John Klicka

Phylogenetics and diversification of tanagers (Passeriformes: Thraupidae), the largest radiation of Neotropical songbirds

Thraupidae is the second largest family of birds and represents about 4% of all avian species and 12% of the Neotropical avifauna. Species in this family display a wide range of plumage colors and patterns, foraging behaviors, vocalizations, ecotypes, and habitat preferences. The lack of a complete phylogeny for tanagers has hindered the study of this evolutionary diversity. Here, we present a comprehensive, species-level phylogeny for tanagers using six molecular markers. Our analyses identified 13 major clades of tanagers that we designate as subfamilies. In addition, two species are recognized as distinct branches on the tanager tree. Our topologies disagree in many places with previous estimates of relationships within tanagers, and many long-recognized genera are not monophyletic in our analyses. Our trees identify several cases of convergent evolution in plumage ornaments and bill morphology, and two cases of social mimicry. The phylogeny produced by this study provides a robust framework for studying macroevolutionary patterns and character evolution. We use our new phylogeny to study diversification processes, and find that tanagers show a background model of exponentially declining diversification rates. Thus, the evolution of tanagers began with an initial burst of diversification followed by a rate slowdown. In addition to this background model, two later, clade-specific rate shifts are supported, one increase for Darwins finches and another increase for some species of Sporophila. The rate of diversification within these two groups is exceptional, even when compared to the overall rapid rate of diversification found within tanagers. This study provides the first robust assessment of diversification rates for the Darwins finches in the context of the larger group within which they evolved.

New insights into New World biogeography: An integrated view from the phylogeny of blackbirds, cardinals, sparrows, tanagers, warblers, and allies

ABSTRACT Understanding the biogeographic origins and temporal sequencing of groups within a region or of lineages within an ecosystem can yield important insights into evolutionary dynamics and ecological processes. Fifty years ago, Ernst Mayr generated comprehensive—if limited—inferences about the origins of the New World avifaunas, including the importance of pre-Isthmian dispersal between North and South America. Since then, methodological advances have improved our ability to address many of the same questions, but the phylogenies upon which such analyses should be based have been incompletely sampled or fragmentary. Here, we report a near-species-level phylogeny of the diverse (~832 species) New World clade Emberizoidea—the group that includes the familiar sparrows, cardinals, blackbirds, wood-warblers, tanagers, and their close relatives—to our knowledge the largest essentially complete (≥95%) phylogenetic hypothesis for any group of organisms. Biogeographic analyses based on this tree suggest initial dispersal into the New World via Beringia, with rapid subsequent diversification, including early dispersal of 1 lineage (the tanagers, Thraupidae) into South America. We found substantial dispersal between North and South America prior to closure of the Isthmus of Panama, but with a notable increase afterward, with a directional bias from north to south. With much greater detail and historical rigor, these analyses largely confirm Mayrs speculations based on taxonomy, resolving outstanding ambiguity regarding the continental origins of some groups such as the Emberizidae and Icteridae. The phylogeny reported here will be a resource of broad utility for addressing additional evolutionary and ecological questions with this diverse group.

How Many Kinds of Birds Are There and Why Does It Matter

Estimates of global species diversity have varied widely, primarily based on variation in the numbers derived from different inventory methods of arthropods and other small invertebrates. Within vertebrates, current diversity metrics for fishes, amphibians, and reptiles are known to be poor estimators, whereas those for birds and mammals are often assumed to be relatively well established. We show that avian evolutionary diversity is significantly underestimated due to a taxonomic tradition not found in most other taxonomic groups. Using a sample of 200 species taken from a list of 9159 biological species determined primarily by morphological criteria, we applied a diagnostic, evolutionary species concept to a morphological and distributional data set that resulted in an estimate of 18,043 species of birds worldwide, with a 95% confidence interval of 15,845 to 20,470. In a second, independent analysis, we examined intraspecific genetic data from 437 traditional avian species, finding an average of 2.4 evolutionary units per species, which can be considered proxies for phylogenetic species. Comparing recent lists of species to that used in this study (based primarily on morphology) revealed that taxonomic changes in the past 25 years have led to an increase of only 9%, well below what our results predict. Therefore, our molecular and morphological results suggest that the current taxonomy of birds understimates avian species diversity by at least a factor of two. We suggest that a revised taxonomy that better captures avian species diversity will enhance the quantification and analysis of global patterns of diversity and distribution, as well as provide a more appropriate framework for understanding the evolutionary history of birds.

