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Genetic, phenotypic and ecological differentiation suggests incipient speciation in two Charadrius plovers along the Chinese coast

Wang, Xuejing; Que, Pinjia; Heckel, Gerald; Hu, Junhua; Zhang, Xuecong; Chiang, Chung-Yu; Zhang, Nan; Huang, Qin; Liu, Simin; Martinez, Jonathan; Pagani-Núñez, Emilio; Dingle, Caroline; Yan Leung, Yu; Székely, Tamás; Zhang, Zhengwang; Liu, Yang


Xuejing Wang

Pinjia Que

Gerald Heckel

Junhua Hu

Xuecong Zhang

Chung-Yu Chiang

Nan Zhang

Qin Huang

Simin Liu

Jonathan Martinez

Caroline Dingle

Yu Yan Leung

Tamás Székely

Zhengwang Zhang

Yang Liu


Speciation with gene flow is an alternative to the nascence of new taxa in strict allopatric separation. Indeed, many taxa have parapatric distributions at present. It is often unclear if these are secondary contacts, e.g. caused by past glaciation cycles or the manifestation of speciation with gene flow, which hampers our understanding of how different forces drive diversification. Here we studied genetic, phenotypic and ecological aspects of divergence in a pair of incipient shorebird species, the Kentish (Charadrius alexandrinus) and the White-faced Plovers (C. dealbatus), shorebirds with parapatric breeding ranges along the Chinese coast. We assessed divergence based on molecular markers with different modes of inheritance and quantified phenotypic and ecological divergence in aspects of morphometric, dietary and climatic niches.

Our integrative analyses revealed small to moderate levels of genetic and phenotypic distinctiveness with symmetric gene flow across the contact area at the Chinese coast. The two species diverged approximately half a million years ago in dynamic isolation with secondary contact occurring due to cycling sea level changes between the Eastern and Southern China Sea in the mid-late Pleistocene. We found evidence of character displacement and ecological niche differentiation between the two species, invoking the role of selection in facilitating divergence despite gene flow.

These findings imply that ecology can indeed counter gene flow through divergent selection and thus contributes to incipient speciation in these plovers. Furthermore, our study highlights the importance of using integrative datasets to reveal the evolutionary history and assist the inference of mechanisms of speciation.


Wang, X., Que, P., Heckel, G., Hu, J., Zhang, X., Chiang, C., Zhang, N., Huang, Q., Liu, S., Martinez, J., Pagani-Núñez, E., Dingle, C., Yan Leung, Y., Székely, T., Zhang, Z., & Liu, Y. (2019). Genetic, phenotypic and ecological differentiation suggests incipient speciation in two Charadrius plovers along the Chinese coast. BMC Evolutionary Biology, 19(1), Article 135.

Journal Article Type Article
Acceptance Date May 29, 2019
Online Publication Date Jun 27, 2019
Publication Date 2019
Deposit Date Nov 2, 2022
Publicly Available Date Nov 2, 2022
Journal BMC Evolutionary Biology
Publisher BMC
Peer Reviewed Peer Reviewed
Volume 19
Issue 1
Article Number 135
Keywords Parapatry, Character displacement, Gene flow, Hybridization, Stable isotope analysis, Ecological niche
Public URL


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