Article

A phylogenetic analysis of lizards of the Liolaemus chiliensis group (Iguania: Tropiduridae)

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Abstract

The lizard genus Liolaemus includes over 160 species of which almost half are in the chiliensis group. Although some researchers have attempted to define smaller species groups within this large clade, the relationships among the taxa within the group as a whole remain enigmatic. The objectives of this study were to (1) identify characters that will be useful for present and future phylogenetic studies of this group, and (2) generate preliminary phylogenetic hypotheses for taxa within this large clade of lizards. I examined more than 800 specimens of 73 taxa belonging to the chiliensis group from which I identified 55 phylogenetically informative morphological characters. Additional characters (6) were derived from published and unpublished data on chromosomes, life history, and ecology. Four species considered basal for the genus were taken as outgroups. A tree-building program (PAUP 4.062) recovered three trees of length 11.516 (Retention index: 0.59). Differences found among these topologies were restricted to the relationships of species of the elongatus group, in which monophyly was recovered in only one tree. Results from PAUP's analysis support the monophyly of several previously proposed species groups: alticolor, altissimus, gravenhorstii, hellmichi, kriegi, leopardinus, monticola, nigromaculatus, nigroviridis, pictus and tenuis. Interestingly, most of the groups indicated above are endemic to areas that have recently been described as areas of high endemism for southern South America.

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... Subsequently, Lobo (2005) and Díaz Gómez and Lobo (2006) proposed a new clade within the L. elongatus group, named L. capillitas group, which included the species of the L. petrophilus group by Morando et al. (2003), distributed in north-western Argentina. Based on a consensus of previous studies (Etheridge, 1995;Schulte et al., 2000;Lobo, 2001Lobo, , 2005Espinoza et al., 2004), Lobo et al. (2010) defined the L. kriegi group (four species) and the L. elongatus group, which includes the L. capillitas clade (15 species), grouping the species in the L. elongatus and L. petrophilus groups by Morando et al. (2003). More recently, Avila et al. (2010) recovered four clades: the L. elongatus, L. kriegi, L. petrophilus (including the L. capillitas subclade), and L. punmahuida clades. ...
... Table 1 summarises the ages of the group and clades estimated by different studies. Very few systematic studies on Liolaemus incorporated morphological evidence (Lobo, 2001;Abdala, 2005;Lobo, 2005;Abdala, 2007;Lobo et al., 2010;Quinteros, 2013) and even fewer studies incorporated genital morphology, which proved to be very informative within Liolaemus (Quipildor et al., 2018). The main aim of this study was to describe a new species of the Liolaemus elongatus group, which we recognized as distinct from other members based on morphological characters (scalation, hemipenis, color pattern), and DNA sequences. ...
... Field studies did not involve endangered species. We studied the morphological characters commonly included in Liolaemus taxonomic studies, such as those described in Laurent (1985), Frost (1992), Etheridge (1993Etheridge ( , 1995Etheridge ( , 2000, Cei (1986), Lobo and Espinoza (1999), Lobo (2001Lobo ( , 2005, Abdala (2007), Quinteros (2013) and Quipildor et al. (2018), which altogether conform a record of more than 200 morphological characters, including scale counts, shape, ornamentation, variation in the disposal of the scales (imbrications), precloacal pores, neck folding, color pattern, life colors, hemipenis, among others. The description of colors in life was made based on photographs taken in the field immediately after capture of individuals. ...
... In 2014, Abdala and Quinteros (2014) reviewed the family Liolaemidae, and later presented another proposal to avoid the spread of publications that lacked scientific rigor. Several proposals of group rearrangements have been published (e.g., Lobo 2001Lobo , 2005Avila et al. 2006;Abdala 2007a;Fontanella et al. 2012;Breitman et al. 2013;Olave et al. 2014Olave et al. , 2015Troncoso-Palacios et al. 2015a); here, we use the arrangement shown in Table 7.3. ...
... Cei (1986) proposed this group to include L. gravenhorstii and L. cyanogaster. Lobo (2001) presented a morphological phylogeny expanding this group, to include L. chiliensis, L. nitidus, L. robertmertensi, and L. schroederi. Lobo later (2005) updated the morphological phylogeny and found that these species form a single or two separate groups, depending on the type of analysis. ...
... Given the known vs potential Andean distribution of this clade, its diversity and the number of candidate species, additional fieldwork should target remote Andean cordilleras and further phylogeographic and phylogenetic research. Medina et al. (2017) presented a multilocus phylogeny that is largely congruent with relationships inferred from morphological characters (Lobo 2001;and Lobo et al. 2010b, bibliographical review). These studies inferred the L. kriegi complex as sister to the (L. ...
Chapter
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The diversity of the Patagonian lizard fauna is a combination of low diversity at higher levels, e.g., families, some of them almost marginal to the region, coupled with very high species diversity concentrated in two genera Liolaemus and Phymaturus, and a high number of endemics. The number of described species almost tripled since Cei’s last and only monograph on Patagonian herpetofauna in 1987. But changes were not limited to species numbers; taxonomy changed as studies of this fauna improved with the use of modern taxonomic techniques and dedicated field surveys made to some previously unknown areas. The purpose of this chapter is to provide a taxonomic summary of Patagonian lizards. We include a number of plates with pictures of poorly known and/or endemic Patagonian species probably never included in any book or publication.
... (Etheridge, 1995;Schulte II et al., 2000;Lobo et al., 2010;among others). Subsequently, many new groups were proposed for each of these subgenera (Ortiz, 1981;Cei, 1986Cei, , 1993Schulte II et al., 2000;Lobo, 2001Lobo, , 2005Avila et al., 2006Avila et al., , 2007Abdala, 2007;Lobo et al., 2010;Quinteros, 2012;Abdala & Juárez Heredia, 2013;Quinteros, 2013;Portelli & Quinteros, 2018; among many others), including 13 monophyletic groups for the subgenus Liolaemus s.s. (Lobo, 2001(Lobo, , 2005Lobo et al., 2010;Portelli & Quinteros, 2018). ...
... Subsequently, many new groups were proposed for each of these subgenera (Ortiz, 1981;Cei, 1986Cei, , 1993Schulte II et al., 2000;Lobo, 2001Lobo, , 2005Avila et al., 2006Avila et al., , 2007Abdala, 2007;Lobo et al., 2010;Quinteros, 2012;Abdala & Juárez Heredia, 2013;Quinteros, 2013;Portelli & Quinteros, 2018; among many others), including 13 monophyletic groups for the subgenus Liolaemus s.s. (Lobo, 2001(Lobo, , 2005Lobo et al., 2010;Portelli & Quinteros, 2018). others) three populations without taxonomic status. ...
... We studied the classical morphological characters traditionally used in Liolaemus taxonomy, including those of Laurent (1985), Cei (1986Cei ( , 1993, Etheridge (1993Etheridge ( , 1995Etheridge ( , 2000, Lobo (2001Lobo ( , 2005, Abdala (2002Abdala ( , 2003Abdala ( , 2007 and Quinteros (2012Quinteros ( , 2013. Terminology follows Smith (1946) for squamation and Frost (1992) for neck folds. ...
Article
We redescribe Liolaemus bibronii and describe three new species of Liolaemus, a genus of lizards distributed across South America. These species belong to the L. alticolor–bibronii group, which are included in the subgenus Liolaemus s.s. Liolaemus bibronii was previously proposed as a species complex, but many populations initially assigned to this complex were described as valid species. The three new species described here were populations denominated under L. bibronii. In order to validate the new species, we apply an integrative approach, including molecular and morphological evidence. Also, we perform phylogenetic analyses applying parsimony and Bayesian inference. The three new species described here show a set of character states that allow them to be distinguished from L. bibronii, from each other and from all other species of Liolaemus. Our phylogenies show that the newly described species are more related to other species than to L. bibronii. With this study, we are closer to solving the taxonomic puzzle that L. bibronii represents.
... A H S I li A (: 'I -"Phylogenellc Relationships In the chiliens~s Group (Iguania: Liolarnidae: Liolaernus): Adding new Characters and Taxa". In this study a new analysis of the chiliensis group (L~olaem~dae) is made a d d~n g twenty four new characters to the previous matr~x (now 83 characters) of Lobo (2001a) and seven species (now 79). The frequency values were updated for more than 50% of species Included previously. ...
... Recientemente, Lobo y Abdala (2002) en el analisis de 35 caracteres osteol6gicos en solamente 2 4 especies del genero encontraron soporte para 6 grupos (lineomaculatus, tenuis, alticolor, nitidus, signifer, montanus) propuestos en el pasado en base a otras fuentes de evidencia, como morfologia externa, enzimas y ADN. Continuando el estudio morfologico iniciado en el trabajo anterior con el grupo chiliensis (Lobo, 2001a), fue posible aumentar la muestra de especimenes para mas de la mitad de las especies y sumar nuevos caracteres y nuevas especies a la matriz original. Los objetivos principales de este trabajo son presentar la descrip-ci6n de nuevos caracteres, la re-definici6n y re-codificaci6n de otros y presentar las hltimas hip6tesis obtenidas (en este caso aplicando tambien pesos implicados) . ...
... Se sumaron 24 nuevos caracteres a la matriz original (caracteres 8-12, 21, 27, 29, 49-59, 61, 67, 71, 73 y 76). En com-paraci6n con el estudio previo (Lobo, 2001a) se ampli6 la muestra de especimenes para aproximadamente el 60% de las especies y se sumaron siete especies: L. heliodermis. L. dicktracyi, L. urnbrqer, L. petrophilus, L. ceii, L. sp. 1 y L. sp. 3. Las especies aqui llamadas L. sp. ...
Article
Full-text available
Phylogenetic Relationships In the chiliensis Group (Iguania: Liolamidae: Liolaemus): Adding new Characters and Taxa". In this study a new analysis of the chiliensis group (Liolaemidae) made adding twenty four new characters to the previous matrix (now 83 characters) of Lobo (2001) and seven species (now 79). The frequency values were updated for more than 50% of species Included previously. Analyses were made considering "equally weighted characters" criteria and also using the implied weights of Goloboff (1993). A total of six analyses were made, following the "equally weighted characters" criteria and the weighting scheme (five runs, changing each time the value of the weighting c nstant K, from K : 2 through K= 6). In general, results of the present study are in agreement with those obtained in a previous study In this study the nigromaculatus group is monophylethic while the elongatus and monticola groups are paraphylethic; the following monophylethic groupings are recognized: alticolor, altissimus, capillitas (including capillitas and three recently described new species), gravenhorsti, kriegi, leopardinus, nigromaculatus, nigroviridis, pictus, robertmertensi including chilienss, nitidus and robertmertensi) and tenuis.
... We included 45 terminal taxa: 41 species of Liolaemus representing our in-group, and three species of Phymaturus and Ctenoblepharys adspersa used as the out-group (see Supplementary Tables S1 and S2). 1 In addition, we optimized the characters in the most recent phylogenetic topology within the family. Because there is no phylogeny for Liolaemus that includes all terminal taxa analyzed here, we reconstructed a metatree following Schulte et al. (2000), Espinoza et al. (2004), and Pyron et al. (2013) for the entire genus, Avila et al. (2006) and Abdala (2007) for the L. boulengeri series, and Lobo (2001Lobo ( , 2005, Avila et al. (2015), and Quinteros (2013) for the Liolaemus subgenus and the groups within. ...
... Our topology recovered the L. elongatus-petrophilus group as paraphyletic. The location of L. coeruleus as basal to the L. elongatus-petrophilus species plus the L. alticolor-bibronii group species is inconsistent with other proposed phylogenies (Schulte et al. 2000;Lobo 2001Lobo , 2005Lobo and Abdala 2002;Morando et al. 2007;Schulte 2013;Pyron et al. 2013). The monophyly of the L. alticolor-bibronii group recovered here is consistent with previous studies (Lobo 2001(Lobo , 2005Morando et al. 2007;Pyron et al. 2013;Quinteros 2013;Schulte 2013). ...
