On the taxonomy of the desert lizard Liolaemus stolzmanni (Steindachner, 1891): A third point of view (Squamata: Liolaemidae).

Recently, two publications, Langstroth (2011) and Valladares-Faúndez et al. (2018), have provided relevant information on the taxonomy of the poorly know South American lizard, Liolaemus stolzmanni (Steindachner 1891).. However, these publications contradict each other, in regards to L. reichei (Werner 1907) as junior synonym of L. stolzmanni and did not solve the problem of the type locality of L. stolzmanni. In fact, currently the type locality of L. stolzmanni is thought to be imprecise because the description (Steindachner 1891), written in German, stated it as "Hoch Peru" (High Peru), which has been erroneously understood as "northern Peru" or "high altitudes from Peru", a zone in which L. stolzmanni has been never found. Here, we used the book of memories of Jan Stanislaw Sztolcman, the Polish collector, a historical data never taken in account before, to restrict the type locality and provide comments on its taxonomy.

Liolaemus jamesi (Boulenger, 1891): restriction of the type locality and holotype characterization (Squamata: Liolaemidae).

Liolaemus is one of the most diverse genus of lizards in the world (Esquerré et al. 2013), with 257 species listed in the last review focusing on its diversity (Abdala Quinteros 2014). Certain species within this genus, especially the earlier ones, were described in very little detail, even lacking an appropriate description of the holotype and/or type locality. This has created uncertainties in the taxonomic identity of several Liolaemus species (Espinoza et al. 2011; Quinteros et al. 2008; Troncoso-Palacios Garin 2013; Langstroth 2011).

Speciation across mountains: Phylogenomics, species delimitation and taxonomy of the Liolaemus leopardinus clade (Squamata, Liolaemidae).

Organisms rapidly diversifying across unstable environments such as mountain tops provide substantial challenges for resolving evolutionary histories and delimiting species. The Liolaemus leopardinus clade is a group of five species of lizards adapted to high altitudes in central Chile, with most species found in the Andes, but one species, L. frassinettii is found in the independent Costa Cordillera. Despite their allopatric distributions, they display shallow mitochondrial divergences, making phylogenetics and species delimitation of this clade hard to resolve. We use an integrative approach to delimit species by considering morphological data (linear and landmark-based), mitochondrial DNA (mtDNA), and nuclear DNA (Sequences and SNPs collected with ddRADseq). We find strong conflicting signals between phylogenetic analyses of the nuclear and mtDNA data. While mtDNA places L. frassinettii as sister to the rest of the clade, the SNPs support a south to north order of divergences, with southernmost species (new taxon described here) as sister to the rest of the clade. Moreover, species delimitation using mtDNA only supports two species (one in the Costa and one in the Andes), whereas combined analyses using the nuclear data and morphology support multiple Andean taxa, including a new one we describe here. Based on these results, population structure analyses and our knowledge of the geological and climatic history of the Andes, we argue that this mito-nuclear discordance is explained by past introgression among the Andean taxa, likely during glacial periods that forced these lizards to lower altitudes where they would hybridize. The complete isolation between the Costa and Andes cordilleras has prevented any further contact between taxa on either mountain chain. Our study highlights the importance of using multiple lines of evidence to resolve evolutionary histories, and the potential misleading results from relying solely on mtDNA.

Without a body of evidence and peer review, taxonomic changes in Liolaemidae and Tropiduridae (Squamata) must be rejected.

In his recent self-published book "Reptiles en Chile", Diego Demangel Miranda presented 13 taxonomic changes for liolaemid and tropidurid lizards. While these could be considered validly published according to the International Code of Zoological Nomenclature, we show that these taxonomic propositions lack the necessary scientific rigor in terms of replicability, specimen work, lack of peer review and that they do not follow best practices accepted by the herpetological community. Therefore, we hereby invalidate all 13 taxonomic changes proposed in this book, leaving the taxonomy unaffected. Finally, we call attention to the potentially negative consequences of using these taxonomic changes in conservation and environmental impact studies as incorrect decisions might be taken in relation to the species involved.

An updated phylogeny and morphological study of the Phymaturus vociferator clade (Iguania: Liolaemidae).

Species delimitation in Phymaturus has been a difficult task due to the highly conserved morphological and ecological features present in this genus. Almost all species of Phymaturus have been described without DNA data or lacking statistical analyses which makes even more difficult to compare species. Although two molecular phylogenetic studies have been recently published, here we provide the first multilocus phylogenetic reconstruction including all Chilean species, with samples from all type localities and some previously unsampled populations. We also estimate pairwise distances among the Chilean species of Phymaturus (P. vociferator and P. mallimaccii clades) and compare our results with the P. payuniae clade, where previous studies have used multiple lines of evidence. Additionally, we performed univariate and multivariate morphological analyses and skeletal comparisons (clavicle) for the species of the P. vociferator clade. As a result of this integrative approach, we describe a new species.

