Search

This is an open access article distributed under the terms of the Creative Commons Attribution License [Attribution 4.0 International (CC BY 4.0): https://creativecommons.org/licenses/by/4.0/], which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. The offcial and authorized publication credit sources, which will be duly enforced, areas follows: offcial journal title Amphibian & Reptile Conservation; offcial journal website: amphibian-reptile-conservation.org., [EN] Integrative evidence of several external morphological characters and molecular phylogenetic analyses of mitochondrial DNA (12S, cyt-b) are used to place a new species of Andean lizard of the genus Liolaemus (Iguania: Liolaemidae) in the Liolaemus montanus group and as sister group of the clade formed by L. signifer. The new species is characterized by a unique combination of morphometric characteristics, scalation, and color pattern. The L. montanus group now contains seventeen species in southern Peru, distributed along the eastern and western slopes of the Andes., [ES] Utilizamos evidencia integradora de varios caracteres morfológicos externos y análisis filogenéticos moleculares de ADN mitocondrial (12S, cyt-b) que ubican una nueva especie del género Liolaemus (Iguania: Liolaemidae) en el grupo de Liolaemus montanus y como grupo hermano del clado formado por L. signifer. La nueva especie se caracteriza por una combinación única de patrón morfométrico, escamación y color. El grupo montanus del género Liolaemus en Perú contiene diecisiete especies, distribuidas a lo largo de la vertiente oriental y occidental de los Andes en el sur del país.

By courtesy of Heinz Grillitsch (Natural History Museum of Vienna) we reviewed photographs of the three types of L. stolzmanni. We examined one not type specimen and take photographs of a live individual, being to our knowledge all specimen currently placed in herpetological collections and the only and first photographic record of a live individual of this species. By courtesy of Cesar Aguilar (Museo de Historia Natural de San Marcos) we reviewed photographs of one paratype of L. insolitus Cei & P��faur 1981. We also examined five adult specimens of L. poconchilensis Valladares 2004, 10 adult specimens of L. reichei and six adult specimens of L. torresi (N����ez, Navarro, Gar��n, Pincheira-Donoso & Meriggio 2003). All examined specimens are listed in the Appendix I. Additional data for L. insolitus was taken from description (Cei & P��faur 1981). Characters were taken according to Langstroth (2011) and Valladares-Fa��ndez et al. (2018), but supralabial scales were counted in two different ways: between the rostral scale and the mouth��s rictal and between the rostral scale to the posterior level of the subocular scale. Lorilabial scales were counted between the rostral scale (above the supralabials) to the posterior level of the subocular scale. Scales were examined under magnification lenses. Midbody scales count of L. stolzmanni types was taken from Steindachner (1891). Historical evidence was taken from the published notes of the collector of C. stolzmanni types (Stolzmann 1912, in Polish). In regards to the type locality, although Langstroth (2011) brings light on several aspect of L. stolzmanni taxonomy, we partially disagree about his statements on the L. stolzmanni type locality. First, we agree in that northern Peru can be discarded because no Liolaemus occurs there (Langstroth 2011). But we think that there is no need to suggest that Stolzmann acquired the type specimens as gifts or purchased because he carry out field campaigns to the northern Chile (southern Bolivia and Peru at 1875) and in the current southern Peru, in places in which species of ���prhynosaurid��� lizards have been recorded and then it can be concluded that specimens were collect by himself as was pointed out by Steindachner (1891). Jan Stolzmann went to South America two times, one (1875���1881) traveled through Chile, Bolivia and Peru and another (1882���1884) traveled through Ecuador (Mierzwa-Szymkowiak & Breure 2017). This second travel can be discarded because no Liolaemus occurs in Ecuador (Abdala & Quinteros 2014). Stolzmann (1912) described in detail his first South American travel. He declares that arrived to the Valpara��so port (Chile) from Warsaw in 1875 and after that he traveled by sea to the Antofagasta port (currently Chile but formerly Bolivia) touching land in October 26 (p. 27). It is not stated how many days he was in Antofagasta, but he provides a detailed description of geography and industrial activities (smelting). Remarkably, Stolzmann (1912) clarify as footnote in the page 28, that Antofagasta port is placed in Bolivia (currently in Chile), and stated (in Spanish) that it was formerly known as ���Alto Peru��� (translated as High Peru in English or Hoch Peru in German), the type locality of L. stolzmanni according to Steindachner (1891). Later, he traveled by sea to Arica port (currently Chile but formerly Peru) performing a basic description of the environment. However, Stolzmann (1912, p. 29) stated that he was in Arica only few hours, and then the collection of Liolaemus seems to be unlikely. Finally, he traveled by sea to Mollendo port in Peru, noted as Islay in the map by Stolzmann (1912), and go to Lima port (Peru), where he began his journey by northern Peru (Fig. 1). At this point, it is clear that the type locality of L. stolzmanni is in the surroundings of Antofagasta, because the reference to ���Alto Peru��� (High Peru) made by the collector (Stolzmann 1912) and the mention of Hoch Peru (High Peru) as the type locality by the species descriptor (Steindachner 1891). In the aim of confirm this, we carried out a field campaign to the surroundings of Antofagasta in Marsh 7���8, 2018, and find one specimen assignable to L. stolzmanni in the desert near Mejillones (23��05`S ��� 70��21`W) at 60 km NE from Antofagasta (Figs. 2, 3) and additionally we reviewed a another specimen of L. stolzmanni from Hornitos, MNHNCL 4620 (22��56`S ��� 70��18`W, N����ez et al. 1998; N����ez & G��lvez 2015) at 75 km NE from Antofagasta. We propose to restrict the type locality of L. stolzmanni to transect between Antofagasta and Mejillones, Chile, because Mejillones is the record most geographically closer to Antofagasta. We remark that at date, this species has been not recorded in Antofagasta itself, then, probably Stolzmann collected L. stolzmanni in an excursion carry out from Antofagasta. Moreover, we discard that Stolzmann`s collection of L. stolzmanni specimens was made in other place of his field trip, because although there are three places of his journey in which ���prhynosaurid��� lizards can be found, all them can be discarded: 1. Although L. torresi occurs in Mantos Blancos at 35 NE km from Antofagasta (Riveros-Riffo & Torres-Mura 2015), it can be easily differentiated from L. stolzmanni, because L. torresi has more scales around midbody (60���68) than L. stolzmanni (approximately 50 on the types, Steindachner 1891; 45 on MNHNCL 4620); and L. torresi has the tail larger than the body, whereas the tail is shorter than the body on L. stolzmanni. 2. Although L. poconchilensis, occurs in Poconchile, 25 E km from Arica (Valladares 2004), both can be easily differentiated because L. poconchilensis has more scales around midbody (56���62) than L. stolzmanni, and has bluish spots dispersed on the dorsum, whereas this trait is absent in L. stolzmanni. 3. Although L. insolitus occurs in the surroundings of Mollendo (Cei & P��faur 1981), females of L. insolitus have precloacal pores, whereas the female paralectotype of L. stolzmanni has no precloacal pores. Moreover, as was noted by Langstroth (2011), both species have different color pattern, because L. insolitus has series of large dorsal dark patches with lighter shades at the center and bluish spots dispersed on the dorsum, whereas L. stolzmanni has series of small dorsal dark spots without lighter shades at the center and without bluish spots dispersed on the dorsum. In regards to L. reichei we found that both species are very similar and can be differentiated mainly by the snout vent length (SVL). Whereas L. stolzmanni has a maximum SVL = 57.0 mm (n = 4, all known specimens), L. reichei has a maximum SVL = 50.8 mm (n = 3, Valladares-Fa��ndez et al. 2018; n = 10, our data). Two hypotheses arise: 1. Both species are cryptic and can be diagnosed based solely in their sizes or, 2. Both species are conspecific and their different sizes can be explained by a geographic variation, then, L. stolzmanni is a senior synonym. This should be tested through DNA phylogenetic analysis in a further study, but for the moment, and given that both species are allopatric with more than 360 km of distance between type localities, we recommend keep both as valid species to maintain the taxonomic stability as was proposed by Valladares-Fa��ndez et al. (2018). On the other hand, we did not found differences in the dorsal color pattern between L. stolzmanni and L. reichei. Both have light brown dorsal color with 5���7 series of small dorsal black spots without lighter shades at the center, running from the shoulders to the base of the tail, with yellowish or orange inconspicuous transversal stripes between these, and absence of bluish spots dispersed on the dorsum (Fig. 4). We disagree with Valladares-Fa��ndez et al. (2018) in regards to some of the characters that they proposed as diagnostic between these species: 1. According to Valladares-Fa��ndez et al. (2018) both species differ on the number of supralabials scales (9 on L. reichei vs 6 on L. stolzmanni). Apparently, this difference is due to the use of different methodologies for the scale counts. The neotype of L. reichei has nine supralabials between the rostral and the mouth��s rictal whereas has six between the rostral and the posterior level of the eye (Valladares-Fa��ndez et al. 2018, Figure 3B). Besides, there are 9���10 supralabials between the rostral and the mouth rictal on L. stolzmanni and six between the rostral and the posterior level of the eye; in similar way, there are 8���10 and six in our sample of L. reichei; then this feature cannot be considered as diagnostic (Fig. 5) because both species overlap these scale counts if the supralabials are counted in the same way. 2. Specimens L. reichei reviewed by Valladares-Fa��ndez et al. (2018) have the interparietal and both parietal scales large and well formed, as in our sample of L. reichei. However, Valladares-Fa��ndez et al. (2018) point out that these scales are relatively small, fragmented and irregular in L. stolzmanni, but it is not observed in none of the three type specimens of L. stolzmanni, and then it is not a diagnostic character (Fig. 5). 3. According to Valladares-Fa��ndez et al. (2018) both species differ in the number of lorilabials scales (eight in L. reichei vs five in L. stolzmanni). We found 7���8 lorilabials counted between the rostral and the level of the posterior border of the subocular scale in L. stolzmanni (not counted in NMW 13580:3) and the same variation was found in our sample of L. reichei, so this character cannot be stated as diagnostic (Fig. 5). 4. According to Valladares-Fa��ndez et al. (2018) the ventral side is strongly variegated on L. reichei while is white, immaculate or lightly variegated in L. stolzmanni. This character cannot be considered as diagnostic because we found specimens of L. reichei with whitish immaculate ventral color. 5. According to Valladares-Fa��ndez et al. (2018) the lateral nuchal skin folds is well-developed and complex in L. reichei whereas this feature is not well developed on L. stolzmanni. However, our sample of L. reichei do not shows a difference for this feature in regards to L. stolzmanni., Published as part of Troncoso-Palacios, Jaime & Escobar-Gimpel, V��ctor, 2020, On the taxonomy of the desert lizard Liolaemus stolzmanni (Steindachner, 1891) A third point of view (Squamata: Liolaemidae), pp. 138-144 in Zootaxa 4763 (1) on pages 139-141, DOI: 10.11646/zootaxa.4763.1.12, http://zenodo.org/record/3974752, {"references":["Valladares, J. P. (2004) Nueva especie de lagarto del genero Liolaemus (Reptilia: Liolaemidae) del norte de Chile, previamente confundido con Liolaemus (= Phrynosaura) reichei. Cuadernos de Herpetologia, 18, 41 - 51.","Langstroth, R. P. (2011) On the species identities of a complex Liolaemus fauna from the Altiplano and Atacama Desert: insights on Liolaemus stolzmanni, L. reichei, L. jamesi pachecoi, and L. poconchilensis (Squamata: Liolaemidae). Zootaxa, 2809, 20 - 32. https: // doi. org / 10.11646 / zootaxa. 2809.1.2","Valladares-Faundez, J. P., Etheridge, R. & Abdala, C. S. (2018) Resurrection and redescription of Liolaemus reichei, proposal of a neotype to stabilize its taxonomy. Revista Mexicana de Biodiversidad, 89, 393 - 401. http: // dx. doi. org / 10.22201 / ib. 20078706 e. 2018.2.2086","Steindachner, F. (1891) Uber einige neue und seltene Reptilien- und Amphibien- arten. Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften, Mathematisch-Naturwissenschaftliche Classe, 100, 289 - 313.","YDiaquez, E. (1810) Mapa de las posesiones territoriales del Alto y Bajo Peru en 1810. [map]","Mierzwa-Szymkowiak, D. & Breure, A. S. H. (2017) Inside and outside the Neotropics: three Polish naturalists during the late nineteenth and early twentieth centuries. Archives of natural history, 44, 151 - 158.","Abdala, C. S. & Quinteros, A. S. (2014) Los ultimos 30 anos de estudios de la familia de lagartijas mas diversa de Argentina. Actualizacion taxonomica y sistematica de Liolaemidae. Cuadernos de Herpetologia, 28, 55 - 82.","Nunez, H., Navarro, J., Sufan, J. & Galaz, J. L. (1998) Distribucion geografica de Phrynosaura (Reptilia, Tropiduridae). Noticiario Mensual del Museo Nacional de Historia Natural de Chile, 334, 11 - 14.","Nunez, H. & Galvez, O. (2015) Catalogo de la Coleccion Herpetologica del Museo Nacional de Historia Natural y nomenclator basado en la Coleccion. Publicacion Ocasional del Museo Nacional de Historia Natural de Chile, 64, 1 - 203.","Riveros-Riffo, E. & Torres-Mura, J. C. (2015) Distribucion de Liolaemus torresi (Squamata: Liolaemidae), lagarto endemico del desierto de Chile. Gayana, 79, 201 - 222. http: // dx. doi. org / 10.4067 / S 0717 - 65382015000200012"]}

Phymaturus loboi sp. nov. (Fig. 5) 1966 Phymaturus palluma palluma (in part), Donoso-Barros, Rept. Chile: 349. 2005 Phymaturus flagellifer (in part), Pincheira-Donoso & N����ez, Pub. Ocas. Mus. Nac. Hist. Nat. 59: 24. 2005 Phymaturus cf. palluma (in part), Lobo & Quinteros, Pap. Avul. Zool. 45: 155. 2007 Phymaturus cf. palluma (in part), Lobo & Abdala, Cuad. Herpetol. 21: 113. 2012 Phymaturus maulense (in part), Troncoso-Palacios & Lobo, Cuad. Herpetol. 26: 73. 2013 Phymaturus maulense, Esquerr��, N����ez & Scolaro, Zootaxa 3619: 439. 2013 Phymaturus maulense (in part), Troncoso-Palacios, Lobo, Etheridge, Acosta & Laspiur, Zootaxa 3652: 561. 2013 Phymaturus cf. palluma, Lobo & Etheridge, South Am. J. Herpetol. 8: 100. 2013 Phymaturus maulense, Garin, Troncoso-Palacios, Ferri-Y����ez & Lobos. Cuad. Herpetol. 27: 67. 2013 Phymaturus sp. 2 Morando, Avila, P��rez, Hawkins, Sites, Mol. Phyl. Evol. 66: 701. 2013 Phymaturus sp. 6 Morando, Avila, P��rez, Hawkins, Sites, Mol. Phyl. Evol. 66: 701. 2016 Phymaturus maulense Lobo, Barrasso, Hibbard & Basso. Zool. J. Linn. Soc. 176: 649. 2016 Phymaturus sp. 6 Lobo, Barrasso, Hibbard & Basso. Zool. J. Linn. Soc. 176: 653. Holotype. SSUC Re 691. Adult male. Collected at National Route CH 115, 9 km NW of Maule Lagoon (35˚57' S, 70˚34' W), 1900 m asl, Maule Region, Chile by J. Troncoso-Palacios, A. Laspiur and R. G��mez. January 25, 2013 (Fig. 5a,b). Paratypes. SSUC Re 692, male and 693, 694 females (Fig. 5). Same data as the holotype. SSUC Re 410���11 (male and juvenile male). Collected road to Maule Lagoon, Maule Region, Chile, by F. Ferri-Y����ez. November 11, 2011 Other specimens referred. MNHNCL 2353, 2460���61, males. Ba��os del Campanario, 1500 m asl, San Clemente, Maule Region, Chile. J.C. Torres-Mura coll. February 2-3, 1992. Etymology. The species is named after our colleague, the herpetologist Fernando Lobo, a prominent specialist in the Phymaturus genus, who has participated in the description of 16 species in this genus (Lobo and Quinteros 2005, Lobo and Abdala 2007, Lobo and Nenda 2015, Lobo et al. 2010, 2012b, 2012c, 2013, Troncoso-Palacios and Lobo 2012) and the identification of the true type locality and redescription of P. palluma (Lobo and Etheridge 2013). The suggested English common name for this species is "Lobo���s Rocky Lizard" and in Spanish, "Matuasto de Lobo���. Diagnosis. Phymaturus loboi belongs to the palluma group (Etheridge 1995) because it has short, nonimbricate superciliary scales; fragmented subocular scale; and strongly spiny tail scales. Within the palluma group, P. loboi belongs to the P. vociferator clade (Figs. 1 and 2) and differs from the species within this clade in the following characteristics. Phymaturus loboi differs from P. dorsimaculatus, in having higher number of dorsal scales (161.0��7.5 vs. 135.0��4.0, Tables 4 and 6), and a higher number of scales around the interparietal (9.1��1.2 vs. 6.2��2.0, Tables 4 and 6). Moreover, the males of P. loboi have green spotted shoulders and the area behind the shoulders, with an inconspicuous scapular spot (Fig 5), whereas the males of P. dorsimaculatus have black color on the shoulder and in the area behind the shoulder with a bold scapular spot. Differs from P. vociferator, in having higher number of scales around the interparietal scale (9.1��1.2 vs 7.2��1.2, Tables 4 and 6). Males of P. loboi have a green spotted shoulder with an inconspicuous scapular spot (Fig. 5) whereas the males of P. vociferator have black color on the shoulders and in the area behind the shoulders, and a marked scapular spot. Additionally, P. loboi does not vocalize. Differs from P. maulense, in having higher number of ventral scales (184.0��7.6 vs 169.8.0��7.7, Tables 4 and 6). Males of P. loboi have an inconspicuous scapular spot and females have no precloacal pores, whereas the males of P. maulense have a marked scapular spot and all females have precloacal pores. Differs from Phymaturus damasense, in having higher number of ventral scales (184.0��7.6 vs 166.8��6.5, Tables 4 and 6) and fewer subocular scales (2.0��0.9 vs 3.6��0.5, Tables 4 and 6). Additionally, females of P. loboi never have precloacal pores, while 40% of the females of P. damasense have it. Description of the holotype. SSUC Re 691. Adult male. Measurements in mm: snout���vent length (SVL): 102.5; trunk width: 41.7; head length: 20.7; head width: 21.1; head height: 10.9; interorbital distance: 8.4; distance between the nares: 3.9; distance between the naris and the eye: 4.9; distance between rostral and interparietal scale: 12.7; axilla-groin distance: 51.7; arm length (from tip of toe of the 3rd toe to elbow): 29.7; right foot length: 24.1 (from tip of toe of the 4th toe to heel); and tail length: 105.8 (with autotomized tip). Ten scales in contact with interparietal scale; 16 scales between interparietal and rostral (excluding both). Parietal eye inconspicuous and whitish. Supraorbital semicircles incomplete in both sides. Superciliary scales, 12/12 (left/right) juxtaposed and flat. Subocular scales, two. Posterior subocular vertically separated from supralabials by 2 row of lorilabials. Preocular scale smaller than canthal. Preocular and supralabials separated by 4 loreal scales. Canthal separated from nasal by 2 scales. Nasal in contact with 8 scales and separated from rostral by 3 scales. Rostral scale undivided. Nine rhomboidal or pentagonal, slightly keeled temporal scales. Three scales slightly projected on the anterior border of auditory meatus. Eleven supralabial scales and 10 infralabial scales. Mental pentagonal, in contact with 6 scales. Fifty six gular scales, counted transversally between mouth��s commissures. Secondary chin shields separated by 2 scales. Well-developed antehumeral and neck fold. Conspicuous gular and antegular folds. No enlarged scales on posterior margin of gular fold or on chest. Dorsal scales rounded, smooth and juxtaposed. Dorsal scales (counted between the occiput and the level of the anterior border of the hindlimbs), 157. Middorsal scales larger than those on flanks. Ventral scales larger than dorsals, quadrangular, smooth, and juxtaposed. Ventral scales (counted from mental scale to the anterior margin of cloacal opening), 175. Scales around of midbody, 216. Precloacal pores, 11. Supernumerary pores, 7 (1 above and 6 below precloacal pores). No enlarged pigal scales. Scales of the dorsal surface of the femoral area smooth, rounded to pentagonal and juxtaposed or sub-imbricated. Scales of the ventral surface of the femoral area smooth, pentagonal or hexagonal, and juxtaposed. Scales of the dorsal surface of forearm rounded, smooth, and juxtaposed or sub-imbricated. Scales of the ventral surface of forearm rounded, smooth, and juxtaposed. Subdigital lamellae of fingers: I: 9, II: 14, III: 19, IV: 21 and V: 13. Subdigital lamellae of the toes: I: 9, II: 15, III: 20, VI: 24 and V: 15. Tarsal scales smooth or with 2/3 keels. Caudal scales arranged in spiny whorls, slightly keeled with projected mucrons. Color in life of the holotype. Head with black background color, spotted with green in nasal and temporal areas, without spots on the cheeks. Dorsum green, with faint black reticulation, without ocelli. Neck green with black reticulation to the level of the lower meatus (transversally), but is black with very few green spots below the level of the lower meatus (transversally). Scapular spot inconspicuous. Limbs light green, with faint black reticulation. Flanks yellowish green. Base of the tail green, the rest yellowish green (lighter than the body). Throat and chest black, extending ventrally to the insertion of the forelimbs. Abdomen with a �������� yellowish green area (limited by the chest and flanks) with scattered black spots. Center of the belly light green with no appreciable yellow shading on the cloaca. Ventral surface of the limbs light green with few black spots. Ventral surface of the tail light green. Precloacal pores intense orange. Variation. Measurements in mm. Adult males (n = 6): SVL, 102.1���102.8 (102.5 �� 0.3); head length, 20.3���21.8 (20.9 �� 0.8); head width, 20.3���21.5 (21.0 �� 0.4); head height, 10.9���13.2 (11.9 �� 0.9); axilla���groin distance, 50.9��� 51.7 (51.4 �� 0.4); arm length, 29.7���33.0 (31.2 �� 1.7); foot length, 24.1���27.5 (25.7 �� 1.8); tibia length, 14.8���17.8 (16.3 �� 1.5); distance between the interparietal and the rostral, 11.9���12.7 (12.3 �� 0.4); distance between the nares and the eye, 4.9���5.7 (5.2 �� 0.4); and distance between the nares, 3.2���3.9 (3.6 �� 0.3). Adult females (n = 3): SVL, 93.1���98.6 (96.6 �� 3.0); head length, 19.1���19.8 (19.4 �� 0.3); head width, 17.8���18.5 (18.2 �� 0.4); head height, 9.1��� 11.0 (10.1 �� 0.1); axilla���groin distance, 45.4���50.7 (48.5 �� 2.8); arm length, 28.5���30.1 (29.0 �� 0.9); foot length, 23.2���24.3 (23.8 �� 0.6); tibia length, 14.3���15.1 (14.7 �� 0.4); distance between the interparietal and the rostral, 12.3��� 12.4 (12.4 �� 0.1); distance between the nares and the eye, 5.0���5.2 (5.0 �� 0.1); and distance between the nares, 3.5��� 3.8 (3.6 �� 0.2). Variation in the meristic variables (scale count) is provided in the Table 6. Parietal eye inconspicuous, whitish and opaque. No enlarged scales on posterior margin of gular fold or on chest. In males, 8��� 11 (9.7 �� 1.5) precloacal pores, 0���6 supernumerary pores, with no enlarged postcloacal scales. Tarsal scales smooth or bearing 2���3 keels. Males exhibit the same color pattern as holotype, with variation in color shades. One male has the limbs of the same shade of the dorsum and less yellowish on the flanks than other males. Head brown in females, slightly darker than the body. Occipital area with several dark brown spot, surrounding the interparietal (one female), on the supraocular area (���O��� shaped spot), the superciliaries, the snout and temporal area (except postocular area). Scattered light brown scales on the head, between the occipital area and the snout, on the postocular area, and between the subocular and the rictal. Dorsum light brown, with a thin dark brown reticulation forming three short stripes on the neck, but not forming dark crossbars on the dorsum. A �������� shaped dark brown spot on the shoulder. Dorsolateral area is darker than the occipital area. The limbs, lateral neck and flanks are light brown with scattered dark brown spots. Tail light brown with diffuse dark brown rings. Throat variegated, and one specimen has dark brown color only to the level of the meatus. The belly and ventral surfaces of the limbs and the tail are whitish with some scattered dark brown spot (especially on the chest). Flanks with oxide color. The juvenile male (SVL = 80.4 mm) has a color pattern similar to the females, but the dorsal reticulation is thicker than in females, the dark brown spot on the shoulder is ���O��� shaped, the head is noticeable more darker than in females, the throat is black and the flanks are light brown. Distribution and natural history. In Chile Phymaturus loboi is known from the type locality and adjacent areas in the Region del Maule. This is an abundant lizard that inhabits rocky environments from the road to Maule Lagoon (35˚57' S, 70˚34' W, 1900 m asl, Fig. 6) and the Termas del Campanario (35˚55' S, 70˚35' W, 1800 m), at 3 km NW from type locality. In the type locality Phymaturus loboi occurs syntopically with Liolaemus buergeri, L. carlosgarini and L. flavipiceus and the vegetation is dominated by Hypochaeris tenuifolia and Festuca acanthophylla. In Argentina, P. loboi is distributed near Domuyo Volcano (36˚39' S, 70˚32' W, 2200 m) at 68 km SE from type locality. At the date of capture (January), one female had one embryo and several small oocytes. Another female had several oocytes. The intestines of one individual (SSUC Re 410) contained plant remnants and a large number of nematodes of an unidentified species, probably of the genus Spauligodon or Parapharyngodon (C. Garin pers. comm.)., Published as part of Troncoso-Palacios, Jaime, Ferri-Y����ez, Francisco, Laspiur, Alejandro & Aguilar, C��sar, 2018, An updated phylogeny and morphological study of the Phymaturus vociferator clade (Iguania: Liolaemidae), pp. 447-466 in Zootaxa 4441 (3) on pages 456-460, DOI: 10.11646/zootaxa.4441.3.2, http://zenodo.org/record/1301546, {"references":["Lobo, F. & Quinteros, S. (2005) A morphological approach on the phylogenetic relationships within the genus Phymaturus (Iguania: Liolaemidae) with the description of four new species from Argentina. Papeis Avulsos de Zoologia, 45, 143 - 177. https: // doi. org / 10.1590 / S 0031 - 10492005001300001","Lobo, F. & Abdala, C. S. (2007) Descripcion de una nueva especie de Phymaturus del grupo de P. palluma de la provincia de Mendoza, Argentina. Cuadernos de Herpetologia, 21, 103 - 113.","Lobo, F. & Nenda, S. J. (2015) Discovery of two new species of Phymaturus (Iguania: Liolaemidae) from Patagonia, Argentina, and occurrence of melanism in the patagonicus group. Cuadernos de Herpetologia, 29, 5 - 25.","Lobo, F., Abdala, C. S. & Valdecantos, S. (2010) Taxonomic studies of the genus Phymaturus (Iguania: Liolaemidae): description of four new species. South American Journal of Herpetology, 5, 102 - 126. https: // doi. org / 10.2994 / 057.005.0205","Lobo, F., Nenda, S. J. & Slodki, D. (2012 b) A new lizard of Phymaturus (Iguania: Liolaemidae) from Argentina. Herpetologica, 68, 121 - 133. https: // doi. org / 10.1655 / HERPETOLOGICA-D- 11 - 00044.1","Lobo, F., Espinoza, R. E., Sanabria, E. & Quiroga, L. (2012 c) A new Phymaturus (Iguania: Liolaemidae) from the southern extreme of the Argentine puna. Copeia, 2012, 12 - 22. https: // doi. org / 10.1643 / CH- 11 - 086","Troncoso-Palacios, J. & Lobo, F. (2012) A new species of Phymaturus (Iguania: Liolaemidae) of the palluma group from Central Chile. Cuadernos de Herpetologia, 26, 69 - 78.","Etheridge, R. E. (1995) Redescription of Ctenoblepharys adspersa Tschudi, 1845, and the taxonomy of Liolaeminae (Reptilia: Squamata: Tropiduridae). American Museum Novitates, 3142, 1 - 34."]}

Liolaemus tirantii sp. nov. 1996 Liolaemus donosobarrosi Avila, 1996 Cuadernos de Herpetolog��a 9(2):109���110. 2006 Liolaemus donosobarrosi Avila et al., Biological Journal of the Linnean Society 89(2):241���275. Holotype. LJAMM-CNP 1 3464 (Figs. 6, 7), an adult male from a small unnamed sedimentary hill 2 km S Cerro Bandera, 1 km S National Road 22, 18.5 km W Cutral-C��, Zapala Department, Neuqu��n Province, Argentina (38�� 58��� 06.0��� S, 69�� 27��� 08.7��� W, 970 m), collected by J.A. Avila, I. Minoli, C. Medina and M. Kozykariski, 12 November 2010. Paratypes. LJAMM-CNP 13465 (male),13467���13469 (females) from same locality as holotype, collected by J.A. Avila, I. Minoli, C. Medina and M. Kozykariski, 12 November 2010. MLP. S 2625 (male), BYU 52427���52429 (juveniles), eastern outskirts La Amarga village, on the road to Petrified Forest, Zapala Department, Neuqu��n Province, Argentina (39��06���39.7��� S, 69��34���29.2��� W, 803 m), collected by L.J. Avila, M. Morando, D. Perez and J. Perez, 2 May 1998. LJAMM-CNP 2542 (female) from 2 km SE La Amarga village, Zapala Department, Neuqu��n Province, Argentina (39��06���39���S, 69��34���09���W), collected by L.J. Avila and M. Morando, 7 March 2000. LJAMM-CNP 10409 (female), from 2 km E La Amarga village, 11 km E junction Provincial Road 34, Cerro Atravesado, Zapala Department, Neuqu��n Province, Argentina (39��06���39.70��� S, 69��34���9.20��� W, 803 m), collected by L.J. Avila, M. Morando and D. Perez, 26 February 2008. LJAMM-CNP 11292���11293, 11339 (females), from 2 km E La Amarga village, 11 km E junction Provincial Road 34, Zapala Department, Neuqu��n Province, Argentina (39��06���39.70���S, 69��34���9.20���W, 803 m), collected by L.J. Avila, N. Feltrin, M. Nicola, and L.E. Martinez, 17 December 2008. LJAMM-CNP 11297 (female) from National Road, 1 km W junction Provincial Road 34, Cerro Atravesado, Zapala Department, Neuqu��n Province, Argentina (38��55���14���S, 69��42���13.2���W, 788 m), collected by L.J. Avila, N. Feltrin, M. Nicola, and L.E. Martinez, 17 December 2008. LJAMM-CNP 247 (male), 248 (female) from 40 km E Zapala Yacimiento Cerro Bandera, Zapala Department, Neuqu��n Province, Argentina (38��57���22.47��� S, 69��33���11.50��� W, 811 m), collected by S. Tiranti. Diagnosis. Liolaemus tirantii sp. nov. is a robust but small sized member of the clade of Liolaemus referred to as the boulengeri group by Etheridge (1995), and is a member of the Liolaemus donosobarrosi group (Avila et al. 2006, 2007, Olave et. al 2014, 2016) that includes L. cuyanus, L. donosobarrosi, L. mapuche, L. josei, L. puelche and several other potential species still undescribed. Liolaemus tirantii sp. nov. can be distinguished from all other species of its group of Liolaemus by a combination of genetic and morphological characteristics. Liolaemus tirantii sp. nov. can be differentiated from L. cuyanus by its smaller size (64.3 mm vs 92.5 mm maximum SVL), higher number of midbody scales (X = 73.5 vs X = 66.7; min.���max. 66���86 vs 60���76, Blum 2012) with overlap, higher number of dorsal scales (X = 87.2 vs 63.6; min.���max. 79���97 vs 57���74, Blum 2012) with no overlap, four scales in contact with mental vs six in L. cuyanus, lack of prescapular marks and ventral melanism, and a very different general coloration and coloration patterns. Liolaemus tirantii sp. nov. is smaller than Liolaemus mapuche (64.3 mm vs 82.7 mm maximum SVL), has higher number of dorsal scales (X = 87.2 vs 76.2; min.���max. 79���97 vs 70���86) with some overlap, only four scales in contact with mental vs six in L. cuyanus (more common state, Abdala 2002), and lack of scapular marks, ventral melanism and a very different dorsal coloration characterized by a light blue head and scattered blue scales on a light green-blue background (Abdala 2002). Liolaemus tirantii sp. nov. is smaller than L. puelche (64.3 mm vs 89 mm maximum SVL), has a higher number of midbody scales (X = 73.5 vs 70.7; min.���max. 66���86 vs 67���76) with overlap and a higher number of dorsal scales (X = 87.2 vs 76.7; min.���max. 79���97 vs 74���80) with little overlap. Liolaemus tirantii sp. nov. is slightly smaller than L. josei (64.3 mm vs 73.1 mm maximum SVL), has a higher number of midbody scales (X = 73.5 vs 69.1; min.���max. 66���86 vs 62���76), have higher number of dorsal scales (X = 87.2 vs X = 73.1; min.���max. 79���97 vs 67���81) with slight overlap, lacks the well-marked gular melanism of L. josei males as well as its more marked dorsal color pattern. Liolaemus tirantii sp. nov. has a lower average number of scales around midbody than L. donosobarrosi (X = 73.5 vs 82.1; min.���max. 66���86 vs 74���89), middorsal scales (X = 87.21 vs 90.4; min.���max. 79���97 vs 86���101), and ventral scales (X = 94.0 vs 99.5; min.���max. 87���107 vs 88���109). Dorsal color pattern in L. tirantii sp. nov. is a series of rounded to kidney-shaped transversal marks bordered of white, usually fused along the body but more frequently on the limbs insertions, giving it a ¨wavy¨ appearance, whereas L. donosobarrosi has a series of halfmoon dots surrounding white spots, only in a few cases fused to each other, usually in the dorsal neck region; L. tirantii sp. nov. lacks of the black transversal marks observed on the head sides of L. donosobarrosi. Liolaemus tirantii sp. nov. can be distinguished from L. donosobarrosi sp. 3 by its lack of a transverse yellow and brown bands along the dorsum in life, and presence of a series of rounded to kidney-shaped transversal marks bordered of white, usually fused along the body but more frequently on the limbs insertions, given it a ¨wavy¨ appearance. Description of holotype. Adult male (Fig. 6, 7) 64.1 mm snout-vent length (SVL); tail length 97.4 mm. Axilla-groin distance 30.0 mm. Head length 13.3 mm (from anterior border of tympanum to tip of snout), 11.8 mm wide (at anterior border of tympanum), 7.9 mm high (at anterior border of tympanum). Snout length 4.2 mm (posterior margin of canthal to tip of snout). Interorbital distance 5.7 mm. Eye-nostril distance 2.5 mm. Orbitauditory meatus distance 4.2 mm. Forelimb length 17.1 mm. Tibial length 11.6 mm. Foot length 16.9 mm (ankle to insertion of claw on fourth toe). Dorsal head scales something bulged, smooth, 13 between occiput at level of anterior border of tympanum to rostral, pitted with numerous scale organs in the anterior region, and reducing to a single organ, or lack, in the posterior half of the head. Rostral scale wider (2.7 mm) than high (0.9 mm). Two postrostrals, together with anterior lorilabial, separate nasal scales from rostral. Nasal scales longer than wide, irregularly hexagonal; nostril one-half length of nasal, posterior in position. Scales surrounding nasals 8���7. Four elongated internasals. Frontonasals three, irregularly pentagonal. Prefrontals 5, a small pentagonal scale in the center (0.7 mm), and two pairs scales: a pair dorso-lateral larger scales, irregularly hexagonal (1.9 mm), and a pair lateral medium-sized scales (1.4 mm), approximately pentagonal. Three frontal scale, two anterior scales and one posterior scale. Interparietal pentagonal, surrounded by 7 scales; four smaller and irregular in front and sides, two larger in back. Parietal eye evident. Parietals some bulged, irregularly shaped, largest in size to interparietal (2.0 vs. 1.2 mm). Circumorbitals: 12���13. Transversally expanded supraoculars 6���7. Smaller lateral supraoculars: 18���17. Anterior canthal wider than long, separate from nasal by one postnasals. Posterior canthal longer than wide. Loreal scales bulged (3���5). Lorilabials longer than wide (9���7). Superciliaries eight on each side, flattened and elongated, anterior five broadly overlapping dorsally. Orbit with sixteen upper on each side and 16���14 lower ciliaries. Orbit diameter 3.4 x 1.8 mm. Preocular small, unfragmented, longer than wide. Subocular scale elongated, nine times longer than wide (4.1 x 0.6 mm). A well-marked longitudinal ridge along upper margin of preocular and subocular scales. Postocular small, slightly bulged, quarter superimposed to subocular, with a marked longitudinal ridge. Palpebral scales small granular and bulged. Supralabials 6���8, convex. Temporals smooth, convex, juxtaposed with one scale organ in the tip. Anterior auriculars smaller than adjacent posterior temporals, three slightly projecting outward. Posterior auriculars small and granular. External auditory meatus conspicuous, higher (2.0 mm) than wide (1.3 mm). Lateral scales of neck granular with inflated skin. Mental scale wider (2.2 mm) than high (1.3 mm), in contact with four scales. Mental followed posteriorly by two postmentals, and two rows of four chinshields on each side. Six infralabials on each side, first on each side quadrangular, two times wider than supralabials; all others stretched, slightly smaller than supralabials. Gular scales smooth, flat, imbricate, with rounded posterior margins, with melanophores. Scales of throat between chinshields slightly juxtaposed, becoming slightly imbricate toward auditory meatus. Thirty-four gulars between tympanum openings. Infralabials separated from chinshields by one to three rows of scales. Antehumeral well developed; longitudinal neck and postauricular distinctive; gular incomplete; rictal, dorsolateral, oblique unconspicuous. Scales of dorsal neck region rhombals, imbricate, smooth. Eighty-nine dorsal scales between occiput and anterior surface of thighs. Dorsal body scales rhomboidal, imbricate, smooth to very slightly keeled towards the posterior half of body. Dorsal scales grade laterally into slightly smaller, smooth scales at midbody. Scales immediately anterior and posterior to forelimb and hindlimb insertion small, smooth, granular, and nonoverlapping. Body lateral scales grading smaller to larger at midbody. Ventral body scales rhombals, smooth, flat, imbricate, larger than dorsal scales. Seventy-one scales around midbody; scales between mental and precloacal pores 95. Scales of cloacal region about equal in size to ventral body scales. Eight conspicuous precloacal pores. Anterior suprabrachials rhomboidal, imbricate, smooth, slightly larger to dorsal body scales. Postabrachials smaller, smooth, becoming granular near axilla. Supra-antebrachials similar to suprabrachial. Infra-antebrachials rhombals, imbricate, and smooth. Supracarpals imbricated, rhomboidal, smooth. Infracarpals strongly imbricate, rhomboidal, keeled, and slightly mucronate. Subdigital lamellae with three keels, each terminating in a short mucron, numbering: I: 9, II: 14, III: 18, IV: 19, V: 9. Claws robust, curved and sharp, opaque brown. Suprafemorals as large as dorsal body scales, rhomboidal, imbricated, smooth. Postfemorals small, granular shape. Supratibial rhomboidal, imbricated, smooth, some very slightly keeled. Infrafemorals scale small, granular and smooth with enlarged and bulged scales patch, with thirty-six scales (femoral patch), mucronate. Supratarsal rhombals imbricated and smooth. Infratarsal small, rhomboidal, imbricate, smooth. Subdigital scales keeled, each terminating in 1���3 short mucrons, numbering: I: 10, II: 14, III: 18, IV: 23, V: 14. Claws robust, curved and sharp, opaque brown. Tail complete, non-regenerated. Dorsal and lateral caudal scales, quadrangular and moderately keeled. Ventral subtriangular and smooth. Coloration. In life, general dorsal background color on head, neck, trunk, tail, and limbs Robin Rufous (30); ventral background coloration Cream White (52), in both cases at sunlight. Dorsal head surface uniform with some Cream White (52)/Burnt Umber (48) spots irregularly distributed on some scales, becoming more numerous on the nuchal region. A conspicuous solid transversal line of Burnt Umber (48) color in the fourth-sixth line of neck dorsal scales. Between occiput and rump, nine transversal series of four rounded, half-moon to roughly kidneyshaped Burth Umber (48) spots (approximately 7 scales x 3 scales). Each spots bordered posteriorly by a Cream White (52) irregular spot. Dorsal spots turn into an oval shape on rump and fuse on tail forming ¨V¨ shaped marks in the first third part of the tail, with lateral and posterior borders Cream White (52). On the tail, marks fade progressively becoming irregular rings to the tip of the tail. On the lateral sides of the head, clear areas are larger and Cream White (52), with some transversal Robin Rufous (30) marks. On the limbs, Robin Rufous (30) transversal bands intermix with slightly clear bands. Gular region variegated with Mikado Brown (42) marks, mandibular area without any mark. Upper belly and lateral midtrunk with some Dusky Brown (285) speckles. Gular region, lower belly and femoral-tibial hind limbs Cream Yellow (82). All bright coloration disappears after capture, and after several years in preservative, all coloration became faded. Color in preservative. In preservative the general brightly color disappear or fade, but color pattern of whites, blacks and greys persist for several years. Variation. Table 3 summarizes the variation of males and females of the type series for some scale and morphometric characters. Precloacal pores are present only in males. In some males, a narrow Robin Rufous (30) vertebral band became apparent in life. In few individuals dorsal transversal marks connected longitudinally, becoming lines of clear dots on the dorsum. Dorsal spots shape varies, from half-moon shaped, usually in the middle of the trunk to quadrangular, usually near rump. Half-moon spots usually fused in the first in the upper scapular and first half of the trunk. Females (Fig. 8) have a Drab (19) dorsal background coloration, and dorsal spots are usually more half-moon shaped than in males, but usually the first two-three rows are wavy-fused marks bordered with White Cream (52) scales. Each spots have in front a Robin Rufous (30) area, and connecting laterally with dorsolateral spots, a Cream Color (12) zone. Ventral areas of females lack of any dark mark or speckle. A few females have a Robin Rufous (30) longitudinal dorsolateral band connecting spots between scapular region to rump. On regenerated tails, a conspicuous narrow dark dorsal line extends from the cut area to almost the tip of the tail. Etymology. To honor Sergio Igor Tiranti, an enthusiastic biologist, colleague and friend, who first discovered and collected the species in early 1990 at the type locality, and to his father Ivan Nicolas Tiranti, professor of genetics at the National University of R��o Cuarto, Argentina, who helped the first author on its doctoral studies and shared its experiences and friendship in the academic world. Geographic distribution. Liolaemus tirantii sp. nov. is known from three localities in a small area in central Neuqu��n province (Fig. 1): (1) the type locality (Fig. 9), (2) outskirts of La Amarga village, and (3) flatlands near National Road 22 and Provincial Road 34 junction, all in the Zapala Department. Scrocchi et al. (2010) includes a photography epigraph of an adult male of Liolaemus tirantii sp nov. with the name Liolaemus cf. donosobarrosi from Cutral-Co, Neuqu��n, a locality 40 East of our type locality. In the text, they said that L. cf. donosobarrosi ¨Se distribuye en el centro este de Neuqu��n, hasta Zapala y en Rio Negro fue hallada en el Cerro Polic��a, meseta de Renteria¨. Scrocchi et al. (2010) did not mention any voucher specimens from these localities and we cannot confirm above statements with any other reference material. Natural history. The holotype and the paratypes were found basking on small rocks or running between bushes. All localities are in an ecotonal area between Patagonian Steppe (Erial Patag��nico) and Southern Patagonian Monte (Roig 1998). This ecotone is an arid region in central Neuqu��n province, between 800 to 1000 m above sea level, characterized by small hills, folded sedimentary hills, small alluvial plains, basaltic plateaus, and sedimentary piedmonts plateau-shaped. Annual rain usually is lower than 200 mm and medium temperatures are between 10���12�� C. Soils are poor, deep, with very few organic matter that limits the vegetation grow and have a very high natural erosion. Vegetation is typical of ecotone of Patagonian Steppe (Erial Patag��nico) and Southern Patagonian Monte (Roig 1998); predominant vegetation formation is a low to medium shrubs steppe (bushes between 0.50 to 2.00 m). Soils are heavily eroded by rain and wind with intermixed gravel and sand areas or small stony outcrops without vegetation. Common plants are Grindelia chiloensis, Senecio filaginoides, Acantholippia seriphioides, Ephedra ochreata, Schinus roigii, Stillingia patagonica, Chuquiraga avellanedae, Larrea nitida, L. divaricata, L. cuneifolia, L. ameghinoi, Atriplex lampa, Prosopis denudans, Bougainvillea spinosa and endemic species such as Prosopis catellanosii. All sites where L. tirantii sp. nov. was collected are similar in physiography, normally small barren areas, with sandy soil mixed with gravel areas or more compacted soil. Clumps of shrub appear in areas with some degree of protection from the strong winds typical of Patagonia; these clumps are composed mainly of Prosopis denudans and smaller shrubs of Atriplex sp., Lycium chiliensis, and Chuquiraga avellanedae, all of which grow under the protection of this spiny mesquite. These clumps reach a maximum height of 1.8 m, and cover an area between two to four m2 and separated from each other by distances of two or three meters. This region is characterized by highly degraded soils (due to overgrazing or natural erosion), and shows a tendency towards desertification. Clumps seem to be the safest areas for lizards, as well as for small mammals and birds. The majority of the lizards were observed only inside or in the immediate vicinity of the spiny shrub clumps and in few cases basking on the top of stones or open areas. Usually lizards bask under sun in early morning (09:00 to 10:00) and in late afternoon (18:00 to 19:00) in the mid-summer, when they show a bimodal type of activity, but in early spring and late-summer they usually show continuous activity without a mid-day break. When days are cloudy or cold in summer they show an all-day round activity. In the type locality we collected other species of lizards living in syntopy with L. tirantii sp. nov., including L. darwinii, L. gracilis, L. gununakuna, and L. aff. mapuche, as well as a teiid lizard, Aurivela longicauda and a gecko, Homonota darwinii. In nearby localities we collected Bothrops ammodytoides, Philo, Published as part of Avila, Luciano Javier, Perez, Cristian Hern��n Fulvio, Minoli, Ignacio, Medina, Cintia Debora, Sites, Jack W. & Morando, Mariana, 2017, New species of Liolaemus (Reptilia, Squamata, Liolaemini) of the Liolaemus donosobarrosi clade from northwestern Patagonia, Neuqu��n province, Argentina, pp. 535-563 in Zootaxa 4362 (4) on pages 543-549, DOI: 10.11646/zootaxa.4362.4.4, http://zenodo.org/record/1095342, {"references":["Avila, L. J. (1996) Liolaemus donosobarrosi: Ampliacion de su distribucion geografica y primera cita para la provincia de Neuquen. Cuadernos de Herpetologia, 9, 109 - 110.","Avila, L. J., Morando, M. & Sites, J. W. Jr. (2006) Congeneric phylogeography: hypothesizing species limits and evolutionary processes in Patagonian lizards of the Liolaemus boulengeri group (Squamata: Liolaemini). Biological Journal of the Linnean Society, 89, 241 - 275. https: // doi. org / 10.1111 / j. 1095 - 8312.2006.00666. x","Olave, M., Martinez, L. E., Avila, L. J., Sites, J. W. Jr. & Morando, M. (2014) Multilocus phylogeny of the widely distributed South American lizard clade Eulaemus (Liolaemini, Liolaemus). Zoologica Scripta, 43, 323 - 337. https: // doi. org / 10.1111 / zsc. 12053","Blum, R. (2012) Morphometric analyses and intraspecific comparison of two geographically separated populations of Liolaemus cuyanus (Iguania: Liolaemidae) from Northwestern Argentina. Unpublished Master Thesis Georg-August- Universitat Gottingen. Germany, 101 pp.","Abdala, C. S. (2002) Nuevo Liolaemus (Iguania: Liolaemidae) perteneciente al grupo boulengeri de la provincia de Neuquen, Argentina. Cuadernos de Herpetologia, 16, 3 - 13.","Scrocchi, G. J., Abdala, C. S., Nori, J. & Zaher, H. (2010) Reptiles de la provincial de Rio Negro, Argentina. Fondo Editorial Rionegrino, Viedma. 252 pp.","Roig, A. F. (1998) La vegetacion de la Patagonia. In: Correa, M. N. (Ed.), Flora Patagonica. INTA, Buenos Aires, Argentina. pp. 48 - 174."]}

