With 36 species and at least nine potentially independent lineages (not formally described yet) occurring mostly in theAndes and adjacent Patagonia and Puna plateau areas, Phymaturus lizards represent one of the most endemic vertebrategroups of the arid southwestern region of South America. Phylogenetic relationships among species of Phymaturus areinferred using mainly a morphological data set of 206 characters. Also available sequences of mitochondrial DNA for sev-en terminals were added for a total evidence analysis. Most information is included in the discrete characters block; mostcharacters involve color pattern, osteology and squamation. Continuous characters were taken from body proportions,squamation and skeletons. Among morphological data, binary polymorphic characters were analyzed applying the scaledcoding criteria. Continuous characters were entered in the analysis using standardized ranges, a method that allows a sim-ple optimization to estimate distances/costs avoiding the arbitrary coding as discrete characters. For our parsimony anal-yses we chose the implied weights method, which underweights homoplastic characters. Several runs were madeanalyzing all the information combined and also separating morphological from molecular datasets. Binary polymor-phisms were also analyzed as missing data. All characters affected by sexual dimorphism were analyzed separating thesexes; female information was more congruent with the total evidence analysis. Characters involving continuous and poly-morphic information are relevant for searching and building phylogenetic hypotheses in Phymaturus. There exists signif-icant congruence between the molecular information analyzed in this study and previous published analyses. Within bothmain clades of Phymaturus, northern subgroups are those more recently originated during the genus diversification. Spe-cies belonging to the puna subclade of the palluma group are arranged in two natural groups, one distributed in the north(Catamarca and La Rioja provinces), and the other in the south, La Rioja and San Juan provinces. Within the patagonicusgroup, the majority of the species are arranged in a south-central Chubut clade, eastern-central Chubut clade, central Rio Negro clade and a Payunia clade.

after the study of a diverse collection of Phymaturus from three argentinian institutions and additional samples collected in the last two years we discovered several populations of uncertain taxonomic status. Based on 93 morphological characters of squamation, color pattern, gular and nuchal folds, precloacal pores, and morphometric data, we conclude that at least four of those are independent lineages which require formal description. characters related to sexual dimorphism and dichromatism as well as the ontogenetic change of several others, from juvenile to adult specimens, are described. according to the most recent revision of the genus (Lobo and Quinteros, 2005a) and considering the descriptions made in the last four years, the taxonomic composition of the genus was raised to 23 species. in this study we provide the formal description of four additional new taxa, including their diagnosis and detailed comparisons with other members of their species groups. Two new species belong to the patagonicus group (provinces of chubut and rio negro, in patagonia between 46° and 41° of latitude) while the other two belong to the palluma group (neuquén and catamarca provinces, western argentina, 39° and 27-26°30' of latitude respectively). with the exception of one case for which four localities are reported, all new species are restricted to their type localities. This fact confirms a common historical distributional pattern for most species of the genus.

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The behavior of lizards can be highly influenced by chemical senses. The most studied pheromonal sources in lizards has been the femoral and precloacal gland secretions, although studies have been focused on male secretions, probably because these glands are usually only present in males or are poorly developed in females when they are present. Here, we aimed to study inLiolaemuschiliensis, one of the fewLiolaemusspecies in which females have precloacal glands, if female precloacal secretions convey information. We recorded the response of both sexes to secretions from females and males, as well as to control (solvent). The lizards started to explore the secretions sooner than the control. Both sexes moved more when exposed to female secretions than to the control, and males, but not females, explored female secretions more than the other scents. These results suggest that volatile compounds of the secretions allow lizards to recognize the presence of conspecifics, and, at least for males, these trigger the exploration of non-volatile compounds of the secretions that may reveal the sex of the individual that deposited them. This is the first study that explores the response to female precloacal secretions inLiolaemus, and data indicate that the female secretions ofL. chiliensiscontain relevant information for social interactions.

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Abstract Animals employ a wide variety of communication tactics that rely on distinct sensory modalities. Lizards are characterized by their heightened dependence on chemical and visual communication. Some authors have proposed that a reduced number of chemical secretory pores may be associated with an increased visual dependence in some species. Here, we study two species of Liolaemus lizards with different chemical features to compare their visual and chemical communication. The first species, L. coeruleus, does not have precloacal pores in either sex, while L. albiceps has precloacal pores in both sexes. We expect that L. coeruleus will principally adhere to the visual modality, while L. albiceps will show greater chemical responses. We filmed the lizard’s response to different chemical and visual stimuli. In the trials, both species demonstrated chemical self-recognition, L. albiceps exhibited less total time in motion but more behavioural displays in the presence of conspecific scents, suggesting conspecific chemical recognition too. On the other hand, visuals results showed that L. coeruleus reacted more to the presence of conspecifics than L. albiceps. These observations suggest that L. coeruleus relies more on visual signalization, while L. albiceps has a greater dependence on chemical communication. Our results may indicate a correspondence between precloacal secretions and the response to these by conspecifics in both species studied.

Studies about copulatory organs in Squamata were restricted to the morphology of hemipenes until Bo ¨hme reported homologous paired structures in females of a species of Varanus, which he called hemiclitores. We report the presence of hemiclitores in females of Phymaturus and of two species of Liolaemus and describe observations on the interspecific variation in hemipenis morphology in Phymaturus. Phymaturus and Liolaemus belong to Liolaemidae, a species-rich family of lizards; research about hemipenis morphology is scarce and limited to a few species of those genera. We found the retractor clitoridis magnus in all of the species analyzed; however, the transverse penis was not present in all of them. The general structure of hemiclitores of Phymaturus and Liolaemus females resembled that described for other squamate species; they were smaller than hemipenes and exhibited a sulcus spermaticus. The variation found in different features of these organs (shape, size, pigmentation), as well as the general morphology of hemipenes, should be studied more extensively in more species. These results contribute to the description of structures that are still poorly known in the large groups of Squamata and support the idea that hemiclitores should be considered an apomorphy of Squamata.RESUMEN.-Los estudios de los ´organos copuladores de Squamata estuvieron restringidos a la morfologı ´a de los hemipenes hasta que Bo ¨hme report ´o la presencia de estructuras pareadas homologas a la de los machos, a las que llam ´o hemiclitoris. Nosotros reportamos la presencia de hemiclitoris en las hembras del ge ´nero Phymaturus y en dos especies del ge ´nero Liolaemus ası ´como observaciones sobre la variaci ´on interespecı ´fica en la morfologı ´a de los hemipenes de Phymaturus. Phymaturus y Liolaemus pertenecen a Liolaemidae, una familia de lagartijas muy diversa, para los cuales las investigaciones en la morfologı ´a de los hemipenes son escasas y limitadas a unas pocas especies. En todas las especies revisadas encontramos el m ´usculo retractor clitoridis magnus pero no el transversus penis. La estructura general de los hemiclitoris de las hembras de Phymaturus y Liolaemus, se asemeja a la descrita para otras especies de Squamata; son ma ´s pequen ˜as que los hemipenes y exhiben un sulcus spermaticus. La variaci ´on encontrada en diferentes caracterı ´sticas de estos ´organos (forma, taman ˜o, pigmentaci ´on), ası ´como la morfologı ´a general de hemipenes, debe ser estudiada ma ´s extensamente incluyendo un mayor n ´umero de especies. Estos resultados contribuyen con el reporte y la descripci ´on de estructuras que son a ´un poco conocidas en los grandes grupos de Squamata, y soporta la idea que los hemiclitoris deben ser consideramos una apomorfia de Squamata.