The larger, family-level phylogenetic analyses indicated that both WG lineages diverged early within their respective clades (Fig. 1). Each of these two lineages is a distinct clade not closely related to the genus historically or currently grouped within. Additionally, both were found to be discrete clades deeply divergent from their closest relatives. The WG Laughingthrushes did not group with any of the other traditional laughingthrush clades (including ones designated as Trochalopteron or Strophocincla), but instead were sister to a clade composed of Heterophasia, Minla, Actinodura, Leiothrix, Liocichla and Crocias (Fig. 1a). Similarly, we reconstructed the WG Shortwings as sister to a newly uncovered clade comprised of several genera of mainly Asian blue flycatchers, the Niltavinae [28, 35], and not closely related to traditional shortwings in the genera Myiomela or Brachypteryx (Fig. 1b).
Both WG lineages are not only divergent from their closest relatives, they are also not sister to any single genus as defined under current taxonomic treatments. Thus, they cannot easily be lumped or subsumed under an existing clade without an extensive reorganization of multiple genera. As a better alternative, we designated new genus names: Western Ghats Laughingthrushes as Montecincla and Western Ghats Shortwings as Sholicola (see below for descriptions), to recognize their distinctiveness and to avoid confusion with older names associated with other taxa that are not part of these new groups.
The ancestral area reconstruction showed dispersal into the WG for Montecincla as most likely from the Himalayas and for Sholicola from the Himalayas + Southeast Asia (Additional file 1: Figures. 1, S2, S3). The two WG lineages have a similar estimated timing of separation from their closest relatives (Table 1). These dates also coincide with climatic events that led to the drying of peninsular India (Table 1), perhaps leading to discontinuing gene flow and a differentiation between peninsular forms from Himalayan / Southeast Asian forms. Furthermore, the divergence of populations within both lineages across the sky islands exhibits similar timing of diversification in response to the same biogeographic barriers (see below).
Phylogenetic analysis of both groups showed clades corresponding to four major biogeographic regions in the WG, north to south: A- Bababudan & Banasura hills; B- Nilgiri hills; C- Anamalai, Palani, and Meghamalai hills; D- Ashambu hills (Fig. 2). Montecincla was comprised of four reciprocally monophyletic clades (Fig. 2b). Bayesian species delimitation using BPP also showed that these four clades were distinct species. Elevating former subspecies names, these are: Montecincla jerdoni (range A), M. cachinnans (B), M. fairbanki (C), M. meridionalis (D). Similarly, analysis of Sholicola revealed three reciprocally monophyletic clades (Fig. 2c) that were distinct species according to BPP. While two existing subspecies names that match these units can be elevated, namely Sholicola major (range A, B) and S. albiventris (C), the third (D) is a new species we designate below. Analyses of fixed characters of plumage, morphometrics and song data (below) match the distinct units uncovered in the phylogenetic and species delimitation analyses. Therefore multiple lines of data support four species in the Montecincla complex and three species in the Sholicola complex.
Our analysis of plumage variation for Montecincla supports four distinct groups (Table 2). All four species have distinguishing features in multiple color patches that separate these evolutionary units. Within-species variation is low and populations with similar features occur in geographically clustered sky islands (shown in Fig. 2). In Sholicola, S. major (A, B) showed striking differences from the other two species south of the Palghat Gap. The differences in plumage of between S. albiventris (C) and Sholicola sp. nov. (D; see below) are mainly in terms of the extent of white coloration on the belly. The newly described species has a considerably larger white belly patch (~24mm in length) than S. albiventris (~19mm) (Table 3).
The four species of Montecincla and the three species of Sholicola differed significantly from each other in morphospace occupied (Fig. 3). This differentiation within both Montecincla and Sholicola was largely driven by tarsus length, although the direction of increase was reversed – Montecincla in the southernmost region of Ashambu Hills (D) is larger than the other species further north, while in Sholicola the species in the southern hills were the smallest (Additional file 1: Tables S3, S4).
Spectrograms reveal the presence of unique song types in all four species of Montecincla and all three Sholicola species (Fig. 4). All four species of Montecincla show distinct features in song. Montecincla fairbanki sings at a higher bandwidth and song rate than the other populations, whereas M. meridionalis sings at a lower bandwidth than other species, while M. jerdoni has a higher song complexity with longer phrases than the other populations (Additional file 1: Table S5). Songs of all Sholicola species are also quantitatively distinct, with differences in song length and frequency across different species (more details in Purushotham and Robin ). The song of the newly described species of Sholicola is shorter in length and higher in frequency from S. albiventris (Additional file 1: Table S6).
