Cryptic species in a colorful genus: Integrative taxonomy of the bush robins (Aves, Muscicapidae, Tarsiger) suggests two overlooked species.

Several cryptic avian species have been validated by recent integrative taxonomic efforts in the Sino-Himalayan mountains, indicating that avian diversity in this global biodiversity hotspot may be underestimated. In the present study, we investigated species limits in the genus Tarsiger, the bush robins, a group of montane forest specialists with high species richness in the Sino-Himalayan region. Based on comprehensive sampling of all 11 subspecies of the six currently recognized species, we applied an integrative taxonomic approach by combining multilocus, acoustic, plumage and morphometric analyses. Our results reveal that the isolated north-central Chinese populations of Tarsiger cyanurus, described as the subspecies albocoeruleus but usually considered invalid, is distinctive in genetics and vocalisation, but only marginally differentiated in morphology. We also found the Taiwan endemic T. indicus formosanus to be distinctive in genetics, song and morphology from T. i. indicus and T. i. yunnanensis of the Sino-Himalayan mountains. Moreover, Bayesian species delimitation using BPP suggests that both albocoeruleus and formosanus merit full species status. We propose their treatment as 'Qilian Bluetail' T. albocoeruleus and 'Taiwan Bush Robin' T. formosanus, respectively.

Complete subspecies-level phylogeny of the Oriolidae (Aves: Passeriformes): Out of Australasia and return.

Old World orioles (Oriolidae) are medium-sized passerine birds confined largely to forested areas of Africa, Eurasia and Australasia. We present a new complete molecular (mtDNA) subspecies level phylogeny of the Oriolidae including all 113 taxa (35 species) together with a backbone phylogeny of 19 taxa from the main Oriolus clades based on (i) 21 nuclear genes, (ii) whole mito-genomes, and (iii) genome-wide ultraconserved elements. We use this phylogeny to assess systematic relationships and the biogeographical history of this avian family. Furthermore, we use morphological measurements to investigate the relationship between size and shape axes and upstream or back-colonization of this extensive island region from Asia. We show that several subspecies or groups of subspecies may warrant species rank and we find a continental example of two morphologically distinct species (O. mellianus/O. traillii) being genetically (mtDNA) very similar. Biogeographically, we confirm previous findings that members of the Oriolidae originated in Australo-Papua. Dispersal out of this area took place around 15 Mya to southeast Asia and Africa, and from Africa to the Palearctic followed by recolonization of the Indonesian and Philippine island region during the Plio-Pleistocene. Recolonisation of the Indonesian and Philippine islands coincided with an increase in body size, which may have facilitated the ability to co-exist with other congenerics.

The evolution of mimicry of friarbirds by orioles (Aves: Passeriformes) in Australo-Pacific archipelagos.

Observations by Alfred Wallace and Jared Diamond of plumage similarities between co-occurring orioles (Oriolus) and friarbirds (Philemon) in the Malay archipelago led them to conclude that the former represent visual mimics of the latter. Here, we use molecular phylogenies and plumage reflectance measurements to test several key predictions of the mimicry hypothesis. We show that friarbirds originated before brown orioles, that the two groups did not co-speciate, although there is one plausible instance of co-speciation among species on the neighbouring Moluccan islands of Buru and Seram. Furthermore, we show that greater size disparity between model and mimic and a longer history of co-occurrence have resulted in a stronger plumage similarity (mimicry). This suggests that resemblance between orioles and friarbirds represents mimicry and that colonization of islands by brown orioles has been facilitated by their ability to mimic the aggressive friarbirds.

The published complete mitochondrial genome of Blue-fronted Redstart (Phoenicurus frontalis) is a chimera and includes DNA from Pink-rumped Rosefinch Carpodacus waltoni eos (Aves: Passeriformes).

The complete mitochondrial genome of Blue-fronted Redstart (Phoenicurus frontalis), GenBank accession number MT360379 (NC_053917), was published by Li and colleages in 2020. Here we show that this mitogenome is actually a chimera containing DNA fragments of both P. frontalis (15,518 bp, 92.5%) and Pink-rumped Rosefinch (Carpodacus waltoni eos, 1258 bp, 7.5%). This mitogenome has been re-used in at least three phylogenies. Our study confirms that mitogenomes are best verified with multiple gene trees, and that any anomalies should be investigated by direct comparison of sequences.

Complete mitochondrial genome MK992912 of Great Knot (Calidris tenuirostris) is a chimera with DNA from Pacific Golden Plover Pluvialis fulva (Aves: Charadriiformes).