... The location of L. coeruleus as basal to the L. elongatus-petrophilus species plus the L. alticolor-bibronii group species is inconsistent with other proposed phylogenies (Schulte et al. 2000;Lobo 2001Lobo , 2005Lobo and Abdala 2002;Morando et al. 2007;Schulte 2013;Pyron et al. 2013). The monophyly of the L. alticolor-bibronii group recovered here is consistent with previous studies (Lobo 2001(Lobo , 2005Morando et al. 2007;Pyron et al. 2013;Quinteros 2013;Schulte 2013). ...
Article
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The structure of copulatory organs is widely used in systematics for both differentiating species and for studying phylogenetic relationships. We describe the hemipenes of 42 species belonging to the genus Liolaemus, representing most of their internal groups. We reported 42 characters, the majority not published previously. We constructed a metatree based on previously proposed phylogenetic studies and optimized the hemipenial characters in this topology. Among the most informative characters are presence or absence of flounces or calyces on the sulcate face, ornamentation of the apex, presence or absence of an asulcate face prominence, and presence of a thickening on the proximal region of the asulcate face. Furthermore, we performed a phylogenetic analysis exclusively with the hemipenial characters, not with the intention of making a phylogeny based on this single set of characters, but rather to demonstrate their significance for the reconstruction of relationships in Liolaemus. The obtained results show that the main clades are recovered. We also compared the hemipenial morphology between closely related species to evaluate its taxonomic importance. We conclude that in Liolaemus, the hemipenes can be used both for the differentiation of species and to provide additional evidence for establishing their phylogenetic relationships.
... ALMOST half of the species of Liolaemus are in the chiliensis group (Lobo, 2001), including the small and slender lizard Liolaemus gracilis (Bell, 1843). This species was cited in one of the categorizations of the reptile habitat associations of Buenos Aires province as the only Liolaemus species inhabiting the rocky grounds of the Tandilia System (Gallardo, 1977). ...
... This finding is not surprising because similar differences have been found for other groups (i.e., alticolor group, Espinoza, 1999, 2004;Martínez Oliver and Lobo, 2002), which in the same manner as this case but unlike other groups of Liolaemus, may have recently been isolated and differentiated. Characters of the general pattern provide important information about phylogenetic affinities and have been used recently in a morphology-based phylogenetic analysis of the chiliensis group (Lobo, 2001(Lobo, , 2005. Liolaemus tandiliensis shares with L. gracilis, L. saxatilis, L. sanjuanensis, L. robertmertensi, L. chiliensis and L. nitidus the extreme reduction of lateral neck folds, and longitudinal and antegular folds are absent in these species. ...
... The lateral surface of the neck is formed by large, lanceolate, keeled, and imbricate scales (equal or nearly equal to the size of the dorsal scales at this level). In adittion, L. tandiliensis has a differentiated auricular scale and two or three enlarged scales on the anterior margin of ear, as do the chilean species L. lemniscatus and L. fuscus (Lobo, 2001(Lobo, , 2005. Morphologically, these species are similar to small forms of the alticolor group Espinoza, 1999, 2004), although relationships between these species and that group are not yet very clear. ...
... The genus Liolaemus Wiegmann, 1834 comprises 283 species (Abdala et al., 2021) distributed in Argentina, southern Brazil, Bolivia, Chile, Paraguay, Peru, and Uruguay. Several taxonomic and phylogenetic studies have divided the genus into two subgenera: Eulaemus Girard, 1858 and Liolaemus sensu stricto (Girard, 1858;Laurent, 1983;Etheridge, 1995;Schulte et al., 2000;Lobo, 2001;Abdala and Quinteros, 2014). Recently, Esquerré et al. (2019;2022) recovered some species of the L. walkeri group as sister clade of both subgenera, but this hypothesis requires more study. ...
... Recently, Esquerré et al. (2019;2022) recovered some species of the L. walkeri group as sister clade of both subgenera, but this hypothesis requires more study. Within these subgenera, numerous groups and subgroups have been proposed (e.g., Cei, 1986;1993;Lobo, 2001;2005;Ávila et al., 2006;Abdala, 2007;Abdala and Juárez Heredia, 2013;Fontanella et al., 2012;Quinteros, 2012;Bell, 1843, L. koslowskyi Etheridge, 1993, L. ornatus Koslowsky, 1898, and L. fitzingerii Duméril and Bibron, 1837, while others are endemic, e.g., L. abaucan Etheridge, 1993 Lobo and Kretzschmar, 1996, L. tehuelche Abdala, 2003, and L. telsen Cei and Scolaro, 2003 (for details see Andrade Díaz et al., 2017). Also, species that were once considered widely distributed, such as L. darwinii and L. boulengeri Koslowsky, 1898, are species complexes (Etheridge, 1993Abdala, 2003;2005;Morando et al., 2004;Abdala et al., 2011;Camargo et al., 2012a, b;, and the distribution of the composite species might be more restricted. ...
Article
The Liolaemus boulengeri group is part of the subgenus Eulaemus, genus Liolaemus. This group is widely distributed in Argentina, Bolivia, Chile, and Paraguay, as well as in the Peruvian Titicaca basin and the coasts of Brazil and Uruguay. Here, we combined the revision of a fossil record of Liolaemus, dated at 20 million years (Myr), with relaxed molecular clock analysis to provide a time-calibrated, molecular-based phylogenetic hypothesis including 90% of the group’s known species. We found the Liolaemus boulengeri group (= L. boulengeri section) formed by three main groups, the L. wiegmannii, L. anomalus, and L. darwinii-melanops groups. We performed biogeographic analyses applying Bayesian Binary (BBM), Dispersion-Extinction-Cladogenesis (DEC), and Statistical-Dispersion-Vicariance (S-DIVA) and found that the ancestral area of the L. boulengeri group was likely located in central-west Argentina and reached its current distribution after a series of dispersal and vicariance events. These processes may have been favored by a period of climatic stasis which occurred at the beginning of the group’s diversification, around 41 Myr. The congruence of the results of all three biogeographic analyses evidences new hypothetical historical distributions and events which led to the current species distribution of the L. boulengeri group.
... Very few of systematic studies on Liolaemus incorporate morphological evidence (Lobo, 2001(Lobo, , 2005Abdala, 2005Abdala, , 2007Lobo et al., 2010;Quinteros, 2013), and even less studies incorporate genital morphology (Quipildor et al., 2018a(Quipildor et al., , 2018bRuiz et al., 2019). In the present study, our main aim is to describe a new species of the L. ornatus group, that we recognize as distinct from other members based on morphological characters (scalation, hemipenes, colour pattern), and DNA sequences. ...
... Field studies did not involve endangered species. We studied the morphological characters commonly included in Liolaemus taxonomic studies, such as those described in Laurent (1985), Frost (1992), Etheridge (1993Etheridge ( , 1995Etheridge ( , 2000, Cei (1986), Lobo and Espinoza (1999), Lobo (2001Lobo ( , 2005 Abdala (2007), Quinteros (2013) and Quipildor et al. (2018a), which altogether conform a record of more than two hundred morphological characters including scale counts, shape, ornamentation, imbrications, precloacal pores, neck foldings, colour pattern, life colours, etc. The description of colours in life was made based on photographs made on the field immediately after the capture of the individuals. ...
Article
Andean orogeny is one of the main factors that promoted the diversification of many groups of animals. Among them are the lizards of the Liolaemidae family and. within this, Liolaemus is the genus with the most described species up to date, with approximately 277 species. One of the groups within the genus is the Liolaemus ornatus group, distributed in the south of Central Andes mountain chain. In this work, we describe a new species belonging to the L. ornatus group, using an integrative approach, analyzing both morphological (lepidosis, morphometric, coloration and hemipenes characters) and molecular evidence. We made a divergence analysis of the representatives of the L. ornatus group. Furthermore, we correlated the ages of the geological events with the distribution areas for the L. ornatus group. This newly described species shows character states (morphological and molecular) that allow its clear distinction from the other members of the L. ornatus group, as well as from the rest of Liolaemus. The divergence analysis places the new species in the L. ornatus group and allows us to infer the successive geological events that served as physical barriers for the diversification of the group, besides explaining the current distribution of the group.
... An example of these issues occurs in species related to Liolaemus robertmertensi, a species belonging traditionally to a like-named species group of the subgenus Liolaemus. Previous phylogenies included species such as L. robertmertensi as L. chiliensis, L. nitidus, and L. sanjuanensis in this group (Cei, 1993, Lobo, 2001Lobo et al, 2010;Abdala and Quinteros, 2014). Quinteros (2013) performed a morphological phylogeny of the L. alticolor-bibronii group and recovered the L. robertmertensi group nested within the more inclusive group, formed by L. nitidus, L. robertmertensi, and L. saxatilis. ...
... We analyzed the morphological characters traditionally used in Liolaemus taxonomy including those from Laurent (1985), Etheridge (1993Etheridge ( , 1995Etheridge ( , 2000, Cei (1986Cei ( , 1993, Lobo (2001Lobo ( , 2005, Abdala (2007) and Quinteros (2012Quinteros ( , 2013. Description of color patterns follow those proposed by Lobo and Espinoza (1999) and Quinteros (2012Quinteros ( , 2013. ...
Article
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We redescribe Liolaemus robertmertensi and describe a new species of Liolaemus, the second most speciose lizard genus. These species belong to the L. robertmertensi group of the subge-nus Liolaemus sensu stricto. In recent years, some newly identified populations were assigned to Liolaemus robertmertensi. We applied an integrative approach using morphological and phylogenetic evidence to determine the relationships among these populations and found one to belong to a new species described here. We performed statistical analyses to differentiate L. robertmertensi and the new species from other species of the alticolor-bibronii group. The new species shows a set of character states that allows it to be distinguished from L. robertmertensi and from all other species of Liolaemus. Despite the description of this new species, the taxo-nomic status of many populations still remain unknown.
... This recent study was based on samples from almost all of the species and candidate species recognized in this group, coupled with extensive geographic sampling. The new taxon we describe here is part of the northernmost group of these lizards, the Liolaemus capillitas clade (Lobo 2001), that is inferred with strong support and includes L. capillitas Hulse 1979, L. heliodermis Espinoza, Lobo & Cruz 2000, L. dicktracyi Espinoza & Lobo 2003, L. umbrifer Espinoza & Lobo 2003, L. talampaya Sites 2004 andL. tulkas Quinteros, Abdala, Díaz Gómez &Scrocchi 2008, a set of species distributed in several mountain ranges far away from the majority of the other species of the Liolaemus elongatuskriegi group, which is mostly distributed in Patagonian and southern Andes ranges. ...
... Specimens examined: For purposes of diagnosing the new species we examined a series of each of the six species currently considered to be members of the Liolaemus capillitas clade (e.g. Lobo 2001;Medina et al. 2017; Appendix 1). Specimens examined, including the new species described herein are deposited in the herpetological collections of the Carnegie Museum of Natural History (CM), Pittsburgh, Pennsylvania, USA; Museum of Vertebrate Zoology (MVZ), University of California, Berkeley, California, USA; Monte L. Bean Life Sciences Museum-Brigham Young University (BYU), Provo, Utah, USA; Kansas University (KU), Lawrence, Kansas, USA, Museo de La Plata, La Plata, Argentina, and mainly on specimens deposited in the collection LJAMM-CNP of the Instituto Patagónico para el Estudio de los Ecosistemas Continentales (IPEEC-CONICET) in Puerto Madryn, Chubut, Argentina. ...
Article
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A new species of the Liolaemus capillitas clade is described. Liolaemus galactostictos sp. nov. differs from other members of its group by a combination of morphological and molecular traits, in particular its black dorsal coloration pattern not found in any other Liolaemus species. Liolaemus galactostictos sp. nov. is only known from its type locality. This new species is found in rocky fields surrounded by grasslands on the top of the Velasco Mountains, a ¨sky island environment¨, in northwestern Argentina. As well as other members of its clade this species seems to be strictly saxicolous, viviparous and feeds on insects.