Liolaemus parthenos, new species urn:lsid:zoobank.org:act: 53 CC 2 C 59 - 2 E 4 D- 41 AA- 8804 - 462 B 90 ABAECC Figures 1���5; Table 1 Liolaemus boulengeri Cei (1973: 464, in part). Liolaemus boulengeri Cei and Roig (1976:71, 87, Map 5, in part). Liolaemus boulengeri Cei and Castro (1978:9, 21, Map 16, in part). Liolaemus boulengeri Cei (1986: 220���221, in part). Liolaemus boulengeri Schulte et al. (2000:79, 87; table 1; figs. 4���6). Liolaemus sp. nov. Morando et al. (2004: 845; table 1���3; figs. 2, 8). Liolaemus sp. Abdala and D��az G��mez (2006: 29; fig. 3). Liolaemus sp. 3. Abdala (2007:51, 55, 57, 60, 77; figs. 32���36). Liolaemus cf. darwinii Pincheira-Donoso et al. (2007: 32; fig. 3). Liolaemus cf. darwinii Schulte (2013: 5; fig. 1). Liolaemus sp. 3 Olave et al. (2014: 329, 331; figs. 4, 6). Holotype.��� FML 16221, adult female, Argentina, Mendoza Province, San Rafael Department, collected on the dunes next to El Nihuil Dam on Provincial Route 180, 35�� 2 ' 19.77 ''S, 68 �� 40 ' 12.60 ''W, 1305 m, C. Abdala, J. Abdala, and E. Malovini, January 2001 (Fig. 1). Paratypes.��� FML 16222 ��� 24, same data as holotype; IBAUNC 9772 ���73, 9775��� 77, Argentina, Mendoza Province, San Rafael Department, 10 to 60 km S of El Nihuil, 35 �� 7 ' 0.71 ''S, 68 �� 41 ' 4.28 ''W to 35 �� 33 ' 49.47 '' S, 68 �� 41 ' 15.14 ''W, 1400���1600 m, 25 November 1973; IBAUNC 11431 ��� 35, Argentina, Mendoza Province, San Rafael Department, Pampa del Diamante, 34 �� 54 ' 44.39 '' S, 68 �� 51 ' 38.58 ''W, 1400 m, 17 February 1975; MHNSR 78���82, Argentina, Mendoza Province, San Rafael Department, Club de Pescadores, El Nihuil, 35 �� 2 ' 10.54 ''S, 68 �� 42 ' 33.84 ''W, 1325 m, 1 January 1975. Diagnosis.��� Liolaemus parthenos is the only known triploid unisexual iguanian lizard with the karyotype 3 n = 49 (19 M + 30m). The new species is a member of the L. boulengeri group (Etheridge, 1995; Abdala, 2007), with which it shares a femoral patch of enlarged scales on the posterior thigh. The L. boulengeri group includes the following subclades (Abdala, 2007): L. anomalus, L. darwinii, L. wiegmannii, and L. melanops (= L. telsen group + L. goetschi group), each of which includes several species as described below. Within the L. boulengeri group, L. parthenos is a member of the L. darwinii group (Abdala, 2007), with which it shares the following eight synapomorphies: (1) 14���18 dorsal head scales; (2) 24���28 gulars; (3) tail length/SVL ratio: 1.40���1.65; (4) femur length/ SVL ratio: 0.16���0.20; (5) 1���3 differentiated scales along the lower half of the anterior border of the external auditory meatus; (6) weakly developed longitudinal fold; (7) dark line passing vertically through superciliaries, eye, and subocular; and (8) small prescapular spots. Liolaemus parthenos can be distinguished from species in the L. anomalus group (L. acostai, L. anomalus, L. ditadai, L. lentus, L. millcayac, L. pipanaco, and L. pseudoanomalus) because the new species has a tail that is longer than its body, a head that is longer than wide, and lacks both a palpebral ������comb������ and pterygoid teeth. Liolaemus parthenos can be distinguished from species in the L. wiegmannii group (L. arambarensis, L. azarai, L. cuyumhue, L. lutzae, L. multimaculatus, L. occipitalis, L. rabinoi, L. riojanus, L. salinicola, L. scapularis, and L. wiegmannii) because the new species has a single row of lorilabials between the subocular and the supralabials (2���3 rows in members of the L. wiegmannii group) and four scales around the mental (6 scales in the L. wiegmannii group species). The new species, can be distinguished from species of the L. goetschi group (L. camarones, L. canqueli, L. casamiquelai, L. chehuachekenk, L. cuyanus, L. dumerili, L. fitzingerii, L. goetschi, L. mapuche, L. melanops, L. morenoi, L. puelche, L. rothi, L. sagei, L. shehuen, L. tromen, and L. xanthoviridis) based on its shorter SVL (maximum SVL 63.4 mm vs. 77���106 mm for the aforementioned taxa) and considerably less ventral melanism (except L. rothi and L. sagei). The new species also lacks the cephalic melanism typical of L. canqueli and L. melanops. With respect to L. casamiquelai, L. chehuachekenk, L. fitzingerii, L. morenoi, L. tromen, and L. xanthoviridis, the new species lacks the gular and antihumeral melanic collar that is typical of the aforementioned species. Liolaemus parthenos is distinct from L. cuyanus, L. goetschi, L. josei, L. mapuche, and L. puelche because the new species has four scales in contact with the mental, whereas L. cuyanus has six scales and L. goetschi, L. josei, L. mapuche, and L. puelche have from 4���6 scales in contact with the mental. The new species differs from L. donosobarrosi because the former has fewer scales around the midbody (mean = 57.1; range = 53���61 vs. mean = 85.4; range = 79���95, respectively). Also, the new species has a distinct dorsal pattern of quadrangular paravertebral markings, which distinguishes it from L. rothi, which has irregular markings that are not quadrangular. The new species also has pre- and postscapular spots that are not present in L. rothi. Liolaemus parthenos can be distinguished from species in the L. telsen group (L. boulengeri, L. hermannunezi, L. inacayali, L. loboi, L. martorii, L. purul, L. senguer, L. sitesi, and L. tehuelche) by its lack of gular melanism and dorsolateral bands that are more well defined. The new species differs from L. telsen because the former has less gular melanism, paravertebral markings are more well defined, well-defined dorsolateral bands, and fewer middorsal scales on the trunk (counted from occiput to anterior thigh: mean = 65; range = 61���68 vs. mean = 88.8; range = 83���96, respectively). The new species also differs from L. boulengeri, L. hermannunezi, L. loboi, L. martorii, L. purul, and L. tehuelche because the former has less conspicuous scapular spots. Within the L. darwinii group, L. parthenos is the only species with strongly cuspidate coronas and distally expanded cheek teeth, as well as a unique color pattern for the group. The new species can be further distinguished from members of the L. darwinii group (L. abaucan, L. albiceps, L. calchaqui, L. chacoensis, L. cinereus, L. crepuscularis, L. darwinii, L. diaguita, L. espinozai, L. grosseorum, L. irregularis, L. koslowskyi, L. laurenti, L. lavillai, L. montanezi, L. olongasta, L. ornatus, L. pacha, L. quilmes, and L. uspallatensis) because L. parthenos has a unique pattern of large dark brown or black quadrangular or subquadrangular paravertebral and lateral markings, white dorsolateral stripes, and a white abdomen with small dark spots. Liolaemus parthenos can be further distinguished from L. albiceps, L. calchaqui, L. crepuscularis, L. irregularis, L. lavillai, and L. ornatus because females of the latter three species possess precloacal pores and adult L. albiceps and L. irregularis are considerably larger than L. parthenos (L. parthenos: maximum SVL 63.4 mm vs. 82.5���86.1 mm, respectively). Finally, L. parthenos can be differentiated from L. darwinii because the new species lacks an antehumeral arch and abdominal and femoral melanism. Description of holotype.��� Adult female (Fig. 1). SVL: 59.8 mm. Tail complete, not regenerated: 92.7 mm. Rostral surrounded by 6 scales. Nasal dorsolateral in orientation. Supraoculars: 8. Interparietal smaller than parietals. Lorilabials: 7. Supralabials: 8. Mental surrounded by 4 scales. Infralabials: 5. Temporals: 8, not keeled. Posterior teeth with expanded cusps. Anterior border of auditory meatus with one differentiated scale. Snout blunt in lateral view. Midbody scales: 58. Dorsal scales: 68 from occiput to anterior thigh, laminar, imbricate, and keeled. Lateral neck folds well developed. Gulars: 27. Ventrals: in 21 rows, imbricate, same size as dorsals. Infratarsals and infracarpals imbricate and trifid. Femoral patch well developed. No precloacal pores. Head light brown with two rows of quadrangular dark brown paravertebral marks with white posterior margins. Background of vertebral region dark brown with four lateral markings of same color. Anterior and posterior limbs same color as dorsum. Ventrally, throat variegated with dark brown or black spots; spot density increases posteriorly toward chest, abdomen, limbs, and tail. Measurements of holotype.��� Head 1.2 times longer (11.9 mm) than wide (9.6 mm). Head height: 8.2 mm. Eye diameter: 2.9 mm. Subocular: 3.7 mm. Auditory meatus height: 2.2 mm; width: 1.8 mm. Torso length: 26.8 mm; width: 12.9 mm. Thigh length: 9.9 mm. Upper arm length: 8.2 mm; width: 2.1 mm. Forearm length: 8.3 mm. Hand length: 9.2 mm. Tail length: 92.7 mm. Variation.��� Data from 70 females, unless otherwise noted. Mental in contact with four scales. Infralabials: 5���6. Lorilabials: 6���7, in one row. Interparietal smaller than parietal. Temporals: 8���12. Auditory meatus with 1���3 scales on anterior edge and 0���2 on posterior edge. Neck folds well developed. Gulars: 23���28. Head narrow, longer (mean = 12.1 mm) than wide (mean = 9.4 mm). Neck narrower than head and trunk. Body slender, SVL 36.1���63.4 mm (mean = 56.7 mm). Scales around midbody: 53���61 (mean = 57.1). Dorsal scales between occiput and anterior thigh: 61���68 (mean = 65.0). Dorsal scales laminar, keeled, and imbricate. Infratarsals and infracarpals with imbricate, trifid scales. Femoral patch with 23���30 cone-shaped, mucronate scales. Ventrals: 91���105 (mean = 97.6), larger than dorsals. Precloacal pores absent. Complete tail: 64.3���92.7 mm long (n = 48; mean = 83.3 mm); 1.4���1.6 times longer than SVL. Color variation in life.��� Dorsal head light brown or gray with small black, irregularly distributed scales. Background dorsal color of body usually light brown, but light gray specimens not uncommon. Paravertebral markings square or rectangular, arranged longitudinally to long axis of body and dark brown or dark gray with white posterior margins. Lateral markings same color and form as paravertebral markings, but smaller than paravertebrals in some specimens. Vertebral region dark gray or light brown. Prescapular or postscapular spots less well defined, generally darker than lateral markings, but sometimes indistinguishable from them. Throat, abdomen, and cloaca region white with small irregularly shaped black spots. Ventral spots usually most dense along sides of abdomen. Mental and shield scales usually dark brown, but occasionally dark green. Longitudinal line extending along lateral tail (25���50 % total tail length) splits into small black or dark brown rectangular bars. Color becomes less vibrant, but otherwise does not change appreciably after> 6 mo in preservative (70 % ethanol). Fig. 5. Etymology.��� The specific epithet parthenos is a Greek noun meaning ������virgin������ or ������maiden,������ in reference to the presumed reproductive strategy of this all-female species. Distribution.��� Liolaemus parthenos is known only from the outskirts of El Nihuil, Salinas del Diamante, and from south of El Nihuil on Provincial Route 180 near Sierra El Nevado, San Rafael Department, Mendoza Province, Argentina (1305��� 1600 m; Fig. 2). Natural history.��� Liolaemus parthenos is apparently an allfemale parthenogenetic species. Despite 24 collecting expeditions since 1995 for a total of approximately 650 combined person hours of observing and collecting this species in nature, and after examining more than 300 specimens (note only 65 specimens were collected and preserved because of restrictions imposed by collecting permits), no males were encountered. Thus, the probability of not encountering a male, if this was a bisexual species with an equal population sex ratio, would be 0.5 65 or 2.7 20, an infinitesimally small likelihood. Field observations of active L. parthenos were made from October through February, and again in April. During summer months (December to February), lizards were active only in the morning and afternoon. The shift to bimodal activity apparently helps them avoid the extreme heat of midday. In April (fall), only juveniles were active at midday, indicating seasonal or ontogenetic variation in diel activity. Body temperatures (T bs) from four adult L. parthenos (SVL = 51.8 �� 3.4 mm) collected in the late afternoon of 11 February 1995 along the sandy shore of El Nihuil reservoir averaged 34.6 �� 3.4 ��C, which is similar to the field T bs reported for other low-elevation Liolaemus lizards (Espinoza et al., 2004). Liolaemus parthenos is oviparous and reproduces in summer. Reproductively active females (well-developed eggs in oviducts) were observed in January (n = 5), and nonreproductive females were found in December and February (n = 13). However, in a different year, we found females with oviducts full of developing eggs in December (n = 11). These observations suggest that the reproductive period begins in November and lasts until late January. Liolaemus parthenos is sympatric with the lizards Aurivela longicauda (Teiidae), Diplolaemus sexcinctus and Leiosaurus bellii (Leiosauridae), and two congeneric species: L. gracilis and L. grosseorum. Several species of snakes, which are potential predators of the new species, have also been observed in the area: the dipsadine colubrid snakes Oxyrhopus rhombifer, Philodryas trilineata, and Xenodon semicinctus, as well as the viperid Bothrops ammodytoides. Cytogenetic analysis.��� Liolaemus parthenos has a triploid karyotype with 3 n = 49 chromosomes (19 macrochromosomes + 30 microchromosomes): 17 of the macrochromosomes are biarmed metacentric or submetacentric and two are acrocentric. All the microchromosomes appear to be acro- or telocentric (Fig. 3). Because of their size and morphology, the macrochromosomes clustered into four groups: (1) the two largest pairs almost equal in size, but differing in centromere position (the first metacentric and the second submetacentric); (2) pairs 3 and 4 almost equal in size and morphology (with a centric fission in the haploid homologue of pair 3); (3) pair 5 (submetacentric in two homologues and metacentric in the third); and (4) pair 6, the smallest of the macrochromosomes with metacentric morphology. Only two of the homologous regions of chromosome pair 2 show secondary constrictions and areas with NORs in the telomere of their long arms. Phylogenetic analysis.��� The trees based on Bayesian inference and maximum parsimony were topologically very similar, with sources of conflict occurring solely at nodes that were weakly supported in one or both analyses (Fig. 4). With respect to the new species, both trees strongly supported the placement of L. parthenos in the L. grosseorum clade of the L. boulengeri group: L. boulengeri group (L. chacoensis group (L. laurenti group (L. darwinii group (L. grosseorum clade)))) (Fig. 4). The L. grosseorum clade is also supported by nine morphological synapomorphies (Abdala, 2007). Within the L. grosseorum clade, L. parthenos is in a well-supported yet unresolved clade with L. darwinii (R��o Negro Province) and L. laurenti (Fig. 4 A) or nested within a weakly supported clade containing populations of L. darwinii from the more southern provinces of Mendoza and Ŕıo Negro and those occurring farther north in La Rioja Province, with L. laurenti strongly supported as sister to these four lineages (Fig. 4 B). DISCUSSION Based on previous studies (Schulte et al., 2000; Morando et al., 2004; Pincheira-Donoso et al., 2007; Schulte, 2013) and the phylogenetic hypotheses presented here (Fig. 4), L. parthenos is closely related to L. darwinii and L. laurenti, species that are morphologically typical of the L. darwinii group (Etheridge, 1993; Abdala, 2007). The closest known locality for L. laurenti is approximately 300 km to the north of the type locality of L. parthenos, whereas L. darwinii has a much wider distribution (Abdala, 2007) including a small population that is sympatric with L. parthenos at the type locality of the latter species. It is noteworthy that a similarly proximate distributional association has been reported for a parthenogenetic species of Leiolepis and its matrilineal ancestor (Grismer and Grismer, 2010; Grismer et al., 2014). A close phylogenetic association between L. parthenos and L. darwinii has been reported in previous phylogenetic analyses based on mtDNA. Schulte et al. (2000) recovered L. parthenos (as ������ L. boulengeri ������) in the clade: L. grosseorum (L. darwinii (L. parthenos (L. laurenti))). A phylogeographic analysis of the L. darwinii group conducted by Morando et al. (2004) recovered L. parthenos (as ������ L. sp. nov.������) as the sister taxon to L. laurenti + L. darwinii. Finally, Pincheira-Donoso et al. (2007) recovered L. parthenos (as L. cf. darwinii; SDSU 3469, AF099275) as the sister taxon to L. darwinii in the clade (L. grosseorum (L. darwinii (L. parthenos + L. darwinii from Chubut))). Although L. parthenos is superficially similar to L. boulengeri, these three molecular studies each strongly support the relationships among L. parthenos, L. darwinii, L. laurenti, and L. grosseorum. The most parsimonious interpretation of the triploid karyotype of L. parthenos (Fig. 5) is that the species arose via hybridization. The karyotype of L. parthenos (Fig. 4) has a clear diploid component (12 M + 20m) contributed by one parental species and an additional haploid set (7 M + 10m) provided by the other parental genome (Fig. 5). In the haploid component the two acrocentric macrochromosomes are interpreted as homologs of the two arms (3 p and 3 q) of pair 3 (3 a) in the 2 n set. The 30 microchromosomes of L. parthenos are best interpret, Published as part of Cristi��n S. Abdala, Diego Baldo, Ricardo A. Ju��rez & Robert E. Espinoza, 2016, The First Parthenogenetic Pleurodont Iguanian: A New All-female Liolaemus (Squamata: Liolaemidae) from Western Argentina, pp. 487-497 in Copeia 104 (2) on pages 489-494, DOI: 10.1643/CH-15-381, http://zenodo.org/record/269580, {"references":["Cei, J. M. 1973. Herpetologia patagonica VII. Notas ecologicas y morfologicas sobre Liolaemus bibroni [sic] y L. boulengeri (Sauria, Iguanidae). Physis 32: 459 - 469.","Cei, J. M., and V. G. Roig. 1976. Fauna y ecosistemas del oeste arido Argentino. I. Reptiles de la Provincia de Mendoza. Deserta 4: 69 - 91.","Cei, J. M., and L. P. Castro. 1978. Atlas de los vertebrados inferiores de la region de Cuyo. Publicaciones Ocasionales del Instituto de Biologia Animal, Universidad Nacional de Cuyo, Serie Cientifica 2: 1 - 38.","Cei, J. M. 1986. Reptiles del Centro, Centro-oeste y Sur de la Argentina. Museo Regionale di Scienze Naturali, Torino: Monografie 4: 1 - 527.","Schulte, J. A., II, J. R. Macey, R. E. Espinoza, and A. Larson. 2000. Phylogenetic relationships in the iguanid lizard genus Liolaemus: multiple origins of viviparous reproduction and evidence for recurring Andean vicariance and dispersal. Biological Journal of the Linnean Society 69: 75 - 102.","Morando, M., L. J. Avila, J. Baker, and J. W. Sites, Jr. 2004. Phylogeny and phylogeography of the Liolaemus darwinii complex (Squamata: Liolaemidae): evidence for introgression and incomplete lineage sorting. Evolution 58: 842 - 861.","Abdala, C. S., and J. M. Diaz Gomez´. 2006. A new species of the Liolaemus darwinii group (Iguania: Liolaemidae) from Catamarca Province, Argentina. Zootaxa 1317: 21 - 33.","Abdala, C. S. 2007. Phylogeny of the boulengeri group (Iguania: Liolaemidae, Liolaemus) based on morphological and molecular characters. Zootaxa 1538: 1 - 84.","Pincheira-Donoso, D., J. A. Scolaro, and J. A. Schulte, II. 2007. The limits of polymorphism in Liolaemus rothi: molecular and phenotypic evidence for a new species of the Liolaemus boulengeri clade (Iguanidae, Liolaemini) from boreal Patagonia of Chile. Zootaxa 1452: 25 - 42.","Schulte, J. A., II. 2013. Undersampling taxa will underestimate molecular divergence dates: an example from the South American lizard clade Liolaemini. International Journal of Evolutionary Biology 2013: 1 - 12.","Olave, M., L. J. Avila, J. W. Sites, Jr., and M. Morando. 2014. Multilocus phylogeny of the widely distributed South American lizard clade Eulaemus (Liolaemini, Liolaemus). Zoologica Scripta 43: 323 - 337.","Espinoza, R. E., J. J. Wiens, and C. R. Tracy. 2004. Recurrent evolution of herbivory in small, cold-climate lizards: breaking the ecophysiological rules of reptilian herbivory. Proceedings of the National Academy of Sciences of the United States of America 101: 16819 - 16824.","Etheridge, R. 1993. Lizards of the Liolaemus darwinii complex (Squamata: Iguania: Tropiduridae) in northern Argentina. Bollettino del Museo Regionale di Scienze Naturali, Torino 11: 137 - 199.","Grismer, J. L., and L. L. Grismer. 2010. Who's your mommy? Identifying maternal ancestors of asexual species of Leiolepis Cuvier, 1829 and the description of a new endemic species of asexual Leiolepis Cuvier, 1829 from southern Vietnam. Zootaxa 2433: 47 - 61.","Grismer, J. L., A. M. Bauer, L. L. Grismer, K. Thirakhupt, A. Aowphol, J. R. Oaks, P. L. Wood, Jr., C. K. Onn, N. Thy, M. Cota, and T. Jackman. 2014. Multiple origins of parthenogenesis, and a revised species phylogeny for the Southeast Asian butterfly lizards, Leiolepis. Biological Journal of the Linnean Society 113: 1080 - 1093.","Aiassa, D., R. Martori, and N. Gorla. 2005. Citogenetica de los lagartos del genero Liolaemus (Iguania: Liolaemidae) de A merica del sur. Cuadernos de Herpetologia 18: 23 - 36.","Vrijenhoek, R. C., R. M. Dawley, C. J. Cole, and J. P. Bogart. 1989. A list of the known unisexual vertebrates, p. 19 - 23. In: Evolution and Ecology of Unisexual Vertebrates. R. M. Dawley and J. P. Bogart (eds.). New York State Museum Bulletin 466, New York.","Kraus, F. 1995. The conservation of unisexual vertebrate populations. Conservation Biology 9: 956 - 959.","Schmidt, B. R. 1996. Sexual and asexual reproduction in vertebrates. Trends in Ecology & Evolution 11: 253 - 254.","Avise, J. C. 2008. Clonality: The Genetics, Ecology, and Evolution of Sexual Abstinence in Vertebrate Animals. Oxford University Press, New York.","Lamborot, M. 2008. Evolucion cromosomica en reptiles, p. 161 - 193. In: Herpetologia de Chile. M. A. Vidal Maldonado and A. Labra Lillo (eds.). Science Verlag, Santiago, Chile.","Neaves, W. B., and P. Baumann. 2011. Unisexual reproduction in vertebrates. Trends in Genetics 27: 81 - 88.","Lamborot, M., and M. Vasquez. 1998. A triploid lizard (Liolaemus gravenhorsti) [sic] from Chile. Journal of Herpetology 32: 617 - 620.","Lamborot, M., M. E. Manzur, and E. Alvarez-Sarret. 2006. Triploidy and mosaicism in Liolaemus chiliensis (Sauria: Tropiduridae). Genome 49: 445 - 453.","Wright, J. W., and C. H. Lowe. 1968. Weeds, polyploids, parthenogenesis, and geographical and ecological distribution of all-female species of Cnemidophorus. Copeia 1968: 128 - 138.","Darevsky, I. S., L. A. Kupriyanova, and T. Uzzell. 1985. Parthenogenesis in reptiles, p. 411 - 526. In: Biology of the Reptilia, Volume 15. C. Gans and F. Billett (eds.). J. Wiley & Sons, New York.","Beukeboom, L. W., and R. C. Vrijenhoek. 1998. Evolutionary genetics and ecology of sperm-dependent parthenogenesis. Journal of Evolutionary Biology 11: 755 - 782."]}

Liolaemus pacha sp. nov. (Figs. 2 to 4). 1843. Liolaemus darwinii, Bell, T., Smith, Elder & Co., London, pp. 1���55. 1993. Liolaemus quilmes, Etheridge, R., Boll. Mus. Reg. di Sc. Naturali, Torino, 11: 137���199. 1993. Liolaemus darwinii (partim), Cei, J. M., Mus. Reg. di Sc. Naturali, Torino, Monografie XIV, 949 pp. 1996. Liolaemus quilmes, Halloy, M., Bull. of the Maryland Herp. Soc., 32: 43���57. 1998. Liolaemus quilmes, Halloy, M., Etheridge, R. and Burghardt, G., Herpetological Monographs, 12: 1���37. 2002. Liolaemus quilmes, Halloy, M. and Robles, C., Bull. of the Maryland Herp. Soc., 38: 118���129. 2003. Liolaemus quilmes, Halloy, M. and Robles, C., Cuad. Herp., 17: 67���73. 2004. Liolaemus quilmes, Halloy, M., Acta Zool. Lilloana, 48 (1���2): 103���121. 2004. Liolaemus quilmes, Martins E. P., Labra, A., Halloy, M. and Tolman Thompson J., Animal Behaviour, 68 (3): 453���463. 2005. Liolaemus quilmes (partim), Abdala, C. S., Rev. Esp. de Herp., 19: 5���17. 2006. Liolaemus quilmes (partim), Abdala, C. S. and Lobo, F., Cuad. Herp., 19 (2): 3���18. 2006. Liolaemus quilmes (partim), Abdala, C. S. and D��az G��mez, J., Zootaxa, 1317: 21���33. 2006. Liolaemus quilmes (partim), Avila, L. J., Morando, M. and Sites, J. W. Jr., Biol. Journal of the Linnean Society, 89: 241��� 275. 2006. Liolaemus quilmes, Halloy, M., Herp. Review, 37: 88���89. 2006. Liolaemus quilmes, Halloy, M. and Castillo, M., Herp. Nat. History, 9 (2): 127���133. 2006. Liolaemus quilmes, Halloy, M., Robles, C. and Cuezzo, F., Rev. Esp. Herp., 20: 47���56. 2007. Liolaemus quilmes (partim), Abdala, C. S., Zootaxa, 1538: 1���84. 2007. Liolaemus quilmes, Halloy, M., Guerra, C. and Robles, C., Bull. of the Maryland Herp. Soc., 43: 110���118. 2008. Liolaemus quilmes, Robles, C. and Halloy, M., Cuad. de Herpetolog��a, 22: 73���79. 2009. Liolaemus quilmes, Robles, C. and Halloy, M., South Am. Journal of Herp., 4: 253���258. 2009 a. Liolaemus quilmes, Salica, M. J. and Halloy, M., Acta Zool. Lilloana, 53: 89���96. 2009 b. Liolaemus quilmes, Salica, M. J. and Halloy, M., Rev. Esp. Herp., 23: 141���149. 2010. Liolaemus quilmes, Halloy, M., Robles, C. and Salica, M. J. Anales de la Academia Nac. Ciencias, Buenos Aires, Argentina, 44: 3���16. 2010. Liolaemus quilmes, Robles, C. and Halloy, M., Herpetological Journal, 20: 243���248. 2011. Liolaemus quilmes, Salica, M. J. and Halloy, M., Herp. Review, 42 (4): 606. 2012. Liolaemus quilmes, Halloy, M., Herpetological Journal, 22 (4): 267���270. 2012. Liolaemus quilmes, Robles, C. and Halloy, M., Salamandra, 48 (2): 115���121. 2013. Liolaemus quilmes, Halloy, M., Robles, C., Salica, M. J., Semhan, R., Ju��rez Heredia, V. and Vicente, N., Cuadernos de Herpetolog��a, 27 (1): 15���26. Holotype. Fundaci��n Miguel Lillo (FML) 02448/ 1. Km 98, Ruta Provincial 307 (Los Cardones), Taf�� del Valle Department, Tucum��n Province, 26 �� 40 ��� 1.5 ��� S 65 �� 49 ��� 5.1 ��� W, datum: WGS84, 2725 m, collector, O. Pagaburo, 21 / 04/ 1999. Paratypes. FML 2448 / 3���4 / 6���9 Km 98, Ruta Provincial 307, Amaicha del Valle, Taf�� del Valle Department, Tucum��n Province. O. Pagaburo, Col. 21 /04/ 1999. Etymology. Pacha comes from the Aimara and Quechua indigenous languages from northwestern Argentina. It means ���earth��� and by extension ���world��� or ���cosmos���. In this part of the country, social gatherings called Pachamama are celebrated every year in the autumn to thank and bless the earth and its products. Diagnosis. Liolaemus pacha sp. nov., belongs to the L. boulengeri group because it has a patch of enlarged scales on the posterior region of the thigh (Etheridge 1995; Abdala et al. 2006). Within this group, it belongs to the L. laurenti group (Abdala 2007) because its posterior teeth have straight edged crowns and sexual dichromatism is evident. Within the L. laurenti group, Liolaemus pacha sp nov. is a member of the L. darwinii group (Abdala 2007) or the darwinii complex (Etheridge 1993) because it has a black line which crosses the eye vertically (except L. chacoensis) and pre-scapular spots in both males and females (Abdala 2007). Within the L. darwinii group (Abdala 2007), L. pacha differs from the members of the L. ornatus group (L. albiceps, L. calchaqui, L. crepuscularis, L. irregularis, L. lavillai, and L. ornatus) because it has an oviparous reproductive mode. It also differs from L. albiceps and L. irregularis because it is smaller (max SVL 67.9 mm vs 82.5 and 86.1 mm, respectively). Liolaemus pacha differs from L. abaucan and L. uspallatensis in having imbricate and keeled dorsal scales, whereas in those two species the dorsal scales are smooth or weakly keeled. Liolaemus chacoensis exhibits keeled temporal scales and mucronate dorsal scales, absent in L. pacha. The presence of a large prescapular spot distinguishes L. pacha from L. abaucan, L. darwinii and L. koslowskyi, which have large and evident postscapular spots. It differs from L. cinereus in having a black stripe in the lateral field and in lacking melanism on the throat. Liolaemus darwinii, L. grosseorum, L. laurenti, L. montanezi and L. olongasta have a black antehumeral arch, absent in L. pacha. Liolaemus pacha sp. nov. differs from L. quilmes mainly because of its size (see below). It has a larger prescapular spot (Fig. 5, left) than L. quilmes (Fig. 6, left) and numerous light blue scales on the sides of the body and tail (Fig. 2). It has a Y-shaped mark on the snout (Fig. 7), reduced in L. quilmes (Table 1). In L. pacha, scales on dorsum are uniformly colored (Fig. 8) whereas in L. quilmes scales on dorsum are irregularly variegated with diffuse spots (Fig. 9). Both species present alternating light and dark spots on supra and infralabials, more diffuse on infralabials in L. pacha sp. nov. (Fig. 10, top), well marked in L. quilmes (Fig. 10, bottom). Anterior limbs with lightly visible bands and posterior limbs without these (Fig. 2 and 5, left), whereas in L. quilmes both anterior and posterior limbs with visible bands (Fig. 6, left). The new species presents yellow dorsolateral spotted bands (Fig. 2), different from L. quilmes which are continuous. Liolaemus pacha sp. nov. has ventrally scattered spots in the jaw area, light yellow on the throat and thigh areas (Fig. 3 and 5, right), in L. quilmes throat with spots that reach to the neck, ventral area white (Fig. 6, right, Table 1). Moreover, among its distinctive characteristics, L. pacha sp. nov. differs significantly from 3 populations of L. quilmes in 10 characters (Table 2, P Description of the holotype. Adult male. SVL 63.1 mm. Trunk length 33.8 mm. Head longer (15.1 mm) than wide (11.5 mm). Head height 7.8 mm. Eye diameter 3.8 mm. Interorbital distance 8.6 mm. Orbit���auditory meatus distance 5.8 mm. Auditory meatus 2.5 mm high, 1.3 mm wide. Orbit���commissure of mouth distance 3.3 mm. Internares distance 2.6 mm. Subocular scale length 3.1 mm. Femur length 13.5 mm, tibia 10.4 mm, and foot 17.3 mm. Humerus length 9.4 mm. Forearm length 7.6 and hand 9.1 mm. Tail length 95.5 mm. Length of the pigal region 6.4 mm and cloaca width 8.5 mm. Dorsal surface of the head smooth, with 17 scales. Rostral wider than long, bordered by six scales. Mental larger than rostral, trapezoidal, bordered by four scales. Nasal not in contact with rostral. Nasal surrounded by six scales, separated from canthal by one scale. Five scales between frontal and supercilliaries. Five scales between frontal and rostral. Frontal not divided. Two postrostrals. Interparietal smaller than parietals, in contact with eight scales. Orbital semicircles complete. Preocular separated from lorilabial row by one scale. Three scales in anterior margin of auditory meatus. Fifteen smooth temporals. Four lorilabials in contact with subocular. Seven supralabials, none in contact with subocular. Seven supraoculars. Eight lorilabials. Six infralabials, second ventrally in contact with two scales. Seventy scales around midbody. Seventy-five round, slightly imbricated, slightly keeled dorsals from occiput to hind limbs. Twenty-four rows of scales in dorsum. Scales of flank same size and shape as dorsals. Ninety-three ventrals same size as dorsals, flat, imbricate. Thirty smooth, weakly imbricated gulars. Four precloacal pores. Antehumeral scales flat, larger than or equal in size to dorsals. Postauricular, rectal, and longitudinal folds present. Scales on the longitudinal fold granular and smooth. Fourth finger with 17 subdigital lamellae; fourth toe with 25. Infracarpals flat, imbricate, trifid; infratarsals flat, imbricate, trifid. Without gular fold. With 36 scales on the neck, from the posterior border of the ear to the shoulder, through the longitudinal fold. With femoral parch. Dorsal scales of the tail without mucro and with keel, ventrals smooth. Color in alcohol. Dorsum and sides of the head brown colored with numerous scales and small spots dark brown. Subocular light brown colored with dark spots. Lorilabials and supralabials alternate between light and dark color, infralabials with alternating diffuse spots (Fig. 10 top). Snout with diffuse inverted Y shaped mark (Fig. 7). Body dorsally with 8 series of paravertebral spots, subquadrangular shaped, with white-colored, spotted, dorsolateral bands. Paravertebral spots with white posterior elongated spots. Dorsum and sides of the body grey colored to light brown, scattered irregularly with numerous white colored scales (Fig. 5 left). Scales on dorsum, uniformly colored (Fig. 8). With white-colored spotted vertebral line. Presence of prescapular spot. With many scattered light blue spots on the sides of the body (Fig. 2). Anterior and posterior limbs light brown colored, with white spots distributed irregularly. Femur with obvious light blue spots. Tail grey to brown-colored, with elongated spots and numerous light blue spots, mainly distributed at the base. Ventrally the mandibular region is white colored with diffuse spots from infralabials (Fig. 3 and 5, right). Ventrum, anterior and posterior limbs, cloaca and tail completely white. Color in life (Table 1). Dorsum and sides of the head light chestnut brown, with dark and/or black chestnut brown scales and spots. Supralabials with alternating dark and light chestnut brown spots, infralabials with light and dark spots diffuse. Chestnut-brown inverted Y-mark, on the tip of the snout (Fig. 7). Body with 8 pairs of paravertebral spots of subquadrangular shape, black colored, with elongated posterior white spot (Fig. 5, left). With prescapular black spot. Lateral region of the body beige reddish with numerous white and light blue spots (Fig. 2). With vertebral line and dorsolateral bands discontinuous yellow colored. With diffuse subquadrandular dorsolateral spots. Neck with yellow band from above the ear to the anterior limb. Scales on the dorsum with a homogeneous colored pattern. Limbs and tail chestnut-brown colored, scattered with white, yellow light blue and dark brown scales. Arm and forearm with two diffuse rings (Figs. 2 and 5, left). Posterior members without rings. Tail with numerous light blue scales. Ventrally, the mandibular area, anterior limbs, ventrum, cloaca and tail are white (Fig. 3 and 5, right). In the mandibular region, with disseminated spots from the infralabials to the anterior region of the neck (Fig. 3 and 5, right). Gular region, sides and femur with diffuse yellow spots (Fig. 3). Obvious orange-colored precloacal pores. Variation (Table 2). Based on 25 specimens (15 males, 10 females) (Appendix 1). Head longer (10.9���15.1 mm; mean = 13.3 mm) than wide (9.1���13.4 mm; mean = 11.4 mm). Head height 6.4���9.9 mm (mean = 7.9 mm). Neck narrower than head. SVL 53.6���67.9 mm (mean = 61.2 mm), averaging 1.7 times longer than trunk. Tail length 63.9���103.8 mm (mean = 85.1 mm). Humerus length 5.9���9.7 mm (mean = 8.1 mm). Dorsal surface of head smooth, with 12���20 scales. One row of lorilabials, same size as supralabials. Seven to ten lorilabials. Five to eight supralabials. Frontal not divided. Interparietal always smaller than parietals, surrounded by 5���7 scales. Nasal surrounded by 6���9 scales. Four to seven infralabials. Mental in contact with four scales. Ten to 13 smooth round temporals. Longitudinal, postauricular, and antehumeral folds present. Horizontal fold Y-shaped between shoulder and auditory meatus. Scales around midbody 46���64 (mean = 58.5). Gulars 26���44 (mean = 31.7). Dorsal scales between occiput and hind limbs 62���77 (mean = 69.0). Ventrals 79���97 (mean = 90.2). Males with 4���7 (mean = 5.5) precloacal pores. Females with 1���2 (mean = 1.7) precloacal pores. Dorsals flat, slightly imbricate, slightly keeled. Twelve to 20 infradigital lamellae on fourth finger and 21���28 on fourth toe. CHARACTER / SPECIES L. pacha sp. nov. Animan�� El Tio Ruinas P Variation in color pattern. With clear sexual dichromatism. Background color of the head may vary, from grey to light chestnut-brown to dark. Dorsum of the head presents occasionally a greater number of irregular black spots. Laterally the head may present conspicuous yellow spots. On the sides of the head, alternating clear and dark spots, diffuse on the infralabials and well marked on the supralabials (Fig. 10, top), stand out. The inverted Yshaped mark on the snout diffuse or absent (Fig. 7). The color of the body may vary between brownish and reddish tones. Without a black antehumeral arc. With discontinuous or absent vertebral line. Paravertebral spots shaped as horseshoe with posterior outline or subquadrangulars, black colored, with posterior white spot elongated or circular shaped. Dorsal body scales with a homogeneous pattern (Fig. 8). With diffuse dorsolateral subquadrangular spots, smaller than paravertebrals and sometimes absent. White line discontinuous on the sides of the body. Conspicuous black colored prescapular spot present. Limbs vary from grey to brown. Arm and forearm with two diffuse or absent rings (Fig. 5, left). The tail is lighter colored than the dorsal region of the body, with numerous light blue scales. With discontinuous spots. Ventrally white colored occasionally the mandibular, gular and femur regions, light yellow colored. Jaw with diffuse or absent spots. Precloacal pores conspicuous orange color. Head and body of females varying from grey to dark brown. Inverted Y-spot on snout diffuse. Paravertebral lines light ochre which unite on the tail. Paravertebral and subquadrandular dorsolateral spots black with thin posterior white spot, occasionally very diffuse. Lateral white line diffuse. Laterally body with white and light blue rare or absent spots. Anterior limbs with diffuse or absent rings. Posterior limbs without rings. Laterally the neck varying between yellow to intense orange. Ventrally white colored. Precloacal pores rare, barely visible. Distribution. The new species is known from the type locality Km 98, Ruta Provincial 307 (Los Cardones), Taf�� del Valle Department, Tucum��n Province (26 �� 40 ��� 1.5 ��� S 65 �� 49 ��� 5.1 ��� W, datum: WGS 84; 2725 m). Natural history. Liolaemus pacha is found on rocky to sandy subtrates. The type locality is characterized by firm substrate, large rocks, scattered shrubs and tall cacti (Thrichocereus pasacana, Cactaceae). It corresponds to the Monte and Prepuna ecoregions (Cabrera & Willink 1980 ). They are mainly insectivorous occasionally eating small plant parts depending on the season and availability (Halloy et al. 2006). It is an oviparous species (Ramirez Pinilla 1992). Males defend territories which generally overlap with two to three females (Halloy 1996; Halloy & Robles 2002; Robles & Halloy 2009; Robles 2010). Both males and females use visual displays, such as headbobs, forelimb waves, and tail movement, during encounters as well as during courtship (Halloy 1996; Martins et al. 2004; Halloy & Castillo 2006; Halloy 2012). It is a dimorphic species (Etheridge 1993; Halloy 1996; this study), males being slightly larger and more colorful than females. Females present during the reproductive season nuptial coloration on the side of the neck, going from yellowish to intense orange (Halloy et al. 2007; Salica 2008; Salica & Halloy 2009 b). Males emerge from hibernation in September whereas females emerge one to two months later (Halloy & Robles 2003). Matings occur end of October, beginning of November. Neonates are observed by the end of December and in January (Ram��rez Pinilla 1992; pers. obs.). Males and females remain active through March and April when they return to hibernation., Published as part of Heredia, Viviana Ju��rez, Robles, Cecilia & Halloy, Monique, 2013, A new species of Liolaemus from the darwinii group (Iguania: Liolaemidae), Tucum��n province, Argentina, pp. 524-538 in Zootaxa 3681 (5) on pages 526-533, DOI: 10.11646/zootaxa.3681.5.2, http://zenodo.org/record/217441

Liolaemus shehuen sp. nov. (Figs. 4���7) 1999 Cei and Scolaro Liolaemus fitzingerii, Revue francais de Aquariologie, 26, 79- 82. 2007 Liolaemus sp 7. Abdala C. S., Zootaxa, 1538, 84 pp. Holotype. ― FML 22191. Adult male. 60 km west of Telsen, Telsen Department, Chubut Province, Argentina. C. S. Abdala, J. S. Abdala and M. Juarez cols. January 23, 2004. (Figs. 4, 5). Paratypes. ― FML 22192 - 195.4 individuals. 1 male, 3 females. Same data as holotype. FML 22204 - 205.2 individuals. 2 males, same data as holotype. FML 22217 - 219.3 individuals. 1 male, 2 females. 80 km west of Telsen, Telsen Department, Chubut Province, Argentina. C. S. Abdala, J. S. Abdala and M. Juarez cols. January 23, 2004. Diagnosis.― Liolaemus shehuen belongs to the L. boulengeri group, characterized by having a patch of enlarged scales on the posterior medial surface of the thigh (Etheridge 1995; Abdala 2007) and hypertrophy of the flexor tibialis internus muscle (Abdala et al. 2006); tail long relative to body length, head proportionally longer than wide, palpebral ���comb��� not developed, and more precloacal pores both in males and females than in the remaining species of the clade. Liolaemus shehuen differs from the species of the L. darwinii group (Table 2) in having posterior teeth with expanded crowns and different body proportions, and in lacking a transversal black line on the eye. It differs from the species of the L. wiegmannii group (Table 2) in having a row of lorilabial scales (never two or three). Liolaemus shehuen differs from L. boulengeri, L. donosobarrosi, L. goetschi, L. hermannunezi (Pincheira-Donoso, Scolaro & Schulte), L. inacayali (Abdala), L. josei (Abdala), L. loboi (Abdala), L. martorii (Abdala), L. rothi (Koslowskyi 1898) L. sagei (Etheridge & Christie), L. senguer (Abdala), L. tehuelche (Abdala), and L. telsen (Cei & Scolaro) in having an evident black antehumeral arch that is absent in the aforementioned species. The species differs from all the species of the L. telsen group, except L. rothi and L. sagei, in having longer snout-vent length (max SVL 90.8 mm vs 32.3���77.1 mm). It differs from L. mapuche (Abdala) in having different dorsal body coloration, lacking light blue scales and having more marked ventral melanism. It differs from L. cuyanus in having more marked ventral melanism and different dorsal coloration. Group Species L. anomalus L. anomalus, L. ditadai, L. lentus and L. pseudoanomalus L. abaucan, L. albiceps, L. calchaqu��, L. chacoensis, L. cinereus, L. crepuscularis, L. darwinii, L. L. darwinii diaguita, L. espinozai, L. grosseorum, L. irregularis, L. koslowskyi, L. montanezi, L. laurenti, L. lavillai, L. olongasta, L. ornatus, L. quilmes and L. uspallatensis L. boulengeri, L. inacayali, L. josei, L. loboi, L. martorii, L. puelche, L. rothi, L. sagei, L. senguer, L. telsen L. tehuelche, and L. telsen L. arambarensis, L. azarai, L. cuyumhue, L. lutzae, L. multimaculatus, L. occipitalis, L. rabinoi, L. L. wiegmannii riojanus, L. salinicola, L. scapularis and L. wiegmannii L. goetschi L. cuyanus, L. donosobarrosi, L. goetschi and L. mapuche L. camarones, L. canqueli, L. casamiquelai, L. chehuachekenk, L. fitzingerii, L. melanops, L. L. fitzingerii morenoi L. shehuen and L. xanthoviridis Liolaemus shehuen sp. nov. belongs to the L. fitzingerii clade (Abdala, 2007), which is characterized by its tricuspidate posterior teeth with expanded crowns, neck equal to or wider than head, cylindrical body, short limbs relative to the trunk, black antehumeral arch present and ventrally expanded, and males with ventral melanism. Within the Liolaemus fitzingerii clade, L. shehuen sp.nov. has four to six scales in contact to mental (as the southern populations of L. fitzingerii). In L. shehuen the head melanism characteristic of L. canqueli and L. melanops is absent. Liolaemus shehuen lacks the intense green dorsum with irregular, transverse black paravertebral spots of L. melanops, and the transverse black and yellow-orange bands of L. canqueli. Liolaemus shehuen differs from L. chehuachekenk in lacking a conspicuous vertebral band, having fewer individuals with pre- and postscapular spots, fewer ventral scales (109-121, mean= 115.6 vs. 99-116, mean= 106), and shorter snout-vent length (SVL 90.8 mm vs. 98.3 mm); it differs from L. fitzingerii in having shorter snout-vent length (SVL 90.8 mm vs. 106 mm), and lacking the bluish green or light blue dorsum coloration with red paravertebral spots and red scales speckled, as L. fitzingerii has. Liolaemus shehuen differs from L. morenoi in lacking evident scapular markings and having more ventral scales (109-121, mean= 115.6 vs. 98-113, mean= 107.6). L. morenoi never presents yellow-red dorsal coloration either. Liolaemus shehuen differs from L. xanthoviridis in having different dorsal and ventral coloration, with paravertebral spots and more uniform vertebral region without irregular markings. Description of the holotype. ― Adult male. SVL 90.6 mm. Trunk length 45.1 mm. Head longer (17.4 mm) than wide (14.8 mm). Head height 11.6 mm. Eye diameter 4.3 mm. Interorbital distance 8.7 mm. Orbit���auditory meatus distance 6.6 mm. Auditory meatus 3.1 mm high, 1.6 mm wide. Orbit���commissure of mouth distance 2.5 mm. Internares distance 3.0 mm. Subocular scale length 5.4 mm. Femur length 13.5 mm, tibia 13.7 mm, and foot 25.6 mm. Humerus length 13.1 mm. Tail length 120.2 mm. Dorsal surface of the head smooth, with 15 scales. Rostral wider than long, bordered by six scales. Mental larger than rostral, trapezoidal, bordered by four scales. Nasal not in contact with rostral. Four internasals. Nasal surrounded by eight scales, separated from canthal by two scales. Five scales between frontal and supercilliaries. Seven scales between frontal and rostral. Frontal not divided. Two postrostrals. Interparietal smaller than parietals, in contact with six scales. Orbital semicircles complete. Preocular separated from lorilabial row by one scale. Three scales in anterior margin of auditory meatus. Eleven smooth temporals. Three lorilabials in contact with subocular. Eight supralabials, none in contact with subocular. Eight supraoculars. Eight lorilabials. Five infralabials, second ventrally in contact with two scales. Six chinshields, second pair separated by three scales. Seventy-six scales around midbody. Eighty-six round, slightly imbricate, slightly keeled dorsals from occiput to hind limbs. Thirty-six scale rows in dorsum. Scales of flank same size and shape as dorsals. Ventrals same size as dorsals, flat, imbricate. Thirty-five smooth, weakly imbricate gulars. Eight precloacal pores. Antehumeral scales flat, larger than or equal in size to dorsals. Postauricular, rectal, and longitudinal folds present. Scales on the longitudinal fold granular and smooth. Fourth finger with 19 subdigital lamellae; fourth toe with 26. Infracarpals flat, imbricate, trifid; infratarsals flat, imbricate, not trifid. C olor in life.― Head grey with scattered red spots and scales on the temporal and occipital region. Conspicuous antehumeral arch. Dorsum and flanks yellow-red with faint, small paravertebral spots. Some of these spots on the posterior region have a lighter spot inside. Vertebral region orange, with two light grey stripes on each side of vertebral region. Dorsal fore- and hind limbs same color as trunk. Dorsal tail greenish grey with yellow and orange scales on flanks and at tail base. Throat pale white-yellow. Gular region, chest to cloacal region and ventral hind limbs melanic. Flanks, forelimbs and ventral tail same color as throat. Variation.― (Table 3). Based on 10 specimens. Head longer (14.8���17.9 mm; mean = 16.4 mm) than wide (11.9���14.8 mm; mean = 13.4 mm). Head height 9.8���11.6 mm (mean = 10.7 mm). Neck wider than head. SVL 72.7���90.8 mm (mean = 79.9 mm), averaging 2.1 times longer than trunk. Tail length 94.9���120.3 mm (mean = 109.6 mm). Humerus length 9.7���13.1 mm (mean = 11.4 mm). Dorsal surface of head smooth, with 14���18 scales. One row of lorilabials, same size as supralabials. Six to nine lorilabials. Seven to 10 supralabials. Frontal unique or divided horizontally, into two or four scales. Interparietal always smaller than parietals, surrounded by 6���8 scales. Nasal surrounded by 8���9 scales. Five to seven infralabials. Mental in contact with four or six scales. Ten to 11 smooth round temporals. Longitudinal, postauricular, and antehumeral folds present. Horizontal fold Y-shaped between shoulder and auditory meatus. Scales around midbody 69���80 (mean = 74). Gulars 28���35 (mean = 31.5). Dorsal scales between occiput and hind limbs 63���88 (mean = 77.6). Ventrals 110���119 (mean = 117.3). Males with 8���9 (mean = 8.2) precloacal pores. Females with 0���3 (only one female) (mean = 0.5) precloacal pores. Dorsals flat, slightly imbricate, slightly keeled. Seventeen to 20 infradigital lamellae on fourth finger and 22���29 on fourth toe. Variation in color pattern. ― No sexual dichromatism evident (Figs. 6���7). Head usually grey or light brown with small yellow-orange spots or scales. Black arch ventrally expanded over antehumeral fold. Dorsal trunk and limbs yellow-orange or yellow-red, more intense in some specimens. Faint and small round paravertebral spots, generally dark, but sometimes preceded by a lighter spot. Some specimens have one or two dorsolateral whiteyellow strips and/or vertebral line of same color as paravertebral spots. Some specimens have diffuse white-yellow bars transversal to the body axis. Lateral region of the body same color as dorsum. Tail uniformly bluish grey or greenish grey with scattered spots. Gular region, chest to cloacal region, anterior forelimbs and hind limbs have ventral melanism. Males more melanic than females. In juveniles melanism expands from gular region to cloacal region as the individual grows. Natural history.― Liolaemus shehuen sp. nov. was found in the Telsen plateau and nearby regions, along National Route 4, an area with sand dunes and abundant vegetation, an ecotone between the phytogeographic regions of Monte and Patagonia (Cabrera & Willink 1980). The soil is mostly sandy with basaltic rocks, surrounding a lowland with red soils called ���Bajo de la tierra colorada���. The vegetation consists mainly of Stipa, Verbena, Mulinum, Nassauvia and Lycium (Cei & Scolaro 1999). The Patagonian plateaus (Telsen, Somuncur�� in R��o Negro and Chubut provinces, and Buenos Aires Lake, in Santa Cruz Province) are areas with high herpetofauna diversity. Liolaemus shehuen sp. nov. is sympatric with Diplolaemus darwinii (Bell), Homonota darwinii (Boulenger), L. bibronii (Bell), L. petrophilus (Donoso-Barros & Cei), L. rothi, L. telsen, Leiosaurus belli (Dum��ril & Bibron), Pristidactylus nigroingulus (Cei, Scolaro & Videla), the snakes Philodryas patagoniensis (Girard), Philodryas trilineata (Burmesteir), and the viperid Bothrops ammodytoides (Leybold). Liolaemus shehuen is oviparous and insectivorous. It is active between 12.00 AM and 15:00 PM, when the sand temperature is highest. There is no more information about its biology. Distribution.― Liolaemus shehuen is distributed on the Telsen plateau and east of R��o Negro Province, Argentina (Fig. 8). Etymology.― shehuen is a Tehuelche word that means 'Sun', and makes reference to the dorsal coloration of Liolaemus shehuen. The tehuelches, also known as Patagones, are ancient indigenous people that have inhabited Patagonia for more than 10000 years., Published as part of Abdala, Cristian Sim��n, G��mez, Juan Manuel D��az & Heredia, Viviana Isabel Juarez, 2012, From the far reaches of Patagonia: new phylogenetic analyses and description of two new species of the Liolaemus fitzingerii clade (Iguania: Liolaemidae), pp. 34-60 in Zootaxa 3301 on pages 39-44, DOI: 10.5281/zenodo.246399, {"references":["Etheridge, R. E. (1995) Redescription of Ctenoblepharys adspersa Tschudi, 1845, and the taxonomy of Liolaeminae (Reptilia: Squamata: Tropiduridae). American Museum Novitates, 3142, 1 - 34.","Abdala, C. S. (2007) Phylogeny of the boulengeri group (Iguania: Liolaemidae, Liolaemus) based on morphological and molecular characters. Zootaxa, 1538, 1 - 84.","Abdala, V. Abdala, C. & Tulli, M. J. (2006) Muscular characters in the phylogeny of Liolaemus (Squamata: Iguania: Liolaemidae): a reappraisal. Zootaxa, 1205, 55 - 68.","Cabrera, A. L. & Willink, A. (1980) Biogeografia de America Latina. Secretaria General de la OEA. Prog. Reg. de Desarrollo Cientifico y Tecnologico, Washington D. C.","Cei, J. M. & Scolaro, J. A. (1999) Speciation of the \" darwinii complex \" (genus Liolaemus, \" patch group \") in the southernmost area of its distribution (Reptilia: Tropiduridae). Revue francais de. Aquariologie, 26, 79 - 82."]}

Liolaemus diaguita sp. nov. (Fig. 1���5) 1993. Liolaemus quilmes ���Etheridge, Boll. Mus. Reg. Sci. Nat. Torino, 11: 137���199. 2008. L. cf. quilmes 1 ���Morando et al., Zoologica Scripta, 37: 603���618. Holotype. MCN 3247. 61 Km N of Cafayate, on National Road 34, 400 m S of R��o Las Conchas, Quebrada de las Conchas; Guachipas Department, Salta Province. 25 �� 49 '07.1''S; 65 �� 41 ' 26.1 ''W. 1328 m. Arias, F., Palavecino, A., Portelli, S. & Quinteros, S. cols. 27 /01/ 2010. Paratypes. FML 23821 - 825. MCN 3145 - 50; 3243; 3248 - 50. Same data as Holotype. Diagnosis. Liolaemus diaguita sp nov., belongs to the L. boulengeri group because it has a patch of enlarged scales on posterior region of thigh (Etheridge, 1995; Abdala et al. 2006). Within this group, it belongs to the L. laurenti group (Abdala, 2007) because its posterior teeth have straight edged crowns and sexual dichromatism is evident. Within the L. laurenti group, Liolaemus diaguita sp nov. is a member of the L. darwinii group (Abdala, 2007) or the darwinii complex (Etheridge, 1993) because it has a black line which crosses the eye vertically (except L. chacoensis) and pre-scapular spots in both males and females (Abdala, 2007). Tooth shape and the line crossing the eye distinguish the new taxon from all members of the L. melanops group [Abdala, 2007 (L. boulengeri, L canqueli, L. cuyanus, L. donosobarrosi, L. fitzingerii, L. inacayali, L. josei, L. mapuche, L. martorii, L. melanops, L. morenoi, L. loboi, L. sagei, L. rothi, L. senguer, L. tehuelche, L. telsen, and L. xanthoviridis)]; also the presence of sexual dichromatism distinguish L. diaguita from L canqueli, L. cuyanus, L. chehuachekenk, L. goestchi, L. fitzingerii, L. martorii, L. morenoi, L. sagei, and L. xanthoviridis. It differs from L. canqueli and L. melanops, because L. diaguita lacks cephalic melanism, and from L. canqueli, L. cuyanus, L. mapuche, L. morenoi, L. fitzingerii and L. xanthoviridis because those species show gular and antehumeral melanism, absent in L. diaguita. The new species has four scales contacting the mental scale, differing from L. cuyanus and all the members of the L. wiegmannii group (L. arambarensis, L. azarai, L. cranwelli, L. cuyumhue, L. lutzae, L. multimaculatus, L. occipitalis, L. rabinoi, L. riojanus, L. salinicola, L. scapularis and L. wiegmannii), which have six scales contacting the mental and from L. josei and L. mapuche where the mental scale can contact 4 to 6 scales. Also, the members of the L. wiegmannii group have two rows of lorilabial scales, while L. diaguita has one row. The new taxon differs from species of the L. anomalus group (L. anomalus, L. ditadai, L. lentus, and L. pseudoanomalus - sensu Abdala, 2007) because its tail is longer than snout-vent length (same length in L. anomalus group), head larger than wide (wider than long in L. anomalus group), enlarged superior cilliaries absent (present in L. anomalus group), larger number of precloacal pores in males and females, and lack of pterigoid teeth. Within the L. darwinii group (Abdala, 2007), L. diaguita differs from the members of the L. ornatus group (L. albiceps, L. calchaqui, L. crepuscularis, L. irregularis, L. lavillai, and L. ornatus) because it has an oviparous reproductive mode and it lacks precloacal pores in females. Also it differs from L. albiceps and L. irregularis because it is smaller (Max SVL 49.7mm vs 82.5 and 86.1 mm respectively). Liolaemus diaguita differs from L. abaucan and L. uspallatensis in having imbricate and keeled dorsal scales, while in those two species the dorsal scales are smooth or weakly keeled. Liolaemus chacoensis exhibits keeled temporal scales and mucronate dorsal scales, absent in L. diaguita. The presence of a small postscapular spot distinguishes L. diaguita from L. abaucan, L. darwinii and L. koslowskyi which have a large and evident postscapular spots. It differs from L. cinereus in having a black stripe in the lateral field and in lacking melanism on the throat. Liolaemus darwinii, L. grosseorum, L. laurenti, L. montanezi and L. olongasta have a black antehumeral arch, absent in L. diaguita. Liolaemus espinozai shows spots located posterior to the postscapular spot and a lateral line in males and females, while both are absent in L. diaguita specimens. Liolaemus diaguita differs from L. quilmes because it has a small prescapular spot and a black stripe in the lateral field, both absent in L. quilmes. Males and females of Liolaemus diaguita have a unique dorsal color pattern formed by paravertebral and lateral spots which can join together forming paravertebral and lateral stripes respectively. Additionally, the specimens lack spots on throat and belly. These character states distinguish the new taxon from all members of the L. laurenti group. Description of the holotype. Adult male. Snout vent length (SVL) 49.68. Head 1.3 times longer (12.48 mm) than wide (9.21mm). Head height 6.71 mm. Interorbital distance 6.75 mm. Eye-auditory meatus distance 4.71 mm. Auditory meatus height 2.1 mm; wide 1.61 mm. Supralabials���eye 2.3 mm; distance between nares 1.53 mm. Subocular legnth 0.72 mm. Trunk length 22.31 mm; tail length 70.32 mm. Tibia length 4.84 mm. Foot length 14.98 mm. Humerous length 3.62 mm. Dorsal surface of head smooth, with five enlarged supraoculars. Eight scales between rostral and frontal scales. Mental scale trapezoidal, larger than rostral, in contact with four scales. Four chinshields scales. There is no contact between nasal and rostral scales, separated by one scale. Seven scales surround the nasal scale, which is separated from canthal scale by four scales. Two postrostral scales, with seven scale organs. Interparietal scale larger than paritetals, in contact with six scales. Hellmich index (dorsal scales in head, from occiput to mental scale) 17. Preocular separated from lorilabial row by one scale. Nine supercilliaries. Sixteen upper cilliaries. Anterior edge of the auditory meatus with three scales projecting. Auricular scale (located in antero-superior edge of the auditory meatus) present. Eleven smooth temporals. Subocular scale in contact with four lorilabials. Five supraocular scales. Nine supralabial and six infralabial scales. One row of nine lorilabials. Scales around midbody 57. Sixty-three round, keeled and imbricated dorsal scales (from occiput to forelimbs). Sixty-eight ventral scales from mental scale to cloaca, larger than dorsals, laminar and inbricate. Twenty-four imbricate and smooth gulars. Antegular fold present. Forty-five granular and smooth neck scales (counted from posterior margin of the auditory meatus to sholuder, along the longitudinal fold). Neck folds (auricular, antehumeral and longitudinal) evident. Seventeen infradigital lamellae on fourth finger and 28 on fourth toe. Femoral patch evident. Seven precloacal pores. Dorsal caudal scales mucronate and smooth; ventral smooth. Color of the holotype in ethanol. Dorsal background color dark brown. Head dorsally brown with dark brown spots. Lateral zone of head brown with black lines. Temporal region with black spots. A black line born in the posterior margin of the eye, extending and reaching the lateral field. A vertical black line crosses the eye (like all other members of the L. darwinii group���except L. chacoensis). Dark brown vertebral zone, without vertebral line. In the region close to the occiput there are light brown to white lines. Eight pairs of paravertebral white spots are evident. Two dorsolateral white stripes, bordered in light brown. Lateral fields dark brown to black, with white spots speckled. Ventrolateral line white and evident, bordered by a brown line ventrally. Ventrolateral field brown with rounded white spots. Fore and hind limbs brown (lighter than dorsum) speckled with dark brown and white spots. Ventrally throat, chest, belly, limbs and tail are immaculate, with a creamy white color. Tail dorsally, brown with black spots forming a ring pattern. Color in life of the holotype. Dorsal background. Head with same color of dorsum with dark brown to black lines. Temporal region same color than dorsum, with a black line which begins on the posterior margin of the eye, and reaches the lateral region of body. Pre-scapular spot present. Paravertebral field black with ten pairs (between fore and hind limbs) of white irregular spots. Vertebral zone without vertebral line. Dorsolateral stripes light brown with white spots. Lateral field brownish yellow. In the lateral field is evident a dark brown line bordered with white spots. Ventrolateral line white and distinct. Ventrolateral field light gray with white spots speckled. Fore and hind limbs grayish brown with light gray and dark brown spots. Tail dorsally, same color of dorsum of body. The black paravertebral field of the body extends to the proximal region of tail. In distal region of tail there are black and white lines, forming a ringed pattern. Ventrally, throat, chest, belly, fore and hindlimbs, and tail, immaculate white cream. Variation. Based on fifteen paratypes. Head longer (10.2���12.6 mm; Mean = 11,5 mm) than wide (8.3���12.5 mm; Mean = 9.7 mm). Head height 6.0��� 7.4 mm (Mean = 6.5 mm). SVL 41.5���49.6 mm (Mean = 47.2 mm), averaging 2 times longer than trunk. Tail length 64.3���81.7 mm (Mean = 70.7 mm). Width of tail base 4.6���6.5 mm (Mean = 5.9 mm). Humerus length 3.1���6.2 mm (Mean = 4.4 mm). Tibia length 4.4���6.5 mm (Mean = 5.1). Foot length 11.3���17.9 mm (Mean = 14.0 mm). Fourth finger length 4.9���7.9 mm (Mean = 6.2 mm). Fourth toe length 8.9���11.6 mm (Mean = 9.9 mm). Dorsal surface of head smooth, with 14���20 scales (Hellmich index). Seven to eight supercilliaries. Ten to sixteen upper cilliaries. One row of lorilabials, same size as supralabials. Eight to ten lorilabials. Seven to eleven supralabials. Frontal divided into two scales. Interparietal always larger than parietals, surrounded by 4���8 scales. Nasal surrounded by 6���8 scales. Four to six infralabials. Mental in contact with four scales. Ten to thirteen smooth and round temporals. Longitudinal, postauricular and antehumeral folds present. Horizontal fold Yshaped between the shoulder and the auditory meatus. Scales around midbody 52���58 (Mean = 55.5). Gulars 20���29 (Mean = 22.8). Dorsal scales between occiput and hind limbs 57���68 (Mean = 62.6). Ventrals 58���70 (Mean = 64.6). Males with 5���7 (Mean = 5.8) precloacal pores. Females without precloacal pores. Dorsals flat, imbricate and keeled. Sixteen to twenty infradigital lamellae on fourth finger and 24���30 on fourth toe. Variation in color pattern. (Fig 1���4) Sexual dimorphism evident. Males exhibit the same color pattern as the holotype. The only difference is in the intensity of the yellow color in the lateral fieldis more intense in males.Females with the dorsal background color gray to light brown. Head of females are similar in males and females. The vertebral region is bordered by black rounded spots. The size and number of those spots vary and they are absent in some specimens. Pre-scapular spot present. The lateral field exhibits two stripes, a black on the top and white cream below, without spots. Light brown fore and hind limbs with dark brown lines and spots. Ventrally females show the same color of the holotype. Distribution. The new species is known only from its type locality. (Fig. 5) Natural history. Liolaemus diaguita was found in a sandy area where vegetation is predominated by shrubs (Thrichocereus sp, and Prosopis sp). The area belongs to the phytoegegraphical region of Monte (Cabrera and Willink, 1980). It can be found basking on rocks along the road. Etymology. The epithet diaguita comes from native people denominated Diaguitas, who inhabit Northwestern Argentina, especially in Valles Calchaquies, since the year 850. These communities developed a rich culture, and at present there are more than 60,000 people in Northwestern Argentina who belong to this ethnic group. SA D PPM PPF Max. SVL L. diaguita 52���58 (55.5) 57���68 (62.6) 5���7 (5.8) 0 49.6 mm continued on the next page... Table 1 continued, Published as part of Abdala, Cristian Sim��n, Quinteros, Andr��s Sebasti��n, Arias, Federico, Portelli, Sabrina & Palavecino, Antonio, 2011, A new species of the Liolaemus darwinii group (Iguania: Liolaemidae) from Salta Province, Argentina, pp. 26-38 in Zootaxa 2968 on pages 27-32, DOI: 10.5281/zenodo.204489, {"references":["Etheridge, R. E. (1995) Redescription of Ctenoblepharys adspersa Tschudi, 1845, and the taxonomy of Liolaeminae (Reptilia: Squamata: Tropiduridae). American Museum of Natural History Novitates, 3142, 1 - 34.","Abdala, V., Abdala, C. S. & Tulli, M. J. (2006) Three traditional muscular characters in the phylogeny of Liolaemus (Squamata: Tropiduridae) a reappraisal. Zootaxa, 1205, 55 - 68.","Abdala, C. S. (2007) Phylogeny of the boulengeri group (Iguania: Liolaemidae, Liolaemus) based on morphological and molecular characters. Zootaxa, 1538, 1 - 84.","Etheridge, R. E. (1993) Lizards of the Liolaemus darwinii complex (Squamata: Iguania: Tropiduridae) in northern Argentina. Bollettino del Museo Regionale di Scienze Naturali, Torino, 11, 137 - 199.","Cabrera, A. L. & Willink, A. (1980) Biogeografia de America Latina. Secretaria General de la Organizacion de los Estados Americanos (OEA), Departamento de Asuntos Cientificos y Tecnologicos, Monografia 13 (Serie de Biologia), 122 pp."]}