The shadow of the past: Convergence of young and old South American desert lizards as measured by head shape traits.

Convergence is a pervasive phenomenon in the Tree of Life, and evolution of similar phenotypes sharing the same environmental conditions is expected in phylogenetically closely related species. In contrast, contingent factors are probably more influential in shaping phenotypic diversity for distantly related taxa. Here, we test putative convergent evolution of lizard head morphologies among relatively closely related desert dwelling Liolaemus species, and the very distantly related Ctenoblepharys adspersa. We estimated a multilocus time-calibrated phylogeny of 57 species of South American liolaemus lizards, based on seven molecular markers. We collected head shape data for 468 specimens, and used three phylogenetic comparative methods (SURFACE, CONVEVOL, and WHEATSHEAF index) to test for and estimate the strength of convergence. We found strong evidence for convergence among Pacific desert lizard C. adspersa, Liolaemus audivetulatus, Liolaemus insolitus, Liolaemus poconchilensis, Liolaemus stolzmanni, and a candidate species (Liolaemus "Moquegua"). Our results suggest that, despite the long divergence and phylogenetic distance of C. adspersa with respect to convergent Liolaemus species, natural selection was probably more important than historical contingency in shaping phenotypic evolution in these desert lizards.

The True Identity of the New World Iguanid Lizard Liolaemus chillanensis Müller and Hellmich 1932 (Iguania: Liolaemidae) and Description of a New Species in the Liolaemus elongatus Group.

Jaime Troncoso-Palacios, Damien Esquerré, Félix A. Urra, Hugo A. Díaz, Carlos Castro-Pastene, and María Soledad Ruiz (2018) Liolaemus is a particularly species-rich radiation of New World iguanid lizards from southern South America. Thanks to intense systematic and taxonomic research, the knowledge on its species- level diversity has increased rapidly over the past several years. The L. elongatus species-complex has received considerable attention and a remarkable case is Liolaemus chillanensis, a name that has been used for two different species that are sympatric in Termas de Chillán, central Chile. Since the holotype is lost, we propose that the first step to identify the true L. chillanensis is through the analysis of the original description. Then we provide a morphological and molecular characterization of L. chillanensis based on topotypes and a description of the taxon previously confused with it.

Two new Liolaemus lizards from the Andean highlands of Southern Chile (Squamata, Iguania, Liolaemidae).

Liolaemus is a diverse genus of lizards, subdivided into two subgenera: Liolaemus (sensu stricto) and Eulaemus, distributed mainly in Chile and Argentina. The Liolaemus elongatus-kriegi complex is the most diverse group within Liolaemus (sensu stricto), especially the species closely related to Liolaemus elongatus, which form a clade currently comprising nine species. Several Chilean species of this group have been recently described, mainly from volcanoes and poorly explored mountains. Here molecular and morphological evidence are provided for a new species of the Liolaemus elongatus clade, which is characterized by its small size and lack of dorsal pattern, unusual features for the species of this group of lizards. Additionally, the lack of precloacal pores in males of Liolaemus (sensu stricto) is a trait found in few species, which do not constitute a monophyletic group. A second new southern Chilean species is also described, without precloacal pores and supported by molecular phylogenetics to be related to Liolaemus villaricensis. Both new species were found in the same locality, near a lake located in a pre-Andean zone with Araucaria and Nothofagus forest. The two species are dedicated to prominent Lonkos (tribal chiefs) of the Mapuche and Pehuenche people: Janequeo and Leftraru. Additionally, the phylogenetic results suggest that Liolaemus lonquimayensis is a synonym of Liolaemus elongatus.

A new species of Liolaemus related to L. nigroviridis from the Andean highlands of Central Chile (Iguania, Liolaemidae).

The Liolaemus nigroviridis group is a clade of highland lizards endemic to Chile. These species are distributed from northern to central Chile, and currently there are no cases of sympatric distribution. This study describes a new species, Liolaemus uniformis sp. n., from this group, and provides a detailed morphological characterization and mitochondrial phylogeny using cytochrome-b. Liolaemus uniformis was found in sympatry with Liolaemus nigroviridis but noticeably differed in size, scalation, and markedly in the color pattern, without sexual dichromatism. This new species has probably been confused with Liolaemus monticola and Liolaemus bellii, both of which do not belong to the nigroviridis group. The taxonomic issues of this group that remain uncertain are also discussed.

Two new species of the Liolaemuselongatus-kriegi complex (Iguania, Liolaemidae) from Andean highlands of southern Chile.