... Likewise, phylogenies have been proposed for different groups and subgroups of the genus; for example, Liolaemus s.s. (Morando, 2004;Medina et al., 2014), Eulaemus (Fontanella et al., 2012;Olave et al., 2014), the Liolaemus boulengeri group (Avila et al., 2006;Abdala, 2007), the Liolaemus lineomaculatus group (Breitman et al., 2011(Breitman et al., , 2013, the Liolaemus elongatus-kriegi group (Lobo, 2001(Lobo, , 2005, the Liolaemus alticolor-bibronii group (Morando et al., 2007;Quinteros, 2013;Portelli & Quinteros, 2018), the Liolaemus anomalus group (Abdala & Juarez Heredia, 2013) and the Liolaemus archeforus-kingii group (Breitman et al., 2015). The most inclusive phylogenies of Squamata (Pyron et al., 2013;Zheng & Wiens, 2016) or Liolaemus broadly (Schulte et al., 2000;Valladares et al., 2002;Espinoza et al., 2004;Schulte, 2013;Olave et al., 2014) have included some species of the L. montanus Koslowsky, 1898, group, but never > 25% of the members of this group. ...
... But there is also evidence of homoplasious molecular data (Engstrom et al., 2004;Castoe et al., 2009;Jarvis et al., 2014). Nevertheless, the use of morphological character states has remained widespread for the Liolaemidae (Lobo, 2001(Lobo, , 2005Lobo & Quinteros, 2005;Abdala, 2007;Quinteros, 2013;Abdala & Juárez Heredia, 2013). Continuous characters analysed as such were used in many different studies (e.g. ...
Article
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.
... Likewise, phylogenies have been proposed for different groups and subgroups of the genus; for example, Liolaemus s.s. (Morando, 2004;Medina et al., 2014), Eulaemus (Fontanella et al., 2012;Olave et al., 2014), the Liolaemus boulengeri group (Avila et al., 2006;Abdala, 2007), the Liolaemus lineomaculatus group (Breitman et al., 2011(Breitman et al., , 2013, the Liolaemus elongatus-kriegi group (Lobo, 2001(Lobo, , 2005, the Liolaemus alticolor-bibronii group (Morando et al., 2007;Quinteros, 2013;Portelli & Quinteros, 2018), the Liolaemus anomalus group (Abdala & Juarez Heredia, 2013) and the Liolaemus archeforus-kingii group (Breitman et al., 2015). The most inclusive phylogenies of Squamata (Pyron et al., 2013;Zheng & Wiens, 2016) or Liolaemus broadly (Schulte et al., 2000;Valladares et al., 2002;Espinoza et al., 2004;Schulte, 2013;Olave et al., 2014) have included some species of the L. montanus Koslowsky, 1898, group, but never > 25% of the members of this group. ...
... But there is also evidence of homoplasious molecular data (Engstrom et al., 2004;Castoe et al., 2009;Jarvis et al., 2014). Nevertheless, the use of morphological character states has remained widespread for the Liolaemidae (Lobo, 2001(Lobo, , 2005Lobo & Quinteros, 2005;Abdala, 2007;Quinteros, 2013;Abdala & Juárez Heredia, 2013). Continuous characters analysed as such were used in many different studies (e.g. ...
... Likewise, phylogenies have been proposed for different groups and subgroups of the genus; for example, Liolaemus s.s. (Morando, 2004;Medina et al., 2014), Eulaemus (Fontanella et al., 2012;Olave et al., 2014), the Liolaemus boulengeri group (Avila et al., 2006;Abdala, 2007), the Liolaemus lineomaculatus group (Breitman et al., 2011(Breitman et al., , 2013, the Liolaemus elongatus-kriegi group (Lobo, 2001(Lobo, , 2005, the Liolaemus alticolor-bibronii group (Morando et al., 2007;Quinteros, 2013;Portelli & Quinteros, 2018), the Liolaemus anomalus group (Abdala & Juarez Heredia, 2013) and the Liolaemus archeforus-kingii group (Breitman et al., 2015). The most inclusive phylogenies of Squamata (Pyron et al., 2013;Zheng & Wiens, 2016) or Liolaemus broadly (Schulte et al., 2000;Valladares et al., 2002;Espinoza et al., 2004;Schulte, 2013;Olave et al., 2014) have included some species of the L. montanus Koslowsky, 1898, group, but never > 25% of the members of this group. ...
... But there is also evidence of homoplasious molecular data (Engstrom et al., 2004;Castoe et al., 2009;Jarvis et al., 2014). Nevertheless, the use of morphological character states has remained widespread for the Liolaemidae (Lobo, 2001(Lobo, , 2005Lobo & Quinteros, 2005;Abdala, 2007;Quinteros, 2013;Abdala & Juárez Heredia, 2013). Continuous characters analysed as such were used in many different studies (e.g. ...
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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.
... Relationships of Liolaemus nitidus have long been difficult to establish (Pincheira-Donoso & Núñez, 2005). Most recent morphological studies agree on its affinities to L. robertmertensi and L. chiliensis (Lobo, 2001(Lobo, , 2005Pincheira-Donoso & Núñez, 2005;Quinteros, 2013). However, molecular analyses indicate completely different phylogenetic position, particularly its close relationship to L. monticola (Troncoso-Palacios et al., 2015a;Panzera et al., 2017;Torres-Pérez et al., 2017). ...
... The fact that precloacal pores were not observed in any of the specimens suggests that all of these individuals are females. However, members of the subgenus Liolaemus (a group to which all rediscovered lizards belong) tend to have significantly fewer precloacal pores than do members of Eulaemus (Laurent, 1992) and there are several Liolaemus species in which both females and males completely lack these pores (Lobo, 2001;Pincheira-Donoso & Scolaro, 2007;Troncoso-Palacios et al., 2015b). Also, the 'taphonomical' factor must be taken into consideration, that is, all the processes acting on a specimen after it was collected. ...
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Johann Ludwig Christian Gravenhorst’s herpetological collection at the Museum of Natural History, University of Wrocław included numerous important specimens of amphibians and reptiles. The majority, if not the entirety, of this collection has long been thought to be lost. However, we were able to rediscover some type specimens of lizards. The rediscovered specimens include the holotypes of Liolaemus conspersus and L. hieroglyphicus, one syntype of Callopistes maculatus (here designated as the lectotype) and two syntypes of L. lineatus (one of which is herein designated as the lectotype). Reexamination of these specimens indicates that previous synonymies proposed for L. conspersus and two syntypes of L. hieroglyphicus are problematic; furthermore, more complex taxonomic work is needed to resolve this issue. Two rediscovered syntypes of L. lineatus differ in several scalation traits and are possibly not conspecific. The type specimens of several other species of lizards from Gravenhorst’s collection (Liolaemus marmoratus, L. unicolor and two other syntypes of L. lineatus, Leiocephalus schreibersii and Chalcides viridanus) were not found and are probably lost.
... The Liolaemus elongatus -kriegi complex as defined by Cei (1979) on the basis of several diagnostic morphological characters includes widespread polytypic species for which species recognition rests on an inadequate taxonomy . Different taxonomic groupings for the species included in this complex have been proposed by Cei (1974Cei ( , 1979, but also see Cei 1975Cei , 1986Cei , 1993, Ortiz (1981), Etheridge (1995), Espinoza et al. (2000), Schulte et al. (2000), Lobo (2001Lobo ( , 2005, Espinoza and Lobo (2003), Pincheira Donoso and Nuñez (2005), and Scolaro and Pincheira Donoso (2007). Morando et al. (2003) used the name Liolaemus elongatus-kriegi complex to be consistent with the original hypothesis of group content by Cei (1979), but resolved three well-supported species groups on the basis of mtDNA sequences. ...
... Also, in Morando (2004) in a phylogenetic tree including many species from the chiliensis group and several genes, L. neuquensis was not recovered as part of the elongatus clade. Other species such as L. leopardinus, L. ramonensis, L. valdesianus, L. curis, L.cristiani, as well as some "subspecies" of the L. monticola clade, may also be related to the Liolaemus elongatus clade (Lobo, 2001, Torres Perez et al. 2009), here we find L. chillanensis recovered within the elongatus clade, but a test of this hypothesis must await a more extensive study. In a recent publication, Lobo et al. (2010) includes as part of the elongatus clade, the following species: austromendocinus, elongatus, flavipiceus, gununakuna, parvus, petrophilus, punmahuida, thermarum, tregenzai, and all other species previously suggested as related with the elongatus clade are included in new created groups (leopardinus, chillanensis, monticola and kriegi) of the subgenus Liolaemus. ...
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A new species of lizard of the genus Liolaemus from Cordillera del Viento, northwestern Neuquén Province, Argentina is described. The new species is a member of the Andean-Patagonian Liolaemus elongatus clade and is known only from a single locality above 3000 m in the eastern slopes of Domuyo Volcano. Liolaemus antumalguen sp. nov. is a stout species, saxicolous, probably viviparous and omnivorous, and seems to have low population density. It is easily differentiated from all other species of the elongatus complex for a highly variable but characteristic dorsal pattern and a completely black ventral coloration.
... Whitin Liolaemus, several phylogenetic hypotheses have been proposed (Laurent 1985;Etheridge 1995;Schulte et al. 2000;Lobo 2001Lobo , 2005Morando et al. 2004;Avila et al. 2006;Abdala 2007). The Liolaemus boulengeri group of species was characterized by the presence of a patch of enlarged scales on the posterior medial surface of the thigh (Etheridge 1995) and by the hypertrophy of the puboischiotibialis muscle . ...
... The characters considered were those currently studied in Liolaemus, which were described or cited mainly by Laurent (1985), Etheridge (1993Etheridge ( , 1995Etheridge ( , 2000, Cei (1986), Lobo (2001), and Abdala (2002Abdala ( , 2003Abdala ( , 2005. Neck fold terminology follows Frost (1992). ...
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The genus Liolaemus is a widely distributed and very diverse natural group of lizards. The L. boulengeri group is characterized by the presence of a patch of enlarged scales on the posterior medial surface of the thigh. Liolaemus goetschi, which belongs to this group, was described by Müller and Hellmich in 1938 based on few specimens collected from Laguna Playa (Río Negro Province, Argentina). Those specimens were deposited in museums in Munich (Germany) and Breslau (Poland). The type material of L. goetschi deposited in Poland was lost and because the material in Germany is of difficult access, many populatios of other species have been confused with L. goetschi in subsequent works. In this work, L. goetschi was redescribed by using the characters currently designated for the group and the known distribution of the species was extended. The taxonomic status and phylogenetic relationships of L. goetschi within the L. boulengeri clade was also analyzed, especially with respect to L. melanops and L. martorii.
... Liolaemus comprises 255 species distributed in southern South America, from Tierra del Fuego (south- ernmost distribution) in Argentina to La Libertad (north- ernmost distribution) in northern Peru. The L. alticolor- bibronii group is a member of the Liolaemus sensu stricto subgenus (Ortiz, 1981;Lobo, 2001Lobo, , 2005Espinoza et al., 2004;Lobo et al., 2010;Martinez et al., 2011;Quinteros, 2012Quinteros, , 2013. Study of the taxonomy of the L. alticolor- bibronii group began with Ortiz (1981) and Cei (1986), and the group has grown to include 26 described species, including the new species described herein (Quinteros, 2013). ...