Liolaemus scrocchii sp. nov. (Fig. 7���8) Liolaemus dorbignyi (non Koslowsky, 1898): Laurent (1991), N����ez (1992), N����ez and Jacksic (1992), Laurent (1995), Etheridge (1995), Laurent (1998), Schulte et al. (2000), Valladares et al. (2002), Espinoza et al. (2004), Pincheira- Donoso and N����ez (2005), Cruz et al. (2005), and Abdala (2007). Holotype��� FML 17228. Male. Two Km SE of Susques on National road 52. Dto. Susques, Jujuy, Argentina. Abdala, C., Barrionuevo, S., Quinteros, S., and G., Scrocchi, Cols. 25 -Feb-05. Paratypes ��� FML 17229. Male, MCN 2136���2137, male and female; same data as the holotype. FML 0 1763, 5 females, 4 males; Santa Rosa de los Pastos Grandes. Dto. Los Andes, Salta. O. Pagaburo col. 06- May- 86. FML 0 1759 male and female; Agua de los Sapos. (Puesto de Cruz Soriano), 20 Km from Mina Tincalayo. Dto. Los Andes, Salta; O.Pagaburo, col. 06-May- 86. FML 0 1817. Male, El Quemado, Abra de Pives. Dto. Tumbaya, Jujuy, 4000 m. E. Lavilla, A. Marcus, and R. Montero cols. 20 -Feb- 87. FML 0 6481. Male. La Mesada, 64 km from La Poma to Abra del Acay. Dto. La Poma, Salta. 4665 m. S. Torres col. 21 -Jul- 97. FML 0 0 992. male and 2 females. Around base of the Pa��o mountain. Serran��a de Cha��i. 4200���4300 m. Dto. Belgrano, Jujuy. G. Scrocchi col. MCN 714; MCN 898; MCN 961. Females. Casa Mocha, climbing from NW of the Nevado del Cha��i. 4500���4700 m. C. Bracamonte col. 29 -Mar- 2002. MCN 899. Male. Abra del Cerro Toro. Sierra de Aguilar. R. Seggiaro col. 10 -Oct- 2002. MCN 964. Male. Road to Abra del Acay from national road 51. Near 4900 m. I. Mart��nez Oliver, S. Lobo and F. Lobo cols. 19 -Jan-03. MCN 1922���1925. Female and 2 males. Km 210, Nacional road 51. 0.6 Km from nacional road 51. 6.4 Km to S of Olacapato. 24.2411475 S; 66.69711 W. 4070 m. B. Casimiro, Espinoza, R., Lobo, F. and S. Quinteros cols. 17 -Jan-06. Diagnosis��� Liolaemus scrochii belongs to the L. montanus group (Etheridge, 1995), diagnosed by a blade-like process on posterior distal tibia, associated with a greatly hypertrophied M. tibialis anticus (Abdala, et al. 2006). It has scales of equal size on the posterior face of the femur, distinguishing it from the L. boulengeri group (Abdala, 2007). Within the L. montanus group, L. scrocchii is a robust lizard, with a large size (max. SVL 95 mm) which distinguishes it from L. andinus, L. audituvelatus, L. disjunctus, L. eleodori, L. erroneus, L. etheridgei, L. fabiani, L. famatinae, L. fittkaui, L. griseus, L. huacahuasicus, L. insolitus, L. islugensis, L. molinai, L. montanus, L. multicolor, L. ortizi, L. pantherinus, L. poconchilensis, L. poecilochromus, L. pulcherrimus, L. rechei, L. rosenmanni, L. ruibali, L. signifier, and L. vallecurensis, which are smaller (SVL 50��� 76 mm). Liolaemus scrocchii has flat, imbricated and slightly keeled dorsal scales which distinguish it form L. andinus, L. audituvelatus, L. erguetae, L. erroneus, L. forsteri, L. insolitus, L. islugensis, L.molinai, L. patriciaiturrae, L. poecilochromus L. robertoi, L. rosenmanni, and L. schmidti, (smooth and juxtaposed dorsal scales) and from L. aymararum, L. disjunctus, L. etheridgei, L. fittkaui, L. huacahuasicus, L. montanus, L. ortizi, L. polystictus, L. stolzmanni, L. thomasi, and L. williamsi (imbricate and strongly keeled scales). Liolaemus chlorostictus, L. jamesi, L. orientalis, L. poconchilensis, L. robustus, L. signifer and L. stolzmanni have slightly keeled but juxtaposed scales. The number of scales around midbody in L. scrocchii is 55���68, distinguishing it from L. andinus, L. disjunctus, L. duellmani, L. eleodori, L. erguetae, L. foxi, L. islugensis, L. molinai, L. multicolor, L. nigriceps, L. patriciaiturrae L. pleopholis,, L. poecilochromus, L. pulcherrimus, L. robertoi, L. rosenmanni, L. ruibali, L. signifer, and L. vallecurensis (between 64 and 110). The females of L. scrocchii have precloacal pores 54.14 % (eight of fourteen females), differing from L. andinus, L. duellmani, L. fabiani, L. islugensis, L. jamesi, L. melanogaster, L. pantherinus, L. puritamensis, L. robertoi, L. rosenmanni, L. signifer, L. stolzmanni, and L vallecurensis in which females lack precloacal pores. The dorsal color pattern is composed of transverse paravertebral spots. These spots are band-shaped, never reach the lateral region, and are never in contact in the vertebral zone (Fig. 7). This is the main difference between L. dorbignyi, L. griseus and other members of the L. montanus group. Also, the number of scales around midbody in L. scrocchi is generally higher 55���68 (x = 61.8) than L. dorbignyi 48���59 (x = 54.5). The presence of juxtaposed or subjuxtaposed dorsal scales and the absence of keeled or slightly keeled dorsal scales further differentiate L. scrocchii from L. dorbignyi. Description of the Holotype ���Snout-vent-length 87.3 mm. Trunk length 41.7. Head larger (19.2 mm) than wider (17.2 mm). Head height 11.1 mm. Eye diameter 7.1 mm. Interorbital distance 11 mm. Orbit-auditory meatus distance 7.1 mm. Auditory meatus height 2.8 mm, wide 0.9 mm. Orbit-comisure of mouth distance 3.1 mm. Internares 3 mm. Sub-ocular scale 4.3 mm long. Femur length 14.6, tibia 17.3, and foot 25.8 mm. Humerus length 11.8 mm. Dorsal surface of the head smooth, with 14 dorsal scales. Rostral wider than high, bordered by six scales. Mental larger than rostral, bordered by four scales, trapezoidal. Nasals do not contact rostral. Two internasal scales. Seven scales surrounding nasal separated from canthal by two scales. Five scales between rostral and supercilliaries, six between rostral and frontal. Frontal scale divided. Two postrostrals. Interparietal smaller than parietals, in contact with five scales. Fourteen upper cilliaries. Supraorbital semicircles incomplete. Preocular scale separated from lorilabials by one scale. Two scales on anterior margin of auditory meatus and three on upper margin. Six smooth temporals (from upper margin of the auditory meatus to postocular scale) Three lorilabials contact subocular. Seven supralabials, the fourth of equal in size to others, without scales organs, and not in contact with the subocular. Six supraoculars. Eight lorilabials. Six infralabials, the second ventrally in contact with two scales. Five chinshields, the second separated by two scales. Fifty seven scales around midbody. Forty eight, rounded, sub-juxtaposed, smooth dorsals, from occiput to hindlimbs. Twenty nine scale rows on dorsum. Ventrals scales same size as dorsals, flat and juxtaposed. Forty eight sub-juxtaposed and smooth gulars. Five precloacal pores. Antehumeral scales flat, larger or equal to dorsals. Postauricular, rictal and longitudinal folds present. Scales on the longitudinal fold, granular and smooth. Fourth finger with 17 subdigital lamellae and 26 on fourth toe. Flank scales, rounded, smooth with some small granular intercalated scales. Without femoral patch. Flat, imbricate and triphid infracarpal scales. imbricated and keeled infratarsal scales. Color in ethanol��� Head dorsally dark brown, with scattered dark gray scales. Head darker than rest of body. Trunk reddish-brown, with seven conspicuous, rhomboidal, black paravertebral spots, arranged transverse to body axis; posteriorly spots in contact in vertebral zone. A black spot present on posterior region of trunk, owing, probably, to scale damage. Fore and hind limbs same color as trunk, but without dark spots. Vertebral line, scapular spots and dorsolateral stripes absent. Lateral region of body with same color trunk, but light gray in ventrolateral zone. Iridescent or brightly colored scales, absents. Dorsally tail with same color as trunk. Cloacal region gray. Ventrally, from mental to cloacal region light gray, variegated with dark and light spots. On thighs a light yellow color. Tail exhibits a light gray, ringed pattern. Variation��� Based on 27 paratypes. Head larger (15.1���20.6 mm; x = 17,9 mm, than wider (13.0��� 19.4 mm; x = 16.4 mm). Head height 9.1���12.3 mm (x = 10.8 mm). Neck wider than head. Snout-vent length, 2.1 times longer than trunk. Snout-vent length 72.8���94.5 mm (x = 83.9 mm). Tail length 85.1���101.3 mm, (x = 92.3 mm). Tail base width 8.9���14.1 mm, (x = 11.3 mm). Humerus length 5.5���12.7 mm, (x = 9.3 mm). Tibia length 12.2���18.5 mm, (x = 16.3 mm). Foot length 21.8���27.9 mm, (x = 24.0 mm). Fourth finger length 7.1��� 10.1 mm, (x =9.0 mm) and fourth toe 11.1���17.3 mm, (x = 14.1 mm). Dorsal surface of head, slightly rugose, with 14���19 scales. Six to 7 supercilliaries. Twelve to 16 upper cilliaries. One row of lorilabials same size as supralabials. Seven to 10 lorilabials. Six to 8 supralabials. Frontal scale divided in two, three or four. Interparietal scale larger, smaller or equal to parietals, and surrounded by 5���8 scales. Nasal scale surrounded by 5���8 scales. Four to 6 infraliabals. Mental in contact with 4 scales. Four to 8 smooth horizontal temporals. Up to 4 auriculars scales on anterior margin of auditory meatus. Neck folds present; horizontal fold is ���Y��� shaped between shoulder and auditory meatus. Scales around midbody 55���68 (x = 61.8). Gulars 34���46 and 36���48 scales on neck (from posterior margin of auditory meatus to shoulder, along horizontal fold). Dorsal scales between occiput and hind limbs 42���53 (x = 49.6). Ventrals 72���98 (x = 86.4). Males with 2���10 (x = 5.6) precloacal pores. Among fourteen females, eight exhibits 3���6 precloacal pores. Flat, juxtaposed or subjuxtaposed, smooth dorsal scales. Sixteen to 21 infradigital lamellae on fourth finger and 23���28 in fourth toe. Without femoral patch. Color in life��� (Fig. 7���8) Without sexual dichromatism. Dorsal head design uniform, light brown or reddish-brown, with yellowish shades and dark or black scales. Body background light brown or reddish-brown, in some specimens orange tones are present. Large dark brown or black rhomboidal or rectangular paravertebral spots, always transverse to body axis (Fig. 7). These spots never fused in vertebral zone of trunk, but are fused at base of tail. Lateral fields yellow, orange or reddish-brown, darker than dorsum. Light blue or white scales irregularly spread on laterals fields and on fore and hind limbs. Without vertebral line, dorsolateral bands, or scapulars spots. Tail light brown, white or yellow shaded. Paravertebral spots disappear toward end of tail. Ventrally intense yellow from mental to cloacal region, variegated with dark and light brown scales (Fig. 8). Some specimens with gular region orange or reddish. Tail pattern same as that of body. Distribution��� Liolaemus scrocchii occurs in the Puna region of Jujuy and in the northwest of Salta province (Fig. 5). In Jujuy province it is known from Susques, Abra de Pives, Cerran��as de Cha��i, Chorcan, Abd��n Castro Tolay, Sey and Sierra de Aguilar, and in Salta province it is found at Abra del Acay, Olacapato, Santa Rosa de los Pastos Grandes, Huaytiquina and Mina Tincalayo (Fig 3). Natural History��� Liolaemus scrocchii, is known from localities in the phytogeographic region of the Puna (Cabrera and Willink, 1980), always above the 3000 m. It is commonly found basking on rocks and on the sand banks nearby (Fig. 9). It is viviparous, omnivorous with a high degree of plant material in its diet (Espinoza et al. 2004). It lives in sympatry with L. multicolor, L. ornatus Koslowsky and L. puna Lobo and Espinoza. Etymology��� We name this new taxon scrocchii to acknowledge our colleague and friend Gustavo Scrocchi for his extensive and dedicated work on the systematics and biology of Argentine reptiles, and in particular for his invaluable kindness and companionship. Remarks��� The species described here belongs to the L. montanus group (Etheridge, 1995) within the subgenus Eulaemus (Laurent, 1983). The squamation and color pattern distinguish it from L. dorbignyi and the other species of the genus. The species here described inhabits many localities of the Puna in Jujuy Province and in northwestern Salta Province, between 23 ����� 25 �� S and 65 ����� 67 �� W. This species is viviparous, saxicolous and lives on rocky mountain slopes. Individuals of L. scrocchii from the Nevado del Cha��i (Jujuy province) and from Abra del Acay (Salta, Province), in captivity, exhibit a particular behavior to stay, part of the day, in water., Published as part of Quinteros, Andr��s Sebasti��n, Abdala, Cristian Sim��n & Lobo, Fernando Jos��, 2008, Redescription of Liolaemus dorbignyi Koslowsky, 1898 and description of a new species of Liolaemus (Iguania: Liolaemidae), pp. 51-67 in Zootaxa 1717 on pages 59-62, DOI: 10.5281/zenodo.274176, {"references":["Koslowsky, J. (1898) Enumeracion sistematica y distribucion geografica de los reptiles argentinos. Revista del Museo de La Plata, 8, 161 - 200.","Laurent, RF. (1991) Sobre algunas especies nuevas y poco conocidas de Liolaemus (Iguanidae) de la provincia de Jujuy, Argentina. Acta Zoologica Lilloana, XL, 2, 91 - 108.","Laurent, R. F. (1995) Sobre una pequena coleccion de lagartos del genero Liolaemus (Tropiduridae) proveniente del extremo suroeste de Bolivia. Cuadernos de Herpetologia, 9 (1), 1 - 6.","Etheridge, R. E. (1995) Redescription of Ctenoblepharys adspersa Tschudi, 1845, and the taxonomy of Liolaeminae (Reptilia: Squamata: Tropiduridae). American Museum of Natural History Novitates, 3142, 34 pp.","Laurent, RF. (1998) New forms of lizards of the subgenus Eulaemus of the genus Liolaemus (Reptilia: Squamata: Tropiduridae) from Peru and northern Chile. Acta Zoologica Lillooana, 44 (1), 1 - 26.","Schulte J. A., Macey, J. R., Espinoza, R. E. & Larson, A. (2000) Phylogenetic relationships in the iguanid lizard genus Liolaemus: multiple origins of viviparous reproduction and evidence for recurring andean vicariance and dispersal. Biological Journal of the Linnean Society, 69, 75 - 102.","Valladares, J. P., Etheridge, R., Schulte, II J. Manriquez, G. & Spotorno, A. (2002) Nueva especie de lagartija del norte de Chile, Liolaemus molinai (Reptilia: Liolaeminae). Revista Chilena de Historia Natural, 75, 473 - 489.","Pincheira - Donoso, D. & Nunez, H. (2005) Las especies del Genero Liolaemus Wiegmann, 1834 (Iguania: Tropiduridae: Liolaeminae) Taxonomia, Sistematica y Evolucion. Publicacion Ocasional del Museo Nacional de Historia Natural, Chile, 59, 7 - 486.","Cruz, F. B., Fitzgerald, F. B., Espinoza, R. E. & Schulte II J. A .. (2005) The importance of phylogenetic scale in tests of Bergmann's and Rapoport's rules: lessons from a clade of South American lizards. Journal of Evolutionary Biology, 18 (6), 1559 - 1578.","Abdala, C. S. (2007) Phylogeny of the boulengeri group (Iguania: Liolaemidae, Liolaemus) based on morphological and molecular characters. Zootaxa, 1538, 84 pp.","Abdala, V., Abdala C. S & Tulli M. J. (2006) Muscular characters in the phylogeny of Liolaemus (Squamata: Iguania: Liolaemidae): a reappraisal. Zootaxa, 1205, 5 - 68","Cabrera, A. L. & Willink A. (1980) Biogeografa de Amrica Latina. Secretara General de la Organizacin de los Estados Americanos (OEA), Departamento de Asuntos Cientficos y Tecnolgicos, Monografa 13 (Serie de Biologa), 122 pp.","Laurent, RF. (1983) Contribucion al conocimiento de la estructura taxonomica del genero Liolaemus Wiegmann (Iguanidae). Boletin de la Asociacion Herpetologica Argentina, 1 (3), 16 - 18."]}

Liolaemus dorbignyi Koslowsky 1898 (Fig. 1���4) Designation of a lectotype��� (Fig. 1) Among the specimens of Liolaemus dorbignyi collected by Koslowsky (in his original description there is no mention of the number of specimens collected) which he used to write his description of this species, five adult specimens (three males and two females) are deposited in the Museo de La Plata (MLP). Koslowsky (1898) included a plate with three photographs of adult specimens (his Lam. VIII). Both ventral and dorsal pictures corresponded to two male specimens, but none seem to correspond to the preserved males extant in the MLP collection. There is a picture of a lateral view of a head, but this picture does not allow us to compare with the specimens preserved. The description also does not provide enough information to allow a determination of which specimens were used for the type description. Ferraro and Williams (2006) studied the type material of amphibians and reptiles deposited in the Museo de La Plata, which includes the species described by Koslowsky. In relation to L. dorbignyi, they identified a syntype (MLP.S 844), and argued that it corresponded to the ventral picture of a specimen in Koslowsky���s plate. According to Art 73.2 of the ICZN, all specimens belonging to the type series are automatically considered syntypes, when no holotype or lectotype was designed, so the five specimens of the Koslowsky���s series are syntypes. We don���t agree with Ferraro and Williams (2006), and we consider that none of the specimens of Koslowsky���s series match the specimens illustrated, and therefore we designate as lectotype the individual catalogued as MLP.S 0 57 (in compliance with art 74, of ICZN), a specimen which better represent the species. Also, the specimens catalogued as MLP.S 057; 844 ��� 45; 990 ��� 91 are designated paralectotypes. Designtion of the type locality��� Since Koslowsky (1898) described the type locality of Liolaemus dorbignyi as ���Catamarca province,��� the identity of this species has been problematic. Unfortunately, Koslowsky���s field notes, which would be very helpful, are lost (Jorge Williams pers. comm.). During the revision of materials deposited at the FML (Fundaci��n Miguel Lillo, Tucum��n) and MCN (Museo de Ciencias Naturales, Salta), we found specimens corresponding to different populations assigned to L. dorbignyi. Among these samples, only one individual (FML 1855) is similar to those collected by Koslowsky. However, these specimens had an incorrect label about their provenance (Saujil, National Road 60, 1550 m. Department Tinogasta, Catamarca Province). This incorrect label was confirmed during field trips, and O. Pagaburo (collector of FML 1855), told us that the specimens were collected in the Sierra de Fiambal��, over 3000 m. Later, from those mountains we collected specimens, which are similar to those collected by Pagaburo and Koslowsky. Koslowsky (1898), in the original description of L. dorbignyi, designated as type locality ���Catamarca Province.��� To determine where Koslowsky could have collected the specimens, preserved specimens were compared to specimens of different populations collected in several localities of Catamarca province, Argentina. Also we analyzed the pictures from Koslowsky���s publication; because of the five preserved specimens only two retain the original color pattern, while the others have this pattern obscure and inconspicuous owing to their preservation conditions. From the populations studied, those from Sierra de Fiambal��, Catamarca Province exhibit the same color pattern (with dark bands transverse to the body axis), flat and keeled dorsal scales, and better match with the preserved specimens, the pictures and the original description. There are no known records from Koslowsky���s journeys in Catamarca. However, the comparisons between the different populations of Catamarca with the preserved material (Fig. 1), allows us to restrict the type locality of L. dorbignyi to Sierra de Fiambal��, Catamarca province. Diagnosis��� Liolaemus dorbignyi belongs the L. montanus group (Etheridge, 1995), because it has a blade-like process on posterior distal tibia, associated with a greatly hypertrophied M. tibialis anticus (Abdala, et al. 2006). Within this group, the presence of scales of equal size on the posterior face of the femur differentiates it from the L. boulengeri group (Abdala, 2007). Among the species that have not a patch of enlarged scales on the posterior face of the femur, L. dorbignyi is a robust and large sized Liolaemus (max SVL 98.3 mm) which distinguishes it from L. andinus Koslowsky, L. audituvelatus N����ez and Y����ez, L. disjunctus Laurent, L. eleodori Cei et al., L. erroneus N����ez and Y����ez, L. etheridgei Laurent, L. fabiani Y����ez and N����ez, L. famatinae Cei, L. fittkaui Laurent, L. griseus Laurent, L. huacahuasicus Laurent, L. insolitus Cei and P��faur, L. islugensis Ortiz and Marquet, L. molinai Valladares et al., L. montanus Koslowsky, L. multicolor Koslowsky, L. ortizi Laurent, L. pantherinus Pellegrin, L. poconchilensis Valladares, L. poecilochromus Laurent, L. pulcherrimus Laurent, L. reichei Werner, L. rosenmanni N����ez and Navarro, L. ruibali Donoso- Barros, L. signifier Dumerill and Bibr��n, and L. vallecurensis Pereyra, (max SVL 50���70 mm). Liolaemus dorbignyi has flat dorsal scales, imbricate, and slightly keeled, character states that distinguish it from L. andinus, L. audituvelatus, L. erguetae Laurent, L. erroneus N����ez and Y����ez, L. forsteri Laurent, L. insolitus, L. islugensis, L. molinai, L. patriciaiturrae Navarro and N����ez, L. poecilochromus, L. robertoi Pincheira-Donoso and N����ez, L. rosenmanni, and L. schmidti Marx (smooth and juxtaposed dorsal scales). It may also be distinguished from L. aymararum Ve lo s o et al., L. disjunctus, L. etheridgei, L. fittkaui, L. huacahuasicus, L. montanus, L. ortizi, L. polystictus Laurent, L. stolzmanni Steindachner, L. thomasi Laurent, and L. williamsi Laurent, which have dorsal scales imbricate but strongly keeled. Liolaemus chlorostictus Laurent, L. jamesi Boulenger, L. orientalis M��ller, L. poconchilensis, L. robustus Laurent, L. signifier, and L. stolzmanni, have weakly keeled but juxtaposed dorsal scales, characteristic that distinguishes it from L. dorbignyi. The number of scales around midbody in L. dorbignyi is 48 to 59, fewer than in L. andinus, L. disjunctus, L. duellmani Cei, L. eleodori, L. erguetae, L. foxi N����ez et al., L. islugensis, L. molinai, L. multicolor, L. nigriceps Philippi, L. patriciaiturrae, L. pleopholis Laurent, L. poecilochromus, L. pulcherrimus, L. robertoi, L. rosenmanni, L. ruibali, L. signifer, and L. vallecurensis, which have 64 to 110. Females of L. dorbignyi have precloacal pores, differing from following species in which females lack precloacal pores, L. andinus (Koslowsky, 1895), L. duellmani (Cei, 1986), L. fabiani (Pincheira-Donoso and N����ez, 2005), L. islugensis (Ortiz & Marquet, 1987), L. jamesi (Pincheira-Donoso and N����ez, 2005), L. melanogaster (Lurent, 1998), L. pantherinus ((Pincheira-Donoso and N����ez, 2005), L. puritamensis N����ez and Fox, L. robertoi, L. rosenmanni, L. signifer and L. stolzmanni (Pincheira-Donoso & N����ez, 2005), and L vallecurensis. The presence of precloacal pores in females could be problematic for the diagnosis, because some species are known for few specimens, but we used this character because the new species show a high percentage of females with precloacal pores and in all the samples (from different localities), at least one female exhibit pores. The dorsal color pattern is formed by dark bands transverse to the longitudinal body axis, and which searate the laterals and the ventral color, distinguishing it from the others species of the L. montanus group. Description of the Lectotype ��� MLP.S 057���Male, SVL 95.0. Trunk length 45.8. Head length 20.7. Head width 20.3. Head height (at parietal) 15.2. Eye diameter 3.2. Interorbital distance 9.7. Orbit-auditory meatus distance 8.9. Auditory meatus height 3.7, width 1.9. Orbit-commissure of mouth distance 2.7, distance between nares 3.2. Subocular scale 4.9. Femur 14.2, tibia 17.8, foot 25.0 and humerus 9.1. Dorsal surface of the head smooth, with sixteen scales (From occiput to rostral sclae). Rostral scale wider than higher, bordered by six scales. Mental scale larger than rostral, trapezoidal, bordered by four scales. Rostral scale not in contact with nasal. Two internasals. Nasal scale surround by seven scales, separated from canthal by two scales. Five scales between rostral and supercilliaries. Six scales between rostral and frontal. Frontal horizontally divided. Two postrostrals. Interparietal smaller than or equal to parietals, surrounded by six scales. Twelve upper ciliaries. Supraorbital semicircles complete. Five supraoculars. Preocular scale separated from lorilabials row by one scale. Two projecting scales on anterior and superior margin of auditory meatus. Eight smooth horizontal temporals and six verticals. Nine lorilabials, five in contact with subocular scale. Seven supralabials, fourth larger than the others, without sense organs, not in contact with subocular scale. Four infralabials, second ventrally in contact with two scales. Four chinshields; second imbricate separated by two scales. Forty eight scales around midbody. Forty nine dorsal scales, from occiput to forelimbs, with rounded anterior margins, and slightly keeled. Twenty seven rows of scales on dorsum. Ventrals larger than dorsals, flat, juxtaposed in abdominal region, and subjuxtaposed on chest and laterals regions. Thirty five subjuxtaposed and smooth gulars, between auditory meatus. Seven precloacal pores. Antehumeral scales flat and larger than dorsals. Postauricular, rictal and longitudinal nuchal folds present. Scales on the longitudinal fold smooth. Seventeen subdigital lamellae on IV finger, and twenty four on toe IV. Third and fourth left toes absent. Flanks scales triangulars, smooth with several small granular intercalated scales. Femoral patch absent. Infracarpals scales flat, imbricate and trifid. Infratarsals scales flat, imbricate and keeled. Color in ethanol��� Head gray dorsally, with dark brown tonalities, darker than the body. Trunk, lead-gray with seven to nine dark transverse bands, faded in vertebral and paravertebral regions. Fore and hind limbs with same pattern and color as trunk. Vertebral line absent, scapular spots and dorsolateral stripes absent. Lateral region dark gray, like the dorsum, showing projections of the dorsal transverse bands. Scales iridescent or with bright color, absent. Tail autotomyzed near the base. Ventrally, from mental to cloacal region light gray, variegated, with dark and light spots. In belly region predominate dark brown spots. Variation��� Based on 4 paralectotypes and 17 additional specimens. Head longer (16.4���29.3; x = 19.8 than wider (14.7���20.3; x = 17.6). Head height 8.9���15.2, (x = 11.0). Neck wider than the head. Snout Vent Length 2.2 times longer than trunk length. Snout Vent Length 73.5���98.3, (x = 89.5). Tail length 78.0��� 117.4, (x = 96.6). Width of tail base 8.9���14.5, (x = 11.6). Humerus length 7.3���14.1, (x = 10.4). Tibial length 11.0��� 18.7, (x = 15.5). Foot length 21���27.2, (x = 24.4). Fourth finger length 8.3���11.4, (x = 9.6) and IV toe 11.4 ���16.0, (x = 13.2). Dorsal surface of the head slightly rugose, with 14���19 (x = 16.5) scales. Six to 8 supercilliaries. Twelve to 14 upper cilliaries. Eight to 10 lorilabials in a single row equal in size to supralabials. Six to 8 supralabials scales. Frontal scale azygous or horizontally divided in 2. Interparietal scale smaller, equal to, or larger than parietals, surrounded by 5���9 scales. Nasal scale surrounded by 6���8 scales. Four to 6 infralabials. Four scales in contact with the mental scale. Five to 9 smooth horizontal temporals and 4���7 verticals. Up to 4 auriculars scales on the anterior margin of the auditory meatus. Horizontal fold ���Y��� shaped, between shoulder and auditory meatus. Scales around midbody 48���59 (x = 54.7). Thirty two to 52 gulars scales and 35-45 on neck (from posterior margin of auditory meatus to shoulder, along horizontal fold). Dorsal scales between occiput and hindlimbs 45���56 (x = 50.6). Seventy seven to 94 (x = 85.9) ventrals scales. Precloacal pores in males 5���8 (x = 6.5). Three of six females with 5 to 6 precloacals pores. Gulars scales 32���52 (x = 40.3). Dorsal scales flat, imbricated and slightly keeled or unkeeled. Ventral scales same size as dorsal scales. Seventeen to 22 subdigital lamellae on the fourth finger and 21���26 on the fourth toe. Without femoral patch. FIGURE. 6. Habitat of L. dorbignyi from Sierra de Fiambal��, Catamarca, Argentina. Photo: S. Quinteros. Color in life��� Sexual dichromatism present (Fig. 2���4). Dorsal head in males, black or dark brown; gray or brown in females, but never darker than males. Dorsum and sides of the body intense yellow or dark orange, with large paravertebral spots transverse to the body axis, divided on the sides of the body (Fig. 2). Paravertebral spots black or dark brown, band-shaped. In females the dorsal background is gray or reddishbrown. Without vertebral line, dorsolateral stripes nor scapulars spots. In males the tail is dorsally light gray or greenish-gray. Ventrally yellow or intense orange from the mental scale to the cloacal region, the tail ventrally exhibit same color of dorsum (Fig. 3). Females exhibit the same pattern but lighter (Fig. 4). Distribution��� The main problem with the identification of Liolaemus dorbignyi, was the lack, in the original description, of the designation of a precise type locality. According to the results obtained in this work, the distribution of L. dorbignyi is restricted to Sierra de Fiambal, Catamarca Province, Argentina. Specifically we found populations in the localities known as Puesto la Lagunita, between 35 and 38 Km NW of Medanitos, at 4350 m, and in La Cienaga, Campo El Potrerito, 3900 m. However it is probable that L. dorbignyi is distributed along the entire Sierra de Fiambal��, over the 3500 m. (Fig. 5.) Natural History��� Liolaemus dorbignyi, inhabits large rocky areas of the Sierra de Fiambal�� (Fig. 6). It is a viviparous species. No more data about its biology are known. It lives in syntopy with Phymaturus cf. antofagastensis, L. cf. andinus and L. cf. puna., Published as part of Quinteros, Andr��s Sebasti��n, Abdala, Cristian Sim��n & Lobo, Fernando Jos��, 2008, Redescription of Liolaemus dorbignyi Koslowsky, 1898 and description of a new species of Liolaemus (Iguania: Liolaemidae), pp. 51-67 in Zootaxa 1717 on pages 52-59, DOI: 10.5281/zenodo.274176, {"references":["Koslowsky, J. (1898) Enumeracion sistematica y distribucion geografica de los reptiles argentinos. Revista del Museo de La Plata, 8, 161 - 200.","Ferraro, D. P. & Williams J. D. (2006) Material tipo de la coleccion herpetologica del Museo de La Plata, Buenos Aires, Argentina. Cuadernos de Herpetologia, 19 (2), 19 - 36.","Etheridge, R. E. (1995) Redescription of Ctenoblepharys adspersa Tschudi, 1845, and the taxonomy of Liolaeminae (Reptilia: Squamata: Tropiduridae). American Museum of Natural History Novitates, 3142, 34 pp.","Abdala, V., Abdala C. S & Tulli M. J. (2006) Muscular characters in the phylogeny of Liolaemus (Squamata: Iguania: Liolaemidae): a reappraisal. Zootaxa, 1205, 5 - 68","Abdala, C. S. (2007) Phylogeny of the boulengeri group (Iguania: Liolaemidae, Liolaemus) based on morphological and molecular characters. Zootaxa, 1538, 84 pp.","Cei, J. M. (1986) Reptiles del centro, centro-oeste y sur de la Argentina. Museo Regionale di Scienze Naturali, Torino, Monografie IV, 527 pp","Pincheira - Donoso, D. & Nunez, H. (2005) Las especies del Genero Liolaemus Wiegmann, 1834 (Iguania: Tropiduridae: Liolaeminae) Taxonomia, Sistematica y Evolucion. Publicacion Ocasional del Museo Nacional de Historia Natural, Chile, 59, 7 - 486.","Ortiz, J. C. & Marquet, P. (1987) Una nueva especie de lagarto altoandino: Liolaemus islugensis (Reptilia-Iguanidae). Guyana, Zoologia, 51, 59 - 63.","Laurent, RF. (1998) New forms of lizards of the subgenus Eulaemus of the genus Liolaemus (Reptilia: Squamata: Tropiduridae) from Peru and northern Chile. Acta Zoologica Lillooana, 44 (1), 1 - 26."]}

Liolaemus lavillai Abdala & Lobo Liolaemus ornatus Etheridge 1993: 171 (in part); Morando et al. 2004: 842 (in part). Liolaemus lavillai Abdala & Lobo 2006a: 5 (type locality: Cuesta del Acay, 3800 m, La Poma Department, Salta Province, Argentina); Abdala 2007: 44; Díaz 2007: 116., Published as part of Pincheira-Donoso, Daniel, Scolaro, J. Alejandro & Sura, Piotr, 2008, A monographic catalogue on the systematics and phylogeny of the South American iguanian lizard family Liolaemidae (Squamata, Iguania), pp. 1-85 in Zootaxa 1800 on page 51, DOI: 10.5281/zenodo.6789337, {"references":["Etheridge, R. (1993) Lizards of the Liolaemus darwinii Complex (Squamata: Iguania: Tropiduridae) in Northern Argentina. Bollettino del Museo Regionale di Scienze Naturali di Torino, 11, 137 - 199.","Morando, M., Avila, L. J., Baker, J. & Sites, J. W. (2004) Phylogeny and phylogeography of the Liolaemus darwinii complex (Squamata: Liolaemidae): evidence for introgression and incomplete lineage sorting. Evolution, 58, 842 - 861.","Abdala, C. S. & Lobo, F. (2006 a) Nueva especie del grupo de Liolaemus darwinii (Iguania: Liolaemidae) del noroeste de Argentina. Cuadernos de Herpetologia, 19, 3 - 18.","Abdala, C. S. (2007) Phylogeny of the boulengeri group (Iguania: Liolaemidae, Liolaemus) based on morphological and molecular characters. Zootaxa, 1538, 1 - 84.","Diaz, J. M. (2007) Reptilia, Iguania, Liolaeminae, Liolaemus, Puna, Prepuna, and mountain ranges, northwestern Argentina. Check List, 3, 105 - 118."]}

Liolaemus boulengeri Koslowsky Liolaemus boulengeri Koslowsky 1898: 176 (type locality: “Territories of Chubut and Neuquén, Argentina ”, restricted to the area between Esquel, Trevelín, Tecka and Gobernador Costa, Chubut Province, Argentina, by Abdala 2005a); Liebermann 1939: 71; Donoso-Barros 1970a: 180; Cei 1973a: 464, 1986: 220; Laurent 1992: 31; Núñez & Jaksic 1992: 71; Martori & Aun 1994: 97; Etheridge 1995: 34; Avila et al. 1998: 11; Etheridge & Espinoza 2000: 5; Abdala 2005a: 5; Chebez et al. 2005: 29; Scolaro 2005: 28; Sura 2005: 428; Abdala 2007: 44. Liolaemus micropholis Werner 1909 (synonym fide Donoso-Barros 1970a); Etheridge & Espinoza 2000: 6., Published as part of Pincheira-Donoso, Daniel, Scolaro, J. Alejandro & Sura, Piotr, 2008, A monographic catalogue on the systematics and phylogeny of the South American iguanian lizard family Liolaemidae (Squamata, Iguania), pp. 1-85 in Zootaxa 1800 on page 48, DOI: 10.5281/zenodo.6789337, {"references":["Koslowsky, J. (1898) Enumeracion sistematica y distribucion geografica de los reptiles argentinos. Revista del Museo de La Plata, 8, 161 - 200.","Abdala, C. S. (2005 a) Dos nuevas especies del genero Liolaemus (Iguania: Liolaemidae) y redescripcion de Liolaemus boulengeri (Koslowsky, 1898). Cuadernos de Herpetologia, 19, 3 - 33.","Liebermann, J. (1939) Catalogo sistematico y zoogeografico de los Lacertilidos argentinos. Physis, 16, 61 - 82.","Donoso-Barros, R. (1970 a) Liolaemus. In: Peters, J. A. & Donoso-Barros, R. (Ed.), Catalogue of the Neotropical Squamata. Part II. Lizards and Amphisbaenians. Bulletin United States National Museum, 297, 170 - 195.","Cei, J. M. (1973 a) Herpetologia patagonica. VI. Los Liolaemus del grupo fitzingeri en Santa Cruz y Chubut (Sauria, Iguanidae). Physis, Buenos Aires, 32, 447 - 458.","Cei, J. M. (1986) Reptiles del centro, centro-oeste y sur de la Argentina. Herpetofauna de las zonas aridas y semiaridas. Museo Regionale di Scienze Naturali di Torino, Monografie 4, 528 pp.","Laurent, R. F. (1992) On some overlooked species of the genus Liolaemus Wiegmann (Reptilia-Tropiduridae) from Peru. Breviora, 494, 1 - 33.","Nunez, H. & Jaksic, F. M. (1992) Lista comentada de los reptiles terrestres de Chile continental. Boletin del Museo Nacional de Historia Natural de Chile, 43, 63 - 91.","Martori, R. & Aun, L. (1994) Analisis comparativo de la composicion de tres comunidades de Squamata de la Sierra Grande de Cordoba, Argentina. Cuadernos de Herpetologia, 8, 97 - 103.","Etheridge, R. (1995) Redescription of Ctenoblepharys adspersa Tschudi, 1845, and the taxonomy of Liolaeminae (Reptilia: Squamata: Tropiduridae). American Museum Novitates, 3142, 1 - 34.","Avila, L. J., Acosta, J. C. & Murua, A. F. (1998) Herpetofauna de la provincia de San Juan, Argentina: lista comentada y distribucion geografica. Cuadernos de Herpetologia, 12, 11 - 29.","Etheridge, R & Espinoza, R. E. (2000) Taxonomy of the Liolaeminae (Squamata: Iguania: Tropiduridae) and a semiannotated bibliography. Smithsonian Herpetological Information Service, 126, 1 - 64.","Chebez, J. C., Rey, N. R. & Williams, J. D. (2005) Reptiles de los parques nacionales de la Argentina. Literatura of Latin America, Buenos Aires, 37 pp.","Scolaro, J. A. (2005) Reptiles patagonicos: Sur. Una guia de campo. Ediciones Universidad Nacional de la Patagonia San Juan Bosco, Trelew, 80 p.","Sura, P. 2005. Encyclopedia of extant amphibians and reptiles, Wydawnictwo Fundacja, Nowy Sacz, 544 pp.","Abdala, C. S. (2007) Phylogeny of the boulengeri group (Iguania: Liolaemidae, Liolaemus) based on morphological and molecular characters. Zootaxa, 1538, 1 - 84.","Werner, F. (1909) Neue oder seltenere Reptilien des Musee Royal d'Histoire Naturelle de Belgique in Brussel. Zoologischen Jahrbunchern, 28, 263 - 288."]}

Liolaemus crepuscularis sp. nov 1993, Liolaemus ornatus (part), Etheridge, R. Boll. Mus. Reg. Sci. nat. Torino. 11 (1): 137–199. 2005, Liolaemus sp. 5. Abdala, C. S. Doctoral Thesis, UNT. 277 pp. 2006, Liolaemus sp. Mina Capillitas, Abdala C. S. & Lobo F. Cuad. Herpetol. 19 (2) 3­18 Holotype. FML 16622 (FBC 415). Adult male, from Puesto Flores, 2 km South from Mina Capillitas, Ruta Provincial 47 (km 50 a 52), 3100 m Departamento Andalgalá, provincia de Catamarca, Argentina. Collected by C. Abdala, Quinteros S., Scrocchi G. and S. Barrionuevo. March 3, 2005. Paratypes. FML 16623 (FBC 414; 416–418). Three males, one female, same data as holotype. FML 16627 – 29 (CSA 527–528; 530). Two males, one female, from Puesto Flores, 2 km South of Mina Capillitas, Ruta Provincial 47 (km 50 a 52), 3100 m, Departamento Andalgalá, provincia de Catamarca, Argentina. Collected by C. Abdala and S. Barrionuevo. November 12, 2004. FML 3611. One male, two females and one juvenile, form Mina Capillitas, 2800 m, Departamento Andalgalá, provincia de Catamarca, Argentina. Collected by F. Lobo, Marcus A. and R. Montero. December 18, 1995. MCN 1937–1939 (CSA 529; 531–532). Two males, one female, from Puesto Flores, 2 km South of Mina Capillitas, Ruta Provincial 47 (km 50 a 52), 3100 m, Departamento Andalgalá, provincia de Catamarca, Argentina. Collected by C. Abdala and S. Barrionuevo. November 12, 2004. Etymology. The specific epithet crepuscularis refers to the brown­red mixed with blue coloration of the flanks, reminiscent of the colours of nightfall. This word derives from the Latin “crepusculum” which means “twilight.” Diagnosis. Liolaemus crepuscularis is a member of the L. boulengeri group diagnosed by a patch of enlarged scales on the posterior thigh (Etheridge, 1995). Within this group it is a member of the L. laurenti group (Abdala, 2005), diagnosed by posterior marginal teeth with straight­side tooth crowns, a dark stripe through the eye, and marked sexual dichromatism. Tooth morphology and the dark stripe through the eye distinguish this new species from the species in the L. melanops group (Abdala, 2005) (L. boulengeri Koslowsky, L canqueli Cei & Scolaro, L. cuyanus Cei & Scolaro, L. donosobarrosi Cei, L. fitzingerii (Dumeril & Bibrón), L. inacayali Abdala, L. josei Abdala, L. mapuche Abdala, L. martorii Abdala, L. melanops (Bursmeister), L. morenoi Etheridge & Christie, L. loboi Abdala, L. sagei Etheridge & Christie, L. rothi Koslowsky, L. senguer Abdala, L. tehuelche Abdala, L. telsen Cei & Scolaro and L. xanthoviridis Cei & Scolaro, and sexual dichromatism distinguishes L. crepuscularis from L canqueli, L. cuyanus, L. fitzingerii, L. martorii, L. morenoi, L. sagei, and L. xanthoviridis. The lack of cephalic melanism distinguish L. crepuscularis from L. canqueli and L. melanops, and the absence of melanic gular or antehumeral collar separates it from L. canqueli, L. cuyanus, L. mapuche, L. morenoi, L. fitzingerii and L. xanthoviridis. Liolaemus crepuscularis has four scales in contact with the mental, not six as L. cuyanus and the species of the L. wiegmannii group Etheridge (L. arambarensis Verrastro et al, L. azarai Avila, L. lutzae Mertens, L. multimaculatus (Dumeril & Bibrón), L. occipitalis Boulenger, L. rabinoi (Cei), L. riojanus Cei, L. salinicola Laurent, L. wiegmannii (Dumeril & Bibrón) y L. scapularis Laurent), or variation from four to six scales as in L. josei and L. mapuche. One row of lorilabials, not two, distinguishes the new species from the species of the L. wiegmannii group. Liolaemus crepuscularis differs from the species of the L. anomalus group (L. anomalus Koslowsky, L. pseudoanomalus Cei, L. ditadai Cei, and L. lentus Gallardo) (Abdala, 2005) by having caudal autotomy, tail longer than body, head longer than wider, palpebral ‘comb’ not developed, a larger number of precloacal pores in both males and females, and the absence of pterygoid teeth. Within the L. laurenti group (Abdala, 2005), L. crepuscularis is part of the L. ornatus group (with L. albiceps Lobo & Laurent, L. calchaqui Lobo & Kretzschmar, L. irregularis Laurent, L. ornatus and L. lavillai). These species differs from L. abaucan Etheridge, L. chacoensis Shreve, L. darwinii Bell, L. grosseorum Etheridge, L. koslowskyi Etheridge, L. laurenti Etheridge, L. olongasta Etheridge, L. quilmes Etheridge and L. uspallatensis Macola & Castro, by having a larger number of precloacal pores in females and because of being viviparous. Within the L. ornatus group, it differs from L. albiceps and L. irregularis in having a shorter maximum snout­vent length (L. crepuscularis mean SVL 64.0 mm in comparison to 82.5 mm of L. albiceps and 86.1 mm of L. irregularis), and females with fewer precloacal pores (= 6.5 vs. = 7.5 y = 8.4 respectively). Liolaemus crepuscularis is distinguished from L. ornatus in having males with background coloration of dorsum light brown to ferrous red and head coloration darker than the rest of the body, paravertebral markings more rounded and never in contact, dorsolateral strips less marked, body and tail with more light blue scales, little dark brown spots on the throat, belly white or light red, and lateral fields of intense ferrous red. Liolaemus crepuscularis is distinguished from L. calchaqui in having a larger body size (maximum SVL 64.0 mm vs. 57.8 mm) and different color pattern, with rounded or quadrangular paravertebrals markings well defined, light blue scales on the lateral body and tail, and lateral fields red, a pattern not present in L. calchaqui. L. crepuscularis is distinguished from L. lavillai by the dorsal coloration (brown in L. lavillai) and the absence of light blue scales in L. lavillai. Liolaemus crepuscularis males have a characteristic dorsal color pattern, with dorsal field light brown or ferrous red, dorsolateral stripes with yellow and light blue scales, and dark brown prescapular spot similar in size and color to the lateral markings. This set of characters of the color pattern differentiates it from the remaining species of the L. laurenti group. The presence of the prescapular spot similar to the lateral markings distinguish L. crepuscularis from species where this mark better defined (L. abaucan, L. darwinii, L. espinozai Abdala, L. grosseorum, L. koslowskyi and L. quilmes) and from species where it is absent (L. albiceps, L. calchaqui, L. irregularis, L. ornatus). In L. crepuscularis the black antehumeral arch typical of L. darwinii, L. laurenti, L. grosseorum and L. olongasta is absent. Description of holotype. Adult male. Snout­vent length 56.7 mm. Head 1.2 times longer (12.4 mm) than wider (10.5 mm). Head height 6.7 mm. Neck 8.5 mm. Eye diameter 3.2 mm. Orbit­auditory meatus distance 4.7 mm. Auditory meatus oval shaped (height 2.7 mm, width 1.6 mm). Orbit­comisure of mouth distance 1.4 mm, distance between nares 2.4 mm. Subocular 3.6 mm. long. Axilla­groin distance 24.5 mm, tail length 96.6 mm. Body width 13.2 mm. Humerus length 8.3 mm, forearm length 6.5 mm, manus 9.3 mm. Pygal region 6.9 mm, vent width 7.4 mm. Dorsal surface of the head scales slightly rugose, with fifteen dorsal scales. Rostral wider than higher and surrounded by six scales. Mental trapezoidal in shape, larger than rostral, and surrounded by four scales. Nasal and rostral scales not in contact. Four internasals longer than wide. Six scales surrounding nasal. Canthal scale separated from nasal by two scales, one very small in contact with nasal, one larger in contact with canthal. Five scales between rostral and supercilliares, seven between rostral and frontal. Frontal divided in four. Four postrostrals. Intreparietal in contact with seven scales, and slightly smaller than adjacent right parietal, larger than left parietal. Preocular scale separated from lorilabials by one scale. Three scales on the anterior margin of the auditory meatus, superior margin with one differentiated scale, upper anterior margin with one scale projected outwards. Eight horizontal temporals and seven verticals, unkeeled. Subocular in contact with four lorilabials, not in contact with fourth supralabial. Six supraoculars. Eight supralabials, fourth equal or slightly larger than third, fifth and sixth, without scale organs. Nine lorilabials. Five infralabials, the second in contact with two scales. Four chinshields, the second separated by two scales. Scales around midbody: 57. Dorsal scales between occiput and groin: 65, keeled, imbricated, with round posterior margin. Twenty six rows of scales on dorsum. Ventral scales imbricated, laminar, larger than dorsals. Twenty nine gulars, imbricated, unkeeled. Without gular fold. Eight precloacal pores. Antehumeral scales subtriangular, enlarged, well differentiated. Postauricular, rictal, longitudinal and antehumeral folds well marked. Longitudinal fold scales flat, unkeeled. Subdigital lamellae of fourth finger, 15. Subdigital lamellae of fourth toe, 22. Scales of the flanks with a little apical notch. Femoral patch formed by 23 conic, pointed scales. Infracarpals and infratarsals laminar, imbricated, trifid. Tricuspid teeth, with straight crowns. Color in life. Dorsal background color of head light brown, with dark spots irregularly distributed. Head not darker than body. Black stripe transverse to orbit. Wide black spot from the posterior margin of orbit to occiput. Dorsal background color ferrous red, two series of well marked, black paravertebral spots in form of crescent, with yellow scales in the posterior margin of spots. Light yellow vertebral field, barely evident. Black prescapular spot well defined, anterior to it, on right side of the body there is a smaller black spot, on left side these spots are united by a small longitudinal black spot. Three diffuse spots posterior to prescapular spot. Background color of flanks equal to dorsum, with light blue, yellow and white scales distributed irregularly. Limbs grey with black spots. Forelimbs with a black stripe, similar to a bracelet. Tail light gray dorsally, paravertebral spots become diffuse and fused at base of tail, flanks with numerous light blue scales. Ventral coloration light yellow turning to light brown with dark brown marks on throat. Forelimbs, vent, and tail same color as throat. Intense yellow colored hindlimbs, especially femoral region. Two rows of small dark grey markings on tail. Variation. Based on 41 individuals, 23 males, 17 females. Squamation. Dorsal surface of head slightly rugose. Nasal surrounded by 6–7 scales; 6–9 supralabials; 5–8 lorilabials in one row; 6–8 supraoculars. Frontal undivided or divided in 2, 3 or 4. Parietals larger than or equal to interparietals, surrounded by 5–9 scales. Scales between rostral and occiput 13–17 scales (= 15.6). Mental scale in contact with four scales. Infralabials 4–6. Gulars 23–30 (= 25.7). Temporals unkeeled; 7–11 longitudinal temporals; 7–10 transversal. Auditory meatus always higher (= 2.5 mm) than wider (= 1.6 mm), with distinctive scales on the margins, 1–3 on anterior margin and 0–2 in posterior margin. One scale projecting outwards at posterior upper margin of auditory meatus. Neck folds well developed. Head slender, longer (= 11.5 mm) than wider (= 9.7 mm), height (= 7.5 mm). Neck narrower than head and trunk. Trunk smaller (= 22.6 mm) than half SVL (= 27.3 mm). Body slender. SVL in males and females 56–64 mm (= 54.5 mm). Humerus length (= 7.8 mm), width (= 2.8 mm). Radium length (= 7.1 mm). Manus length (= 8.0 mm). Femur length (= 9.5 mm). Scales around midbody in males and females, 56–64 (= 59.7). Dorsal scales between occiput and hindlimb 59–72 (= 65.8). Dorsal scales laminar, imbricate and keeled. Subdigital lamellae of fourth finger 13–18 (= 15.1). Subdigital lamellae of fourth toe 22–25 (= 23.1), with two or three keels. Infracarpals and infratarsals scales laminar, trifid, imbricated. Femoral patch with 20–27 conic, pointed scales. Ventral scales larger than dorsals. Tail 1.5–1.7 times longer than SVL. Males with 6–9 (= 7.4) precloacal pores. Females with 5–7 (= 6.5) precloacal pores. Coloration (Fig. 1 a). Head light brown or grey, with small dark markings on the dorsal surface. As in all the members in the L. laurentii group, both males and females have a dark transverse stripe across the orbits (Abdala, 2005). Liolaemus crepuscularis has marked sexual dicromatism, with the males having distinctive and evident coloration characters that distinguishes it from related species. The characteristic dorsal coloration of this new species is a brown to ferrous red background coloration that in some individuals may change to yellow. Flanks usually have the same color as the dorsum; some individuals have intense yellow or light brown flanks. In most specimens the markings in the form of crescent posterior to the prescapular spot are diffuse. In females the background dorsal color is grey or light brown, paravertebral markings dark grey or dark brown, in the form of a crescent, with white scales on the posterior margin. During the reproductive season, in females, the flanks and neck turn yellow. Ventrally are usually white or light yellow on the flanks, with fewer small markings on the throat than males. Belly, vent and hindlimbs are immaculate. Distribution (Fig. 2). Liolaemus crepuscularis is known only from the type locality and near localities, always above 2800 m. Natural History. Liolaemus crepuscularis inhabits the southwestern slopes of the Nevados del Aconquija in Catamarca, always above 2800 m. This region presents an arid climate, with annual precipitation less than 300 mm, most of which is snow. The habitat is typical of the Prepuna region (Cabrera and Willink, 1980), Festuca sp., cola de zorro (Pappophorum sp.) and muña muña (Satureja parvifolia) is the dominant flora. At the site there are numerous rocks, which are used by L. crepuscularis as refuge. Liolaemus crepuscularis is sympatric with L. ramirezae, and L. capillitas. Usually L. ramirezae is associated with Festuca sp, L. capillitas with larger rocks, and L. crepuscularis with the smaller ones. At this site also is found the snake Tachymenis peruviana,. Liolaemus crepuscularis is viviparous, but nothing more is known of its biology.

