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The Golden-spectacled Warbler Seicercus burkii - a species swarm (Aves: Passeriformes: Sylviidae), Part I

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... Determination of species limits typically becomes problematic in the case of cryptic species where species have a highly similar phenotype and with widely overlapping breeding ranges. Several of the " new " leaf-warblers (Phylloscopidae sensu Alström et al. 2006) are examples of cryptic speciation (Irwin et al. 2001aIrwin et al. , 2001bIrwin et al. , 2005), and integrative studies of morphology, song, and genetics have resulted in a number of taxonomic recommendations: i) upgrading of former subspecies to full species (Olsson et al. 2004Olsson et al. , 2005 Päckert et al. 2009), ii) descriptions of species new to science (Alström & Olsson, 1999; Alström et al. 2010; Martens et al. 1999 Martens et al. , 2008, and iii) revalidation of historical taxon names not in use (Martens et al. 2004Martens et al. , Päckert et al. 2014). Splitting one polytypic species into several clear-cut and diagnosable species might require a re-evaluation of species distributions because distributional data is lacking for many of these newly-erected species. ...
... Golden-spectacled warblers are a perfect example. It has been known since 1999 that these birds do not represent a single species with a continuous distribution from the Himalayas across large parts of China and Southeast Asia (Alström & Olsson 1999, Martens et al. 1999). Instead, these birds embrace what we now know to be six well defined biological species: two sister species pairs resulting from vicariant events are present across a Sino-Himalayan East-West disjunction at different elevational levels (S. whistleri and S. valentini at high elevations, S. burkii and S. tephrocephalus at low elevations). ...
... Morphology: All specimens backing our occurrence data were examined and determined to species level according to diagnostic phenotypic characters detailed in earlier studies: coloration and shape of the median and lateral crownstripes, shape of white spots on tail feathers, etc. (Fig. 1; further drawings in Martens et al. 1999, Alström & Olsson 1999, Alström 2006 photographs in Martens et al. 2003). Many specimens had already been examined by the late Siegfried Eck (Martens et al. 1999Martens et al. , 2003). We relied on identifications in his original notes. ...
... Two papers on the systematics of the Golden-spectacled Warbler Seicercus burkii complex were published almost simultaneously in autumn 1999 (Alstrom & Olsson 1999, Martens et al. 1999. Both propose that what was previously considered to be one species, S. burkii, is in fact a complex of sibling species, and both describe a new species from central China! ...
... This is the only taxon for which there appears to be complete agreement between the two papers. . Tree comparing molecular data sets between taxa as named in Martens et a/. (1999) and in Alstrom and Olsson (1999). Our DNA samples (unpubl. data) are in all but one case from the same localities as those of Martens et a/., i.e. Emei Shan (soror, tephrocephalus 'group 6', valentinb and Nepal (nemoralis); however, tephrocephalus 'group 4' is from northern Vietnam, as we lack material from this taxon from Taibai Shan. ...
... S. whist& Ticehurst, 1925 (1% Ith subspecies whzstlen Ticehurst, 1925 andnewfordis KO&, 1954) S. tephrocephaZus (Anderson, 1871) S. omeiensis Martens, Eck, Packert and Sun, 1999 S. valentini (Nartert, 1907) (with subspecies valentrni (Hartert, 1907) and latouchet (Bdngs, 1929)) s. soyor lilstrom and Olsson 1999 ...
... The number of species accepted has increased steadily, mainly by the splitting of long-established polytypic species. Papers such as those of Helbig et al. (1996), Irwin et al. (2001a), Johansson et al. (2007), Martens et al. (1999, Olsson et al. (2004Olsson et al. ( , 2005, Päckert et al. (2004Päckert et al. ( , 2009a, and Richman & Price (1992), to mention just a few, are milestones in this respect. The various authors treating Phylloscopus and Seicercus in del Hoyo et al. (2006), on the other hand, made only partial use of these results and upheld a quite conservative view. ...