... A small (snout-vent length [SVL] <50 mm), slen- der-bodied and long-tailed Liolaemus belonging to the subgenus Liolaemus (sensu Laurent, 1985;Schulte et al., 2000;Lobo et al., 2010), with which it shares the follow- ing synapomorphies: supralabials narrow, width equal to or less than that of lorilabials, usually four, the pos- terior one elongate and usually upturned posteriorly (Etheridge, 1995;Lobo, 2001Lobo, , 2005. Within this subge- nus, L. chungara belongs to the L. alticolor-bibronii group (sensu Lobo et al., 2010;Quinteros, 2012Quinteros, , 2013, with which it shares the following characteristics: small body size (SVL < 60 mm), a distinct dorsal color pattern formed by dorsolateral stripes, paravertebral spots, vertebral and ventrolateral lines, and a fine gray to black markings on the ventral surface of the tail (Lobo, 2005, Quinteros, 2012). ...
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We describe a new slender species of Liolaemus of the L. alticolor-bibronii group of the subgenus Liolaemus. The new species is phe-netically and biogeographically close to L. alticolor, L. paulinae, and L. puna but presents a combination of character states that differentiates it from all other species of Liolaemus. The new taxon is the first species of subgenus Liolaemus sensu stricto recorded as having supernumerary pores. The new species inhabits places where Parastrephia lucida is the predominant flora and is distributed in areas close to Putre, in the Arica y Parinacota region, northern Chile. Data on its biology, complete distribution, and conservation status are lacking. Resumen. Describimos una nueva especie de Liolaemus, perteneciente al grupo de L. alticolor-bibronii del subgénero Liolaemus sensu stricto. La nueva especie es fenética y biogeográficamente cercana a L. alticolor, L. paulinae y L. puna. Pero muestra una combinación de estados de carácter que permiten su diferenciación con estas y con todos los demás Liolaemus. El nuevo taxon es la única especie del subgénero Liolaemus sensu stricto que exhibe poros supernumerarios. La nueva especie habita en regiones donde Parastrephia lucida es la planta predomiante y está distribuida en áreas cercanas a la localidad de Putre, en la Región de Arica y Parinacota, en el norte de Chile. No existen datos sobre su biología, distribución completa ni de su estado de conservación.
... The phylogenetic analyses included typical morphological characters (Lobo, 2001;Lobo, 2005;Quinteros, 2012;Quinteros, 2013) and sequences of Cytb and 12S genes taken from the literature (Espinoza, Wiens & Tracy, 2004;Morando et al., 2007;Victoriano et al., 2008;Aguilar et al., 2013;Troncoso-Palacios et al., 2016;Olave et al., 2011; see File S1 for specimens' GenBank accession numbers). Sequences were aligned and edited with MEGA v.7.0.26 (Kumar, Stecher & Tamura, 2016). ...
... See Fig. 5 (Quinteros, 2013) who detected the L. lemniscatus and the L. robertmertensi groups nested inside the L. alticolor-bibronii group (Parsimony tree, Fig. 3). Previous studies found both groups outside of the L. alticolor-bibronii group (Cei, 1986;Cei, 1993;Lobo, 2001;Lobo, 2005;Díaz Gómez & Lobo, 2006). Schulte et al. (2000), Schulte (2013), Pyron, Burbrink & Wiens (2013), while Zheng & Wiens (2016) found L. robertmertensi settled inside the L. alticolor-bibronii group, but recovered the L. lemniscatus group outside. ...
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The genus Liolaemus comprises more than 260 species and can be divided in two subgenera: Eulaemus and Liolaemus sensu stricto . In this paper, we present a phylogenetic analysis, divergence times, and ancestral distribution ranges of the Liolaemus alticolor-bibronii group ( Liolaemus sensu stricto subgenus). We inferred a total evidence phylogeny combining molecular ( Cytb and 12S genes) and morphological characters using Maximum Parsimony and Bayesian Inference. Divergence times were calculated using Bayesian MCMC with an uncorrelated lognormal distributed relaxed clock, calibrated with a fossil record. Ancestral ranges were estimated using the Dispersal-Extinction-Cladogenesis (DEC-Lagrange). Effects of some a priori parameters of DEC were also tested. Distribution ranged from central Perú to southern Argentina, including areas at sea level up to the high Andes. The L. alticolor-bibronii group was recovered as monophyletic, formed by two clades: L. walkeri and L. gracilis , the latter can be split in two groups. Additionally, many species candidates were recognized. We estimate that the L. alticolor-bibronii group diversified 14.5 Myr ago, during the Middle Miocene. Our results suggest that the ancestor of the Liolaemus alticolor-bibronii group was distributed in a wide area including Patagonia and Puna highlands. The speciation pattern follows the South-North Diversification Hypothesis, following the Andean uplift.
... In the last 30 years, several authors (Lobo, 2001(Lobo, , 2005Lobo et al., 2010;Nuñez et al., 1991;Videla and Cei, 1996) have suggested further species additions to the L. elongatus-kriegi group, such as the L. cristiani clade (Esquerré et al., 2013). Most of its component species were described based on morphological characters (Abdala et al., 2010;Cei, 1974;Donoso Barros and Cei, 1971;Espinoza and Lobo, 2003;Espinoza et al., 2000;Esquerré et al., 2013;Hulse, 1979;Koslowsky, 1896;Müller and Hellmich, 1939;Nuñez et al., 1991;Pincheira-Donoso and Scolaro, 2007;Quinteros et al., 2008;Werner, 1907), and a few of these have been corroborated by molecular (mostly mtDNA) characters (Avila et al., , 2010Escobar Huerta et al., 2015;Torres-Pérez et al., 2009;Troncoso-Palacios et al., 2015). ...
... The L. elongatus and L. petrophilus complexes were inferred as strongly supported clades in the mitochondrial gene tree (Appendix D) and in the SVDquartet species tree (Fig. 4), and the L. kriegi complex was inferred as monophyletic in all of our analyses, in agreement with other authors (Lobo, 2001(Lobo, , 2005Lobo et al., 2010;Pyron et al., 2013;Schulte II et al., 2000). Support for a monophyletic L. kriegi complex was high to moderate with the four approaches we used (Appendix D, E and Figs. 2 and 3), and included four described (L. ...
Article
We present different approaches to a multi-locus phylogeny for the Liolaemus elongatus-kriegi group, including almost all species and recognized lineages. We sequenced two mitochondrial and five nuclear gene regions for 123 individuals from 35 taxa, and compared relationships resolved from concatenated and species tree methods. The L. elongatus-kriegi group was inferred as monophyletic in three of the five analyses (concatenated mitochondrial, concatenated mitochondrial + nuclear gene trees, and SVD quartet species tree). The mitochondrial gene tree resolved four haploclades, three corresponding to the previously recognized complexes: L. elongatus, L. kriegi and L. petrophilus complexes, and the L. punmahuida group. The BEAST species tree approach included the L. punmahuida group within the L. kriegi complex, but the SVD quartet method placed it as sister to the L. elongatus-kriegi group. BEAST inferred species of the L. elongatus and L. petrophilus complexes as one clade, while SVDquartet inferred these two complexes as monophyletic (although with no statistical support for the L. petrophilus complex). The species tree approach also included the L. punmahuida group as part of the L. elongatus-kriegi group. Our study provides detailed multilocus phylogenetic hypotheses for the L. elongatus-kriegi group, and we discuss possible reasons for differences in the concatenation and species tree methods.
... In the case of smaller structures, the measurements were taken using a micrometer eyepiece. Nomenclature of bones, processes and foramina follow de Queiroz (1982), Keller and Krause (1986), Frost (1992), Etheridge (2000, Lobo and Abdala (2001), Lobo (2001Lobo ( , 2005) and Torres-Carvajal, (2004). In total, 55 characters, 19 continuous and 36 discrete, of the cranial and postcranial skeleton were examined. ...
... Maxillary tooth morphology II: (0) crowns without differentiated cusps, (1) three conspicuous cusps (all species studied here). Species of the L. nigromaculatus group (subgenus Liolaemus sensu stricto) were reported as having broad maxillary teeth without secondary cusps ( Lobo, 2001). Non-polymorphic binary. ...
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Fifty-five skeletal characters (continuous and discrete) were analyzed for species of the L. darwinii group: L. albiceps, L. chacoensis, L. grosseorum, L. irregularis, L. koslowskyi, L. lavillai, L. ornatus, L. quilmes, plus L. inacayali (L. telsen group) and L. scapularis (L. wiegmannii group). We report polymorphic intraspecific variation that has not previously been taken into account and we describe 21 new characters that provide original information across the group. We detected several morphological synapomorphies for the darwinii group and subclades. The enclosure of Meckel’s cartilage by a dentary outgrowth on lingual side of lower jaw (a synapomorphy of the subgenus Liolaemus sensu stricto and of the Phymaturus patagonicus group) also occurs within the L. darwinii group. The morphology of maxillary teeth with three conspicuous cusps may be a potential synapomorphy of the subgenus Eulaemus. The morphology of maxillary teeth may have adaptive value. Characters that were studied in other groups of lizards were informative for Liolaemus.
... Runs were performed using implicit enumeration (Branch and Bound) under equal weights. Considering the most current phylogenies of Squamata (Conrad, 2008;Gauthier et al., 2012;Pyron et al., 2013;Simões & Pyron, 2021;Zheng & Wiens, 2016), Leiosauridae and Liolaemidae (Abdala, 2007;Abdala & Juarez Heredia, 2013;Abdala et al., 2020;Breitman et al., 2013Breitman et al., , 2015Lobo, 2001Lobo, , 2005Lobo et al., 2010;Olave et al., 2014;Portelli & Quinteros, 2018;Quinteros, 2013;Quinteros et al., 2014Quinteros et al., , 2020, a metatree was reconstructed including all the taxa studied in this work. On that tree, we optimized the character states and searched for potential synapomorphies. ...
Article
Few studies considered the anatomy of the nerve plexuses and musculature associated with them in ectothermic sauropsids. Based on differentiated Sudan Black B staining and conventional dissections, we describe the neuroanatomy of the brachial plexus, its main associated nerves, and muscles. For that, representatives of the genera Diplolaemus, Liolaemus, Phymaturus , and Tropidurus were selected. Based on this, potentially useful characters for phylogenetic analysis were described. Our results show that the brachial plexus can be formed by four, five, or six nerve branches. The brachial flexor trunk, circumflex, interosseous, median, radial, subscapulocoracoid, supracoracoid , and ulnar nerves were identified. Regarding the muscles innervated by the main nerves, the following muscles were identified: biceps brachii, deltoideus scapularis, latissimus dorsi, levator scapulae, pectoralis, serratus thoracis, trapezius, triceps longus caudalis , and triceps longus lateralis . Phylogenetic analyzes revealed 31 potential synapomorphies. There exists evidence that neuroanatomy studies in a phylogenetic context could provide useful information helping to elucidate the relationships between taxonomic groups.
... Algunas formas están restringidas a un tipo de hábitat como las especies saxícolas (que viven en rocas), arborícolas, arenícolas; otras en cambio, pueden encontrarse en un amplio rango: las llamadas especies generalistas. Recientemente hubo un creciente interés en estudios con enfoques ecomorfológicos en este género que podría atribuirse a su accesibilidad y al progresivo conocimiento que se fue generando en las últimas décadas, tanto de su morfología (Abdala y Moro 2003), ecología (Schulte et al., 2004Cruz et al., 2009, entre otros) y sobre su filogenia (Laurent, 1983;Schulte et al., 2000;Lobo, 2001Lobo, , 2005Morando et al., 2004, Lobo y Quinteros, 2005Lobo et al., 2007;Ávila et al., 2006;Abdala, 2007;Abdala y Quinteros, 2008;Pyron et al., 2013, entre otros). En el caso particular de nuestro grupo de trabajo, luego de la obtención de caracteres morfológicos de la anatomía interna, externa y uñas de numerosas especies de Liolaemus, y mediante la aplicación de la metodología arriba descrita, hemos abordado una amplia gama de preguntas, de las cuales seleccionamos algunas. ...
