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The Long-tailed Rosefinch Carpodacus sibiricus breeds in two widely disjunct areas in East Asia. Five subspecies are usually recognized. The plumage differences between different subspecies groups within C. sibiricus are notable, suggesting it may actually comprise more than one species. However, proper taxonomic assessment has not been carried out, probably in part due to the scarcity and lack of museum specimens of some of these subspecies. In the current study, we re-evaluated the taxonomy of the C. sibiricus complex using molecular, acoustic and morphological data. Using mitochondrial DNA, the samples formed two clades, a northern clade (C. s. sibiricus, C. s. ussuriensis and C. s. sanguinolentus) and a southern clade (C. s. henrici and C. s. lepidus). We also revealed that C. s. ussuriensis and C. s. sanguinolentus are mostly indistinguishable based on mitochondrial markers. This north/south split pattern is congruent with the pattern of plumage differences among north and south taxa. However, no diagnosable differences were found in their songs. We propose synonymizing C. s. ussuriensis with C. s. sanguinolentus. At the species level, we suggest splitting the complex into two species, with Siberian Long-tailed Rosefinch C. sibiricus comprising C. s. sibiricus and C. s. sanguinolentus, and Chinese Long-tailed Rosefinch C. lepidus comprising C. l. henrici and C. l. lepidus.

Aim: Variation in species diversity among different geographical areas may result from differences in speciation and extinction rates, immigration and time for diversification. An area with high species diversity may be the result of a high net diversification rate, multiple immigration events from adjacent regions, and a long time available for the accumulation of species (known as the 'time-for-speciation effect'). Here, we examine the relative importance of the three aforementioned processes in shaping the geographical diversity patterns of a large radiation of passerine birds.Location: Global.Taxon: Babblers (Aves: Passeriformes).Methods: Using a comprehensive phylogeny of extant species (similar to 90% sampled) and distributions of the world's babblers, we reconstructed their biogeographical history and analysed the diversification dynamics. We examined how species richness correlates with the timing of regional colonization, the number of immigration events and the rate of speciation within all 13 geographical distribution regions.Results: We found that babblers likely originated in the Sino-Himalayan Mountains (SHM) in the early Miocene, suggesting a long time for diversification and species accumulation within the SHM. Regression analyses showed the regional diversity of babblers can be well explained by the timing of the first colonization within of these areas, while differences in rates of speciation or immigration have far weaker effects. Nonetheless, the rapid speciation of Zosterops during the Pleistocene has accounted for the increased diversification and accumulation of species in the oceanic islands.Main Conclusions: Our results suggest that the global diversity patterns of babblers have predominantly been shaped by the time-for-speciation effect. Our findings also support an origin centred in tropical and subtropical parts of the SHM, with a cradle of recent diversification in the oceanic islands of the Indo-Pacific and Indian Ocean regions, which provides new insights into the generation of global biodiversity hotspots.

We re‐evaluated the taxonomy of the Striated Prinia Prinia crinigera–Brown Prinia P. polychroa complex using molecular, morphological and vocal analyses. The extensive seasonal, sexual, age‐related, geographical and taxon‐specific variation in this complex has never before been adequately studied. As no previous genetic or vocal analyses have focused on this group, misinterpretation of taxonomic signals from limited conventional morphological study alone was likely. Using mitochondrial and nuclear DNA, we found that P. crinigera sensu lato (s.l.) comprises two non‐sister groups of taxa (Himalayan crinigera and Chinese striata groups) that differ substantially morphologically and vocally and that are broadly sympatric in Yunnan Province, China. Prinia polychroa cooki (Myanmar) and P. p. rocki (southern Vietnam) are each morphologically, vocally and genetically distinct. Thai, Cambodian and Laotian populations formerly ascribed to P. p. cooki are morphologically and vocally most similar to and most closely related to Javan P. p. polychroa, and require a new name, proposed here. Prinia p. bangsi of Yunnan is part of the crinigera group rather than of P. polychroa, and hence there is no evidence for sympatry between P. polychroa s.l. and P. crinigera s.l., nor of the occurrence of P. polychroa in mainland China or Taiwan. We recommend the recognition of five species in the complex, with the following suggestions for new English names: Himalayan Prinia P. crinigera sensu stricto (s.s.; with subspecies striatula, crinigera, yunnanensis and bangsi); Chinese Prinia P. striata (subspecies catharia, parumstriata and striata); Burmese Prinia P. cooki (monotypic); Annam Prinia P. rocki (monotypic) and Deignan's Prinia P. polychroa s.s. (subspecies Javan polychroa and the new Southeast Asian taxon). This study underlines the importance of using multiple datasets for the elucidation of diversity of cryptic bird species and their evolutionary history and biogeography.

In the absence of nuclear-genomic differentiation between two populations, deep mitochondrial divergence (DMD) is a form of mito-nuclear discordance. Such instances of DMD are rare and might variably be explained by unusual cases of female-linked selection, by male-biased dispersal, by “speciation reversal” or by mitochondrial capture through genetic introgression. Here, we analyze DMD in an Asian Phylloscopus leaf warbler (Aves: Phylloscopidae) complex. Bioacoustic, morphological, and genomic data demonstrate close similarity between the taxa affinis and occisinensis, even though DMD previously led to their classification as two distinct species. Using population genomic and comparative genomic methods on 45 whole genomes, including historical reconstructions of effective population size, genomic peaks of differentiation and genomic linkage, we infer that the form affinis is likely the product of a westward expansion in which it replaced a now-extinct congener that was the donor of its mtDNA and small portions of its nuclear genome. This study provides strong evidence of “ghost introgression” as the cause of DMD, and we suggest that “ghost introgression” may be a widely overlooked phenomenon in nature.

