Abstract:
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. Westress 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.