... The morphological characters traditionally used in Liolaemus taxonomy were examined in this study, including those of Laurent (1985), Cei (1986Cei ( , 1993, Etheridge (1993Etheridge ( , 1995Etheridge ( , 2000, Lobo (2001), Abdala (2002Abdala ( , 2003Abdala ( , 2007, and Abdala et al. (2019. The terminology of Smith (1946) was followed for descriptions of squamation, and that of Frost (1992) for descriptions of neck-folding. ...
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A new lizard of the genus Liolaemus is described from the Tacna region of southern of Peru. This species belongs to the L. montanus group and was initially thought to be L. poconchilensis and L. insolitus. However, a series of diagnostic characters differentiate it consistently from these two species and all other species of the genus. To determine the taxonomic status of these lizards, their phylogenetic relationships were analyzed, as well as their morphological and ecological characteristics. The results of the analysis support the conclusion that this population of lizards represents a new species to science, and that the new species is related to L. nazca and L. chiribaya. The new species has sexual dimorphism and is known from elevations of ca. 1,000 m above sea level in the hyperarid Pacific deserts, which are populated by scattered Ephedra americana and Poissonia sp. Due to its highly restricted range and observed habitat loss, we recommend this species be categorized as Critically Endangered. Resumen.—Una nueva especie de lagarto del género Liolaemus es descrita para la Región Tacna, sur de Perú. Esta especie pertenece al grupo L. montanus, la que fue inicialmente confundida con L. poconchilensis y L. insolitus. Sin embargo, una serie de caracteres diagnósticos la diferencian consistentemente de estas y otras especies del género. Para determinar su estatus taxonómico, nosotros analizamos sus relaciones filogenéticas, así como sus características morfológicas y ecológicas. Nuestros resultados sustentan la conclusión que esta población es una nueva especie para la Ciencia, e indica que esta nueva especie está relacionada a L. nazca and L. chiribaya. La nueva especie presenta dimorfismo sexual, y es conocida en elevaciones cercanas a los 1,000 m sobre el nivel del mar, en el hiperárido desierto del Pacífico con matorral de Ephedra americana y Poissonia sp. Debido a su distribución restringida y la pérdida de hábitat observada, nosotros proponemos que sea incluida en la lista de especies amenazadas como En Peligro Crítico.
... The morphological characters traditionally used in Liolaemus taxonomy were examined in this study, including those of Laurent (1985), Cei (1986Cei ( , 1993, Etheridge (1993Etheridge ( , 1995Etheridge ( , 2000, Lobo (2001), Abdala (2002Abdala ( , 2003Abdala ( , 2007, and Abdala et al. (2019. The terminology of Smith (1946) was followed for descriptions of squamation, and that of Frost (1992) for descriptions of neck-folding. ...
Article
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A new lizard of the genus Liolaemus is described from the Tacna region of southern of Peru. This species belongs to the L. montanus group was initially thought to be L. poconchilensis and L. insolitus. However, a series of diagnostic characters differentiate it consistently from these two species and all other species of the genus. To determine the taxonomic status of these lizards, their phylogenetic relationships were analyzed, as well as their morphological and ecological characteristics. The results of the analysis support the conclusion that this population of lizards represents a new species to science, and that the new species is related to L. nazca and L. chiribaya. The new species has sexual dimorphism and is known from elevations of ca. 1,000 m above sea level in the hyperarid Pacific deserts, which are populated by scattered Ephedra americana and Poissonia sp. Due to its highly restricted range and observed habitat loss, we recommend this species be categorized as Critically Endangered.
... El avance y desarrollo de los aspectos taxonómicos y filogenéticos en el género Liolaemus, en los últimos años ha sido significativo (Abdala 2003(Abdala , 2005(Abdala , 2007Abdala y Lobo 2006;Abdala y Juárez 2013;Abdala y Quinteros 2008Abdala et al., , 2012Abdala et al., , 2019Aguilar et al., 2017Aguilar et al., , 2018Aguilar et al., , 2019Ávila et al., 2006, 2008, 2013Breitman et al., 2011;Lobo 2001Lobo , 2005Lobo et al., 2010a;Quinteros 2013;Quinteros et al., 2008Quinteros et al., , 2019Schulte et al., 2000;Valladares et al., 2002). Este avance en el conocimiento taxonómico, filogenético, estuvo durante muchos años pausado en el grupo de Liolaemus montanus y recién en los últimos años ha tomado un impulso acentuado (Avila et al., 2013, Gutierrez et al., 2018Aguilar Puntriano et al., 2017Abdala et al., 2019Chaparro et al., 2020;Huamani Valderrama, 2020). ...
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ABSTRACT Due to lack of information about the precise type locality of Liolaemus andinus, and the subsequent loss of the type material, for a long-time various population of divergent morphological characteristics were assigned to this taxon, which caused the identification of the true L. andinus to become problematic. The present study started in 2004 and after sixteen years of research of the various populations assigned or related to this species, L. andinus is redescribed, a neotype is assigned, and considerations on its probable provenance, and data on its biology and distribution are provided. Also, a hypothesis is presented on the provenance of L. andinus based on biological and historical support evidence. The taxonomic position of populations that have been assigned to and confused with L. andinus in the literature is determined, several of which have affinity with L. poecilochromus and L. rosenmanni. Morphology and molecular-based analyses performed in this study allow us to describe two new Liolaemus species, previously confused and identified as Liolaemus andinus. These new taxa inhabit the high Andes phytogeographic region in Argentina, at altitudes above 3000 m a.s.l. One of these species is distributed in the southwest of Catamarca province and the other inhabits the central-western portion of La Rioja province. The most significant morphological differences between these new species and Liolaemus andinus are mainly the coloration pattern and some lepidosis characters related to number of scales. RESUMEN Debido a la falta de información sobre la procedencia exacta de colecta de los ejemplares de la serie tipo Liolaemus andinus y posterior pérdida, durante mucho tiempo se asignaron a esta especie varias poblaciones con características morfológicas dispares, lo que trajo como pro-blema la identificación del verdadero L. andinus. Este estudio comenzó en el año 2004 y luego de dieciséis años de investigación de las poblaciones asignadas o relacionadas a L. andinus, se la redescribe, se designa un neotipo, se considera su probable procedencia y se aportan datos de su biología y distribución. Asimismo, se presenta una hipótesis sobre la procedencia y dis-tribución de L andinus basada en argumentos biológicos e históricos. También se determina la posición taxonómica de las poblaciones asignadas y confundidas con L. andinus, varias de las cuales tienen afinidad con Liolaemus poecilochromus y Liolaemus rosenmanni. Los análisis morfológicos y moleculares desarrollados en este estudio, nos permiten describir dos nuevas especies de Liolaemus, que anteriormente eran identificadas y confundidas con Liolaemus andinus. Estos nuevos taxones habitan en la región fitogeográfica altoandina de Argentina, en altitudes mayores a 3000 m s.n.m; una se distribuye en el suroeste de la provincia de Catamarca y la restante en el centro oeste de la provincia de La Rioja. Las diferencias morfológicas más significativas entre estas nuevas especies y Liolaemus andinus se encuentran principalmente en el patrón de coloración y algunos caracteres de lepidosis relacionados al número de escamas.
... Características Morfologicas. Para realizar la diagnosis y variaciones en la nueva especie se estudiaron los caracteres habitualmente utilizados en Liolaemus, descriptos principalmente por Laurent (1985); Etheridge (1993Etheridge ( , 1995Etheridge ( , 2000; Cei (1986); Lobo (2001); Abdala (2007); Abdala y Juarez Heredia (2013); Paz (2012) y Quinteros (2012). La descripción de los colores en vida se realizó a partir de fotografías tomadas al cap turar los individuos. ...
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The conservation of the biodiversity of the Patagonian steppe and monte has received scarce attention in comparison with other biomes. In order to advance specific conservation actions, it is important to provide a severe basis for the biodiversity of the area. For this, it is essential to solve multiple taxonomic challenges, including the description of new species. This taxo-nomic resolution is crucial for categorizing the conservation status of the taxa studied. In this study, a new Patagonian species of the genus Liolaemus is formally described and categorized. The species was found in arid, hostile areas in the phytogeographic region of the Monte, in Río Negro province, Argentina. Characters of external morphology, morphometry, color pattern and folidosis were employed to describe the new species. In addition, a phylogenetic hypothesis based on morphological characters is presented, recovering a clade formed by the new Liolaemus species, L. donosobarrosi and L. tirantii as a monophyletic group, maintaining the same relationships in most of the analysis performed. The conservation category assigned to the new species was based on the modified SUMIN index, used by Asociación Herpetológica Argentina, resulting in the vulnerable species category. The population of this new species is very scarce and despite having conducted a thorough search for years, the number of registered specimens is extremely low. RESUMEN La conservación de la biodiversidad del monte y la estepa Patagónica ha recibido escasa atención en comparación con otros biomas. Para poder avanzar en acciones específicas de conservación es importante proporcionar una base sólida acerca de la biodiversidad de la zona. Para esto, es fundamental resolver múltiples desafíos taxonómicos, entre los cuales se incluye la descripción de nuevas especies. Esta resolución taxonómica es crucial para poder categorizar el estado de conservación de los taxones estudiados. En este trabajo se describe y categoriza formalmente una nueva especie del género Liolaemus. La misma fue hallada en áreas áridas y hostiles de la región fitogeográfica del Monte, en la Provincia de Río Negro, Argentina. Para su descripción, se utilizaron caracteres de morfología externa, morfometría, patrón de coloración y folidosis. Además, se presenta una hipótesis filogenética basada en caracteres morfológicos, que recu-pera un clado conformado por la nueva especie, L. donosobarrosi y L. tirantii como grupo monofilético, manteniendo las mismas relaciones de parentesco en la mayoría de los análisis realizados. La categoría de conservación de la nueva especie se hizo en base al índice SUMIN modificado, utilizado por la Asociación Herpetológica Argentina, dando como resultado la categoría de especie Vulnerable. La población de esta nueva especie es muy escasa y a pesar de haber realizado una búsqueda exhaustiva durante años, el número de ejemplares registrados es extremadamente bajo.
... Later, Schulte et al. (2000) performed a molecular based phylogeny of Liolaemus, where they arrive at several general conclusions on the genus, for example that subsequent events of dispersals across the Andes mountain range followed by vicariances have shaped the genus over its history. Lobo (2001), in a phylogeny of the L. chiliensis group assigned to the species studied the distribution areas defined by Roig-Juñent (1994), found that major groups correspond to these areas, although in that analysis, the L. elongatus group is not recovered as monophyletic. Díaz Gómez & Lobo (2006) were the first to perform formal biogeographical analyses in which an ancestral area is assigned to the L. elongatus group, although the focus of this work is more general, that is, the L. sensu stricto group. ...
Article
Published by the British Herpetological Society The genus Liolaemus includes 268 species, classified in two subgenera, Eulaemus and Liolaemus sensu stricto. The latter is formed by 12 monophyletic groups; one of them being the Liolaemus elongatus group, distributed in South America. We studied the biogeographic history of the L. elongatus group. We obtained a phylogenetic hypothesis recovering five main clades: the L. punmahuida, L. elongatus sensu stricto, L. kriegi, L. petrophilus and L. capillitas clades. Based on that hypothesis we obtained a time calibrated tree. The ancestral ranges were estimated applying three methodologies: DEC, DEC+j (using predefined areas) and GEM (using explicit geographical data). Our results show that the ancestral area of the L. elongatus group was located in central Argentina, and its divergence began around 11.5 Mya. From here, a combination of events (founder events and/or vicariances) led the species to their current distribution. Despite their differences, DEC+j and GEM show congruent results.