Taxa classified as subspecies may in fact be cryptic species. We assessed the taxonomic status of the Blue‐throated Flycatcher Cyornis rubeculoides complex in India, which consists of several forms with similar plumages and song. We used mitochondrial and nuclear DNA, plumage traits, and detailed song analysis to ascertain the taxonomic status of the different forms. The molecular data identified three primary clades: (1) in the west Himalayan foothills, (2) at higher elevations in the northeast hill states of Meghalaya, Nagaland and Mizoram, and (3) at lower elevations in the northeastern hills of Meghalaya and the east Himalayas of Arunachal Pradesh. The western clade represents nominate C. rubeculoides rubeculoides. The high‐elevation eastern clade was considered to be C. rubeculoides rogersi, because it included a sample from this subspecies from near the type locality in southwest Myanmar. These two sister clades had an estimated divergence time of 1.5 million years (my). The low‐elevation east clade has previously been assigned to C. rubeculoides, but we showed it is closely related to the Hainan Blue Flycatcher Cyornis hainanus, formerly thought to breed only further east, with an estimated divergence time of only ~0.8 my. This clade may represent a subspecies of C. hainanus or, given reports of widespread sympatry with C. hainanus in Thailand, a distinct species, Cyornis dialilaemus. However, more research is advocated, including molecular data, from the area of overlap. Songs were remarkably similar across all taxa. In playback experiments, C. r. rubeculoides in the west responded to all taxa. This is in agreement with recent work demonstrating that song differences and responses to songs are not always a good indicator of the progress of reproductive isolation.

Despite the ongoing taxonomic revolution incorporating multiple species delimitation methods, knowledge gaps persist in the taxonomy of comparatively well-studied animal groups such as birds. Morph ...

As species serve as basic units of study in many fields of biology, assessments of species limits are fundamental for such studies. Here, we used a multilocus dataset and different coalescent-based methods to analyze species delimitation and phylogenetic relationships in the Brownish-flanked Bush Warbler Horornis fortipes complex, which is widespread in the Sino-Himalayan region. We also examined the vocal and morphometric divergence within this complex. Our genetic results suggested that Horornis fortipes is composed of at least three independently evolving lineages, which diverged 1.1–1.8 million years ago. However, these lineages have hardly diverged in song or morphometrics and only very slightly in plumage. Our result indicate that there are three incipient species in Horonis fortipes complex diverged in central Himalayas and Hengduan Mountains, but not between the continent and Taiwan island.

Prinias (Cisticolidae:Prinia) are resident warblers of open areas across Africa and Asia and include many polytypic species whose species limits have not been seriously reevaluated recently. Based on an integrative taxonomic analysis of morphology, song, and mitochondrial DNA (mtDNA), we suggest that 2 species should be recognized in the Graceful Prinia (Prinia gracilis) complex. In addition, our morphological analyses show the existence of a well-marked undescribed form in southeastern Somalia, which we name herein as a new subspecies. Prinia gracilisis a small, drab, long-tailed species with streaking above and plain pale underparts that has been suggested to fall into 2 groups: the southwestern nominate group (from Egypt to Oman) and the northeastern lepida group (from Turkey through India). However, the characters presented to justify this grouping are variable and show a mosaic pattern, and whether genetic and vocal differences exist is unknown. We found consistent between-group song differences, with the nominate group giving consistently longer inter-phrase intervals, whereas the members of the lepida group sing an essentially continuous reel. An mtDNA tree suggests a deep split between the nominate and lepida groups, with a coalescence time between these clades of similar to 2.2 million years ago. Vocal and mtDNA analyses provided evidence that the northeastern Arabian Peninsula taxon carpenteri belongs to the lepida group. We found that, of all the morphological characters proposed, only proportions and tail barring and spotting relatively consistently distinguish the 2 groups. However, these characters strongly suggest that the eastern Arabian Peninsula is populated by taxa of both the gracilis and lepida groups, in different areas, but we lack genetic and bioacoustic data to corroborate this. Although further study is needed in potential contact zones, we suggest that 2 species should be recognized in the P. gracilis complex, and we propose the retention of the English name Graceful Prinia for P. gracilis sensu stricto, while we suggest that P. lepida be known as Delicate Prinia. Shared first authorship: Per Alström and Pamela C Rasmussen

The eastern subspecies of Greater Short-toed Lark Calandrella brachydactyla dukhunensis has recently been considered a separate species, Calandrella dukhunensis, by several authors based on molecular data. We present supporting evidence for this treatment based on studies of morphology, vocalisations and song-flight, and also present new data on other aspects of its biology based on field studies. We show that its breeding distribution is considerably smaller than previously thought, and is restricted to the eastern half of Mongolia and, marginally, neighbouring parts of China and perhaps Russia.

Background The White-browed Shortwing (Brachypteryx montana) is widespread from the central Himalayas to the southeast Chinese mainland and the island of Taiwan, the Philippines and Indonesia. Multiple subspecies are recognised, and several of these have recently been suggested to be treated as separate species based on differences in morphology and songs. Methods We here analyse plumage, morphometrics, songs, two mitochondrial and two nuclear markers, and geographical distributions of the two mainland Asian taxa B. m. cruralis and B. m. sinensis and the Taiwanese B. m. goodfellowi. Results We conclude that these differ congruently in morphology, songs and DNA. Male B. m. goodfellowi is the most divergent in plumage (sexually monomorphic, unlike the two others; male similar to female), and B. m. cruralis and B. m. sinensis differ in male plumage maturation. The song of B. m. cruralis is strongly divergent from the others, whereas the songs of B. m. sinensis and B. m. goodfellowi are more similar to each other. Brachypteryx m. sinensis and B. m. goodfellowi are sisters, with an estimated divergence time 4.1 million years ago (mya; 95% highest posterior distribution [HPD] 2.8–5.5 mya), and B. m. cruralis separated from these two 5.8 mya (95% HPD 4.1–7.5 mya). We also report notable range extensions of B. m. sinensis as well as sympatry between this taxon and B. m. cruralis in Sichuan Province, China. Brachypteryx m. montana from Java is found to be more closely related to Lesser Shortwing (B. leucophris) and Rusty-bellied Shortwing (B. hyperythra) than to the mainland Asian and Taiwanese taxa. Conclusion Our data support a recent proposal to treat the three mainland Asian and Taiwanese taxa as three species, separate from B. montana sensu stricto: B. cruralis (central Himalayas to south central China and south Vietnam), B. sinensis (north central to southeastern part of mainland China) and B. goodfellowi (Taiwan Island).

