Intra- and interspecific variation in self-control capacities of parrots in a delay of gratification task.

Forgoing immediate satisfaction for higher pay-offs in the future (delayed gratification) could be adaptive in situations that wild animals may encounter. To explain species-differences in self-control, hypotheses based on social complexity, feeding ecology, brain size and metabolic rate have been proposed. To explore these hypotheses in a comparative setting, we tested three macaw species (neotropical parrots)-great green macaws (N = 8), blue-throated macaws (N = 6), blue-headed macaws (N = 6)-and the distantly related African grey parrots (afrotropical parrots; N = 8) in a modified rotating tray task, in which subjects are required to inhibit consuming a constantly available low-quality reward in favour of a high-quality reward that becomes available only after an increasing delay (min. 5 s, max. 60 s). All four species successfully waited for a minimum of 8.3 s ± 11.7 s (group level mean ± SD) with African greys reaching a delay of 29.4 ± 15.2 s, and great green macaws-as best performing macaw species-tolerating delays of 20 s ± 8 s. The best performing African grey individual reached a maximum delay of 50 s, whereas, a great green and a blue-throated macaw tolerated a delay of 30 s max. Females tolerated higher maximum delays than males. Engaging in distraction behaviours enhanced waiting performance across species and all birds were able to anticipate the waiting duration. Our results suggest that both feeding and socio-ecological complexity may be a factor in self-control, but further systematically collected comparative data on self-control of different (parrot) species are required to test the evolutionary hypotheses rigorously.

Development of polymorphic microsatellite markers for the Trichoglossus haematodus and cross-species amplification in Trichoglossus moluccanus.

BACKGROUD: Trichoglossus haematodus is the most popular parrots globally and one of the most bred species in Korea's zoos. However, despite its popularity, there are limited studies on the population genetics of this species. METHODS AND RESULTS: In this study, 10 polymorphic microsatellite markers were developed for T. haematodus. The number of alleles ranged from 6 to 9 (mean 7.9). Null alleles were present in two loci (TH-07 and TH-08). The observed heterozygosity ranged from 0.4444 to 1.0000 (mean 0.7000). One locus (TH-08) indicated a significant deviation from the Hardy-Weinberg equilibrium after Bonferroni correction (p < 0.005). The mean inbreeding coefficient (FIS) of the 10 loci was positive, suggesting that there is inbreeding in the population. Since the polymorphism information content (PIC) values were more than 0.7 in all loci, all markers developed in this study were classified as informative. The parentage exclusion probabilities considering all loci were higher than 0.99 in all three cases (P1, P2, and P3). The cross-species amplification of the 10 markers was tested in T. moluccanus, a close relative species of T. haematodus. These markers were also informative for T. moluccanus with PIC values higher than 0.7 in all loci. Additionally, the parentage exclusion probabilities (P1, P2 and P3) for T. moluccanus were above 0.99. However, due to the small number of T. haematodus and T. moluccanus investigated in this study, the 10 microsatellite markers should be analyzed with more individuals of these two species in future studies. CONCLUSIONS: The markers developed in this study might be helpful for investigations of genetic diversity and parentage analysis of T. haematodus and T. moluccanus.

Characterization and Comparative Analysis of Complete Mitogenomes of Three Cacatua Parrots (Psittaciformes: Cacatuidae).

Cacatua alba, Cacatua galerita, and Cacatua goffiniana are parrots of the family Cacatuidae. Wild populations of these species are declining with C. alba listed by the International Union for the Conservation of Nature and Natural Resources (IUCN) as Endangered. In this study, complete mitogenomes were sequenced for a comparative analysis among the Cacatua species, and a detailed analysis of the control region. Mitogenome lengths of C. alba,C. galerita, and C. goffiniana were 18,894, 18,900, and 19,084 bp, respectively. They included 13 protein coding genes, two ribosomal RNA genes, 24 transfer RNA genes, three degenerated genes, and two control regions. Ten conserved motifs were found in three domains within each of the two control regions. For an evolution of duplicated control regions of Cacatua, domain I and the 3' end of domain III experienced an independent evolution, while domain II and most of the regions of domain III was subjected to a concerted evolution. Based on a phylogenetic analysis of 37 mitochondrial genes, the genus Cacatua formed a well-supported, monophyletic, crown group within the Cacatuidae. Molecular dating results showed that Cacatua diverged from other genera of Cacatuinae in the middle of Miocene.

