Carl H. Oliveros

Phylogeny and biogeography of the core babblers (Aves: Timaliidae).

The avian family Timaliidae is a species rich and morphologically diverse component of African and Asian tropical forests. The morphological diversity within the family has attracted interest from ecologists and evolutionary biologists, but systematists have long suspected that this diversity might also mislead taxonomy, and recent molecular phylogenetic work has supported this hypothesis. We produced and analyzed a data set of 6 genes and almost 300 individuals to assess the evolutionary history of the family. Although phylogenetic analysis required extensive adjustment of program settings, we ultimately produced a well-resolved phylogeny for the family. The resulting phylogeny provided strong support for major subclades within the family but extensive paraphyly of genera. Only 3 genera represented by more than 3 species were monophyletic. Biogeographic reconstruction indicated a mainland Asian origin for the family and most major clades. Colonization of Africa, Sundaland, and the Philippines occurred relatively late in the familys history and was mostly unidirectional. Several putative babbler genera, such as Robsonius, Malia, Leonardina, and Micromacronus are only distantly related to the Timaliidae.

Origin and diversification of Philippine bulbuls

We examine the origin and diversification of Philippine bulbuls using a phylogenetic framework. Maximum likelihood and Bayesian methods are used to construct trees from DNA sequences of two mitochondrial and two nuclear genes obtained from 11 Philippine bulbul species as well as 32 other Asian and African taxa. The study finds eight independent colonization events of bulbuls to the Philippines, including one clade comprising Philippine members of the genus Ixos that underwent extensive diversification within the archipelago. Each Philippine clade of bulbuls invaded either the Palawan region or the oceanic islands of the Philippines, but not both. Genetic data reveal at least five lineages that warrant recognition as full species. This study underscores how Philippine avian diversity is currently underestimated and highlights the need for further phylogenetic studies in other Philippine bird groups.

Tectonic collision and uplift of Wallacea triggered the global songbird radiation

Songbirds (oscine passerines) are the most species-rich and cosmopolitan bird group, comprising almost half of global avian diversity. Songbirds originated in Australia, but the evolutionary trajectory from a single species in an isolated continent to worldwide proliferation is poorly understood. Here, we combine the first comprehensive genome-scale DNA sequence data set for songbirds, fossil-based time calibrations, and geologically informed biogeographic reconstructions to provide a well-supported evolutionary hypothesis for the group. We show that songbird diversification began in the Oligocene, but accelerated in the early Miocene, at approximately half the age of most previous estimates. This burst of diversification occurred coincident with extensive island formation in Wallacea, which provided the first dispersal corridor out of Australia, and resulted in independent waves of songbird expansion through Asia to the rest of the globe. Our results reconcile songbird evolution with Earth history and link a major radiation of terrestrial biodiversity to early diversification within an isolated Australian continent.

Phylogeny of Gekko from the Northern Philippines, and Description of a New Species from Calayan Island

Abstract We use mitochondrial gene sequences to estimate relationships among Gekko populations from the northern Philippines. These data, plus morphological and biogeographical evidence, suggest that the Babuyan and Batanes island groups (north of Luzon Island) are inhabited by a minimum of six distinct evolutionary lineages, only two of which (Gekko porosus Taylor from the Batanes and Gekko crombota from Babuyan Claro) have been formally recognized as distinct species. In this paper, we provide a description of another new species, the endemic Gekko from Calayan Island. This geographically isolated species is diagnosed on the basis of a distinct color pattern, body size, scalation, and significant divergence in mitochondrial gene sequences. The new species has been found on rocky outcrops and limestone caves in forested areas and on trunks of mature forest trees ≤500 m above sea level, away from the islands coast. Given the history of geological isolation of Calayan Island and the distinctiveness of the endemic Gekko populations there, we are confident in diagnosing this gecko as a unique evolutionary lineage; it is unlikely that this species will be found on neighboring land masses. The remaining major islands of the Babuyans group (Camiguin Norte, Fuga, and Dalupiri islands) each contain similarly distinct endemic species that await description; additional surveys throughout the Batanes and Babuyan islands will be necessary to arrive at an estimate of total species diversity for this isolated gekkonid radiation.

