Multilocus phylogeny of the avian family Alaudidae (larks) reveals complex morphological evolution, non-monophyletic genera and hidden species diversity

Per Alström, Keith N. Barnes, Urban Olsson, F. Keith Barker, Paulette Bloomer, Aleem Ahmed Khan, Masood Ahmed Qureshi, Alban Guillaumet, Pierre André Crochet, Peter G. Ryan

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47 Scopus citations


The Alaudidae (larks) is a large family of songbirds in the superfamily Sylvioidea. Larks are cosmopolitan, although species-level diversity is by far largest in Africa, followed by Eurasia, whereas Australasia and the New World have only one species each. The present study is the first comprehensive phylogeny of the Alaudidae. It includes 83.5% of all species and representatives from all recognised genera, and was based on two mitochondrial and three nuclear loci (in total 6.4. kbp, although not all loci were available for all species). In addition, a larger sample, comprising several subspecies of some polytypic species was analysed for one of the mitochondrial loci. There was generally good agreement in trees inferred from different loci, although some strongly supported incongruences were noted. The tree based on the concatenated multilocus data was overall well resolved and well supported by the data. We stress the importance of performing single gene as well as combined data analyses, as the latter may obscure significant incongruence behind strong nodal support values. The multilocus tree revealed many unpredicted relationships, including some non-monophyletic genera (Calandrella, Mirafra, Melanocorypha, Spizocorys). The tree based on the extended mitochondrial data set revealed several unexpected deep divergences between taxa presently treated as conspecific (e.g. within Ammomanes cinctura, Ammomanes deserti, Calandrella brachydactyla, Eremophila alpestris), as well as some shallow splits between currently recognised species (e.g. Certhilauda brevirostris-C. semitorquata-C. curvirostris; Calendulauda barlowi-C. erythrochlamys; Mirafra cantillans-M. javanica). Based on our results, we propose a revised generic classification, and comment on some species limits. We also comment on the extraordinary morphological adaptability in larks, which has resulted in numerous examples of parallel evolution (e.g. in Melanocorypha mongolica and Alauda leucoptera [both usually placed in Melanocorypha]; Ammomanopsis grayi and Ammomanes cinctura/deserti [former traditionally placed in Ammomanes]; Chersophilus duponti and Certhilauda spp.; Eremopterix hova [usually placed in Mirafra] and several Mirafra spp.), as well as both highly conserved plumages (e.g. within Mirafra) and strongly divergent lineages (e.g. Eremopterix hova vs. other Eremopterix spp.; Calandrella cinerea complex vs. Eremophila spp.; Eremalauda dunni vs. Chersophilus duponti; Melanocorypha mongolica and male M. yeltoniensis vs. other Melanocorypha spp. and female M. yeltoniensis). Sexual plumage dimorphism has evolved multiple times. Few groups of birds show the same level of disagreement between taxonomy based on morphology and phylogenetic relationships as inferred from DNA sequences.

Original languageEnglish (US)
Pages (from-to)1043-1056
Number of pages14
JournalMolecular Phylogenetics and Evolution
Issue number3
StatePublished - Dec 2013

Bibliographical note

Funding Information:
A close relationship between + b) is supported by all loci. Galerida , Alauda and Melanocorypha leucoptera (clade A2a Melanocorypha leucoptera is firmly nested in this clade, and hence far removed from the other Melanocorypha (A1c). The sister relationship with Alauda receives high PP and moderate bootstrap support, although this is only supported by ODC in the SLAs. This is further supported by a closer resemblance to Alauda than to Melanocorypha or Galerida in morphology, vocalisations, behaviour and ecology ( de Juana et al., 2004 ; P.A. and Krister Mild, unpublished), although – as has repeatedly been revealed by the present study – morphological similarity can be an extremely poor indicator of relationship among larks (see also Section 4.4 ). Galerida magnirostris and G. modesta have been placed in the monotypic genera Calendula ( Pätzold, 2003; Wolters, 1979 ) and Heliocorys ( Wolters, 1979 ), respectively.


  • Morphological evolution
  • Nodal support
  • Phylogeny
  • Taxonomy


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