Genomes of Three Closely Related Caribbean Amazons Provide Insight for Species History and Conservation.

Islands have been used as model systems for studies of speciation and extinction since Darwin published his observations about finches found on the Galapagos. Amazon parrots inhabiting the Greater Antillean Islands represent a fascinating model of species diversification. Unfortunately, many of these birds are threatened as a result of human activity and some, like the Puerto Rican parrot, are now critically endangered. In this study we used a combination of de novo and reference-assisted assembly methods, integrating it with information obtained from related genomes to perform genome reconstruction of three amazon species. First, we used whole genome sequencing data to generate a new de novo genome assembly for the Puerto Rican parrot (Amazona vittata). We then improved the obtained assembly using transcriptome data from Amazona ventralis and used the resulting sequences as a reference to assemble the genomes Hispaniolan (A. ventralis) and Cuban (Amazona leucocephala) parrots. Finally, we, annotated genes and repetitive elements, estimated genome sizes and current levels of heterozygosity, built models of demographic history and provided interpretation of our findings in the context of parrot evolution in the Caribbean.

Dissection by genomic and plumage variation of a geographically complex hybrid zone between two Australian non-sister parrot species, Platycercus adscitus and Platycercus eximius.

The study of hybrid zones advances understanding of the speciation process, and approaches incorporating genomic data are increasingly used to draw significant conclusions about the impact of hybridisation. Despite the progress made, the complex interplay of factors that can lead to substantially variable hybridisation outcomes are still not well understood, and many systems and/or groups remain comparatively poorly studied. Our study aims to broaden the literature on avian hybrid zones, investigating a potentially geographically and temporally complex putative hybrid zone between two native Australian non-sister parrot species, the pale-headed and eastern rosellas (Platycercus adscitus and Platycercus eximius, respectively). We analysed six plumage traits and >1400 RADseq loci and detected hybrid individuals and an unexpectedly complex geographic structure. The hybrid zone is larger than previously described due to either observer bias or its movement over recent decades. It comprises different subregions where genetic and plumage signals of admixture vary markedly in their concordance. Evidence of contemporary hybridisation (later generation and backcrossed individuals) both within and beyond the previously defined zone, when coupled with a lack of F1 hybrids and differential patterns of introgression among potentially diagnostic loci, indicates a lack of post-zygotic barriers to gene flow between species. Despite ongoing gene flow, species boundaries are likely maintained largely by strong pre-mating barriers. These findings are discussed in detail and future avenues for research into this system are proposed, which would be of benefit to the speciation and hybrid zone literature.

The influence of molecular markers and methods on inferring the phylogenetic relationships between the representatives of the Arini (parrots, Psittaciformes), determined on the basis of their complete mitochondrial genomes.

BACKGROUND: Conures are a morphologically diverse group of Neotropical parrots classified as members of the tribe Arini, which has recently been subjected to a taxonomic revision. The previously broadly defined Aratinga genus of this tribe has been split into the 'true' Aratinga and three additional genera, Eupsittula, Psittacara and Thectocercus. Popular markers used in the reconstruction of the parrots' phylogenies derive from mitochondrial DNA. However, current phylogenetic analyses seem to indicate conflicting relationships between Aratinga and other conures, and also among other Arini members. Therefore, it is not clear if the mtDNA phylogenies can reliably define the species tree. The inconsistencies may result from the variable evolution rate of the markers used or their weak phylogenetic signal. To resolve these controversies and to assess to what extent the phylogenetic relationships in the tribe Arini can be inferred from mitochondrial genomes, we compared representative Arini mitogenomes as well as examined the usefulness of the individual mitochondrial markers and the efficiency of various phylogenetic methods. RESULTS: Single molecular markers produced inconsistent tree topologies, while different methods offered various topologies even for the same marker. A significant disagreement in these tree topologies occurred for cytb, nd2 and nd6 genes, which are commonly used in parrot phylogenies. The strongest phylogenetic signal was found in the control region and RNA genes. However, these markers cannot be used alone in inferring Arini phylogenies because they do not provide fully resolved trees. The most reliable phylogeny of the parrots under study is obtained only on the concatenated set of all mitochondrial markers. The analyses established significantly resolved relationships within the former Aratinga representatives and the main genera of the tribe Arini. Such mtDNA phylogeny can be in agreement with the species tree, owing to its match with synapomorphic features in plumage colouration. CONCLUSIONS: Phylogenetic relationships inferred from single mitochondrial markers can be incorrect and contradictory. Therefore, such phylogenies should be considered with caution. Reliable results can be produced by concatenated sets of all or at least the majority of mitochondrial genes and the control region. The results advance a new view on the relationships among the main genera of Arini and resolve the inconsistencies between the taxa that were previously classified as the broadly defined genus Aratinga. Although gene and species trees do not always have to be consistent, the mtDNA phylogenies for Arini can reflect the species tree.