Liolaemus crepuscularis sp. nov 1993, Liolaemus ornatus (part), Etheridge, R. Boll. Mus. Reg. Sci. nat. Torino. 11 (1): 137���199. 2005, Liolaemus sp. 5. Abdala, C. S. Doctoral Thesis, UNT. 277 pp. 2006, Liolaemus sp. Mina Capillitas, Abdala C. S. & Lobo F. Cuad. Herpetol. 19 (2) 3��18 Holotype. FML 16622 (FBC 415). Adult male, from Puesto Flores, 2 km South from Mina Capillitas, Ruta Provincial 47 (km 50 a 52), 3100 m Departamento Andalgal��, provincia de Catamarca, Argentina. Collected by C. Abdala, Quinteros S., Scrocchi G. and S. Barrionuevo. March 3, 2005. Paratypes. FML 16623 (FBC 414; 416���418). Three males, one female, same data as holotype. FML 16627 ��� 29 (CSA 527���528; 530). Two males, one female, from Puesto Flores, 2 km South of Mina Capillitas, Ruta Provincial 47 (km 50 a 52), 3100 m, Departamento Andalgal��, provincia de Catamarca, Argentina. Collected by C. Abdala and S. Barrionuevo. November 12, 2004. FML 3611. One male, two females and one juvenile, form Mina Capillitas, 2800 m, Departamento Andalgal��, provincia de Catamarca, Argentina. Collected by F. Lobo, Marcus A. and R. Montero. December 18, 1995. MCN 1937���1939 (CSA 529; 531���532). Two males, one female, from Puesto Flores, 2 km South of Mina Capillitas, Ruta Provincial 47 (km 50 a 52), 3100 m, Departamento Andalgal��, provincia de Catamarca, Argentina. Collected by C. Abdala and S. Barrionuevo. November 12, 2004. Etymology. The specific epithet crepuscularis refers to the brown��red mixed with blue coloration of the flanks, reminiscent of the colours of nightfall. This word derives from the Latin ���crepusculum��� which means ���twilight.��� Diagnosis. Liolaemus crepuscularis is a member of the L. boulengeri group diagnosed by a patch of enlarged scales on the posterior thigh (Etheridge, 1995). Within this group it is a member of the L. laurenti group (Abdala, 2005), diagnosed by posterior marginal teeth with straight��side tooth crowns, a dark stripe through the eye, and marked sexual dichromatism. Tooth morphology and the dark stripe through the eye distinguish this new species from the species in the L. melanops group (Abdala, 2005) (L. boulengeri Koslowsky, L canqueli Cei & Scolaro, L. cuyanus Cei & Scolaro, L. donosobarrosi Cei, L. fitzingerii (Dumeril & Bibr��n), L. inacayali Abdala, L. josei Abdala, L. mapuche Abdala, L. martorii Abdala, L. melanops (Bursmeister), L. morenoi Etheridge & Christie, L. loboi Abdala, L. sagei Etheridge & Christie, L. rothi Koslowsky, L. senguer Abdala, L. tehuelche Abdala, L. telsen Cei & Scolaro and L. xanthoviridis Cei & Scolaro, and sexual dichromatism distinguishes L. crepuscularis from L canqueli, L. cuyanus, L. fitzingerii, L. martorii, L. morenoi, L. sagei, and L. xanthoviridis. The lack of cephalic melanism distinguish L. crepuscularis from L. canqueli and L. melanops, and the absence of melanic gular or antehumeral collar separates it from L. canqueli, L. cuyanus, L. mapuche, L. morenoi, L. fitzingerii and L. xanthoviridis. Liolaemus crepuscularis has four scales in contact with the mental, not six as L. cuyanus and the species of the L. wiegmannii group Etheridge (L. arambarensis Verrastro et al, L. azarai Avila, L. lutzae Mertens, L. multimaculatus (Dumeril & Bibr��n), L. occipitalis Boulenger, L. rabinoi (Cei), L. riojanus Cei, L. salinicola Laurent, L. wiegmannii (Dumeril & Bibr��n) y L. scapularis Laurent), or variation from four to six scales as in L. josei and L. mapuche. One row of lorilabials, not two, distinguishes the new species from the species of the L. wiegmannii group. Liolaemus crepuscularis differs from the species of the L. anomalus group (L. anomalus Koslowsky, L. pseudoanomalus Cei, L. ditadai Cei, and L. lentus Gallardo) (Abdala, 2005) by having caudal autotomy, tail longer than body, head longer than wider, palpebral ���comb��� not developed, a larger number of precloacal pores in both males and females, and the absence of pterygoid teeth. Within the L. laurenti group (Abdala, 2005), L. crepuscularis is part of the L. ornatus group (with L. albiceps Lobo & Laurent, L. calchaqui Lobo & Kretzschmar, L. irregularis Laurent, L. ornatus and L. lavillai). These species differs from L. abaucan Etheridge, L. chacoensis Shreve, L. darwinii Bell, L. grosseorum Etheridge, L. koslowskyi Etheridge, L. laurenti Etheridge, L. olongasta Etheridge, L. quilmes Etheridge and L. uspallatensis Macola & Castro, by having a larger number of precloacal pores in females and because of being viviparous. Within the L. ornatus group, it differs from L. albiceps and L. irregularis in having a shorter maximum snout��vent length (L. crepuscularis mean SVL 64.0 mm in comparison to 82.5 mm of L. albiceps and 86.1 mm of L. irregularis), and females with fewer precloacal pores (= 6.5 vs. = 7.5 y = 8.4 respectively). Liolaemus crepuscularis is distinguished from L. ornatus in having males with background coloration of dorsum light brown to ferrous red and head coloration darker than the rest of the body, paravertebral markings more rounded and never in contact, dorsolateral strips less marked, body and tail with more light blue scales, little dark brown spots on the throat, belly white or light red, and lateral fields of intense ferrous red. Liolaemus crepuscularis is distinguished from L. calchaqui in having a larger body size (maximum SVL 64.0 mm vs. 57.8 mm) and different color pattern, with rounded or quadrangular paravertebrals markings well defined, light blue scales on the lateral body and tail, and lateral fields red, a pattern not present in L. calchaqui. L. crepuscularis is distinguished from L. lavillai by the dorsal coloration (brown in L. lavillai) and the absence of light blue scales in L. lavillai. Liolaemus crepuscularis males have a characteristic dorsal color pattern, with dorsal field light brown or ferrous red, dorsolateral stripes with yellow and light blue scales, and dark brown prescapular spot similar in size and color to the lateral markings. This set of characters of the color pattern differentiates it from the remaining species of the L. laurenti group. The presence of the prescapular spot similar to the lateral markings distinguish L. crepuscularis from species where this mark better defined (L. abaucan, L. darwinii, L. espinozai Abdala, L. grosseorum, L. koslowskyi and L. quilmes) and from species where it is absent (L. albiceps, L. calchaqui, L. irregularis, L. ornatus). In L. crepuscularis the black antehumeral arch typical of L. darwinii, L. laurenti, L. grosseorum and L. olongasta is absent. Description of holotype. Adult male. Snout��vent length 56.7 mm. Head 1.2 times longer (12.4 mm) than wider (10.5 mm). Head height 6.7 mm. Neck 8.5 mm. Eye diameter 3.2 mm. Orbit��auditory meatus distance 4.7 mm. Auditory meatus oval shaped (height 2.7 mm, width 1.6 mm). Orbit��comisure of mouth distance 1.4 mm, distance between nares 2.4 mm. Subocular 3.6 mm. long. Axilla��groin distance 24.5 mm, tail length 96.6 mm. Body width 13.2 mm. Humerus length 8.3 mm, forearm length 6.5 mm, manus 9.3 mm. Pygal region 6.9 mm, vent width 7.4 mm. Dorsal surface of the head scales slightly rugose, with fifteen dorsal scales. Rostral wider than higher and surrounded by six scales. Mental trapezoidal in shape, larger than rostral, and surrounded by four scales. Nasal and rostral scales not in contact. Four internasals longer than wide. Six scales surrounding nasal. Canthal scale separated from nasal by two scales, one very small in contact with nasal, one larger in contact with canthal. Five scales between rostral and supercilliares, seven between rostral and frontal. Frontal divided in four. Four postrostrals. Intreparietal in contact with seven scales, and slightly smaller than adjacent right parietal, larger than left parietal. Preocular scale separated from lorilabials by one scale. Three scales on the anterior margin of the auditory meatus, superior margin with one differentiated scale, upper anterior margin with one scale projected outwards. Eight horizontal temporals and seven verticals, unkeeled. Subocular in contact with four lorilabials, not in contact with fourth supralabial. Six supraoculars. Eight supralabials, fourth equal or slightly larger than third, fifth and sixth, without scale organs. Nine lorilabials. Five infralabials, the second in contact with two scales. Four chinshields, the second separated by two scales. Scales around midbody: 57. Dorsal scales between occiput and groin: 65, keeled, imbricated, with round posterior margin. Twenty six rows of scales on dorsum. Ventral scales imbricated, laminar, larger than dorsals. Twenty nine gulars, imbricated, unkeeled. Without gular fold. Eight precloacal pores. Antehumeral scales subtriangular, enlarged, well differentiated. Postauricular, rictal, longitudinal and antehumeral folds well marked. Longitudinal fold scales flat, unkeeled. Subdigital lamellae of fourth finger, 15. Subdigital lamellae of fourth toe, 22. Scales of the flanks with a little apical notch. Femoral patch formed by 23 conic, pointed scales. Infracarpals and infratarsals laminar, imbricated, trifid. Tricuspid teeth, with straight crowns. Color in life. Dorsal background color of head light brown, with dark spots irregularly distributed. Head not darker than body. Black stripe transverse to orbit. Wide black spot from the posterior margin of orbit to occiput. Dorsal background color ferrous red, two series of well marked, black paravertebral spots in form of crescent, with yellow scales in the posterior margin of spots. Light yellow vertebral field, barely evident. Black prescapular spot well defined, anterior to it, on right side of the body there is a smaller black spot, on left side these spots are united by a small longitudinal black spot. Three diffuse spots posterior to prescapular spot. Background color of flanks equal to dorsum, with light blue, yellow and white scales distributed irregularly. Limbs grey with black spots. Forelimbs with a black stripe, similar to a bracelet. Tail light gray dorsally, paravertebral spots become diffuse and fused at base of tail, flanks with numerous light blue scales. Ventral coloration light yellow turning to light brown with dark brown marks on throat. Forelimbs, vent, and tail same color as throat. Intense yellow colored hindlimbs, especially femoral region. Two rows of small dark grey markings on tail. Variation. Based on 41 individuals, 23 males, 17 females. Squamation. Dorsal surface of head slightly rugose. Nasal surrounded by 6���7 scales; 6���9 supralabials; 5���8 lorilabials in one row; 6���8 supraoculars. Frontal undivided or divided in 2, 3 or 4. Parietals larger than or equal to interparietals, surrounded by 5���9 scales. Scales between rostral and occiput 13���17 scales (= 15.6). Mental scale in contact with four scales. Infralabials 4���6. Gulars 23���30 (= 25.7). Temporals unkeeled; 7���11 longitudinal temporals; 7���10 transversal. Auditory meatus always higher (= 2.5 mm) than wider (= 1.6 mm), with distinctive scales on the margins, 1���3 on anterior margin and 0���2 in posterior margin. One scale projecting outwards at posterior upper margin of auditory meatus. Neck folds well developed. Head slender, longer (= 11.5 mm) than wider (= 9.7 mm), height (= 7.5 mm). Neck narrower than head and trunk. Trunk smaller (= 22.6 mm) than half SVL (= 27.3 mm). Body slender. SVL in males and females 56���64 mm (= 54.5 mm). Humerus length (= 7.8 mm), width (= 2.8 mm). Radium length (= 7.1 mm). Manus length (= 8.0 mm). Femur length (= 9.5 mm). Scales around midbody in males and females, 56���64 (= 59.7). Dorsal scales between occiput and hindlimb 59���72 (= 65.8). Dorsal scales laminar, imbricate and keeled. Subdigital lamellae of fourth finger 13���18 (= 15.1). Subdigital lamellae of fourth toe 22���25 (= 23.1), with two or three keels. Infracarpals and infratarsals scales laminar, trifid, imbricated. Femoral patch with 20���27 conic, pointed scales. Ventral scales larger than dorsals. Tail 1.5���1.7 times longer than SVL. Males with 6���9 (= 7.4) precloacal pores. Females with 5���7 (= 6.5) precloacal pores. Coloration (Fig. 1 a). Head light brown or grey, with small dark markings on the dorsal surface. As in all the members in the L. laurentii group, both males and females have a dark transverse stripe across the orbits (Abdala, 2005). Liolaemus crepuscularis has marked sexual dicromatism, with the males having distinctive and evident coloration characters that distinguishes it from related species. The characteristic dorsal coloration of this new species is a brown to ferrous red background coloration that in some individuals may change to yellow. Flanks usually have the same color as the dorsum; some individuals have intense yellow or light brown flanks. In most specimens the markings in the form of crescent posterior to the prescapular spot are diffuse. In females the background dorsal color is grey or light brown, paravertebral markings dark grey or dark brown, in the form of a crescent, with white scales on the posterior margin. During the reproductive season, in females, the flanks and neck turn yellow. Ventrally are usually white or light yellow on the flanks, with fewer small markings on the throat than males. Belly, vent and hindlimbs are immaculate. Distribution (Fig. 2). Liolaemus crepuscularis is known only from the type locality and near localities, always above 2800 m. Natural History. Liolaemus crepuscularis inhabits the southwestern slopes of the Nevados del Aconquija in Catamarca, always above 2800 m. This region presents an arid climate, with annual precipitation less than 300 mm, most of which is snow. The habitat is typical of the Prepuna region (Cabrera and Willink, 1980), Festuca sp., cola de zorro (Pappophorum sp.) and mu��a mu��a (Satureja parvifolia) is the dominant flora. At the site there are numerous rocks, which are used by L. crepuscularis as refuge. Liolaemus crepuscularis is sympatric with L. ramirezae, and L. capillitas. Usually L. ramirezae is associated with Festuca sp, L. capillitas with larger rocks, and L. crepuscularis with the smaller ones. At this site also is found the snake Tachymenis peruviana,. Liolaemus crepuscularis is viviparous, but nothing more is known of its biology., Published as part of Abdala, Cristian Sim��n & G��mez, Juan Manuel D��az, 2006, A new species of the Liolaemus darwinii group (Iguania: Liolaemidae) from Catamarca Province, Argentina, pp. 21-33 in Zootaxa 1317 on pages 22-29, DOI: 10.5281/zenodo.173968

We examined the Liolaemus jamesi holotype (NHMUK 1946.8.12.39) and 20 additional specimens from the Tarapac�� Region (Appendix I). The characteristics for scalation were taken according to Etheridge (1995), Lobo (2005) and Quinteros et al. (2008). The lateral head scalation was taken from the left side. Historical data on the L. jamesi type locality was taken from published field notes of the Ambrose A. Lane field trip (Lane & Sclater 1897); the elevation and locality coordinates were estimated using Google Earth. In regards to the type locality, according to Lane and Sclater (1897, p. 10) the journey of Lane in the Tarapac�� Province began at Iquique Port (20��13`S ��� 70��08`W, 30 masl) and then he travelled by the following trajectory outward and by way of return (Fig. 1): Pica (20��28`S ��� 69��18`W, 1300 masl), Pampa de Huasco (20��21`S ��� 69��01`W, 4000 masl), Laguna (Salar) de Huasco (20��17`S ��� 68��54`W, 3900 masl), Pampa de Sacaya (20��00`S ��� 68��35`W, 4300 masl) and Cancosa (19��51`S ��� 68��36`W, 3950 m). Neither Boulenger (1891) nor Lane and Sclater (1897) stated the specific location from which the holotype L. jamesi was collected and there had been no new records of this species recorded from this area for more than 100 years. In fact, the first and only mention of L. jamesi specimens collected from this area is the record from Salar del Huasco (Moreno et al. 2001), which is at 3900 masl, Boulenger (1891) did however estimate the altitude in which the holotype was collected at 3050-3660 masl. Ruiz De Gamboa and Ortiz (2016) reviewed specimens of L. jamesi from all known distributions, among them one voucher specimen from Altos de Pica (20��17`S ��� 69��05`W, 3545 mals), road to Salar del Huasco, which matched Boulenger`s (1891) altitude range. Moreover, we collected specimens of L. jamesi (MUAP 122���24, Fig. 2) at 17 km W from Laguna del Huasco at 3800 masl (20��17`S ��� 69��03`W). Hence, we are certain that this species occurs in this region at higher elevations. We propose a restriction of the type locality of L. jamesi to: road to Salar del Huasco, between 3545 and 3660 masl, Tarapac�� Region, Chile, because 1) this transect was explored by the holotype collector Ambrose A. Lane; 2) the other places visited by Lane are out of the elevation range provided by Boulenger (1891); 3) There are records of this species from the Tarapac�� Region in Salar del Huasco (Moreno et al. 2001) and the road to this place (Ruiz De Gamboa and Ortiz, 2016); and 4) we found specimens at this altitude range matching with the data provided by Boulenger (1891). In regards to the holotype (Fig. 2), the measurements and scalation according to Boulenger (1891) are: Adult male. SVL: 100 mm. Tail length (end lost): 90 mm. Head length: 23 mm. Head width: 20 mm. Forelimb length: 47 mm. Hindlimb length: 66 mm. Midbody scales: 52. The features observed from digital photographs are as follows: Two postrostrals. Two internasals. Hexagonal interparietal scale, with a central, small, and opaque ������parietal eye������ in the center. The interparietal scale is smaller than the parietals and is surrounded by seven scales. Ten scales between the interparietal scale and the rostral scale. Twenty one scales between the occiput and the rostral. Orbital semicircle incomplete on the right side and formed by 16 scales on the left side; nine supraoculars (enlarged) on right side and seven on the left side; seven supercilliaries. Frontal divided into five scales (1���1���1���2, anterior to posterior). Two scales between the nasal and the canthal. Unfragmented preocular, separated from supralabials by one row of lorilabials. The nasal is separated from the rostral by one scale and is surrounded by seven scales. There is one row of lorilabials between the supralabials and the subocular. There are nine supralabials and seven infralabial scales. The mental shape is irregular. There are five pairs of postmental shields; but the first postmental scale on the right side is not discernible. Temporal scales are juxtaposed, smooth and with some interstitial granules. There are nine temporal scales between the level of the supercilliaries and commissure of the mouth. There are two small scales slightly projected to the auditory meatus, which do not cover it. The auricular scale is not differentiated. There is a well-developed ���Y��� shaped lateral neck fold, well-developed antehumeral fold and dorso-lateral fold. No gular fold. Dorsal scales are rounded, subimbricate, smooth or slightly keeled, with interstitial granules. The interstitial granules are more abundant on the flanks. Dorsal scales are larger than the ventrals. Dorsal scales: 45, 46 according to Boulenger (1891). Ventral scales are rounded to rhomboidal, smooth, imbricated, with some interstitial granules. Ventral scales: 88. There are six precloacal pores. There is a ventral incision, probably for organ examination, running from the chest to the midpoint of the belly. Suprafemoral and infrafemoral scales are rounded, imbricate, and smooth. Supraantebrachials scales are rounded, imbricate, and smooth. Infraantebrachials scales are rounded, imbricate, and smooth to conical towards the axilla. The dorsal scales of the tail are rounded, subimbricate, weakly keeled, with interstitial granules. The ventral scales of the tail vary from rounded to rhomboidal or triangular, and are imbricate and smooth. Lamellae of fourth finger 23 and lamellae of fourth toe 29 (right hand). Light brown head, with dark brown spots on the snout, occipital and circumorbital zones. The loreal and temporal zones are light grey. The anterior and middle zones of the subocular scale are grey, but there is a dark brown vertical stripe running from the eigth supralabial to the postocular scale (passing over the posterior zone of the subocular). The dorsal colour is light brown on the right hand side and grey on the left hand side (by deterioration). Some dorsal scales have black spots on the anterior border and there is a black coloured space between the dorsal scales. The keel on the dorsal scales (when it is present) is not black spotted. The flanks are light grey, with a black in between the scales. The dorsal surface of the limbs and the dorsal surface of the tail are both light grey, with black in between the scales. Some scales on the tail (especially in the middle of the tail) is brown shaded in colour. The tip of the tail is brown. The throat, chest and belly are dark brown, with a few light grey scales dispersed. The ventral surface of the hind limbs is light grey in colour with some brown spots. The cloaca is light grey with dark brown in the middle. Precloacal pores are orange. The ventral surface of the tail is light grey with some dark brown spots and a complete dark brown vertebral stripe. While it is not possible determine the exact type locality, this has also been the case for many other old descriptions of Liolaemus and other South American lizards, the reason being because the descriptions commonly tended to be incomplete or imprecise. Recent publications have been successful in restricting the type locality of some poorly known southernmost South American lizards to bounded geographical ranges, based on the historical data of the field notes, with the goal of producing characterizations and delimitations of the species and their ranges (Lobo & Etheridge 2013; Quinteros et al. 2008; Troncoso-Palacios & Garin 2013; Troncoso-Palacios et al. 2013; Troncoso-Palacios 2018). In this case, the restriction of the type locality of L. jamesi will allow determination of further morphological and molecular characterization of this taxon, such that it can be compared to species which show some morphologically similarity, especially with the taxa previously considered to be subspecies of L. jamesi or currently considered synonyms of it: L. aymararum Veloso, Sallaberry, Navarro, Iturra, Valencia, Penna & D��az, 1982, considered to be a junior synonym of L. jamesi by Pincheira-Donoso and N����ez (2005) and L. jamesi pachecoi Laurent, 1995, later raised to full species by Langstroth (2011). Moreover, Langstroth (2011), based on the mtDNA data by Schulte and Moreno-Roark (2010), pointed out that mtDNA divergence between L. aymararum and L. pachecoi is enough to recognize both as full species. Furthermore, a recent mt + nuclear DNA phylogeny (Aguilar-Puntriano et al. 2018) found that L. jamesi is neither the sister species of L. aymararum nor L. pachecoi. We hope that our proposed restriction of the type locality of L. jamesi and the characterization of additional material from its vicinity will facilitate further research on these issues., Published as part of Troncoso-Palacios, Jaime, Gamboa, Margarita Ruiz De & Campbell, Patrick D., 2019, Liolaemus jamesi (Boulenger, 1891): restriction of the type locality and holotype characterization (Squamata: Liolaemidae), pp. 442-446 in Zootaxa 4612 (3) on pages 442-444, DOI: 10.11646/zootaxa.4612.3.12, http://zenodo.org/record/3993643, {"references":["Etheridge, R. E. (1995) Redescription of Ctenoblepharys adspersa Tschudi, 1845, and thetaxonomy of Liolaeminae (Reptilia: Squamata: Tropiduridae). American Museum Novitates, 3142, 1 - 34.","Lobo, F. (2005) Las relaciones filogeneticas dentro grupo chiliensis (Iguania: Liolaemidae: Liolaemus): sumando nuevos caracteres y taxones. Acta Zoologica Lilloana, 49, 65 - 87.","Quinteros, A. S., Abdala, C. S. & Lobo, F. J. (2008) Redescription of Liolaemus dorbignyi Koslowsky, 1898 and description of a new species of Liolaemus (Iguania: Liolaemidae). Zootaxa, 1717, 51 - 67.","Lane, A. A. & Sclater, P. L. (1897) Field-notes on the birds of Chili. By Ambrose A. Lane. With an introduction and remarks by P. L. Sclater. The Ibis, January, 1897, 8 - 51. https: // doi. org / 10.1111 / j. 1474 - 919 X. 1897. tb 01261. x","Boulenger, G. A. (1891) Description of a new lizard of the genus Ctenoblepharis from Chili. Proceedings of the Zoological Society of London, 1891, 3. https: // doi. org / 10.1111 / j. 1469 - 7998.1891. tb 06802. x","Moreno, R., Moreno, J., Torres-Perez, F., Ortiz, J. C. & Breskovic, A. (2001) Herpetological catalogue of Museo del Mar of Arturo Prat University, Iquique, Chile. Gayana, 65, 149 - 153. https: // doi. org / 10.4067 / S 0717 - 65382001000200006","Lobo, F. & Etheridge, R. (2013) The type locality of Phymaturus palluma (Molina, 1782) (Reptilia: Iguania: Liolaemidae) and the status of Phymaturus adrianae Pereyra, 1992 and other unnamed populations. South American Journal of Herpetology, 8, 89 - 101. https: // doi. org / 10.2994 / SAJH-D- 12 - 00033.1","Troncoso-Palacios, J. & Garin, C. F. (2013) On the identity of Liolaemus nigromaculatus Wiegmann, 1834 (Iguania, Liolaemidae) and correction of its type locality. ZooKeys, 294, 37 - 56. https: // doi. org / 10.3897 / zookeys. 294.4399","Troncoso-Palacios, J. (2018) Propuesta de restriccion de la localidad tipo de Microlophus tarapacensis (Donoso-Barros, 1966) (Squamata: Tropiduridae) en base a evidencias historicas. Cuadernos de Herpetologia, 32, 123 - 127. https: // doi. org / 10.31017 / CdH. 2018. (2018 - 008)","Veloso, A., Sallaberry, M., Navarro, J., Iturra, P., Valencia, J., Penna, M. & Diaz, N. (1982) Contribucion sistematica al conocimiento de la herpetofauna del extremo norte de Chile. In: El Hombre y Los Ecosistemas de Montana. Oficina Regional de Ciencias y Tecnologia de la UNESCO para America Latina y el Caribe, Montevideo, pp. 135 - 268.","Pincheira-Donoso, D. & Nunez, H. (2005) Las especies chilenas del genero Liolaemus (Iguanidae Tropiduridae, Liolaeminae). Taxonomia, sistematica y evolucion. Publicacion Ocasional del Museo Nacional de Historia Natural, Chile, 59, 7 - 486.","Langstroth, R. P. (2011) On the species identities of a complex Liolaemus fauna from the Altiplano and Atacama Desert: insights on Liolaemus stolzmanni, L. reichei, L. jamesi pachecoi, and L. poconchilensis (Squamata: Liolaemidae). Zootaxa, 2809 (1), 20 - 32. https: // doi. org / 10.11646 / zootaxa. 2809.1.2","Schulte II, J. A. & Moreno-Roark, F. (2010) Live birth among Iguanian lizards predates Pliocene-Pleistocene glaciations. Biologyletters, 6, 216 - 218. https: // doi. org / 10.1098 / rsbl. 2009.0707","Aguilar-Puntriano, C., Avila, L. J., De la Riva, I., Johnson, L., Morando, M., Troncoso-Palacios, J., Wood Jr., P. L. & Sites Jr., J. W. (2018) The shadow of the past: Convergence of young and old South American desert lizards as measured by head shape traits. Ecology and Evolution, 8, 11399 - 11409. https: // doi. org / 10.1002 / ece 3.4548"]}

Liolaemus calliston sp. nov. Holotype. LJAMM-CNP 10994 (Figs. 10, 11), an adult male from 4.4 km SE Aguada Pichana Main Camp Total Oil Company, ~ 30 km S junction Provincial Road 7, A��elo Department, Neuqu��n province, Argentina (38��25���43.7���S, 69��09���31.3���W, 431 m), collected by C.H.F. Perez, J. Nittmann, F. Farinaccio and D.R. Perez, 28 October 2008. Paratypes. LJAMM-CNP 10995���10996, MLP.S 2624 (females) from same locality as holotype, collected by C.H.F. Perez, J. Nittmann, F. Farinaccio and D.R. Perez, 28 October 2008. LJAMM-CNP 12420���12422 (females) from same locality as holotype, collected by L.J. Avila and C.H.F. Perez, 4 December 2009. LJAMM-CNP 12417��� 12419 (males) from same locality as holotype, collected by L.J. Avila and C.H.F. Perez, 4 December 2009. Diagnosis. Liolaemus calliston sp. nov. is a robust but small sized member of the clade of Liolaemus referred to as the boulengeri group by Etheridge (1995) and is a member of the Liolaemus donosobarrosi group (Avila et al. 2006, 2007, Olave et. al 2014, 2016), that includes L. cuyanus, L. donosobarrosi, L. mapuche, L. josei, L. puelche and several other potential species still undescribed. Liolaemus tirantii sp. nov. can be distinguished from all other species of its group of Liolaemus by a combination of genetic and morphological characteristics. Liolaemus calliston sp. nov. can be differentiated from L. cuyanus for having smaller size (61.0 mm vs 92.5 mm maximum SVL), have higher number of midbody scales (X = 76.8 vs X = 66.7; min.���max. 70���84 vs 60���76, Blum 2012) with overlap, higher number of dorsal scales (X = 95.3 vs 63.6; min.���max. 86���100 vs 57���74, Blum 2012) with no overlap, only four scales in contact with mental vs six in L. cuyanus, lack of prescapular marks and ventral melanism, and a very different general coloration and coloration patterns. Liolaemus calliston sp. nov. is smaller than Liolaemus mapuche (61.0 mm vs 82.7 mm maximum SVL), have higher number of dorsal scales (X = 95.3 vs 76.2; min.���max. 86���100 vs 70���86) with almost no overlap, only four scales in contact with mental vs six in L. cuyanus (more common state, Abdala 2002), and lacks of the scapular marks, ventral melanism and lacks of the very different dorsal coloration characterized by a light blue head and scattered blue scales on a light green-blue background, (Abdala, 2002). Liolaemus calliston sp. nov. is smaller than L. puelche (61.0 mm vs 89 mm maximum SVL), have higher number of midbody scales (X = 76.8 vs 70.7; min.���max. 70���84 vs 67���76) with slight overlap and higher number of dorsal scales (X = 95.3 vs 76.7; min.���max. 86���100 vs 74���80) with no overlap. Liolaemus calliston sp. nov. is smaller than L. josei (61.0 mm vs 73.1 mm maximum SVL), have higher number of midbody scales (X = 76.8 vs 69.1; min.���max. 70���84 vs 62���76), have higher number of dorsal scales (X = 95.3 vs X = 73.1; min.���max. 86���100 vs 67���81) with no overlap, lacks of the well-marked gular melanism of L. josei males as well as its more marked dorsal color pattern. Liolaemus calliston sp. nov. can be only be differentiate from L. donosobarrosi in its particular dorsal coloration of colorful transversal bands in live animals, lack of transversal dark bands on the sides of the head, an oval-shaped tympanic opening vs a more quadrangular in L. donosobarrosi, and supralabials and lorilabials scales quadrangular shaped in L. calliston sp. nov. than in L. donosobarrosi. Dorsal coloration pattern in L. tirantii lacks of the banded pattern of L. calliston and the colorful general coloration; series of transversal half-moon dots almost disappear in live animal in L. calliston sp. nov. and are fused each other laterally along the body, given it a ¨wavy¨ appearance. L. calliston sp. nov. have a higher average number of dorsal scales than in L. tirantii sp. nov. (X = 95.3 vs 87.2). Infralabials, supralabial and lorilabials scales more quadrangular shaped in L. calliston sp. nov. than in L. tirantii sp. nov. Description of holotype. Adult male (Fig. 10, 11) 60.4 mm snout-vent length (SVL); tail length 103.4 mm. Axilla-groin distance 27.2 mm. Head length 13.1 mm (from anterior border of tympanum to tip of snout), 11.5 mm wide (at anterior border of tympanum), 7.5 mm high (at anterior border of tympanum). Snout length 4.9 mm (orbittip of snout distance). Interorbital distance 6.0 mm. Eye-nostril distance 3.9 mm. Orbit-auditory meatus distance 4.7 mm. Forelimb length 17.4 mm. Tibial length 12.6 mm. Foot length 18.5 mm (ankle to insertion of claw on fourth toe). Dorsal head scales something bulged, smooth, 16 between occiput at level of anterior border of tympanum to rostral, pitted with numerous scale organs in the anterior region, and reducing to a single organ, or lack, in the posterior half of the head. Rostral scale wider (2.5 mm) than high (0.8 mm). Two postrostrals, together with anterior lorilabial, separate nasal scales from rostral. Nasal scales longer than wide, irregularly heptagonal; nostril one-half length of nasal, posterior in position. Scales surrounding nasals 7���7. Two internasals. Frontonasals four, irregular in size and position. A row with three scales between frontonasals and prefrontals. Prefrontals 5, a small pentagonal scales in the center (1.2 mm), and two pairs scales: a pair dorso-lateral larger scales, irregularly hexagonal (1.6 mm), and a pair lateral medium-sized scales (1.4 mm), aproximatly pentagonal. Two frontal scale. Frontoparietals in two rows, two anterior and posterior scales. Interparietal subpentagonal, surrounded by six scales; four smaller and irregular in front and sides, two larger in back. Parietal eye evident. Parietals some bulged, irregularly shaped, similar in size to interparietal (1.8 vs. 1.6 mm). Circumorbitals: 13���12. Transversally expanded supraoculars 6���6. Smaller lateral supraoculars: 14���19. Anterior canthal wider than long, separate from nasal by one postnasals. Posterior canthal longer than wide. Loreal scales, bulged (6���7). Lorilabials longer than wide (7���8), smaller than labials. Superciliaries 6���7, flattened and elongated, anterior four broadly overlapping dorsally. Orbit with 14���15 upper and thirteen lower ciliaries on each side. Orbit diameter 3.3 x 2.1 mm. Preocular small, unfragmented, longer than wide. Subocular scale elongated, nine times longer than wide (4.2 x 0.4 mm). A wellmarked longitudinal ridge along upper margin of preocular and subocular scales. Postocular small, slightly bulged, quarter superimposed to subocular, with a marked longitudinal ridge. Palpebral scales small granular and bulged. Supralabials six on each side, convex. Temporals smooth, convex, juxtaposed with one scale organ in the tip. Anterior auriculars smaller than adjacent posterior temporals, something projecting outward (3���2). Posterior auriculars small and granular. External auditory meatus conspicuous, higher (1.9 mm) than wide (1.1 mm). Lateral scales of neck granular with inflated skin. Mental scale wider (2.3 mm) than high (1.4 mm), in contact with five scales. Mental followed posteriorly by two postmentals, and two rows of four chinshields on right side (left side with scar damage). Five infralabials on each side, stretched, slightly smaller than supralabials. Gular scales smooth, flat, imbricate, with rounded posterior margins, with melanophores. Scales of throat between chinshields slightly juxtaposed, becoming slightly imbricate toward auditory meatus. Thirty-three gulars between tympanum openings. Infralabials separated from chinshields by one to three rows of scales. Antehumeral and longitudinal neck well developed; postauricular very distinctive; gular incomplete; rictal, dorsolateral, oblique unconspicuous. Scales of dorsal neck region rhombals, imbricate, smooth. Ninety-nine dorsal scales between occiput and anterior surface of thighs. Dorsal body scales rhomboidal, imbricate, smooth to very slightly keeled towards the posterior half of body. Dorsal scales grade laterally into slightly smaller, smooth scales at midbody. Scales immediately anterior and posterior to forelimb and hindlimb insertion small, smooth, granular, and nonoverlapping. Body lateral scales grading smaller to larger at midbody. Ventral body scales rhombals, smooth, flat, imbricate, larger than dorsal scales. Eighty-four scales around midbody; scales between mental and precloacal pores 94. Scales of cloacal region about equal in size to ventral body scales. Six conspicuous precloacal pores. Anterior suprabrachials rhomboidal, imbricate, smooth, slightly larger in size to dorsal body scales. Postabrachials smaller, smooth, becoming granular near axilla. Supra-antebrachials similar to suprabrachial. Infraantebrachials rhombals, smooth. Supracarpals imbricated, rhomboidal, smooth. Infracarpals strongly imbricate, rhomboidal, keeled, 1���3 mucronate. Subdigital lamellae with 2���3 keels, each terminating in a short mucron, numbering: I: 9, II: 14, III: 19, IV: 20, V: 12. Claws robust, curved and sharp, opaque brown. Suprafemorals as large as dorsal body scales, rhomboidal, imbricated, smooth, some very slightly keeled. Postfemorals small, granular shape. Supratibial rhomboidal, imbricated, smooth. Infrafemorals scale small, granular and smooth with enlarged and bulged scales patch, with thirty scales (femoral patch), mucronate. Supratarsal rhombals, imbricated and smooth. Infratarsal small, rhomboidal, imbricate, smooth. Subdigital scales keeled, each terminating in 1���3 short mucron, numbering: I: 11, II: 16, III: 20, IV: 25, V: 15. Claws robust, curved and sharp, opaque brown. Tail complete, non-regenerated. Dorsal and lateral caudals scales, quadrangular and moderatly keeled. Ventral subtriangular and smooth, toward posterior half moderatly keeled. Color in life. Head dorsal coloration darker than body coloration, homogenously Prout��s Brown (47) with darker areas Burnt Umber (48); all this pattern, extend to the sides of the head. Dorsal background coloration between occiput to rump with five Dark Salmon Color (59) transverse bands separated by similar wide bands of Light Yellow Orcher (13). Each Dark Salmon Color (59) band posteriorly bordered by slightly evident half-moon Tawny (60) dots barely encircling Cream White (52) scales. Two more Dark Salmon Color (59) bands continuous along the first quarter of the tail, then bands became smaller and encircle the tail as incomplete and broken rings, and almost disappear in the last third to the tip. From the posterior area of the tympanum to the cloacal region of the tail, a lateral band of Dark Salmon Color (59), sometimes dotted of Cream White (52). From the lateral sides of the neck to the first third of the tail, iridescent Caribbean Blue (168) dots are homogeneously distributed, reaching some middorsal areas of the trunk and becoming faded and less evident in the last two thirds of the tail. Limbs with incomplete rings of Burnt Umber (48) on a Dark Salmon Color (59) background. Ventral areas Cream White (52) with a reticulated dark on the mandibular, gular and ventral neck regions. Some Sulphur Yellow (80) speckles on the chest and upper belly becoming less evident on the lower belly; continuous Spectrum Yellow (79) coloration along the infrafemoral region. Color in preservative. In preservative the general brightly color pattern disappear or fade, all dorsal bands of Light Yellow Orcher (13) and Sulphur/Spectrum Yellow (80/79) coloration disappear, Sulphur Yellow (80) speckles fade but are evident. Variation. Table 4 summarizes the variation of males and females of the type series for some scale and morphometric characters. Precloacal pores are present only in males. In males, dorsal series of roughly half-moon shaped dots became more evident in some specimens. Almost in all individuals observed, dots fused with the nearest on the dorsolateral area given and appearance of three spots, with a posterior Pale Sulphur Yellow (92) border. A male lacks of symmetrically arranged left-right half-moon dots series, with left series side missing. Patterns of transversally arranged dots are not regularly distributed. Density pattern of Caribbean Blue (168) dots varies between individuals. Ventral dark reticulation varies in density and arrangement but are limited to gular and neck region. Dark dots are dispersed on chest and belly, with color variation between Maroon (39) to Jet Black (300). Sulphur Yellow (80)/Spectrum Yellow (79) coloration is limited to gular/upper chest region and femoral region, in some individuals extended to the foot. Females (Fig. 12) usually have wider bands of Light Yellow Orcher on the dorsal area; ventral surface of one female was completely and homogeneously cover with Salmon Color (83), except in mandibular and gular region. Iridescent Caribbean Blue (168) dots are absent in females. Etymology. The specific epithet is an adjective of Greek origin (calliston = very beautiful), in reference to the distinct and brightly colorful pattern of dorsal and lateral body of this new species. Distribution. This species is restricted to the type locality. Natural history. No studies on the basic biology or ecology of this new species are available; we have only anecdotal data from the original collecting trip. The type locality of Liolaemus calliston sp. nov. is characterized by sedimentary rocks separated by barren soil and sparse clumps of low bushes that have been heavily impacted by heavy road machinery. Despite our efforts, we were unable to observe any animal in less disturbed environments that surround the type locality. Lizards were collected late in the afternoon, and only a few were basking on rocks. Lizards were wary and dropped into burrows below the stones when the intruder is still at a large distance. If pressed, they ran from rock to rock across patches of bare soil or between rocks accumulated on the edges of the disturbed area, sometimes seeking cover between roots or basal root clumps. They are fast and heavy runners and tend to return to their burrow or seek shelter under rocks. Liolaemus calliston sp. nov. is oviparous; adult females captured in December had large eggs that extended their abdominal walls, and had conspicuously slow escape runs. They appear to exist at a very low density but ongoing studies in the area will be crucial to elucidate some biological and ecological characteristics of this species. Vegetation is typical of Monte Sur Patag��nico (Roig 1998); predominant vegetation formation is a low to medium shrubs steppe characterized by bushes between 0.50 to 2.00 m. Soils are subject to wind erosion which form dwarf dunes interspersed between rock outcrops separated by bare sand dunes (Fig. 13). Common plants include: Grindelia chiloensis, Senecio filaginoides, Acantholippia seriphioides, Ephedra ochreata, Chuquiraga erinacea, Larrea nitida, L. divaricata, L. cuneifolia, Atriplex lampa, Prosopis alpataco, Fabiana peckii and Adesmia guttulifera, Cercidium praecox, Sporobolus rigens, Hyalis arg��ntea, Monttea aphylla, Neosparton ephedroides and Senna aphylla. Lizard fauna in this general region include up to six congeners: Liolaemus darwinii, L. grosseorum, L. gracilis, L. cuyumhue, L. goetschi, L. austromendocinus, the three phyllodactylid geckos Homonota darwinii, H. horrida, H. underwoodi, two leiosaurids, Leiosaurus belli and Pristidactylus fasciatus, and the teiid Aurivela longicauda. Snakes observed at the type locality include Philodryas trilineata, Bothrops diporus, Xenodon semicinctus, Siagonodon borrichianus; at least the first two of these are likely predators of L. calliston sp. nov., Published as part of Avila, Luciano Javier, Perez, Cristian Hern��n Fulvio, Minoli, Ignacio, Medina, Cintia Debora, Sites, Jack W. & Morando, Mariana, 2017, New species of Liolaemus (Reptilia, Squamata, Liolaemini) of the Liolaemus donosobarrosi clade from northwestern Patagonia, Neuqu��n province, Argentina, pp. 535-563 in Zootaxa 4362 (4) on pages 551-555, DOI: 10.11646/zootaxa.4362.4.4, http://zenodo.org/record/1095342, {"references":["Avila, L. J., Morando, M. & Sites, J. W. Jr. (2006) Congeneric phylogeography: hypothesizing species limits and evolutionary processes in Patagonian lizards of the Liolaemus boulengeri group (Squamata: Liolaemini). Biological Journal of the Linnean Society, 89, 241 - 275. https: // doi. org / 10.1111 / j. 1095 - 8312.2006.00666. x","Olave, M., Martinez, L. E., Avila, L. J., Sites, J. W. Jr. & Morando, M. (2014) Multilocus phylogeny of the widely distributed South American lizard clade Eulaemus (Liolaemini, Liolaemus). Zoologica Scripta, 43, 323 - 337. https: // doi. org / 10.1111 / zsc. 12053","Blum, R. (2012) Morphometric analyses and intraspecific comparison of two geographically separated populations of Liolaemus cuyanus (Iguania: Liolaemidae) from Northwestern Argentina. Unpublished Master Thesis Georg-August- Universitat Gottingen. Germany, 101 pp.","Abdala, C. S. (2002) Nuevo Liolaemus (Iguania: Liolaemidae) perteneciente al grupo boulengeri de la provincia de Neuquen, Argentina. Cuadernos de Herpetologia, 16, 3 - 13.","Roig, A. F. (1998) La vegetacion de la Patagonia. In: Correa, M. N. (Ed.), Flora Patagonica. INTA, Buenos Aires, Argentina. pp. 48 - 174."]}