A complete mitochondrial genome of Great Knot (Calidris tenuirostris), MK992912, was published by He and colleagues in 2020. Here we show that this mitogenome is actually a chimera containing DNA fragments of both C. tenuirostris (15,567 bp, 92.8%) and Pacific Golden Plover (Pluvialis fulva, 1208 bp, 7.2%). Detecting such errors is possible before publication if each sequenced fragment is separately analyzed phylogenetically before assembling the fragments into a single mitogenome. This mitogenome has been re-used in at least four phylogenies. The error is documented to avoid the perpetuation of erroneous sequence information in the literature.

The spatio-temporal colonization and diversification across the Indo-Pacific by a 'great speciator' (Aves, Erythropitta erythrogaster).

The Indo-Pacific region has arguably been the most important area for the formulation of theories about biogeography and speciation, but modern studies of the tempo, mode and magnitude of diversification across this region are scarce. We study the biogeographic history and characterize levels of diversification in the wide-ranging passerine bird Erythropitta erythrogaster using molecular, phylogeographic and population genetics methods, as well as morphometric and plumage analyses. Our results suggest that E. erythrogaster colonized the Indo-Pacific during the Pleistocene in an eastward direction following a stepping stone pathway, and that sea-level fluctuations during the Pleistocene may have promoted gene flow only locally. A molecular species delimitation test suggests that several allopatric island populations of E. erythrogaster may be regarded as species. Most of these putative new species are further characterized by diagnostic differences in plumage. Our study reconfirms the E. erythrogaster complex as a 'great speciator': it represents a complex of up to 17 allopatrically distributed, reciprocally monophyletic and/or morphologically diagnosable species that originated during the Pleistocene. Our results support the view that observed latitudinal gradients of genetic divergence among avian sister species may have been affected by incomplete knowledge of taxonomic limits in tropical bird species.

Systematic revision of the avian family Cisticolidae based on a multi-locus phylogeny of all genera.

The avian taxon Cisticolidae includes c. 110 species which are distributed throughout the tropical and subtropical parts of the Old World. We estimated the phylogeny of 47 species representing all genera assumed to be part of Cisticolidae based on sequence data from two mitochondrial and two nuclear markers, in total 3495bp. Bayesian inference and maximum likelihood analyses resulted in a generally well-supported phylogeny which clarified the position of several previously poorly known taxa. The placement of Drymocichla, Malcorus, Micromacronus, Oreophilais, Phragmacia, Phyllolais, Poliolais and Urorhipis in Cisticolidae is corroborated, whereas Rhopophilus and Scotocerca are removed from Cisticolidae. Urorhipis and Heliolais are placed in the genus Prinia whereas Prinia burnesii is shown to be part of Timaliidae, and is placed in the genus Laticilla. Although not recovered by all single loci independently, four major clades were identified within Cisticolidae, and one of these is here described as a new taxon (Neomixinae).

The importance of voucher specimens: misidentification or previously unknown mtDNA diversity in Phalacrocorax capillatus (Aves: Phalacrocoracidae)?

A recently published complete mitochondrial genome of Japanese or Temminck's cormorant (Phalacrocorax capillatus) was the first of this species (GenBank accession number LC714913). Comparison of COI sequences shows that this mitogenome clustered with great cormorant (Phalacrocorax carbo) rather than with its sister taxon P. capillatus. This suggests that the mitogenome was either a misidentified P. carbo or represents previously unknown intraspecific diversity in P. capillatus overlapping with that of P. carbo. Unfortunately, no voucher specimen was retained so it remains impossible to distinguish between these alternatives. We suggest that great restraint should be exercised using this mitogenome as a reference for P. capillatus. We reiterate previous pleas to retain voucher specimens for mitogenome sequences to enable re-analysis of the identity of the material.

The published complete mitochondrial genome of Spotted Greenshank (Tringa guttifer) is a chimera with DNA from Red-necked Stint (Calidris ruficollis) (Aves: Charadriiformes).

A recently published complete mitochondrial genome of Spotted Greenshank (Tringa guttifer) was the first DNA sequence of this species (GenBank accession number MK905885, RefSeq number NC_044665; Liu et al. 2019, The complete mitochondrial genome of the Spotted Greenshank Tringa guttifer (Charadriiforemes [sic]: Charadriidae), Mitochondrial DNA Part B. 4:2353-2354). Here we show that this mitogenome is actually a chimera containing DNA fragments of both a Tringa sandpiper (presumably T. guttifer) and the Red-necked Stint (Calidris ruficollis). This mitogenome has been re-used in at least three phylogenies. The error is documented to avoid the perpetuation of erroneous sequence information in the literature.

A new genus for Philydor erythrocercum and P. fuscipenne (Aves: Furnariidae).

N/A.

A new genus for Dasycrotapha plateni and D. pygmaea (Aves: Zosteropidae).

N/A.

A new species of scops-owl (Aves, Strigiformes, Strigidae, Otus) from Príncipe Island (Gulf of Guinea, Africa) and novel insights into the systematic affinities within Otus.