Inference of Evolutionary Jumps in Large Phylogenies using Lévy Processes.

Although it is now widely accepted that the rate of phenotypic evolution may not necessarily be constant across large phylogenies, the frequency and phylogenetic position of periods of rapid evolution remain unclear. In his highly influential view of evolution, G. G. Simpson supposed that such evolutionary jumps occur when organisms transition into so-called new adaptive zones, for instance after dispersal into a new geographic area, after rapid climatic changes, or following the appearance of an evolutionary novelty. Only recently, large, accurate and well calibrated phylogenies have become available that allow testing this hypothesis directly, yet inferring evolutionary jumps remains computationally very challenging. Here, we develop a computationally highly efficient algorithm to accurately infer the rate and strength of evolutionary jumps as well as their phylogenetic location. Following previous work we model evolutionary jumps as a compound process, but introduce a novel approach to sample jump configurations that does not require matrix inversions and thus naturally scales to large trees. We then make use of this development to infer evolutionary jumps in Anolis lizards and Loriinii parrots where we find strong signal for such jumps at the basis of clades that transitioned into new adaptive zones, just as postulated by Simpson's hypothesis. [evolutionary jump; Lévy process; phenotypic evolution; punctuated equilibrium; quantitative traits.

Revising the phylogenetic position of the extinct Mascarene Parrot Mascarinus mascarin (Linnaeus 1771) (Aves: Psittaciformes: Psittacidae).

The phylogenetic position of the extinct Mascarene Parrot Mascarinus mascarin from La Réunion has been unresolved for centuries. A recent molecular study unexpectedly placed M. mascarin within the clade of phenotypically very different Vasa parrots Coracopsis. Based on DNA extracted from the only other preserved Mascarinus specimen, we show that the previously obtained cytb sequence is probably an artificial composite of partial sequences from two other parrot species and that M. mascarin is indeed a part of the Psittacula diversification, placed close to P. eupatria and P. wardi.

The status of three little known names proposed by Miranda-Ribeiro (1926) and the synonymization of Pyrrhura snethlageae Joseph & Bates, 2002 (Psittaciformes: Psittacidae: Arinae).

The presence of complex geographic variation among species allied to the absence of clear morphological breaks among populations has led to many poorly defined taxa in the genus Pyrrhura (Psittacidae: Arinae: Arini). This article addresses nomenclature of the Pyrrhura picta species complex, particularly the identity of three little-known names introduced by Miranda-Ribeiro (1926): Pyrrhura luciani ochrotis, P. l. pallescens and P. l. melanoides. We show that these names represent subjective synonyms of Pyrrhura snethlageae Joseph & Bates, 2002. Furthermore, we argue that Pyrrhura pallescens (Miranda-Ribeiro, 1926) n. comb. should be considered as the valid senior name because: (1) there is no doubt about the type locality (near the type locality of P. snethlageae) and, as a consequence, it "will best serve stability and universality of nomenclature" as recommended by the International Code of Zoological Nomenclatural, and; (2) it corresponds to the most common phenotype usually referred to as P. snethlageae, the species name that has been used for bird specimens from this region. Those considering P. snethlageae lucida Arndt, 2008 as a valid taxon should refer to this population as Pyrrhura pallescens lucida (Arndt, 2008) n. comb. In a near future P. pallescens melanoides (Miranda-Ribeiro, 1926) n. comb. is likely to be considered the best senior synonym of P. p. lucida given its type locality (Teles Pires River, MT).

Considerations on the type specimens and type locality of Pyrrhura roseifrons (Gray, 1859) (Psittacidae).

The Rose-fronted Parakeet Pyrrhura roseifrons (Gray, 1859) is a poorly known species that occurs in lowland forest of western Amazonia, from eastern Peru to western Brazil and northern Bolivia (Collar 1997; Forshaw 2010). Like many Pyrrhura species, it is mostly green, and has a pale auricular patch, red tail and belly, but presents a bright red head, distinct from any other close relatives within the genus. It was long considered a subspecies of P. picta (Statius Miller, 1776) (Arndt 1983; Arndt 1996; Collar 1997; Juniper & Parr, 1998) and only recently has its specific rank been restored (Joseph 2000, 2002; Ribas et al. 2006) and accepted in the literature (Dickinson & Remsen 2013; del Hoyo & Collar 2014; Remsen et al. 2015).