Genomic approaches to understanding population divergence and speciation in birds

ABSTRACT The widespread application of high-throughput sequencing in studying evolutionary processes and patterns of diversification has led to many important discoveries. However, the barriers to utilizing these technologies and interpreting the resulting data can be daunting for first-time users. We provide an overview and a brief primer of relevant methods (e.g., whole-genome sequencing, reduced-representation sequencing, sequence-capture methods, and RNA sequencing), as well as important steps in the analysis pipelines (e.g., loci clustering, variant calling, whole-genome and transcriptome assembly). We also review a number of applications in which researchers have used these technologies to address questions related to avian systems. We highlight how genomic tools are advancing research by discussing their contributions to 3 important facets of avian evolutionary history. We focus on (1) general inferences about biogeography and biogeographic history, (2) patterns of gene flow and isolation upon secondary contact and hybridization, and (3) quantifying levels of genomic divergence between closely related taxa. We find that in many cases, high-throughput sequencing data confirms previous work from traditional molecular markers, although there are examples in which genome-wide genetic markers provide a different biological interpretation. We also discuss how these new data allow researchers to address entirely novel questions, and conclude by outlining a number of intellectual and methodological challenges as the genomics era moves forward.

Molecular phylogeny and insular biogeography of the lowland tailorbirds of Southeast Asia (Cisticolidae: Orthotomus)

The lowland tailorbirds of Southeast Asia (Orthotomus) offer an excellent opportunity for comparative biogeography because of their diversity in the Greater Sunda and Philippine islands. We reconstructed the phylogeny of all species in the genus using maximum likelihood, Bayesian, and coalescent methods on DNA sequences of three gene segments: an autosomal intron (TGF), a Z-linked intron (MUSK), and a mitochondrial coding gene (ND2). Although resolution is low in parts of the phylogeny, several well defined clades emerge. When considered in light of distribution, these clades indicate that the Greater Sunda and Philippine islands were occupied early in Orthotomus history by the ancestors of O. sericeus in the Greater Sundas and O. frontalis in the Philippines. Subsequently, tailorbirds diversified further in each island group: O. atrogularis, O. ruficeps, and O. sepium arose in the Greater Sundas, and O. castaneiceps castaneiceps, O. c. chloronotus, O. derbianus, O. samarensis, O. nigriceps, and O. cinereiceps in the Philippines. Among the continental taxa (including Sundaic birds), the older lineages (O. sutorius and O. sericeus) are habitat generalists and the recently evolved taxa are more specialized. In the Philippines, several taxa once considered conspecific with O. atrogularis turn out to be highly divergent species (>9% in ND2). Indeed, all Philippine allospecies are well diverged from one another. This finding supports the recent assertion of higher-than-appreciated bird endemicity in the Philippines.

Phylogeography of the Variable Dwarf-Kingfisher Ceyx lepidus (Aves: Alcedinidae) Inferred from Mitochondrial and Nuclear DNA Sequences

ABSTRACT. We reconstructed the phylogeographic relationships of the Variable Dwarf-Kingfisher (Ceyx lepidus) using DNA sequence data. Maximum likelihood and Bayesian analysis methods were used to reconstruct trees from a multilocus data set of all 15 named subspecies of the Ceyx lepidus species complex. The concatenated data-set length was 2,471 base pairs and included two mitochondrial genes and two noncoding nuclear introns. Support for the monophyly of C. lepidus was equivocal. We instead found support for a clade including all C. lepidus subspecies plus two endemic Philippine taxa: C. argentatus and C. cyanopectus. Relationships among subspecific taxa were not well resolved, and many nodes were collapsed into polytomies suggesting a rapid and widespread colonization. In situ diversification likely played a role in generating current diversity within four archipelagos: the Philippines, Malukus, Bismarcks, and Solomons. Some biogeographic patterns recovered for the Solomon Islands taxa match those seen in other bird species, such as the close relationship of taxa on Bougainville, Choiseul, and Isabel. By contrast, the sister relationship between populations on Guadalcanal and the New Georgia Group is novel. We discuss species limits and make taxonomic recommendations to treat all 15 subspecies of C. lepidus as species.

