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Integrative taxonomy of the Plain-backed Thrush (Zoothera mollissima) complex (Aves, Turdidae) reveals cryptic species, including a new species
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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.
We revise the Z. mollissima-Z. dixoni complex by integrating morphological, acoustic, genetic (two mitochondrial and two nuclear markers), ecological and distributional datasets.
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.
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.
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Integrative taxonomy of the Plain-backed Thrush (Zoothera mollissima) complex (Aves, Turdidae) reveals cryptic species, including a new species
),
Abstract
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.
We revise the Z. mollissima-Z. dixoni complex by integrating morphological, acoustic, genetic (two mitochondrial and two nuclear markers), ecological and distributional datasets.
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.
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.
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Acknowledgements
We gratefully acknowledge the staff of the following museums for allowing access to the collections and in many cases loan of specimens under their care and/or collected by them: Paul Sweet, AMNH; Robert Prŷs-Jones and Mark P. Adams, NHMUK; Moe Flannery, CALAS; John Bates, David Willard, and Ben Marks, FMNH; He Peng, IOZ; Yang Xiaojun and Liu Luming, KIZ; Jeremiah Trimble, MCZ; Jérôme Fuchs and Patrick Bousses, MNHN; Laura Abraczinskas, MSUM; Helen James and Gary Graves, NMNH; Steven van der Mije, NNM; Claudia Kamcke and Bettina Borges-Naumer, SNMB; Diarmaid Ó Foighil and Janet Hinshaw, UMMZ; and Sylke Frahnert and Jurgen Fiebig, ZMB. In addition, DNA samples, including toepad samples, were provided by Joel Cracraft, Thomas Trombone and Paul Sweet, AMNH; Robert Prŷs-Jones and Mark P. Adams, NHMUK; Chen Jun and He Peng, IOZ; Yang Xiaojun, KIZ; Jérôme Fuchs, MNHN; and Diarmaid Ó Foighil and Janet Hinshaw, UMMZ. We are most grateful to Craig Brelsford, Liang Chen, James Eaton, Sid Francis, Paul Holt, Xiaonong Yang and Lei Zhou for providing sound recordings; and to Björn Anderson, Steve Bale, Garima Bhatia, Craig Brelsford, Vijay Cavale, George Chapman, Li Cheng, Ranjan Kumar Das, Augusto Faustino, John and Jemi Holmes, Yann Muzika, Sachin Rai, Pranjal Saikia, Takema Saitoh, Subrato Sanyal, Adesh Shivkar, Nitin Srinivasmurthy, Rajneesh Suvarna, Ulrich Weber and Huaming Zhou for providing photos. We gratefully acknowledge Robert Prŷs-Jones for providing considerable information on types; Edward C. Dickinson for providing extensive background information on collections and specifically on some of the types involved; Effie Warr for information on types history; Richard Schodde for providing further advice on typification and nomenclature; Tom Versluijs for providing help on statistical analysis; Peng Li, Jian Zhao and Min Zhao for field assistance; Sid Francis and Roland Zeidler for various information; René Pop for editing some photos for figures; Yang Liu for various assistance, including collecting of photos and locality information and for help with data collection at KIZ; Gobinathan Maheswaran, Zoological Survey of India and Pramod Nair, Salim Ali Centre for Ornithology and Natural History for providing information on specimens; and Wu Fei of KIZ for translating and providing information on localities of specimens. We gratefully acknowledge partial financial support from the Ministry of Science and Technology of China (Grant No. 2014FY210200, to. T.C. and Y.G.), the Russian Science Foundation (Project No. 14-50-00029, to M.K.), the Delia Koo Global Faculty Endowment of the Asian Studies Center, Michigan State University (to P.C.R.), The Sound Approach and Jornvall Foundation (both to P.A. and U.O.), the Chinese Academy of Sciences Visiting Professorship for Senior International Scientists (No. 2011T2S04, to P.A.) and Swarovski Optik Greater China (to P.A.).
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