The widespread Old World avian family Locustellidae ('grassbirds and allies') comprises 62 extant species in 11 genera. In the present study, we used one mitochondrial and, for most species, four nuclear loci to infer the phylogeny of this family. We analysed 59 species, including the five previously unsampled genera plus two genera that had not before been analysed in a densely sampled dataset. This study revealed extensive disagreement with current taxonomy; the genera Bradypterus, Locustella, Megalurus, Megalurulus and Schoenicola were all found to be non-monophyletic. Non-monophyly was particularly pronounced for Megalurus, which was widely scattered across the tree. Three of the five monotypic genera (Amphilais, Buettikoferella and Malia) were nested within other genera; one monotypic genus (Chaetornis) formed a clade with one of the two species of Schoenicola; whereas the position of the fifth monotypic genus (Elaphrornis) was unresolved. Robsonius was confirmed as sister to the other genera. We propose a phylogenetically informed revision of genus-level taxonomy, including one new generic name. Finally, we highlight several non-monophyletic species complexes and deep intra-species divergences that point to conflict in taxonomy and suggest an underestimation of current species diversity in this group.

DNA phylogenies have gradually shed light on the phylogenetic relationships of the large babbler group. We focus in this study on the family Leiothrichidae (laughingthrushes and song babblers), whi ...

Generally, genotypes and phenotypes are expected to be spatially congruent; however, in widespread species complexes with few barriers to dispersal, multiple contact zones, and limited reproductive isolation, discordance between phenotypes and phylogeographic groups is more probable. Wagtails (Motacilla) are a genus of birds with striking plumage pattern variation across the Old World. Up to 13 subspecies are recognized within a single species, yet previous studies using mitochondrial DNA have supported polyphyletic phylogeographic groups that are inconsistent with subspecies plumage characteristics. In this study, we investigate the link between phenotypes and genotype by taking a phylogenetic approach. We use genome-wide SNPs, nuclear introns, and mitochondrial DNA to estimate population structure, isolation by distance, and species relationships. Together, our genetic sampling includes complete species-level sampling and comprehensive coverage of the three most phenotypically diverse Palearctic species. Our study provides strong evidence for species-level patterns of differentiation, however population-level differentiation is less pronounced. SNPs provide a robust estimate of species-level relationships, which are mostly corroborated by a combined analysis of mtDNA and nuclear introns (the first time-calibrated species tree for the genus). However, the mtDNA tree is strongly incongruent and is considered to misrepresent the species phylogeny. The extant wagtail lineages originated during the Pliocene and the Eurasian lineage underwent rapid diversification during the Pleistocene. Three of four widespread Eurasian species exhibit an east-west divide that contradicts both subspecies taxonomy and phenotypic variation. Indeed, SNPs fail to distinguish between phenotypically distinct subspecies within the M. alba and M. flava complexes, and instead support geographical regions, each of which is home to two or more different looking subspecies. This is a major step towards our understanding of wagtail phylogeny compared to previous analyses of fewer species and considerably less sequence data.

Patterns of heterogeneous genomic differentiation have been well documented between closely related species, with some highly differentiated genomic regions ("genomic differentiation islands") spread throughout the genome. Differential levels of gene flow are proposed to account for this pattern, as genomic differentiation islands are suggested to be resistant to gene flow. Recent studies have also suggested that genomic differentiation islands could be explained by linked selection acting on genomic regions with low recombination rates. Here, we investigate genomic differentiation and gene-flow patterns for autosomes using RAD-seq data between two closely related species of long-tailed tits (Aegithalos bonvaloti and A. fuliginosus) in both allopatric and contact zone populations. The results confirm recent or ongoing gene flow between these two species. However, there is little evidence that the genomic regions that were found to be highly differentiated between the contact zone populations are resistant to gene flow, suggesting that differential levels of gene flow is not the cause of the heterogeneous genomic differentiation. Linked selection may be the cause of genomic differentiation islands between the allopatric populations with no or very limited gene flow, but this could not account for the heterogeneous genomic differentiation between the contact zone populations, which show evidence of recent or ongoing gene flow.

Aspects of bird song have been shown to correlate with morphological and ecological features, including beak and body size, and habitat. Here we study evolution of song length and song frequency among 30 species belonging to the Cettiidae. Frequency is negatively correlated with body size, and song length increases with latitude. Although migration distance correlates with latitude, the association of song length with latitude is only present within the non-migratory species, implying the association is not a consequence of migration. We place these correlations in a historical framework to show that the body size-frequency association arose early in the group, but the latitude-song length association is more evolutionarily labile. We suggest that latitudinal correlates of song length may reflect increased importance of sexual selection by female choice.

Diversification of avifaunas associated with savannah and steppes appears to correlate with open habitats becoming available, starting in the Miocene. Few comparative analyses exist for families fo ...

