A new species of Phymaturus of the P. mallimaccii Group from the Andes of central Chile (Iguania: Liolaemidae)

Nós descrevemos uma nova espécie de Phymaturus da Província Mountain na Cordilheira San Ramón, Região Metropolitana, Chile. Essa nova espécie pertence ao grupo palluma e ao subgrupo de P. mallimaccii. No entanto, esse novo táxon possui características incomuns para o subgrupo de P. mallimaccii: grande fragmentação da escama subocular (4–5), escama pré-ocular sempre menor que a cantal, machos com coloração dorsal oliva e padrão difuso formado por faixas claras nas áreas paravertebrais e um triângulo claro entre os ombros, com a extremidade orientada em direção ao rostro. Além disso, essa nova espécie é completamente isolada e é a espécie do subgrupo de P. mallimaccii que se encontra distribuída mais ao sul. Também é notável sua baixa abundância. Adicionalmente, fornecemos informações sobre sua história natural e discutimos alguns aspectos de populações ainda não-classificadas de Phymaturus chilenos.A new species of Phymaturus from the Provincia Mountain in the San Ramón Mountains in the Metropolitan Region of Chile is described. It is a member of the P. palluma Group and the P. mallimaccii subgroup. The new species can be distinguished from other members of the P. mallimaccii subgroup by having a highly fragmented subocular scale (4 or 5), a preocular scale that is smaller than the canthal, males having a olive dorsum with a diffuse pattern of light bands on the paravertebral fields and a light triangle between the shoulders, which has the apex of the triangle oriented toward the snout. The distribution of new species lies south of that of other members of the P. mallimaccii subgroup, from which it is completely isolated. The low abundance of the new species is noteworthy. Information about the natural history of the lizard is provided and some aspects of unnamed populations of Chilean Phymaturus are discussed.Se describe una nueva especie de Phymaturus para el Cerro Provincia de la Sierra de San Ramón en la Región Metropolita de Chile. Esta pertenece al grupo de P. palluma y al subgrupo de P. mallimaccii. La nueva especie puede ser diferenciada de los otros miembros del subgrupo de P. mallimaccii por presentar una alta fragmentación de la escama subocular (4 o 5), la escama preocular más pequeña que la cantal, machos con color dorsal oliva y un patrón difuso de bandas claras en los campos paravertebrales con un triángulo claro entre los hombros, el cual tiene el ápice orientado hacia el hocico. Esta nueva especie se distribuye más al sur que las otras especies del subgrupo de P. mallimaccii, de las cuales está completamente aislada. La baja abundancia de esta nueva especie es destacable. Se provee información sobre la historia natural del lagarto y se discuten ciertos aspectos sobre las poblaciones chilenas de Phymaturus que no han sido denominadas

Ontogenetic drivers of morphological evolution in monitor lizards and allies (Squamata: Paleoanguimorpha), a clade with extreme body size disparity

Background Heterochrony, change in the rate or timing of development, is thought to be one of the main drivers of morphological evolution, and allometry, trait scaling patterns imposed by size, is traditionally thought to represent an evolutionary constraint. However, recent studies suggest that the ontogenetic allometric trajectories describing how organisms change as they grow may be labile and adaptive. Here we investigated the role of postnatal ontogenetic development in the morphological diversification of Paleoanguimorpha, the monitor lizards and allies, a clade with extreme body size disparity. We obtained linear and geometric morphometric data for more than 1,600 specimens belonging to three families and 60 species, representing ~ 72% of extant paleoanguimorph diversity. We used these data to undertake one of the largest comparative studies of ontogenetic allometry to date. Results Heterochrony is likely dictating morphological divergence at shallow evolutionary scales, while changes in the magnitude and direction of ontogenetic change are found mainly between major clades. Some patterns of ontogenetic variation and morphological disparity appear to reflect ontogenetic transitions in habitat use. Generally, juveniles are more similar to each other than adults, possibly because species that differ in ecology as adults are arboreal as juveniles. The magnitude of ontogenetic change follows evolutionary models where variation is constrained around an optimal value. Conversely, the direction of ontogenetic change may follow models with different adaptive optima per habitat use category or models where interspecific interactions influence its evolution. Finally, we found that the evolutionary rates of the ontogenetic allometric trajectories are phylogenetically variable. Conclusions The attributes of ontogenetic allometric trajectories and their evolutionary rates are phylogenetically heterogeneous in Paleoanguimorpha. Both allometric constraints and ecological factors have shaped ontogeny in the group. Our study highlights the evolutionary lability and adaptability of postnatal ontogeny, and teases apart how different evolutionary shifts in ontogeny contribute to the generation of morphological diversity at different evolutionary scales

The true identity of the new world iguanid lizard liolaemus chillanensis Müller and Hellmich 1932 (Iguania: Liolaemidae) and description of a new species in the liolaemus elongatus group

Liolaemus is a particularly species-rich radiation of New World iguanid lizards from southern South America. Thanks to intense systematic and taxonomic research, the knowledge on its species-level diversity has increased rapidly over the past several years. The L. elongatus species-complex has received considerable attention and a remarkable case is Liolaemus chillanensis, a name that has been used for two different species that are sympatric in Termas de Chillán, central Chile. Since the holotype is lost, we propose that the first step to identify the true L. chillanensis is through the analysis of the original description. Then we provide a morphological and molecular characterization of L. chillanensis based on topotypes and a description of the taxon previously confused with it.Fil: Troncoso Palacios, Jaime. Universidad de Chile; ChileFil: Esquerré, Damien. The Australian National University; AustraliaFil: Urra, Félix A.. Universidad de Chile; ChileFil: Díaz, Hugo A.. Universidad de Chile; ChileFil: Castro Pastene, Carlos. Universidad de Concepción; ChileFil: Ruiz, Maria Soledad. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Salta. Instituto de Bio y Geociencias del NOA. Universidad Nacional de Salta. Facultad de Ciencias Naturales. Museo de Ciencias Naturales. Instituto de Bio y Geociencias del NOA; Argentin

