Liolaemus huacahuasicus, Laurent, 1985

Liolaemus huacahuasicus View in CoL clade

This clade includes 14 terminals ( Fig. 3 View Figure 3 ), all medium-sized lizards (maximal SVL = 85 mm) with triangular, imbricate, keeled dorsal body scales, with the dark scales being more strongly keeled than the lighter scales. Body colour in males varies between chestnut and yellow, and there are dark paravertebral blotches with light borders and a generally yellowish venter. This clade is supported by one discrete morphological character, four discrete coloration characters and eight molecular characters (Supporting Information, Table S1). The lizards of the L. huacahuasicus clade are distributed in the extra-Andean sierras of Argentina ( L. montanus , L. aff. montanus , Liolaemus griseus Laurent, 1984 , L. orko , L. huacahuasicus and Liolaemus pulcherrimus Laurent, 1982 ), Puna and high Amazonian slopes of Bolivia ( L. signifer , L. fittkaui and L. aff. signifer 2), Chilean Puna ( L. aff. signifer 1) and the Puna and western ranges of Peru ( Liolaemus annectens Boulenger, 1901 , L. aff. annectens , Liolaemus etheridgei Laurent, 1998 , L. evaristoi Gutiérrez et al., 2018 and L.signifer ) ( Fig. 6 View Figure 6 ). The composition of the L. huacahuasicus clade was recovered in all analyses except the hypotheses where K = 3–8, in which L. annectens , L. evaristoi , L. signifer , L. etheridgei , L. aff. annectens and L. aff. signifer 1 formed a group separate from the rest of the L. huacahuasicus clade and sister to the L. robustus clade, whereas the monophyletic group formed by L. montanus , L. aff. montanus , L. griseus , L. fittkaui , L. orko , L. huacahuasicus , L. pulcherrimus and L. aff. signifer 2 were related to part of the L. ortizi clade.

MIXED BAYESIAN HYPOTHESIS

This hypothesis was highly congruent with that obtained with ‘parsimony’ ( Fig. 7). The clades and groups recovered within the L. montanus group were the same, with changes in composition in only a few instances. The L. chlorostictus , L. multicolor , L. reichei , L. forsteri and L. robustus clades had the same species composition, with only some changes in the internal relationships ( Fig. 7). The clade with the greatest differences with regard to the ‘parsimony’ hypothesis was the L. huacahuasicus clade, which included members of the L. andinus clade ( L. robertoi Pincheira-Donoso & Nuñez, 2003 , L. vallecurensis and L. aff. poecilochromus ), the L. dorbignyi clade ( L. aff. jamesi 3, L. aff. jamesi 1 and L. aff. jamesi 2), the L. jamesi clade ( L. aymararum ) and the L. ortizi clade ( L. aff. multiformis ) ( Fig. 7).

MORPHOLOGY-BASED HYPOTHESIS (MH)

In all runs performed with the morphology-only matrix, the L. montanus group was never recovered as monophyletic; instead, the outgroup species L. abaucan , L. inacayali , L. kingii , L. kolengh , L. pseudoanomalus , L. robertmertensi and L. tiranti were included within the L. montanus group, always as terminals of the L. ortizi group. Nevertheless, owing to the evidence for monophyly provided by the TEH and molecular hypotheses in the present study and by other authors ( Schulte et al., 2000, 2013; Espinoza et al., 2004; Aguilar-Puntriano et al., 2018) and understanding that the outgroup and the characters proposed to separated them are not sufficient to demonstrate monophyly, we have decided to present only the MH without the conflicting terminals under the assumption of a monophyletic L. montanus group.

Various hypotheses with different typologies and compositions were obtained for the clades within the L. montanus group, as a function of the K values applied. Based on the criteria proposed by Mirande (2009), we selected the most stable phylogenetic hypothesis, in which K = 14, as our proposed hypothesis ( Fig. 8 View Figure 8 ). This same hypothesis was also recovered with K = 12–16. Thus, the proposed hypothesis was highly congruent with the hypothesis obtained when K = 7–11, recovering the same groups, albeit with some differences in their relationships and composition. In the proposed MH, the L. montanus group was supported by 31 characters, of which nine were continuous, 11 were lepidosis characters and the remaining 11 were miscellaneous.

In the proposed MH, ten clades were recovered, of which eight were congruent with those proposed by the TEH ( Fig. 3 View Figure 3 ). The species L. nigriceps and L. patriciaiturrae were placed basally and outside of the L. andinus clade, as also occurred in the TEH. The following clades were recovered ( Fig. 8 View Figure 8 ): ( L. nigriceps ( L. patriciaiturrae ( L. chlorostictus clade ( L. jamesi clade + L. dorbignyi clade) + ( L. forsteri clade