The first fossil parrot (Aves, Psittaciformes) from Siberia and its implications for the historical biogeography of Psittaciformes.

Modern parrots (crown Psittaciformes) are a species-rich group of mostly tropical and subtropical birds with a very limited fossil record. A partial tarsometatarsus from the late Early Miocene of Siberia (Baikal Lake) is the first pre-Quaternary find of crown Psittaciformes in Asia (and Siberia in particular) and is also the northern-most find of this bird order worldwide. This find documents a broad geographical distribution of parrots during the warmest phase of the Miocene (the so-called 'Miocene Climatic Optimum'), which has implications for the historical biogeography of Psittaciformes. The presence of parrots on both sides of the Pacific Ocean at the end of the Early Miocene implies a (most probably eastwards) trans-Beringian dispersal which likely took place about 16-18 Ma. The broad Eurasian distribution of parrots in the past further supports a hypothesis that ancestors of modern genera Coracopsis and Agapornis could reach Africa from Eurasia.

Rearrangement and evolution of mitochondrial genomes in parrots.

Mitochondrial genome rearrangements that result in control region duplication have been described for a variety of birds, but the mechanisms leading to their appearance and maintenance remain unclear, and their effect on sequence evolution has not been explored. A recent survey of mitochondrial genomes in the Psittaciformes (parrots) found that control region duplications have arisen independently at least six times across the order. We analyzed complete mitochondrial genome sequences from 20 parrot species, including representatives of each lineage with control region duplications, to document the gene order changes and to examine effects of genome rearrangements on patterns of sequence evolution. The gene order previously reported for Amazona parrots was found for four of the six independently derived genome rearrangements, and a previously undescribed gene order was found in Prioniturus luconensis, representing a fifth clade with rearranged genomes; the gene order resulting from the remaining rearrangement event could not be confirmed. In all rearranged genomes, two copies of the control region are present and are very similar at the sequence level, while duplicates of the other genes involved in the rearrangement show signs of degeneration or have been lost altogether. We compared rates of sequence evolution in genomes with and without control region duplications and did not find a consistent acceleration or deceleration associated with the duplications. This could be due to the fact that most of the genome rearrangement events in parrots are ancient, and additionally, to an effect of body size on evolutionary rate that we found for mitochondrial but not nuclear sequences. Base composition analyses found that relative to other birds, parrots have unusually strong compositional asymmetry (AT- and GC-skew) in their coding sequences, especially at fourfold degenerate sites. Furthermore, we found higher AT skew in species with control region duplications. One potential cause for this compositional asymmetry is that parrots have unusually slow mtDNA replication. If this is the case, then any replicative advantage provided by having a second control region could result in selection for maintenance of both control regions once duplicated.

Complete mitochondrial genome of Military Macaw (Ara militaris): its comparison with mitogenomes of two other Ara species.

The Military Macaw is one of the eight species of the genus Ara. The genus is one of six genera, which form morphologically diverse group termed as Macaws. Parrots of this group differ in body size on demand of the genus and species. Six of Ara species are classified as large Macaws. Based on morphological similarities and differences, these species can be segregated into three pairs according to their plumage coloration. Representative mitochondrial genomes were sequenced only for A. glaucogularis (blue and yellow coloration) and A. macao (predominantly red/scarlet). Ara militaris is one of two predominantly green species and full mitochondrial genome of considered species was sequenced in this study. It's comparison with A. glaucogularis and A. macao mitogenomes revealed higher degree of identity between militaris and macao sequences than between militaris and glaucogularis mtDNAs. Ara militaris mitogenome will be indispensable to refine the phylogenetic relationships within Macaw group.

Complete mitochondrial genome of Mitred Conure (Psittacara mitratus): its comparison with mitogenome of Socorro Conure (Psittacara brevipes).

The Mitred Conure (Psittacara mitratus) is a species native to the forests and woodlands in the Andes in central Peru, south through west-central Bolivia, to northwestern Argentina. The genus Psittacara have recently been distinguished on the basis of molecular revision of the Aratinga species. In consequence, it became one of the nine genera, which form morphologically diverse group termed as Conures. Although, the number of species/subspecies belonging to Psittacara remains controversial, it is now believed that the genus is represented by 11 species. Taxonomic position of Mitred Conure was determined by molecular research and makes the species crucial for examination of evolutionary diversification of the genus. Therefore, we sequenced complete mitochondrial genome of P. mitratus mitogenome to gain a source of comprehensive molecular data. It will be indispensable to refine the phylogenetic relationships within "Psittacara group" as well as within the tribe Arini.