The elongatus-kriegi complex is one of the most diverse clades of the Liolaemus (sensu stricto) subgenus of lizards. There are currently 29 species recognized in this group distributed between Chile and Argentina. Based on molecular evidence, there seem to be five main clades nested within this complex: the elongatus, leopardinus, kriegi, petrophilus and punmahuida clades. Liolaemusbuergeri and Liolaemuskriegi, both of the kriegi clade, were believed to inhabit the surroundings of the Laja Lagoon, in the Biobío Region of Chile. Moreover, this Chilean population of Liolaemuskriegi was recently recognized as an undescribed taxon called "Liolaemus sp. A" based on molecular phylogenetics. In this work, we studied these two populations of the Laja Lagoon and provided the morphological diagnosis to describe them as two new species: Liolaemusscorialis sp. n. and Liolaemuszabalai sp. n., previously considered Liolaemusbuergeri and "Liolaemuskriegi/Liolaemus sp. A" respectively. Additionally, we identified another population of Liolaemusscorialis in the vicinity of La Mula Lagoon in the Araucanía Region of Chile. Liolaemusscorialis differs from almost all of the species of the elongatus-kriegi complex by its considerably smaller size. Nevertheless, without molecular data we cannot assign it to any particular subclade. Liolaemuszabalai belongs to the kriegi clade based on published molecular phylogenies. Finally, we provide some natural history data on both species and we document for the first time the presence of Liolaemusneuquensis in Chile from a museum specimen from La Mula Lagoon.

The missing leopard lizard: Liolaemus ubaghsi sp. nov., a new species of the leopardinus clade (Reptilia: Squamata: Liolaemidae) from the Andes of the O'Higgins Region in Chile.

Liolaemus is an extremely species rich genus of iguanid lizards from southern South America. Most of the diversity though is found in the Andes Cordillera, between Argentina and Chile. Here we describe Liolaemus ubaghsi sp. nov., from El Teniente Mine, in the Andean mountains of the O'Higgins Region in Chile. This species presents scalation and pattern traits that belong to the leopardinus clade, a group of viviparous, high altitude lizards that inhabit the mountain ranges surrounding Santiago City. The species of this clade in turn belong to the Andean and Patagonian elongatus-kriegi complex. Liolaemus ubaghsi sp. nov. has been previously recognized as L. leopardinus and L. elongatus, nevertheless we present diagnostic traits that allow us to describe it as a new species. It mainly differs from the rest of the leopardinus clade (L. leopardinus, L. ramonensis, L. valdesianus and L. frassinettii) by having the following unique combination of traits: ochre background coloration, a wide dark occipital stripe, dark flanks, white dots dispersed on the dorsum, absence of leopard-like spots and enlarged infralabial scales. 

On the identity of Liolaemus nigromaculatus Wiegmann, 1834 (Iguania, Liolaemidae) and correction of its type locality.

In the current study, we review the taxonomic status of Liolaemus nigromaculatus. Despite being the nominal species of the nigromaculatus group and being the second species of the genus Liolaemus that was described, this species is of uncertain type locality and its true identification is a matter of discussion. After carefully analyzing several digital pictures of the holotype (juvenile male), reviewing all of the literature concerning the issue, examining specimens of nearly all recognized species of the nigromaculatus group, and determining the locations visited by the specimen collector, we are able to point out the following: 1) Liolaemus nigromaculatus was collected between Puerto Viejo and Copiapó of the Atacama region in Chile, and not in Huasco 2) Liolaemus bisignatus is a nomen nudum, and populations attributed to Liolaemus bisignatus should be referred to as Liolaemus nigromaculatus. 3) There is agreement that Liolaemus copiapoensis is indistinguishable from populations currently referred to as Liolaemus bisignatus (= Liolaemus nigromaculatus), 4) Populations found in Huasco (currently considered the type locality of Liolaemus nigromaculatus) are very similar to those found in Caldera (currently considered Liolaemus bisignatus) and should be designated as Liolaemus nigromaculatus, and 5) Liolaemus oxycephalus and Liolaemus inconspicuus are not synonymous with Liolaemus nigromaculatus, although their true identities are difficult to determine. We also detail several characteristic based on the holotype of Liolaemus nigromaculatus, in addition to drawing diagnostic comparisons between this species and others belonging to the nigromaculatus group.

The rediscovery of Oplurus bibronii Guichenot, 1848 a valid species of the liolaemid genus Phymaturus (Iguania: Liolaemidae).

Oplurus bibronii was a species described more than 160 years ago from the highlands of Ovalle, Chile. The species was earlier synonymized with Phymaturus palluma and since then its taxonomic status has not been re-examined. In February of 2011, we were successful in capturing specimens of Phymaturus in the highlands of Ovalle and additionally we examined high quality digital pictures of the type series of O. bibronii. Our analysis shows that the type series is composed of two different species, one of which overlaps their diagnostic characters with the characters of the specimens collected in the highlands of Ovalle, and therefore they correspond to Phymaturus bibronii (new combination). In addition, our review of the holotype, and some paratypes and topotypes of the recently described P. paihuanense shows that this is a junior synonym of P. bibronii.