... Because Phylloscopus is morphologically so extremely homogeneous yet at the same time so astonishingly species-rich, it has been the focus of evolutionary studies attempting to better understand ecological and evolutionary traits in the leaf warbler assemblage. Sample themes relate to the high number of sympatric breeding species (Alexander 1950, Martens & Eck 1995, Price & Jamdar 1991b; the evolution of breeding distributions (Price et al. 1997), with special emphasis on the Himalayan avifauna (Johansson et al. 2007); molecular genetic aspects of evolution and ecological differentiation (Price 2010, Richman & Price 1992; species swarms (Alström & Olsson 1999, Martens et al. 1999; and cryptic species (Olsson et al. 2005), together resulting in a revolution, especially of Asian leaf warbler systematics (Rheindt 2006). In addition, the association of habitat and song characteristics has been investigated (Badyaev & Leaf 1997) as have links between vocalisations and phylogeny (Martens 1980), radiation versus molecular genetics and bioacoustics (Päckert et al. 2004), phylogeography and the evolutionary timescale of Phylloscopus radiations in the Sino-Himalayan region (Päckert et al. 2009b), determinants of northern and southern range limits (Gross & Price 2000), morphology and ecology along an altitudinal gradient (Price 1981), evolution of ecological differences in breeding and non-breeding seasons (Price & Gross 2005), correlates of wing morphology with foraging behaviour and migration distance (Marchetti et al. 1995), phylogeny versus ecological diversification and community Martens. ...
... With one exception (S. poliogenys), they have vivid green upperparts and bright yellow underparts, and a yellowish (in rare cases whitish) eye ring occurs throughout. Most taxa were disentangled only recently, from vocalisation and molecular genetic data (Alström & Olsson 1999, 2000, Martens et al. 1999, Olsson et al. 2004, Päckert et al. 2004. The currently recognised eight species of the complex are distributed all along the Himalayan chain including SE Tibet, to large stretches of mountainous country from the eastern rim of the Tibetan plateau all across the southern half of China and large parts of eastern Indochina. ...
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Recent and proposed taxonomic changes to the warbler genera Phylloscopus and Seicercus are reviewed within the geographic scope of this series. Their reliability relative to secondary contacts, hybridization, introgression, bioacoustics and molecular genetics is discussed in terms of current species concepts, mainly the Biological Species Concept, and certain previously unpublished data are added in support of taxonomic decisions made. Within the area under consideration, this review records an increase in the number of Phylloscopus species from the 33 in Mayr et al. (1986) and 44 in del Hoyo et al. (2006) to 50. In Seicercus, S. burkii, a previously merged taxon swarm, has been split into six partly allopatric, partly sympatric species. This enlarging of species number emerges from due application of the results of bioacoustic and molecular genetic analyses to passerine systematics. Homogeneous external morphology largely hid the phylogenetic complexity and richness of these genera, but their species totals are not yet fixed; several populations presently accepted at species level need further substantiation.
... Our example (Seicercus) is a case typical of cryptic species, in that the decisive morphological characters only "take shape" when considered in the light of acoustic and molecular data, and hence demand especially careful data acquisition. Here we report on the Seicercus burkii complex (Sylviidae: standing close to, if not within Phylloscopus; see MARTENS et al. 1999: 283) of the Himalayas, China and countries bordering them to the south. Using many methods this group has been shown to be a species swarm. ...
... Despite certain differences the analyses of the "burkii diversity" by the Swedish group (ALSTRÖM & OLSON 1999) and the German group (MARTENS et al.1999) are extremely similar. MARTENS et al. (1999) were more reluctant to combine vicariant taxa into polytypic species. They named a total of eight taxa, the combination of which to form biospecies was not clearly discernible in all cases. ...
... ALSTRÖM & OLSSON (1999) also describe these populations ("group 4", "group 5") as distinctly short-winged (maximum 57 mm). MARTENS et al. (1999) ANDERSON (1871ANDERSON ( , 1878, was found to have an omeiensis haplotype (cyt b). Hence the northern (omeiensis; W China) and southern populations (tephrocephalus sensu ANDER-SON, non sensu ALSTRÖM & OLSSON; Burma), which differ only slightly in colour, should be united in a single species. ...
... Furthermore, we used DNA barcoding as a control for correct identification of cryptic species in the field with a broader choice of target species from our total sampling (143 samples from 45 species). Due to their subtle phenotypic distinctiveness, cryptic species are not easily distinguished from their closest relatives without additional data from vocalisations or genetics (Irwin et al. 2001, Bickford et al. 2007, such as in the case of golden-spectacled warblers of the former Seicercus burkii complex (Alström and Olsson 1999, Martens et al. 1999, Päckert et al. 2004, Olsson et al. 2005, today Phylloscopus burkii and allies (del Hoyo and Collar 2016). For the eight species-level taxa of this group, DNA barcode reference sequences were recently published . ...