A new species of scops-owl (Aves, Strigiformes, Strigidae, Otus) is described from Príncipe Island, São Tomé and Príncipe (Gulf of Guinea, Africa). This species was discovered for science in 2016, although suspicions of its occurrence gained traction from 1998, and testimonies from local people suggesting its existence could be traced back to 1928. Morphometrics, plumage colour and pattern, vocalisations, and molecular evidence all support the species status of the scops-owl from Príncipe, which is described here as Otusbikegila sp. nov. Phylogenetic analyses suggest that this species descended from the first colonisation of the Gulf of Guinea islands, being sister to the clade including the mainland African Scops-Owl O.senegalensis, and the island endemics Sao Tome Scops-Owl O.hartlaubi and Pemba Scops-Owl O.pembaensis. The most diagnostic trait in the field is its unique call which, curiously, is most similar to a distantly related Otus species, the Sokoke Scops-Owl O.ireneae. The new species occurs at low elevations of the old-growth native forest of Príncipe, currently restricted to the south of the island but fully included within Príncipe Obô Natural Park. Otusbikegila sp. nov. takes the number of single-island endemic bird species of Príncipe to eight, further highlighting the unusually high level of bird endemism for an island of only 139 km2.

ResumoDescrevemos uma nova espécie de mocho-de-orelhas ou kitóli (Strigiformes: Strigidae: Otus) da Ilha do Príncipe, São Tomé e Príncipe (Golfo da Guiné, África). Esta espécie foi descoberta para a ciência apenas em 2016, embora suspeitas da sua existência tenham ganho força a partir de 1998, e testemunhos de habitantes locais sobre a sua ocorrência já estarem documentados em 1928. A morfometria, a cor e padrão da plumagem, as vocalizações e dados moleculares demonstram que esta população de mocho no Príncipe é uma espécie nova, que foi batizada de mocho-do-príncipe (lista mundial) ou kitóli-do-príncipe (nome nacional), Otusbikegila sp. nov. As análises filogenéticas indicam que esta espécie descende da primeira colonização das ilhas do Golfo da Guiné, sendo irmã do clado que inclui o mocho-d'ore­lhas-africano O.senegalensis, do continente, o mocho-de-são-tomé (ou kitóli-de-são-tomé) O.hartlaubi e o mocho-de-pemba O.pembaensis, ambos endémicos das ilhas que lhes dão o nome. No campo, a característica mais diagnóstica é o seu canto único que, curiosamente, é mais parecido com o da espécie de Otus mais afastada, o mocho-de-sokoke O.ireneae. A nova espécie ocorre nas zonas baixas da floresta nativa do Príncipe, atualmente restrita ao sul da ilha, mas totalmente inserida no Parque Natural do Obô do Príncipe. Otusbikegila sp. nov. eleva o número de espécies de aves endémicas restritas ao Príncipe para oito, sublinhando ainda mais o nível extremamente elevado de aves endémicas para uma ilha de apenas 139 km2.

The published complete mitochondrial genome of the milk shark (Rhizoprionodon acutus) is a misidentified Pacific spadenose shark (Scoliodon macrorhynchos) (Chondrichthyes: Carcharhiniformes).

The recently published mitogenome of milk shark Rhizoprionodon acutus (MN602076/NC_046016) was fully resolved in an unexpected phylogenetic position in the original mitogenome announcement, which rendered the genus Scoliodon paraphyletic. Here, we show that this mitogenome is actually that of a misidentified Pacific spadenose shark (Scoliodon macrorhynchos). The error is documented to avoid the perpetuation of erroneous sequence information in the literature.

Sharp Increase of Problematic Mitogenomes of Birds: Causes, Consequences, and Remedies.

Authentic DNA sequences are crucial for reliable evolutionary inference. Concerns about the identification of DNA sequences have been voiced several times in the past but few quantitative studies exist. Mitogenomes play important roles in phylogenetics, phylogeography, population genetics, and DNA identification. However, the large number of mitogenomes being published routinely, often in brief data papers, has raised questions about their authenticity. In this study, we quantify problematic mitogenomes of birds and their reusage in other papers. Of 1,876 complete or partial mitogenomes of birds published until January 1, 2020, the authenticity of 1,559 could be assessed with sequences of conspecifics. Of these, 78 (5.0%) were found to be problematic, including 45 curated reference sequences. Problems were due to misidentification (33), chimeras of two or three species (23), sequencing errors/numts (18), incorrect sequence assembly (1), mislabeling at GenBank but not in the final paper (2), or vice versa (1). The number of problematic mitogenomes has increased sharply since 2012. Worryingly, these problematic sequences have been reused 436 times in other papers, including 385 times in phylogenies. No less than 53% of all mitogenomic phylogenies/networks published until January 1, 2020 included at least one problematic mitogenome. Problematic mitogenomes have resulted in incorrect phylogenetic hypotheses and proposals for unwarranted taxonomic revision, and may have compromised comparative analyses and measurements of divergence times. Our results indicate that a major upgrade of quality control measures is warranted. We propose a comprehensive set of measures that may serve as a new standard for publishing mitogenome sequences.