Phymaturus rahuensis sp. nov. (Fig. 4, 5, 6) Phymaturus sp. 16 Morando, M., Avila, L.J., Perez, C.H.F., Hawkins, M. and Sites, Jr. J.W., 2013, Molecular Phylogenetics and Evolution, 66, 698. Type material. Holotype: LJAMM-CNP 16126, adult male collected on rocky hills of Provincial Road 46, 25 km E Rahue, 6 km junction Provincial Road 24, Bajada de Rahue, La Jardinera stream (39 �� 23 ��� 15.5 ���S, 70 �� 43 ��� 57.1 ���W, 1127 m, datum = WGS 84), Cat��n Lil department, Neuqu��n province, Argentina, A. Gonz��lez Mar��n, L.J. Avila and C.H.F. P��rez, collectors. Paratypes: LJAMM-CNP 16117 -16120, 16123, 16127 (adult males) and LJAMM-CNP 16121 -16122, 16124- 16125, 16128- 16129 (adult female), same data as holotype. LJAMM-CNP 5378 (adult male) and 5379 (adult female) collected on rocky cliffs on Provincial Road 46, 25 km E Rahue, 6 km junction Provincial Road 24, Bajada de Rahue, La Jardinera stream (39 �� 23 ��� 15.5 ���S, 70 �� 43 ��� 57.1 ���W, 1127 m, datum = WGS 84), Cat��n Lil department, Neuqu��n province, Argentina, L.J. Avila, C.H.F. P��rez, K. Dittmar, M. Morando & J.W. Sites, Jr. collectors. Diagnosis. Phymaturus rahuensis sp. nov. is a member of the Phymaturus patagonicus group because it exhibits synapomophies found on this species��� group, bellies light-orange or pink, a set of enlarged scales projected over the auditory meatus and external margins of postmental scales dark pigmented (sensu Lobo et al. 2012 b) and (sensu Etheridge 1995) flat imbricate superciliaries, non-rugose dorsal scales on tail, and subocular scale usually not fragmented. Within the Phymaturus patagonicus group, P. rahuensis sp. nov., differs from P. cacivioi, P. castillensis, P. ceii, P. desuetus, P. excelsus, P. felixi, P. indistictus, P. manuelae, P. spectabilis, P. spurcus, P. tenebrosus, P. videlai, in the presence of a dorsal pattern formed by dispersed white spots. Scales around midbody are non-overlapping among P. rahuensis (216���261), P. patagonicus (162���192) and P. yachanana (160���194). Number of ventral scales in P. rahuensis (167���193) is higher and slightly overlapping with P. patagonicus (142��� 171) and P. yachanana (135���168). Average number of scales around midbody in P. rahuensis (X��= 236.7) is higher than in P. somuncurensis (X��= 218.29), P. sinervoi (X��= 227.9), P. calcogaster (X��= 221.20), P. etheridgei (X��= 221.55) and with P. camilae (range 205-228, Scolaro et al. 2013). Phymaturus rahuensis sp. nov. has a marked sexual dichromatism not present in P. nevadoi (according to original description) and P. sitesi. Phymaturus rahuensis sp. nov. has a dorsal pattern coloration (well marked in females) formed by a paravertebral band with two dorsolateral series of quadrangular to irregular marks fused to the paravertebral band but usually separated each other by dark lines, a pattern not observed in any other species of the clade. Phymaturus rahuensis sp. nov. has a dorsal coloration with small rounded white spots scattered on its body (4���12 scales size each) similar to P. nevadoi (4���9 scales) and P. delheyi (1���10 scales), larger than in P. sitesi (1���2 scales), but smaller than in P. payuniae (4���40 scales, sometimes fused forming irregular marks) and P. zapalensis (5���14 scales). Phymaturus rahuensis sp. nov. has a tail with vanishing rings as P. zapalensis but not spotted as in P. delheyi, P. nevadoi, P. payuniae, and P. sitesi. Average scales around midbody number is higher in Phymaturus rahuensis sp. nov. (X��= 236.7, SD = 12.8) than all other species of the group: P. nevadoi (X��= 208.3, SD = 8.7), P. payuniae (X��= 219.0, SD = 11.7), P. sitesi (X��= 221.5, SD = 7.9), P. delheyi (X��= 216.7, SD = 15.7) and P. zapalensis (X��= 221.2, SD = 13.8). Average ventral scale number is lower in P. rahuensis (X��= 179.6, SD = 7.2) than in P. payuniae (X��= 183.3, SD = 10.4), P. delheyi (X��= 185.5, SD = 9.2) and P. zapalensis (X��= 186.3, SD = 13.0) but higher than P. nevadoi (X��= 164.0, SD = 8.4) and P. sitesi (X��= 176.2, SD = 9.5). Average supralabials scales number is lower in P. rahuensis (X = 8.0, SD = 0.5) than in the other species of the group: P. nevadoi (X��= 9.2, SD = 0.5), P. payuniae (X��= 11.1, SD = 1.1), P. sitesi (X��= 9.2, SD = 1.2), P. delheyi (X��= 9.0, SD = 0.8) and P. zapalensis (X��= 8.8, SD = 0.8). Phymaturus rahuensis sp. nov. has lower finger lamellae number (X��= 19.8, SD = 1.1) than the P. nevadoi (X��= 22.6, SD = 1.4), P. payuniae (X��= 22.1, SD = 1.1), P. sitesi (X��= 21.8, SD = 0.9), P. delheyi (X��= 22.4, SD = 1.1) and P. zapalensis (X��= 21.7, SD = 1.8) and lower toe lamellae number (X��= 25.8, SD = 1.1) vs P. nevadoi (X��= 28.8 SD = 1.2), P. payuniae (X��= 27.8, SD = 1.9), P. sitesi (X��= 28.8, SD = 1.2), P. delheyi (X��= 29.6, SD = 2.0) and P. zapalensis (X = 28.4, SD = 1.8). The new species can be distinguished from P. nevadoi, P. payuniae, P. zapalensis, P. delheyi and P. sitesi by differences in mitochondrial and nuclear genes (exclusive haplotypes with cytochrome b and 12 S mitochondrial genes and with PNN, Phy 38 and Phy 41 nuclear markers (Morando et al. 2013). Based on cyt-b data (Morando et al., 2013), uncorrected genetic distances between previously morphologically described species of Phymaturus are low (e.g. for a species pair within the patagonicus group: P. nevadoi vs. P. payuniae = 0.81 %, in Avila et al. 2014); some species pairs even have smaller values, but they may be conspecific (Morando et al., 2013). The smallest uncorrected genetic distances between P. rahuensis sp. nov. and other taxa of the payuniae clade are: P. rahuensis vs P. zapalensis = 1.7 %, P. rahuensis vs P. sp. 17 = 1.44 %, while with the rest of the species of this clade, genetic distances are> 3.6 %. Description of the holotype. Adult male 88.1 mm (SVL). Axilla-groin distance 45.5 mm. Tail length 101.9 mm. Head length 15.5 mm; head width 16.8 mm; head depth 9.4 mm; snout length 5.6 mm (from anterior border of the orbit to tip of snout), horizontal diameter of the orbit 6.6 mm. Arm length 26.5 mm; tibial length 17.9 mm; foot length 24.4 mm. Upper head scales smooth, convex, bulged, pitted with scale organs in postrostral, internasals, frontonasals, frontals, and prefrontals. Rostral flat, two times wide as long (2.7 x 1.1 mm). Two postrostrals on the left side, the outermost smaller than the inner, in the right side abnormally small and numerous, with conspicuous scale organs (1-6); rostral and nasal not in contact, separated by the anterior lorilabial scale and a small prenasal. Nasal scales almost rounded (1.3 x 1.5 mm). Nostril rounded, occupying almost all the nasal scale. Nasal scales in contact with 9 - 8 scales. Internasal, frontonasals, medium in size, irregular in shape; prefrontals and frontals, medium in size, hexagonal; supraorbital semicircles complete, 13 ��� 11 enlarged supraoculars; frontoparietals, parietals, and circumorbitals irregular in size and shape from each other. Interparietal hexagonal, only distinguishable by a large and conspicuous white cream ���eye��� in the middle, occupying most of the scale. Eighteen dorsal head scales between rostral and nuchals. Two scales between nasal and first canthal. First canthal small, higher than wide, convex with two scale organs. Posterior canthal larger, longer than wide. Posterior canthal slightly overlaps first supercilliary. Supercilliaries 7 ��� 5 (left-right) irregular flattened and elongated, on left sides overlapped the first five, and the right side the three first scales. Loreal region flat, three irregular scales on both sides. Upper ciliary scales in two rows, those of inner rows flat and quadrangular, those of outer row granular and compressed. Lower and upper ciliaries similar in size and shape. Palpebral scales small, irregular, slightly granular. One preocular, small, square; one elongate subocular (4.5 x 0.6 mm), unfragmented, one small postocular; a well marked longitudinal ridge along upper margin of subocular and preocular scale, and not marked in postocular. Two rows of lorilabials becoming only one in half subocular. Lorilabials convex, 12 / 6���10 / 6, approximately rectangular, slightly narrow than supralabials, pitted with 1���5 scale organs. Supralabials 7, flat. Temporal scales conical, smooth, swollen, juxtaposed, with a scale organ at the inferior side. Auditory meatus higher than wide (4.1 x 2.0 mm) with 3 outwardly projecting scales along anterior border; posterior border surrounded by granular scales. Mental pentagonal, wider than long (2.2 x 1.4); in contact with anterior infralabials, anterior sublabials and postmentals. Infralabials 6. Chinshields 5 - 4, irregular, slightly quadrangular, separated from infralabials by series of 1���2 ��� 3 irregular scales, first equal in size but becoming smaller to back. One- 4 scale organ present in some supralabials and infralabials. Gular scales round, flat, and juxtaposed. Sixty-six between auditory meatus. Antehumeral, gular and lateral fold well developed, antegular fold notorious, and postauricular folds few developed. Ante-humeral pocket well developed. Fifty-seven scales between auditory meatus and scapula. In ventral view, gular fold incomplete with its anterior margins delimited with enlarged scales on their borders. Dorsal body scales rounded, smooth, juxtaposed. Forty-four dorsal scales along midline of the trunk in a distance equivalent to head length. Scales around midbody 216. Scales on flanks conical. Ventral scales larger that dorsals, rhombals and moderately imbricated. Ventral scales between mental and precloacal pores 178. Scales of the cloacal apron equal in size than body scales, flat, rounded, moderately imbricate. Precloacal pores eight. Supra-brachial and antebrachial scales smooth, rhombals and imbricated, larger than dorsal body scales. Supracarpals laminar, rhombals, smooth. Supradigital lamellae convex, imbricate. Infra-brachial and ante-brachial scales small, granular. Subdigital finger lamellae with 3���4 keels (more conspicuous in proximal lamellae), 3���4 mucronate. Subdigital finger lamellae numbering I: 11; II: 15; III: 19; IV: 21; V: 15. Claws moderately long. Infracarpals with round margins and 1-3 obtuse keels, 1-3 mucronate. Supra-femorals smooth, imbricate, rhomboidal to round. Infra-femorals slightly larger and imbricate, smooth, rhomboidal. Supra-tarsals rhombals, smooth and imbricate. Post-femoral scales smaller and granular. Supra-tibials imbricate, conical, and mucronate. Infra-tibials larger than infra-femorals, rhombals, smooth, imbricate. Infra-tarsals with 1���3 obtuse keels, mucronate. Sub-digital toe lamellae numbering I: 12; II: 16; III: 22; IV: 27; V: 18. Caudal scales arranged in spinose annuli, scales larger than body and limbs scales, slightly keeled, imbricated, out-projecting. Holotype coloration. Dorsal background coloration brown/dark drab, more intense on trunk becoming cinnamon-drab on head, limbs, and tail. Dorsal coloration pattern on body is a wide dorsal band with a clear and continuous vertebral band and two dorsolateral series of irregular rounded to roughly quadrangular spot wide joined to the vertebral band and separated each other by a narrow dark transversal line that disappear in some areas becoming like dark irregular small spots along paravertebral line sides. On each side of the trunk, between neck to anterior insertion of the hind limbs two conspicuous dusky brown/black lateral bands. Trunk, forelimbs and head speckled with cream white (4���12 scales) more evident on lateral and dorsolateral bands becoming less conspicuous on head and forelimbs and almost not evident on vertebral band and neck. Limbs with cream white reticulate. First two thirds of the tail with cinnamon-drab vanishing rings becoming non-conspicuous in the distal portion. Throat, gular region, malar region, neck lateral region and upper chest pale neutral grey with dusky brown reticulate. Lower chest and upper belly yellow ocher, more intense in cloacal apron and adjacent femoral region. Ventrolateral region drab with dusky brown reticulated. Ventral region of anterior forelimbs pale neutral grey and posterior forelimbs cream white. Ventral region of tail pale neutral grey. Color in preservative. After preservation, head, dorsum, body flanks and tail maintain original coloration but darken and lose some tones. Chest, belly and femoral regions lost the distinctive ventral yellow-ocher. Variation. Based on eight adult males (Table 4, Fig. 4, 6): SVL 83.6���88.11 mm. Axilla-groin distance 38.9��� 45.5 mm. Foot length 21.8���24.4 mm. Tibial length 15.5���18.5 mm. Arm length 24.4���26.8 mm. Head length 15.1��� 15.9 mm. Head width 15.0��� 16.8 mm. Head height 9.0��� 9.5 mm. Midbody scales 216���261. Dorsal scales in head length 44���49. Ventral scales 167���186. Supralabials 7���9. Infralabials 6���9. Scales around nasal 6���8. Third finger lamellae 18���22. Fourth toe lamellae 24���28. Precloacal pores 8���11. In seven adult females (Table 4, Fig. 5, 6): SVL 72.2���95.1 mm. Axilla-groin distance 37.6 ���51.0 mm. Foot length 21.6 ���23.0 mm. Tibial length 15.2���17.6 mm. Arm length 23.3 ���25.0 mm. Head length 13.0��� 15.9 mm. Head width 13.8���16.6 mm. Head height 7.4���9.4 mm. Midbody scales 221���244. Dorsal scales in head length 40���46. Ventral scales 176���193. Supralabials 8���9. Infralabials 7���8. Scales around nasal 6���8. Third finger lamellae 19���21. Fourth toe lamellae 25���27. Coloration in females differs from the males but follow the same coloration pattern but it is more noticeable, with larger dorsolateral spots (10��� 12), and well distinct. Dorsal band is very distinctive from the lateral band. In female LJAMM-CNP 16128, dorsal background of body, head, forelimbs and tail is beige. In same females, dorsolateral spots are smaller and irregular, from rounded to like a four point star, then transversal separating lines expand its size becoming rounded along the vertebral band giving the appearance of a series of dark ocelli. Lateral region of body, from the neck region to the groin, dusky brown/black, speckled with cream color smudges (5���30 scales), postauricular region only reticulated. Dorsal pattern of limbs with white cream reticulate and a few scattered dusky brown spots (1-2 scales). Tail with white cream reticulate, transforming in rings toward the posterior half region. Ventral surface pale neutral grey. Throat, gular region, malar region and neck lateral region with dusky brown reticulate. Lower belly, cloacal apron, femoral, tibial and tarsal region orange yellow. Ventrolateral region pale neutral grey with dusky brown reticulated. Ventral tail beige. The other female of the type series has the same color pattern. Eight males have the same basic color pattern with greater or lesser degree of development of the dorsal cream white spots or extension of the dusky brown area. In the specimen with completely dusky brown dorsum, the spots in the dorsolateral region are denser. In some specimens in the dorsal band, the spots are absent and in other, the dusky brown spots form transverse stripes. In males the yellow ocher color in the lower belly, cloacal and femoral adjacent region is similar in all the specimens but LJAMM 16127 that also has an orange yellow color in the first third of the tail. Males (n= 8) Females (n= 7) Mean SD Range Mean SD Range SVL 85.6 1.7 83.6���88.11 86.8 8.0 72.2���95.1 AGD 42.3 2.3 38.9���45.5 44.9 4.5 37.6 ���51.0 HL 15.3 0.2 15.1���15.9 14.8 1.0 13.0��� 15.9 HW 16.0 0.5 15.0��� 16.8 15.3 0.9 13.8���16.6 HH 9.3 0.2 9.0��� 9.5 8.7 0.7 7.4���9.4 FL 23.2 0.9 21.8���24.4 22.4 0.4 21.6 ���23.0 TL 17.2 0.8 15.5���18.5 16.6 0.9 15.2���17.6 AL 25.4 0.9 24.4���26.8 24.1 0.5 23.3 ���25.0 SAMB 240.6 15.5 216���261 232.2 7.8 221���244 DS 46.3 1.8 44���49 42.8 2.6 40���46 VS 177 7.2 167���186 182.5 6.6 176���193 LM 20 1.5 18���22 19.7 0.7 19���21 LP 26 1.3 24���28 25.7 0.9 25���27 SLS 7.8 0.6 7���9 8.1 0.3 8���9 ILS 7.6 0.9 6���9 7.5 0.5 7���8 PC 9.3 0.9 8���11 ��� ��� ��� Etymology. The specific name, ��� rahuensis ���, refers to the region where the species was collected "Bajada del Rahue". The Rahue toponym in Mapudungun means "place where there is gray clay". Geographic distribution. Phymaturus rahuensis sp. nov. was only collected on rocks 1127 m above sea level in Bajada del Rahue at La Jardinera streams, in the mountain range formation known as Cat��n Lil mountains. This location has rocky marine sediments from middle Jurassic; the area was occupied by marine ingression and regression that deposited the sedimentary landscape, named as Los Molles Formation (Riccardi 1993; Garc��a Morabito 2010). All range was affected by Holocenic glaciations but details about its effects on the landscape (and the biota) are still unknown (Rabassa et al. 2011) (Fig. 7). Natural history. Phymaturus rahuensis sp. nov. is a common and easy to observe species. Little information about natural history and biology of this new species is available. Phymaturus rahuensis sp. nov. was usually found surrounded by vegetation characteristic of the Patagonica Phytogeographic Province, Payunia District, dominated by communities of Senna arnottiana and Stillingia patagonica inter alia (Roig 1998), also species as Haploppapus sp., Mulinum spinosum and several species of grasses (Stipa spp. and Festuca spp.) were observed. The holotype and paratypes were found by active search, but usually they were spotted basking on rocky outcrops or hiding in crevices. Liolaemus elongatus and L. kriegi are the only observed lizards that shared the same microhabitat. One species of amphibian, Pleurodema bufoninum, was observed on a little stream. No data about reproduction, diet or other natural history characteristics are available, but as in other related species of Phymaturus, Phymaturus rahuensis sp. nov. probably is viviparous and feeds on plant matter, and possibly some arthropods. Discussion, Published as part of Mar��n, Andrea Gonz��lez, P��rez, Cristian Hern��n Fulvio, Minoli, Ignacio, Morando, Mariana & Avila, Luciano Javier, 2016, A new lizard species of the Phymaturus patagonicus group (Squamata: Liolaemini) from northern Patagonia, Neuqu��n, Argentina, pp. 412-430 in Zootaxa 4121 (4) on pages 420-426, DOI: 10.11646/zootaxa.4121.4.3, http://zenodo.org/record/261343, {"references":["Lobo, F., Abdala, C. S. & Valdecantos, S. (2012 b) Morphological diversity and phylogenetic relationships within a South- American clade of iguanian lizards (Liolaemidae: Phymaturus). Zootaxa, 3315, 1 - 41.","Etheridge, R. (1995) Redescription of Ctenoblepharys adspersa Tschudi, 1845, and the taxonomy of Liolaeminae (Reptilia: Squamata: Tropiduridae). American Museum of Natural History Novitates, 3142, 1 - 34.","Scolaro, J. A., Jara, M. & Pincheira-Donoso, D. (2013) The sexual signals of speciation? A new sexually dimorphic Phymaturus species of the patagonicus clade from Patagonia Argentina. Zootaxa, 3722 (3), 317 - 332. http: // dx. doi. org / 10.11646 / zootaxa. 3722.3.2","Morando, M., Avila, L. J. Perez, C. H. F. Hawkins, M. & Sites, J. W. (2013) Molecular phylogeny of Phymaturus (Squamata, Liolaemini): too many lineages and few names. Molecular Phylogenetics and Evolution, 66, 694 - 714. http: // dx. doi. org / 10.1016 / j. ympev. 2012.10.019","Riccardi, A. C. (1993) Formacion Los Molles. In: Riccardi, A. C. & Damborenea, S. E. (Eds.), Lexico estratigrafico de la Argentina. Volumen 9, Jurasico. Asociacion Geologica Argentina, Serie B, Didactica y Complementaria 21, Buenos Aires, Argentina, pp. 239 - 243.","Garcia Morabito, E. (2010) Tectonica y estructura del retroarco andino entre los 38 ° 15 ' y los 40 ° S. Tesis Doctoral Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires. Avaliable from: http: // digital. bl. fcen. uba. ar / Download / Tesis / Tesis _ 4699 _ GarciaMorabito. pdf (Accessed 8 Jun. 2016)","Rabassa, J., Coronato, A., Ponce, J. F., Schlieder, G. & Martinez, O. (2011) Depositos glacigenicos (Cenozoico Tardio - Cuaternario) y geoformas asociadas. Relatorio XVIII Congreso Geologico Argentino, Neuquen.","Roig, A. F. (1998) La vegetacion de la Patagonia. In: Correa, M. N. (Ed.), Flora Patagonica. INTA, Buenos Aires, Argentina, pp. 48 - 174.","Lobo, F. & Nenda, S. J. (2015) Discovery of two new species of Phymaturus (Iguania: Liolaemidae) from Patagonia, Argentina, and occurrence of melanism in the patagonicus group. Cuadernos de Herpetologia, 29 (1), 5 - 25.","Ocampo, M., Aguilar-Kirigin, A. & Quinteros, S. (2012) A New Species of Liolaemus (Iguania: Liolaemidae) of the alticolor group from La Paz, Bolivia. Herpetologica, 68 (3), 410 - 417. http: // dx. doi. org / 10.1655 / HERPETOLOGICA-D- 12 - 00001.1","Breitman, M. F., Parra, M. & Perez, C. H. F. (2011 a) Two new species of lizards from the Liolaemus lineomaculatus section (Squamata: Iguania: Liolaemidae) from southern Patagonia. Zootaxa, 3120, 1 - 28.","Breitman, M. F., Perez, C. H. F., Parra, M., Morando, M., Sites, J. & Avila, L. J. (2011 b) New species of lizard from the magellanicus clade of the Liolaemus lineomaculatus section (Squamata: Iguania: Liolaemidae) from southern Patagonia. Zootaxa, 3123, 32 - 48.","Aguilar, C., Wood, P. Jr., Cusi, J., Guzman, A., Huari, F., Lundberg, M., Mortensen, E., Ramirez, C., Robles, D., Suarez, J., Ticona, A., Vargas, V., Venegas, P. & Sites, J. (2013) Integrative taxonomy and preliminary assessment of species limits in the Liolaemus walkeri complex (Squamata, Liolaemidae) with descriptions of three new species from Peru. ZooKeys, 364, 47 - 91. http: // dx. doi. org / 10.3897 / zookeys. 364.6109","Claude, J. (2008) Morphometrics with R. Morphometrics with R XVIII. Springer, New York, 318 pp.","Abdi, H., Williams, L. J. & Valentin, D. (2013) Multiple factor analysis: principal component analysis for multitable and multiblock data sets. Wiley Interdisciplinary Reviews: Computational Statistics, 5 (2), 149 - 179. http: // dx. doi. org / 10.1002 / wics. 1246","Bookstein, F. L. (1991) Morphometric tools for landmark data. Geometry and biology. New York: Cambridge University Press.","Adams, D. C., Rohlf, F. J. & Slice, D. E. (2004) Geometric morphometrics: ten years of progress following the ' revolution'. Italian Journal Zoology, 71, 5 - 16. http: // dx. doi. org / 10.1080 / 11250000409356545","Kaliontzopoulou, A. (2011) Geometric morphometrics in herpetology: modern tools for enhancing the study of morphological variation in amphibians and reptiles. Basic y Applied Herpetology, 25, 5 - 32. http: // dx. doi. org / 10.11160 / bah. 11016","Florio, A. M., Ingram, C. M., Rakotondravony, H. A., Louis, E. E. & Raxworthy, C. J. (2012) Detecting cryptic speciation in the widespread and morphologically conservative carpet chameleon (Furcifer lateralis) of Madagascar. Journal of Evolutionary Biology 25 (7), 1399 - 1414. http: // dx. doi. org / 10.1111 / j. 1420 - 9101.2012.02528. x","Kaliontzopoulou, A., Carretero, M. A. & Llorente, G. A. (2012) Morphology of the Podarcis wall lizards (Squamata: Lacertidae) from the Iberian Peninsula and North Africa: patterns of variation in a putative cryptic species complex. Zoological Journal of the Linnean Society, 164, 173 - 193. http: // dx. doi. org / 10.1111 / j. 1096 - 3642.2011.00760. x","Dayrat, B. (2005) Towards integrative taxonomy. Biological Journal of the Linnean Society, 85 (3), 407 - 415. http: // dx. doi. org / 10.1111 / j. 1095 - 8312.2005.00503. x","Padial, J. M. & De La Riva, I. (2007) Integrative taxonomists should use and produce DNA barcodes. Philosophical Transactions of the Royal Society of London, 68, 67 - 68."]}

Liolaemus yatel sp. nov. (Fig. 1���3) 1986, Liolaemus lineomaculatus, Cei (partim); Museo Regionale di Scienze Naturali, Torino. Monografia IV: 527 pp. 2011, Liolaemus lineomaculatus, Breitman, Parra, P��rez, Sites (partim); Zootaxa. 3120: 1���28. 2014, Liolaemus lineomaculatus Breitman, Minoli, Avila, Medina, Sites, Morando (partim); Cuadernos de Herpetolog��a. 28 (2): 83���110. Holotype (Fig. 1): FML 24646 (CIPD 628). Adult female. Monumento Natural Bosques Petrificados, Puerto Deseado department, Santa Cruz province, 47 �� 41��21 ���� S; 68 �� 01��03���� W. D. Procopio Col. 03/03/ 2009. Paratypes All individuals were collected in the Monumento Natural Bosques Petrificados, Puerto Deseado department, Santa Cruz province, Argentina. FML 24647 (CIPD 329). Adult male. 47 �� 41��11 ���� S; 68 �� 00��36 ���� W. D. Procopio and O. Stellatelli cols. 27 / 10 / 2006. FML 24648 (CIPD 629) Adult female. 47 �� 41��15 ���� S; 68 �� 00��38 ���� W. D. Procopio col. 06/03/ 2009. FML 24649 (CIPD 394). Adult male. 47 �� 41��39 ���� S; 68 �� 00��13 ���� W. D. Procopio col. 11 /03/ 2007. FML 24650 (CIPD 396): Adult female. 47 �� 41��42 ���� S; 68 �� 00��33 ���� W. D. Procopio col. 11 /03/ 2007. MLP.R. 5704 (CIPD 395). Adult male. 47 �� 41��48 ���� S; 68 �� 00��22 ���� W. D. Procopio col. 11 /03/ 2007. MLP.R 5705 (CIPD 643): Adult female. 47 �� 41��34 ���� S; 68 �� 00��21 ���� W. D. Procopio col. 11 /03/ 2007. MLP.R 5706 -07 (CIPD 644 - 45): Juveniles. 47 �� 41��40 ���� S; 68 �� 00��11 ���� W. D. Procopio col. 11 /03/ 2007. Diagnosis. Liolaemus yatel sp. nov. belongs to the L. lineomaculatus section (Breitman et al., 2011 b; 2013) and, within this section, it belongs to the L. lineomaculatus group (Etheridge, 1995; Abdala & Lobo, 2006), along with the L. kingii and L. magellanicus groups (Breitman et al., 2013). The most striking differences between Liolaemus yatel sp. nov. and species of the L. kingii group (L. archeforus, L. baguali, L. chacabucoense, L. escarchadosi, L. gallardoi, L. kingii, L. sarmientoi, L. scolaroi, L. somuncurae, L. tari, L. tristis, L. uptoni, and L. zullyi) are the absence of precloacal pores in males, a shorter snout-vent length (max SVL 61.1 mm vs. range between 67 and 112 mm, respectively; an exception is L. scolaroi: max SVL: 61 mm), and clearly contrasting dorsal and ventral coloration patterns. Further, L. yatel sp. nov. and the species of the L. kingii group also differ in scalation patterns and morphometry (Breitman et al., 2013). Among other diagnostic traits Liolaemus yatel sp. nov. differs from L. magellanicus and L. caparensis, that belong to the L. magellanicus group (Breitman et al., 2013), by the absence of precloacal pores in males. Morphology showed differences between the new species and the other taxa of the L. lineomaculatus group (Tables 1, 2). Within this group, the absence of trifid scales distinguish the new taxon from all members of the L. lineomaculatus group and L. magellanicus (trifid scales present in: 58 % of the examined specimens of L. magellanicus (Fig. 4), 67 % of L. lineomaculatus, and 100 % in L. avilae, L. hatcheri, L. kolengh L. morandae, and L. silvanae) (Table 1). Liolaemus yatel sp. nov. also differs from L. hatcheri, L. kolengh and L. silvanae (L. silvanae group) for lacking either keeled nuchal scales or imbricate and subimbricated postfemorals (Table 1). In addition, body dorsal scales are subimbricated, slightly keeled and without mucron in L. yatel sp. nov., in contrast to L. magellanicus (Fig. 4), L. caparensis, and species of the L. lineomaculatus group whose scales are imbricated, strongly keeled and mucronated. Table 1 shows further differences in scalation and color patterns of body spots with other species of that group. Morphological tests showed significant differences between the new species and the other taxa of the L. lineomaculatus group. Means and ranks for meristic and qualitative characters are summarized in Table 1. Univariate tests showed that the number of scales around midbody could be used to tell between L. yatel sp. nov. and all other species but L. silvanae, whereas the number of dorsal scales differed significantly between L. yatel sp. nov. and L. silvanae (Table 2). The DFA indicated that the first two discriminant functions were statistically significant (Table 3). The first discriminant function accounted for 63.70 % of the total variance; this function was significantly correlated with the number of scales around midbody, as well as the number of dorsal and ventral scales (Table 3; Fig. 5). The second discriminant function was significantly correlated with the number of nuchal scales (Table 3; Fig. 5). Based on both discriminant functions, the number of scales around midbody, and the number of dorsal, ventral and nuchal scales contributed significantly to separate the centroids of most species (Table 3; Fig. 5). Our analysis allowed the identification of L. yatel sp. nov. based on meristic traits. Table 4 indicates that the nine species (i.e. L. avilae, L. caparensis, L. hatcheri, L. kolengh, L. lineomaculatus, L. magellanicus, L. morandae, L. silvanae, and L. yatel sp. nov.) were each correctly classified with 95.86 % accuracy. All specimens of L. yatel sp. nov. were correctly classified (Table 4). These results indicate that this species possesses morphological characteristics which distinguish it from the other known species of the L. lineomaculatus group. Description of the holotype. Adult female. Snout vent length (SVL) 60.3 mm. Head 1.14 times longer (11.4 mm) than wide (10.0 mm). Head height 6.9 mm. Interorbital distance 7.9 mm. Eye-auditory meatus distance 4.0 mm. Auditory meatus height 1.6 mm; width 1.2 mm. Trunk length 32.4 mm; width 17.1 mm. Tail length 48.9 mm, and not regenerated. The tail is shorter than the SVL. Width tail base 7.0 mm. Arm length 8.0 mm; forearm length 6.6 mm; and hand 7.6 mm. Thigh length 10.2 mm; leg 10.1 mm; Foot length 12.9 mm; IV toe 9.2 mm. DISCRIMINANT FUNCTION Dorsal surface of head smooth. Rostral scale wider (2.7 mm) than high (1.6 mm), in contact with six scales. Mental scale trapezoidal, wider (3.1 mm) than high (2.8 mm), in contact with four scales. Lateral postrostral scale does not contact the first supralabial scale. There is no contact between nasal and rostral scales. Distal end of frontal scale separated from superciliaries by six scales. Six scales between rostral and frontal scales. Frontal scale is divided into three parts. Two postrostral scales, with one and two scale organs respectively. Interparietal scale shorter than paritetals, in contact with seven scales. Eight smooth, juxtaposed or subjuxtaposed temporals. Subocular (length 4.2 mm) is white, with the posterior end and the upper edge dark, and in contact with four lorilabials. Eye diameter 3.1 mm. The postocular scale is not divided. Five supraocular scales. The supraorbital scales form an incomplete semicircle. Six supralabial scales, the fifth is the largest and is curved upwards at its posterior end, without contacting the subocular scale. Four infralabial scales, the second one contacts three scales. Seven lorilabial scales. One scale between preocular and lorilabials. Seven scales surround the nasal scale, which is separated from the canthal scale by two scales. Four internasal scales. Two postmental scales. Six superciliaries. Twelve upper cilliaries. Hellmich index (dorsal scales in head, from occiput to mental scale) 12. Scales around midbody 60. Seventy eight round, slightly keeled or without keels, without mucron and juxtaposed or subjuxtaposed dorsal scales (from occiput to forelimbs). Thirty-four rows of scales on the dorsum. Thirty-seven granular and smooth neck scales (counted from the posterior margin of the auditory meatus to the shoulder, along the longitudinal fold). Antehumeral scales subtriangular and differentiated. Neck folds (auricular, antehumeral and longitudinal) evident. Scales of longitudinal fold granular, juxtaposed and without keel. Thirty gular scales. Eightyeight larger than dorsals, laminar and imbricated ventral scales (from mental scale to cloaca). Thirteen pigal scales. Without precloacal pores. Sixteen infradigital lamellae on fourth finger and 22 on fourth toe. The scales of sides of the body are laminar and without keel. Anterior edge of the auditory meatus with one projected scale. Auricular scale (located in the antero-superior edge of the auditory meatus) absent. The central and lateral nuchal scales are undifferentiated, granular and without keel. Without trifid scales between nuchal areas and lateral cephalic. Coloration (Fig. 1): The dorsal pattern of the head is a pale brown background with hints of gray and a few dark, irregularly scattered spots. The body back is light brown with 11 pairs of paravertebral, dark brown, crescentshaped spots that show an anterior indentation. Flanks have spots that follow the same pattern, color and shape than that of paravertebral design. The contact between paravertebral and lateral spots forms streaks transverse to the body axis. Paravertebral spots do not contact in the vertebral region. Vertebral line, dorsolateral stripes, antehumeral arch and scapular spots are absent. Some fuzzy spots appear along the lateral body midline. The background color of the body back continues in the dorsal region of limbs and tail. A few ring-shaped dark spots appear on dorsal areas of forelimbs and hindlimbs. In the distal region of the tail, lateral and dorsal spots touch and form pseudo-rings. Ventral regions of the head and abdomen are white, with scattered black spots and scales that become denser towards the center of the ventral region, which also shows a very light orange background color. A few dark scales in the limbs and the gular region. The holotype preserved in 70 % ethanol maintains the color pattern observed in life, but shows a more grayish and less intense coloration. Variation. Based on eight paratypes. Liolaemus small, with SVL in adult males and females from 45.7 to 61.1 mm (x��= 55.2 mm). The tail length is shorter than or equal to the SVL, from 48.9 to 56.4 mm (x��= 53.1 mm, n = 5). Body length 18.3���32.4 mm (x��= 24.3 mm) and body width 14.1���18.5 mm (x��= 16.2 mm). The head is almost as wide as long (length: 10.4���12.8 mm, x��= 11.7 mm; width: 9.3���10.9 mm, x��= 10.2 mm). Head height 6.9���8.4 mm (x = 7.6 mm). Dorsal surface of head smooth, with 12���15 (x��= 13.5) scales (Hellmich index). Six to nine (x��= 7.0) scales between the frontal and rostral scales. One or two (x��= 1.7) scales between the nasal and canthal scales. Nasal surrounded by seven scales. Minimal contact between nasal and rostral scales observed in only two individuals. Interparietal lesser than or equal to parietals, surrounded by 6���8 (x��= 7.1) scales. Three to five (x��= 3.5) supraocular scales. Eight to nine (x��= 8.8) smooth and round temporal scales. Nine to ten (x��= 9.5) superciliar scales. The ear is always higher than wide (high: 1.6���2.5, x��= 1.9; width: 0.9���1.6, x��= 1.2). One to three auricular scales. Without differentiated supero-posterior auricular scale and supero-anterior auricular scale. Eleven to fourteen upper cilliaries (x��= 12.5). Preocular scale separated by one scale from the loreolabial scales. Postocular not divided. Six to seven (x��= 6.5) loreolabials. Three to four loreolabials (x��= 3.7) are in contact with the subocular. Five to six (x��= 5.8) supralabial scales. Mental scale in contact with four scales. Three to four (x��= 3.7) infralabial scales. The second infralabials are in contact with 2���3 (x��= 2.7) scales. Neck with 29���37 (x��= 33.5) granular scales and without keel. Neck folds (auricular, antehumeral and longitudinal) evident. The longitudinal fold contains 20���29 (x��= 24.5) scales. Antehumeral scales are imbricated, without keel, subtriangular and differentiated. Gulars 29���31 (x��= 30). Without gular fold. Nuchal central scales are cone-shaped, are granular and without keel, like lateral nuchal scales. Body dorsal scales are laminar, slightly keeled or without keel, without mucron, juxtaposed or subjuxtaposed. Scales around midbody 59���66 (x��= 62.3). Dorsal scales between occiput and hind limbs 60���78 (x��= 65.2). Ventral scales 85���92 (x��= 87.1). Pigal scales 11���14 (x��= 12.6). Males and females without precloacal pores. Sixteen to seventeen infradigital lamellae on fourth finger and 17���22 on fourth toe. Scales of the dorsum of the tail are slightly keeled, laminar, imbricated and the ventral scales are without keel, laminar and imbricated. Without sexual dichromatism in Liolaemus yatel sp. nov. (Fig. 2). Dorsal and lateral regions of the head are uniformly light brown to light gray although some specimens show dark specks. The background color of the dorsal regions of the body, limbs and tail is light brown or light gray. Vertebral line absent. Dark brown or faded black paravertebral and lateral spots of varied shapes occur. In most specimens paravertebral spots and lateral spots have a crescent shape with an anterior indentation whereas these spots have a subquadrangular shape in other specimens. In most individuals paravertebral spots join lateral spots forming irregular streaks or lines, transverse to the major body axis, that vary considerably in thickness across specimens. Several individuals show reddish-brown specks anterior to each paravertebral or lateral spot. Some specimens have pale yellow or pale reddish brown faint and discontinuous dorsolateral stripes with diffuse contours. Some dark rosettes, or discontinuous ring-shaped spots, occur in the dorsal regions of the limbs. Only a few specimens show a reddish brown coloration on the posterior flank of the thigh. The color pattern of the body back continues along most of the dorsal region of the tail. Near the end of the tail paravertebral and lateral spots approach without contact and form incomplete rings or pseudo-rings. In most cases the ventral region is pristine white in females, while males exhibit a few black scales against white background irregularly scattered across the gular region, abdomen or cloaca. Some males show a reddish and / or slightly yellowish tone in the ventral areas of the body, the hind limbs and the cloaca. Distribution (Fig. 3): Liolaemus yatel sp. nov. has been found only in its type locality, the Monumento Nacional Bosques Petrificados National Park, Puerto Deseado county, Santa Cruz Province, Argentina. Natural history. From a biogeographical point of view, the area inhabited by Liolaemus yatel belongs to the Patagonian province, Central Patagonian District (Cabrera & Willink, 1973), which is characterized by a mixed steppe of grasses and low-lying thorny shrubs (Soriano, 1983) with cover 60 % in the valleys and lowlands (Bertiller & Bisigato, 1998). In arid sites, vegetation is dominated by the shrubs Chuquiraga avellanedae, Nassauvia glomerulosa, and Junellia tridens and by Stipa spp. grasses. In protected and relatively mesic lowland sites, vegetation is characterized by meadows and dense grasslands dominated by Distichlis spicata and Schoenoplectus spp., and shrubs such as Prosopis denudans, Berberis heterophylla, Schinus spp., Junellia tridens and Colliguaja integerrima. Climate is cold, dry and very windy. Winter frosts are frequent and the mean summer temperature is 17 ��C. Annual rainfall ranges between 100 and 300 mm, and concentrates during autumn and winter; snowfall is rare. Prevailing western winds are often strong. Topography is characterized by plateaus defined by cliffs and steep slopes, narrow valleys and flat or rolling depressions, sometimes quite extensive. In the locality where Liolaemus yatel sp. nov. was found soil is made of sandy clay interspersed with small basalt clasts. The scarce woody vegetation of this open landscape consists of isolated individuals of Atriplex lampa and Suaedea divaricata. The rolling ground becomes remarkably muddy in the rain and large mud crusts are typical during dry periods. This type of habitat (locally known as "guadal") is representative of the type locality (47 �� 41 ' 415 " S, 68 �� 00' 06" W), placed between an intermittent lagoon (Laguna Grande) and a barren geological formation (Bajo Pobre). Specimens of Liolaemus yatel were collected between 11: 30 and 15:00. In the spot where specimen ICPD 329 was found, air temperature at ground level ranged between 23 ��C and 33 ��C in the mid-day hours. At the beginning of March, when the remaining individuals were collected, temperatures ranged from 15 ��C to 20 ��C. Most individuals were found on bare ground, where they perfectly camouflaged against the background. Escape behavior consisted in seeking shelter under bushes or in the cracks of the dry mud. We saw some individuals inside burrows dug in small mounds of sand and mud, or in the walls of small dry streams. We observed other lizard species, apparently in low density, near the spot where Liolaemus yatel sp. nov. was found. Species that coexist with L. yatel sp. nov. include Liolaemus boulengeri, L. fitzingeri, L.kingii, L. bibronii, Diplolaemus bibronii, D. darwinii, and Homonota darwini. All of them were found in the ecotone between guadal and shrub-steppe. Etymology. The species name refers to the term that the native Tehuelche people uses to name the rocky ground that surrounds the sites where the specimens were collected., Published as part of Abdala, Cristian Sim��n, Procopio, Diego Esteban, Stellatelli, Oscar An��bal, Travaini, Alejandro, Rodr��guez, Alejandro & Monachesi, Mario Ricardo Ruiz, 2014, New Patagonian species of Liolaemus (Iguania: Liolaemidae) and novelty in the lepidosis of the southernmost lizard of the world: Liolaemus magellanicus, pp. 526-542 in Zootaxa 3866 (4) on pages 528-537, DOI: 10.11646/zootaxa.3866.4.4, http://zenodo.org/record/225761, {"references":["Breitman, M. F., Avila, L. J., Sites, J. W. Jr. & Morando, M. (2011 b) Lizards from the end of the world: Phylogenetic relationships of the Liolaemus lineomaculatus section (Squamata: Iguania: Liolaemini). Molecular Phylogenetics and Evolution, 59 (2), 364 - 376. http: // dx. doi. org / 10.1016 / j. ympev. 2011.02.008","Breitman, M. F., Morando, M. & Avila, L. J. (2013) Past and present taxonomy of the Liolaemus lineomaculatus section (Liolaemidae): is the morphological arrangement hypothesis valid?. Zoological Journal of the Linnean Society, 168 (3), 612 - 668. http: // dx. doi. org / 10.1111 / zoj. 12037","Etheridge, R. E. (1995) Redescription of Ctenoblepharys adspersa Tschudi, 1845, and the taxonomy of Liolaeminae (Reptilia: Squamata: Tropiduridae). American Museum Novitates, 3142, 1 - 34.","Abdala, C. S. & Lobo, F. (2006) Description of a new Patagonian lizard species of the Liolaemus silvanae group (Iguania: Liolaemidae). South American Journal of Herpetology, 1 (1), 1 - 8. http: // dx. doi. org / 10.2994 / 18089798 (2006) 1 [1: DOANPL] 2.0. CO; 2","Cabrera, A. L. & Willink, A. (1973) Biogeografia de America Latina, Monografia 13, Serie de Biologia. Secretaria General de la Organizacion de los Estados Americanos, Washington DC., USA, 120 pp.","Soriano, A. (1983) Deserts and semi-deserts of Patagonia. In: West, N. E. (Ed.), Temperate deserts and semi-deserts. Elsevier, Amsterdam, pp. 440 - 454.","Bertiller, M. B. & Bisigato, A. (1998) Vegetation dynamics under grazing disturbance. The state-and-transition model for the Patagonian steppes. Ecologia Austral, 8 (2), 191 - 199."]}

Liolaemus camarones sp. nov. (Figs. 8���12) 1977, Liolaemus fitzingerii fitzingerii, Cei J. M. and J. A. Scolaro, Physis 36 (92): 225���226. 2006, Liolaemus fitzingerii, ��vila, L. J., M. Morando, and J. W. Sites, Jr. Biological Journal of the Linnean Society, 89, 241 ��� 275. 2008, Liolaemus fitzingerii, ��vila, L. J., M. Morando, and J. W. Sites, Jr. Journal of Herpetology 42: 186���196. 2007, Liolaemus sp. 9, Abdala, C. S. Zootaxa, 1538, 84 pp. Holotype. ― FML 23301. Adult male. Beach 3 km south of Camarones, Bahia Camarones, Florentino Ameghino Department, Chubut Province, Argentina. Collected by C. S. Abdala, J. S. Abdala and E. Malovini. January 22 2002. (Figs. 9���10). Paratypes. ― FML 23302 -23308. Seven individuals, 5 males, 2 females. Same data as holotype. FML 24140 - 144. 5 individuals, 3 males, 2 females. Beach 1 km south of Camarones, road to Elola beach, Bah��a Camarones, Florentino Ameghino Department, Chubut Province, Argentina. S 44 �� 48��33.0���� W 69 �� 44��8.9 ����. Collected by C. S. Abdala, F. Cruz, G. Perotti and R. Semhan. December 18 2009. Diagnosis.― Liolaemus camarones sp. nov. belongs to the Liolaemus boulengeri group, which is characterized by having a patch of enlarged scales on the posterior medial surface of the thigh (Etheridge 1995; Abdala 2007) and a hypertrophy of the flexor tibialis internus muscle (Abdala et al. 2006). It differs from the species of the Liolaemus anomalus group (Table 2) reported in Abdala (2007), in having tail long relative to body length, head proportionally longer than wide, palpebral ���comb��� not developed, and more precloacal pores in males and females. It differs from the species of the L. darwinii (Table 2) in having posterior teeth with expanded crowns and different body proportions, and in lacking a transversal black line on the eye. It differs from the species of the L. wiegmannii group (Table 2) in having a row of lorilabial scales (never two or three) and four scales in contact with mental (six in the L. wiegmannii group). L. camarones sp. nov. differs from L. boulengeri, L. donosobarrosi, L. goetschi, L. hermannunezi, L. inacayali, L. josei, L. loboi, L. martorii, L. rothi, L. sagei, L. senguer, L. tehuelche, and L. telsen in having an evident black antehumeral arch, which is lacking in the aforementioned species. It differs from all the species from the L. telsen group, except L. rothi and L. sagei, in having longer snout-vent length (max SVL 95.6 mm vs. 62.3���77.1 mm). It differs from L. mapuche in having four scales in contact with mental (four to six in L. mapuche) and more marked ventral melanism. It differs from L. cuyanus in having four scales in contact with mental (six in L. cuyanus) and more marked ventral melanism. Liolaemus camarones sp. nov. belongs to the L.fitzingerii clade (Abdala 2007), which is characterized by its tricuspidate posterior teeth with expanded crowns, neck equal to or wider than head, cylindrical body, short limbs relative to the trunk, black antehumeral arch present and ventrally expanded, and males with ventral melanism. Within the Liolaemus fitzingerii clade, L. camarones sp.nov. differs from L. canqueli and L. melanops in lacking head melanism and in the dorsal coloration pattern; it does not present the intense green dorsum with irregular, transversal black paravertebral spots of L. melanops or the transversal black and yellow-orange bands of L. canqueli. Liolaemus camarones differs from L. chehuachekenk and L. morenoi in lacking pre- and postescapular spots and having different coloration pattern; it differs from L. fitzingerii in having shorter snout-vent length (maximum SVL 95.6 mm vs. 106 mm), evident sexual dichromatism with different dorsal coloration pattern in males and females, and evident paravertebral spots, and in lacking dorsolateral markings and red scales in the vertebral region. It differs from L. xanthoviridis in the dorsal coloration pattern, with paravertebral spots and vertebral region more uniform, without irregular markings and antehumeral arch narrower, never reaching forelimbs. Description of the holotype. ― (Figs. 9,10) Adult male. SVL 92.7 mm. Trunk length 42.5 mm. Head longer (18.1 mm) than wide (15.5 mm). Head height 12.2 mm. Eye diameter 4.1 mm. Interorbital distance 8.4 mm. Orbit��� auditory meatus distance 5.9 mm. Auditory meatus 2.9 mm high, 1.7 mm wide. Orbit���commissure of mouth distance 3.2 mm. Internares 2.9 mm. Subocular scale 4.6 mm. Femur length 15.5 mm, tibia 18.1 mm, and foot 24.8 mm. Humerus length 13.1 mm. Tail length 125.8 mm. Dorsal surface of the head smooth, with 15 scales. Rostral wider than tall, bordered by six scales. Mental larger than rostral, trapezoidal, bordered by four scales. Nasal not in contact with rostral. Four internasals. Nasal surrounded by eight scales, separated from canthal by two scales. Five scales between frontal and supercilliaries. Six scales between frontal and rostral. Frontal divided into two scales. Two postrostrals. Interparietal smaller than parietals, in contact with six scales. Orbital semicircles complete. Preocular separated from lorilabial row by one scale. Four scales in anterior margin of auditory meatus and two in superior margin of auditory meatus. Nine smooth temporals. Three lorilabials in contact with subocular. Ten supralabials, none in contact with subocular. Nine supraoculars. Eight lorilabials. Seven infralabials, second in contact ventrally with two scales. Six chinshields, second pair separated by two scales. Seventy-two scales around midbody. Seventy-eight round, imbricate, keeled dorsals from occiput to hind limbs. Thirty-six scale rows in dorsum. Scales of flank same size and shape as dorsals. A hundred-sixteen ventrals, same size as dorsals, flat, imbricate. Thirty-seven smooth, weakly imbricate gulars. Seven precloacal pores. Antehumeral scales flat, of equal size to or larger than dorsals. Postauricular, rectal and longitudinal folds present. Scales on the longitudinal fold granular and smooth. Fourth finger with 18 subdigital lamellae; fourth toe with 24. Infracarpals flat, imbricate, trifid and infratarsals flat, imbricate, not trifid. Color in life. ―Head greenish-grey with scattered red scales on the frontal and parietal regions. Conspicuous, black antehumeral arch. Dorsum and lateral region of the body dark green with small white-yellowish paravertebral spots, surrounded by a black line, thicker on the anterior margin of the spot. Vertebral region brown with two brown-orange longitudinal stripes. Small orange spots scattered between paravertebral spots and dorsolateral stripes. Body size with several white-yellowish scales and spots. Fore- and hind limbs same color as dorsum and lateral region of body with white-yellowish scales. Dorsal tail light green with scattered white-yellowish spots. Throat bluish green. Gular region with an incomplete melanic collar. Chest to cloacal region, ventral forelimbs and base of tail melanic; ventrolateral region, thighs and ventral tail bluish green. Variation.― (Table 3) Based on 12 specimens. Head longer (14.5���18.5 mm; mean = 16.8 mm) than wide (11.8���15.5 mm; mean = 14.0 mm). Head height 11.2���13.7 mm (mean = 13.3 mm). Neck wider than head. SVL 67.8���95.6 mm (mean = 84.7 mm), averaging 2.3 times longer than trunk. Tail length 91.1 ��� 132.0 mm (mean = 116.1 mm). Humerus length 10.7���15.1 mm (mean = 13.9 mm). Dorsal surface of head smooth, with 15���17 scales. One row of lorilabials, same size as supralabials. Seven to eight lorilabials. Eight to 11 supralabials. Frontal divided horizontally, into two to four scales. Interparietal always smaller than parietals, surrounded by 5���7 scales. Nasal surrounded by eight scales. Six to seven infralabials. Mental in contact with four scales. Nine to ten smooth or slightly keeled round temporals. Longitudinal, postauricular, and antehumeral folds present. Horizontal fold Yshaped between the shoulder and the auditory meatus. Scales around midbody 71���79 (mean = 73.6). Gulars 33���37 (mean = 34.8). Dorsal scales between occiput and hind limbs 71���78 (mean = 74.1). Ventrals 103���120 (mean = 110.9). Males with 7���10 (mean = 8.6) precloacal pores. Females without precloacal pores. Dorsals flat, imbricate, keeled. 1720 infradigital lamellae on fourth finger and 24���26 on fourth toe. Variation in color pattern.― With sexual dichromatism. Males with head greenish grey or brown with reddish scales. Black antehumeral arch present. Dorsum and lateral region of the body dark green or brownish green, small yellowish white paravertebral spots, surrounded by a black line, thicker on the anterior margin of the spot. Some individuals with a diffuse white or yellow vertebral line. Paravertebral spots almost quadrangular with anterior margin diagonal and posterior rounded. Vertebral region brown. Two thin, faint orange-brown dorsolateral stripes. In some specimens, these stripes are discontinuous or absent. Between paravertebral spots and dorsolateral stripes, several orange, yellow or brown scales or markings. Lateral region of the body, tail and fore- and hind limbs with numerous yellowish white and light blue scales. Fore- and hind limbs same color as dorsum. Dorsal tail green or brown, lighter than dorsum. Throat greenish blue or yellowish with grey markings. Gular region with a melanic, sometimes incomplete collar. Chest to cloacal region, hind limbs and base of tail melanic, greenish grey or whitish. When melanic, lateral region of the body, forelimbs and tail greenish grey or light green ventrally. Females with head grey or reddish orange (Figs. 11���12). Dorsum reddish brown or bright brown. Antehumeral arch light grey or absent. Vertebral line discontinuous, yellowish white. Paravertebral spots similar to males. Faint dorsolateral stripes with yellowish white markings. Lateral region of the body always lighter than dorsum, with spots similar to paravertebral. Forelimbs grey with few markings dorsally. Throat to cloacal region and tail bluish white with some black scales, more intense on sides of the body. Natural history.― Liolaemus camarones sp. nov. inhabits sand dunes near Bah��a Camarones. The predominant vegetation is the shrub Chuquiraga sp. (shrub with orange flowers and light green leaves) or two plants adapted to saline soils: Atriplex lampa (zampa) and Suaeda divaricata (vidriera). Liolaemus camarones is insectivorous and oviparous. Adults live in small holes under shrubs and usually bask during the hours of highest solar radiation. There is no more information about its biology. Distribution.― Liolaemus camarones sp nov. inhabits the beaches of Bah��a Camarones and nearby areas, Chubut Province, Argentina (Fig. 8). Etymology.― The word camarones refers to the locality where this new species occurs, Bahia Camarones., Published as part of Abdala, Cristian Sim��n, G��mez, Juan Manuel D��az & Heredia, Viviana Isabel Juarez, 2012, From the far reaches of Patagonia: new phylogenetic analyses and description of two new species of the Liolaemus fitzingerii clade (Iguania: Liolaemidae), pp. 34-60 in Zootaxa 3301 on pages 44-48, DOI: 10.5281/zenodo.246399

Liolaemus camarones sp. nov. (Figs. 8–12) 1977, Liolaemus fitzingerii fitzingerii, Cei J. M. and J. A. Scolaro, Physis 36 (92): 225–226. 2006, Liolaemus fitzingerii, Ávila, L. J., M. Morando, and J. W. Sites, Jr. Biological Journal of the Linnean Society, 89, 241 – 275. 2008, Liolaemus fitzingerii, Ávila, L. J., M. Morando, and J. W. Sites, Jr. Journal of Herpetology 42: 186–196. 2007, Liolaemus sp. 9, Abdala, C. S. Zootaxa, 1538, 84 pp. Holotype. ― FML 23301. Adult male. Beach 3 km south of Camarones, Bahia Camarones, Florentino Ameghino Department, Chubut Province, Argentina. Collected by C. S. Abdala, J. S. Abdala and E. Malovini. January 22 2002. (Figs. 9–10). Paratypes. ― FML 23302 -23308. Seven individuals, 5 males, 2 females. Same data as holotype. FML 24140 - 144. 5 individuals, 3 males, 2 females. Beach 1 km south of Camarones, road to Elola beach, Bahía Camarones, Florentino Ameghino Department, Chubut Province, Argentina. S 44 ° 48´33.0´´ W 69 ° 44´8.9 ´´. Collected by C. S. Abdala, F. Cruz, G. Perotti and R. Semhan. December 18 2009. Diagnosis.― Liolaemus camarones sp. nov. belongs to the Liolaemus boulengeri group, which is characterized by having a patch of enlarged scales on the posterior medial surface of the thigh (Etheridge 1995; Abdala 2007) and a hypertrophy of the flexor tibialis internus muscle (Abdala et al. 2006). It differs from the species of the Liolaemus anomalus group (Table 2) reported in Abdala (2007), in having tail long relative to body length, head proportionally longer than wide, palpebral ‘comb’ not developed, and more precloacal pores in males and females. It differs from the species of the L. darwinii (Table 2) in having posterior teeth with expanded crowns and different body proportions, and in lacking a transversal black line on the eye. It differs from the species of the L. wiegmannii group (Table 2) in having a row of lorilabial scales (never two or three) and four scales in contact with mental (six in the L. wiegmannii group). L. camarones sp. nov. differs from L. boulengeri, L. donosobarrosi, L. goetschi, L. hermannunezi, L. inacayali, L. josei, L. loboi, L. martorii, L. rothi, L. sagei, L. senguer, L. tehuelche, and L. telsen in having an evident black antehumeral arch, which is lacking in the aforementioned species. It differs from all the species from the L. telsen group, except L. rothi and L. sagei, in having longer snout-vent length (max SVL 95.6 mm vs. 62.3–77.1 mm). It differs from L. mapuche in having four scales in contact with mental (four to six in L. mapuche) and more marked ventral melanism. It differs from L. cuyanus in having four scales in contact with mental (six in L. cuyanus) and more marked ventral melanism. Liolaemus camarones sp. nov. belongs to the L.fitzingerii clade (Abdala 2007), which is characterized by its tricuspidate posterior teeth with expanded crowns, neck equal to or wider than head, cylindrical body, short limbs relative to the trunk, black antehumeral arch present and ventrally expanded, and males with ventral melanism. Within the Liolaemus fitzingerii clade, L. camarones sp.nov. differs from L. canqueli and L. melanops in lacking head melanism and in the dorsal coloration pattern; it does not present the intense green dorsum with irregular, transversal black paravertebral spots of L. melanops or the transversal black and yellow-orange bands of L. canqueli. Liolaemus camarones differs from L. chehuachekenk and L. morenoi in lacking pre- and postescapular spots and having different coloration pattern; it differs from L. fitzingerii in having shorter snout-vent length (maximum SVL 95.6 mm vs. 106 mm), evident sexual dichromatism with different dorsal coloration pattern in males and females, and evident paravertebral spots, and in lacking dorsolateral markings and red scales in the vertebral region. It differs from L. xanthoviridis in the dorsal coloration pattern, with paravertebral spots and vertebral region more uniform, without irregular markings and antehumeral arch narrower, never reaching forelimbs. Description of the holotype. ― (Figs. 9,10) Adult male. SVL 92.7 mm. Trunk length 42.5 mm. Head longer (18.1 mm) than wide (15.5 mm). Head height 12.2 mm. Eye diameter 4.1 mm. Interorbital distance 8.4 mm. Orbit– auditory meatus distance 5.9 mm. Auditory meatus 2.9 mm high, 1.7 mm wide. Orbit–commissure of mouth distance 3.2 mm. Internares 2.9 mm. Subocular scale 4.6 mm. Femur length 15.5 mm, tibia 18.1 mm, and foot 24.8 mm. Humerus length 13.1 mm. Tail length 125.8 mm. Dorsal surface of the head smooth, with 15 scales. Rostral wider than tall, bordered by six scales. Mental larger than rostral, trapezoidal, bordered by four scales. Nasal not in contact with rostral. Four internasals. Nasal surrounded by eight scales, separated from canthal by two scales. Five scales between frontal and supercilliaries. Six scales between frontal and rostral. Frontal divided into two scales. Two postrostrals. Interparietal smaller than parietals, in contact with six scales. Orbital semicircles complete. Preocular separated from lorilabial row by one scale. Four scales in anterior margin of auditory meatus and two in superior margin of auditory meatus. Nine smooth temporals. Three lorilabials in contact with subocular. Ten supralabials, none in contact with subocular. Nine supraoculars. Eight lorilabials. Seven infralabials, second in contact ventrally with two scales. Six chinshields, second pair separated by two scales. Seventy-two scales around midbody. Seventy-eight round, imbricate, keeled dorsals from occiput to hind limbs. Thirty-six scale rows in dorsum. Scales of flank same size and shape as dorsals. A hundred-sixteen ventrals, same size as dorsals, flat, imbricate. Thirty-seven smooth, weakly imbricate gulars. Seven precloacal pores. Antehumeral scales flat, of equal size to or larger than dorsals. Postauricular, rectal and longitudinal folds present. Scales on the longitudinal fold granular and smooth. Fourth finger with 18 subdigital lamellae; fourth toe with 24. Infracarpals flat, imbricate, trifid and infratarsals flat, imbricate, not trifid. Color in life. ―Head greenish-grey with scattered red scales on the frontal and parietal regions. Conspicuous, black antehumeral arch. Dorsum and lateral region of the body dark green with small white-yellowish paravertebral spots, surrounded by a black line, thicker on the anterior margin of the spot. Vertebral region brown with two brown-orange longitudinal stripes. Small orange spots scattered between paravertebral spots and dorsolateral stripes. Body size with several white-yellowish scales and spots. Fore- and hind limbs same color as dorsum and lateral region of body with white-yellowish scales. Dorsal tail light green with scattered white-yellowish spots. Throat bluish green. Gular region with an incomplete melanic collar. Chest to cloacal region, ventral forelimbs and base of tail melanic; ventrolateral region, thighs and ventral tail bluish green. Variation.― (Table 3) Based on 12 specimens. Head longer (14.5–18.5 mm; mean = 16.8 mm) than wide (11.8–15.5 mm; mean = 14.0 mm). Head height 11.2–13.7 mm (mean = 13.3 mm). Neck wider than head. SVL 67.8–95.6 mm (mean = 84.7 mm), averaging 2.3 times longer than trunk. Tail length 91.1 – 132.0 mm (mean = 116.1 mm). Humerus length 10.7–15.1 mm (mean = 13.9 mm). Dorsal surface of head smooth, with 15–17 scales. One row of lorilabials, same size as supralabials. Seven to eight lorilabials. Eight to 11 supralabials. Frontal divided horizontally, into two to four scales. Interparietal always smaller than parietals, surrounded by 5–7 scales. Nasal surrounded by eight scales. Six to seven infralabials. Mental in contact with four scales. Nine to ten smooth or slightly keeled round temporals. Longitudinal, postauricular, and antehumeral folds present. Horizontal fold Yshaped between the shoulder and the auditory meatus. Scales around midbody 71–79 (mean = 73.6). Gulars 33–37 (mean = 34.8). Dorsal scales between occiput and hind limbs 71–78 (mean = 74.1). Ventrals 103–120 (mean = 110.9). Males with 7–10 (mean = 8.6) precloacal pores. Females without precloacal pores. Dorsals flat, imbricate, keeled. 1720 infradigital lamellae on fourth finger and 24–26 on fourth toe. Variation in color pattern.― With sexual dichromatism. Males with head greenish grey or brown with reddish scales. Black antehumeral arch present. Dorsum and lateral region of the body dark green or brownish green, small yellowish white paravertebral spots, surrounded by a black line, thicker on the anterior margin of the spot. Some individuals with a diffuse white or yellow vertebral line. Paravertebral spots almost quadrangular with anterior margin diagonal and posterior rounded. Vertebral region brown. Two thin, faint orange-brown dorsolateral stripes. In some specimens, these stripes are discontinuous or absent. Between paravertebral spots and dorsolateral stripes, several orange, yellow or brown scales or markings. Lateral region of the body, tail and fore- and hind limbs with numerous yellowish white and light blue scales. Fore- and hind limbs same color as dorsum. Dorsal tail green or brown, lighter than dorsum. Throat greenish blue or yellowish with grey markings. Gular region with a melanic, sometimes incomplete collar. Chest to cloacal region, hind limbs and base of tail melanic, greenish grey or whitish. When melanic, lateral region of the body, forelimbs and tail greenish grey or light green ventrally. Females with head grey or reddish orange (Figs. 11–12). Dorsum reddish brown or bright brown. Antehumeral arch light grey or absent. Vertebral line discontinuous, yellowish white. Paravertebral spots similar to males. Faint dorsolateral stripes with yellowish white markings. Lateral region of the body always lighter than dorsum, with spots similar to paravertebral. Forelimbs grey with few markings dorsally. Throat to cloacal region and tail bluish white with some black scales, more intense on sides of the body. Natural history.― Liolaemus camarones sp. nov. inhabits sand dunes near Bahía Camarones. The predominant vegetation is the shrub Chuquiraga sp. (shrub with orange flowers and light green leaves) or two plants adapted to saline soils: Atriplex lampa (zampa) and Suaeda divaricata (vidriera). Liolaemus camarones is insectivorous and oviparous. Adults live in small holes under shrubs and usually bask during the hours of highest solar radiation. There is no more information about its biology. Distribution.― Liolaemus camarones sp nov. inhabits the beaches of Bahía Camarones and nearby areas, Chubut Province, Argentina (Fig. 8). Etymology.― The word camarones refers to the locality where this new species occurs, Bahia Camarones.