Elevational Goldilocks zone underlies the exceptional diversity of a large lizard radiation (Liolaemus; Liolaemidae)

Mountains are among the most biodiverse regions on the planet, and how these landforms shape diversification through the interaction of biological traits and geo-climatic dynamics is integral to understanding global biodiversity. In this study, we investigate the dual roles of climate change and mountain uplift on the evolution of a hyper-diverse radiation, Liolaemus lizards, with a spatially explicit model of diversification using a reconstruction of uplift and paleotemperature in central and southern South America. The diversification model captures a hotspot for Liolaemus around 40°S in lineages with low-dispersal ability and narrow niche breadths. Under the model, speciation rates are highest in low latitudes (35°S) and higher elevations (>2,000 m). Temperature change through the Cenozoic explained variation in speciation and extinction rates through time and across different elevational bands. Our results point to the conditions of mid elevations being optimal for diversification (i.e., Goldilocks Zone), driven by the combination of (1) a complex topography that facilitates speciation during periods of climatic change, and (2) a relatively moderate climate that enables the persistence of ectothermic lineages and buffers species from extinction

Parallel selective pressures drive convergent diversification of phenotypes in pythons and boas

Pythons and boas are globally distributed and distantly related radiations with remarkable phenotypic and ecological diversity. We tested whether pythons, boas and their relatives have evolved convergent phenotypes when they display similar ecology. We collected geometric morphometric data on head shape for 1073 specimens representing over 80% of species. We show that these two groups display strong and widespread convergence when they occupy equivalent ecological niches and that the history of phenotypic evolution strongly matches the history of ecological diversification, suggesting that both processes are strongly coupled. These results are consistent with replicated adaptive radiation in both groups. We argue that strong selective pressures related to habitat-use have driven this convergence. Pythons and boas provide a new model system for the study of macro-evolutionary patterns of morphological and ecological evolution and they do so at a deeper level of divergence and global scale than any well-established adaptive radiation model systems

A new species of Phymaturus of the P. mallimaccii Group from the Andes of central Chile (Iguania: Liolaemidae)

© 2014 Departamento de Ciências Biológicas - ESALQ - USP. A new species of Phymaturus from the Provincia Mountain in the San Ramón Mountains in the Metropolitan Region of Chile is described. It is a member of the P. palluma Group and the P. mallimaccii subgroup. The new species can be distinguished from other members of the P. mallimaccii subgroup by having a highly fragmented subocular scale (4 or 5), a preocular scale that is smaller than the canthal, males having a olive dorsum with a diffuse pattern of light bands on the paravertebral fields and a light triangle between the shoulders, which has the apex of the triangle oriented toward the snout. The distribution of new species lies south of that of other members of the P. mallimaccii subgroup, from which it is completely isolated. The low abundance of the new species is noteworthy. Information about the natural history of the lizard is provided and some aspects of unnamed populations of Chilean Phymaturus are discussed

Liolaemus riodamas Esquerré, Núñez & Scolaro, 2013 (Reptilia: Squamata: Liolaemidae): new record for the Maule Region in Chile

Liolaemus riodamas is a recently described lizard, only known from a very restricted river canyon in the Andes of the O’Higgins Region in Chile. The species is particular for being one of the few Liolaemus lacking the pheromone secreting glands known as precloacal pores. Here we document a second locality for this species located 30 km southwest of the type locality. This is also 500 m higher than the type locality, therefore a considerable altitudinal extension. This is a very positive discovery since the type locality of L. riodamas is intervened by a hydroelectric project, and gives hope to the conservation of this unique taxon

Liolaemus riodamas Esquerré, Núñez & Scolaro, 2013, sp. nov.