Naming new taxa
As outlined above, the results of our analysis warrant new names for two genera and one species, in addition to elevating six subspecies to species level.
Western Ghats laughingthrushes
The laughingthrushes of the Western Ghats have been treated as part of the genus Garrulax Lesson, 1831 by Ali and Ripley  and as Strophocincla Wolters, 1981 by Rasmussen and Anderton , which was followed by others [22, 26]. However, in most global avian checklists, they are currently placed in Trochalopteron [8, 21] following phylogenetic insights . Genus-level treatments in laughingthrushes have clearly been very unstable given extensive morphological variation and labile trait evolution in these birds. Our phylogenetic results reveal that all previous genus designations are inappropriate for Western Ghats Laughingthrushes and here we erect a new genus to recognize their distinctiveness.
Furthermore, we propose elevation of the four subspecies to full species level. This fits assertions by others: Rasmussen and Anderton  tentatively delimited two species (one north and the other south of the Palghat Gap with two subspecies each) but suggested that the four allopatric populations may each warrant species-level status with more evidence. Additionally, Praveen and Nameer  conducted an analysis of plumage differences and suggested that this complex was best represented as four distinct species. In line with our results of multiple character systems, we hereby propose elevation of these subspecies as four full allopatric species (Additional file 1: Figure S4). This is essentially a revival of the taxonomic treatment at the time of the original descriptions of these taxa.
Montecincla genus novum
Order Passeriformes: Family Leiothrichidae
Suggested common name: Chilappan
Type species: C. [=Crateropus] cachinnaus [sic] Jerdon, 1839 = Strophocincla cachinnans = Montecincla cachinnans comb. nov.
Additional species included: Garrulax (?) Jerdoni [sic] Blyth, 1851 = Strophocincla cachinnans jerdoni = Montecincla jerdoni comb. nov.; Trochalopteron Fairbanki Blanford, 1869 = Strophocincla fairbanki = Montecincla fairbanki comb. nov.; and Trochalopterum meridionale Blanford, 1880 = Strophocincla fairbanki meridionalis = Montecincla meridionalis comb. nov.
ZooBank Registry: urn:lsid:zoobank.org:act:88B21D13-2639-4BFD-8729-74308F609E4E
Diagnosis: Montecincla gen. nov. can be differentiated from other genera of traditional laughingthrushes - Garrulax; Ianthocincla Gould, 1835; Trochalopteron; and from Turdoides babblers by the combination of a prominent white supercilium, olive brown upperparts, small size, brown wings, rufous flanks and dark bill. Montecincla lacks the complex upperwing colouration (consisting of strong barring or conspicuous wing panels) of Actinodura Gould, 1836, and of Liocichla Swinhoe, 1877. Montecincla also lacks the slender greyish body and long tail of sibias of the genus Heterophasia Blyth, 1842. Laniellus Swainson, 1832 (=Crocias Temminck, 1836) has a diagnostic whitish underside with bold, streaked flanks that are absent in Montecincla. Minla Hodgson, 1837 is a genus of small birds (14cm) with a colourful reddish tail or rump, while Montecincla is larger (20cm) with a dull olive-brown tail and rump. Leiothrix Swainson, 1832, is characterized by a colourful bill and a yellow throat, contrasting with the black bill and greyish-white or black throats of Montecincla.
Description: Montecincla chilappans are medium-sized (~20cm) songbirds with rounded tails. Their upperparts are olive grey and they have a prominent white supercilium. The breast is greyish-white in three species and rufous in one (Table 2).
Etymology: Montecincla, of feminine gender, is a combination of Latin “mons” (gen. “montis”), with the meaning “mountain”, and Greek “kinklos”, denoting an unidentified type of songbird (often assumed to be a thrush). We chose this appropriate moniker because the genus is confined to the higher mountains of the Western Ghats. Our suggested vernacular name “Chilappan” stems from the local name of this genus (in Malayalam language), denoting their joyful cackling calls.
Comments: We suggest Banasura Chilappan as a common name for Montecincla jerdoni in recognition of Mt. Banasura, one of its strongholds and its type locality. Similarly, we suggest the common names Nilgiri Chilappan for Montecincla cachinnans, Palani Chilappan for Montecincla fairbanki, and Ashambu Chilappan for Montecincla meridionalis.
Western Ghats blue robins or shortwings
The species were historically placed in the genus Phoenicura Swainson, 1831, later in Callene Blyth, 1847, then in Brachypteryx Horsfield, 1821 and more recently in Myiomela. None of these generic names are appropriate given our phylogenetic analysis. We thus erect a new genus to recognize the distinctness of Western Ghats Blue Robins (also known as Western Ghats Shortwings). In addition, we recognize three distinct species in this genus, including one new, previously unrecognized species.
