Within clade A1, the unexpected sister relationships between the two monotypic genera Chersophilus and Eremalauda (A1b) and between this clade and the Calandrella rufescens–cheleensis–raytal–athensis complex (A1a) are well supported by the data. The strongly supported sister relationship between the Calandrella cinerea–brachydactyla–acutirostris complex (A1d) and Eremophila (A1e) is equally surprising. All of these relationships are recovered in SLAs of two unlinked loci and are not contradicted by any other SLAs, and the A1d + A1e clade also receives support from an indel in the ODC alignment. Accordingly, these relationships all seem robust. Eremalauda dunni often has been placed in Ammomanes (Meinertzhagen, 1951; Pätzold, 2003; Peters, 1960; Wolters, 1979 [subgenus Eremalauda]), but a close relationship with the type species of this genus (A. cinctura; clade C1b) is strongly refuted by the present study. Meinertzhagen’s (1951) placement of Chersophilus in Certhilauda (together with e.g. Alaemon and Chersomanes), based on especially bill structure and behaviour, is strongly rejected by our data.
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The similarities in size, structure and plumage between the two distantly related clades of traditional Calandrella (here recognised as Calandrella and Alaudala; cf. de Juana et al., 2004; Fig. 3) are likely the result of either retained plesiomorphies or parallel evolution. The similarity between the north African/west Asian Eremalauda dunni and Afrotropical Spizocorys starki, between the Western Palearctic Chersophilus and Afrotropical Certhilauda, and between the north African/west Asian Ammomanes and Afrotropical Ammomanopsis (cf. de Juana et al., 2004; Fig. 3) provide examples of close morphological similarity evolving independently in similar environments. In contrast, the dissimilarity between Ammomanopsis and its closest relatives, Chersomanes and Certhilauda, suggests strong divergence in the former.
The sister relationship between the genera Calandrella (as redefined here) and Eremophila suggests remarkable plumage divergence in the latter lineage (which is one of the most aberrant of all larks; cf. de Juana et al., 2004 and Fig. 3). Similarly, the close relationship between Alaudala (as redefined here; clade A1a) and the two monotypic genera Eremalauda and Chersophilus reveal extraordinary changes in both structure (especially bill) and plumage among sister taxa (cf. de Juana et al., 2004; Fig. 3). Meinertzhagen’s (1951) inappropriate placement of Chersophilus, Pseudalaemon, Calendulauda, Alaemon, Chersomanes and Certhilauda in one genus based on bill structure and behaviour (notably strong digging with the bill when feeding, and fast running) is a striking example of a misclassification caused by the strong lability and adaptability of bill morphology in larks.