Liolaemus riodamas sp. nov. Figures 6 to 10 Holotype. MNHNCL- 4684, male. Collected by Damien Esquerré and Herman Núñez at Las Damas River, Termas del Flaco, Libertador Bernardo O’Higgins Region; S 34 ° 56 ' 714-887 ´´ W 70 ° 23´634-682 ´´ 2012-2057 masl, between February 1 and 2, 2011. Paratypes. MNHNCL- 1962 to 1966 and MNHNCL-4682, 4683, 4685 and 4686. MNHNCL-1962, 4682 and 4686 are males and MNHNCL-1963, 1964, 1965, 1966 and 4685 are females. MNHNCL 4682-4686 same data as holotype. MNHNCL- 1962-1966 same locality, collected by Herman Núñez and Antonieta Labra, February 24, 1985. MNHNCL- 1964 and 4683 were used for diaphanisation. Etymology. The name comes from the locality they were found: Las Damas River (In Spanish known as “Río Las Damas”). The common names “Las Damas River Lizard” in English and “Lagarto del Rio Las Damas” in Spanish are proposed. Diagnosis. Medium to large-sized lizard, with a mean SVL of 80.9 mm and a maximum of 85.5 mm. Sturdy aspect, robust limbs and a strongly folded neck. Tail about 1.5 times the length of the body. 83–96 scales around midbody. General background coloration brown to dark gray, with no defined pattern. Dorsal coloration of the head dark brown to black. Some specimens have individual yellow scales scattered on the posterior region of the dorsum. A light-yellow coloration is observed at the ventral region of some live specimens that extends to the sides of the tail. This species presents the characteristic dorsal squamation pattern of the kriegi group, with small rounded and juxtaposed scales, and tiny granular scales in the interstitial region. The dorsal scales are weakly keeled and only around the vertebral line. It completely lacks precloacal pores, which makes it distinct from almost all of the species of the elongatus-kriegi complex. It is important, therefore, to differentiate this species from all of the species lacking these glands. First, because of this trait, is can be said that this species is not L. ceii as was previously suggested (Núñez & Torres-Mura 1992). Furthermore L. ceii has 102–115 scales around midbody, in contrast with the 83–96 scales in L. riodamas. From the original description of L. thermarum (Videla & Cei 1996), where it is described lacking precloacal pores, it differs in that L. thermarum would exhibit a dark bilateral stripe on the flanks, absent in the uniform pattern of L. riodamas. According to Avila et al. (2010), L. thermarum has two to three precloacal pores and a dorsal stripe, both traits absent in L. riodamas, however in a recent article (Avila et al. 2012) that sample of lizards is assigned to L. smaug. L. riodamas differs from L. neuquensis and L. coeruleus because these species have a lower number of scales around midbody (68–74 and 63–69 respectively, after Videla & Cei (1996) and Scolaro et al. (2007)), presence of dorsal pattern and a blue/greenish ventral coloration. From L. cristiani it differs because this species has a black stripe along the sides of the trunk, and a smaller body size much smaller, body size (mean SVL of 70.7 vs. 80.9 mm in L. riodamas). The species differs from L. flavipiceus because the latter has a lower number of scales around midbody (73–81). In addition L. riodamas lacks the body melanism of L. flavipiceus, and the red ventral coloration in some specimens of L. flavipiceus. From L. punmahuida it differs because this species is larger (maximum SVL of 95 mm in L. punmahuida vs. 85 mm in L. riodamas); furthermore L. punmahuida has an ochre background coloration, and bright red-yellowish coloration in the cloacal region, in contrast to the brown to gray dorsal coloration, and the occasional and slight yellow pigmentation in the cloacal region of L. riodamas. L. riodamas differs from L. tregenzai because this species has a green-bluish ventral coloration, and an evident sexual dichromatism, both traits not observed in L. riodamas. Some L. leopardinus males lack precloacal pores, but this species is clearly distinct from L. riodamas because of the leopard-like dorsal pattern of L. leopardinus. L. capillitas males also lack precloacal pores sometimes, but this species has only 58–67 scales around midbody, and a red cloacal coloration in females, not observed in L. riodamas. Description of the holotype. Male, with an SVL of 81.83 mm. Head slightly longer than wide: 17.4 mm long (from the anterior edge of the ear opening to the point of the snout), 15.75 mm wide (between the anterior edges of the ear openings) and 10.78 mm high (at the level of the anterior edge of the ear openings). Snout length (from the anterior margin of the eye to the rostral scale): 6.21 mm. Neck wider than head, effect of a prominent longitudinal fold on the neck. Hind limb extended forward barely reaches the armpit. The tail is regenerated. It lacks precloacal pores. Rostral scale rectangular, 3.2 times wider than high and in contact with eight scales, including the nasals, which touch the rostral with only the anterior corner of the scale. Nasals polygonal, the nostril located posteriorly, occupying half of the scale’s surface. Two postrostrals irregularly squared, followed by four internasal scales, the two medial ones about two times larger than the other two. Three frontal azygos, the most anterior the largest, and the most posterior the smallest. Eight frontonasals in contact with the frontal azygos. Two squared prefrontal scales, the frontal undivided. Two longitudinally elongated postfrontals, the left one transversally divided in half. Interpatietal irregularly pentagonal, in contact with six scales. A gray pineal eye in the middle of interparietal scale. Four small and squared parietal scales, slightly larger than the interparietal, although they resemble two longitudinally elongated parietal scales transversally divided in half. The scales in the nuchal and supratemporal regions are medium-sized, polymorphic, juxtaposed and smooth. 4-5 large supraoculars, accompanied by 22 - 20 small supraoculars. 8 - 8 very elongated and imbricate superciliary scales, excluding the canthal. Numerous scale organs on all the dorsal region of the head, increasing in numbers towards anterior part of the head. Canthal scales sharpened towards the anterior end. 6 - 6 loreal scales, excluding the canthal. A large and elongated subocular scale, divided at the posterior end, forming a small scale, and both together spanning the entire length of the eye. 8 - 8 lorilabial scales, in one row. 7-8 supralabials, the fourth-fifth one more elongated and curved upwards posteriorly. It has a slightly elongated snout, with the loreal distance (distance between the anterior margin of the eye and the rostral scale) barely larger than the ocular diameter. The scales in the anterior border of the eye are elongated and overlapping, each one with a scale organ on it. 15 - 15 upper and 14 - 14 lower palpebrals, all squared and each one with a scale organ on it. Temporal scales rounded, smooth and juxtaposed. Auricular scale barely distinguishable. Timpanic scales poorly developed, small and convex. Scale organs present in all the lateral region of the head, being especially numerous in the lorilabial and loreal scales. Mental scale barely wider than rostral, although twice its height, and in contact with four scales. Four pairs of postmentals, the second pair separated by two scales. 