Liolaemus burmeisteri sp. nov. Holotype. ��� MLP.S 2612 (Figs. 1, 2), an adult male from Ruta Provincial 41, 7 km S Caepe Malal, Chos Malal Department, Neuqu��n, Argentina (37 �� 13 ��� 51.4 ���S, 70 �� 22 ��� 24.3 ���W, 1037 m), collected 19 January 2007. Paratypes. ��� MLP.S 2613, LJAMM 5241���5242 (male) and MLP.S 2614-2617 (female) from same locality as holotype. Collected 20 January 2003. LJAMM 7639, 7641, 7646 (male) and 7637 ���7638, 7640, 7642 ���7645, 7647 (female) from same locality as holotype, collected 19 January 2007. Liolaemus bur- Liolaemus Liolaemus Liolaemus Liolaemus Liolaemus Liolaemus shi- Character meisteri antumalguen chillanensis elongatus smaug choique tan sp.nov. (n = (n = 11) (n = 99) (n = 21) (n = 22) 19) Max. SVL 85.2 107.7 70.3 94.7 68.3 90.7 94.7 (mm) SAM 76.2 �� 2.7 (70 - 77.0�� 3.6 (68- 81 - 95 77.5 ��4.0(68- 74.1 �� 2.6 (70- 74 - 88 78.1 �� 3.9 (71- 81) 82) 87) 79) 85) Diagnosis. ��� Liolaemus burmeisteri can be easily distinguished from all other members of the Liolaemus elongatus clade, by its light brown/ochre general coloration not found in any of the other species and by its homogeneous dorsal pattern without bands, stripes or spotted areas (Table 1). Liolaemus burmeisteri lacks of a well-defined bodybanding pattern and dark ochre/green, or black general coloration, typical of L. elongatus. Liolaemus smaug has darker coloration and fewer numbers of dorsal scales without overlapping (64���75, 0 = 70.3 vs 76���85, 0 = 81.1 in L. burmeisteri) and higher numbers of ventral scales (107���119, 0 = 111.9 vs 99���110, 0 = 104.7 in L. burmeisteri) with little overlap. Liolaemus smaug has a dorsal banding pattern characterized by a very variable but always present vertebral band ranging from a well marked gray-ochre zone to a dark spotted area, and a very wide and dark lateral band, all traits absent in live specimens of L. burmeisteri. Liolaemus chillanensis has a different dorsal coloration pattern, with a vertebral band formed by dark spots, more ochre dark general coloration, more darker lateral bands, and more scales around midbody (81-95 vs 70-81). Liolaemus antumalguen is a larger species (maximum SVL 107.7, 0 = 94.4 vs 85.2 mm, 0 = 74.5), with more enlarged neck pouches, and with fewer scales along the dorsum and little overlap with L. burmeisteri (70-78, 0 = 73.0 vs 76-85, 0 = 81.1). General coloration is darker in L. antumalguen, usually with a dorsal pattern of black dots, and black head and belly, and tail without any pattern of tail rings, characteristics never found in L. burmeisteri. Liolaemus shitan has fewer numbers of dorsal scales (66���78, 0 = 70.6 vs 76���85, 0 = 81.1 in L. burmeisteri) and higher numbers of ventral scales (106���129, 0 = 120.6 vs 99���110, 0 = 104.7 in L. burmeisteri) with little overlap. Liolaemus shitan has a general melanic coloration without any apparent pattern, a coloration trait absent in L. burmeisteri. Liolaemus choique has higher numbers of ventral scales (118���135 vs 99���110 in L. burmeisteri) without overlapping. General coloration is yellow in Liolaemus choique with a dorsal pattern of vertebral and lateral black melanic bands never found in L. burmeisteri. Description of holotype. ��� Adult male. SVL 85.2 mm, total length 210.2 mm; tail incomplete, regenerated (125 mm length). Axilla���groin distance 35.2 mm. Head 19.2 mm long (from anterior border of auditory meatus to tip of snout), 16.6 mm wide (at anterior border of auditory meatus), 9.9 mm high. Arm length 23.1 mm. Tibia length 17.7 mm. Foot length 23.2 mm (ankle to tip of claw on fourth toe). Dorsal head scales irregular, some smooth and others with small holes (not scale organs), irregularities appearing more frequently in parietal���temporal areas. Scale organs more abundant in the anterior head region than in the parietal and temporal region. Nineteen scales between rostral and occiput (at level of anterior border of auditory meatus). Rostral scale wider (4.0 mm) than high (1.3 mm). Two postrostrals. Four internasals, irregular. Five frontonasals, irregular. Four prefrontals, irregular. Frontal scale divided longitudinally, forming two scale rows between circumorbitals. Four scales between frontal and rostral. Interparietal pentagonal, similar size to parietals. Interparietal surrounded by seven scales, with a central, small, and obscure ������parietal eye������ in the scale center. Supraorbital semicircles complete. Seven enlarged supraoculars. Five scales between frontal and supercilliaries. Seven supercilliaries, irregular flattened and elongated. Nineteen temporals on each side, smooth and protruding with 1���3 scales organs. Canthal scales separated from nasal by one scale. Loreal region concave. Seven scales surrounding nasals on each side. Nasal in slight contact with rostral on each side. Orbit with 16 upper and 14 lower ciliaries. Orbit diameter 2.5 x 4.2 mm (measured between upper and lower ciliaries). Subocular scale elongate. Preocular fragmented on two scales. Six lorilabials, three in contact with subocular. Six supralabials on each side. Fourth supralabial curved upward posteriorly but fourth not in contact with subocular. Infralabials, four, first scale twice as high as posterior infralabials. Postrostrals, internasals, frontonasals, prefrontals, loreal, lorilabials, supra��� and infralabials with conspicuous scale organs. Three outwardly projecting scales along anterior border of auditory meatus. Auditory meatus about twice as high (3.9 mm) as wide (1.7 mm). Lateral scales of neck granular with skin below appearing slightly inflated. Antehumeral, longitudinal, and postauricular neck folds distinct, oblique less conspicuous, gular incomplete, rictal not present. Thirty���seven scales between auditory meatus and antehumeral fold (counted along longitudinal fold). Scales of dorsal neck region similar to dorsals. Mental wider (3.9 mm) than long (2.1 mm), followed posteriorly by two rows of chinshields with four scales in the left side and six on right side. Chinshield rows four scales in contact with mental. Throat scales between chinshields slightly juxtaposed, strongly imbricated toward auditory meatus. Fourty���nine gulars between auditory meatus. Eighty���two dorsal scales between occiput and anterior surface of thighs. Thirty���six longitudinal keeled scales rows. Scales become larger and less keeled through flanks. Flank scales with one scale organ at the tip. Scales small and granular around limb insertions. Eighty scales around midbody. Ventral scales of similar size to dorsals, but smooth and round, 104 between mental and cloacal aperture. Precloacal scales slightly larger than ventrals. Five precloacal pores. Tail quadrangular in cross section near cloacal area, becoming oval to round in the rest. Caudal scales in discernible annuli. Dorsal and upper lateral caudals scales keeled, imbricate, mucronate, and outward projecting. Lower lateral scales moderately keeled and mucronated, and ventral scales smooth. Suprabrachials, imbricate, moderately keeled; prebrachials imbricate, weakly keeled, grading into smaller subimbricate or granular infrabrachials. Supra���antebrachials imbricate, very weakly keeled; post���antebrachials imbricate, moderately keeled with 1���3 mucronated keeled toward the ventral region; pre���antebrachials imbricate, smooth; and infra���antebrachials imbricate, becoming of smooth in the posterior region to moderately keeled and mucronate at the anterior region. Suprametacarpals imbricate, smooth; inframetacarpals imbricate, keeled, 2���3 mucronate. Supradigitals of manus smooth, wider than long; subdigitals with three keels, each terminating in a short blunt mucron, numbering: I: 10, II: 15; III: 22; IV: 22; V: 16. Claws robust, moderately curved, opaque brown. Suprafemorals imbricate, moderately keeled; prefemorals and infrafemorals imbricate, smooth. Postfemorals small, granular. Supra��� and pretibials imbricate, moderately blunty keeled; infratibials imbricate, smooth. Supratarsals imbricate moderately keeled; and infratarsals imbricate, 3 ��� keeled, mucronate. Supradigitals of foot smooth, wider than long; Subdigital scales keeled, becoming of 4 ���keeled in the posterior region of digit to 3 ���keeled in the tip, mucronate, numbering: I: 13; II: 18; III: 25; IV: 30; V: 19. Claws robust, moderately curved, opaque brown. Color of holotype in life. ��� General body coloration uniform, light brown or light ochre. Dorsal and lateral head scales, with scattered few dark brown smudges on postrostral, internasal, frontonasal, prefrontal, temporals and frontal scales. A thin black line on the superior border of the subocular scale. Infralabial scales whitish with small dark brown areas. Dorsal scales between occiput and cloacal region light brown/ochre without definite pattern, speckled with very small (one scale) white or black spots. Tail light brown, weakly ringed with ochre bands. Body lateral region with a dark brown band between axilla and groin, darker than dorsal scales, speckled with a few white spots (one scale). Upper limb surfaces ochre with reticulated of dark brown. Ventral scales of throat, neck, chest, belly and forelimbs light gray with some melanic sectors, more marked in belly midline. Ventral scales of lower belly and femoral region, bright yellow. Ventral scales of cloacal region and tail whitish. Color of holotype in preservative. ��� After five years in preservative, the dorsal coloration of the head, dorsum, body flanks and tail become darker while maintaining the contrast, but the white spots largely disappeared. Ventral scales of throat, neck, chest, belly and forelimbs darkened, and distinctive yellow ventral coloration of the femoral region and belly turned dark gray. General pattern of coloration become more marked and general coloration disappeared. Variation. ��� Liolaemus burmeisteri adults ranging from 60.5���85.2 mm SVL. As in other members of this group, no obvious body size sexual dimorphism was observed (except tail expansion in cloacal areas of males and slightly smaller head width in females). In seven males (Table 2, Fig. 3): SVL: 77.8���85.2 mm. Head length: 17.3��� 19.25 mm. Head width: 13.6���16.6 mm. head height: 8.15���9.95 mm. Foot length: 23.2���24.9 mm. Tibial length: 17.4���18.9 mm. Arm length: 21.9���25.4 mm. Midbody scales: 73���81. Dorsal scales (between occiput at the anterior margin of auditory meatus and anterior surface of thighs): 76���83. Ventral scales 101���109. Fourth toe lamellae: 29��� 31. Supralabial scales: 6���8. Infralabial scales: 5. Cloacal pores: 0���5. In twelve females (Table 2, Fig. 4): SVL: 58.3���78.9 mm. Head length: 13.6���16.7 mm. Head width: 10.9���13.8 mm. head height: 6.30���8.15 mm. Foot length: 20.5 ���23.0 mm. Tibial length: 13.9���17.6 mm. Arm length: 18.7���23.4 mm. Midbody scales: 70���79. Dorsal scales: 76���85. Ventral scales: 99���110. Fourth toe lamellae: 27���31. Supralabial scales: 7���9. Infralabial scales: 4���5. For nineteen individuals (males and females): dorsal scales: 81.15 ��2,89 (76���85). Interparietal scale usually pentagonal shaped, bordered by 6���8 scales. Supralabial scales 6���9. Infralabial scales 4���5. Third finger lamellae ranges 19���23. Fourth toe lamellae ranges 27���31. Dorsal and lateral coloration in life is almost identical in all individuals and varies only in intensity. Yellow ventral coloration of males in femoral and lower belly region is variable in extent and intensity in the areas indicated. In preservative, dorsal coloration of all individuals fades to a darker, although all retained the contrast between back and head and flanks and in some individuals scattered dorsal white spots disappeared. All distinctive femoral areas and lower belly coloration fades in preservative from yellow to dark gray. Etymology. ��� The specific name is to honor Karl Hermann Konrad Burmeister, a German naturalist, paleontologist and zoologist. Carlos Germ��n Conrado Burmeister (as he was known in Argentina) was born in Stralsund, Germany in 1807 and past away in Argentina in 1892 after a prolific work with near 300 publications about animals, plants, geology, and paleontology of South America, including its Description Physique de la R��publique Argentine d���apr��s des observations personnelles et ��trang��res (with a version in German). He was director of the Museo P��blico de Buenos Aires (now Museo Argentino de Ciencias Naturales Bernardino Rivadavia) for 30 years and was in charge of the organization of the National Academy of Sciences in C��rdoba, founded by the Argentinean president Domingo F. Sarmiento. Distribution. ��� Liolaemus burmeisteri is known only from the type locality (Fig. 5), Ruta Provincial 41, 7 km S Caepe Malal (37 �� 13 ��� 51.4 ���S, 70 �� 22 ��� 24.3 ���W, 1037 m), northwestern Patagonia, Chos Malal Department, Neuqu��n Province, Argentina. Natural history. ��� The collection area was on a small plateau with sedimentary rocks and sparse vegetation. The type locality is a plateau with rocky soils and with medium to big���sized rocks scattered or rocky outcrops, alternating with areas of loose sand (Fig. 6). This area is only sparsely vegetated, with soil erosion resulting from grazing by goats and horses. However, in undisturbed areas, typical vegetation is composed of plants of the genera Stillingia, Colliguaja, Nassauvia, Haploppapus, Fabiana, Schinus and Stipa, which are representatives of the Payunia District, Patag��nica Province, Andino���Patag��nico Domain (Roig 1998). Liolaemus burmeisteri appears to be restricted to rocky areas, similar to many other members of this clade (Cei 1974, 1979, 1986, 1993; Espinoza & Lobo 2003; Espinoza et al. 2000; Hulse 1979). When active, lizards move across the rocky substratum and bask on horizontal surfaces or on top of medium���sized stones. When disturbed, L. burmeisteri usually attempts to escape into the cracks and crevices in the rocks. Based on analyses of stomach contents, L. burmeisteri feeds on a variety of insects and other small arthropods. In two digestive tracts, we found Orthoptera, Coleoptera, Formicidae, Solifugae, Araenae, sand grains and indeterminate parts of arthropods. These contents are very similar to those found by Videla (1983) in L. elongatus from Mendoza Province. Detailed biological information has been published only for a few phylogenetically related lizards (e.g. Ibarg��engoytıa & Cussac 1998; Quatrini et al. 2001; Videla 1983), with observations in Cei (1986), Espinoza & Lobo (2003), Espinoza et al. (2000), Hulse (1979), and Schulte et al. (2000). No conclusive evidence of viviparity can be offered, but all related species of the elongatus clade share this reproductive mode (Cei 1986; Espinoza & Lobo 2003; Espinoza et al. 2000; Hulse 1979; Schulte et al. 2000). At the type locality, L. burmeisteri was found in syntopy with Homonota darwinii, and no other species of lizards were observed. Remarks ���A mtDNA gene tree analysis, including the new described species as well as other related species and candidate species of this and other related clades is depicted in Fig. 7. This tree is based on the mitochondrial gene fragment cyt-b (805 bp). Liolaemus burmeisteri is recovered as the sister taxon of L. smaug but with very low support. The petrophilus and kriegi clades also include candidate species and are recovered with strong support (0.95 and 0.99 respectively). Recently a typographic error named this clade as buergeri in the remarks section of the description of L. antumalguen (Avila et al. 2010). The objective of this tree is to show the position of this new species in relation with these three related clades. Phylogenetic relationships for these clades, including several lines of evidence, are under study by our research group and a detailed analysis will be published elsewhere. Males (N= 7) Females (N= 12) Mean SD Range Mean SD Range antumalguen austromendocinus buergeri capillitas ceii chillanensis curis cristiani dicktracy elongatus flavipiceus gununakuna heliodermis kriegi parvus leopardinus petrophilus punmahuida talampaya thermarun tregenzai tulkas umbrifer villaricensis Cei, 1974 E E E Cei, 1975 E B K E K E Cei, 1979 E K K E K E Ortiz, 1981 E K K E K E Cei, 1986 E K K E K E Cei, 1993 E E E E Espinoza et al., 2000 E E E E E E Schulte et al., 2000 X X X X X X X Lobo, 2001 E K E E E K X Morando et al., 2003 P K P K E K P P P Avila et al., 2004 P K P P E P K P P P P P Lobo, 2005 C C C C D��az G��mez & Lobo, 2006 Torres-P��rez et al., 2009 E E K Lobo et al., 2010 E X C X X C E E E C X E E X C X E C C Avila et al., 2010 E E E E E Species composition of the elongatus clade has changed during the last thirty years (Table 3), but based on available knowledge and ongoing systematic studies of the petrophilus (Feltrin et al. in prep.) and the elongatus (Medina et al. in prep.) clades; we consider the elongatus clade composition to include: L. elongatus Koslowsky 1896, L. antumalguen Avila et al. 2010, L. chillanensis (Torres-Perez et al. 2009), three recently described species, Liolaemus smaug, L. choique and L. shitan (Abdala et al. 2010), and L. burmeisteri. The three described species by Abdala et al. (2010) were considered in previous studies as populations of Liolaemus elongatus (Morando et al. 2003; Morando et al. 2004). Liolaemus thermarum was described from a type locality named Termas del Azufre, Malarg��e, in Mendoza Province, 10 km from the Peteroa volcano (Videla & Cei 1996); according to the description of the type locality all lizards were collected in a glacial valley named ���Ba��os del Azufre��� along a volcanic relief Azufre-Planchon- Peteroa. Later, Espinoza et al. (2000) cited this species for Laguna Ni��a Encantada, close to Las Le��as, but Cei & Videla (2003) point out that those samples do not correspond to L. thermarum. Morando et al. (2003) suggested that their sample named Liolaemus sp. 5, from an area close to L. thermarum type locality, could correspond to this species. However, adequate comparison to type specimens deposited in the MACN was not possible due to the poor preservation conditions, and the few characters that could be observed showed some discordance (e.g., absence of precloacal pores in L. thermarum whereas two to four pores are present in Liolaemus sp. 5). Later, we analyzed sample, Published as part of Avila, Luciano Javier, Perez, Cristian Hernan Fulvio, Medina, Cintia Debora, Sites, Jack Walter & Morando, Mariana, 2012, A new species of lizard of the Liolaemus elongatus clade (Reptilia: Iguania: Liolaemini) from Curi Leuvu River Valley, northern Patagonia, Neuqu��n, Argentina, pp. 37-52 in Zootaxa 3325 on pages 38-49, DOI: 10.5281/zenodo.213183, {"references":["Roig, A. F. (1998) La vegetacion de la Patagonia. In: Correa, M. N. (Ed.), Flora Patagonica. INTA, Buenos Aires, Argentina, pp. 48 - 174.","Cei, J. M. (1974) Revision of the Patagonian iguanids of the Liolaemus elongatus complex. Journal of Herpetology, 8, 219 - 229.","Cei, J. M. (1979) The Patagonian Herpetofauna. In: Duellman, W. E. (Ed.), The South American Herpetofauna: Its origin, Evolution, and Dispersal. Museum of Natural History, University of Kansas, Lawrence, Kansas, pp. 309 - 329.","Cei, J. M. (1986) Reptiles del centro, centro-oeste y sur de la Argentina. Museo Regionale di Scienze Naturali di Torino Monografie, 4, 1 - 527.","Cei, J. M. (1993) Reptiles del noroeste, nordeste y este de la Argentina. Museo Regionale di Scienze Naturali di Torino, Monografie 14, 1 - 949.","Espinoza, R. E. & Lobo, F. (2003) Two new species of Liolaemus lizards from northwestern Argentina: speciation within the northern subclade of the elongatus group (Iguania: Liolaemidae). Herpetologica, 59, 89 - 105.","Espinoza, R. E., Lobo, F. & Cruz, F. B. (2000) Liolaemus heliodermis, a new lizard from northwestern Argentina with remarks on the content of the elongatus group (Iguania: Tropiduridae). Herpetologica, 56, 507 - 516.","Hulse, A. C. (1979) A new Liolaemus (Sauria, Iguanidae) from the high Andes of Argentina, with ecological comments. Annals of Carnegie Museum, 48, 203 - 209.","Ibarguengoytia, N. R. & Cussac, V. E. (1998) Reproduction of the viviparous lizard Liolaemus elongatus in the highlands of southern South America: plastic cycles in response to climate? Herpetological Journal, 8, 99 - 105.","Quatrini, R., Albino, A. & Barg, M. (2001) Variacion morfologica y dieta en dos poblaciones de Liolaemus elongatus Koslowskyi, 1896 (Iguania: Tropiduridae) del noroeste patagonico. Revista Chilena de Historia Natural, 74, 639 - 651.","Schulte, J. A. II, Macey, J. R. Espinoza, R. E. & Larson, A. (2000) Phylogenetic relationships in the iguanid lizard genus Liolaemus: multiple origins of viviparous reproduction and evidence for recurring Andean vicariance and dispersal. Biological Journal of Linnean Society, 69, 75 - 120.","Avila, L. J., Morando, M., Perez, D. R. & Sites, J. W. Jr. (2010) A new species of the Liolaemus elongatus clade (Reptilia: Iguania: Liolaemini) from Cordillera del Viento, northwestern Patagonia, Neuquen, Argentina Zootaxa, 2667, 28 - 42.","Lobo, F. (2001) A phylogenetic analysis of lizards of the Liolaemus chiliensis group (Iguania: Tropiduridae). Herpetological Journal, 11, 137 - 150.","Morando, M., Avila, L. J., Baker, J. & Sites, J. W. Jr. (2004) Phylogeny and phylogeography of the Liolaemus darwinii complex (Squamata: Liolaemidae): evidence for introgression and incomplete lineage sorting. Evolution, 58, 842 - 861.","Lobo, F. (2005) Las relaciones filogeneticas dentro del grupo chiliensis (Iguania, Liolaemidae, Liolaemus): sumando nuevos caracteres y taxones. Acta Zoologica Lilloana, 49, 67 - 89.","Torres-Perez, F., Mendez, M. A., Benavides, E., Moreno, R. A., Lamborot, M., Palma, R. E. & Ortiz, J. C. (2009) Systematics and evolutionary relationships of the mountain lizard Liolaemus monticola (Liolaemini): how morphological and molecular evidence contributes to reveal hidden species diversity. Biological Journal of the Linnean Society, 96, 635 - 650.","Lobo, F., Espinoza, R. E. & Quinteros, S. (2010) A critical review and systematic discussion of recent classification proposals for liolaemid lizards. Zootaxa, 2549, 1 - 30.","Koslowsky, J. (1896) Sobre algunos reptiles de Patagonia y otras regiones argentinas. Revista del Museo de La Plata, 7, 447 - 457.","Abdala, C. S., Quinteros, A. S., Scrocchi, G. J. & Stazzonelli, J. C. (2010) Three new species of the Liolaemus elongatus group (Iguania: Liolaemini) from Argentina. Cuadernos de Herpetologia, 24, 93 - 109.","Videla, F. & Cei, J. M. (1996) A new peculiar Liolaemus species of the '' chiliensis '' phyletic group from the volcanic Cordilleran landscapes of southern Mendoza Province, Argentina (Iguania, Lacertilia, Reptilia). Bolletino Museo Regionale di Scienze Naturali di Torino, 14, 505 - 516.","Cei, J. M. & Videla, F. (2003). A new species of Liolaemus lacking precloacal pores in males from the Andean south-eastern mountains of Mendoza Province, Argentina. (Liolaemidae, Iguania, Lacertilia, Reptilia). Bolletino del. Museo Regionale di Scienze Naturali di Torino, 20, 275 - 290"]}

Liolaemus shehuen sp. nov. (Figs. 4–7) 1999 Cei and Scolaro Liolaemus fitzingerii, Revue francais de Aquariologie, 26, 79- 82. 2007 Liolaemus sp 7. Abdala C. S., Zootaxa, 1538, 84 pp. Holotype. ― FML 22191. Adult male. 60 km west of Telsen, Telsen Department, Chubut Province, Argentina. C. S. Abdala, J. S. Abdala and M. Juarez cols. January 23, 2004. (Figs. 4, 5). Paratypes. ― FML 22192 - 195.4 individuals. 1 male, 3 females. Same data as holotype. FML 22204 - 205.2 individuals. 2 males, same data as holotype. FML 22217 - 219.3 individuals. 1 male, 2 females. 80 km west of Telsen, Telsen Department, Chubut Province, Argentina. C. S. Abdala, J. S. Abdala and M. Juarez cols. January 23, 2004. Diagnosis.― Liolaemus shehuen belongs to the L. boulengeri group, characterized by having a patch of enlarged scales on the posterior medial surface of the thigh (Etheridge 1995; Abdala 2007) and hypertrophy of the flexor tibialis internus muscle (Abdala et al. 2006); tail long relative to body length, head proportionally longer than wide, palpebral ‘comb’ not developed, and more precloacal pores both in males and females than in the remaining species of the clade. Liolaemus shehuen differs from the species of the L. darwinii group (Table 2) in having posterior teeth with expanded crowns and different body proportions, and in lacking a transversal black line on the eye. It differs from the species of the L. wiegmannii group (Table 2) in having a row of lorilabial scales (never two or three). Liolaemus shehuen differs from L. boulengeri, L. donosobarrosi, L. goetschi, L. hermannunezi (Pincheira-Donoso, Scolaro & Schulte), L. inacayali (Abdala), L. josei (Abdala), L. loboi (Abdala), L. martorii (Abdala), L. rothi (Koslowskyi 1898) L. sagei (Etheridge & Christie), L. senguer (Abdala), L. tehuelche (Abdala), and L. telsen (Cei & Scolaro) in having an evident black antehumeral arch that is absent in the aforementioned species. The species differs from all the species of the L. telsen group, except L. rothi and L. sagei, in having longer snout-vent length (max SVL 90.8 mm vs 32.3–77.1 mm). It differs from L. mapuche (Abdala) in having different dorsal body coloration, lacking light blue scales and having more marked ventral melanism. It differs from L. cuyanus in having more marked ventral melanism and different dorsal coloration. Group Species L. anomalus L. anomalus, L. ditadai, L. lentus and L. pseudoanomalus L. abaucan, L. albiceps, L. calchaquí, L. chacoensis, L. cinereus, L. crepuscularis, L. darwinii, L. L. darwinii diaguita, L. espinozai, L. grosseorum, L. irregularis, L. koslowskyi, L. montanezi, L. laurenti, L. lavillai, L. olongasta, L. ornatus, L. quilmes and L. uspallatensis L. boulengeri, L. inacayali, L. josei, L. loboi, L. martorii, L. puelche, L. rothi, L. sagei, L. senguer, L. telsen L. tehuelche, and L. telsen L. arambarensis, L. azarai, L. cuyumhue, L. lutzae, L. multimaculatus, L. occipitalis, L. rabinoi, L. L. wiegmannii riojanus, L. salinicola, L. scapularis and L. wiegmannii L. goetschi L. cuyanus, L. donosobarrosi, L. goetschi and L. mapuche L. camarones, L. canqueli, L. casamiquelai, L. chehuachekenk, L. fitzingerii, L. melanops, L. L. fitzingerii morenoi L. shehuen and L. xanthoviridis Liolaemus shehuen sp. nov. belongs to the L. fitzingerii clade (Abdala, 2007), which is characterized by its tricuspidate posterior teeth with expanded crowns, neck equal to or wider than head, cylindrical body, short limbs relative to the trunk, black antehumeral arch present and ventrally expanded, and males with ventral melanism. Within the Liolaemus fitzingerii clade, L. shehuen sp.nov. has four to six scales in contact to mental (as the southern populations of L. fitzingerii). In L. shehuen the head melanism characteristic of L. canqueli and L. melanops is absent. Liolaemus shehuen lacks the intense green dorsum with irregular, transverse black paravertebral spots of L. melanops, and the transverse black and yellow-orange bands of L. canqueli. Liolaemus shehuen differs from L. chehuachekenk in lacking a conspicuous vertebral band, having fewer individuals with pre- and postscapular spots, fewer ventral scales (109-121, mean= 115.6 vs. 99-116, mean= 106), and shorter snout-vent length (SVL 90.8 mm vs. 98.3 mm); it differs from L. fitzingerii in having shorter snout-vent length (SVL 90.8 mm vs. 106 mm), and lacking the bluish green or light blue dorsum coloration with red paravertebral spots and red scales speckled, as L. fitzingerii has. Liolaemus shehuen differs from L. morenoi in lacking evident scapular markings and having more ventral scales (109-121, mean= 115.6 vs. 98-113, mean= 107.6). L. morenoi never presents yellow-red dorsal coloration either. Liolaemus shehuen differs from L. xanthoviridis in having different dorsal and ventral coloration, with paravertebral spots and more uniform vertebral region without irregular markings. Description of the holotype. ― Adult male. SVL 90.6 mm. Trunk length 45.1 mm. Head longer (17.4 mm) than wide (14.8 mm). Head height 11.6 mm. Eye diameter 4.3 mm. Interorbital distance 8.7 mm. Orbit–auditory meatus distance 6.6 mm. Auditory meatus 3.1 mm high, 1.6 mm wide. Orbit–commissure of mouth distance 2.5 mm. Internares distance 3.0 mm. Subocular scale length 5.4 mm. Femur length 13.5 mm, tibia 13.7 mm, and foot 25.6 mm. Humerus length 13.1 mm. Tail length 120.2 mm. Dorsal surface of the head smooth, with 15 scales. Rostral wider than long, bordered by six scales. Mental larger than rostral, trapezoidal, bordered by four scales. Nasal not in contact with rostral. Four internasals. Nasal surrounded by eight scales, separated from canthal by two scales. Five scales between frontal and supercilliaries. Seven scales between frontal and rostral. Frontal not divided. Two postrostrals. Interparietal smaller than parietals, in contact with six scales. Orbital semicircles complete. Preocular separated from lorilabial row by one scale. Three scales in anterior margin of auditory meatus. Eleven smooth temporals. Three lorilabials in contact with subocular. Eight supralabials, none in contact with subocular. Eight supraoculars. Eight lorilabials. Five infralabials, second ventrally in contact with two scales. Six chinshields, second pair separated by three scales. Seventy-six scales around midbody. Eighty-six round, slightly imbricate, slightly keeled dorsals from occiput to hind limbs. Thirty-six scale rows in dorsum. Scales of flank same size and shape as dorsals. Ventrals same size as dorsals, flat, imbricate. Thirty-five smooth, weakly imbricate gulars. Eight precloacal pores. Antehumeral scales flat, larger than or equal in size to dorsals. Postauricular, rectal, and longitudinal folds present. Scales on the longitudinal fold granular and smooth. Fourth finger with 19 subdigital lamellae; fourth toe with 26. Infracarpals flat, imbricate, trifid; infratarsals flat, imbricate, not trifid. C olor in life.― Head grey with scattered red spots and scales on the temporal and occipital region. Conspicuous antehumeral arch. Dorsum and flanks yellow-red with faint, small paravertebral spots. Some of these spots on the posterior region have a lighter spot inside. Vertebral region orange, with two light grey stripes on each side of vertebral region. Dorsal fore- and hind limbs same color as trunk. Dorsal tail greenish grey with yellow and orange scales on flanks and at tail base. Throat pale white-yellow. Gular region, chest to cloacal region and ventral hind limbs melanic. Flanks, forelimbs and ventral tail same color as throat. Variation.― (Table 3). Based on 10 specimens. Head longer (14.8–17.9 mm; mean = 16.4 mm) than wide (11.9–14.8 mm; mean = 13.4 mm). Head height 9.8–11.6 mm (mean = 10.7 mm). Neck wider than head. SVL 72.7–90.8 mm (mean = 79.9 mm), averaging 2.1 times longer than trunk. Tail length 94.9–120.3 mm (mean = 109.6 mm). Humerus length 9.7–13.1 mm (mean = 11.4 mm). Dorsal surface of head smooth, with 14–18 scales. One row of lorilabials, same size as supralabials. Six to nine lorilabials. Seven to 10 supralabials. Frontal unique or divided horizontally, into two or four scales. Interparietal always smaller than parietals, surrounded by 6–8 scales. Nasal surrounded by 8–9 scales. Five to seven infralabials. Mental in contact with four or six scales. Ten to 11 smooth round temporals. Longitudinal, postauricular, and antehumeral folds present. Horizontal fold Y-shaped between shoulder and auditory meatus. Scales around midbody 69–80 (mean = 74). Gulars 28–35 (mean = 31.5). Dorsal scales between occiput and hind limbs 63–88 (mean = 77.6). Ventrals 110–119 (mean = 117.3). Males with 8–9 (mean = 8.2) precloacal pores. Females with 0–3 (only one female) (mean = 0.5) precloacal pores. Dorsals flat, slightly imbricate, slightly keeled. Seventeen to 20 infradigital lamellae on fourth finger and 22–29 on fourth toe. Variation in color pattern. ― No sexual dichromatism evident (Figs. 6–7). Head usually grey or light brown with small yellow-orange spots or scales. Black arch ventrally expanded over antehumeral fold. Dorsal trunk and limbs yellow-orange or yellow-red, more intense in some specimens. Faint and small round paravertebral spots, generally dark, but sometimes preceded by a lighter spot. Some specimens have one or two dorsolateral whiteyellow strips and/or vertebral line of same color as paravertebral spots. Some specimens have diffuse white-yellow bars transversal to the body axis. Lateral region of the body same color as dorsum. Tail uniformly bluish grey or greenish grey with scattered spots. Gular region, chest to cloacal region, anterior forelimbs and hind limbs have ventral melanism. Males more melanic than females. In juveniles melanism expands from gular region to cloacal region as the individual grows. Natural history.― Liolaemus shehuen sp. nov. was found in the Telsen plateau and nearby regions, along National Route 4, an area with sand dunes and abundant vegetation, an ecotone between the phytogeographic regions of Monte and Patagonia (Cabrera & Willink 1980). The soil is mostly sandy with basaltic rocks, surrounding a lowland with red soils called ‘Bajo de la tierra colorada’. The vegetation consists mainly of Stipa, Verbena, Mulinum, Nassauvia and Lycium (Cei & Scolaro 1999). The Patagonian plateaus (Telsen, Somuncurá in Río Negro and Chubut provinces, and Buenos Aires Lake, in Santa Cruz Province) are areas with high herpetofauna diversity. Liolaemus shehuen sp. nov. is sympatric with Diplolaemus darwinii (Bell), Homonota darwinii (Boulenger), L. bibronii (Bell), L. petrophilus (Donoso-Barros & Cei), L. rothi, L. telsen, Leiosaurus belli (Duméril & Bibron), Pristidactylus nigroingulus (Cei, Scolaro & Videla), the snakes Philodryas patagoniensis (Girard), Philodryas trilineata (Burmesteir), and the viperid Bothrops ammodytoides (Leybold). Liolaemus shehuen is oviparous and insectivorous. It is active between 12.00 AM and 15:00 PM, when the sand temperature is highest. There is no more information about its biology. Distribution.― Liolaemus shehuen is distributed on the Telsen plateau and east of Río Negro Province, Argentina (Fig. 8). Etymology.― shehuen is a Tehuelche word that means 'Sun', and makes reference to the dorsal coloration of Liolaemus shehuen. The tehuelches, also known as Patagones, are ancient indigenous people that have inhabited Patagonia for more than 10000 years.

Liolaemus casamiquelai sp. nov. (Figure 1, 2) Type material. Holotype: MACN 39042, an adult male collected along the south side of Ruta Nacional 23, 14 km W Aguada de Guerra (41 �� 09��� S, 68 �� 30 ��� W, 866 m, Fig. 5), 25 de Mayo Department, R��o Negro province, Argentina, 27 November 2000, by L. J. Avila, and C. H. F. Perez. Paratypes: MACN 39043, MLP.S 2603, LJAMM-CNP 2986, LJAMM-CNP 4878 adult males; MACN 39044 - 5, MLP.S 2601 - 3, LJAMM-CNP 2977, LJAMM-CNP 2979 - 80 adult females. Same data as the holotype. MLP.S 2600, LJAMM-CNP 2948, adult males; MACN 39041, MLP.S 2599, LJAMM-CNP 2949 adult females. Shrubby margins of Cari Lafquen Lagoon, 7 km N Ingeniero Jacobacci (41 �� 13 ��� S, 69 �� 24 ��� W, 855 m), 25 de Mayo Department, Rio Negro Province, Argentina, 28 November 2000 by L. J. Avila and C. H. F. Perez. LJAMM-CNP 11090 - 92 adult males. Cari Lafquen Chica lagoon and Ruta Provincial 6, 10 km N junction Ruta Nacional 23 (41 �� 13 ��� S, 69 �� 24 ��� W, 834 m), 25 de Mayo Department, R��o Negro province, Argentina, 6 November 2008, by L. J. Avila, and M. Nicola. LJAMM-CNP 11102 adult males; LJAMM-CNP 11101 adult females. Ruta Nacional 23, 22.3 km E Ingeniero Jacobacci (41 �� 18 ��� S, 69 �� 17 ��� W, 860 m), 25 de Mayo Department, R��o Negro province, Argentina, 7 November 2008, by L. J. Avila, and M. Nicola. LJAMM-CNP 11099, 11100 adult females. Ruta Provincial 74, 33.2 km N Chasic�� (road to Aguada Guzman), 33.2 km N junction Ruta Provincial 6 (40 ��02���S, 68 �� 54 ��� W, 876 m), El Cuy Department, R��o Negro province, Argentina, 6 November 2008, by L. J. Avila, and M. Nicola. LJAMM-CNP 2973 - 6 adult males. Ruta Provincial 6, 64 Km NE Ingeniero Jacobacci (40 �� 53 ���, 69 �� 17 ���, 854 m), 25 de Mayo Department, R��o Negro province, Argentina, 29 November 2000, by L. J. Avila, and C. H. F. P��rez. Diagnosis. Liolaemus casamiquelai is a robust and medium sized member of the clade of Liolaemus referred to as the boulengeri group by Etheridge (1995), and is a member of the Liolaemus melanops complex that includes L. canqueli, L. goetschi, L. martorii, L. melanops, L. morenoi, and several other candidate species still in need of detailed morphological study (Morando 2004; Avila et al. 2006 a). Liolaemus casamiquelai can be distinguished from all other species of its group of Liolaemus by a combination of characteristics: its larger size, stout body, intense melanic venter, lack of melanic head, and bright green dorsal coloration with a pattern of transversal irregular black stripes (Figs. 1, 2, 3). Liolaemus casamiquelai differs from L. canqueli and L. melanops in lacking a dark brown or black hood on the head in adults of either sex, and from L. goestchi and L. martorii in lacking large, black pre-and postcapular spots and these species are smaller. Liolaemus casamiquelai differs from L. morenoi in having a green background color with transverse black bands rather than a dorsal background color of light tan with brown spots or wavy with white posterior borders. Description of the holotype. Adult male 92.7 mm (SVL), tail 131.0 mm. Axilla-groin distance 43.7 mm. Head length 17.7 mm; head width 14.9 mm; head depth 11.0 mm. Arm length 26.2 mm; tibial length 19.0 mm; foot length 26.4 mm. Upper head scales smooth, convex, bulged, pitted with scale organs. Rostral pentagonal, twice as wide as high (3.17 x 1.4 mm). Two postrostrals, wider than high, nasal scales separate from rostral. Nasal scales subpentagonal in shape (1.8 x 1.7 mm). Nostrils roughly oval in shape, occupying almost half of the nasal scale. Nasal scales in contact with eight scales on each side. Internasal scales slightly convex. Four internasals, two in contact with postrostral, left almost rectangular, right quadrangular; two separate from postrostral by first internasal, almost oval in shape. Four frontonasals, two medials, hexagonal, in tandem, smaller than laterals. Left lateral frontonasals larger than right lateral scale. Nine prefrontals, regularly arranged, first row with a small medial, two large mediolateral and two smaller lateral; medial and laterals rhomboidal and similar in size, mediolaterals symmetric but roughly rectangular. Second row with two symmetric scales, regular and roughly pentagonal in shape, last row with two symmetric but smaller scales. Four small frontal scales almost similar in size than last prefrontals, in two rows. Eight frontoparietals irregular in shape, larger than frontal and parietal; arranged in three rows. Interparietal pentagonal with a large and conspicuous white cream ���eye��� in the middle, surrounded by five scales. Two large parietals, regularly arranged. Circumorbital circles incomplete: 13���14. Supraoculars 8���9, laterally expanded. Two rows of irregular, small scales between supraoculars and external circumorbitals, 11-15. Supercilliaries 7 - 7, first five strongly overlapped. Nuchal scales small, irregularly rounded, slightly bulged. Two internasals. First canthal higher than wide, separating first and second loreals, contacting lorilabials. Posterior canthal longer than wide. Posterior canthal overlaps only a small part of first supercilliary. Loreals: three irregular in size but symmetrically arranged on each side, two between first canthal and preocular. Canthals, preocular and anterior subocular forming a slight concavity. One preocular, slightly longer than wide; one elongate subocular (5.1 x 0.8 mm), one small postocular; a very evident keel in preocular and subocular, postocular with a less marked keel. Lorilabials convex to flat, 7 - 7, all roughly quadrangular, pitted with conspicuous and numerous scale organs. Upper ciliary scales in two rows, rectangular, compressed, and moderately projecting. Lower and upper ciliaries similar in size and shape. Palpebral scales small, irregular, convex. Supralabials 8 - 8, slighlty convex. Temporal scales rounded, smooth, convex, juxtaposed, with a scale organ in the tip. Auditory meatus higher than wide (3.4 x 1.2 mm) surrounded by granular scales. Mental pentagonal wider than high (2.9 x 2.0); in contact with anterior infralabial and postmental but not in contact with anterior sublabials. Infralabials 6 - 5. Chinshields 5 - 5, transversally expanded, separated from infralabials by series of 2-3 elongated but smaller sublabials. Only a few scale organs present in supralabials and infralabials. Gular scales smooth, flat, imbricate, rhomboidal, immediately after postmental, gradually becoming rounded posteriorly and with a distinct posterior apical notch. Lateral neck folds (longitudinal, oblique, antehumeral and post auricular) well developed. Antegular and gular folds not developed. Dorsal body scales subtriangular, slightly to not imbricated. Eighteen to 24 longitudinal rows with scales with distinct but smooth keels, a few with an apical scale organ. At midbody, dorsal scales laterally smooth, imbricate. Scales anterior to, above, posterior, and close to posterior forelimb and hind limb insertions, small, smooth, slightly overlapping, becoming granular. Ventral body scales smooth, flat, imbricate, with an apical notch in gulars; subtriangular to obovate, same size than dorsal body scales. Scales around midbody 72; scales between occiput an anterior margin of hind-limb articulations 76. Scales of cloacal apron more equal in size than ventral body scales. Preocloacal pores 7. Suprabrachials smooth, rhomboidal, imbricated, with a small notch in the tip. Infrabrachials small, convex, and not overlapping. Supra-antebrachials smooth, imbricate, rhomboidal to obovate, some with a small notch in the tip, others larger than suprabrachials. Infra-antebrachials smooth, rhomboidal, distally becoming with one to three small but distinct spines near the hand insertion. Supracarpals smooth, strongly imbricate. Infracarpals keeled, imbricate. Supradigital lamellae smooth, strongly imbricate. Subdigital lamellae tricarinate, imbricate, numbering: I: 12, II: 20, III: 20, IV: 16, V; 10. Suprafemorals smooth, imbricate, rhomboidal, few with an apical scale organ. Infrafemorals smooth, imbricate, rhomboidal. Supratibials smooth, imbricate, becoming rounded distally. Twenty six to 45 spinose scales in a postfemoral patch. Infratibials smooth, imbricate. Supratarsals, smooth, imbricate, a few with an apical notch. Infratarsals strongly imbricate, with one keel, and one to three small but distinct spines near the digits insertion. Supradigital smooth, imbricate. Infradigital bi- to tri-carinate, numbering I: 11, II: 16, III: 23, IV: 29, V: 18. Ventral caudal scales smooth, a few slightly keeled, imbricated. Dorsal caudal scales keeled, imbricated. Coloration. In life, general background color on neck, body, tail, and limbs, brightly green-yellow (in sunlight) becoming gray/light gray in the shade. Dorsal head surface irregularly brown intermixed with smaller, irregular gray areas, grading to light tan in azygous and nuchal areas. Between occiput and rump, a series of twelve more or less regular, transversal black bands, 2-5 scales wide, more regular in the midbody between limbs. First two marks on the nuchal area irregularly divided and merged, limiting clear spots with background coloration. Markings continue on the lateral sides of the head and neck as irregular well-marked lines 1-2 scales wide, becoming faded and gray on the ventral surface. On the lateral sides of the head, clear areas are larger and clear-white. On the trunk dorsum, transversal marks are broken, or slightly connected along vertebral midline. Some marks between limbs become forked on the lateral regions of the trunk. All marks between limbs extend, usually well defined, to the ventral scales. Between legs and first third of the tail, transverse marks merge and become more regular. Along the tail transverse marks merge longitudinally and become almost indistinguishable in the last third of the tail. Background coloration becomes light-gray on distal two-thirds of the tail. Lower lateral sides of the neck, trunk, and first third of the tail orange-yellow when lizard is in the shade without bright coloration. Otherwise, no lateral color differs from the general background. On dorsal limb areas an irregular pattern of black lines, continuous from the dorsum, delimit rounded or irregular, clear spots with a light gray background. Ventral surfaces of the gular, chest, belly, interfemoral, femoral and apron areas black. Ventral surface of the limbs and tail white with some gray areas. All bright coloration disappears after capture, and after several years in preservative, all coloration became faded. Variation. Based on the paratypes (Table 2, Fig. 4): in 15 males: SVL 78.8���106.2 mm. Axilla groin distance 37.8���54.5 mm. Foot length 15.6���27.1 mm. Tibial length 16.2���19.6 mm. Arm length 22���28.5 mm. Head length 15.1���18.8 mm. Head width 13.1���16.2 mm. Head depth 9.5���13.8 mm. Midbody scales 64���78. Dorsal scales between occiput to rump 66���87. Ventral scales 92���109. Supralabials 6���9. Infralabials 5���7. Fourth toe lamellae 27���32. Precloacal pores 7���11. In 15 females: SVL 74.1���103.5 mm. Axilla-groin distance 38.3���56.4 mm. Foot length 21.8 ���27.0 mm. Tibial length 15.3���18.2 mm. Arm length 22.1���26.6 mm. Head length 13.9���18.2 mm. Head width 12 ���15.0 mm. Head depth 9.1���11.6 mm. Midbody scales 66���76. Dorsal scales between occiput to rump 75���84. Ventral scales 96���115. Supralabials 5���9. Infralabials 5���7. Fourth toe lamellae 25���33. Dorsal head scales variable in size, some irregular in shape, most convex but a few almost flat. Cloacal apron larger in males that in females. Scales of the cloacal apron slightly smaller that ventral scales in males, same size or slightly smaller in females. Postfemoral patch not present in females and variable in size in males between 18���22 scales. Males (N= 15) Females (N= 15) In some individuals melanic coloration is more extensive than the holotype. In a few individuals dorsal transversal marks are connected longitudinally, becoming lines of clear dots on the dorsum. In some individuals melanic coloration extends on lateral sides of the neck and head, leaving only a few clear scales; others have the same areas with a very reticulated pattern of anastomosed gray lines. All individuals show a bright yellow-green coloration in full sun, which immediately fades in the shade and disappears completely when lizards are kept in captivity. Dorsal background color bright green in recently captured adult males and females, few individuals (20-30 %) have a yellowish-green sheen. Individuals from locality Ruta Nacional 23 10 km E Ingeniero Jacobacci have a grey background coloration on the head and tail. Etymology. To honor Rodolfo Mag��n Casamiquela, a recently deceased vertebrate paleontologist and anthropologist, born in Ingeniero Jacobacci (formerly Huahuel Niyeu) in 1932, a small town very close to L. casamiquelai type locality. R. M. Casamiquela was a researcher (retired) of Consejo Nacional de Investigaciones Cient��ficas y T��cnicas of Argentina (CONICET), and developed a large and prolific scientific and cultural literature ranging from amphibians fossils to historic tales of Patagonia. Geographic distribution. Liolaemus casamiquelai is known from five localities (Fig. 6) in western Rio Negro province: (1) the type locality, (2) in the eastern edge of Cari Lafquen Chica Lagoon, (3) along the National Road 23 eastern of Ingeniero Jacobacci town, (4) 64 km north of Ingeniero Jacobacci along the Provincial Road 6, and (5) south of Aguada Guzman town, along the Provincial Road 74. Natural history. The holotype and the paratypes were found basking on the edges or walking inside of clumps of spinose shrubs. All localities except (5) are in Meseta Central ecoregion (Bran et al. 2000), the most arid region in R��o Negro province, with an annual rain lower than 200 mm and temperatures between 10-12 �� C. This ecoregion is characterized by the presence of lower plateaus, small hills, and peneplains covered by lower-shrub steppe vegetation with dominant plants as Nassauvia axillaris, N. glomerulosa, and Chuquiraga avellanedae or shrub-grass steppes with clumps of Prosopis denudans and Lycium sp, surrounded by grassy areas with several species of grass, more common is Stipa humilis. The fifth locality is an ecotonal area between southern Monte and the Meseta Central ecoregion, with the same vegetation plus some individuals of Larrea spp. and clumps shrubs of Schinus johnstonni. According to Cabrera (1994) phytogeographic classification, all the localities are included in the Dominio Andino Patag��nico, Provincia Patag��nica. All of the sites where L. casamiquelai were collected are similar in physiography, normally flat, with sandy soil mixed with gravel areas or more compacted soil. Clumps of shrub appear in areas with some degree of protection from the strong winds typical of Patagonia; these clumps are composed mainly of Prosopis denudans (or Schinus johnstonii in locality 5) and smaller shrubs of Atriplex sp., Lycium chiliensis, and Chuquiraga avellanedae, all of which grow under the protection of this spiny mesquite. These clumps reach a maximum height of 1.8 m, and cover an area between two to four m 2 and separated from each other by distances of two or three meters. This region is characterized by highly degraded soils (due to overgrazing), and shows a tendency towards desertification. Clumps seem to be the safest areas for lizards and rodents, as well for some birds. The majority of the lizards were observed only inside or in the immediate vicinity of the spiny shrub clumps. We never observed a lizard in small shrubs or between grasses surrounding this clumps. All lizards observed during several field trips were seen in the immediate edges or inside the clumps. Usually lizards bask under sun in early morning (09:00 to 10:00) and in late afternoon (18:00 to 19:00) in the midsummer, when they show a bimodal type of activity, but in early spring and late-summer they usually show continuous activity without a mid-day break. In the type locality we collected other species of Liolaemus living in syntopy with L. casamiquelai, including L. inacayali, L. bibronii, and L. aff. melanops, as well as two leiosaurid species Leiosaurus bellii and Pristidactylus nigroiugulus Avila et al. (2003). In other localities we collected Liolaemus elongatus, L. petrophilus, Phymaturus aff. ceii, and Homonota darwinii usually in nearby rocky outcrops but not sharing the same microhabitat than L. casamiquelai (Acosta et al. 1996 a, b; Avila et al. 2003; Avila et al. 2004; Avila et al. 2006 a, b). In nearby localities we collected Bothrops ammodytoides and Philodryas patagoniensis (Avila et al. 2001), both are known to eat lizards and are probable predators of L. casamiquelai. Some other likely predators, such as the hawks Falco sparverius and Buteo polyosoma, are common in this area. Liolaemus casamiquelai is probably oviparous as well others members of its clade, and feeds mainly in arthropods and some plant matter as a dissected stomach content show. Remarks. Phylogenetic analysis based on the mitochondrial gene fragment cyt-b (805 bp), using Maximum Parsimony and Bayesian approaches of all described species of the fitzingerii group (melanops + fitzingerii complexes) and two outgroups (L. boulengeri and L. cuyanus) recovered the tree shown in Fig. 7. This topology presents evidence to support the monophyly of the melanops complex with L. casamiquelai as the sister taxon of L. martorii (MP Bootstrap Support [MPB]: 90; Posterior Probability [PP]: 0.51) and closely related to L. goestchi. Liolaemus morenoi is the basal species of this clade we call melanops N (MPB: 83, PP: 0.99). Liolamus canqueli is the sister taxon of L. melanops, clade we refer to as melanops S (MPB: 96, PP: 0.94). The uncorrected pairwise distances within species of the melanops N clade range from 0.62 % (L. goestchi vs. L. casamiquelai) to 2.73 % (L. morenoi vs. L.martorii); the distance between L. canqueli vs. L. melanops is 1.74 %; and species distances between melanops N and S clades range from 3.11 % to 3.85 %. There is evidence of incomplete lineage sorting for this marker within and between these clades (Morando et al., in prep.), and species from this complex (as in other Liolaemus complexes) exhibit strong variation in background and pattern coloration that difficult an easy recognition of species limits without a proper sampling effort and an adequate knowledge of contact zones. A poor morphological knowledge of large series, lack of extensive sampling, absence of revision of type specimens, fail to collect samples in the right type locality made difficult an adequate study of the species of the complex and originated a complicated nomenclatural status for some of them. But a discussion of t, Published as part of Avila, Luciano Javier, Perez, Cristian Hernan Fulvio, Morando, Mariana & Sites, Jack Walter, 2010, A new species of Liolaemus (Reptilia: Squamata) from southwestern Rio Negro province, northern Patagonia, Argentina, pp. 47-59 in Zootaxa 2434 on pages 49-56, DOI: 10.5281/zenodo.194791, {"references":["Etheridge, R. (1995) Redescription of Ctenoblepharys adspersa Tschudi, 1845, and the taxonomy of Liolaeminae (Reptilia: Squamata: Tropiduridae). American Museum Novitates, 3142, 1 - 34.","Avila, L. J., Morando, M. & Sites, Jr., J. W. (2006 a) Inferring evolutionary processes in \" inertial species \" - an example in Patagonian lizards of the Liolaemus fitzingerii complex (Squamata: Liolaemidae). Biological Journal of Linnean Society, 89, 241 - 275.","Bran, D., Ayesa, J. & Lopez, C. (2000) Regiones ecologicas de Rio Negro. Comunicacion Tecnica Nº 59. INTA (Instituto Nacional de Tecnologia Agropecuaria), Bariloche, pp. 1 - 8. Available from http: // www. inta. gov. ar / bariloche / info / catalog / Teledeteccion / AreasEcologicas % 20 RN. pdf (accessed 13 May 2009.","Cabrera, A. L. (1994) Regiones Fitogeograficas Argentinas. In: Kluger, W. F. (Ed.), Enciclopedia Argentina de Agricultura y Jardineria. Editorial ACME S. A. C. I., Buenos Aires, pp. 1 - 85.","Acosta, J. C., Blanco, G. M. & Avila, L. J. (1996 a) Organizacion espacial de una comunidad de saurios del Sur de la Provincia de Rio Negro - Argentina. FA C E N A, 12, 145 - 149.","Acosta, J. C., Avila, L. J. & Blanco, G. M. (1996 b) Ecologia de Liolaemus boulengeri (Sauria: Tropiduridae) en el Noroeste de la meseta patagonica (Ing. Jacobacci, Rio Negro, Argentina). Cuadernos de Herpetologia, 9, 40 - 47.","Avila, L. J., Frutos, N., Morando, M., Perez, C. H. F. & Kozykariski, M. (2006 b) Reptilia, Liolaemini, Liolaemus petrophilus & Liolaemus pictus: Filling distribution gaps, geographic distribution. Check List, 2, 66 - 69.","Avila, L. J., Morando, M. & Perez, D. R. (2001) New records and natural history notes for lizards and snakes from Patagonia, Argentina. Herpetological Review, 32, 64 - 65.","Cei, J. M. & Scolaro, J. A. (2003) Rectificacion taxonomica y nomenclatural del presente status del taxon Liolaemus melanops Burmeister 1888. FA CE N A, 19, 163 - 164.","Abdala, C. S. (2003) Cuatro nuevas especies del genero Liolaemus (Iguania: Liolaemida), pertenecientes al grupo boulengeri, de la Patagonia, Argentina. Cuadernos de Herpetologia, 17, 3 - 32."]}