Complete mitochondrial genome of Red-throated Conure (Psittacara rubritorquis): its comparison with mitogenome of Socorro Conure (Psittacara brevipes).

According to some taxonomists the Red-throated Conure (Psittacara rubritorquis) is considered a subspecies of Green Conure (Psittacara holochlora). Some other classifications treat rubritorquis as a separate species based on relatively minor morphological differences between both species/subspecies. So far, taxonomic position of P. rubritorquis was determined by molecular researches using only ND2 gene sequence or incomplete combined mitochondrial ND2, COI and CYTB gene sequences. Obtained outcomes found that P. rubritorquis should be treated as a subspecies of P. holochlora. However, the lack of P. h. brewsterii and P. h. strenua samples as well as incompleteness of combined mitochondrial sequence do not exclude opposite scenario. Therefore, we sequenced P. rubritorquis mitogenome to gain a source of molecular data appropriate for future examination of evolutionary diversification of the P. holochlora group.

Molecular systematics of the Cape Parrot (Poicephalus robustus): implications for taxonomy and conservation.

The taxonomic position of the Cape Parrot (Poicephalus robustus robustus) has been the focus of much debate. A number of authors suggest that the Cape Parrot should be viewed as a distinct species separate from the other two P. robustus subspecies (P. r. fuscicollis and P. r. suahelicus). These recommendations were based on morphological, ecological, and behavioural assessments. In this study we investigated the validity of these recommendations using multilocus DNA analyses. We genotyped 138 specimens from five Poicephalus species (P. cryptoxanthus, P. gulielmi, P. meyeri, P. robustus, and P. rueppellii) using 11 microsatellite loci. Additionally, two mitochondrial (cytochrome oxidase I gene and 16S ribosomal RNA) and one nuclear intron (intron 7 of the ß-fibrinogen gene) markers were amplified and sequenced. Bayesian clustering analysis and pairwise FST analysis of microsatellite data identified P. r. robustus as genetically distinct from the other P. robustus subspecies. Phylogenetic and molecular clock analyses on sequence data also supported the microsatellite analyses, placing P. r. robustus in a distinct clade separate from the other P. robustus subspecies. Molecular clock analysis places the most recent common ancestor between P. r. robustus and P. r. fuscicollis / P. r. suahelicus at 2.13 to 2.67 million years ago. Our results all support previous recommendations to elevate the Cape Parrot to species level. This will facilitate better planning and implementation of international and local conservation management strategies for the Cape Parrot.

DNA Barcoding Identifies Illegal Parrot Trade.

Illegal trade threatens the survival of many wild species, and molecular forensics can shed light on various questions raised during the investigation of cases of illegal trade. Among these questions is the identity of the species involved. Here we report a case of a man who was caught in a Brazilian airport trying to travel with 58 avian eggs. He claimed they were quail eggs, but authorities suspected they were from parrots. The embryos never hatched and it was not possible to identify them based on morphology. As 29% of parrot species are endangered, the identity of the species involved was important to establish a stronger criminal case. Thus, we identified the embryos' species based on the analyses of mitochondrial DNA sequences (cytochrome c oxidase subunit I gene [COI] and 16S ribosomal DNA). Embryonic COI sequences were compared with those deposited in BOLD (The Barcode of Life Data System) while their 16S sequences were compared with GenBank sequences. Clustering analysis based on neighbor-joining was also performed using parrot COI and 16S sequences deposited in BOLD and GenBank. The results, based on both genes, indicated that 57 embryos were parrots (Alipiopsitta xanthops, Ara ararauna, and the [Amazona aestiva/A. ochrocephala] complex), and 1 was an owl. This kind of data can help criminal investigations and to design species-specific anti-poaching strategies, and demonstrate how DNA sequence analysis in the identification of bird species is a powerful conservation tool.

Does vocal learning accelerate acoustic diversification? Evolution of contact calls in Neotropical parrots.