5 - 5 infralabials, with scale organs scattered on them. Scales in the gular region rounded, smooth and imbricate. Lateral region of neck with a pronounced longitudinal fold, ending with an antehumeral fold. Neck scales small, convex, granular and with very small granular scales in the interstitial space. Dorsal scales small, smaller than the ventral scales, rounded and juxtaposed. Poorly developed keels on the mid-dorsal scales, and disappearing to the sides. The scales on the flanks therefore are smooth, but in form and arrangement very similar to the dorsal scales, being only slightly larger. Tiny granular scales are scattered in interstitial space in the dorsal region and flanks. No folds are observed in the lateral region of the trunk. Ventral scales are rounded, smooth and imbricate. 85 scales around midbody. Dorsal scales of the arm are subtriangular, smooth and imbricate. Dorsal scales of forearm are very similar but more rounded, as they are in the dorsal region of the hand. On the ventral region of the arm the scales are small, granular, juxtaposed, and surrounded by even smaller granular scales in the interstitial space. The scales on the ventral region of the forearm are as large as the dorsal ones, subtriangular, imbricate, and start showing keels towards the palm, where the scales are completely keeled, triangular, imbricate and with a slightly jagged edge. The third finger of the left hand has 16 rectangular and transversally disposed lamellae, each one provided with three keels. The scales in the dorsal femoral region are rounded, smooth, subimbricate and with few small granular scales on the interstitial region. In the dorsal tibial region they are subtriangular, juxtaposed, slightly keeled and with visible tiny granular scales in the intertstitial region. In the dorsal region of the foot the scales are rounded, smooth and imbricate. The scales on the ventral femoral and tibial regions are rounded, smooth and imbricate. In the plantar region scales are triangular, slightly keeled and imbricate. Some scales in the plantar region are slightly jagged on the edges. The fourth toe has 25 rectangular and transversally disposed lamellae, each one provided with three keels. The dorsal scales on the tail are quadrangular, imbricate and keeled. Ventral scales on the tail are triangular, smooth and imbricate. Color and pattern in preservative. It lacks pattern. The dorsal region is uniformly brownish, with the dorsal region of the head darker than the rest of the body. From the groin, towards the posterior dorsal part of the body, a few white scales are scattered, not going further than the middle of the trunk. The ventral coloration is light gray. Variation in the paratypes. Body measurements were divided into males and females (holotype is included within the males): Mean and extreme values, in mm, for the males (five specimens) are: snout-vent length: 80.17 (72.71–85.5); axilla-groin distance: 38.12 (34.71–41.11); left forelimb length: 31.59 (27.43–33.16); left hindlimb length: 46.18 (42.7–48.85); tail length: 123.5 (112–135); head length: 17.19 (15.42–18.09); head width: 15.21 (13.44–16.24); and head height: 10.26 (9.57–10.78). For the females (six specimens) the measurements are: snoutvent length: 81.61 (78.7–84.8); axilla-groin distance: 38.92 (36.1–41.9); left forelimb length: 31.72 (29.8–34.4); left hindlimb length: 46.1 (42.8–50.2); tail length: 124 (value of only non-regenerated tail); head length: 17.82 (16.18–19.53); head width: 15.34 (14.15–16.64); and head height: 9.95 (9.52–10.3). No evident sexual dimorphism is observed. In the five males (including the holotype), the mean and extreme values of the number of scales around midbody are 86.4 (83–90). None has precloacal pores. Two transverse gular folds, including the antehumeral fold, instead of a longitudinal neck fold, are observed in two specimens. The male paratypes exhibit the following variation in squamation with respect to the holotype. Two frontal azygos, with the posterior one being larger, in two specimens. Two longitudinally elongated parietal scales in three specimens, 3 to 5 large supraoculars and 13 to 18 small supraoculars, 5 to 7 superciliaries, 4 to 7 loreal scales, 5 to 6 lorilabials, 5 to 6 supralabials and 5 to 6 infralabials. Slight keels in the dorsal region of the forearm in two specimens. Ventral scales of forearm rounded in two specimens. 18 to19 lamellae on the third finger of the left hand. Scales in the dorsal femoral region are imbricate and with no small granular scales in the interstitial region in one specimen, and imbricate but with granular scales in interstitial region in another specimen. Scales in the dorsal femoral region are slightly keeled in one specimen. Scales in the dorsal tibial region are subimbricate in one specimen and completely imbricate in another. 24 to 27 lamellae on the fourth toe of the left foot. In the females, the mean and extreme values of the number of scales around midbody are 84 (80–93). None has precloacal pores. Two transverse gular folds, including the antehumeral fold, instead of a longitudinal neck fold, are observed in one specimen. In comparison with the holotype the five female paratypes exhibit the following variation. Two frontal azygos, with the posterior one bigger, in three specimens. Six frontonasal scales in one specimen. Longitudinally enlarged frontal scale and short squared postfrontals in two specimens. Squared interparietal scale in one specimen. Two longitudinally elongated parietal scales in four specimens. 4 to 5 large supraocular and 14 to 19 small supraoculars, 6 to 8 superciliaries, 4 to 7 loreals, 6 to 8 lorilabials, 6 to 7 supralabials and 5 to 6 infralabials. Five pairs of postmentals in one specimen. Dorsal scales subimbricate in one specimen. Dorsal scales of the forearm very slightly keeled in three specimens. Ventral scales of the forearm rounded in two specimens. 18 to 21 lamellae in the third finger of the left hand. Dorsal femoral scales imbricate in two specimens, and with poorly developed keels in another specimen. Dorsal tibial scales subimbricate in two specimens. Dorsal scales of the foot slightly keeled in one specimen. 