The Rufous-headed Robin Larvivora ruficeps is one of the world's rarest and least known birds. We summarize the known records since it was first described in 1905 from Shaanxi Province, central China. All subsequent Chinese records are from seven adjacent localities in nearby Sichuan Province. We studied its phylogenetic position for the first time using mitochondrial and nuclear markers for all species of Larvivora and a broad selection of other species in the family Muscicapidae. Our results confirmed that L. ruficeps is appropriately placed in the genus Larvivora, and suggested that it is sister to the Rufous-tailed Robin Larvivora sibilans, with these two forming a sister clade to a clade comprising both the Japanese Robin Larvivora akahige and Ryukyu Robin Larvivora komadori. Siberian Blue Robin Larvivora cyane and Indian Blue Robin Larvivora brunnea form the sister clade to the other Larvivora species. In contrast, song analyses indicated that the song of L. ruficeps is most similar to that of L. komadori, whereas the song of L. sibilans is relatively more similar to that of L. akahige, and songs of L. cyane and L. brunnea closely resemble each other. We used ecological niche modelling to estimate the suitable habitats of L. ruficeps based on the records from breeding grounds, suggesting that north and central Sichuan, south Gansu, south Shaanxi and south-east Tibet are likely to contain the most suitable habitats for this species.

A tree based on the mitochondrial cyt b gene for 278 samples from throughout the range of the Eurasian Reed Warbler Acrocephalus scirpaceus ¿ African Reed Warbler A. baeticatus complex shows wellsupported geographically structured divergence for eight distinct lineages. The phylogenetic structuring together with the clarification of priority, provided by sequence data from seven type specimens, suggests that both taxonomy and distribution boundaries are in need of revision. The Iberian and Moroccan populations form a well-supported clade, and we propose that these are treated as taxonomically distinct, under the name ambiguus (Brehm, 1857). We propose that the names scirpaceus, fuscus, avicenniae, ambiguus, minor, cinnamomeus, hallae and baeticatus are used for the well supported clades in the complex, which we recommend to treat as one polytypic species, A. scirpaceus, pending studies of gene flow and assortative mating in the contact zones., We thank the Ministry of Environment and Tourism, Namibia for granting the necessary permits, and Ursula Franke-Bryson for kindly providing feathers for dna analyses from Namibia.

Niche segregation plays a critical role in the speciation process, but determining the extent to which taxa are geographically or ecologically isolated is challenging. In this study, we use stable ...

The Wallcreeper Tichodroma muraria is usually placed in a monotypic family or subfamily within the superfamily Certhioidea, with assumed close relationships to Certhia (treecreepers), Sitta (nuthatches) and Salpornis (spotted creepers). Previous studies have suggested that Tichodroma is most closely related to Sitta, alternatively to Salpornis. We analysed the relationships of Tichodroma using two mitochondrial and five nuclear loci. The tree based on concatenated sequences strongly supported a sister relationship between Tichodroma and Sitta, as well as between Salpornis and Certhia. However, species tree analysis (MP-EST) was unable to resolve these relationships, and although the concatenation tree remains the best hypothesis, more data are needed to corroborate this.

We studied the phylogeography and population history of the white wagtail Motacilla alba, which has a vast breeding range, covering areas with different Pleistocene climatic histories. The mitochon ...

The taxonomy of the Narcissus Flycatcher Ficedula narcissina-Yellow-rumped Flycatcher Ficedula zanthopygia complex from East Asia has long been debated. Most authors recognize two species: F. narcissina, with the subspecies narcissina (most of Japan and Sakhalin Island), owstoni (south Japanese islands) and elisae (northeast China) and F. zanthopygia (monotypic), although species status has been proposed for elisae and sometimes for owstoni. Here, we revise the taxonomy of this complex based on an integrative approach utilizing morphology, songs and mitochondrial and nuclear DNA for all taxa. All taxa were diagnosably different in plumage, and there were also structural differences among them, although the northernmost populations of owstoni (sometimes recognized as jakuschima and shonis) were somewhat intermediate in plumage, structure and male plumage maturation between southern populations of owstoni and narcissina. All taxa had different songs, and a discriminant function analysis of four song variables correctly classified 100% of all songs. A strongly supported phylogeny was recovered based on three mitochondrial genes and three nuclear introns (total of 3543 bp), revealing a sister relationship between F. zanthopygia and the other taxa, between F. n. narcissina and F. n. owstoni, and between F. n. elisae and F. n. narcissina + F. n. owstoni. The corrected COI distances among the three F. narcissina subspecies ranged from 2.8% (narcissina-owstoni) to 8.2% (narcissina-elisae). We suggest that the congruent differences in multiple independent traits and the deep genetic divergences among the four taxa in the F. narcissina-F. zanthopygia complex support treatment of all of these taxa as separate species. However, we acknowledge the paucity of data for F. owstoni and recommend further studies of this taxon. We suggest listing both F. elisae and F. owstoni, which have small and fragmented populations, as globally threatened.

The leaf warbler radiation (Aves: Phylloscopidae) has undergone a c. 50% increase in the number of recognised species over the last three decades, mainly as a result of analyses of vocalisations and DNA. Using a multilocus dataset for all of the species in this family, and multispecies coalescent-based as well as concatenation methods, we provide the first complete species-level phylogeny for this important group, as well as an estimate of the timing of diversification. The most recent common ancestor for the family was dated at 11.7 million years ago (mya) (95% highest posterior density 9.8–13.7 mya), and divergence times between sister species ranged from 0.5 mya (0.3–0.8 mya) to 6.1 mya (4.8–7.5 mya). Based on our results, we support synonymising Seicercus with Phylloscopus, which results in a monogeneric Phylloscopidae. We discuss the pros and cons of this treatment, and we argue against proliferation of taxonomic names, and conclude that a large monogeneric Phylloscopidae leads to the fewest taxonomic changes compared to traditional classifications. We briefly discuss morphological evolution in the light of the phylogeny. The time calibrated phylogeny is a major improvement compared to previous studies based on a smaller number of species and loci and can provide a basis for future studies of other aspects of phylloscopid evolution.