Liolaemus cuyumhue sp. nov. Figure 1. Type material. Holotype: MACN 38981 (Fig. 1), an adult male from sand dunes near Ruta Provincial 7, 28.7 km NW A��elo, A��elo Basin, A��elo Department, Neuqu��n Province, Argentina (38 �� 11 ��� S, 69 �� 01��� W, 259 m), collected 2 February 2003 by L. J. Avila, M. Morando, C. H. F. Perez, and K. Dittmar. Paratypes (Fig. 2): FML 17592, LJAMM 4520 ��� 1 (males), from sand dunes, southern edge of Ruta Provincial 7, (38 �� 13 ' S, 68 �� 57 ' W, 260 m), A��elo Department, Neuqu��n Province; 15 November 2003; collected by D. R. Perez, J. Perez, M. Perez, and M. Perez Carri��; FML 17594, MACN 38982 ��� 3, MLP.S 2587, 2591 (males), FML 17593, LJAMM 3692, MACN 38984, MLP.S 2586, 2588 / 9���90 (females) from sand dunes near Ruta Provincial 7, (38 �� 13 ' S, 68 �� 57 ' W, 258 m), A��elo Department, Neuqu��n Province; 10 February 2006; collected by D. R. Perez, C. de la Vega and D. Zu��iga. Diagnosis. Liolaemus cuyumhue is a member of the wiegmannii group of Liolaemus lizards, characterized by the presence of two or more rows of lorilabials scales rather than one between subocular and supralabilals, smaller in size than in other Liolaemus; flat or concave rather than convex infralabials, mental scale narrower anteriorly than posteriorly; and with the exception of L. cuyanus and L. mapuche, six scales in contact with the mental (Etheridge, 1995, 2000; Abdala, 2002). Liolaemus cuyumhue is distinguished from L. riojanus because its background coloration is yellow-cream to a light red-brick tonality rather than orange-brick or light ochre; dorsal spots are larger than in L. riojanus, and usually have few blue iridescent scales distributed on the dorsolateral areas of the body, rather than mostly grouped into small clumps of 2���4 scales on the sides of the body. L. cuyumhue never have suprascapular spots and series of dorsolateral yellow spots as is observed in some populations of L. riojanus. Another difference from L. riojanus, females of L. cuyumhue always lack precloacal pores. L. cuyumhue males are smaller than L. multimaculatus males (61.7 mm vs 72 mm SVL), with smaller dorsal scales, and fewer scales around midbody (62���71 vs 68���92). L. cuyumhue lack the dark brown/black scapular spots always present in L. multimaculatus. L. cuyumhue have smaller circular brown spots, speckles of smaller white dots, and a low density of blue iridescent scales distributed on the dorsolateral areas of the body instead of rhomboidal to circular gray, tan, or brown spots, speckles of white scales, and blue iridescent scales scattered irregularly on a gray or brown background, observed in L. multimaculatus. L. cuyumhue never have the general coloration of grey and blue scales observed in L. rabinoi, and never have iridescent blue scales clumped into small patches of 3���11 scales each on the sides of the body. Liolaemus cuyumhue lack the sexual dichromatism observed in L. wiegmannii, have smooth or slightly keeled dorsal head scales instead of rugose and protruded, dorsal narines instead of lateral, more supralabial (8���11 vs 4���6) and infralabial scales (7���10 vs 5���7), and smaller body scales (midbody scale counts 64���71 vs 38���60, dorsal body scale counts 78���90 vs 42���62). L. wiegmannii females sometimes have precloacal pores (0���6) but these are completely absent in L. cuyumhue females. Dorsal scales in L. cuyumhue are smooth to slightly keeled and juxtaposed to slightly imbricated instead of strongly keeled and imbricated as in L. wiegmannii. L. cuyumhue males lack dorsolateral and lateral dark spots, paravertebral lines, numerous and larger blue scales in lateral areas, and bright yellow or orange scales, all characteristics of L. wiegmannii. L. cuyumhue always lack pre and postscapular spots or paravertebral lines observed in L. azarai, body size is larger (maximum SVL in males 61.7 vs 54.3 mm; in females 59.9 vs 48.7 mm), and have smaller body scales (midbody scale counts 64���71 vs 32���42, dorsal body scales 78���90 vs 42���60). L. cuyumhue males have higher number of precloacal pores (7���9 vs 5���6) but in females these always are absent (2���3 in L. azarai females). L. cuyumhue is slightly larger than L. arambarensis (maximum SVL 60 vs 56 mm in males, 54.3 vs 48.7 mm in females), have smaller dorsal scales, with more scales around midbody (64���71 vs 60���66), and along the trunk (78���90 vs 57��� 64). L. cuyumhue lack a mid-dorsal white line and two dorsolateral stripes, with two series of paravertebral brown marks resembling triangles bordered by a white bar, as observed in L. arambarensis. L. cuyumhue is considerably smaller than L. lutzae (males 61.7 mm vs 84.0 mm, females 59.9 mm vs 69.0 mm), dorsal coloration lacks the conspicuous wide vertebral band bordered with dark paravertebral spots, and wide gray dorsolateral stripes observed in L. lutzae. L. cuyumhue is smaller (males 61.7 mm vs 70.0 mm, females 59.9 mm vs 60.0 mm), lacks strongly keeled and imbricate dorsal scales, lacks a dark longitudinal bar above the forelimb insertion, and lacks the ventral sexual dimorphism observed in L. occipitalis. L. cuyumhue is smaller than L. salinicola and L. scapularis (61.7 vs 76 vs 77 mm in males, 59.9 vs 68 vs 65 mm in females) and lacks stripes and gray throat observed in L. salinicola and the scapular mark characteristic of L. scapularis. Description of the holotype. Adult male. SVL 51.2 mm, total length 111.2 mm. Axilla groin distance 23.4 mm. Head width 10.8 mm, head length 12.9 mm, head height 6.6 mm, snout length 4.8 mm, horizontal diameter of orbit 2.5 mm, internarinal distance 3.3 mm; eye-nostril distance 1.9. Arm length 15.1 mm, tibial length 11.5 mm, foot length 15 mm (all from the right side). Dorsal head scales smooth, flat to slightly concave, a few pitted with scale organs. Eighteen dorsal head scales (from a line drawn horizontally between anterior margin of external auditory meatus to anterior margin of rostral). Rostral pentagonal, wider that high (1.9 x 0.8 mm). Two postrostrals, wider than high; together with anterior lorilabials separate nasal scales from rostral. Nasal scales longer than wide (1.4 x 0.9 mm), dorsolateral in position. Nostril almost rounded in shape, occupying slightly less than half of nasal scale. Nasal scales in contact with nine scales (left) and eight scales (right). Internasal scales in two rows; two anterior, small, and pentagonal preinternasals; three posterior post-internasals, longer than wide, medial slightly larger than laterals. Two small postnasals behind each nasal. Ten frontonasals, irregular in shape, in two irregular rows. Nine prefrontals in two rows, five scales in front, four scales in back. Two scales conspicuously larger in each row, lateral in position in the first row, medial in the second row. Three frontal scales, median longer than wide, almost equal in size to prefrontals. Nine frontoparietals, irregular. Interparietal with a conspicuous parietal eye, oblong, surrounded by eight smaller irregular scales. Parietals smooth and irregular, variable in size. Eighteen circumorbitals on left side, twenty two on right side. Supraoculars small, irregular, numerous, 59 left, 46 right, five conspicuously larger, wider than high on each side. First canthal higher than wide, posterior canthal longer than wide, non overlapping first superciliary. A well marked but blunt canthal ridge. Six superciliaries on each side, overlapped. Loreals 9 ��� 8; flat to slightly convex. Loreals, preocular, and lorilabials forming a conspicuous concavity. Lorilabials in two-three rows, slightly bulged, becoming smaller below subocular, none wider than supralabials. Upper ciliary scales in two rows, those of inner row flat and quadrangular; those of outer row rectangular and conspicuously projecting. Lower ciliaries more similar in size and shape than upper ciliaries, some conspicuously projecting. Palpebral scale small, irregular, flat. Preocular small, quadrangular, with a keel. Subocular elongated (4.4 x 0.4 mm), with a distinct and sharp keel. Supralabials 8 - 8, last three on right side, distinct, two times longer than high. Temporal scales roughly quadrangular, higher than large. Supratemporals smaller than temporals, roughly rounded, a few almost granular. Nuchals slightly keeled, juxtaposed, rounded. Lateral nuchals small, almost granular, non-imbricated. Occipital scales small, irregular, flat, becoming granular distally. Auditory meatus oval, higher than wide (1.9 x 1.1 mm), surrounded above, behind, and below by granular scales. Pretympanic scales, rounded to obovate, non-projecting. Mental heptagonal (1.6 x 1.2 mm), wider than high, in contact with six scales: infralabials, postmentals, and sublabials. Infralabials flat 7���8, meeting sublabials at an acute angle. Chinshields slightly evident, two postmentals evident but following scales becaming similar to sublabials and gulars in the third row. First sublabials and sublabials in contact with infralabials, smaller than surrounding scales. Gulars imbricate, smooth, becoming rounded and notched distally. Longitudinal oblique and antegular folds distinct, gular and antehumeral well marked. Dorsal body scales obovate, subimbricated, with a blunt keel. Dorsal limb scales 2��� 3 times larger than dorsal body scales, rhomboidal to obovate, imbricate, and bluntly keeled. Suprabrachials nearly two times larger than suprantebrachials, with a more regular rhomboidal shape and keel. Supraantebrachials becoming obovate near the insertion, a few with a small spine. Supracarpals and supradigitals smooth. Infrabrachials small, almost granular. Infraantebrachials 2���4 times larger than infrabrachials, nongranular. Infrabrachials oval to rounded, Juxtaposed near the infrabrachials, becoming progressively imbricated and obovate, some with a blunt keel, near the hand. Infracarpals blunt to well marked keel, imbricate, none tridentate. Subdigital lamellae of manus with three keels, numbering I: 9, II: 13, III: 20; IV: 20; V: 12. Claws long and slender, slightly curved. Supracarpals smooth or with a blunt keel, supradigitals smooth. Infrafemorals and infratibials smooth, imbricated. Infratarsal keeled, strongly imbricated, none tridentate. Infradigitals unidentate, numbering: I: 10; II: 14, II: 18, IV: 24, V: 14. Long and slender claws, slightly curved. Ventral scales lanceolate to rhomboidal, smooth, imbricated. Some scales with an apical notch. Ventral scales slightly larger than dorsal scales. Scales in the flanks between dorsal and ventral distinctly smaller. Scales in front, above and behind limb articulations small, granular to flat. Caudals imbricate, with a blunt but conspicuous keel, same size to the dorsal and ventrals, becoming smaller to the tip of the tail. Ventral caudals with a blunt keel. Scales of the cloacal apron slightly smaller that the ventral scales. Seven insconspicuous precloacal pores. Coloration. Dorsal pattern on head, trunk, tail and limbs with brown rounded spots (4 to 9 scales in size) and speckles of cream, gray, tan, and reddish-brown on reddish background. Between 9 to 10 irregular transversal series of 5 to 7 spots between occiput and rump. Background coloration varies according to ambient light between a yellow-cream (full sun) to a red-brick tonality (in shade). Reddish-brown spots larger than other colored spots, sometimes white bordered anteriorly, posteriorly or both. Some larger creamy spots on dorsolateral areas of trunk. Head with brown spots smaller than in other areas of body; tail brown spots faintly larger than in limbs, head and trunk. Some clear lines faintly indicated on ciliar, supraocular, and longitudinal neck folds. Lateral areas of the body with larger brown spots on white background. Ventral areas immaculate cream-colored except in throat and lateral areas of chest and belly marked with brightly light brown to reddish-brown spots irregularly distributed. Some small, scattered blue iridescent scales are distributed on the dorsolateral areas of the body. In preservative same general coloration but faded, bright ventral spots become light gray, blue iridescent scales disappear. Variation. Morphometric and scale variation is presented in Table 1 and 2 (with all other diagnostic features mentioned above), and dorsal and ventral color variation is depicted in Figure 2. As in other members of the wiegmannii group (except L. wiegmannii), intrapopulation variation in morphology is not conspicuous. Postfemoral patch not present in females and variable in size in males, between 18���25 scales. Dorsal background color usually similar to holotype, recently captured adult males have a brighter coloration. Variation most frequently seen is in the distribution and size of dorsal and ventral spots. Maximum SAM DS PP Scapular Dorsal sexual Iridescen Source SVL marks dichromatism t blue scales F M F M Sexual dimorphism. Females lack of the colored ventral spots characteristic of the males and the iridescent blue scales. The base of tail of males is expanded laterally, and cloacal opening is not rounded as is seen in females. Female lacks of the larger and obvious precloacal pores. Etymology. Named in reference to the characteristics of the substrate of the type locality; cuyumhue is a Mapuche word meaning sandy place. Geographic distribution. Liolaemus cuyumhue is known only from the type locality were the holotype and paratypes were collected, in the A��elo basin close to the Provincial Road 7, in Neuqu��n Province (Figs. 3, 4). Sand dune formations with suitable habitats for L. cuyumhue are restricted to the southern edges of the A��elo basin, but additional field work is necessary to explore other apparently suitable habitats north of this area. Liolaemus cuyumhue lives in marked isolation from other members of the L. wiegmannii group, the closest locality for L. multimaculatus is ~ 580 km SE (Vega and Bellagamba 1994); L. wiegmannii is found more than 300 km NE (Tiranti and Avila 1997), L. rabinoi was described for sandy areas around the Nihuil Dam in Mendoza province, more than 400 km N (Cei 1974, Etheridge 2000), and L. riojanus inhabit in dunes at least 700 km N (Cei 1979, Etheridge 2000). Field surveys carried out from 2000 to 2008 in dunes systems around the A��elo basin revealed other Liolaemus species different from any related to the wiegmanni group. Natural history. Liolaemus cuyumhue is known from an isolated sand dune system in the region known as Bajo de A��elo, in eastern Neuqu��n. This is a depression between the Auca Mahuida Plateau and the Colorado and Neuqu��n rivers, with its lowest point reaching 223 m. The geological origin of the A��elo depression is attributed to karst activity, but sand dunes are probably remnants of old sea shore dunes (Uliana and Dellape 1981). The dunes are sparsely covered by clumps of Sporobolus rigens, Neosparton darwinii, Larrea divaricata, Prosopis flexuosa var depressa, and Atriplex zampa. Liolaemus cuyumhue is syntopic with other species of Liolaemus, including L. grosseorum, L. mapuche, L. gracilis, and in surrounding outcrops L. austromendocinus; other lizard species such as Homonota fasciata, H. darwinii, Cnemidophorus longicaudus, Leiosaurus belli, and Pristidactylus cf fasciatus are also found in sympatry. The holotype was collected at mid afternoon on a cloudy day and was the only specimen of this species observe that day, while individuals of other species were normally active. Paratypes were collected late in the afternoon after the substrate temperature was cooler. We did not observed bimodal activity but probably this type of activity is common in summer on very hot days. L. cuyumhue is insectivorous and presumably oviparous, as are other members of the wiegmannii group. Individuals of L. cuyumhue were observed only on bare or sparsely vegetated dunes with extensive areas of open sand. They appear not to extend out into the more vegetated sandy flats or rocky areas that usually border the dunes. Field observations of L. cuyumhue were made in February and November 2003, February 2006, and January 2007. When first observed, most individuals were close to clumps of vegetation with a few observed basking on open areas between the clumps. Lizard observation was very difficult because their coloration usually matches soil color, and usually we detected them only when they were running from our approach. Their first escape behavior was to flee into the clumps and remain motionless on the surface. If pursuit continued, they would move to another location and again remain immobile on the surface. Only if pursued multiple times would lizards bury into the sand. We did not ever observe such a burial, but we presume that this behavior is likely very similar to the description made for L. multimaculatus (a closely related species) by Halloy et al. (1998). They usually ���disappear��� in a clump of vegetation with no evidence of a burrow entrance. Liolaemus cuyumhue lives in a region were the oil and gas companies conduct some of the more intensive operations in Argentina; new rigs, tracks and roads are opened regularly; frequently modifying some areas with suitable habitats for L. cuyumhue that apparently do not cover large areas in the Bajo de A��elo. How the oil activity will affect this species must await further study. The entire wiegmannii group is threatened by human activities, urban and tourist development are rapidly fragmenting coastal sand dunes in Buenos Aires province, the habitat of L. multimaculatus (Vega et al. 2000); tourist activities also destroy the habitat of L. lutzae (Rocha and Bergallo 1992) and L. occipitalis (Di Bernardo et al. 2000); industrial agriculture (vineyards, olive groves), road construction and deforestation have destroyed extensive areas of habitat for L. salinicola and L. scapularis (Avila, unpublished data); a large dam on the Rio Paran�� probably destroyed large areas inhabited by L. azarai (Avila 2003); and L. rabinoi is considered by some authors as extinct, probably by habitat degradation as a result of human alteration after construction of El Nihuil Dam (Cei 1986, Etheridge 2000). We hope that this destiny will not be the future for our new species. Phylogenetic affinities. Liolaemus cuyumhue is the only species of the wiegmannii group recorded from Neuqu��n province. The analyses of the separate gene partitions (Cyt-b, ND 4, 12 S and C-mos) showed some incongruences, but the majority were between the outgroups, and in most cases these relationships were not resolved and the outgroups formed a polytomy basal to the wiegmannii group. Most of the relationships among the ingroup taxa were consistent with the different partitions, Published as part of Avila, Luciano Javier, Morando, Mariana, Perez, Daniel Roberto & Sites, Jack W., 2009, A new species of Liolaemus from A��elo sand dunes, northern Patagonia, Neuqu��n, Argentina, and molecular phylogenetic relationships of the Liolaemus wiegmannii species group (Squamata, Iguania, Liolaemini), pp. 39-55 in Zootaxa 2234 on pages 41-49, DOI: 10.5281/zenodo.190368, {"references":["Etheridge, R. (1995) Redescription of Ctenoblepharys adspersa Tschudi, 1845, and the taxonomy of Liolaeminae (Reptilia: Squamata: Tropiduridae). American Museum Novitates, 3142, 1 - 34.","Etheridge, R. (2000) A review of lizards of the Liolaemus wiegmannii group (Squamata, Iguania, Tropiduridae), and a history of morphological change in the sand-dwelling species. Herpetological Monographs, 14, 293 - 352.","Abdala, C. S. (2002) Nuevo Liolaemus (Iguania: Liolaemidae) perteneciente al grupo boulengeri de la provincia de Neuquen, Argentina. Cuadernos de Herpetologia, 16, 3 - 13.","Tiranti, S. I. & Avila L. J. (1997) Reptiles of La Pampa Province, Argentina: an annotated checklist. Bulletin of Maryland Herpetological Society, 33, 97 - 117.","Cei, J. M. (1974) Two new species of Ctenoblepharis (Reptilia, Iguanidae) from the arid environments of the central Argentina (Mendoza province). Journal of Herpetology, 8, 71 - 75.","Cei, J. M. (1979) A reassessment of the genus Ctenoblepharis (Reptilia, Sauria, Iguanidae) with description of a new subspecies of Liolaemus multimaculatus from western Argentina. Journal of Herpetology, 13, 297 - 232. Cei, J. M. (1986) Reptiles del centro, centro-oeste y sur de la Argentina. Herpetofauna de las zonas aridas y semiaridas. Museo Regionale di Scienze Naturali Torino, Monografie 4, 527 pp.","Uliana, M. A. & Dellape D. A. (1981) Estratigrafia y evolucion paleoambiental de la sucesion maestrichtiana-eoterciaria del engolfamiento neuquino (Patagonia septentrional). Actas VIII Congreso Geologico Argentino, 3, 673 - 711.","Halloy, M., Etheridge, R. & Burghardt G. M. (1998) To bury in sand: phylogenetic relationships among lizard species of the boulengeri group, Liolaemus (Reptilia: Squamata: Tropiduridae), based on behavioral characters. Herpetological Monographs, 12, 1 - 37.","Vega, L. E., Bellagamba, P. J., & Fitzgerald, L. A. (2000) Long term effects of anthropogenic habitat disturbance on a lizard assemblage inhabiting coastal dunes in Argentina. Canadian Journal of Zoology, 78, 1653 - 1660.","Rocha, C. F. & Bergallo H. (1992) Population decrease: the case of Liolaemus lutzae, an endemic lizard of Southeastern Brazil. Ciencia e Cultura, Sao Paulo, Brazil (Journal of the Brazilian Association for the Advancement of Science), 48, 52 - 54.","Di Bernardo, M., Martins, M. B. & Oliveira, R. B. (2000) Liolaemus occipitalis. In: IUCN 2008. 2008 IUCN Red List of Threatened Species. Available from www. iucnredlist. org (accessed 13 March 2009).","Schulte, J. A., Macey, J. R., Espinoza, R. E. & Larson, A. (2000) Phylogenetic relationships in the iguanid lizard genus Liolaemus: multiple origins of viviparous reproduction and evidence for recurring Andean vicariances and dispersal. Biological Journal of the Linnean Society, 69, 75 - 102.","Cruz, F. B., Fitzgerald, L. A., Espinoza, R. E. & Schulte, J. A. II (2005) The importance of phylogenetic scale in tests of Bergmann's and Rapoport's rules: lessons from a clade of South American lizards. Journal of Evolutionary Biology, 18, 1559 - 1574.","Verrastro, L., Veronese, L., Bujes, C. & Dias Filho, N. M. (2003) A new species of Liolaemus from Southern Brazil (Iguania: Tropiduridae). Herpetologica, 59, 105 - 118."]}

Liolaemus martorii Abdala Liolaemus boulengeri Vega & Bellagamba 1994: 142. Liolaemus martorii Abdala 2003: 15 (type locality: Las Grutas, Western San Antonio Department, Río Negro Province, Argentina), 2007: 44. Observations: The taxonomic status of this species should be reevaluated on the basis of comparative evidence between its type series and specimens of the Patagonian species Liolaemus goetschi. When Abdala (2003) proposed L. martorii as a new species, the taxon L. goetschi was still considered a synonym of L. melanops (e.g. Cei 1986). Nevertheless, Cei & Scolaro (2003) suggested that L. goetschi and L. melanops represent two different species. This might explain why Abdala (2003) did not provide comparisons between the populations identified as L. goetschi and L. martorii. Morphological and field observations between these two taxa carried out by one of us (JAS) suggest that they may be conspecific. Distribution, morphology and coloration exhibit strong overlap between L. goetschi and L. martorii. The problem should be reassessed using appropriate statistical analyses., Published as part of Pincheira-Donoso, Daniel, Scolaro, J. Alejandro & Sura, Piotr, 2008, A monographic catalogue on the systematics and phylogeny of the South American iguanian lizard family Liolaemidae (Squamata, Iguania), pp. 1-85 in Zootaxa 1800 on page 51, DOI: 10.5281/zenodo.6789337, {"references":["Vega, L. E. & Bellagamba, P. J. (1994) Reptiles de la Reserva de Usos Multiples Caleta de los Loros, Rio Negro, Argentina. Cuadernos de Herpetologia, 8, 141 - 145.","Abdala, C. S. (2003) Cuatro nuevas especies del genero Liolaemus (Iguania: Liolaemidae), pertenecientes al grupo boulengeri, de la Patagonia, Argentina. Cuadernos de Herpetologia, 17, 3 - 32.","Cei, J. M. (1986) Reptiles del centro, centro-oeste y sur de la Argentina. Herpetofauna de las zonas aridas y semiaridas. Museo Regionale di Scienze Naturali di Torino, Monografie 4, 528 pp.","Scolaro, J. A. & Cei, J. M. (2003) Una excepcional nueva especie de Phymaturus de la precordillera de Chubut, Argentina (Liolaemidae, Iguania, Lacertilia, Reptilia). Facena, 19, 107 - 112."]}

Liolaemus alticolor Barbour Liolaemus lativittatus Werner 1904: 8 (?) (probably conspecific fide Müller & Hellmich 1938a; Donoso-Barros 1966a; see also Etheridge & Espinoza 2000). Liolaemus alticolor Barbour 1909: 51 (type locality: Tiahuanacu, Bolivia); Burt & Burt 1931: 273; Laurent 1992: 30; Núñez & Jaksic 1992: 69; Etheridge 1995: 33; Carrillo de Espinoza & Icochea 1995: 9; Etheridge & Espinoza 2000: 4; Escobar et al. 2001: 1679; Martínez-Oliver & Lobo 2002: 53; Lobo & Espinoza 2004: 852; Chebez et al. 2005: 28; Sura 2005: 428; Díaz & Lobo 2006: 264. Liolaemus altcolor alticolor Hellmich 1961: 2; Donoso-Barros 1964: 7, 1966a: 198, 1970a: 178, 1970b: 88; Vanzolini 1986: 12; Veloso & Navarro 1988: 499; Baudoin et al. 1990: 429. Liolaemus (Liolaemus) alticolor Pincheira-Donoso & Núñez 2005: 193. Observations: Diverse aspects of the systematics, geographical distribution and reproductive biology of this species and its related taxa have largely been debated. See Lobo & Espinoza (1999, 2004), Pincheira-Donoso (2005), Pincheira-Donoso & Núñez (2005) and Lobo et al. (2007) for recent discussions and additional details., Published as part of Pincheira-Donoso, Daniel, Scolaro, J. Alejandro & Sura, Piotr, 2008, A monographic catalogue on the systematics and phylogeny of the South American iguanian lizard family Liolaemidae (Squamata, Iguania), pp. 1-85 in Zootaxa 1800 on page 19, DOI: 10.5281/zenodo.6789337, {"references":["Werner, F. (1904) Ergebnisse der Hamburger Magalhaensischen Sammelreise. I. Band. Allgemeines, Chordonier, Echinodermen und Coelenteraten. IV. Reptilien und Batrachier, L. Friedrichsen and Co., Hamburg, 21 pp.","Muller, L. & Hellmich, W. (1938 a) Beitrage zur Kenntnis der Herpetofauna Chiles. XI. Uber zwei neue Liolaemus Formen aus der chilenischen Kordillere (Sammlung Schroder). Zoologischer Anzeiger, 122, 225 - 237.","Donoso-Barros, R. (1966 a) Reptiles de Chile. Ediciones de la Universidad de Chile, Santiago, 458 pp.","Etheridge, R & Espinoza, R. E. (2000) Taxonomy of the Liolaeminae (Squamata: Iguania: Tropiduridae) and a semiannotated bibliography. Smithsonian Herpetological Information Service, 126, 1 - 64.","Barbour, T. (1909) Some new South American cold-blooded vertebrates. Proceedings of the New England Zoological Club, 4, 47 - 52.","Burt, C. E. & Burt, M. D. (1931) South American lizards in the collection of the American Museum of Natural History. Bulletin of the American Museum of Natural History, 61, 227 - 395.","Laurent, R. F. (1992) On some overlooked species of the genus Liolaemus Wiegmann (Reptilia-Tropiduridae) from Peru. Breviora, 494, 1 - 33.","Nunez, H. & Jaksic, F. M. (1992) Lista comentada de los reptiles terrestres de Chile continental. Boletin del Museo Nacional de Historia Natural de Chile, 43, 63 - 91.","Etheridge, R. (1995) Redescription of Ctenoblepharys adspersa Tschudi, 1845, and the taxonomy of Liolaeminae (Reptilia: Squamata: Tropiduridae). American Museum Novitates, 3142, 1 - 34.","Carrillo de Espinoza, N. & Icochea, J. (1995) Lista taxonomica preliminar de los reptiles vivientes del Peru. Publicaciones del Museo de Historia Natural \" Javier Prado \", Universidad Nacional Mayor de San Marcos, Serie A, Zoologia, 49, 1 - 27.","Escobar, C. A., Labra, A. & Niemeyer, H. M. (2001) Chemical composition of precloacal secretions of Liolaemus lizards. Journal of Chemical Ecology, 27, 1677 - 1690.","Martinez-Oliver, I. & Lobo, F. (2002) Una nueva especie de Liolaemus del grupo alticolor (Iguania: Liolaemidae) de la Puna Saltena, Argentina. Cuadernos de Herpetologia, 16, 47 - 64.","Lobo, F. & Espinoza, R. E. (2004) Two new Liolaemus from the Puna Region of Argentina and Chile: further resolution of purported reproductive bimodality in Liolaemus alticolor (Iguania: Liolaemidae). Copeia, 2004, 850 - 867.","Chebez, J. C., Rey, N. R. & Williams, J. D. (2005) Reptiles de los parques nacionales de la Argentina. Literatura of Latin America, Buenos Aires, 37 pp.","Sura, P. 2005. Encyclopedia of extant amphibians and reptiles, Wydawnictwo Fundacja, Nowy Sacz, 544 pp.","Diaz, J. M. & Lobo, F. (2006) Historical biogeography of a clade of Liolaemus (Iguania: Liolaemidae) based on ancestral areas and dispersal-vicariance analysis (DIVA). Papeis Avulsos de Zoologia, 46, 261 - 274.","Hellmich, W. (1961) Bemerkungen zur geographischen Variabilitat von Liolaemus alticolor Barbour. Opuscula Zoologica, 58, 1 - 6.","Donoso-Barros, R. (1964) Ecologia herpetologica del Norte Grande de Chile. Noticiario Mensual del Museo Nacional de Historia Natural de Chile, 101, 6 - 7.","Donoso-Barros, R. (1970 a) Liolaemus. In: Peters, J. A. & Donoso-Barros, R. (Ed.), Catalogue of the Neotropical Squamata. Part II. Lizards and Amphisbaenians. Bulletin United States National Museum, 297, 170 - 195.","Donoso-Barros, R. (1970 b) Catalogo herpetologico chileno. Boletin del Museo Nacional de Historia Natural de Chile, 31, 49 - 124.","Vanzolini, P. E. (1986) Addenda and corrigenda to the Catalogue of Neotropical Squamata. Smithsonian Herpetological Information Service, 70, 1 - 25.","Veloso, A. & Navarro, J. (1988) Lista sistematica y distribucion geografica de anfibios y reptiles de Chile. Bollettino del Museo Regionale di Scienze Naturali di Torino, 6, 481 - 539.","Baudoin, W., Pacheco, M. J. Pacheco, L. (1990) Reptiles. In: Forno, E. & Boudoin, M. (Ed.), Historia Natural de un valle en Los Andes: La Paz. Servicio Grafico Quipus, La Paz, 421 - 452.","Pincheira-Donoso, D. & Nunez, H. (2005) Las especies chilenas del genero Liolaemus (Iguania, Tropiduridae, Liolaeminae). Taxonomia, sistematica y evolucion, Publicacion Ocasional del Museo Nacional de Historia Natural de Chile, Santiago, 486 pp.","Lobo, F., & Espinoza, R. E. (1999) Two new cryptic species of Liolaemus (Iguania: Tropiduridae) from nothwestern Argentina: resolution of the purported reproductive biomodality of Liolaemus alticolor. Copeia, 1999, 122 - 140.","Pincheira-Donoso, D. (2005) Anfibios y reptiles de la Provincia de El Loa. In: Ramirez, G. M. & Pincheira-Donoso, D. (Ed.), Fauna del Altiplano y Desierto de Atacama. Vertebrados de la Provincia de El Loa. Phrynosaura Ediciones, Calama, 93 - 150.","Lobo, F. & Abdala, C. S. (2007) Descripcion de una nueva especie de Phymaturus del grupo de P. palluma de la Provincia de Mendoza, Argentina. Cuadernos de Herpetologia, 21, 103 - 113."]}

Liolaemus scrocchii Quinteros, Abdala & Lobo Liolaemus dorbignyi (see references in Quinteros et al. 2008: 59). Liolaemus scrocchii Quinteros et al. 2008: 59 (type locality: Two km SE of Susques on National Road 52, Susques Department, Jujuy Province, Argentina)., Published as part of Pincheira-Donoso, Daniel, Scolaro, J. Alejandro & Sura, Piotr, 2008, A monographic catalogue on the systematics and phylogeny of the South American iguanian lizard family Liolaemidae (Squamata, Iguania), pp. 1-85 in Zootaxa 1800 on page 44, DOI: 10.5281/zenodo.6789337, {"references":["Quinteros, A. S., Abdala, C. S. & Lobo, F. J. (2008) Redescription of Liolaemus dorbignyi Koslowskyi, 1898 and description of a new species of Liolaemus (Iguania: Liolaemidae). Zootaxa, 1717, 51 - 67."]}

Liolaemus morenoi Etheridge & Christie Liolaemus morenoi Etheridge & Christie 2003: 336 (type locality: Sand dunes near National Road 40, 2 km S Cerrito Piñon, 650 m, Collón Curá Department, Neuquén Province, Argentina); Sura 2005: 429; Scolaro 2006: 42; Abdala 2007: 44., Published as part of Pincheira-Donoso, Daniel, Scolaro, J. Alejandro & Sura, Piotr, 2008, A monographic catalogue on the systematics and phylogeny of the South American iguanian lizard family Liolaemidae (Squamata, Iguania), pp. 1-85 in Zootaxa 1800 on page 52, DOI: 10.5281/zenodo.6789337, {"references":["Etheridge, R. & Christie, M. I. (2003) Two new species of the lizard genus Liolaemus (Squamata: Liolaemidae) from northern Patagonia, with comments on Liolaemus rothi. Journal of Herpetology, 37, 325 - 341.","Sura, P. 2005. Encyclopedia of extant amphibians and reptiles, Wydawnictwo Fundacja, Nowy Sacz, 544 pp.","Scolaro, J. A. (2006) Reptiles patagonicos: Norte. Una guia de campo, Ediciones Universidad Nacional de la Patagonia San Juan Bosco, Comodoro Rivadavia, 112 p.","Abdala, C. S. (2007) Phylogeny of the boulengeri group (Iguania: Liolaemidae, Liolaemus) based on morphological and molecular characters. Zootaxa, 1538, 1 - 84."]}

Liolaemus kolengh Abdala & Lobo Liolaemus kolengh Abdala & Lobo 2006b: 2 (type locality: Ceballos Hill way, near to Río Ceballos [47°01.542’S, 71°48.475’W], 1485 m, Lago Buenos Aires Department, Santa Cruz Province, Argentina); Pincheira-Donoso & Scolaro 2007: 57. Liolaemus lineomaculatus Boulenger Proctotretus kingii Bell 1843: 13 (in part, fide Donoso-Barros 1966a); Gray 1845: 215 (in part, fide Donoso-Barros 1966a). Liolaemus lineomaculatus Boulenger 1885a: 149 (type locality: “Patagonia”. Restricted to Puerto Deseado, Patagonia, Argentina, by Donoso-Barros 1966a); Koslowsky 1895: 363, 1896: 450; Andersson 1898: 461; Stejneger 1909: 217; Burt & Burt 1931: 275, 1933: 33; Liebermann 1939: 72; Donoso-Barros & Codoceo 1962: 28; Donoso-Barros 1966a: 275, 1970a: 185; 1970b: 93; Cei 1971b: 417, 1974c: 183, 1986: 264; Porter 1972: 401; Vanzolini 1986: 15; Veloso & Navarro 1988: 506; Núñez & Jaksic 1992: 74; Pereyra et al. 1992: 277; Scolaro 1992: 287; Etheridge 1995: 33; Videla & Cei 1998: 63; Etheridge & Espinoza 2000: 7; Schulte et al. 2000: 78; Lobo 2001: 139; Chebez et al. 2005: 32; Scolaro 2005: 41; Sura 2005: 429; Pincheira-Donoso & Scolaro 2007: 57. Liolaemus hatcheri Stejneger 1909: 218 (see Scolaro & Cei 2002). Vilcunia lineomaculatus Laurent 1985b: 7. Liolaemus (Liolaemus) lineomaculatus Pincheira-Donoso & Núñez 2005: 293., Published as part of Pincheira-Donoso, Daniel, Scolaro, J. Alejandro & Sura, Piotr, 2008, A monographic catalogue on the systematics and phylogeny of the South American iguanian lizard family Liolaemidae (Squamata, Iguania), pp. 1-85 in Zootaxa 1800 on page 38, DOI: 10.5281/zenodo.6789337, {"references":["Abdala, C. S. & Lobo, F. (2006 b) Description of a new Patagonian lizard species of the Liolaemus silvanae group (Iguania: Liolaemidae). South American Journal of Herpetology, 1, 1 - 8.","Scolaro, J. A. & Ibarguengoytia, N. R. (2007) A new species of Phymaturus from rocky outcrops in the central steppe of Rio Negro province, Patagonia Argentina (Reptilia: Iguania: Liolaemidae). Zootaxa, 1524, 47 - 55.","Bell, T. (1843) Reptiles, Part V. In: Darwin, C. (Ed.), The Zoology of the voyage of H. M. S. Beagle, under command of Captain Fitzroy, R. N., during the years 1832 to 1836. Smith, Elder & Co., London, 1 - 55.","Donoso-Barros, R. (1966 a) Reptiles de Chile. Ediciones de la Universidad de Chile, Santiago, 458 pp.","Gray, J. E. (1845) Catalogue of the specimens of lizards in the collection of the British Museum, Trustees of the British Museum, London, 289 pp.","Boulenger, G. A. (1885 a) Catalogue of the lizards in the British Museum (Natural History), London, 497 pp.","Koslowsky, J. (1895) Batracios y reptiles de La Rioja y Catamarca (Republica Argentina) recogidos durante los meses de Febrero a Mayo de 1895 (expedicion del Director del Museo). Revista Museo La Plata, 6, 357 - 370.","Koslowsky, J. (1896) Sobre algunos reptiles de patagonia y otras regiones argentinas. Revista del Museo de La Plata, 7, 447 - 457.","Andersson, L. G. (1898) List of reptiles and batrachians collected by the Swedish Expedition to Tierra del Fuego (1895 - 1896) under direction of Dr. Otto Nordenskiold. Ofversigt af K. Vetenskaps-Akademiens Forhanfingar, 1898, 457 - 462.","Stejneger, L. (1909) Batrachians and reptiles. Part II. In: Scott, W. B. (Ed.), Reports of the Princeton University expeditions to Patagonia, 1896 - 1899. Zoology, Princeton University, Princeton, 211 - 224.","Burt, C. E. & Burt, M. D. (1931) South American lizards in the collection of the American Museum of Natural History. Bulletin of the American Museum of Natural History, 61, 227 - 395.","Burt, C. E. & Burt, M. D. (1933) A preliminary check list of the lizards of South America. Transactions of the Academy of Science of St. Louis, 28, 1 - 104.","Liebermann, J. (1939) Catalogo sistematico y zoogeografico de los Lacertilidos argentinos. Physis, 16, 61 - 82.","Donoso-Barros, R. & Codoceo, M. (1962) Reptiles de Aysen y Magallanes. Boletin del Museo Nacional de Historia Natural de Chile, 28, 1 - 45.","Donoso-Barros, R. (1970 a) Liolaemus. In: Peters, J. A. & Donoso-Barros, R. (Ed.), Catalogue of the Neotropical Squamata. Part II. Lizards and Amphisbaenians. Bulletin United States National Museum, 297, 170 - 195.","Donoso-Barros, R. (1970 b) Catalogo herpetologico chileno. Boletin del Museo Nacional de Historia Natural de Chile, 31, 49 - 124.","Cei, J. M. (1971 b) Herpetologia patagonica. I. Liolaemus del grupo magellanicus. Caracteristicas taxonomicas y geneticas. Physis, Buenos Aires, 30, 417 - 424.","Cei, J. M. (1974 c) Herpetologia patagonica VIII. La altiplancie volcanica entre Primos Pinos y Rio Kilka, Neuquen: notas Herpetologicas. Physis, Buenos Aires, 33, 183 - 185.","Cei, J. M. (1986) Reptiles del centro, centro-oeste y sur de la Argentina. Herpetofauna de las zonas aridas y semiaridas. Museo Regionale di Scienze Naturali di Torino, Monografie 4, 528 pp.","Porter, K. R. (1972) Herpetology, W. B. Saunders Co, Philadelphia, 523 pp.","Vanzolini, P. E. (1986) Addenda and corrigenda to the Catalogue of Neotropical Squamata. Smithsonian Herpetological Information Service, 70, 1 - 25.","Veloso, A. & Navarro, J. (1988) Lista sistematica y distribucion geografica de anfibios y reptiles de Chile. Bollettino del Museo Regionale di Scienze Naturali di Torino, 6, 481 - 539.","Nunez, H. & Jaksic, F. M. (1992) Lista comentada de los reptiles terrestres de Chile continental. Boletin del Museo Nacional de Historia Natural de Chile, 43, 63 - 91.","Pereyra, E. A., Mella, J. & Diaz, N. F. (1992) Analisis meristico preliminar de la estructura taxonomica del genero Liolaemus Wiegmann. Acta Zoologica Lilloana, 41, 277 - 286.","Scolaro, J. A. (1992) Morphological differences between Vilcunia and Liolaemus magellanicus and L. lineomaculatus: a discriminant analysis. Acta Zoologica Lilloana, 41, 287 - 293.","Etheridge, R. (1995) Redescription of Ctenoblepharys adspersa Tschudi, 1845, and the taxonomy of Liolaeminae (Reptilia: Squamata: Tropiduridae). American Museum Novitates, 3142, 1 - 34.","Videla, F. & Cei, J. M. (1998) Especes de Liolaemus sans pores precloacaux chez les males, le long des Andes meridionales (Reptilia: Squamata: Iguania: Tropiduridae). Revue Francais de Aquariologie, 25, 63 - 68.","Etheridge, R & Espinoza, R. E. (2000) Taxonomy of the Liolaeminae (Squamata: Iguania: Tropiduridae) and a semiannotated bibliography. Smithsonian Herpetological Information Service, 126, 1 - 64.","Schulte II, J. A., Macey, J. R., Espinoza, R. E. & Larson, A. (2000) Phylogenetic relationships in the iguanid lizard genus Liolaemus: multiple origins of viviparous reproduction and evidence for recurring Andean vicariance and dispersal. Biological Journal of the Linnean Society, 69, 75 - 102.","Lobo, F. (2001) A phylogenetic analysis of lizards of the Liolaemus chiliensis group (Iguania: Tropiduridae). Herpetological Journal, 11, 137 - 150.","Chebez, J. C., Rey, N. R. & Williams, J. D. (2005) Reptiles de los parques nacionales de la Argentina. Literatura of Latin America, Buenos Aires, 37 pp.","Scolaro, J. A. (2005) Reptiles patagonicos: Sur. Una guia de campo. Ediciones Universidad Nacional de la Patagonia San Juan Bosco, Trelew, 80 p.","Sura, P. 2005. Encyclopedia of extant amphibians and reptiles, Wydawnictwo Fundacja, Nowy Sacz, 544 pp.","Scolaro, J. A. & Cei, J. M. (2002) Vilcunia periglacialis Cei & Scolaro, 1982 (currently Liolaemus periglacialis; Reptilia, Sauria): proposed precedence over Liolaemus hatcheri Stejneger, 1909. Bulletin of Zoological Nomenclature, 59, 275 - 277.","Laurent, R. F. (1985 b) Segunda contribucion al conocimiento de la estructura taxonomica del genero Liolaemus Wiegmann (Iguanidae). Cuadernos de Herpetologia, 1, 1 - 37.","Pincheira-Donoso, D. & Nunez, H. (2005) Las especies chilenas del genero Liolaemus (Iguania, Tropiduridae, Liolaeminae). Taxonomia, sistematica y evolucion, Publicacion Ocasional del Museo Nacional de Historia Natural de Chile, Santiago, 486 pp."]}

Liolaemus genus In the recent decades, Laurent (1983, 1985b, 1992) was the first author that formulated an explicit and powerful hypothesis on the phylogenetic relationships of higher groups among Liolaemus lizards. After performing a series of statistical analyses on previously overlooked and underestimated morphological traits, he found that these lizards can be separated into two major groups, one mainly Andean-occidental and one Andean-oriental. Morphologically, the occidental group, called by Laurent (1985b, 1992) “Chilean group”, includes species characterized by narrow supralabial scales, usually four or fewer, the last one elongated and upturned in its posterior margin, and with a low number of precloacal glands (= preanal pores), usually four or fewer (see Etheridge 1995). This group of taxa was placed by Laurent (1985b, 1992) into the subgenus Liolaemus (sensu stricto). Species belonging to the oriental group, called “Argentinean group”, exhibit short supralabial scales, usually five or more, the last one is not upturned posteriorly, a high number of precloacal glands, usually five or more (up to 12), the puboischiotibialis muscle (flexor tibialis internus fide Abdala, V. et al. 2006) hypertrophied, and a conspicuous sharp, blade-like process on the tibia (see Etheridge 1995; also Núñez et al. 2000). This complex of species was recognized under the subgenus Eulaemus (Laurent 1992; see also Etheridge 1995). More recently, additional morphological evidence led Etheridge (1995) to support this classification model, who also recognized that a series of species grouped into the Eulaemus subgenus exhibit a unique patch of enlarged scales on the posterior surface of the thigh. These lizards were then recognized as the boulengeri complex (see Cei 1993; Etheridge 1995, 2000; Halloy et al. 1998; Pincheira-Donoso & Núñez 2005; Abdala 2007, for details). Remarkably, these groups of species have remained largely accepted, and more importantly, have recently been supported by phylogenetic hypotheses constructed on both morphological and molecular evidence (e.g. Etheridge 2000; Schulte et al. 2000; Espinoza et al. 2004; Cruz et al. 2005; Abdala 2007). Even though Laurent (1983, 1985b, 1992) and Etheridge (1995) reported pioneer evidence to hypothesize relationships among higher-level groups within Liolaemus, the first explicit phylogenetic study focusing on the relationships among species within these clades was only recently published (Schulte et al. 2000; see also Schulte et al. 1998). Schulte et al. ’s (2000) molecular phylogeny supported substantially the hypothesis of a major event of Liolaemus diversification into two main clades, the subgenus Liolaemus and the series signifier (indicated by Schulte et al. as subgenus Eulaemus). In addition, this phylogenetic study provided the first empirical evidence to test the hypothesis that austral Liolaemus species recognized as members of the groups archeforus and kingii (Scolaro & Cei 1997) may represent primitive Patagonian states in the evolutionary history of this genus (Laurent 1985b; see also Cei 1986). According to Laurent (1985b), morphological peculiarities observed in these austral species would be indicative of their ancestral position within Liolaemus (see also Etheridge 1995). However, Schulte et al. ’s (2000) results rejected such claims. Indeed, this phylogeny revealed that the archeforus-kingii line (i.e. subgenus Donosolaemus) would be related to the Eulaemus subgenus, and therefore, that it originated after the first main diversification event experienced by this genus, during the Tertiary (e.g. Pincheira-Donoso & Núñez 2003). Later phylogenetic studies have provided, in general, similar results in relation to the relationships among subgenera, among clades within subgenera (e.g. 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). In spite of an increasing number of studies exploring relationships among Liolaemus taxa published over the last few years, many aspects of the phylogeny of this lineage remain still controversial. For example, the position in the phylogenetic structure of the genus Liolaemus of L. pseudoanomalus, a species recognized by Cei (e.g. 1986, 1993; see also Laurent 1984a; Núñez & Yáñez 1984b) as a member of the anomalus group, tends to show substantial discordance among different studies (e.g. Schulte et al. 2004; Espinoza et al. 2004; Abdala 2007; Pincheira-Donoso et al. 2007a). More importantly, the phylogenetic relationships between a large number of Liolaemus species remain to be studied for the first time., Published as part of Pincheira-Donoso, Daniel, Scolaro, J. Alejandro & Sura, Piotr, 2008, A monographic catalogue on the systematics and phylogeny of the South American iguanian lizard family Liolaemidae (Squamata, Iguania), pp. 1-85 in Zootaxa 1800 on pages 15-17, DOI: 10.5281/zenodo.6789337, {"references":["Laurent, R. F. (1983) Contribucion al conocimiento de la estructura taxonomica del genero Liolaemus Wiegmann (Iguanidae). Boletin de la Asociacion Herpetologica Argentina, 1, 15 - 18.","Laurent, R. F. (1985 b) Segunda contribucion al conocimiento de la estructura taxonomica del genero Liolaemus Wiegmann (Iguanidae). Cuadernos de Herpetologia, 1, 1 - 37.","Laurent, R. F. (1992) On some overlooked species of the genus Liolaemus Wiegmann (Reptilia-Tropiduridae) from Peru. Breviora, 494, 1 - 33.","Etheridge, R. (1995) Redescription of Ctenoblepharys adspersa Tschudi, 1845, and the taxonomy of Liolaeminae (Reptilia: Squamata: Tropiduridae). American Museum Novitates, 3142, 1 - 34.","Nunez, H., Navarro, J. & Veloso, A. (2000) Liolaemus foxi, una nueva especie de lagarto para el norte de Chile (Squamata: Reptilia: Sauria). Boletin del Museo Nacional de Historia Natural de Chile, 49, 117 - 130.","Cei, J. M. (1993) Reptiles del noroeste, nordeste y este de la Argentina. Herpetofauna de las selvas subtropicales, Puna y Pampas. Museo Regionale di Scienze Naturali di Torino, Monografie 14, 949 pp.","Etheridge, R. (2000) A review of lizards of the Liolaemus wiegmannii group (Squamata, Iguania, Tropiduridae), and a history of morphological change in the sand-dwelling species. Herpetological Monographs, 14, 293 - 352.","Halloy, M., Etheridge, R. & Burghardt, G. M. (1998) To bury in the sand: phylogenetic relationships among lizard species of the boulengeri group, Liolaemus (Reptilia: Squamata: Tropiduridae), based on behavioral characters. Herpetological Monographs, 12, 1 - 37.","Pincheira-Donoso, D. & Nunez, H. (2005) Las especies chilenas del genero Liolaemus (Iguania, Tropiduridae, Liolaeminae). Taxonomia, sistematica y evolucion, Publicacion Ocasional del Museo Nacional de Historia Natural de Chile, Santiago, 486 pp.","Abdala, C. S. (2007) Phylogeny of the boulengeri group (Iguania: Liolaemidae, Liolaemus) based on morphological and molecular characters. Zootaxa, 1538, 1 - 84.","Schulte II, J. A., Macey, J. R., Espinoza, R. E. & Larson, A. (2000) Phylogenetic relationships in the iguanid lizard genus Liolaemus: multiple origins of viviparous reproduction and evidence for recurring Andean vicariance and dispersal. Biological Journal of the Linnean Society, 69, 75 - 102.","Espinoza, R. E., Wiens, J. J. & Tracy, C. R. (2004) Recurrent evolution of herbivory in small, cold-climate lizards: Breaking the ecophysiological rules or reptilian herbivory. Proceedings of the National Academy of Sciences, USA, 101, 16819 - 16824.","Cruz, F. B., Fitzgerald, L. A., Espinoza, R. E. & Schulte II, J. A. (2005). The importance of phylogenetic scale in tests of Bergmann's and Rapoport's rules: lessons from a clade of South American lizards. Journal of Evolutionary Biology, 18, 1559 - 1574.","Schulte II, J. A., Losos, J. B., Cruz, F. B. & Nunez, H. (2004) The relationship between morphology, escape behaviour and microhabitat occupation in the lizard clade Liolaemus (Iguanidae: Tropidurinae: Liolaemini). Journal of Evolutionary Biology, 17, 408 - 420.","Schulte II, J. A., Macey, J. R., Larson, A. & Papenfuss, T. J. (1998) Molecular tests of phylogenetic taxonomies: a general procedure and example using four subfamilies of the lizard family Iguanidae. Molecular Phylogenetics and Evolution, 10, 367 - 376.","Scolaro, J. A. & Cei, J. M. (1997) Systematic status and relationships of Liolaemus species of the archeforus and kingii groups: a morphological and taxonumerical approach (Reptilia: Tropiduridae). Bolletino Museo Regionale di Scienze Naturali di Torino, 15, 369 - 406.","Cei, J. M. (1986) Reptiles del centro, centro-oeste y sur de la Argentina. Herpetofauna de las zonas aridas y semiaridas. Museo Regionale di Scienze Naturali di Torino, Monografie 4, 528 pp.","Pincheira-Donoso, D. & Nunez, H. (2003) Liolaemus robertoi, una nueva especie de Los Andes del norte de Chile perteneciente al grupo ruibali (Iguania: Tropiduridae: Liolaeminae). Multequina, Latin American Journal of Natural Resources, 12, 1 - 15.","Avila, L. J., Morando, M., Perez, C. H. F. & Sites, J. W, Jr. (2004) Phylogenetic relationships of lizards of the Liolaemus petrophilus group (Squamata, Liolaemidae), with description of two new species from western Argentina. Herpetologica, 60, 187 - 203.","Morando, M., Avila, L. J., Baker, J. & Sites, J. W. (2004) Phylogeny and phylogeography of the Liolaemus darwinii complex (Squamata: Liolaemidae): evidence for introgression and incomplete lineage sorting. Evolution, 58, 842 - 861.","Pincheira-Donoso, D., Scolaro, J. A. & Schulte II, J. A. (2007 a) The limits of polymorphism in Liolaemus rothi: molecular and phenotypic evidence for a new species of the Liolaemus boulengeri clade (Iguanidae, Liolaemini) from boreal Patagonia of Chile. Zootaxa, 1452, 25 - 42.","Lobo, F. (2001) A phylogenetic analysis of lizards of the Liolaemus chiliensis group (Iguania: Tropiduridae). Herpetological Journal, 11, 137 - 150.","Diaz, J. M. & Lobo, F. (2006) Historical biogeography of a clade of Liolaemus (Iguania: Liolaemidae) based on ancestral areas and dispersal-vicariance analysis (DIVA). Papeis Avulsos de Zoologia, 46, 261 - 274.","Laurent, R. F. (1984 a) On some iguanid genera related to or previously confused with Liolaemus Wiegmann. Journal of Herpetology, 18, 357 - 373.","Nunez, H. & Yanez, J. (1984 b) Abas y Velosaura nuevos generos de lagartos Iguanidae y proposiciones sistematicas respecto de los grupos aliados (Reptilia: Squamata). Boletin del Museo Nacional de Historia Natural de Chile, 40, 97 - 105."]}