24 to 29 lamellae in the fourth toe of the left foot. Color variation is poorly visible within the preserved specimens. No kind of sexual dichromatism is observed. Some specimens are clearer, reaching a whitish gray tone, especially in the specimens of the older sample (1985). Perhaps the most pronounced variability is found in the melanism level of the dorsal region of the head, varying from dark brown to almost black. The clear scales scattered towards the dorsum starting from the groin, visible on the holotype, are visible only on the specimens of the newer sample (2011). The ventral region varies from whitish gray to gray. In some specimens a slight yellowish tone on the sides of the posterior half of the ventral region is observed. In all of specimens the ventral region of the head exhibits a pattern consisting of a gray background, mottled with white spots. Distribution. It has only been registered at the type locality, in an area of the narrow canyon of Las Damas River, between 2012 and 2057 masl. The river ends at the origin of the Tinguiririca River, near the locality of Termas del Flaco, in the Andean area west of the city of San Fernando, in the Libertador Bernardo O’Higgins Region, Chile. Natural History. Viviparous species. Saxicolous habits, it is found basking over the large rocks that form its natural habitat, with relatively low vegetation. It can also be seen climbing branches of Berberis sp., possibly to eat their flowers (Núñez & Torres-Mura 1992). In the altitudinally lowest part of its distribution it is sympatric with L. curis, and with Phymaturus damasense (Troncoso-Palacios & Lobo 2012) in all of its distribution. Osteological description. Based on specimen MNHNCL- 1964, female, double-stained skeleton. (Figure 11) Cranium features: cranium length (from cephalic condyle to premaxilla) 17.1 mm; cranium width (between sutures of maxilla and jugal) 11.88 mm; cranium height 6,95 mm; orbit length (between lacrimal and premaxilla) 6.78 mm. Anteriorly rounded premaxilla. The maxillar process is shorter than the nasal process (spine) of the premaxilla of the premaxilla; the premaxilla is pierced by two foramina, one on each side. The margin of these foramina is made by the premaxilla and the maxilla; those tiny holes are the passage for the medial ethmoidal nerves (Oelrich 1956). The widest part of the premaxilla is slightly shorter than the nasal spine of the premaxilla. The nasal process is projected backward and rests beneath the anterior junction of the nasal bones. Ventrally, the premaxillae bear six caniniform and pleurodont teeth. Also, ventrally is the incisive process, an antero-inferior projection, which is long, conspicuous and swollen. Nasal capsules relatively enlarged, medially and anteriorly margined by the nasal process of the premaxilla and by the premaxillar process of the nasal bones; the floor of the nasal capsule is the flat premaxillar process of the maxilla. The nasal capsule has a foramina connected with the choanae. Paired nasal bones, twice as long as wide, medially sutured to each other. The anterior portion of both bones covers the spinal process of the premaxilla as described. Laterally and anteriorly, these are sutured to the ascendant process of the maxilla and, caudally at the wide margin with the prefrontal bones. The nasals, at their posterior margin, broadly diverge to house the anterior tip of the frontal bone. The divergence is located at the level of the anterior margin of the orbit. The dorsal surface of the nasal bone is pierced by five small foramina, which are the passages for the cutaneous branches of the ethmoidal nerve and veins (Oelrich 1956). Also, the surface of the nasal bones is wrinkled, sinuous and carved out by the scales. Paired prefrontal bones triangular, with an acute projection backwards. The medial margin is broadly sutured to the nasals and anteriorly strongly sutured to the ascendant process of the maxilla and to the tiny lacrimal. Ventrally the prefrontals form the anterior ceiling of the ocular orbit. The prefrontals are deflected downwards, forming the descendant process of the prefrontals, sutured solidly to the palatine bones. This process forms the medial wall of the lacrimal foramen, whose external wall is completely margined by the small lacrimal bone. The descendant process of the prefrontal forms a small part of the orbito-nasal fenestra, which is completely formed by the descendant process of the frontal. Unpaired flat frontal bone, with a wrinkled surface, product of the overtopping scales. Anteriorly it is sutured to the nasals and prefrontal bones as described. The frontal forms the anterior margin of the ocular orbit. Backwards the frontal diverges widely to join with the parietal. Together, those bones form the epiphyseal foramen. The frontal is weakly sutured to the tiny postfrontal. Ventrally it has a tubular shape housing the olfactory channel. The paired postfrontal bones are laminar, quite small and forming a bridge, connecting the frontal and the postorbital bones. The parietal is a single bone, sutured anteriorly with the frontal as previously described. It is dorsally flattened with a wrinkled surface. Laterally the parietal is abruptly deflected downwards to form the posterior cranial cage. Anteriorly it has two postorbital processes, sutured to the postorbital bones. Towards the caudal end the parietal is widely spread in the supratemporal processes; these processes are sutured to the respective supratemporal bones and also to the paraoccipital process of the exoccipital. The caudal portion of the supratemporal bones is externally margined by the squamosal bone, medially by the paraoccipital process of the exoccipital, and ventrally by the postero-superior portion of the cephalic condyle of the quadrate bone. The supratemporal bones are projected forward through the inferior margin of the supratemporal process to house the parietal in a groove, as was described by Etheridge (1995). The parietal bone, on its external walls receives the superior tip of the epipterygoid bone, an osseous rod, attached to the pterygoid bone. Length of the temporal fenestra 5.5 mm. Postorbital bones are paired. They are flat bones with three processes, at their anterior margin they form the posterior margin of the orbit, and at their posterior margin, they form the anterior margin of the temporal fenestra. The dorsal (medial) apex of the postorbital joins the postfrontal and parietal as previously described. The anterior process is firmly sutured to the jugal, and at its posterior tip it is weakly sutured to the squamosal bone. Squamosal bones are paired, and are like curved rods, whose anterior processes are joined to the postorbital, as was described. Their caudal tips are widened, and there is a process projected to a notch on the cephalic condyle of the quadrate, which is pierced by this process. Quadrates are auricular-shaped paired bones. Inferiorly they present a grooved condyle attached to the mandible. Dorsally the quadrate has a cephalic condyle, sutured strongly to the paraoccipital process, to the supratemporal, and to the squamosal, as described. The attachment is weak and made through cartilaginous fibers. Posteriorly and medially there is a column that reaches the very summit of the quadrate: the posterior crista (Oelrich 1956). Close to the lower part of this column the pterygoid valve reaches the quadrate, to which it is attached by cartilaginous fibers. Jugals are two long and curved rods, forming the inferior ridge of the orbit, distally they are strongly sutured to the maxilla and to the tiny lacrimal; the lowest part of the jugal is deflected backwards and upwards and weakly attached to the anterior tip of the postorbital and squamosal by ligaments. Also, in the lowest part, the jugal is strongly attached to the ectopterygoid. On the external surface it exhibits four conspicuous foramina. Maxillae are paired bones, roughly triangular. The anterior process is sutured to the premaxilla as described. Medially, and as a part of the floor of the nasal concha it is sutured to the septomaxilla. This maxilla has cartilage covering the nasal concha. The ascendant p