International audience; To better determine the history of modern birds, we performed a genome-scale phylogenetic analysis of 48 species representing all orders of Neoaves using phylogenomic methods created to handle genome-scale data. We recovered a highly resolved tree that confirms previously controversial sister or close relationships. We identified the first divergence in Neoaves, two groups we named Passerea and Columbea, representing independent lineages of diverse and convergently evolved land and water bird species. Among Passerea, we infer the common ancestor of core landbirds to have been an apex predator and confirm independent gains of vocal learning. Among Columbea, we identify pigeons and flamingoes as belonging to sister clades. Even with whole genomes, some of the earliest branches in Neoaves proved challenging to resolve, which was best explained by massive protein-coding sequence convergence and high levels of incomplete lineage sorting that occurred during a rapid radiation after the Cretaceous-Paleogene mass extinction event about 66 million years ago.

The Blackthroat (or Blackthroated Blue Robin) Calliope obscura (previously Luscinia obscura) is one of the world’s rarest “robins”. It is extremely poorly known, with only a handful of records since it was firstly described in the 1890s. In 2011–2012, a series of field investigations were carried out in nature reserves in the Qinling Mountains, Shaanxi Province, China. During these surveys, a total of 14 males were found in 2011 and 24 males and 2 females in 2012 in the national nature reserves of Foping and Changqing. Based on these observations, we here describe the Blackthroat’s vocalizations and habitat choice and review its distribution and conservation status. We also provide some notes on the plumage, especially of the female, and the nesting behaviour of this species. The present report confirms that the Blackthroat breeds on the southern slope of the Qinling Mountains in large, dense expanses of dwarf bamboo with scattered coniferous and broadleaved trees above 2,100 m. Our observations suggest that it is more numerous than previously believed, although it appears to be highly localized. The locally dense populations found in 2011 and 2012 and the vast expanses of suitable habitat suggest that the breeding population might be healthy.

The systematics of the avian superfamily Sylvioidea are reviewed, focusing on studies of relationships among families and within genera, more superficially on taxonomic studies at the species level. For the families Bernieridae and Phylloscopidae, new analyses based on already published sequence data are presented. Our understanding of relationships has been vastly improved in recent years due to a large number of molecular studies. However, the relationships among the different families remain largely obscured, probably mainly as a result of rapid divergence of the different primary lineages (families). Also, species level taxonomy has been much improved in recent years due to a large number of studies applying molecular markers and/or vocalizations and other life-history data. It seems likely that the number of species will continue to increase, as new groups are being studied with modern integrative methods.

Soft songs have been detected in many songbirds, but in most species, research on soft songs has lagged behind studies of broadcast songs. In this study, we describe the acoustic features of a soft song in the brownish-flanked bush warbler Cettia fortipes. Compared with the broadcast song, the warbler's soft song was characterized by a lower minimum frequency and longer duration, and it had a higher proportion of rapid frequency modulation notes. Using playback experiments, some in combination with mounted specimens, we found different responses to soft and broadcast songs, and we found that soft song can predict aggressive escalation (attack). We conclude that the soft song is an aggressive signal in this species.

We estimated a phylogeny for 10 taxa currently placed in four polytypic species that col-lectively encompass the African ‘brown buntings’: Cape Bunting Emberiza capensis,Cinnamon-breasted Bunting Emberiza tahapisi, Lark-like Bunting Emberiza impetuani andHouse Bunting Emberiza striolata. We made use of the mitochondrial cytochrome b geneand the nuclear introns 6–7 of ornithine decarboxylase (ODC), and intron 2 of myoglo-bin. There was substantial cytochrome b sequence divergence between taxa currentlytreated as conspecific: sahari vs. striolata (2.6–3.1% (uncorrected-p); 3.0–3.6%(HKY + I)), and goslingi vs. tahapisi (4.4–4.7% (uncorrected-p); 5.4–5.9% (HKY + I)).The degree of divergence of the nuclear loci among taxa was limited, and these locilacked reciprocal monophyly, most likely as a consequence of incomplete lineage sorting.A single representative of the taxon septemstriata, generally treated as a member of thedark-throated tahapisi group, here appears to be genetically consistent with the grey-throated goslingi, and may be of hybrid origin. All other taxa allocated to E. striolata andE. tahapisi make up four reciprocally monophyletic groups consistent with sahari, striola-ta, tahapisi and goslingi, respectively. The extent of genetic evidence suggests that thesetaxa have been evolving as separate evolutionary lineages for a long time. This is furthermanifested in several morphological and vocal characteristics described previously, andwe propose that these divergent taxa be treated as separate species: Cinnamon-breastedBunting Emberiza tahapisi, Gosling’s Bunting Emberiza goslingi, Striolated Bunting Ember-iza striolata and House Bunting Emberiza sahari. We do not propose any taxonomicchanges regarding Emberiza impetuani or Emberiza capensis.Keywords: classification, concatenation, gene trees, hybridization, M