... Later, Schulte et al. (2000) performed a molecular based phylogeny of Liolaemus, where they arrive at several general conclusions on the genus, for example that subsequent events of dispersals across the Andes mountain range followed by vicariances have shaped the genus over its history. Lobo (2001), in a phylogeny of the L. chiliensis group assigned to the species studied the distribution areas defined by Roig-Juñent (1994), found that major groups correspond to these areas, although in that analysis, the L. elongatus group is not recovered as monophyletic. Díaz Gómez & Lobo (2006) were the first to perform formal biogeographical analyses in which an ancestral area is assigned to the L. elongatus group, although the focus of this work is more general, that is, the L. sensu stricto group. ...
Article
The genus Liolaemus includes 268 species, classified in two subgenera, Eulaemus and Liolaemus sensu stricto. The latter is formed by 12 monophyletic groups; one of them being the Liolaemus elongatus group, distributed in South America. We studied the biogeographic history of the L. elongatus group. We obtained a phylogenetic hypothesis recovering five main clades: the L. punmahuida, L. elongatus sensu stricto, L. kriegi, L. petrophilus and L. capillitas clades. Based on that hypothesis we obtained a time calibrated tree. The ancestral ranges were estimated applying three methodologies: DEC, DEC+j (using predefined areas) and GEM (using explicit geographical data). Our results show that the ancestral area of the L. elongatus group was located in central Argentina, and its divergence began around 11.5 Mya. From here, a combination of events (founder events and/or vicariances) led the species to their current distribution. Despite their differences, DEC+j and GEM show congruent results.
... Several species groups have been recently described based on molecular or morphological characters (e.g. Schulte et al., 2000;Morando 2004;Lobo 2001Lobo , 2005Avila et al. 2006;Abdala 2007), and among these the wiegmannii group has been recognized as monophyletic based on morphological, behavioral, and molecular studies (Etheridge 1995(Etheridge , 2000Halloy et al. 1998;Schulte et al. 2000, Avila et al. 2006. Etheridge (1995Etheridge ( , 2000 morphologically characterized the group by the presence of two or more rows of lorilabial scales rather than one, which are smaller than the supralabial scales; flat or concave rather than convex infralabials; mental scale narrower anteriorly than posteriorly; and six scales in contact with mental. ...
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The wiegmannii group of Liolaemus includes L. arambarensis, L. azarai, L. lutzae, L. occipitalis, L. multimaculatus, L. rabinoi, L. riojanus, L. salinicola, L. scapularis, L. wiegmannii, and the species described here. We used sequences of the mitochondrial cyt–b, 12S, and ND4, and the nuclear C–mos gene regions to infer the phylogeny of the majority of the species of the wiegmannii group. We describe a new species that is closely related to L. multimaculatus and L. riojanus, but can be distinguished by a different dorsal coloration, absence of suprascapular spots, and smaller size. Liolaemus cuyumhue sp. nov. lives in marked geographic isolation with respect to other closely-related species of the wiegmannii group.
... Diagnosis. A large Liolaemus species belonging to the chiliensis clade diagnosed by Etheridge (1995), because it has a low number (4-4) of narrow supralabial scales, the last one elongated and upturned in the posterior margin, without a bladelike process on the posterior distal tibia, without femoral patch of enlarged scales, without tridentate dorsal scales, and with Meckel's groove fused (see also Laurent 1992;Schulte et al. 2000;Lobo 2001;Pincheira-Donoso & Núñez 2005). All these traits differentiate L. tregenzai from L. periglacialis, L. kolengh, L. lineomaculatus and L. silvanae, which exhibit the diagnostic traits detailed for the lineomaculatus clade (see Donoso-Barros & Cei 1971;Cei 1986;Etheridge 1995). ...
Article
Most Liolaemus lizard species are characterized by the presence of precloacal glands in males. Only a few taxa lack these sexual signal emitter structures. Phylogenetic evidence suggests that those species are restricted to the clades chiliensis and lineomaculatus. Within the first lineage, L. coeruleus, L. cristiani, L. flavipiceus, L. neuquensis and L. thermarum lack precloacal glands, which have been considered as members of the neuquensis group. Whereas, in the second one, L. periglacialis (= L. hatcheri), L. kolengh, L. lineomaculatus, and L. silvanae exhibit this characteristic. In the present study we provide the description of Liolaemus tregenzai, an additional new species lacking precloacal glands in both sexes. This new taxon, member of the chiliensis clade, is so far known from the Copahue Volcano, Neuquén Province of Argentina, in boreal Patagonia. Liolaemus tregenzai differs from the remaining species of this clade in having a unique combination of morphologic and chromatic traits, such as a large body size, olive or chest-nut dorsal ground colour, with dark brown or blackish pigment on the flanks, green-bluish with intense black pigment on the ventral surface, and evident sexual dichromatism. The ecology of this new lizard is also remarkably, occurring in Andean antarctandic forests, and being common near thawing snows. Phylogenetic relationships of this species with other members of the chiliensis clade and with the taxa recognized as members of the neuquensis group are still unknown.
... groups alticolor, elongatus-kriegi, boulengeri), and among series of species (e.g. Avila et al. 2004;Espinoza et al. 2004;Morando et al. 2004;Cruz et al. 2005;Abdala 2007;Pincheira-Donoso et al. 2007a; see also Lobo 2001;Díaz & Lobo 2006, for morphological based phylogenies). ...
Article
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Iguanian lizards comprise two of the most species-rich vertebrate genera on Earth (Anolis and Liolaemus). Therefore, studies with the aim of understanding their diversity and phylogenetic relationships may have major significance for ecological and evolutionary research. However, difficulties are often associated with these diverse groups. For example, adaptive radiations may lead to the evolution of conspicuous patterns of intraspecific (interpopulational) variation in response to local environmental conditions, in the absence of real speciation events. This can lead to the taxonomic recognition of new species in the absence of true reproductive isolation. In addition, although diverse taxa are appropriate models to evaluate comparatively the effects of selection on ecological and life-history traits, it is often a major challenge to gather all the available information on the distribution of these characteristics across species. This necessitates the development of synthetic works. Here we present a monographic catalogue of the diversity and phylogenetic structure of the entire South American iguanian family Liolaemidae, based on previously published studies. We also provide a complete table to summarize the distribution by country, elevational range, diet and reproductive mode of each species for which this information is available. The Liolaemidae family currently consists of a total of 229 species and subspecies belonging to the genera Ctenoblepharys, Liolaemus and Phymaturus. Remarkably, the genus Liolaemus alone comprises 209 of these taxa, consisting of 200 species, five of them polytypic, and recognized on the basis of 14 subspecies. Liolaemus species occur in Argentina, Bolivia, Brazil, Chile, Paraguay, Peru and Uruguay, representing the widest range of environments occupied by a single lizard genus. In contrast, the genus Ctenoblepharys is monotypic (Ctenoblepharys adspersa) and endemic to Peru, while 19 species of Phymaturus are distributed in Argentina and Chile. In these lizards, plant consumption and viviparity are strikingly common. Among Liolaemus, dietary information was available for 153 taxa. We found that 76 are arthropofagous, 71 omnivorous and six strictly herbivorous. Reproductive information was gathered for 136 species of this genus: 73 are viviparous and 63 oviparous. In Phymaturus, all species are viviparous and dietary information for 17 species revealed that 16 are herbivorous and only one omnivorous. Ctenoblepharys adspersa is arthropofagous and oviparous. As previously supported both theoretically and empirically, plant consumption and viviparity are associated with high latitudes and elevations. Finally, we suggest that the recently proposed species Phymaturus dorsimaculatus Lobo & Quinteros is conspecific to P. vociferator Pincheira-Donoso, from which the former taxon does not differ in morphology, coloration, patterns of sexual dimorphism or geographical distribution.
... Field studies did not involve endangered species. We studied the morphological characters commonly included in Liolaemus taxonomy, such as described in Laurent (1985), Etheridge (1993) Cei (1986, Lobo (2001Lobo ( , 2005 Abdala (2007), and Quinteros (2013). Color pattern follow Lobo and Espinoza (1999) and neck folds follow Frost (1992). ...
... La descripción se realizó en base a siete ejemplares colectados en dicha localidad, encontrándose depositados y disponibles en la Colección Herpetológica de la Fundación Miguel Lillo (ver Apéndice 1). En base al ejemplar tipo y paratipos se realizó la correspondiente diagnosis de la especie, basándose principalmente en caracteres comúnmente utilizados para estudios taxonómicos de Liolaemus como lo son lepidosis, morfometría y coloración (ver Laurent, 1985;Etheridge,1993Etheridge, , 1995Etheridge, , 2000Cei, 1986;Lobo, 2001;Abdala, 2007). Estos caracteres sirvieron para diferenciar a L. choique con el resto de las especies que conformaban el grupo de L. elongatus de ese entonces. ...
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Revalidation of Liolaemus choique Abdala, Quinteros, Scrocchi & Stazzonelli, 2010(Iguania:Liolaemidae). The description of Liolaemus choique was based on an exhaustive morphological diagnosis. However, a posterior phylogeographic study suggested that it would be a junior synonym of L. smaug. Herein, we present morphological behavioral and distributional data indicating that L. choique would be considered as a valid species. Therefore, we suggest to revalidate L. choique, as a distinct taxon from L. smaug.
... Sixty five morphological characters were measured, principally with reference to squamation characters, coloration pattern and body proportions, usually studied in Liolaemus, described or cited in Laurent (1985Laurent ( , 1992, Etheridge (1995Etheridge ( , 1998Etheridge ( , 2000, Cei (1986) and Lobo (2001). Additional data was taken from literature from Donoso Barros & Cei (1971); Cei & Scolaro (1982b); Cei (1986). ...
Article
A new specie of the Liolaemus genus is described in the present work, it was collected in the north-west of Santa Cruz Province, Argentina, at 1485 m. This new lizard belongs to the silvanae group, previously described as Vilcunia genus, characterized by the absence of precloacal pores in males, the presence of dorsal tridentate scales, lateral keeled nuchal scales overlap and postfemoral scales subimbricated. This Patagonian specie has a small size, with a squat look and shows a dorsal coloration pattern with outstanding dorsal-lateral bands, these bands can be of different colors, such as yellow, orange, brown or grey. The individuals of this specie are melanics in the ventral part of the body. This new specie lives in a place with extreme climatic conditions, low temperatures and with a cloudy sky most of the time. These are insectivorous and viviparous lizards, whose phylogenetic relationship with other members of the group are still unknown.
... La sistemática de los lagartos Iguania que se sigue en este trabajo es la propuesta por Schulte et al. (2003) que acepta la monofilia de los Iguanidae, dentro de los cuales se incluyen ocho subfamilias, distribuidas principalmente del Nuevo Mundo.En particular, la monofilia de la subfamilia Tropidurinae* no ha sido probada, por lo cual se la considera un metataxón, y en consecuencia, se denota con un asterisco. Para la determinación de los restos de lagartijas se consideraron caracteres provistos por análisis filogenéticos (Frost y Etheridge 1989;Etheridge 1995Etheridge , 2000Frost et al. 2001;Lobo 2001) y confrontación directa con material osteológico comparativo de especies actuales depositado en la Colección Herpetológica de la Universidad Nacional de Mar del Plata -Sección Osteología (UNMdP-O). ...