The amphibians and reptiles of Luzon Island, Philippines, VIII: the herpetofauna of Cagayan and Isabela Provinces, northern Sierra Madre Mountain Range

Abstract We provide the first report on the herpetological biodiversity (amphibians and reptiles) of the northern Sierra Madre Mountain Range (Cagayan and Isabela provinces), northeast Luzon Island, Philippines. New data from extensive previously unpublished surveys in the Municipalities of Gonzaga, Gattaran, Lasam, Santa Ana, and Baggao (Cagayan Province), as well as fieldwork in the Municipalities of Cabagan, San Mariano, and Palanan (Isabela Province), combined with all available historical museum records, suggest this region is quite diverse. Our new data indicate that at least 101 species are present (29 amphibians, 30 lizards, 35 snakes, two freshwater turtles, three marine turtles, and two crocodilians) and now represented with well-documented records and/or voucher specimens, confirmed in institutional biodiversity repositories. A high percentage of Philippine endemic species constitute the local fauna (approximately 70%). The results of this and other recent studies signify that the herpetological diversity of the northern Philippines is far more diverse than previously imagined. Thirty-eight percent of our recorded species are associated with unresolved taxonomic issues (suspected new species or species complexes in need of taxonomic partitioning). This suggests that despite past and present efforts to comprehensively characterize the fauna, the herpetological biodiversity of the northern Philippines is still substantially underestimated and warranting of further study.

The phylogenetic position of some Philippine ''babblers'' spans the muscicapoid and sylvioid bird radiations

The Philippines is characterized by a high rate of endemism among its terrestrial vertebrates, including enigmatic genera with uncertain affinities. In a recent comprehensive study of the avian family of Timaliidae (babblers), it was shown that three putative babbler genera endemic to the Philippines (Leonardina, Robsonius, and Micromacronus) are distant relatives of Timaliidae. With additional DNA sequences from new samples and data from Genbank, we attempt to determine the phylogenetic affinities of these three genera and examine the resulting implications for biogeography and avian endemism in the Philippines. Well-supported phylogenies recover the three genera in three different families spanning the sylvioid and muscicapoid radiations of passerine birds. Leonardina groups with Muscicapidae and is most closely related to other isolated montane endemic species in Southeast Asia. Robsonius appears to be an early offshoot of Locustellidae. Micromacronus belongs in Cisticolidae, but its position in the family is unresolved. Contrary to implications based on traditional taxonomy, the Philippine archipelago appears to have played a minor role in the diversification of babblers.

DIVERSIFICATION OF AN ENDEMIC SOUTHEAST ASIAN GENUS: PHYLOGENETIC RELATIONSHIPS OF THE SPIDERHUNTERS (NECTARINIIDAE: ARACHNOTHERA)

ABSTRACT. The phylogeny of spiderhunters (Nectariniidae: Arachnothera) was reconstructed by comparing mitochondrial and nuclear DNA sequences of all currently recognized species and with broad geographic sampling of two particularly variable species complexes, the Little Spiderhunter (Arachnothera longirostra) and the streaky spiderhunters (A. modesta and A. affinis). It appears to be a relatively old group, whose diversification was not caused by recent sea-level changes. However, the modern, highly sympatric distribution of the large species in the Sunda lowlands was probably a result of dispersal via recent land bridges. Within the highly variable A. longirostra group, there are substantially diverged taxa in the Philippines that should be considered different species. Within the A. affinis—modesta complex, there are three distinct species and a closely related fourth, which describe a clear allopatric distribution: A. affinis in Java, A. modesta in the rest of the Sunda lowlands (except Sabah), A. magna in the Malayan highlands and mainland Southeast Asia, and A. everetti in the Bornean highlands and Sabah. Depending on whether mitochondrial or nuclear genes were compared, monophyly of the genus was disrupted by a single outgroup sunbird (Hypogramma hypogrammicum) or by all outgroup sunbirds included in the study. The discrepancy between nuclear and mitochondrial results is probably a case of deep coalescence and will require additional markers for resolution.