The African lark genus Heteromirafra is thought to consist of three threatened species inhabiting mid-altitude grasslands, one in South Africa and two in the Horn of Africa. One of the latter, Archer’s Lark H. archeri of Somaliland, has not been seen with certainty since 1922. We surveyed its type locality as well as a nearby area of grassland east of Jijiga in adjacent north-eastern Ethiopia, where sightings of Heteromirafra larks have recently been made. First, we used a combination of morphological and molecular evidence to show that these recent sightings refer to the same taxon as Archer’s Lark. Second, we used a combination of morphological, molecular and vocal evidence to show that these populations are conspecific with the Liben (Sidamo) Lark H. sidamoensis of southern Ethiopia, but that the Horn of Africa populations are highly distinct from Rudd’s Lark H. ruddi of South Africa. Third, we suggest that the extent and quality of their habitat in north-eastern Ethiopia is small and poor, and that the type locality of Archer’s Lark in Somaliland has been completely transformed. Taken together, these results imply that there is a single species of Heteromirafra in the Horn of Africa (for which the scientific name H. archeri has priority, and which we suggest retains the English name Liben Lark), consisting of two tiny populations separated by 590 km of apparently unsuitable habitats. Environmental niche models suggest that there are no environmentally similar locations elsewhere within the region. Despite the discovery of a second population, the Liben Lark remains a highly threatened species in urgent need of conservation intervention to avert the extinction of both of its populations.

The phylogenetic relationships of the avian family Muscicapidae (Old World chats and flycatchers) have historically been enigmatic and remain an active area of study. Widespread instances of non-monophyly resulting from misleading morphological and behavioral similarities have merited numerous taxonomic revisions to the group. Here we report one such instance with regard to the Rusty-tailed Flycatcher Muscicapa ruficauda, which has recently been placed in the newly proposed monotypic genus Ripleyia, due to inferred sister relationship to the genus Muscicapa and related genera (Voelker et al., 2016a). This name was later replaced by Ripleyornis, as it was realized that Ripleyia is a junior homonym of a genus of Mollusca (Voelker et al., 2016b). Using a Bayesian phylogenetic assessment of the Muscicapidae with near-complete taxon sampling of the genus Ficedula for five loci, along with an acoustic comparison of M. ruficauda to a subset of other flycatcher species, we show that this species should be reassigned to the genus Ficedula and accordingly that the names Ripleyia and Ripleyornis are both junior synonyms of Ficedula.

Background: The Plain-backed Thrush Zoothera mollissima breeds in the Himalayas and mountains of central China. It was long considered conspecific with the Long-tailed Thrush Zoothera dixoni, until these were shown to be broadly sympatric. Methods: We revise the Z. mollissima–Z. dixoni complex by integrating morphological, acoustic, genetic (two mitochondrial and two nuclear markers), ecological and distributional datasets. Results: In earlier field observations, we noted two very different song types of “Plain-backed” Thrush segregated by breeding habitat and elevation. Further integrative analyses congruently identify three groups: an alpine breeder in the Himalayas and Sichuan, China (“Alpine Thrush”); a forest breeder in the eastern Himalayas and northwest Yunnan (at least), China (“Himalayan Forest Thrush”); and a forest breeder in central Sichuan (“Sichuan Forest Thrush”). Alpine and Himalayan Forest Thrushes are broadly sympatric, but segregated by habitat and altitude, and the same is probably true also for Alpine and Sichuan Forest Thrushes. These three groups differ markedly in morphology and songs. In addition, DNA sequence data from three non-breeding specimens from Yunnan indicate that yet another lineage exists (“Yunnan Thrush”). However, we find no consistent morphological differences from Alpine Thrush, and its breeding range is unknown. Molecular phylogenetic analyses suggest that all four groups diverged at least a few million years ago, and identify Alpine Thrush and the putative “Yunnan Thrush” as sisters, and the two forest taxa as sisters. Cytochrome b divergences among the four Z. mollissima sensu lato (s.l.) clades are similar to those between any of them and Z. dixoni, and exceed that between the two congeneric outgroup species. We lectotypify the name Oreocincla rostrata Hodgson, 1845 with the Z. mollissima sensu stricto (s.s.) specimen long considered its type. No available name unambiguously pertains to the Himalayan Forest Thrush. Conclusions: The Plain-backed Thrush Z. mollissima s.l. comprises at least three species: Alpine Thrush Z. mollissima s.s., with a widespread alpine breeding distribution; Sichuan Forest Thrush Z. griseiceps, breeding in central Sichuan forests; and Himalayan Forest Thrush, breeding in the eastern Himalayas and northwest Yunnan (at least), which is described herein as a new species. “Yunnan Thrush” requires further study.

The relationships among Old World chats and flycatchers (Muscicapidae) have recently been clarified in two independent molecular phylogenetic studies (Sangster et al. 2010, Zuccon & Ericson 2010). Both studies recovered a well-supported clade of predominantly blue flycatchers of the genera Niltava, Cyornis, Eumyias and Cyanoptila, some species traditionally included in the genus Rhinomyias and one species traditionally included in Ficedula (F. monileger). The family-group name Niltavinae Sangster, Alström, Forsmark & Olsson, 2010, was introduced for this clade (Sangster et al. 2010). Unfortunately, our description of the new taxon Niltavinae did not include a diagnosis (sensu ICZN article 13.1.1) and thus inadvertently made this name unavailable for nomenclatural purposes (Zuccon 2011). The supplementary data associated with our original paper included evidence by which the taxon Niltavinae can be diagnosed: a 1 base pair (bp) deletion in the ornithine decarboxylase gene, as shown in Figure S1 in the online version (doi:10.1016/j.ympev.2010.07.008), but this was not mentioned in the printed version of our paper. Given that Niltavinae Sangster, Alström, Forsmark & Olsson, 2010 represents a nomen nudum, and the clade for which this name was intended thus remains unnamed, we here provide the following description

Historical geological events and climatic changes are believed to have played important roles in shaping the current distribution of species. However, sympatric species may have responded in different ways to such climatic fluctuations. Here we compared genetic structures of two corvid species, the Azure-winged Magpie Cyanopica cyanus and the Eurasian Magpie Pica pica, both widespread but with different habitat dependence and some aspects of breeding behavior. Three mitochondrial genes and two nuclear introns were used to examine their co-distributed populations in East China and the Iberian Peninsula. Both species showed deep divergences between these two regions that were dated to the late Pliocene/early Pleistocene. In the East Chinese clade of C. cyanus, populations were subdivided between Northeast China and Central China, probably since the early to mid-Pleistocene, and the Central subclade showed a significant pattern of isolation by distance. In contrast, no genetic structure was found in the East China populations of P. pica. We suggest that the different patterns in the two species are at least partly explained by ecological differences between them, especially in habitat preference and perhaps also breeding behavior. These dissimilarities in life history traits might have affected the dispersal and survival abilities of these two species differently during environmental fluctuations.