Liolaemus carlosgarini Esquerré, Núñez & Scolaro, 2013, sp. nov.

Liolaemus carlosgarini sp. nov. Figures 1 to 4 Holotype. MNHNCL- 4563. Male, collected by Carlos Garín at International Road 115, below the Maule Lagoon. 19 H 358560 E– 6018117 S. 1915 masl. WGS 84. On February 22, 2008. Paratypes. MNHNCL- 4562 and one used for diaphanisation, males; MNHNCL-4561, 4564, 4565, 4566, 4567, 4568 and one used for diaphanisation, females. Same collection data as the holotype. Measurements in Table 1. Diagnosis. Small to medium sized lizard, with a mean SVL (Snout-Vent Length) of 60.7 mm and a maximum of 68.8 mm. Slender body, moderately robust limbs, head longer than wide, tail over 1.5 times body length, and 86- 101 scales around midbody. Background dorsal coloration is ochre, with a dark occipital stripe, parietal stripes as the background color, and lateral stripes the same coloration as the occipital stripe. Black spots distributed variably between the dorsal region of the head and limbs. Individual white scales scattered on the dorsal region of the body, and a clear and mottled ventral region. It can be distinguished from almost all of the species of the elongatus-kriegi complex by its extreme reduction of precloacal pores. None of the male samples, despite being only three, had these pores, and it seems that a very low frequency of males has them (C. Garín in. litt. 2011). Because the absence of precloacal pores is not absolute in this species, minimum diagnostic traits to distinguish it from all the species of the elongatus - kriegi complex are given. L. carlosgarini is distinguished from L. austromendocinus, L. elongatus, L. gununakuna (Avila et al. 2004), L. petrophilus, L. choique, L. shitan, L. capillitas, L. dicktracyi, L. heliodermis, L. talampaya, L. antumalguen (Avila et al. 2010), L. umbrifer and L. burmeisteri (Avila et al. 2012) because all of these species have a significantly larger SVL (with a maximal SVL ranging from 82 mm in L. heliodermis to 107.8 mm in L. antumalguen). In addition, all of them exhibit a lower number of scales around midbody. The only species that overlaps that number with L. carlosgarini (86–101), is L. gununakuna (84–97), but the latter has iridescent yellow body coloration, with dark transversal bars, very distinguishable from the ochre coloration and longitudinal stripes in L. carlosgarini. Finally, none of the species mentioned above exhibit the design pattern of L. carlosgarini, with longitudinal stripes. It is worth noting the striking exomorphological similarity L. carlosgarini has with L. smaug, which in addition is found relatively close to the type locality of L. carlosgarini (between Las Loicas and Volcán Peteroa provincial road, Malargüe, Mendoza, Argentina: 35 ° 39 ’51,3’’S; 70 ° 12 ’00,9’’W, 1688 m), but it differs from this species because L. smaug has a lower number of scales around midbody (73–80), in addition to a constant presence of precloacal pores in males (3–4) (Abdala 2010; Abdala in. litt. 2011). Sexual dichromatism has not been observed in L. carlosgarini as in L. smaug, nor have golden yellow specimens of L. carlosgarini been observed, but this trait deserves further analysis of live specimens. From L. cristiani it can be differentiated because L. cristiani has a very dark and pronounced melanistic stripe on the flanks (quite different from the paler stripe in L. carlosgarini), and in the absence of an occipital stripe, present in L. carlosgarini. L. cristiani is larger (mean SVL of 70.67 vs. 60.7 mm), and has fewer number of scales around midbody (83–89 vs 86–101), although these values overlap. It is differentiated from L. ceii, L. kriegi, L. buergeri, L. ramonensis, L. valdesianus and L. leopardinus because all of them are larger, the smallest being L. buergeri from Pichuante in Teno River, Chile, with a mean SVL of 73.2 mm and a maximum of 87.2 mm. All of these species have precloacal pores and differences in pattern, coloration and squamation. Nevertheless, L. leopardinus males lacking pores can be found, although the exomorpohological differences, especially in pattern, are very notorious. From L. coeruleus and L. neuquensis it can be distinguished by the blue or greenish ventral coloration in both species, and in having fewer scales around midbody. From L. thermarum, L. punmahuida, L. flavipiceus, L. tregenzai and L. riodamas it can be differentiated because all of these have a lower number of scales around midbody, larger size, and a uniform pattern, opposed to the well-defined design in L. carlosgarini. Description of the holotype. Male, with SVL of 65.35 mm. Head slightly longer than wide: 15.59 mm long (from the anterior edge of the ear opening to the point of the snout), 12.0 5 mm wide (between the anterior edges of the ear openings) and 9.91 mm high (at the level of the anterior edge of the ear openings). Snout length (from the anterior margin of the eye to the rostral scale): 4.0 2 mm. Neck slightly wider than head, effect of the transversal folds of the neck. Hind limb extended forward barely exceeds the armpit. The tail length is 1.36 times the SVL. It lacks precloacal pores. Rostral scale rectangular, 2.3 times wider than high and in contact with eight scales, including the nasals, which touch the rostral with only the anterior corner of the scale. Nasals polygonal, the nostril located posteriorly, occupying half of the scale’s surface. Two postrostrals elongated laterolaterally, followed by four internasal scales, being the two medial ones about four times larger than the side ones. Two frontal azygos, the anterior two times longer than the posterior. Six frontonasals in contact with the frontal azygos. Two prefrontal scales, and the frontal divided transversely; the anterior larger than the posterior. Four small postfrontals. Interpatietal pentagonal, posteriorly elongated, in contact with five scales. A gray pineal eye in the middle of interparietal scale. The two parietal scales are about twice the size of the interparietal. The scales in the nuchal and supratemporal regions are small, polymorphic, juxtaposed and generally smooth, with small ridges on some scales. 7 - 6 large supraoculars, accompanied by 21 - 18 small supraoculars. 6-8 very elongated and imbricate superciliary scales, excluding the canthal. Numerous scale organs on all the dorsal region of the head, increasing in numbers towards the anterior part of the head. 7 - 6 loreal scales, excluding the canthal, and a large and elongated subocular scale that spans the entire length of the eye. 7 - 7 lorilabial scales, in one row, being the sixth one the most elongated. 7 - 6 supralabials, fifth-fourth one more elongated and curved upwards posteriorly. It has a slightly elongated snout, with the loreal distance (distance between the anterior margin of the eye and the rostral scale) greater than the ocular diameter. The scales in the anterior border of the eye are elongated and overlapping, each one with a scale organ on it. 16–19 upper and 16 – 15 lower palpebrals, all squared and each one with a scale organ on it. Temporal scales rounded, smooth and subimbricate. Auricular scale is poorly developed but distinct. Tympanic scales small and highly convex. Scale organs present in all the lateral region of the head, being especially numerous in the lorilabial and loreal scales. Mental scale barely wider than rostral, but notably higher, and in contact with four scales. Five pairs of postmentals, the second pair separated by two scales. 