Evolutionary History of Passerine Birds (Aves: Passeriformes)
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J Ornithol (2015) 156 (Suppl 1):S355–S365 DOI 10.1007/s10336-015-1185-6 REVIEW Evolutionary history of passerine birds (Aves: Passeriformes) from the Qinghai–Tibetan plateau: from a pre-Quarternary perspective to an integrative biodiversity assessment 1 2 3 4 Martin Pa¨ckert • Jochen Martens • Yue-Hua Sun • Dieter Thomas Tietze Received: 31 October 2014 / Revised: 27 January 2015 / Accepted: 23 February 2015 / Published online: 12 March 2015 Ó Dt. Ornithologen-Gesellschaft e.V. 2015 Abstract As one of the most prominent topographical plateau habitats several times independently. Second, we features on Earth, the Qinghai–Tibetan plateau (QTP) un- discuss younger speciation events corresponding to phy- derwent a long and complex history of the QTP uplift from logeographic east–west divides along the southern QTP the collision of the Indian and the Eurasian plates to the margin. A multidisciplinary approach combining genetic, present. At its southern and southeastern margins, it is bioacoustic and morphological markers shed new light on flanked by the most significant hotspots of organismic di- the phylogenetic relationships of Pnoepyga wren babblers versity of the northern hemisphere (including birds), the and on the intraspecific subdivision of the Buff-barred Sino-Himalayan mountain forests. In contrast, the central Warbler (Phylloscopus pulcher). plateau region itself harbours species-poor communities but also a good number of endemics that presumably Keywords Qinghai–Tibetan plateau Á Passeriformes Á evolved from rather ancient (pre-Pleistocene) phylogenetic Sino-Himalayas Á Alpine habitats Á Carpodacus lineage splits. We discuss the evolutionary history of QTP passerines from a twofold perspective including examples from our own research. First, we provide an overview of Introduction those alpine QTP endemics that represent late Miocene and Pliocene lineage splits, i.e. early colonisations to the cen- The Qinghai–Tibetan plateau (QTP) is one of the most tral alpine QTP region. As an example, true rosefinches prominent topographical features on Earth. The long and (genus Carpodacus) presumably evolved from a forested complex history of the QTP uplift from the collision of the eastern QTP centre of origin and colonised the (semi-)open Indian and the Eurasian plates to the present has been much debated but is still not fully understood to date (reviews: Mulch and Chamberlain 2006; Favre et al. 2014). How- Communicated by E. Matthysen. ever, there is a growing understanding that the discussion of biogeographic history and organismic evolution (faunal & Martin Pa¨ckert [email protected] turnovers, colonisation events, continental radiations) of the QTP region cannot be limited to the central plateau 1 Senckenberg Natural History Collections, Museum of region alone, but is firmly associated with the orogenetic Zoology, Ko¨nigsbru¨cker Landstraße 159, 01109 Dresden, history of the flanking mountain chains: Tian Shan in the Germany 2 west, Himalayas in the south, Hengduan Shan in the Institut fu¨r Zoologie, Johannes Gutenberg-Universita¨t, southeast and Altai and Qilian Shan in the north (review in 55099 Mainz, Germany 3 Favre et al. 2014). At its southern and southeastern fringes, Key Laboratory of Animal Ecology and Conservation, the QTP is flanked by the most significant hotspots of or- Institute of Zoology, Chinese Academy of Science, Beijing 100101, China ganismic and also avian diversity of the northern hemi- sphere, namely the Sino-Himalayan mountain forests 4 Institute of Pharmacy and Molecular Biotechnology, Heidelberg University, Im Neuenheimer Feld 364, (Myers et al. 2000; Wikramanayake et al. 2001; Roselaar 69120 Heidelberg, Germany et al. 2007). The remarkably high species richness of these 123 S356 J Ornithol (2015) 156 (Suppl 1):S355–S365 mountain forests strongly contrasts with low species di- recolonisation of the core plateau region from versity on the central QTP itself. However, some of the (north-)eastern glacial refugia (Qu et al. 2010; Lei et al. cold-adapted high alpine endemics of the plateau itself 2014). But going back in time and evolutionary history, have apparently evolved from rather ancient lineage splits diversification and speciation processes among eight and underwent long-term adaptation to extreme conditions species of snowfinches and among four Holarctic lin- (Qu et al. 2013). In fact, there is evidence from the fossil eages of the Horned Lark were suggested to date back to record and from reconstructed biogeographic histories that even late Pliocene times (in fact with quite similar split the core QTP region was an evolutionary centre of origin ages estimates; Table 1;compareLeietal.2014; (Deng et al. 2011; Tseng et al. 2013; Wang et al. 2014)— Alstro¨metal.2013a). Possible evolutionary scenarios emphatically termed a ‘cradle of evolution’ of cold-adapted and taxonomic consequences are still under lively debate mammals and forest-dwelling ground beetles (Erithra and particularly for Horned Larks because of a low signal of Pterostichus; Schmidt et al. 2012). nuclear markers