Learning has been traditionally thought to accelerate the evolutionary change of behavioural traits. We evaluated the evolutionary rate of learned vocalizations and the interplay of morphology and ecology in the evolution of these signals. We examined contact calls of 51 species of Neotropical parrots from the tribe Arini. Parrots are ideal subjects due to their wide range of body sizes and habitats, and their open-ended vocal learning that allows them to modify their calls throughout life. We estimated the evolutionary rate of acoustic parameters of parrot contact calls and compared them to those of morphological traits and habitat. We also evaluated the effect of body mass, bill length, vegetation density and species interactions on acoustic parameters of contact calls while controlling for phylogeny. Evolutionary rates of acoustic parameters did not differ from those of our predictor variables except for spectral entropy, which had a significantly slower rate of evolution. We found support for correlated evolution of call duration, and fundamental and peak frequencies with body mass, and of fundamental frequency with bill length. The degree of sympatry between species did not have a significant effect on acoustic parameters. Our results suggest that parrot contact calls, which are learned acoustic signals, show evolutionary rates similar to those of morphological traits. This is the first study to our knowledge to provide evidence that change through cultural evolution does not necessarily accelerate the evolutionary rate of traits acquired through life-long vocal learning.

Molecular phylogenetics suggests a New Guinean origin and frequent episodes of founder-event speciation in the nectarivorous lories and lorikeets (Aves: Psittaciformes).

The lories and lorikeets (Aves: Loriinae: Loriini) are a readily recognizable, discrete group of nectarivorous parrots confined to the Indo-Pacific region between Wallace's Line and the Pitcairn Island group in the central-east Pacific Ocean. We present the first phylogenetic analysis of all currently recognized genera in the group using two mitochondrial and five nuclear loci. Our analyses suggest a New Guinean origin for the group at about 10million years ago (95% HPD 4.8-14.8) but this origin must be interpreted within the context of that island's complicated, recent geological history. That is, the origin and early diversification of the group may have taken place as New Guinea's Central Cordillera arose and the final constituent terranes that form present-day New Guinea were accreted. The latter activity may have promoted dispersal as a key element in the group's history. We have detected several instances of dispersal out of New Guinea that we argue constitute instances of founder-event speciation. Some phenotypically cohesive genera are affirmed as monophyletic but other genera are clearly in need of taxonomic dismantlement and reclassification. We recognize Parvipsitta Mathews, 1916 for two species usually placed in Glossopsitta and we advocate transfer of Chalcopsitta cardinalis into Pseudeos Peters, 1935. Other non-monophyletic genera such as Charmosyna, Psitteuteles and, probably, Trichoglossus, require improved taxon sampling and further phylogenetic analysis before their systematics can be resolved. Cursory examination of trait mapping across the group suggests that many traits are ancestral and of little use in determining genus-level systematics.

Phylogenetic analysis of the Australian rosella parrots (Platycercus) reveals discordance among molecules and plumage.

Relationships and species limits among the colourful Australian parrots known as rosellas (Platycercus) are contentious because of poorly understood patterns of parapatry, sympatry and hybridization as well as complex patterns of geographical replacement of phenotypic forms. Two subgenera are, however, conventionally recognised: Platycercus comprises the blue-cheeked crimson rosella complex (Crimson Rosella P. elegans and Green Rosella P. caledonicus), and Violania contains the remaining four currently recognised species (Pale-headed Rosella P. adscitus, Eastern Rosella P. eximius, Northern Rosella P. venustus, and Western Rosella P. icterotis). We used phylogenetic analysis of ten loci (one mitochondrial, eight autosomal and one z-linked) and several individuals per nominal species primarily to examine relationships within the subgenera, especially the relationships and species limits within Violania. Of these, P. adscitus and P. eximius have long been considered sister species or conspecific due to a morphology-based hybrid zone and an early phylogenetic analysis of mitochondrial DNA restriction fragment length polymorphisms. The multilocus phylogenetic analysis presented here supports an alternative hypothesis aligning P. adscitus and P. venustus as sister species. Using divergence rates published in other avian studies, we estimated the divergence between P. venustus and P. adscitus at 0.0148-0.6124MYA and that between the P. adscitus/P. venustus ancestor and P. eximius earlier at 0.1617-1.0816MYA, both within the Pleistocene. Discordant topologies among gene and species trees are discussed and proposed to be the result of historical gene flow and/or incomplete lineage sorting (ILS). In particular, we suggest that discordance between mitochondrial and nuclear data may be the result of asymmetrical mitochondrial introgression from P. adscitus into P. eximius. The biogeographical implications of our findings are discussed relative to similarly distributed groups of birds.