5 - 5 infralabials, with scale organs scattered on them. Scales on the gular region rounded, smooth and imbricate. Lateral region of neck with two prominent transversal folds. Neck scales small, convex, granular and with very small granular scales in the interstitial space. Dorsal scales small, the same size as the ventral scales. They vary from rounded to subtriangular, and from juxtaposed to subimbricate. Moderate keels on the mid-dorsal scales form longitudinal lines along the dorsum, reaching the base of the tail. The keels become less prominent towards the side of the body, having already disappeared on the flanks. The scales on the flanks therefore are smooth, but in form and arrangement very similar to the dorsal scales, only being slightly smaller. Tiny granular scales are scattered in interstitial space in the dorsal region and flanks, where they are more evident. In the lateral region of the body a slight and thin fold extends longitudinally from the armpit to the groin. Ventral scales are rounded, smooth and imbricate. 95 scales around midbody. Dorsal scales of the arm are subtriangular, smooth and imbricate. Dorsal scales of forearm are very similar but more rounded, and then the more triangular shape is observed in the dorsal region of the hand. On the ventral region of the arm the scales are small, granular, juxtaposed, and surrounded by even smaller granular scales in the interstitial space. Then, the scales on the ventral region of the forearm are as large as the dorsal ones, subtriangular, imbricate, with very few granular scales in the interstitial space, and they start showing keels towards the palm, where the scales are completely keeled, triangular, imbricate and with a jagged edge. The third finger of the left hand has 20 rectangular and transversally disposed lamellae, each one provided with three keels. The scales on the dorsal femoral region are subtriangular, smooth and imbricate. In the dorsaltibial region they are more rounded, subimbricate, slightly keeled and with visible tiny granular scales in the intertstitial region. In the dorsal region of the foot the scales are subtriangular, smooth and imbricate, more similar to the scales on the femoral region rather than those on the tibial region. The scales on the ventral femoral and tibial regions are rounded, smooth and imbricate. In the plantar region scales are triangular, keeled and imbricate. Some scales in the plantar region are slightly jagged on the edges. The fourth toe has 26 rectangular and transversally disposed lamellae, each one provided with three keels. The dorsal scales on the tail are quadrangular, imbricate and keeled, with a slight mucron. Ventral scales on the tail are triangular, smooth and imbricate. Color and pattern in preservative. The background dorsal coloration is ochre, with tiny black spots scattered in the dorsal region of the head and limbs. A well-defined black occipital stripe, although with approximately only half of the scales on the stripe being melanistic. This stripe reaches the base of the tail, and from there it continues as a thin line of tiny black spots longitudinally elongated. On the sides of the occipital stripe follows a pair of parietal stripe shaving the background ochre coloration, and then, in the flanks a pair of melanistic temporal stripes, which originate at the posterior edge of the eye, and end at the groin. The back of the body exhibits white scales evenly scattered. Ventral coloration is whitish gray, with dark and inconspicuous dots in the gular region. Variation in the paratypes. Body measurements were divided into males and females (holotype is included within males since there are only two males in the whole sample): Mean and extreme values, in mm, for the males (two specimens) are: snout-vent length: 63.3 (61.25–65.35); axilla-groin distance: 28.36 (27.59–29.12); left forelimb length: 23.41 (21.74–25.07); left hindlimb length: 38.92 (37.23–40.6); tail length: 100 (89–111); head length: 15.13 (14.66–15.59); head width: 12.18 (12.05–12.3); and head height: 9.37 (8.82–9.91). For the females (six specimens) the measurements are: snout-vent length: 59.83 (53.46–68.8); axilla-groin distance: 26.12 (22.65–29.65); left forelimb length: 23.28 (22.65–25.11); left hindlimb length: 36.19 (32.53–39.16); tail length: 100 (85–113); head length: 13.32 (12.82–14.7); head width: 10.23 (9.93–11.25); and head height: 7.29 (6.55–7.54). Therefore we assume there is no evidence of sexual dimorphism, even though we measured only two males, the ranges of males measurements are within the range of the females. Nevertheless, a slightly larger size in the head measurements in males in relation to measurements in females is seen. Qualitatively the two males have a slightly more robust appearance than the six females. In the two males, the mean and extreme values of the number of scales around midbody are 92.5 (90–95). None has precloacal pores. The male paratype exhibits the following variation in squamation with respect to the holotype: Three postfrontal scales instead of four, interparietal in contact with six scales instead of five, 5 - 5 large supraoculars instead of 7 -6, 18- 16 small supraoculars instead of 21 - 18. 6 - 7 lorilabials instead of 7 - 7. Slightly more rounded dorsal scales. 21 lamellae on the third finger of the left hand instead of 20. Scales in the dorsal femoral region are rounded and slightly keeled. Plantar scales do not have a jagged edge. 27 lamellae on the fourth toe of the left foot instead of 26. In the females, the mean and extreme values of the number of scales around midbody are 92.6 (86–95). None has precloacal pores. In comparison with the holotype the females exhibit the following variation: longitudinal neck fold instead of two transversal folds in one specimen. Rostral scale in contact with six scales, and without contact with the nasals in two specimens. Undivided frontal scale in one specimen. Three postfrontals in three specimens, and two in three specimens. 4 to 6 large supraocular and 13 to 20 small supraoculars. 6 to 9 lorilabials. 5 to 8 supralabials. 4 to 6 infralabials. Four pairs of postmentals in two specimens. Dorsal scales rounded, imbricate and without visible small granular scales in the interstitial region in two specimens, and triangular, imbricate and without visible small granular scales in the interstitial region in one specimen. Dorsal scales of the forearm very slightly keeled in one specimen. Ventral scales of the forearm rounded and without visible small granular scales in the interstitial region in four specimens, and completely smooth in one specimen. Scales in the palmar region without a jagged edge in two specimens. 20 to 23 lamellae on the third finger of the left hand. Dorsal femoral scales rounded in one specimen, and slightly keeled in another specimen. Dorsal tibial scales triangular in one specimen, and without visible small granular scales in the interstitial region in four specimens. Plantar scales with a strongly jagged edge in one specimen. 