations by the ability of an organism to navigate the landscape matrix. Robb Brumfield and colleagues have examined patterns of genetic differentiation in co-distributed bird species in tropical Central and South America and find unequivocal support for the latter model. The data are a poor fit to the model invoking landscape change, revealing no direct link to Andes uplift. Rather, diversification times differ from each other widely and depend on how long lineages persist in the landscape and how easily they disperse. Since the recognition that allopatric speciation can be induced by large-scale reconfigurations of the landscape that isolate formerly continuous populations, such as the separation of continents by plate tectonics, the uplift of mountains or the formation of large rivers, landscape change has been viewed as a primary driver of biological diversification. This process is referred to in biogeography as vicariance 1 . In the most species-rich region of the world, the Neotropics, the sundering of populations associated with the Andean uplift is ascribed this principal role in speciation 2 , 3 , 4 , 5 . An alternative model posits that rather than being directly linked to landscape change, allopatric speciation is initiated to a greater extent by dispersal events, with the principal drivers of speciation being organism-specific abilities to persist and disperse in the landscape 6 , 7 . Landscape change is not a necessity for speciation in this model 8 . Here we show that spatial and temporal patterns of genetic differentiation in Neotropical birds are highly discordant across lineages and are not reconcilable with a model linking speciation solely to landscape change. Instead, the strongest predictors of speciation are the amount of time a lineage has persisted in the landscape and the ability of birds to move through the landscape matrix. These results, augmented by the observation that most species-level diversity originated after episodes of major Andean uplift in the Neogene period, suggest that dispersal and differentiation on a matrix previously shaped by large-scale landscape events was a major driver of avian speciation in lowland Neotropical rainforests.

The Rusty-breasted Antpitta (Grallaricula ferrugineipectus) is widely distributed within the tropical Andes of South America. We analyzed 73 study specimens, 25 vouchered tissue samples, and 123 audio recordings to assess geographic variation in genetics, vocalizations, and morphology and evaluate species limits. We found that Grallaricula ferrugineipectus as currently defined is polyphyletic because populations from Colombia and Venezuela form a clade closely related to Andean populations of G. nana, whereas populations from Peru and Bolivia are recovered as sister to G. lineifrons. Birds in Colombia and Venezuela (the northern group) last shared a common ancestor with birds from Peru and Bolivia (the southern group) more than 10 million years ago. Northern and southern groups additionally differ in song, suggesting they may have evolved substantial premating reproductive isolation. Discriminant function analysis reliably distinguished songs from northern and southern groups in multivariate acoustic space, but univariate analyses found non-overlapping acoustic variation between northern and southern groups in only one trait: mean note maximum frequency (and other correlated measures of song pitch). This finding suggests that the ‘three-trait' threshold for using vocalizations to inform species limits, which was developed for another suboscine group, the antbirds (Thamnophilidae), may be conservative when applied to antpittas (Grallariidae). In addition, we document apparent clinal variation in song pace within the southern group, a rare example of a suboscine with geographic clinal variation in a vocal trait. Finally, we show that northern and southern groups differ markedly in morphology. In summary, northern and southern groups of Rusty-breasted Antpittas are divergent in genetics, vocalizations, and morphology, demonstrating that these taxa are best classified as 2 monophyletic, biological species with allopatric distributions.

[This corrects the article DOI: 10.1371/journal.pbio.2001073.].