Multi-locus phylogenetic analysis of Old World chats and flycatchers reveals extensive paraphyly at family, subfamily and genus level (Aves: Muscicapidae).

The chats and flycatchers (Muscicapidae) represent an assemblage of 275 species in 48 genera. Defining natural groups within this assemblage has been challenging because of its high diversity and a paucity of phylogenetically informative morphological characters. We assessed the phylogenetic relationships of 124 species and 34 genera of Muscicapidae, and 20 species of Turdidae, using molecular sequence data from one mitochondrial gene and three nuclear loci, in total 3240bp. Bayesian and maximum likelihood analyses yielded a well-resolved tree in which nearly all basal nodes were strongly supported. The traditionally defined Muscicapidae, Muscicapinae and Saxicolinae were paraphyletic. Four major clades are recognized in Muscicapidae: Muscicapinae, Niltavinae (new family-group name), Erithacinae and Saxicolinae. Interesting relationships recovered by this analysis include: (i) a clade comprising the 'blue' flycatcher genera Niltava, Cyornis, Cyanoptila and Eumyias and some species of Rhinomyias; (ii) the position of Erithacus rubecula in a clade of otherwise exclusively African species; (iii) a close relationship between the shortwing Heinrichia calligyna and the flycatcher Rhinomyias insignis; (iv) a sister-relationship between forktails Enicurus and whistling thrushes Myophonus; and (v) a sister relationship of Ficedula and the 'chats'Monticola, Phoenicurus, Saxicola and Oenanthe. A high number of traditionally defined genera was found to be paraphyletic or polyphyletic.

The published complete mitochondrial genome of Eptesicus serotinus is a chimera of Vespertilio sinensis and Hypsugo alaschanicus (Mammalia: Chiroptera).

The mitogenome of Eptesicus serotinus (Serotine bat) was published in 2013 with GenBank accession number KF111725 and NCBI Reference Sequence number NC_022474. This sequence was placed with Vespertilio sinensis (Asian parti-colored bat) in a COI gene tree but with Hypsugo alashanicus (Alashanian pipistrelle) in a cytochrome b gene tree. Direct comparison of mitogenomes showed that 92.4% of this mitogenome is similar to Vespertilio sinensis, 5.9% to Hypsugo alaschanicus, and that 1.6% of the mitogenome could not be attributed to either species, or any other species. This mitogenome has been re-used in at least 17 phylogenies. Our findings suggest that mitogenomes are best verified with multiple gene trees, followed by direct comparison of sequences. We conclude that greater vigilance is warranted to ensure that problematic sequences do not enter the scientific record, and are not re-used in subsequent studies.

Increasing numbers of bird species result from taxonomic progress, not taxonomic inflation.

The impact and significance of modern taxonomy on other fields in biology have been subjects of much debate. It has been proposed that increasing numbers of vertebrate species are largely owing to 'taxonomic inflation'. According to this hypothesis, newly recognized species result from reinterpretations of species limits based on phylogenetic species concepts (PSCs) rather than from new discoveries. Here, I examine 747 proposals to change the taxonomic rank of birds in the period 1950-2007. The trend to recognize more species of birds started at least two decades before the introduction of PSCs. Most (84.6%) newly recognized species were supported by new taxonomic data. Proposals to recognize more species resulted from application of all six major taxonomic criteria. Many newly recognized species (63.4%) were not based exclusively on PSC-based criteria (diagnosability, monophyly and exclusive coalescence of gene trees). Therefore, this study finds no empirical support for the idea that the increase in species is primarily epistemological rather than data-driven. This study shows that previous claims about the causes and effects of taxonomic inflation lack empirical support. I argue that a more appropriate term for the increase in species is 'taxonomic progress'.

Paraclaravis, a new genus for the Purple-winged and Maroon-chested Ground-doves (Aves: Columbidae).

Previous molecular phylogenetic analyses and new analyses reported here demonstrate that the genus Claravis is not monophyletic and comprises two lineages, one with the species pretiosa Ferrari-Pérez, 1886 (Blue Ground-dove), and the other with two species: geoffroyi Temminck, 1811 (Purple-winged Ground-dove) and mondetoura Bonaparte, 1856 (Maroon-chested Ground-dove). Because the generic name Claravis is typified by C. pretiosa (Ferrari-Pérez, 1886), a new genus, Paraclaravis gen. nov., is described for geoffroyi Temminck, 1811 and mondetoura Bonaparte, 1856.