27 to 30 lamellae on the fourth toe of the left foot. There is no evidence of sexual dichromatism in the preserved specimens. This could not be inferred from the pictures because the sex of the specimens photographed is unknown. In some of the specimens the melanistic occipital stripe is weaker, consisting of only a few black scales between the ochre ones. In the specimens with a more pronounced and filled stripe, little black dots are scattered in the background colored parietal stripes. One of the female paratypes exhibits the whole ventral and gular regions mottled with black dots, while in the rest of the sample, this trait is very weak, or simply not present. Osteological description. Based on specimen MNHNCL- 4560, female, double-stained skeleton. (Figure 5) Cranium features: cranium length 15.26 mm (from occipital condyle to premaxilla); cranium width 9.11 mm (between sutures of jugal and maxilla); cranium height 5.0 9 mm (the highest part of the calvarium); ocular orbit 5.55 (from the lacrimal bone to the postorbital one); rostrum 5.27 mm (from the lacrimal to the premaxilla). Rounded premaxilla with lateral process, maximum width of premaxilla half of its length. Two foramina are evident, these are the passages for the medial ethmoidal nerves (Oelrich 1956). These foramina are limited to the premaxilla and the maxilla. Nasal process of the premaxilla is acutely projected backward, lying beneath the divergence of the nasal bones. Ventrally, the premaxilla bears six acute teeth, chiseled, and caniniform on their free tip. Posterior margins of premaxilla are projected forward, almost horizontally. The premaxilla is strongly sutured to the vomer bones. Nasal capsules (=fenestra exonarina) with medial margins limited by the premaxillar spine, the floor is formed by the septomaxilla and the premaxillar process of the maxilla. The roof is formed by a cartilage. The nasal capsule is pierced and communicates with the fenestra exochoanalis. Paired nasal bones, twice as long than wide, medially sutured, anteriorly they cover the caudal tip of the premaxillar spine as described. Laterally the nasals are sutured to the ascendant process of the maxilla, and solidly to the prefrontal bones. Nasals exhibit small foramina, scattered, variable in diameter. Caudally the nasals diverge to embrace the anterior tip of the frontal bone, where they are solidly sutured. The divergence is located at the anterior orbital level. Dorsal surface of nasal bones appear wrinkled, with depressions and scars of the scales on them. Paired and triangular prefrontal bones; medially they are sutured to the nasal bones; their caudal tips are sutured to the single frontal bone. Prefrontals are sutured to the ascendant process of the maxilla and to the tiny lachrymal bone. Prefrontals are abruptly deflected downward forming the descendant process of the prefrontals. This process is strongly sutured to the palatine bones, forming the anterior wall of the ocular capsule. The lachrymal foramen is completely limited by the descendant process of the prefrontals. Its external wall is completely formed by the lachrymal bone. The frontal bone is an unpaired, flat and tubular bone. The anterior process is sutured to the nasal and prefrontal bones. The frontal is the superior margin of the ocular orbits. Backwards it becomes broadly wider and it sutures to the parietal bone. At least in this specimen, the pineal foramen is completely included in the frontal. At both sides of that foramen there are irregular windows. The external borders of the frontal are sutured to the small postfrontal bones. Ventrally, the frontal has the olfactory channel. The postfrontals are paired bones, bridging the frontal and postorbital bones. The parietal bone is a single one, broad, and widely sutured to the frontal. By its anterior and external process, the parietal is sutured to the postfrontal and postorbital bones. Laterally, the parietal is deflected downward; the epipterygoid bones reach these deflections. The parietal is projected towards caudal and lateral. Those projections are the supratemporal processes. These processes are sutured to the squamosal bones, to the tail of the supratemporal, and to the paraoccipital and exoccipital bones. The caudal portion of the supratemporal bones is surrounded externally by the squamosal and quadrate bones and medially by the paraoccipital and exoccipital processes. The parietal has a notch beneath the supratemporal processes, in which is houses the anterior portion of the supratemporal bones. Supratemporal fenestra measures 3.83 mm in anteroposterior axis. Squamosal bones are paired, stick shaped, curved and with an anterior process weakly joined to the postorbital bones. Posteriad they become broader like an anchor. This caudal portion has a process inserted in a notch of the quadrate. The superior process is sutured to the supratemporal process of the parietal. Quadrates are paired, auricular-shaped bone. Ventrally they have sulcated condyles, supporting the mandibular set of bones. Dorsally they have the cephalic condyles. These condyles have a notch, and in it, is inserted the caudal portion of the squamosal bones, which are joined by cartilaginous fibers. There is a posterior crest rising from the mandibular condyle upwards. On the inferior and internal area there is a union with the posterior valves of the pterygoid bones. Postorbitals are flattened paired bones, with three processes. The anterior one is sutured to the jugal bone, the posterior process is connected to the anterior process of the squamosal bone and the medial process is projected inward, joining the parietal and postfrontal bones. Jugals are paired bones, long and curved, slanting downward and forming the inferior margin of the ocular orbit. On their lowest part they are deflected forward and then upward riding on the maxilla, to which they are sutured, reaching the lachrymal bone. On their external surface, the jugals are pierced. On their lowest part, they are strongly sutured to the ectopterygoid. Maxillae are paired bones, with an anterior process sutured to the premaxilla with a medial process. The ascendant process sutures to the nasal, the prefrontal and the lachrymal ones. The posterior process sutures to the jugal as described. The maxillae bear 18 pleurodont teeth, housed in a basin, the crista dentalis. Teeth 1 to 13 have tricuspidated crowns; 14 to 18 are strongly acute and caniniform. Anteriad, the maxillae are sutured by a little extension to the vomerian bones. Ventrally, the maxilla forms the outer margin of the fenestra exochoanalis, a broad arch. It features a rounded palatal process, which is sutured to the palate. Posteriorly, the maxilla forms the outer margin of the infraorbital fenestra, completed externally by the ectopterigoyd; the internal margins of the infraorbital fenestra are formed by the pterygoid and palatine bones. In dorsal view, the ectopterigoid is prolonged via an acute anterior process that reaches the level of tooth 5. Paired vomers strongly sutured to each other. Anteriorly they are sutured to the premaxilla, and then to the palatine to which they bind obliquely. The vomer bones form the medial margin of the fenestra exochoanalis. Between them there is a shall