Sholicola genus novum
Order Passeriformes: family Muscicapidae
Suggested common name: Sholakili
Type species: Phoenicura major Jerdon, 1841 = Myiomela major = Sholicola major comb. nov.
Additional species included: Callene albiventris Blanford, 1868 = Myiomela albiventris = Sholicola albiventris comb. nov.; Sholicola ashambuensis Robin, Vishnudas, Rheindt, Gupta, Hooper, Ramakrishnan & Reddy (see below).
ZooBank Registry: urn:lsid:zoobank.org:act:66837381-635E-476C-9454-092BB03B0A91
Diagnosis: Sholicola is sexually monomorphic, thus differing from the following closely related and sexually dimorphic genera: Cyornis Blyth, 1843; Eumyias Cabanis, 1850; Niltava Hodgson, 1837; and Cyanoptila Blyth, 1847. Being largely blue, it differs from Anthipes Blyth, 1847, which comprises brown birds with distinctive white throats. Members of Sholicola are resident birds with no known seasonal migration as in members of Niltava and Cyanoptila. The three species of Sholicola are phylogenetically distant from the genera they were previously placed under (Brachypteryx, Callene, Phoenicura, Myiomela). They also differ from these genera in having a sexually monomorphic plumage dominated by blue colouration.
Description: Sholicola is a genus of terrestrial blue flycatchers confined to high altitude (above 1200m) forests in the Shola-grassland complexes of the Western Ghats. Their general plumage is dominated by blue. They have a bluish-white band above the black lores, a slightly curved bill tip, well-developed rictal bristles, a short, nearly square tail, and long tarsi. The birds are mainly restricted to the understory, rarely venturing above three meters and feeding on insects singly or in pairs. There is no known sexual dimorphism in plumage though morphometric differences in wing length are recognized .
Etymology: Sholicola, of masculine gender, is a combination of Shola (the local name for montane forests in the Western Ghats) and the suffix –cola (from the Latin verb “colere”), meaning “dweller". We also suggest the common English name Sholakili, where -kili is the local name for “bird”.
Description of a new species in the genus Sholicola:
Sholicola ashambuensis, species nova
English name: Ashambu Sholakili
Holotype: Trivandrum Museum of Natural History (TMNH) No. 725, collected by H.S. Fergusson on 3 May 1903 in the Chemunji Hills, Travencure (Travancore).
Etymology: The adjective ashambuensis denotes the species’ geographical locality, the Ashambu Hills of southern India.
ZooBank Registry: urn:lsid:zoobank.org:act:CB9F32BC-750B-48AA-8F87-C9727144019D
Diagnosis: The following characters can be used to diagnose Sholicola ashambuensis.
Despite being smaller in general body size, S. ashambuensis has a considerably larger white belly patch (~24mm in length) than S. albiventris (~19mm). Only one S. ashambuensis specimen was available for measurements.
S. ashambuensis is a smaller bird (Additional file 1: Figure S5) with a shorter tarsus and longer bill (Additional file 1: Table S7; Additional file 1: Figure S6) than S. albiventris based on an examination of 76 S. albiventris and 21 S. ashambuensis.
S. ashambuensis has a distinct song (Additional file 1: Figure S7) with a higher mean maximum frequency, but narrower song bandwidth, shorter notes and shorter song bouts than S. albiventris, based on 119 S. ashambuensis songs and 203 S. albiventris songs.
When compared side-by-side, S. ashambuensis is paler blue than S. albiventris.
In addition to these phenotypic diagnostic characters, S. ashambuensis forms a reciprocally monophyletic clade based on DNA data from about 25 individuals. Further population genetic data with 15 microsatellites and 218 individuals (including 17 of S. ashambuensis) also support significant genetic differentiation .
Description of holotype: A small, overall dark-blue flycatcher with a large white belly patch reaching to the vent, black lores, and a bluish-white band above the lores. The holotype measures: bill – 16mm, wing– 81mm, tail – 61mm, tarsus – 21mm.
Distribution: Ashambu hills south of Shenkottah gap, southern India, mostly above 1200m elevation (Additional file 1: Figure S4).
Comments: The subtle plumage differentiation and limited fieldwork in the Ashambu Hills, including an absence of systematic capture-based studies, are perhaps the reason why S. ashambuensis has gone taxonomically unrecognized until now. The only museum specimen to our knowledge, the holotype (described above), was re-discovered (by CKV) after being locked up for about 120 years in a drawer of the Travancore Museum. Other specimens thought to be from the southern range are missing (and perhaps destroyed). There are possibly two more specimens in NHMUK (Tring) based on geography, but these were not examined by the authors.