and fundamental differences among In contrast, the discussion on the evolutionary history, split age estimates inferred from independent studies times and areas of origin of Tibetan endemic bird species (Alstro¨metal.2013a; Drovetski et al. 2014). To date, a has long been limited to young speciation processes centre of origin and diversification has not been hypo- during the final QTP uplift phase, in the course of thesised for either of these two groups, but at least for Quarternary climate change and glacial impact. But be- snowfinches a QTP ‘cradle of evolution’ is a quite likely cause comparative phylogeography is dealing with past hypothetical scenario that can be put to test in future demographic processes within species on a rather short studies (see below). evolutionary time scale—and in the case of previously With this paper, we aim at first providing a perspective studied QTP species only on a rather narrow spatial to the older pre-Pleistocene events and early lineage splits scale, too—this method only allows for reconstruc- within passerine genera and early colonisation events to the tions of the terminal events of colonisations and radia- central QTP from its fringes or from adjacent zoogeo- tions during the biogeographic history of a group of graphic regions. Second, we discuss the younger speciation organisms (Qu et al. 2009, 2010). For example, con- events with a focus on the frequently observed phylogeo- gruent phylogeographic scenarios for snowfinch species graphic east–west divides along the southern QTP margin of genera Montifringilla, Onychostruthus and Pyrgilau- under a multidisciplinary perspective from integrative da (Passeridae) as well as for the Horned Lark (Ere- taxonomy. For both topics, we present some exemplary mophila alpestris, Alaudidae) suggested a post-glacial results from our own research. Table 1 Split age estimates for Family Species/lineage Age References alpine QTP endemic species and species groups (genetic Paridae Pseudopodoces humilisa 7.7–9.9 Qu et al. (2013) lineages; split age from closest Fringillidae Carpodacus roborowskiia *9.0 Tietze et al. (2013) relative in Ma) Fringillidae Carpodacus pulcherrimus/C. waltoni cladea *7.0 Tietze et al. (2013) Aegithalidae Leptopoecile (genus) *9.0 Pa¨ckert et al. (2010, 2013) Aegithalidae Leptopoecile sophiae *4.0 Pa¨ckert et al. (2010, 2013) Phylloscopidae Phylloscpus fuscatus/P. fuligiventer clade *7.5 Pa¨ckert et al. (2012) Phylloscopidae Phylloscopus affinis/P. occisinensisa *4.0 Pa¨ckert et al. (2012) Turdidae Turdus mupinus *4.5 Nylander et al. (2008) Turdidae Turdus kessleria *1.0 Nylander et al. (2008) Muscicapidae Phoenicurus ochruros QTP clade 4.08 Qu et al. (2010) Passeridae Montifringilla/Pyrgilauda cladeb 3.8 Lei et al. (2014) Alaudidae Melanocorypha genus *6.5 Alstro¨m et al. (2013a) Alaudidae Melanocorypha maximaa *3 Alstro¨m et al. (2013a) Alaudidae Alauda gulgula *5 Alstro¨m et al. (2013a) Alaudidae Eremophila genus *9 Alstro¨m et al. (2013a) Alaudidae Eremophila alpestris (basal split) *3.5 Alstro¨m et al. (2013a) Alaudidae Eremophila alpestris QTP clade *2.5 Alstro¨m et al. (2013a) a Alpine endemic species according to Vaurie (1972) p. 139 b Except M. nivalis, all species of this clade are endemic to the QTP 123 J Ornithol (2015) 156 (Suppl 1):S355–S365 S357 Fig. 1 Major periods of faunal interchange between mountain forests of the southern/southeastern QTP margins and adjacent bioregions, colour codes of key forest areas for SE Asian birds, according to Collar et al. (2001); modified from Pa¨ckert et al. (2012) (color figure online) Forests of the southern QTP margins: hotspot relevant trigger of accelerated avian diversification at the of biodiversity southern and southeastern QTP margins during the mid- to late Miocene and first colonisation events to the subtropical Four major biodiversity hotspots at the western, southern forests of the Himalayas and the Hengduan Shan from a and eastern QTP margins have been listed by Favre et al. Southeast Asian/Indoburmese centre of origin were sug- (2014: fig. 1), all of them encompassing mountain forest gested to have occurred during that period (Pa¨ckert et al. ecosystems of (1) Central Asia, (2) the Himalayas, (3) 2012; Fig. 1). This early stage of passerine diversification Indo-Burma and (4) the Hengduan Shan. Species richness in SE Asia is backed by the recent finding that the mean patterns of Palearctic passerine birds almost perfectly re- split age of sister species pairs of the Eastern Himalayas flect these four centres (Roselaar et al. 2007; see review on was estimated as ancient as 7.1 Ma (Price et al. 2014)—the passerine diversity of the Sino-Himalayas in Martens et al. time when earliest lineage splits among subtropical sister 2011). In the south, where Sino-Himalayan mountain for- species are thought to have occurred (see the example of ests constitute the greatest biodiversity hotspot of the re- treecreepers, Certhia, in Fig. 2). Recent ecological com- gion, avian species richness increases along a gradient from munity analyses of Himalayan passerines have shed some the drier Western Himalayas towards highest species light on adaptive processes and niche evolution.