The South American lizard genus Liolaemus comprises > 260 species, of which > 60 are recognized as members of the Liolaemus montanus group, distributed throughout the Andes in central Peru, Bolivia, Chile and central Argentina. Despite its great morphological diversity and complex taxonomic history, a robust phylogenetic estimate is still lacking for this group. Here, we study the morphological and molecular diversity of the L. montanus group and present the most complete quantitative phylogenetic hypothesis for the group to date. Our phylogeny includes 103 terminal taxa, of which 91 are members of the L. montanus group (58 are assigned to available species and 33 are of uncertain taxonomic status). Our matrix includes 306 morphological and ecological characters and 3057 molecular characters. Morphological characters include 48 continuous and 258 discrete characters, of which 70% (216) are new to the literature. The molecular characters represent five mitochondrial markers. We performed three analyses: a morphology-only matrix, a molecular-only matrix and a matrix including both morphological and molecular characters (total evidence hypothesis). Our total evidence hypothesis recovered the L. montanus group as monophyletic and included ≥ 12 major clades, revealing an unexpectedly complex phylogeny.

The genus Liolaemus is a group of lizards with more than 230 recognized species, which have been grouped in differentclades and subgroups. One of the monophyletic groups is the one of Liolaemus boulengeri or “the patch group”; this cladeitself is integrated by several monophyletic groups: the groups Liolaemus anomalus, Liolaemus wiegmanii, Liolaemusdarwinii and Liolaemus melanops. The latter group is constituted almost exclusively by Patagonian lizards, and it is ourfocal group. In the present work we describe three new species belonging to the Liolaemus melanops group. One of thenew taxon described, Liolaemus tromen sp. nov., is related to the Liolaemus fitzingerii clade, inhabits in the Center-Westof Neuquén Province, and was confused initially with Liolaemus hermannunezi, a mountain Chilean species. The secondspecies described, Liolaemus purul sp. nov., belongs to the Liolaemus telsen clade, inhabits in the center and north-westof Neuquén Province, and was confused first with Liolaemus boulengeri and then with Liolaemus loboi. While the thirdspecies, Liolaemus dumerili sp. nov., is basal of the L. goetschi group, inhabits in the center-south of Río Negro provinceand was considered as Liolaemus melanops. In addition, we present a phylogenetic analysis based in the morphology and we contribute with a dichotomic key for males of the Liolaemus melanops group.

Phymaturus is a clade of lizards that occurs at moderate to high elevations in western Argentina and the adjacent central region of Chile, as well as in various volcanic plateaus of the Patagonian region of Argentina. This genus had previously been divided into two groups: the patagonicus and the palluma groups. In this study, we analyzed relationships within the patagonicus group. The data set was built for 23 species plus nine other terminal taxa of undetermined taxonomic status. In total, 10,631 bp (ND4, Cytb, 12S, COI, five protein coding nuclear genes and seven anonymous nuclear loci) and 254 morphological characters were analyzed in a combined data set for 35 ingroup taxa and nine outgroups. We also ran separate DNA sequence and morphological data sets. We identified four main clades, and revealed congruencies and incongruences with previous studies. The indistinctus clade is recovered as the most basal within the patagonicus group in the strict parsimony analysis, while the somuncurensis clade is the most basal under Bayesian inference. The previously recovered calcogaster clade resulted paraphyletic in both analyses and part of their species are included in a redefined somuncurensis clade. We found low support at basal nodes provoked in part by contradictory evidence shown by rogue taxa. We show the phylogenetic information given by each partition/marker and how they contribute to relationships found in the total evidence analysis. We discuss the phylogenetic position of Phymaturus manuelae, Phymaturus tenebrosus, and Phymaturus patagonicus.

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The Liolaemus montanus group is a diverse group of lizards that ranges from central Peru to southwestern Mendoza, Argentina, including much of the Plurinational State of Bolivia (“Bolivia”) and Chile. The species of this group mainly inhabit high elevation areas with cold temperatures. In the last years, several species of this group have been described, mostly in Argentina and Chile. In Bolivia, there are at least thirteen valid species belonging to the L. montanus group. In this study, we describe a new species of the L. montanus group with a marked endemism in the Cordillera de Sama of the Tarija Department, Bolivia, and a combination of unique character states that allows its formal description as a new species. The phylogenetic relationships based on analysis of 159 morphological characters suggest that it belongs to the L. montanus group, and that it is closest to Liolaemus pulcherrimus, which is found allopatrically in a small area of the Jujuy Province, Argentina. The multivariate analyses of 66 morphological characters support the phylogenetic relationships. Statistical analyses of inter-species comparisons of morphological characters are not considered the only methods due to the non-independence of some characters states among species; thus, a phylogenetic analysis is recommended. The detailed revision of specimens of the L. montanus group held in the collections of Bolivia is filling major geographic gaps and improving our understanding of the phylogenetic and biogeographic relationships of this widely distributed group of South American lizards.

New Liolaemus (Iguania: Liolaemidae) with a new morphological character from the border between Argentina and Chile. Liolaemus belongs to the Eulaemus subgenus, that is included in the Liolaemus montanus group or section, and is distributed in Argentina, Bolivia, Chile and Peru. The species members of this clade inhabit arid and high altitude environments, usually above 2 800m, and have a strong tendency to herbivory and a viviparous reproductive type. In a herpetological expedition to the Salta Province highlands, Argentina, in April 2012, we collected individuals of this new species for their description. The specimens were collected with loop, slaughtered by anesthesia and deposited in the herpetological collection of Fundación Miguel Lillo in Tucumán (FML). To characterize this new taxon, we analyzed 144 external morphological characters, referring primarily to lepidosis, color pattern, and body proportions, comparing with data from 66 constituent species of the L. montanus group (45 taxa from collection specimens and 21 from literature). Description of the colors in life was made in the field and based on photographs taken during the capture. The species described in this work inhabits a very narrow mountain range area in Salta Province (Argentina), and the Atacama Region (Chile), always above the 3 600m. This new taxon shows characteristics of lepidosis and color patterns clearly different from the other species members of the L. montanus group: the maximum snout-vent length is 72.9mm; shows 74-96 scales around midbody, 89-104 dorsal scales between the occiput and hind limbs, 92-109 ventrals, precloacal pores are evident in both, males and females, and supernumerary pores in males. Also, the presence of postcloacal pores stands out as unique among all Liolaemus. This is a new character, not only for the genus, but also to all Reptilia, becoming a morphological discovery of great importance, since in reptiles only precloacal and femoral pores are known. Based on our field observations, this new Liolaemus is related to rocky environments, which are used as refuge by this omnivorous species, and when handled, very aggressive males. Rev. Biol. Trop. 61 (4): 1563-1584. Epub 2013 December 01.

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