Article
The Hkakabo Razi region located in northern Myanmar is an Important Bird Area and part of the Eastern Himalayan Biodiversity Hotspot. Within the framework of the World Heritage Convention to enlist the site under criterion (ix) and (x), we conducted a biodiversity assessment for passerine birds using DNA barcoding and other molecular markers. Of the 441 bird species recorded, we chose 16 target species for a comparative phylogeographic study. Genetic analysis was performed for a larger number of species and helped identifying misidentified species. We found phylogeographic structure in all but one of the 16 study species. In 13 species, populations from northern Myanmar were genetically distinctive and local mitochondrial lineages differed from those found in adjacent regions by 3.9–9.9% uncorrected genetic distances (cytochrome- b ). Since the genetic distinctiveness of study populations will be corroborated by further differences in morphology and song as in other South-East Asian passerines, many of them will be candidates for taxonomic splits, or in case an older taxon name is not available, for the scientific description of new taxa. Considering the short time frame of our study we predict that a great part of undetected faunal diversity in the Hkakabo Razi region will be discovered.
... Clade M1b is strongly supported by concatenation but not in the *BEAST analysis. We consider this highly plausible because of the generally close similarities between S. valentini and S. whistleri in morphology, song and breeding habitat/altitude (Alström and Olsson 1999;Alström and Olsson, 2000;Martens et al., 1999;Olsson et al., 2004;Päckert et al., 2004). ...
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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 againstproliferation 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.
... That is, among the 10 taxa treated in this set of studies, differentiation ranged from essentially nil (e.g., Spizixos semitorques) to deep (e.g., Pomatorhinus spp.). The relative frequency of deep mitochondrial differentiation within and among continuously distributed populations (e.g., Cyanoderma ruficeps) likely reflects climatedriven, geographic processes that perhaps characterized the Pleistocene in the region (see, e.g., Alström and Olsson, 1999;Martens et al., 1999). ...
... Many species level taxonomic changes have also been affected recently, due to either lumping or splitting of existing species. These include splitting of Phoenicopterus ruber into P. roseus and P. ruber (Knox et al., 2002); Corvus macrorhynchos into C. levaillantii and C. macrorhynchos (Sibley & Monroe, 1990); Hirundo tahitica into H. domicola and H. tahitica (Sibley & Monroe, 1990); Mirafra assamica into M. affinis, M. assamica, M. erythrocephala and M. microptera (Alström, 1998); Chloropsis cochinchinensis into C. jerdoni, C. cochinchinensis, and C. kinabaluensis (Wells et al., 2003); Sula dactylatra into S. dactylatra and S. granti (AOU, 2000); Spilornis minimus into S. klossi and S. cheela (Rasmussen & Anderton (2005); Spizaetus cirrhatus into S. cirrhatus and S. floris (Gjershaug et al., 2004); Otus scops into O. senegalensis (Dowsett & Forbes-Watson, 1993), O. sunia (AOU, 1998), O. alius (Rassmussen, 1998) and O. scops; Acrocephalus stentoreus into A. stentoreus and A. orinus (Bensch & Pearson, 2002); Seicercus burkii into S. burkii, S. valentini, S. whistleri, S. soror and S tephrocephalus (Alström & Olsson, 1999) and S. omeiensis (Martens et al., 1999); Lonchura malacca into L. malacca and L. atricapilla (AOU, 2000). Further, Pomatorhinus horsfieldii has been split into P. melanurus and P. horsfieldii (Collar, 2006;Rasmussen & Anderton, 2005), of which P. melanurus is listed as endemic to Sri Lanka. ...
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The scientific life of Prof. Dr Jochen Martens (Germany: Mainz) is illuminated on occasion of his 80th birthday. Facts and impressions are given as well as lists of his publications (329), taxa he has described (2 families, 29 genera, 296 species) and that have been dedicated to him (11 genera, 219 species, 1 subspecies). Jochen Martens is a renowned specialist for birds (Aves) and for harvestmen (Opiliones). So far, he travelled to 27 countries in 80 journeys with Nepal and the Himalayas as one geographic focus.
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Molecular phylogeography is an important sub-discipline of biogeography, which utilizes the molecular biological techniques to reconstruct the phylogenies of organisms at intra-species and inter-species levels, thus trace the evolutionary history and understand the historical nature of the biota. The neutral and coalescent theories provide theoretical framework for the phylogeographic studies. Mitochondrial DNA and microsatellites are most broadly used genetic markers. Recent advance has been made for birds in Europe and North America. These studies have revealed that that the various spatial distribution patterns and the centers of origin in the species-specific level, and identified different phylogeographic patterns either as the result of vicariance or dispersal. Moreover, differentiation on the migration behaviors between avian species has been recognized as the considerable factor in influencing the phylogeographic patterns. Compared with these two regions, avian phylogeography in China is newly started and more comparative phylogeographic studies are required for different geographical regions and species. Meantime, the usage of next-generation sequencing methods will potentially accelerate our understanding of avian phylogeography in the near future.
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