135‐142). Here we analyse songs and calls from throughout the range of the Arctic Warbler, and conclude that these differ markedly and consistently among the populations representing the three mitochondrial clades. Kurile populations, for which no sequence data are available, are shown to belong to the second clade. To determine the correct application of available scientific names, mitochondrial DNA was sequenced from three name-bearing type specimens collected on migration or in the winter quarters. Based on the congruent variation in mitochondrial DNA, morphology and vocalizations, we propose that three species be recognized: Arctic Warbler Phylloscopus borealis (sensu stricto) (continental Eurasia and Alaska), Kamchatka Leaf Warbler Phylloscopus examinandus (Kamchatka (at least the southern part), Sakhalin, Hokkaido and Kurile Islands), and Japanese Leaf Warbler Phylloscopus xanthodryas (Japan except Hokkaido).

The Scrub Warbler, which inhabits arid areas from North Africa to western Asia, has long been thought to be closely related to cisticolid warblers. However, analyses based on two mitochondrial and four nuclear loci place this species sister to the mainly Asian Cettiidae (bush warblers, tesias, etc.). Superficial morphological similarity to cisticolid warblers has previously clouded the species true relationship. Detailed morphology, such as facial bristles and claw and footpad structure, also supports a closer relationship to Cettiidae and some other non-cisticolid warblers.

We present molecular evidence that Neumann’s Warbler Hemitesia neumanni is deeply nested within the Cettiidae. The species’ distribution in the Albertine Rift of East Africa is intriguing, as the family Cettiidae is principally an Asian radiation. This disjunct distribution could be a result of colonization of Africa by long-distance dispersal, or the Cettiidae may at some point in the past have had a much larger geographical distribution that also covered parts of Africa.

We present the first study of the warbler family Acrocephalidae based on one mitochondrial and three nuclear DNA loci, in total approximately 2900 bp, including most or all of the species in three (Acrocephalus, Hippolais and Chloropeta) of the four genera and one species in the fourth genus (Nesillas) in this family. All three genera were suggested to be non-monophyletic, although the non-monophyly of Acrocephalus is not fully convincingly demonstrated. Six major clades were found, which agreed largely with the results from two earlier mitochondrial studies, and for which the names Hippolais, Iduna, Acrocephalus, Calamocichla, Notiocichla and Calamodus have been used. However, the results also revealed some new constellations, due to better resolution of deeper nodes and the inclusion of more taxa. The taxonomic implications are discussed.

We demonstrate the importance of using multiple criteria in species delimitations, whatever the conceptual base for species delimitation. We do this by studying plumage, biometrics, egg coloration, song, mitochondrial DNA and habitat/altitudinal distribution in the Spotted Bush Warbler Bradypterus thoracicus (Blyth) complex, and by conducting playback experiments. Taxa that we suggest are best treated as separate species [B. thoracicus (Blyth), B. davidi (La Touche) and B. kashmirensis (Sushkin)] differ in most or all of these aspects, particularly in song and mitochondrial DNA, while those that we treat as subspecies (suschkini) or synonyms (przevalskii) differ slightly and only in morphology.

The phylogeny of the avian genus Emberiza and the monotypic genera Latoucheornis, Melophus and Miliaria (collectively the Old World Emberizini), as well as representatives for the New World Emberizini, the circumpolar genera Calcarius and Plectrophenax and the four other generally recognized tribes in the subfamily Emberizinae was estimated based on the mitochondrial cytochrome b gene and introns 6-7 of the nuclear ornithine decarboxylase (ODC) gene. Our results support monophyly of the Old World Emberizini, but do not corroborate a sister relationship to the New World Emberizini. Calcarius and Plectrophenax form a clade separated from the other Emberizini. This agrees with previous studies, and we recommend the use of the name Calcariini. Latoucheornis, Melophus and Miliaria are nested within Emberiza, and we therefore propose they be synonymized with Emberiza. Emberiza is divided into four main clades, whose relative positions are uncertain, although a sister relation between a clade with six African species and one comprising the rest of the species (30, all Palearctic) is most likely. Most clades agree with traditional, morphology-based, classifications. However, four sister relationships within Emberiza, three of which involve the previously recognized Latoucheornis, Melophus and Miliaria, are unpredicted, and reveal cases of strong morphological divergence. In contrast, the plumage similarity between adult male Emberiza (formerly Latoucheornis) siemsseni and the nominate subspecies of the New World Junco hyemalis is shown to be the result of parallel evolution. A further case of parallel plumage evolution, between African and Eurasian taxa, is pointed out. Two cases of discordance between the mitochondrial and nuclear data with respect to branch lengths and genetic divergences are considered to be the result of introgressive hybridization.

Phylloscopus hainanus is a distinctive new species in the family Sylviidae. The species is endemic to Hainan Island, China, where it is restricted to tropical forest in mountains above 600 m. The most distinctive plumage features are the rather deep yellow underparts and the mainly white outer rectrices.