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p> En la presente contribuci ó n se da a conocer una acumulaci ó n de restos de lagartijas recuperados en el sitio arqueol ó gico Orejas de Burro 1 (OB1), ubicado en el campo volc á nico Pali Aike, sur de la provincia de Santa Cruz (Argentina). Los materiales incluyen restos de coronoides, dentarios, frontales, maxilares, huesos largos, parietales, premaxilares, v é rtebras y fragmentos de otros huesos. Las caracter í sticas osteol ó gicas permiten la asignaci ó n taxon ó mica de estos elementos al g é nero Liolaemus. Hasta el momento se ha contabilizado un n ú mero superior a 100 dentarios izquierdos y 100 dentarios derechos, que permiten sostener la presencia de al menos 100 individuos. La abundancia de bolos de regurgitaci ó n de lechuzas hallados en el sitio explicar í a el ingreso de los restos de lagartijas. El hallazgo en OB1 es el tercer reconocimiento del g é nero Liolaemus en un sitio arqueol ó gico de Argentina, constituye el hallazgo m á s austral conocido, y se destaca por la gran abundancia y diversidad de elementos preservados. Hasta el presente, los sitios arqueol ó gicos de Patagonia se caracterizan por la homogeneidad taxon ó mica de lagartijas, donde el ú nico tax ó n presente es Liolaemus. </p
... We studied the morphological characters traditionally used in Liolaemus taxonomy, including those of Laurent (1985), Cei (1986Cei ( , 1993, Etheridge (1993Etheridge ( , 1995Etheridge ( , 2000, Lobo (2001), and Abdala (2002Abdala ( , 2003Abdala ( , 2007. We follow the terminology of Smith (1946) for descriptions of squamation and Frost (1992) for neck-fold terminology. ...
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Liolaemus reichei was described by Werner and published by Otto Bürger in 1907 on the basis of 1 specimen captured in Iquique, Chile. The taxonomy of this species has been controversial, because it was originally placed in the genus Phrynosaura and was recently synonymized with Liolaemus stolzmanni based on a picture of the syntype of the latter species. We have identified a set of diagnostic characters that distinguish L. reichei from L. stolzmanni. Because there has been persistent confusion regarding the identity and the existence of the taxon described by Werner, we here resurrect, redescribe and designate a neotype of this species to stabilize the taxonomy of Liolaemus reichei. This work is based on specimens that were captured near Iquique, Chile, and is consistent with the original description. © 2018 Universidad Nacional Autonoma de Mexico. All rights reserved.
... The lizard genus Liolaemus is distributed across most of the temperate part of continental South America and is associated with climatic regimes ranging from the extremely arid Atacama desert (southern Peru) to the temperate Nothofagus rainforests (southern Chile) (Cei, 1986;Lobo, 2001). Liolaemus currently includes 267 described species and represents globally the most species-rich temperate zone amniote genus. ...
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Hybridization is likely to occur more often between closely related taxa that have had insufficient time to diverge to the point of reproductive incompatibility; hybridization between deeply divergent lineages is rare. In squamate reptiles, hybridization has been proposed as a possible explanation for the extensive paraphyly observed in mitochondrial gene trees in several species complexes of the South American lizard genus Liolaemus. One of the best‐documented cases is within the L. boulengeri and L. rothi complexes, which diverged ~5.5 million years ago. Here we describe a comprehensive study for approaching the hybridization hypothesis between these lizard species complexes. We explored the level of gene tree discordance using the novel “extra lineage contribution” statistics (XLC, presented in this study), that quantifies the level of gene tree discordance contribution per individual within a species. We included molecular data (12 nuclear and 2 mitochondrial genes) from 127 individuals, and results of a coalescent model‐based analysis show that the most likely explanation for the gene tree‐species tree discordance is interspecific hybridization. Our best‐supported hypothesis suggests current and past hybridization between L. rothi (rothi complex) and L. tehuelche (boulengeri complex), and independently between L. rothi and L. boulengeri and L. telsen (boulengeri complex). The hybrid descendants are characterized by intermediate phenotypes between the parental species, but are more similar to L. rothi in body size. We discuss the possible role of hybridization in Liolaemus evolution. This article is protected by copyright. All rights reserved.
... Morphological analysis. To determine the taxonomic status, we studied the morphological characters traditionally used in Liolaemus taxonomy describing primarily lepidosis, color patterns, and body proportions used in phylogenetic analyses, including those of Laurent (1985), Cei (1986Cei ( , 1993, Etheridge (1993Etheridge ( , 1995Etheridge ( , 2000, Etheridge & Espinoza (2000) Lobo (2001Lobo ( , 2005 and Quinteros (2012Quinteros ( , 2013. Descriptions of color in life were based on visual observations of living animals, freshly collected specimens and the inspection of photographs of several individuals taken in the field. ...
Article
We describe a new species of Liolaemus of the L. alticolor-bibronii group of the subgenus Liolaemus sensu stricto. We studied meristic, morphometric and qualitative pattern characters. Statistical tests were performed in order to evaluate morphological differences among the candidate species and the most closely geographically distributed species. Molecular analyses of Cyt-b mitochondrial gene were performed in order to estimate the position of the new species in relation to other taxa. We also recorded natural history data such as habitat, behavior, reproductive state, diet, and body temperature. Liolaemus absconditus sp. nov. differs from other species of Liolaemus in presenting a distinct combination of morphological character states of lepidosis and color pattern, being phylogenetically close to Liolaemus tandiliensis, Liolaemus gracilis and Liolaemus saxatilis. The new species is a saxicolous and endemic lizard of the Tandilia Mountain Range System of Buenos Aires Province.
... Not surprisingly, Liolaemus has been subject to many ecological and evolutionary studies (e.g., Harmon et al., 2003;Camargo, Sinervo & Sites, 2010;Cianferoni et al., 2013;Sheldon, Leaché & Cruz, 2015). Nevertheless, phylogenetic relationships among species of Liolaemus have been difficult to resolve, due in part to the incomplete taxon sampling of most studies (but see Olave et al., 2014), the discordance between inferences based on molecular and morphological data (Schulte II et al., 2000;Lobo, 2001;Lobo, 2005;Vidal & Labra, 2008), and to the incongruences between nuclear and mitochondrial gene trees caused by, likely, incomplete lineage sorting and hybridization (e.g., Olave et al., 2011). ...
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The genus Liolaemus is one of the most ecologically diverse and species-rich genera of lizards worldwide. It currently includes more than 250 recognized species, which have been subject to many ecological and evolutionary studies. Nevertheless, Liolaemus lizards have a complex taxonomic history, mainly due to the incongruence between morphological and genetic data, incomplete taxon sampling, incomplete lineage sorting and hybridization. In addition, as many species have restricted and remote distributions, this has hampered their examination and inclusion in molecular systematic studies. The aims of this study are to infer a robust phylogeny for a subsample of lizards representing the Chilean clade (subgenus Liolaemus sensu stricto ), and to test the monophyly of several of the major species groups. We use a phylogenomic approach, targeting 541 ultra-conserved elements (UCEs) and 44 protein-coding genes for 16 taxa. We conduct a comparison of phylogenetic analyses using maximum-likelihood and several species tree inference methods. The UCEs provide stronger support for phylogenetic relationships compared to the protein-coding genes; however, the UCEs outnumber the protein-coding genes by 10-fold. On average, the protein-coding genes contain over twice the number of informative sites. Based on our phylogenomic analyses, all the groups sampled are polyphyletic. Liolaemus tenuis tenuis is difficult to place in the phylogeny, because only a few loci (nine) were recovered for this species. Topologies or support values did not change dramatically upon exclusion of L. t. tenuis from analyses, suggesting that missing data did not had a significant impact on phylogenetic inference in this data set. The phylogenomic analyses provide strong support for sister group relationships between L. fuscus , L. monticola , L. nigroviridis and L. nitidus , and L. platei and L. velosoi . Despite our limited taxon sampling, we have provided a reliable starting hypothesis for the relationships among many major groups of the Chilean clade of Liolaemus that will help future work aimed at resolving the Liolaemus phylogeny.
... With about 240 recognized species, Liolaemus is one of the most widely distributed and species rich lizard genera worldwide (Lobo et al., 2010;Etheridge and Frost, 2010;Breitman et al., 2011Breitman et al., , 2013Avila et al., 2013). Liolaemus is distributed exclusively in South America, occurring in Bolivia, Paraguay, Peru, Chile, Argentina, Brazil and Uruguay, spanning different environments from the Andes Mountains to the Atacama Desert and from the Pacific Ocean shores to the Atlantic Ocean coasts (Lobo, 2001;Avila, 2003;Pincheira-Donosso et al., 2008). The systematics of the genus is quite complex, with several sections, series and groups being recognized (Schulte et al., 2000;Espinoza et al., 2004;Cruz et al., 2005;Fontanella et al., 2012;Olave et al., 2014). ...
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Intraspecific morphological variation is a relatively common pattern among lizards, where several selective factors have been suggested as responsible for this phenomenon. For instance, geographic variation could result from natural selection along with historical processes, whereas sexual dimorphism has usually been attributed to sexual selection, natural selection, and niche segregation. Liolaemus wiegmannii is a diurnal lizard distributed in the center, center-east and northwest of Argentina, as well as on the shores of southwest and south Uruguay. Information about morphological variation in this species is almost entirely limited to differences in mid-body scales between populations in the north and center of Argentina and some sex-based morphometric variation. Herein, we studied the geographic and sexual morphological variation of Liolaemus wiegmannii from Uruguay to test the hypothesis of morphological isolation by distance and morphological structuring by geographic barriers (rivers), as well as exploring the occurrence of sexual dimorphism in morphometry and lepidosis. Neither geographic distance nor rivers seem to play an important structuring role on the external morphology of Liolaemus wiegmannii in Uruguay. Multiple multivariate analyses support the hypothesis that most of the external morphological variation is probably due to sexual dimor-phism. Natural and sexual selection acting on females and males, respectively, are the most plausible mechanisms underlying the dimorphism observed in this species.
... Algunas formas están restringidas a un tipo de hábitat como las especies saxícolas (que viven en rocas), arborícolas, arenícolas; otras en cambio, pueden encontrarse en un amplio rango: las llamadas especies generalistas. Recientemente hubo un creciente interés en estudios con enfoques ecomorfológicos en este género que podría atribuirse a su accesibilidad y al progresivo conocimiento que se fue generando en las últimas décadas, tanto de su morfología (Abdala y Moro 2003Moro , 2006, ecología (Schulte et al., 2004;Cruz et al., 2009, entre otros) y sobre su filogenia (Laurent, 1983;Schulte et al., 2000;Lobo, 2001Lobo, , 2005Morando et al., 2004, Lobo y Quinteros, 2005Lobo et al., 2007;Ávila et al., 2006;Abdala, 2007;Abdala y Quinteros, 2008;Pyron et al., 2013, entre otros). En el caso particular de nuestro grupo de trabajo, luego de la obtención de caracteres morfológicos de la anatomía interna, externa y uñas de numerosas especies de Liolaemus, y mediante la aplicación de la metodología arriba descrita, hemos abordado una amplia gama de preguntas, de las cuales seleccionamos algunas. ...
... La morfometría es utilizada en la descripción cuantitativa, análisis, interpretación y variación de las formas. Así, estudios en sistemática basados en la morfología de los organismos requieren técnicas de descripción y comparación de la estructura de las formas (Rohlf 1990, Bernal taxonómicas y filogenéticas entre ellos, por ejemplo, descartando o aceptando diferencias entre grupos de individuos que ocupaban un nivel taxonómico incierto (Barrio 1965, Cei 1980, Heyer et al. 1990, Arikan et al. 1998, Maneyro & Arrieta 2000, Lobo 2001Lobo & Abdala 2001, Oliver & Lobo 2002, Méndez et al. 2004, Arroyo et al. 2005, Fabrezi 2006). Debido a la limitada capacidad de dispersión y filopatría en anfibios (Seppä & Laurila 1999;Lampert et al 2003), es frecuente que poblaciones separadas geográficamente presenten diferencias intraespecíficas en sus estructuras morfológicas, particularmente en el tamaño corporal, generadas por una acumulación de diferencias genéticas y morfológicas debido a este aislamiento geográfico , Laugen et al. 2002, Schäuble 2004. ...
