A Large Evaluation of Passerine Cisticolids (Aves: Passeriformes): More About Their Phylogeny and Diversification

Billy Nguembock*, 1, Corinne Cruaud2, Christiane Denys3, 4
1 Laboratoire de Zoologie, Département de Biologie et Physiologie Animales, Université de Yaoundé I, BP 812 Yaoundé, Cameroun
2 Génoscope, Centre National de Séquençage, 2, rue Gaston Crémieux, CP5706, 91057 Evry Cedex, France
3 UMR 7205 Unité Origine, Structure et Evolution de la Biodiversité, Département Systématique et Evolution, Muséum National d’Histoire Naturelle, 55 Rue Buffon, 75005 Paris, France
4 Service de Systématique Moléculaire, IFR CNRS 101, Muséum National d’Histoire Naturelle, 43, Rue Cuvier, 75005 Paris, France

© 2012 Nguembock et al.

open-access license: This is an open access article distributed under the terms of the Creative Commons Attribution 4.0 International Public License (CC-BY 4.0), a copy of which is available at: ( This license permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

* Address correspondence to this author at the Department of Animals Biology and Physiology, Faculty of Sciences, University of Yaounde I, P.O. Box 812 Yaounde, Cameroon; Tel: 00 237 94144294; Fax: 00 237 22 23 53 86; E-mails:


African warblers (or Cisticolidae family) are small perching song birds with a large number of genera distrib-uted throughout the World. The phylogeny of Cisticolidae is well-supported but their diversification is currently poorly known. To deepen their understanding of phylogeny and investigate their diversification, we sequenced four loci (mito-chondrial ATPase 6, ND2 and ND3, and nuclear myoglobin intron 2) for several new cisticolid taxa and added several other sequences. Our analyses retrieve the monophyly of the African warblers and confirm Neomixis as their deepest branch. A group of taxa appear as their potential sister-taxa with our ND2 analyses but not with our combined analyses. New relationships are well-supported. Thus Scotocerca inquieta nests in the cettid clade whereas Camaroptera super-ciliaris, Cisticola chubbi, Cisticola tinniens, Prinia flavicans and Poliolais lopezi belong to the cisticolid clade. Our re-sults support a splitting of the African warblers in two main clades. The first clade consists of genera Orthotomus, Prinia, Cisticola, Scepomycter, Incana, Bathmocercus, Eminia, Hypergerus and Heliolais while the second includes genera Poli-olais, Camaroptera, Urolais, Artisornis, Oreolais, Apalis, Schistolais, Calamonastes and Spiloptila. Our results confirm the polyphyly of Orthotomus which generates a muddle between some Cettia species and other Asian tailorbirds. Waiting a study with Orthotomus sepium Horsf., 1821, to clarify their taxonomy, we suggest temporarily that the name Phyller-gates cucullatus Temminck, 1836, refers to Orthotomus cucullatus of the present study. Our dating analysis reveals that cisticolid clades began their diversification during the transition Early-Middle Miocene epoch and at the beginning of the Pliocene epoch. The diversification of the “open cisticolid” clade would have occurred during the Middle Miocene but that of the “forest cisticolid” lineage would have taken place during the Upper Miocene.

Keywords: African warblers, maximum-likelihood, Bayesian inference, polyphyly, taxonomy, molecular dating, diversify cation, Miocene, Pliocene.