A multilocus phylogeny of a major New World avian radiation: The Vireonidae

The family Vireonidae represents one of the most widespread and well-known New World avian radiations, but a robust species-level phylogeny of the group is lacking. Here, we infer a phylogeny of Vireonidae using multilocus data obtained from 221 individuals from 46 of 52 vireonid species (representing all four genera) and five core Corvoidea outgroups. Our results show Vireonidae to be monophyletic, consistent with a single colonization of the New World by an Asian ancestor. Cyclarhis and Vireolanius are monophyletic genera that diverged early from the rest of Vireonidae. Hylophilus is polyphyletic, represented by three distinct clades concordant with differences in morphology, habitat, and voice. The poorly known South American species Hylophilus sclateri is embedded within the genus Vireo. Vireo, in turn, consists of several well-supported intrageneric clades. Overall, tropical vireonid species show much higher levels of intraspecific genetic structure than temperate species and several currently recognized species are probably comprised of multiple cryptic species.

Effects of climate change on the evolution of Brown Creeper (Certhia americana) lineages

ABSTRACT Understanding how distributions of species change through time allows evaluation of hypotheses about factors shaping biogeographic patterns and evolutionary trajectories of genetic lineages. Ideally, such studies would assess whether population genetic processes are associated with geographic distribution shifts, loss or gain of distributional area through time, or fragmentation of distributional areas, information that can now be derived via ecological niche modeling. We examined the distributional changes through time in lineages and populations of Brown Creeper (Certhia americana), a widespread North American bird, to test biogeographic and population genetic hypotheses. In two populations with genetic support for population bottlenecks, Monterey County in California and the Sierra Madre Oriental in Mexico, ecological niche models indicated range contractions and increased fragmentation since the Last Glacial Maximum (LGM). Projections of niche models to the future suggested continuation of range contractions and fragmentation. Of the 3 major allopatric lineages of Brown Creeper (eastern North America, western North America, and southern North America and Central America), the most stable through time was the southern lineage, which corresponds with increased genetic diversity. The potential geographic distribution of the western lineage has remained stable in size but not location since the LGM, corresponding with a genetic signal of isolation by distance. The eastern lineage experienced range contractions during the LGM, likely resulting in the contemporary lack of genetic structure within the lineage. Finally, there is limited evidence of potential range overlap during the LGM between the western lineage and the other 2 lineages, although the overlap is limited to the Arizona sky islands between the west and south lineages. These results suggest that ecological niche modeling and population genetic data may be used as mutual predictors when investigating phylogeographic patterns and processes.

Annotated checklist of the birds (Aves) of Cerro Hoya National Park, Azuero Peninsula, Panama

Protected only by the extreme ruggedness of its terrain, the montane regions of Cerro Hoya National Park are among the least biologically known regions of Central America. Here we provide a checklist of 225 bird species recorded from five expeditions to the region over the last 18 years, which represents lower species richness than many comparable areas in Panama and lower Central America. However, we recorded nine IUCN globally threatened species, three species with range extensions over 150 km to the southern Azuero Peninsula, and at least two previously unknown bird taxa. These facts suggest that our list of montane taxa from Cerro Hoya may be incomplete, and that this region represents one of the most important—and neglected—protected areas in Panama.

Polyphyly of Hylophilus and a new genus for the Tawny-crowned Greenlet (Aves: Passeriformes: Vireonidae).

Once a catch-all taxon for various small, greenish passerines (Sclater 1881), today the genus Hylophilus Temminck contains 15 species of Neotropical greenlets in the avian family Vireonidae (Clements et al. 2013). Although Hylophilus species do share some common anatomical proportions and plumage features (Baird 1866; Ridgway 1904), some striking and concordant differences in habitat, voice, and iris color led Ridgely and Tudor (1989) to posit that the genus might contain sufficient diversity to warrant splitting into multiple genera.