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Las diferencias fenotípicas intraespecíficas entre poblaciones separadas geográficamente es común entre los anfibios anuros. En el presente trabajo se realiza una caracterización morfométrica de Hypsiboas cordobae, con el objetivo de estudiar la variación geográfica a lo largo de la distribución de la especie. Se midieron 15 variables un total de 86 individuos, pertenecientes a seis localidades de las provincias de Córdoba y San Luís. Se observó dimorfismo sexual en diez de las variables. El análisis comparativo mostró diferencias significativas entre poblaciones, y el análisis discriminante mostró altos porcentajes de clasificación (87,50%). Ocho variables morfométricas mostraron relación significativa y positiva con la latitud; mientras que diez variables mostraron relación significativa y positica con la altitud. Sin embargo, en ambos casos las correlaciones entre las distancias morfométricas y las latitudinales/altitudinales no resultaron significativas (Test de Mantel). Esta variación geográfica en la morfometría de la especie puede deberse a diferencias ambientales, o podrían deberse también a la edad de los individuos. Por lo tanto, estos datos podían ser complementados con estudios relacionados directamente a las condiciones ambientales de cada localidad, como así también estudios de esqueletocronología que permitan determinar la edad de los individuos. ABSTRACT Morphometric differentiation and geographic variation among populations of Hypsiboas cordobae (Anura: Hylidae). Intraspecific phenotypic differentiation among geographically separated populations is common in anurans. We performed morphometric analyses of the endemic frog Hypsiboas cordobae in order to assess the geographic variation of morphometric variables along of distribution of the species. We measured 15 morphometric variables on 86 individuals from six localities of Córdoba and San Luís provinces. We found sexual dimorphism in ten variables. The comparative analysis showed significant differences between populations and discriminant analyses showed high percentages of classification (87,50%). Eight morphometric variables showed positive correlation with latitude; and ten variables showed positive correlation with altitude. However, the correlations between morphometric distances and latitudinal/altitudinal were not significant. This geographic variation in the morphometry of the species may be due to environmental differences, or may be due to the age of individuals. Therefore, these data could be supplemented by studies directly related to the environmental conditions of each locality, as well as skeleton-chronological study to determine individual age.
... Six subspecies of L. pictus have been described based on morphological traits and geographical distribution (Donoso-Barros 1966, 1970Urbina & Zúñiga 1977;Pincheira & Núñez 2005), and four of these are restricted to the Chiloé Archipelago. A taxonomic study of these subspecies by Young (1998) did not define them clearly, and Lobo (2001) proposed raising the subspecies to full species status. Vidal et al. (2012a), however, suggested that disparities between mtDNA haplotypes and subspecific designations could partly be due to selection on morphology. ...
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Understanding the factors that cause heterogeneity among gene trees can increase the accuracy of species trees. Discordant signals across the genome are commonly produced by incomplete lineage sorting (ILS) and introgression, which in turn can result in reticulate evolution. Species tree inference using the multispecies coalescent is designed to deal with ILS and is robust to low levels of introgression, but extensive introgression violates the fundamental assumption that relationships are strictly bifurcating. In this study, we explore the phylogenomics of the iconic Liolaemus subgenus of South American lizards, a group of over 100 species mostly distributed in and around the Andes mountains. Using mitochondrial DNA (mtDNA) and genome-wide restriction-site associated DNA sequencing (RADseq; nDNA hereafter), we inferred a time-calibrated mtDNA gene tree, nDNA species trees, and phylogenetic networks. We found high levels of discordance between mtDNA and nDNA, which we attribute in part to extensive ILS resulting from rapid diversification. These data also reveal extensive and deep introgression, which combined with rapid diversification, explain the high level of phylogenetic discordance. We discuss these findings in the context of Andean orogeny and glacial cycles that fragmented, expanded, and contracted species distributions. Finally, we use the new phylogeny to resolve long-standing taxonomic issues in one of the most studied lizard groups in the New World.
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Taxonomic lists are important tools for understanding biodiversity and they provide informationat different scales about the species distributed within a region; as such, they allow us tounderstand the identities of the species that compose a given supra-specific taxon. While listsof the species of the genus Liolaemus have been published over the past few decades, providingtaxonomic and systematic information, these lists are now outdated and the number of validspecies within the genus has fluctuated rapidly over time and many taxonomic changes havegone unregistered. Although there are alternative means to consult this information, thesesources have suffered from a number of involuntary errors as well as some arbitrary omissionsor inclusions that have affected the dynamics of species inventories. In this work, we presentthe most complete taxonomic enumeration of the species of Liolaemus to date, including theavailable names for all of the valid and invalid species associated with these taxonomic changes.We recognize a total of 283 valid species in Liolaemus, along with another 64 invalid names.Fifteen taxa are recognized as species inquirenda, eight as nomina dubia, and two as nomina nuda.We hope that the careful review of each case and the exhaustive literature review will make thiswork a primary reference on South America's most speciose lizard genus.
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We provide the description for a new species from Catamarca province, Argentina that belongs to the Liolaemus darwinii group. The new species was previously considered as a population of L. ornatus, but it can be distinguished from that species mainly because of differences on coloration pattern, and its geographic distribution, which is disjunct from the populations of L. ornatus. The new species is characterized for having males with a dorsal background coloration of light brown to ferrous red, the head darker than the rest of the body, paravertebral markings rounded and never in contact, dorsolateral strips lightly marked, small dark brown spots on the throat, and a belly that is white or light red.
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The Liolaemus alticolor group (Iguania: Tropiduridae) currently includes two or three species of morphologically similar, small-bodied lizards distributed from southern Peru and Bolivia to northern Argentina and Chile. Recently, a few populations of L. alticolor from northwestern Argentina were reported to be reproductively bimodal - having both oviparous and viviparous females at the same locality. We reexamined lizards from these putatively bimodal populations and found evidence that these populations include two sympatric yet distinct species - one oviparous, the other viviparous. These species can also be distinguished from L. alticolor sensu stricto from the type locality (Tiahuanaco, Bolivia). Here we describe the two new species, L. ramirezae and L. pagaburoi, from the province of Tucuman, Argentina. Liolaemus ramirezae differs from L. alticolor in having distinct neck folds, precloacal pores in females, and an oviparous reproductive mode, and in lacking both spots on the throat in males and a vertebral line. Liolaemus pagaburoi differs from L. alticolor in having distinct paravertebral markings, slender dorsal stripes, and more rugose head scales. Recognition of these new species as distinct resolves the paradox of reproductive bimodality in L. alticolor. Interestingly, both of the new species appear to be microhabitat specialists that associate with a single species of plant in their respective habitats. As predicted by the cold-climate hypothesis, the oviparous species is distributed in a warmer climate than is the viviparous species.
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Abstract— Data scored for cladistic analyses may be quantitative or qualitative, continuous or discrete, and show overlapping or non-overlapping values between taxa. Quantitative and qualitative are modes of expression of data, while continuous or discrete refer to properties of the set of numbers that express the data; both these pairs of terms have been confused with overlapping and non-overlapping. The degree of overlap of values between taxa is often used to filter characters in cladistic analyses: if a minimum amount of overlap is exceeded, or a minimum amount of disjunction not reached, characters are rejected as “not cladistic". However, this rests on a confusion between features of taxa and features of individual organisms (attributes). Cladistic characters are features of taxa, and comprise frequency distributions of attribute values over individuals of a taxon. Cladistic characters logically cannot overlap, although taxa may have overlapping attribute values. Thus, a priori rejection of characters that have overlapping attribute values is non-sensical. Such data may still be rejected from consideration for cladistic analysis if it could be demonstrated that they contain little recoverable phylogenetic signal. Few published analyses have empirically tested this. An analysis of overlapping morphometric data from three series of Banksia suggests that, at least in these cases, they map phylogeny almost as accurately as more conventional, qualitative morphological data. While more such tests are required, morphometric data should not be rejected a priori from cladistic analyses.
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Comments about reproduction and ecology are given. -from English summary
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A population chromosome polymorphism due to a single centric fission of pair four in the lizard Liolaemus fuscus (2n = 32) from northern Chile, is described. Metaphase II analysis of heterozygote males showed a regular meiotic segregation (97%). This supports hypotheses that centric fission rearrangements are involved in this lizard's karyotypic diversity and that the meiotic effects of fission heterozygotes are nearly neutral in most populations.
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Morphometric studies on Liolaemus and related genera support the validity of Ctenoblepharis Tschudi, 1845, here considered to be monotypic. Ortholaemus Girard, 1857, is probably a natural group which includes all species of Liolaemus with more than one row of scales between labials and subocular, i.e., lutzae Mertens, occipitalis Boulenger, scapularis Laurent, wiegmanni Dumeril and Bibron and Pelusaurus cranwelli Donoso-Barros, in addition to the taxa already mentioned by Cei (1979). Nevertheless, no full generic recognition is recommended for Ortholaemus because the group of Liolaemus fitzingeri, and especially L. cuyanus Cei and Scolaro (formerly considered conspecific), bridge the gap between it and other Liolaemus species. The genus Phrynosaura Werner, 1907, is revived for reichei Werner, stolzmanni Steindachner and audituvelatus Nuñez and Yañez. Two species previously also classified in Ctenoblepharis and returned to Liolaemus by Cei (1979) are actually distinct enough to be placed in a new genus, Ceiolaemus. Helocephalus nigriceps Philippi is a Liolaemus. The status of two other Liolaemus species is enigmatic: donosobarrosi (Cei) and insolitus Cei and Pefaur. Liolaemus modestus Tschudi is not a Liolaemus, but a senior synonym of Stenocercus moestus Boulenger, which must therefore be renamed Stenocercus modestus (Tschudi).
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The recently-developed statistical method known as the "bootstrap" can be used to place confidence intervals on phylogenies. It involves resampling points from one's own data, with replacement, to create a series of bootstrap samples of the same size as the original data. Each of these is analyzed, and the variation among the resulting estimates taken to indicate the size of the error involved in making estimates from the original data, In the case of phylogenies, it is argued that the proper method of resampling is to keep all of the original species while sampling characters with replacement, under the assumption that the characters have been independently drawn by the systematist and have evolved independently. Majority-rule consensus trees can be used to construct a phylogeny showing all of the inferred monophyletic groups that occurred in a majority of the bootstrap samples. If a group shows up 95% of the time or more, the evidence for it is taken to be statistically significant. Existing computer programs can be used to analyze different bootstrap samples by using weights on the characters, the weight of a character being how many times it was drawn in bootstrap sampling. When all characters are perfectly compatible, as envisioned by Hennig, bootstrap sampling becomes unnecessary; the bootstrap method would show significant evidence for a group if it is defined by three or more characters.
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Most of 12 taxa karyotypes retain 6 pairs of metacentric macrochromosomes (primitive), but show reduced numbers of microchromosomes (2n=34, 32 and 30). Others whow increased diploid numbers due to macrochromosomal fissions (up to 4 fissions, 2n=40). One shows a fission polymorphism.
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The karyotypes of Chilean lizardsLiolaemus pictusandLiolaemus cyanogaster is described for the first time. Both species possess 34 chromosomes; 6 pairs of macrochromosomes and 11 pairs of microchromosomes. Karyologically it is possible to differenciate this species because the pair No. 2 is metacentric (m) inL. pictusand submetacentric (sm) inL. cyanogaster. It is shortly discussed the signification of formule 2n=34 for the species ofLiolaemus analized karyologically and its possible mechanism of acquisition.
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Liolaemus monticola , Müller and Hellmich, is a highly variable, endemic montane species distributed along the temperate Andes in Chile. The subspecies (Müller and Hellmich) karyotypes established from their type localities are Liolaemus monticola villaricensis and Liolaemus monticola chillanensis ; they retain a relatively conservative 2 n &equals; 32, with 12 macrochromosomes and 20 microchromosomes. Liolaemus monticola monticola differs strikingly from the former having a diploid number ranging from 38 to 40 in different variants. The increased diploid number can be explained by several possible independent chromosomal centric fissions of 1 or 2 pairs of macrochromosomes, and changes in the chromosomal morphology can be explained by pericentric inversion and simple translocation. Liolaemus monticola monticola is also polymorphic for chromosomal fission in pair no. 3. There is also an increase in the number of microchromosomes.