A new species of leaf warbler (Sylviidae: Phylloscopus) is described from the mountain Emei Shan, Sichuan Province, China. Morphologically it is extremely similar to, yet significantly different from, the sympatric Blyth's Leaf Warbler Phylloscopus reguloides and White-tailed Leaf Warbler Phylloscopus davisoni. Songs and calls are both very different from P. reguloides and P. davisoni, and playback tests confirmed the specific status of the new species.

The Golden-spectacled Warbler, usually treated as a single species, Seicercus burkii, is widely distributed in mountains of southern Asia. We argue that it should be treated as five different species, one of which is described here for the first time. Two species occur in sympatry in the Himalayas. The other three species are found in eastern Asia, where all three are sympatric in China, and two of these breed sympatrically also in Vietnam. In Burma and adjacent parts of India, one of the Himalayan species is sympatric with one of the east Asian species. We describe differences in morphology, vocalizations, altitudinal distribution and habitat between these five species, as well as results from playback tests, which show that all sympatric taxa differ and appear to be reproductively isolated from each other.

Morphological, vocal and genetic divergence in the Cettia acanthizoides complex (Aves: Cettiidae)

The Himalayan mountain range is one of the most species-rich areas in the world, harboring about 8% of the world's bird species. In this study, we compare the relative importance of immigration versus in situ speciation to the build-up of the Himalayan avifauna, by evaluating the biogeographic history of the Phylloscopus/Seicercus warblers, a speciose clade that is well represented in Himalayan forests. We use a comprehensive, multigene phylogeny in conjunction with dispersal-vicariance analysis to discern patterns of speciation and dispersal within this clade. The results indicate that virtually no speciation has occurred within the Himalayas. Instead, several speciation events are attributed to dispersal into the Himalayas followed by vicariance between the Himalayas and China/Southeast Asia. Most, perhaps all, of these events appear to be pre-Pleistocene. The apparent lack of speciation within the Himalayas stands in contrast to the mountain-driven Pleistocene speciation suggested for the Andes and the East African mountains.

The avian genus Calandrella (larks) was recently suggested to be non-monophyletic, and was divided into two genera, of which Calandrella sensu stricto comprises 4-5 species in Eurasia and Africa. We analysed mitochondrial cytochrome b (cytb) and nuclear Restriction-site Associated DNA (RAD) sequences from all species, and for cytb we studied 21 of the 22 recognised subspecies, with the aim to clarify the phylogenetic relationships within the genus and to compare large-scale nuclear sequence patterns with a widely used mitochondrial marker. Cytb indicated deep splits among the currently recognised species, although it failed to support the interrelationships among most of these. It also revealed unexpected deep divergences within C. brachydactyla, C. blanfordi/C. erlangeri, C. cinerea, and C. acutirostris. It also suggested that both C. brachydactyla and C. blanfordi, as presently circumscribed, are paraphyletic. In contrast, most of the many subspecies of C. brachydactyla and C. cinerea were unsupported by cytb, although two populations of C. cinerea were found to be genetically distinct. The RAD data corroborated the cytb tree (for the smaller number of taxa analysed) and recovered strongly supported interspecific relationships. However, coalescence analyses of the RAD data, analysed in SNAPP both with and without an outgroup, received equally strong support for two conflicting topologies. We suggest that the tree rooted with an outgroup - which is not recommended for SNAPP - is more trustworthy, and suggest that the reliability of analyses performed without any outgroup species should be thoroughly evaluated. We also demonstrate that degraded museum samples can be phylogenetically informative in RAD analyses following careful bioinformatic treatment. We note that the genus Calandrella is in need of taxonomic revision.

Background The Russet Bush Warbler Locustella (previously Bradypterus) mandelli complex occurs in mountains in the eastern Himalayas, southern China, Vietnam, the Philippines, and Indonesia. The taxonomy has been debated, with one (L. seebohmi) to four (L. seebohmi, L. mandelli, L. montis and L. timorensis) species having been recognised. Methods We used an integrative approach, incorporating analyses of morphology, vocalizations and a molecular marker, to re-evaluate species limits in the L. mandelli complex. Results We found that central Chinese L. mandelli differed from those from India through northern Southeast Asia to southeast China in plumage, morphometrics and song. All were easily classified by song, and (wing + culmen)/tail ratio overlapped only marginally. Both groups were reciprocally monophyletic in a mitochondrial cytochrome b (cytb) gene tree, with a mean divergence of 1.0 ± 0.2%. They were sympatric and mostly altitudinally segregated in the breeding season in southern Sichuan province. We found that the Mt Victoria (western Myanmar) population differed vocally from other L. mandelli, but no specimens are available. Taiwan Bush Warbler L. alishanensis was sister to the L. mandelli complex, with the most divergent song. Plumage, vocal and cytb evidence supported the distinctness of the south Vietnamese L. mandelli idonea. The Timor Bush Warbler L. timorensis, Javan Bush Warbler L. montis and Benguet Bush Warbler L. seebohmi differed distinctly in plumage, but among-population song variation in L. montis exceeded the differences between some populations of these taxa, and mean pairwise cytb divergences were only 0.5-0.9%. We also found that some L. montis populations differed morphologically. Conclusions We conclude that the central Chinese population of Russet Bush Warbler represents a new species, which we describe herein, breeding at mid elevations in Sichuan, Shaanxi, Hubei, Hunan and Guizhou provinces. The taxonomic status of the other allopatric populations is less clear. However, as they differ to a degree comparable with that of the sympatric L. mandelli and the new species, we elevate L. idonea to species status, and retain L. seebohmi and L. montis as separate species, the latter with timorensis as a subspecies. Further research should focus on different populations of L. montis and the Mt Victoria population of L. mandelli.