Phylogenomic insights into the diversification of salamanders in the Isthmura bellii group across the Mexican highlands

Mountain formation in Mexico has played an important role in the diversification of many Mexican taxa. The Trans-Mexican Volcanic Belt in particular has served as both a cradle of diversification and conduit for dispersal. We investigated the evolutionary history of the Isthmura bellii group of salamanders, a widespread amphibian across the Mexican highlands, using sequence capture of ultraconserved elements. Results suggest that the I. bellii group probably originated in southeastern Mexico in the late Miocene and later dispersed across the Trans-Mexican Volcanic Belt and into the Sierra Madre Occidental. Pre-Pleistocene uplift of the Trans-Volcanic Belt likely promoted early diversification by serving as a mesic land-bridge across central Mexico. These findings highlight the importance of the Trans-Volcanic Belt in generating Mexicos rich biodiversity.

Extreme genetic structure and dynamic range evolution in a montane passerine bird: implications for tropical diversification

Aim Employ phylogeographic analyses of a widespread species complex to examine the role of historical and evolutionary processes in the origin and maintenance of high species diversity in the Neotropical montane region. Location Neotropical highlands. Taxon Henicorhina wood-wrens (Aves, Troglodytidae). Methods We collected mtDNA sequence data for 288 individuals thoroughly covering the range of the Henicorhina leucophrys complex from Mexico to Bolivia. Sequences were employed to characterize population structure, infer phylogenetic relationships among populations and their divergence times, examine lineage accumulation through time, and identify presumptive species using coalescent methods. We also explored the origin of elevational and latitudinal replacements involved in spatial changes in species assemblages in the Andes. Results We found remarkable genetic structure within the complex, which consists of numerous lineages reaching >12% sequence divergence; most divergent populations occur in areas separated by topographic barriers but several of them, typically not sister to each other, co-occur with elevational segregation on mountain slopes or replace each other with latitude along the Andes. Some close relatives occur in areas separated by thousands of kilometers, with more distant relatives occupying intervening areas. The complex likely originated in the Mexican highlands and expanded extensively in South America while diverging rapidly at a constant rate into many different lineages which have persisted for millions of years. Coalescent analyses consistently revealed that the complex may comprise more than 30 species; while we do not suggest these presumptive species should be recognized by taxonomists in the absence of additional data, H. leucophrys is a distant outlier among New World birds in terms of high lineage diversity within a single recognized species. Main Conclusions Our study captured wood-wren lineages in the act of building up diversity via divergence and persistence in allopatry, achievement of secondary sympatry, and coexistence at the landscape scale mediated by ecological and evolutionary divergence. Although dispersal by wood-wrens is restricted at present and this likely accounts for strong population structure across topographic barriers, their ranges have been dynamic, managing to disperse over much of the montane Neotropics. Phases of expansion and contraction of ranges and localized extinctions of populations likely account for phylogeographic patterns which are precursors to the origin of new species and the accumulation of diversity in tropical mountains.

Species delimitation and biogeography of the gnatcatchers and gnatwrens (Aves: Polioptilidae)

The New World avian family Polioptilidae (gnatcatchers and gnatwrens) is distributed from Argentina to Canada and includes 15 species and more than 60 subspecies. No study to date has evaluated phylogenetic relationships within this family and the historical pattern of diversification within the group remains unknown. Moreover, species limits, particularly in widespread taxa that show geographic variation, remain unclear. In this study, we delimited species and estimated phylogenetic relationships using multilocus data for the entire family. We then used the inferred diversity along with alternative taxonomic classification schemes to evaluate how lumping and splitting of both taxa and geographical areas influenced biogeographic inference. Species-tree analyses grouped Polioptilidae into four main clades: Microbates, Ramphocaenus, a Polioptila guianensis complex, and the remaining members of Polioptila. Ramphocaenus melanurus was sister to the clade containing M. cinereiventris and M. collaris, which formed a clade sister to all species within Polioptila. Polioptila was composed of two clades, the first of which included the P. guianensis complex; the other contained all remaining species in the genus. Using multispecies coalescent modeling, we inferred a more than 3-fold increase in species diversity, of which 87% represent currently recognized species or subspecies. Much of this diversity corresponded to subspecies that occur in the Neotropics. We identified three polyphyletic species, and delimited 4-6 previously undescribed candidate taxa. Probabilistic modeling of geographic ranges on the species tree indicated that the family likely had an ancestral origin in South America, with all three genera independently colonizing North America. Support for this hypothesis, however, was sensitive to the taxonomic classification scheme used and the number of geographical areas allowed. Our study proposes the first phylogenetic hypothesis for Polioptilidae and provides genealogical support for the reclassification of species limits. Species limits and the resolution of geographical areas that taxa inhabit influence the inferred spatial diversification history.