Liolaemus chacoensis Shreve Liolaemus chacoensis Shreve 1948: 111 (type locality: Fortin Guachalla, Río Pilcomayo, 580 km W Asunción, Chaco Paraguayo, Paraguay); Hellmich 1960: 38; Donoso-Barros 1970a: 181; Cei 1980a: 936, 1986: 219, 1993: 277; Vanzolini 1986: 13; Alvarez et al. 1988: 85; Ramírez-Pinilla & Cruz 1991: 10; Laurent 1992: 31; Cabrera 1993: 27, 1996: 219; Cruz et al. 1993: 101; Martori & Aun 1994: 97, 1995: 94; Etheridge 1995: 33; Lavilla et al. 1995: 117; Aquino et al. 1996: 331; Cruz & Ramírez-Pinilla 1996: 33; Avila et al. 1998: 11; Fitzgerald et al. 1999: 536; Etheridge & Espinoza 2000: 5; Lobo 2001: 139; Chebez et al. 2005: 30; Sura 2005: 428; Abdala 2007: 44. Liolaemus emmae Donoso-Barros 1970c: 23 (synonym fide Cei 1980a); Cei 1973d: 223. Liolaemus (Eulaemus) chacoensis Cabrera & Bee de Speroni 1986: 5., Published as part of Pincheira-Donoso, Daniel, Scolaro, J. Alejandro & Sura, Piotr, 2008, A monographic catalogue on the systematics and phylogeny of the South American iguanian lizard family Liolaemidae (Squamata, Iguania), pp. 1-85 in Zootaxa 1800 on page 49, DOI: 10.5281/zenodo.6789337, {"references":["Shreve, B. (1948) A new Liolaemus from Paraguay. Copeia, 1948, 111 - 112.","Hellmich, W. (1960) Die Sauria der Gran Chaco und seiner Randgebiete. Abhandlungen Bayerische Akademie der Wissenschaften, 101, 1 - 131.","Donoso-Barros, R. (1970 a) Liolaemus. In: Peters, J. A. & Donoso-Barros, R. (Ed.), Catalogue of the Neotropical Squamata. Part II. Lizards and Amphisbaenians. Bulletin United States National Museum, 297, 170 - 195.","Cei, J. M. (1980 a) The conspecificity of the iguanid lizard Liolaemus chacoensis Shreve, 1948 and Liolaemus emmae Donoso Barros, 1970. Copeia, 1980, 936 - 937.","Cei, J. M. (1986) Reptiles del centro, centro-oeste y sur de la Argentina. Herpetofauna de las zonas aridas y semiaridas. Museo Regionale di Scienze Naturali di Torino, Monografie 4, 528 pp.","Cei, J. M. (1993) Reptiles del noroeste, nordeste y este de la Argentina. Herpetofauna de las selvas subtropicales, Puna y Pampas. Museo Regionale di Scienze Naturali di Torino, Monografie 14, 949 pp.","Vanzolini, P. E. (1986) Addenda and corrigenda to the Catalogue of Neotropical Squamata. Smithsonian Herpetological Information Service, 70, 1 - 25.","Alvarez, B. B., Tedesco, M. E. & Hernando, A. B. (1988) Nota preliminar sobre la composicion y distribucion de la lacertofauna de Corrientes, Chaco y Formosa (Argentina). Revista de la Asociacion de Ciencias Naturales del Litoral, 19, 79 - 89.","Laurent, R. F. (1992) On some overlooked species of the genus Liolaemus Wiegmann (Reptilia-Tropiduridae) from Peru. Breviora, 494, 1 - 33.","Cabrera, M. R. (1993) Los saurios del espinal y su presencia en provincias biogeographicas vencinas. Revista del Museo Argentino de Ciencias Naturales \" Bernardino Rivadavia \" e Instituto Nacional de Investigacion de las Ciencias Naturales, 4, 17 - 34.","Cabrera, M. R. (1996) Lista y distribucion geografica de saurios, anfisbenas y tortugas (Amniota, Reptilia) de la provincia de Cordoba. In: di Tada, I. E. & Bucher, E. H. (Ed.), Biodiversidad de Provincia Cordoba. I. Fauna. Universidad Nacional de Rio Cuarto, Rio Cuarto, Cordoba, 215 - 238.","Cruz, F. B., Perotti, M. G. & Fitzgerald, L. A. (1993) Lista de anfibios y reptiles colectados en una localidad del chaco Salteno. Acta Zoologica Lilloana, 42, 101 - 107.","Martori, R. & Aun, L. (1994) Analisis comparativo de la composicion de tres comunidades de Squamata de la Sierra Grande de Cordoba, Argentina. Cuadernos de Herpetologia, 8, 97 - 103.","Martori, R. & Aun, L. (1995) Relevamiento de la herpetofauna de las Sierras Pampeanas del este en la provincia de Cordoba (Argentina) (registro de taxa y comentarios taxonomicos). Acta Zoologica Lilloana, 43, 87 - 97.","Etheridge, R. (1995) Redescription of Ctenoblepharys adspersa Tschudi, 1845, and the taxonomy of Liolaeminae (Reptilia: Squamata: Tropiduridae). American Museum Novitates, 3142, 1 - 34.","Lavilla, E. O., Cruz, F. B. & Scrocchi, G. J. (1995) Amphibiens et reptiles de la Station Biologique \" Los Colorados \" dan la provincia de Salta, Argentina (2 e partie). Revue Francais de Aquariologie, 22, 117 - 128.","Aquino, A. L., Scott, N. J. & Motte, M. (1996) Lista de anfibios y reptiles del Museo Nacional de Historia Natural del Paraguay (marzo, 1980 - septiembre 1995). Pp. 331 - 400, in: MNHNP, Colecciones de fauna y flora del Museo Nacional de Historia Natural del Paraguay. Ministerio de Agricultura y Ganaderia, Asuncion, Paraguay.","Avila, L. J., Acosta, J. C. & Murua, A. F. (1998) Herpetofauna de la provincia de San Juan, Argentina: lista comentada y distribucion geografica. Cuadernos de Herpetologia, 12, 11 - 29.","Fitzgerald, L. A., Cruz, F. B. & Perotti, G. (1999) Phenology of a lizard assemblage in the dry Chaco of Argentina. Journal of Herpetology, 33, 536 - 535.","Etheridge, R & Espinoza, R. E. (2000) Taxonomy of the Liolaeminae (Squamata: Iguania: Tropiduridae) and a semiannotated bibliography. Smithsonian Herpetological Information Service, 126, 1 - 64.","Lobo, F. (2001) A phylogenetic analysis of lizards of the Liolaemus chiliensis group (Iguania: Tropiduridae). Herpetological Journal, 11, 137 - 150.","Chebez, J. C., Rey, N. R. & Williams, J. D. (2005) Reptiles de los parques nacionales de la Argentina. Literatura of Latin America, Buenos Aires, 37 pp.","Sura, P. 2005. Encyclopedia of extant amphibians and reptiles, Wydawnictwo Fundacja, Nowy Sacz, 544 pp.","Abdala, C. S. (2007) Phylogeny of the boulengeri group (Iguania: Liolaemidae, Liolaemus) based on morphological and molecular characters. Zootaxa, 1538, 1 - 84.","Donoso-Barros, R. (1970 c) Un nuevo reptil de Argentina, Liolaemus emmae (Sauria, Iguanidae). Boletin de la Sociedad de Biologia de Concepcion, 42, 23 - 26.","Cei, J. M. (1973 d) Comentarios sobre una nueva especie Argentina de saurio: Liolaemus emmae Donoso Barros (1970). Boletin Sociedad de Biologia de Concepcion, 46, 223 - 225.","Cabrera, M. R. & Bee de Speroni, N. T. (1986) Composicion y distribucion de la lacertofauna de la provincia de Cordoba, Argentina. II. Amphisbaenidae, Anguidae e Iguanidae. Historia Natural en Corrientes, Argentina, 6, 1 - 12."]}

Liolaemus sagei Etheridge & Christie Liolaemus sagei Etheridge & Christie 2003: 332 (type locality: Northeastern shore of Laguna del Toro [= Laguna Barra Tom Curá], 7 km S, 3.5 km E Cerro Mesa, Collón Curá Department, Neuquén Province, Argentina); Sura 2005: 430; Scolaro 2006: 47; Abdala 2007: 44; Pincheira-Donoso et al. 2007a: 28., Published as part of Pincheira-Donoso, Daniel, Scolaro, J. Alejandro & Sura, Piotr, 2008, A monographic catalogue on the systematics and phylogeny of the South American iguanian lizard family Liolaemidae (Squamata, Iguania), pp. 1-85 in Zootaxa 1800 on page 53, DOI: 10.5281/zenodo.6789337, {"references":["Etheridge, R. & Christie, M. I. (2003) Two new species of the lizard genus Liolaemus (Squamata: Liolaemidae) from northern Patagonia, with comments on Liolaemus rothi. Journal of Herpetology, 37, 325 - 341.","Sura, P. 2005. Encyclopedia of extant amphibians and reptiles, Wydawnictwo Fundacja, Nowy Sacz, 544 pp.","Scolaro, J. A. (2006) Reptiles patagonicos: Norte. Una guia de campo, Ediciones Universidad Nacional de la Patagonia San Juan Bosco, Comodoro Rivadavia, 112 p.","Abdala, C. S. (2007) Phylogeny of the boulengeri group (Iguania: Liolaemidae, Liolaemus) based on morphological and molecular characters. Zootaxa, 1538, 1 - 84.","Pincheira-Donoso, D., Scolaro, J. A. & Schulte II, J. A. (2007 a) The limits of polymorphism in Liolaemus rothi: molecular and phenotypic evidence for a new species of the Liolaemus boulengeri clade (Iguanidae, Liolaemini) from boreal Patagonia of Chile. Zootaxa, 1452, 25 - 42."]}

Liolaemus anomalus Koslowsky Liolaemus anomalus Koslowsky 1896: 452 (type locality: La Rioja Province, Argentina); Liebermann 1939: 72; Cei 1979a: 183; Williams & Cei 1983: 7; Tiranti & Avila 1994: 40; Avila et al. 1998: 11; Chebez et al. 2005: 28. Phrynosaura werneri Müller 1928: 64 (synonym fide Cei 1979a); Donoso-Barros 1966a: 345; Peters & Donoso-Barros 1970: 221. Liolaemus lentus Gallardo 1966: 17 (synonym fide Donoso-Barros 1969a), 1968: 7; Abdala 2007: 44. Ctenoblepharis anomalus Donoso-Barros 1969a: 94; Peters & Donoso-Barros 1970: 103. Ctenoblepharis werneri Cei 1974a: 71. Ceiolaemus anomalus Laurent 1984a: 359. Liolaemus anomalus anomalus Cei 1986: 201; Etheridge 1995: 34; Etheridge & Espinoza 2000: 4. Ceiolaemus anomalus anomalus Cabrera 1993: 27. Liolaemus anomalus Abdala 2007: 39. Observations: In a recent phylogenetic study, Abdala (2007) suggested full species status for L. anomalus anomalus and L. anomalus diatadai (Abdala 2007: 39). Since L. anomalus and the taxon ditadai exhibit clear patterns of divergence in a series of morphological and coloration traits, we accept Abdala’s (2007) suggestions., Published as part of Pincheira-Donoso, Daniel, Scolaro, J. Alejandro & Sura, Piotr, 2008, A monographic catalogue on the systematics and phylogeny of the South American iguanian lizard family Liolaemidae (Squamata, Iguania), pp. 1-85 in Zootaxa 1800 on page 47, DOI: 10.5281/zenodo.6789337, {"references":["Koslowsky, J. (1896) Sobre algunos reptiles de patagonia y otras regiones argentinas. Revista del Museo de La Plata, 7, 447 - 457.","Liebermann, J. (1939) Catalogo sistematico y zoogeografico de los Lacertilidos argentinos. Physis, 16, 61 - 82.","Cei, J. M. (1979 a) Remarks on the South American lizard Liolaemus anomolus Koslowsky, and the synonomy of Phrynosaura werneri Muller (Reptilia, Lacertilia, Iguanidae). Journal of Herpetology, 13, 183 - 186.","Williams, J. D. & Cei, J. M. (1983) Redescubrimiento de los sintipos de Liolaemus anomalus Koslowsky, 1896: comentarios sobre la distribucion de la especie. Boletin de la Asociacion Herpetologica Argentina, 1, 7 - 8.","Tiranti, S. & Avila, L. J. (1994) Aportes a la herpetologia de la provincia de La Pampa. I. Lista de los taxa de presencia segura en la provincia. Boletin de la Asociacion Herpetologica Argentina, 10, 39 - 41.","Avila, L. J., Acosta, J. C. & Murua, A. F. (1998) Herpetofauna de la provincia de San Juan, Argentina: lista comentada y distribucion geografica. Cuadernos de Herpetologia, 12, 11 - 29.","Chebez, J. C., Rey, N. R. & Williams, J. D. (2005) Reptiles de los parques nacionales de la Argentina. Literatura of Latin America, Buenos Aires, 37 pp.","Muller, L. (1928) Herpetologische Mitteilungen. Zoologischer Anzeiger, 77, 61 - 84.","Donoso-Barros, R. (1966 a) Reptiles de Chile. Ediciones de la Universidad de Chile, Santiago, 458 pp.","Peters, J. A. & Donoso-Barros, R. (1970) Catalogue of the Neotropical Squamata. Part II. Lizards and Amphisbaenians. Bulletin United States National Museum, 297, 1 - 293.","Gallardo, J. M. (1966) Liolaemus lentus nov. sp. (Iguanidae) de la pampa y algunas observaciones sobre los saurios de dicha provincia argentina y del oeste de Buenos Aires. Neotropica, 12, 15 - 29.","Donoso-Barros, R. (1969 a) Consideraciones nomenclaturales sobre dos lagartijas Argentinas. Boletin Sociedad de Biologia de Concepcion, 41, 93 - 94.","Abdala, C. S. (2007) Phylogeny of the boulengeri group (Iguania: Liolaemidae, Liolaemus) based on morphological and molecular characters. Zootaxa, 1538, 1 - 84.","Cei, J. M. (1974 a) Two new species of Ctenoblepharis (Reptilia, Iguanidae) from the arid environments of the central Argentina (Mendoza Province). Journal of Herpetology, 8, 71 - 75.","Laurent, R. F. (1984 a) On some iguanid genera related to or previously confused with Liolaemus Wiegmann. Journal of Herpetology, 18, 357 - 373.","Cei, J. M. (1986) Reptiles del centro, centro-oeste y sur de la Argentina. Herpetofauna de las zonas aridas y semiaridas. Museo Regionale di Scienze Naturali di Torino, Monografie 4, 528 pp.","Etheridge, R. (1995) Redescription of Ctenoblepharys adspersa Tschudi, 1845, and the taxonomy of Liolaeminae (Reptilia: Squamata: Tropiduridae). American Museum Novitates, 3142, 1 - 34.","Etheridge, R & Espinoza, R. E. (2000) Taxonomy of the Liolaeminae (Squamata: Iguania: Tropiduridae) and a semiannotated bibliography. Smithsonian Herpetological Information Service, 126, 1 - 64.","Cabrera, M. R. (1993) Los saurios del espinal y su presencia en provincias biogeographicas vencinas. Revista del Museo Argentino de Ciencias Naturales \" Bernardino Rivadavia \" e Instituto Nacional de Investigacion de las Ciencias Naturales, 4, 17 - 34."]}

Liolaemus josei Abdala Liolaemus boulengeri Cei 1973a: 459 (in part), 1986: 220 (in part). Liolaemus josei Abdala 2005a: 19 (type locality: El Zampal Bridge, on Nacional Road 40, medanos of Rio Grande, Malargüe Department, Mendoza Province, Argentina), 2007: 44; Scolaro 2006: 37., Published as part of Pincheira-Donoso, Daniel, Scolaro, J. Alejandro & Sura, Piotr, 2008, A monographic catalogue on the systematics and phylogeny of the South American iguanian lizard family Liolaemidae (Squamata, Iguania), pp. 1-85 in Zootaxa 1800 on page 51, DOI: 10.5281/zenodo.6789337, {"references":["Cei, J. M. (1973 a) Herpetologia patagonica. VI. Los Liolaemus del grupo fitzingeri en Santa Cruz y Chubut (Sauria, Iguanidae). Physis, Buenos Aires, 32, 447 - 458.","Abdala, C. S. (2005 a) Dos nuevas especies del genero Liolaemus (Iguania: Liolaemidae) y redescripcion de Liolaemus boulengeri (Koslowsky, 1898). Cuadernos de Herpetologia, 19, 3 - 33.","Scolaro, J. A. (2006) Reptiles patagonicos: Norte. Una guia de campo, Ediciones Universidad Nacional de la Patagonia San Juan Bosco, Comodoro Rivadavia, 112 p."]}

Liolaemus salinicola Laurent Liolaemus (Ortholaemus) salinicola Laurent 1986: 92 (type locality: Salar de Pipanaco, 950 m, Pomán Department, Catamarca Province, Argentina). Liolaemus salinicola Cei 1993: 274; Etheridge 1995: 34, 2000: 306; Etheridge & Espinoza 2000: 9; Verrastro et al. 2003: 109; Chebez et al. 2005: 35; Sura 2005: 430; Abdala 2007: 44., Published as part of Pincheira-Donoso, Daniel, Scolaro, J. Alejandro & Sura, Piotr, 2008, A monographic catalogue on the systematics and phylogeny of the South American iguanian lizard family Liolaemidae (Squamata, Iguania), pp. 1-85 in Zootaxa 1800 on page 55, DOI: 10.5281/zenodo.6789337, {"references":["Laurent, R. F. (1986) Descripciones de nuevos Iguanidae del genero Liolaemus. Acta Zoologica Lilloana, 38, 87 - 105.","Cei, J. M. (1993) Reptiles del noroeste, nordeste y este de la Argentina. Herpetofauna de las selvas subtropicales, Puna y Pampas. Museo Regionale di Scienze Naturali di Torino, Monografie 14, 949 pp.","Etheridge, R. (1995) Redescription of Ctenoblepharys adspersa Tschudi, 1845, and the taxonomy of Liolaeminae (Reptilia: Squamata: Tropiduridae). American Museum Novitates, 3142, 1 - 34.","Etheridge, R. (2000) A review of lizards of the Liolaemus wiegmannii group (Squamata, Iguania, Tropiduridae), and a history of morphological change in the sand-dwelling species. Herpetological Monographs, 14, 293 - 352.","Etheridge, R & Espinoza, R. E. (2000) Taxonomy of the Liolaeminae (Squamata: Iguania: Tropiduridae) and a semiannotated bibliography. Smithsonian Herpetological Information Service, 126, 1 - 64.","Verrastro, L., Veronese, L., Bujes, C. & Martins Dias Filho, M. (2003) A new species of Liolaemus from southern Brazil (Iguania: Tropiduridae). Herpetologica, 59, 105 - 118.","Chebez, J. C., Rey, N. R. & Williams, J. D. (2005) Reptiles de los parques nacionales de la Argentina. Literatura of Latin America, Buenos Aires, 37 pp.","Sura, P. 2005. Encyclopedia of extant amphibians and reptiles, Wydawnictwo Fundacja, Nowy Sacz, 544 pp.","Abdala, C. S. (2007) Phylogeny of the boulengeri group (Iguania: Liolaemidae, Liolaemus) based on morphological and molecular characters. Zootaxa, 1538, 1 - 84."]}

Liolaemus koslowskyi Etheridge Liolaemus nigromaculatus (?) Koslowsky 1895: 363 (fide Etheridge 1993). Liolaemus darwini Koslowsky 1895: 364. Liolaemus darwinii (sensu lato) Cei 1993: 258 (in part). Liolaemus koslowskyi Etheridge 1993: 159 (type locality: 9.2 km E Plaza Central de Pituíl, Provincial Road 11, Famatina Department, Catamarca Province, Argentina), 1995: 34, 2000: 331; Cei 1993: 736; Etheridge & Espinoza 2000: 6; Chebez et al. 2005: 32; Sura 2005: 429; Abdala 2007: 44; Díaz 2007: 115., Published as part of Pincheira-Donoso, Daniel, Scolaro, J. Alejandro & Sura, Piotr, 2008, A monographic catalogue on the systematics and phylogeny of the South American iguanian lizard family Liolaemidae (Squamata, Iguania), pp. 1-85 in Zootaxa 1800 on page 51, DOI: 10.5281/zenodo.6789337, {"references":["Koslowsky, J. (1895) Batracios y reptiles de La Rioja y Catamarca (Republica Argentina) recogidos durante los meses de Febrero a Mayo de 1895 (expedicion del Director del Museo). Revista Museo La Plata, 6, 357 - 370.","Etheridge, R. (1993) Lizards of the Liolaemus darwinii Complex (Squamata: Iguania: Tropiduridae) in Northern Argentina. Bollettino del Museo Regionale di Scienze Naturali di Torino, 11, 137 - 199.","Cei, J. M. (1993) Reptiles del noroeste, nordeste y este de la Argentina. Herpetofauna de las selvas subtropicales, Puna y Pampas. Museo Regionale di Scienze Naturali di Torino, Monografie 14, 949 pp.","Etheridge, R & Espinoza, R. E. (2000) Taxonomy of the Liolaeminae (Squamata: Iguania: Tropiduridae) and a semiannotated bibliography. Smithsonian Herpetological Information Service, 126, 1 - 64.","Chebez, J. C., Rey, N. R. & Williams, J. D. (2005) Reptiles de los parques nacionales de la Argentina. Literatura of Latin America, Buenos Aires, 37 pp.","Sura, P. 2005. Encyclopedia of extant amphibians and reptiles, Wydawnictwo Fundacja, Nowy Sacz, 544 pp.","Abdala, C. S. (2007) Phylogeny of the boulengeri group (Iguania: Liolaemidae, Liolaemus) based on morphological and molecular characters. Zootaxa, 1538, 1 - 84.","Diaz, J. M. (2007) Reptilia, Iguania, Liolaeminae, Liolaemus, Puna, Prepuna, and mountain ranges, northwestern Argentina. Check List, 3, 105 - 118."]}

Liolaemus espinozai Abdala Liolaemus quilmes Etheridge 1993: 152 (in part). Liolaemus espinozai Abdala 2005b: 6 (type locality: El Ingenio, Campo El Arenal, 67 km S Santa Maria, Provincial Road 47, 2620 m, Andalgala Department, Catamarca Province, Argentina), 2007: 44., Published as part of Pincheira-Donoso, Daniel, Scolaro, J. Alejandro & Sura, Piotr, 2008, A monographic catalogue on the systematics and phylogeny of the South American iguanian lizard family Liolaemidae (Squamata, Iguania), pp. 1-85 in Zootaxa 1800 on page 50, DOI: 10.5281/zenodo.6789337, {"references":["Etheridge, R. (1993) Lizards of the Liolaemus darwinii Complex (Squamata: Iguania: Tropiduridae) in Northern Argentina. Bollettino del Museo Regionale di Scienze Naturali di Torino, 11, 137 - 199.","Abdala, C. S. (2005 b) Una nueva especie del genero Liolaemus perteneciente al complejo darwinii (Iguania: Liolaemidae) de la provincia de Catamarca, Argentina. Revista Espanola de Herpetologia, 19, 5 - 17."]}

Liolaemus fitzingerii (Duméril & Bibron) Proctotretus fitzingerii Duméril & Bibron 1837: 286 (type locality: Puerto Deseado, Santa Cruz Province, Argentina, fide Cei 1986; Chile Chico, Aysén Province, Chile, fide Donoso-Barros 1966a); Bell 1843: 11; Guichenot 1848: 38; Duméril & Duméril 1851: 74. Ptychodeira fitzingerii Fitzinger 1843: 73. Eulaemus affinis Girard 1858a: 198 (see Bell 1843; Etheridge & Espinoza 2000), 1858b: 366. Liolaemus (Eulaemus) fitzingerii Girard 1858b: 365. Liolaemus fitzingerii Boulenger 1885a: 150; Andersson 1898: 461; Berg 1898: 6; Burt & Burt 1931: 274; Liebermann 1939: 70; Hellmich 1950: 346; Donoso-Barros & Codoceo 1962: 9; Donoso-Barros 1966a: 293, 1970a: 182, 1970b: 91; Cei & Scolaro 1977b: 223, 1983a: 15; Scolaro & Cei 1977: 219; Núñez & Jaksic 1992: 72; Pereyra et al. 1992: 277; Etheridge 1995: 34; Etheridge & Espinoza 2000: 6; Chebez et al. 2005: 31; Scolaro 2005: 35; Sura 2005: 428; Abdala 2007: 44. Liolaemus fitzingeri fitzingeri Cei 1973a: 447, 1980b: 317, 1986: 208; Vanzolini 1986: 14; Veloso & Navarro 1988: 503; Brygoo 1989: 46; Laurent 1992: 31. Liolaemus (Eulaemus) fitzingerii Pincheira-Donoso & Núñez 2005: 95., Published as part of Pincheira-Donoso, Daniel, Scolaro, J. Alejandro & Sura, Piotr, 2008, A monographic catalogue on the systematics and phylogeny of the South American iguanian lizard family Liolaemidae (Squamata, Iguania), pp. 1-85 in Zootaxa 1800 on page 50, DOI: 10.5281/zenodo.6789337, {"references":["Dumeril, A. M. C. & Bibron, G. (1837) Erpetologie generale ou histoire naturelle complete des reptiles, Edicion Roret, Paris, 4, 571 pp.","Cei, J. M. (1986) Reptiles del centro, centro-oeste y sur de la Argentina. Herpetofauna de las zonas aridas y semiaridas. Museo Regionale di Scienze Naturali di Torino, Monografie 4, 528 pp.","Donoso-Barros, R. (1966 a) Reptiles de Chile. Ediciones de la Universidad de Chile, Santiago, 458 pp.","Bell, T. (1843) Reptiles, Part V. In: Darwin, C. (Ed.), The Zoology of the voyage of H. M. S. Beagle, under command of Captain Fitzroy, R. N., during the years 1832 to 1836. Smith, Elder & Co., London, 1 - 55.","Guichenot, A. (1848) Reptilia. In: Gay, C. (Ed.), Historia fisica y politica de Chile. Tomo segundo, Zoologia. Casa del Autor, Paris. Museo Nacional de Historia Natural, Santiago, 18 - 136.","Dumeril, A. M. C. & Dumeril, A. (1851) Catalogue methodique de la collection des reptiles (Museum d'Histoire Naturelle de Paris), Gide et Baudry, Paris, 224 pp.","Fitzinger, L. (1843) Systema Reptilium. Fasciculus primus, Amblyglossae, Society for the Study of Amphibians and Reptiles, 282 pp.","Girard, C. (1858 a) Descriptions of some new reptiles collected by the U. S. Exploring Expedition, under the command of Captain Charles Wilkes, U. S. N., Fourth part. Proceedings of the Academy of Natural Sciences of Philadelphia, 9, 195 - 199.","Etheridge, R & Espinoza, R. E. (2000) Taxonomy of the Liolaeminae (Squamata: Iguania: Tropiduridae) and a semiannotated bibliography. Smithsonian Herpetological Information Service, 126, 1 - 64.","Girard, C. (1858 b) United States Exploring Expedition during the years 1838, 1839, 1840, 1841, 1842, under the command of Charles Wilkes, U. S. N., Volume 20, Herpetology, Philadelphia, J. B. Lippincott, 496 pp.","Boulenger, G. A. (1885 a) Catalogue of the lizards in the British Museum (Natural History), London, 497 pp.","Andersson, L. G. (1898) List of reptiles and batrachians collected by the Swedish Expedition to Tierra del Fuego (1895 - 1896) under direction of Dr. Otto Nordenskiold. Ofversigt af K. Vetenskaps-Akademiens Forhanfingar, 1898, 457 - 462.","Berg, C. (1898) Contribuciones al conocimiento de la fauna erpetologica argentina y de los paises limitrofes. Anales del Museo Nacional de Historia Natural de Buenos Aires, 6, 1 - 35.","Burt, C. E. & Burt, M. D. (1931) South American lizards in the collection of the American Museum of Natural History. Bulletin of the American Museum of Natural History, 61, 227 - 395.","Liebermann, J. (1939) Catalogo sistematico y zoogeografico de los Lacertilidos argentinos. Physis, 16, 61 - 82.","Hellmich, W. (1950) Die Eidechsen der Ausbeute Schroder (Gattung Liolaemus, Iguan.) (Beitrage zur Kenntnis der Herpetofauna Chiles XIII). Veroffentlichungen Zoologische Staatsammlung, Munchen, 1, 129 - 194.","Donoso-Barros, R. & Codoceo, M. (1962) Reptiles de Aysen y Magallanes. Boletin del Museo Nacional de Historia Natural de Chile, 28, 1 - 45.","Donoso-Barros, R. (1970 a) Liolaemus. In: Peters, J. A. & Donoso-Barros, R. (Ed.), Catalogue of the Neotropical Squamata. Part II. Lizards and Amphisbaenians. Bulletin United States National Museum, 297, 170 - 195.","Donoso-Barros, R. (1970 b) Catalogo herpetologico chileno. Boletin del Museo Nacional de Historia Natural de Chile, 31, 49 - 124.","Cei, J. M. & Scolaro, J. A. (1977 b) Herpetologia patagonica. XIV. Nuevos datos immunologicos sobre iguanidos argentinos del grupo Liolaemus fitzingeri. Physis, Buenos Aires, 37, 223 - 226.","Cei, J. M. & Scolaro, J. A. (1983 a) Un nuevo arreglo taxonomico para los Liolaemus del grupo fitzingeri. Boletin de la Asociacion Herpetologica Argentina, 1, 15 - 16.","Scolaro, J. A. & Cei, J. M. (1977) Herpetologia patagonica. XII. Los iguanidos del groupo Liolaemus fitzingeri en Chubut: datos serologicos y posicion taxonomica. Physis, Buenos Aires, 36, 219 - 223.","Nunez, H. & Jaksic, F. M. (1992) Lista comentada de los reptiles terrestres de Chile continental. Boletin del Museo Nacional de Historia Natural de Chile, 43, 63 - 91.","Pereyra, E. A., Mella, J. & Diaz, N. F. (1992) Analisis meristico preliminar de la estructura taxonomica del genero Liolaemus Wiegmann. Acta Zoologica Lilloana, 41, 277 - 286.","Etheridge, R. (1995) Redescription of Ctenoblepharys adspersa Tschudi, 1845, and the taxonomy of Liolaeminae (Reptilia: Squamata: Tropiduridae). American Museum Novitates, 3142, 1 - 34.","Chebez, J. C., Rey, N. R. & Williams, J. D. (2005) Reptiles de los parques nacionales de la Argentina. Literatura of Latin America, Buenos Aires, 37 pp.","Scolaro, J. A. (2005) Reptiles patagonicos: Sur. Una guia de campo. Ediciones Universidad Nacional de la Patagonia San Juan Bosco, Trelew, 80 p.","Sura, P. 2005. Encyclopedia of extant amphibians and reptiles, Wydawnictwo Fundacja, Nowy Sacz, 544 pp.","Abdala, C. S. (2007) Phylogeny of the boulengeri group (Iguania: Liolaemidae, Liolaemus) based on morphological and molecular characters. Zootaxa, 1538, 1 - 84.","Cei, J. M. (1973 a) Herpetologia patagonica. VI. Los Liolaemus del grupo fitzingeri en Santa Cruz y Chubut (Sauria, Iguanidae). Physis, Buenos Aires, 32, 447 - 458.","Cei, J. M. (1980 b) L'identite des syntypes de Proctotretus fitzingeri Dumeril et Bibron, 1837. Bulletin Museum National d'Historie Naturelle Paris, Ser. 4, A 2, 317 - 320.","Vanzolini, P. E. (1986) Addenda and corrigenda to the Catalogue of Neotropical Squamata. Smithsonian Herpetological Information Service, 70, 1 - 25.","Veloso, A. & Navarro, J. (1988) Lista sistematica y distribucion geografica de anfibios y reptiles de Chile. Bollettino del Museo Regionale di Scienze Naturali di Torino, 6, 481 - 539.","Brygoo, E. R. (1989) Les types d'Iguanides (Reptiles, Sauriens) du Museum national d'Histoire naturelle Catalogue critique. Bulletin Museum National d'Historie Naturelle Paris, Ser. 4, Sec. A (3): 1 - 112.","Laurent, R. F. (1992) On some overlooked species of the genus Liolaemus Wiegmann (Reptilia-Tropiduridae) from Peru. Breviora, 494, 1 - 33.","Pincheira-Donoso, D. & Nunez, H. (2005) Las especies chilenas del genero Liolaemus (Iguania, Tropiduridae, Liolaeminae). Taxonomia, sistematica y evolucion, Publicacion Ocasional del Museo Nacional de Historia Natural de Chile, Santiago, 486 pp."]}

Liolaemus cuyanus Cei & Scolaro Liolaemus fitzingeri cuyanus Cei & Scolaro 1980: 38 (type locality: 10 km E Caucete, San Juan Province, Argentina). Liolaemus cuyanus Laurent 1982c: 9; Cei & Scolaro 1983a: 15; Cei 1986: 212, 1993: 267; Vanzolini 1986: 13; Pereyra et al. 1992: 277; Cabrera 1993: 27; Etheridge 1995: 34, 2000: 331; Cruz et al. 1999: 182; Etheridge & Espinoza 2000: 5; Schulte et al. 2000: 79; Chebez et al. 2005: 30; Sura 2005: 428; Abdala 2007: 44., Published as part of Pincheira-Donoso, Daniel, Scolaro, J. Alejandro & Sura, Piotr, 2008, A monographic catalogue on the systematics and phylogeny of the South American iguanian lizard family Liolaemidae (Squamata, Iguania), pp. 1-85 in Zootaxa 1800 on page 49, DOI: 10.5281/zenodo.6789337, {"references":["Cei, J. M. & Scolaro, J. A. (1980) Two new subspecies of the Liolaemus fitzingeri complex from Argentina. Journal of Herpetology, 14, 37 - 43.","Laurent, R. F. (1982 c) Sinonimia del genero Pelusaurus Donoso-Barros con Liolaemus (Sauria, Iguanidae). Boletin de la Asociacion Herpetologica Argentina, 1, 9 - 10.","Cei, J. M. & Scolaro, J. A. (1983 a) Un nuevo arreglo taxonomico para los Liolaemus del grupo fitzingeri. Boletin de la Asociacion Herpetologica Argentina, 1, 15 - 16.","Cei, J. M. (1986) Reptiles del centro, centro-oeste y sur de la Argentina. Herpetofauna de las zonas aridas y semiaridas. Museo Regionale di Scienze Naturali di Torino, Monografie 4, 528 pp.","Cei, J. M. (1993) Reptiles del noroeste, nordeste y este de la Argentina. Herpetofauna de las selvas subtropicales, Puna y Pampas. Museo Regionale di Scienze Naturali di Torino, Monografie 14, 949 pp.","Vanzolini, P. E. (1986) Addenda and corrigenda to the Catalogue of Neotropical Squamata. Smithsonian Herpetological Information Service, 70, 1 - 25.","Pereyra, E. A., Mella, J. & Diaz, N. F. (1992) Analisis meristico preliminar de la estructura taxonomica del genero Liolaemus Wiegmann. Acta Zoologica Lilloana, 41, 277 - 286.","Cabrera, M. R. (1993) Los saurios del espinal y su presencia en provincias biogeographicas vencinas. Revista del Museo Argentino de Ciencias Naturales \" Bernardino Rivadavia \" e Instituto Nacional de Investigacion de las Ciencias Naturales, 4, 17 - 34.","Etheridge, R. (1995) Redescription of Ctenoblepharys adspersa Tschudi, 1845, and the taxonomy of Liolaeminae (Reptilia: Squamata: Tropiduridae). American Museum Novitates, 3142, 1 - 34.","Etheridge, R. (2000) A review of lizards of the Liolaemus wiegmannii group (Squamata, Iguania, Tropiduridae), and a history of morphological change in the sand-dwelling species. Herpetological Monographs, 14, 293 - 352.","Cruz, F. B., Schulte II, J. A. & Bellagamba, P. (1999) New distributional records and natural history notes for reptiles from southern Argentina. Herpetological Review, 30, 182.","Etheridge, R & Espinoza, R. E. (2000) Taxonomy of the Liolaeminae (Squamata: Iguania: Tropiduridae) and a semiannotated bibliography. Smithsonian Herpetological Information Service, 126, 1 - 64.","Schulte II, J. A., Macey, J. R., Espinoza, R. E. & Larson, A. (2000) Phylogenetic relationships in the iguanid lizard genus Liolaemus: multiple origins of viviparous reproduction and evidence for recurring Andean vicariance and dispersal. Biological Journal of the Linnean Society, 69, 75 - 102.","Chebez, J. C., Rey, N. R. & Williams, J. D. (2005) Reptiles de los parques nacionales de la Argentina. Literatura of Latin America, Buenos Aires, 37 pp.","Sura, P. 2005. Encyclopedia of extant amphibians and reptiles, Wydawnictwo Fundacja, Nowy Sacz, 544 pp.","Abdala, C. S. (2007) Phylogeny of the boulengeri group (Iguania: Liolaemidae, Liolaemus) based on morphological and molecular characters. Zootaxa, 1538, 1 - 84."]}

Liolaemus scapularis Laurent Liolaemus scapularis Laurent 1982a: 144 (type locality: Nacimiento, Catamarca Province, Argentina); Vanzolini 1986: 16; Cei 1993: 275; Ramírez-Pinilla 1994: 521; Etheridge 1995: 34, 2000: 308; Etheridge & Espinoza 2000: 9; Verrastro et al. 2003: 109; Chebez et al. 2005: 35; Sura 2005: 430; Abdala 2007: 44; Díaz 2007: 116. Liolaemus (Ortholaemus) scapularis Laurent 1984a: 370., Published as part of Pincheira-Donoso, Daniel, Scolaro, J. Alejandro & Sura, Piotr, 2008, A monographic catalogue on the systematics and phylogeny of the South American iguanian lizard family Liolaemidae (Squamata, Iguania), pp. 1-85 in Zootaxa 1800 on page 55, DOI: 10.5281/zenodo.6789337, {"references":["Laurent, R. F. (1982 a) Description de trois especes nouvelles du genre Liolaemus (Sauria, Iguanidae). Spixiana, 5, 139 - 147.","Vanzolini, P. E. (1986) Addenda and corrigenda to the Catalogue of Neotropical Squamata. Smithsonian Herpetological Information Service, 70, 1 - 25.","Cei, J. M. (1993) Reptiles del noroeste, nordeste y este de la Argentina. Herpetofauna de las selvas subtropicales, Puna y Pampas. Museo Regionale di Scienze Naturali di Torino, Monografie 14, 949 pp.","Etheridge, R. (1995) Redescription of Ctenoblepharys adspersa Tschudi, 1845, and the taxonomy of Liolaeminae (Reptilia: Squamata: Tropiduridae). American Museum Novitates, 3142, 1 - 34.","Etheridge, R. (2000) A review of lizards of the Liolaemus wiegmannii group (Squamata, Iguania, Tropiduridae), and a history of morphological change in the sand-dwelling species. Herpetological Monographs, 14, 293 - 352.","Etheridge, R & Espinoza, R. E. (2000) Taxonomy of the Liolaeminae (Squamata: Iguania: Tropiduridae) and a semiannotated bibliography. Smithsonian Herpetological Information Service, 126, 1 - 64.","Verrastro, L., Veronese, L., Bujes, C. & Martins Dias Filho, M. (2003) A new species of Liolaemus from southern Brazil (Iguania: Tropiduridae). Herpetologica, 59, 105 - 118.","Chebez, J. C., Rey, N. R. & Williams, J. D. (2005) Reptiles de los parques nacionales de la Argentina. Literatura of Latin America, Buenos Aires, 37 pp.","Sura, P. 2005. Encyclopedia of extant amphibians and reptiles, Wydawnictwo Fundacja, Nowy Sacz, 544 pp.","Abdala, C. S. (2007) Phylogeny of the boulengeri group (Iguania: Liolaemidae, Liolaemus) based on morphological and molecular characters. Zootaxa, 1538, 1 - 84.","Diaz, J. M. (2007) Reptilia, Iguania, Liolaeminae, Liolaemus, Puna, Prepuna, and mountain ranges, northwestern Argentina. Check List, 3, 105 - 118.","Laurent, R. F. (1984 a) On some iguanid genera related to or previously confused with Liolaemus Wiegmann. Journal of Herpetology, 18, 357 - 373."]}

Liolaemus olongasta Etheridge Liolaemus darwinii (sensu lato) Cei 1993: 258. Liolaemus olongasta Etheridge 1993: 167 (type locality: 51.3 km S Villa Unión, Provincial Road 26, General Lavalle Department, La Rioja Province, Argentina), 1995: 34, 2000: 331; Cei 1993: 741; Avila et al. 1998: 11; Etheridge & Espinoza 2000: 8; Chebez et al. 2005: 33; Sura 2005: 429; Abdala 2007: 44; Díaz 2007: 115., Published as part of Pincheira-Donoso, Daniel, Scolaro, J. Alejandro & Sura, Piotr, 2008, A monographic catalogue on the systematics and phylogeny of the South American iguanian lizard family Liolaemidae (Squamata, Iguania), pp. 1-85 in Zootaxa 1800 on page 52, DOI: 10.5281/zenodo.6789337, {"references":["Cei, J. M. (1993) Reptiles del noroeste, nordeste y este de la Argentina. Herpetofauna de las selvas subtropicales, Puna y Pampas. Museo Regionale di Scienze Naturali di Torino, Monografie 14, 949 pp.","Etheridge, R. (1993) Lizards of the Liolaemus darwinii Complex (Squamata: Iguania: Tropiduridae) in Northern Argentina. Bollettino del Museo Regionale di Scienze Naturali di Torino, 11, 137 - 199.","Avila, L. J., Acosta, J. C. & Murua, A. F. (1998) Herpetofauna de la provincia de San Juan, Argentina: lista comentada y distribucion geografica. Cuadernos de Herpetologia, 12, 11 - 29.","Etheridge, R & Espinoza, R. E. (2000) Taxonomy of the Liolaeminae (Squamata: Iguania: Tropiduridae) and a semiannotated bibliography. Smithsonian Herpetological Information Service, 126, 1 - 64.","Chebez, J. C., Rey, N. R. & Williams, J. D. (2005) Reptiles de los parques nacionales de la Argentina. Literatura of Latin America, Buenos Aires, 37 pp.","Sura, P. 2005. Encyclopedia of extant amphibians and reptiles, Wydawnictwo Fundacja, Nowy Sacz, 544 pp.","Abdala, C. S. (2007) Phylogeny of the boulengeri group (Iguania: Liolaemidae, Liolaemus) based on morphological and molecular characters. Zootaxa, 1538, 1 - 84.","Diaz, J. M. (2007) Reptilia, Iguania, Liolaeminae, Liolaemus, Puna, Prepuna, and mountain ranges, northwestern Argentina. Check List, 3, 105 - 118."]}

Liolaemus abaucan Etheridge Liolaemus abaucan Etheridge 1993: 165 (type locality: 12 km S Palo Blanco, 1900 mts, Provincial Road 34, Tinogasta Department, Catamarca Province, Argentina), 1995: 34; Cei 1993: 738; Etheridge & Espinoza 2000: 4; Chebez et al. 2005: 28; Sura 2005: 428; Abdala 2007: 44; Díaz 2007: 114. Liolaemus darwinii (sensu lato) Cei 1993: 258., Published as part of Pincheira-Donoso, Daniel, Scolaro, J. Alejandro & Sura, Piotr, 2008, A monographic catalogue on the systematics and phylogeny of the South American iguanian lizard family Liolaemidae (Squamata, Iguania), pp. 1-85 in Zootaxa 1800 on page 48, DOI: 10.5281/zenodo.6789337, {"references":["Etheridge, R. (1993) Lizards of the Liolaemus darwinii Complex (Squamata: Iguania: Tropiduridae) in Northern Argentina. Bollettino del Museo Regionale di Scienze Naturali di Torino, 11, 137 - 199.","Cei, J. M. (1993) Reptiles del noroeste, nordeste y este de la Argentina. Herpetofauna de las selvas subtropicales, Puna y Pampas. Museo Regionale di Scienze Naturali di Torino, Monografie 14, 949 pp.","Etheridge, R & Espinoza, R. E. (2000) Taxonomy of the Liolaeminae (Squamata: Iguania: Tropiduridae) and a semiannotated bibliography. Smithsonian Herpetological Information Service, 126, 1 - 64.","Chebez, J. C., Rey, N. R. & Williams, J. D. (2005) Reptiles de los parques nacionales de la Argentina. Literatura of Latin America, Buenos Aires, 37 pp.","Sura, P. 2005. Encyclopedia of extant amphibians and reptiles, Wydawnictwo Fundacja, Nowy Sacz, 544 pp.","Abdala, C. S. (2007) Phylogeny of the boulengeri group (Iguania: Liolaemidae, Liolaemus) based on morphological and molecular characters. Zootaxa, 1538, 1 - 84.","Diaz, J. M. (2007) Reptilia, Iguania, Liolaeminae, Liolaemus, Puna, Prepuna, and mountain ranges, northwestern Argentina. Check List, 3, 105 - 118."]}

Liolaemus dorbignyi Koslowsky Liolaemus d’orbignyi Koslowsky1898: 174 (type locality: Catamarca Province, Argentina). Liolaemus dorbignyi Donoso-Barros 1970a: 182; Vanzolini 1986: 14; Laurent 1992: 31; Etheridge 1995: 34; Etheridge & Espinoza 2000: 5; Chebez et al. 2005: 30; Sura 2005: 428; Díaz 2007: 118; Quinteros et al. 2008: 52. Liolaemus dorbigni Cei 1993: 239. Liolaemus dorbigny Díaz 2007: 117., Published as part of Pincheira-Donoso, Daniel, Scolaro, J. Alejandro & Sura, Piotr, 2008, A monographic catalogue on the systematics and phylogeny of the South American iguanian lizard family Liolaemidae (Squamata, Iguania), pp. 1-85 in Zootaxa 1800 on page 39, DOI: 10.5281/zenodo.6789337, {"references":["Koslowsky, J. (1898) Enumeracion sistematica y distribucion geografica de los reptiles argentinos. Revista del Museo de La Plata, 8, 161 - 200.","Donoso-Barros, R. (1970 a) Liolaemus. In: Peters, J. A. & Donoso-Barros, R. (Ed.), Catalogue of the Neotropical Squamata. Part II. Lizards and Amphisbaenians. Bulletin United States National Museum, 297, 170 - 195.","Vanzolini, P. E. (1986) Addenda and corrigenda to the Catalogue of Neotropical Squamata. Smithsonian Herpetological Information Service, 70, 1 - 25.","Laurent, R. F. (1992) On some overlooked species of the genus Liolaemus Wiegmann (Reptilia-Tropiduridae) from Peru. Breviora, 494, 1 - 33.","Etheridge, R. (1995) Redescription of Ctenoblepharys adspersa Tschudi, 1845, and the taxonomy of Liolaeminae (Reptilia: Squamata: Tropiduridae). American Museum Novitates, 3142, 1 - 34.","Etheridge, R & Espinoza, R. E. (2000) Taxonomy of the Liolaeminae (Squamata: Iguania: Tropiduridae) and a semiannotated bibliography. Smithsonian Herpetological Information Service, 126, 1 - 64.","Chebez, J. C., Rey, N. R. & Williams, J. D. (2005) Reptiles de los parques nacionales de la Argentina. Literatura of Latin America, Buenos Aires, 37 pp.","Sura, P. 2005. Encyclopedia of extant amphibians and reptiles, Wydawnictwo Fundacja, Nowy Sacz, 544 pp.","Diaz, J. M. (2007) Reptilia, Iguania, Liolaeminae, Liolaemus, Puna, Prepuna, and mountain ranges, northwestern Argentina. Check List, 3, 105 - 118.","Quinteros, A. S., Abdala, C. S. & Lobo, F. J. (2008) Redescription of Liolaemus dorbignyi Koslowskyi, 1898 and description of a new species of Liolaemus (Iguania: Liolaemidae). Zootaxa, 1717, 51 - 67.","Cei, J. M. (1993) Reptiles del noroeste, nordeste y este de la Argentina. Herpetofauna de las selvas subtropicales, Puna y Pampas. Museo Regionale di Scienze Naturali di Torino, Monografie 14, 949 pp."]}

Liolaemus wiegmannii (Duméril & Bibron) Proctotretus wiegmannii Duméril & Bibron 1837: 284 (type locality: “ Chili ”; probably Carmen del Patagon, about 35 km northwest of the mouth of the Rio Negro, Rio Negro Province, Argentina; see Etheridge 2000); Bell 1843: 15; Duméril & Duméril 1851: 74; Burmeister 1861: 525. Ptychodeira wiegmanni Fitzinger 1843: 73. Leiolaemus wiegmannii Gray 1845: 216. Rhytidodeira wiegmanni Girard 1858a: 198, 1858b: 352. Ortholaemus fitzroii Girard 1858a: 198 (substitute name for wiegmannii, Bell 1843; see also Etheridge & Espinoza 2000). Liolaemus (Ortholaemus) fitzroyi Girard 1858b: 373. Ortholaemus fitzroyi Cope 1862: 351. Liolaemus (Ptychodeira) wiegmanni Steindachner 1867: 37. Liolaemus wiegmannii Boulenger 1885a: 156; Boettger 1893: 61; Amaral 1938: 179; Loveridge 1959: 69; Gallardo 1969: 75, 1977, 1980: 346; Donoso-Barros 1970a: 195; Müller & Steiniger 1978: 429; Cei 1979c: 1, 1986: 198, 1993: 270; Rand 1982: 173; Pereyra et al. 1992: 277; Cruz et al. 1993: 101; Tiranti & Avila 1994: 40; Etheridge 1995: 34, 2000: 310; Martori & Aun 1995: 94; Acosta et al. 1996a: 74; Cabrera 1996: 219; Fitzgerald et al. 1999: 536; Etheridge & Espinoza 2000: 9; Verrastro et al. 2003: 109; Chebez et al. 2005: 36; Sura 2005: 430; Scolaro 2006: 51; Abdala 2007: 44. Liolaemus wiegmani Liebermann 1939: 71. Liolaemus (Ortholaemus) wiegmanni Laurent 1984a: 370; Cabrera & Bee de Speroni 1986: 6., Published as part of Pincheira-Donoso, Daniel, Scolaro, J. Alejandro & Sura, Piotr, 2008, A monographic catalogue on the systematics and phylogeny of the South American iguanian lizard family Liolaemidae (Squamata, Iguania), pp. 1-85 in Zootaxa 1800 on pages 55-56, DOI: 10.5281/zenodo.6789337, {"references":["Dumeril, A. M. C. & Bibron, G. (1837) Erpetologie generale ou histoire naturelle complete des reptiles, Edicion Roret, Paris, 4, 571 pp.","Etheridge, R. (2000) A review of lizards of the Liolaemus wiegmannii group (Squamata, Iguania, Tropiduridae), and a history of morphological change in the sand-dwelling species. Herpetological Monographs, 14, 293 - 352.","Bell, T. (1843) Reptiles, Part V. In: Darwin, C. (Ed.), The Zoology of the voyage of H. M. S. Beagle, under command of Captain Fitzroy, R. N., during the years 1832 to 1836. Smith, Elder & Co., London, 1 - 55.","Dumeril, A. M. C. & Dumeril, A. (1851) Catalogue methodique de la collection des reptiles (Museum d'Histoire Naturelle de Paris), Gide et Baudry, Paris, 224 pp.","Burmeister, H. (1861) Reise durch die La Plata Staaten mit besonderer Rucksicht auf die physische Beschaffenheit und den Culturzustand der Argentinische Republik. Ausgefuhrt in den Jahren 1857, 1858, 1859 und 1860, Halle, H. W. Schmidt, 538 pp.","Fitzinger, L. (1843) Systema Reptilium. Fasciculus primus, Amblyglossae, Society for the Study of Amphibians and Reptiles, 282 pp.","Gray, J. E. (1845) Catalogue of the specimens of lizards in the collection of the British Museum, Trustees of the British Museum, London, 289 pp.","Girard, C. (1858 a) Descriptions of some new reptiles collected by the U. S. Exploring Expedition, under the command of Captain Charles Wilkes, U. S. N., Fourth part. Proceedings of the Academy of Natural Sciences of Philadelphia, 9, 195 - 199.","Girard, C. (1858 b) United States Exploring Expedition during the years 1838, 1839, 1840, 1841, 1842, under the command of Charles Wilkes, U. S. N., Volume 20, Herpetology, Philadelphia, J. B. Lippincott, 496 pp.","Etheridge, R & Espinoza, R. E. (2000) Taxonomy of the Liolaeminae (Squamata: Iguania: Tropiduridae) and a semiannotated bibliography. Smithsonian Herpetological Information Service, 126, 1 - 64.","Cope, E. D. (1862) Catalogues of the reptiles obtained during the explorations of the Parana, Paraguay, Bermejo and Uruguay Rivers by Capt. Thos. J. Page, U. S. N., and of those procured by Lieut. N. Michler, U. S. Top. Eng., commander of the expedition conducting the survey of the Atrato River. Proceedings of the Academy of Natural Sciences of Philadelphia, 14, 346 - 359.","Steindachner, F. (1867) Reise der osterreichischen Fregatte Novara um die Erde in den Jahren 1857, 1858, 1859. Zoologischer Theil. Erster Band (Wirbelthiere). 3. Reptilien, K. K. Hof und Staatsdruckerei, Wien, 98 pp.","Boulenger, G. A. (1885 a) Catalogue of the lizards in the British Museum (Natural History), London, 497 pp.","Boettger, O. (1893) Katalog der Reptilien-Sammlung im Museum der Senckenbergischen Naturforschended Gesellschaft im Frankfurt am Maim. I. Teir (Rhynchocephalen, Shildkroten, Krocodile, Eidechsen, Chamaleons). Gebruder, Knauer, Frankfurt, 1 - 140.","Amaral, A. (1938) Estudos sobre lacertilios neotropicos. 4. Lista remissiva dos lacertilios do Brasil. Memorias del Instituto Butantan, 11, 167 - 204.","Loveridge, A. (1959) Notes on the present herpetofauna of Ascension Island. Copeia, 1959, 69 - 70.","Gallardo, J. M. (1969) Las especies de saurios (Reptilia) de la provincia de Santa Fe, Argentina, y consideraciones sobre su ecologia y zoogeografia. Neotropica, 15, 73 - 81.","Gallardo, J. M. (1977) Reptiles de los alrededores de Buenos Aires, Editorial Universitaria, Buenos Aires, 213 pp.","Gallardo, J. M. (1980) Estudio ecologico sobre anfibios y reptiles del noroeste de la provincia de Buenos Aires (Argentina). I Reunion Iberoamericano de Zoologia Vertebrado, La Rabida, 1980, 331 - 349.","Donoso-Barros, R. (1970 a) Liolaemus. In: Peters, J. A. & Donoso-Barros, R. (Ed.), Catalogue of the Neotropical Squamata. Part II. Lizards and Amphisbaenians. Bulletin United States National Museum, 297, 170 - 195.","Muller, P. & Steiniger, H. (1978) Chorologie und Differenzierungsgeschwindigkeit brasilienianischer Liolaemus Arten (Sauria-Iguainidae). Verhandlungen Gesselshaft fur Okologie, 7, 429 - 440.","Cei, J. M. (1979 c) Nota preliminar sobre la distribucion geografica de Liolaemus wiegmanni (Dum. et Bibr.) (Sauria, Iguanidae). Publicaciones Ocasionales del Instituto de Biologia Animal, Universidad Nacional de Cuyo, Serie Cientifica, 14, 1 - 4.","Cei, J. M. (1986) Reptiles del centro, centro-oeste y sur de la Argentina. Herpetofauna de las zonas aridas y semiaridas. Museo Regionale di Scienze Naturali di Torino, Monografie 4, 528 pp.","Cei, J. M. (1993) Reptiles del noroeste, nordeste y este de la Argentina. Herpetofauna de las selvas subtropicales, Puna y Pampas. Museo Regionale di Scienze Naturali di Torino, Monografie 14, 949 pp.","Rand, A. S. (1982) Clutch size in Brazilian iguanid lizards. Herpetologica, 38, 171 - 178.","Pereyra, E. A., Mella, J. & Diaz, N. F. (1992) Analisis meristico preliminar de la estructura taxonomica del genero Liolaemus Wiegmann. Acta Zoologica Lilloana, 41, 277 - 286.","Cruz, F. B., Perotti, M. G. & Fitzgerald, L. A. (1993) Lista de anfibios y reptiles colectados en una localidad del chaco Salteno. Acta Zoologica Lilloana, 42, 101 - 107.","Tiranti, S. & Avila, L. J. (1994) Aportes a la herpetologia de la provincia de La Pampa. I. Lista de los taxa de presencia segura en la provincia. Boletin de la Asociacion Herpetologica Argentina, 10, 39 - 41.","Etheridge, R. (1995) Redescription of Ctenoblepharys adspersa Tschudi, 1845, and the taxonomy of Liolaeminae (Reptilia: Squamata: Tropiduridae). American Museum Novitates, 3142, 1 - 34.","Martori, R. & Aun, L. (1995) Relevamiento de la herpetofauna de las Sierras Pampeanas del este en la provincia de Cordoba (Argentina) (registro de taxa y comentarios taxonomicos). Acta Zoologica Lilloana, 43, 87 - 97.","Acosta, J. C., Murua, F. & Ortiz, G. (1996 a) Primera cita de Liolaemus wiegmanni (Sauria: Tropiduridae) para la provincia de San Juan, Argentina. Cuadernos de Herpetologia, 10, 74 - 75.","Cabrera, M. R. (1996) Lista y distribucion geografica de saurios, anfisbenas y tortugas (Amniota, Reptilia) de la provincia de Cordoba. In: di Tada, I. E. & Bucher, E. H. (Ed.), Biodiversidad de Provincia Cordoba. I. Fauna. Universidad Nacional de Rio Cuarto, Rio Cuarto, Cordoba, 215 - 238.","Fitzgerald, L. A., Cruz, F. B. & Perotti, G. (1999) Phenology of a lizard assemblage in the dry Chaco of Argentina. Journal of Herpetology, 33, 536 - 535.","Verrastro, L., Veronese, L., Bujes, C. & Martins Dias Filho, M. (2003) A new species of Liolaemus from southern Brazil (Iguania: Tropiduridae). Herpetologica, 59, 105 - 118.","Chebez, J. C., Rey, N. R. & Williams, J. D. (2005) Reptiles de los parques nacionales de la Argentina. Literatura of Latin America, Buenos Aires, 37 pp.","Sura, P. 2005. Encyclopedia of extant amphibians and reptiles, Wydawnictwo Fundacja, Nowy Sacz, 544 pp.","Scolaro, J. A. (2006) Reptiles patagonicos: Norte. Una guia de campo, Ediciones Universidad Nacional de la Patagonia San Juan Bosco, Comodoro Rivadavia, 112 p.","Abdala, C. S. (2007) Phylogeny of the boulengeri group (Iguania: Liolaemidae, Liolaemus) based on morphological and molecular characters. Zootaxa, 1538, 1 - 84.","Liebermann, J. (1939) Catalogo sistematico y zoogeografico de los Lacertilidos argentinos. Physis, 16, 61 - 82.","Laurent, R. F. (1984 a) On some iguanid genera related to or previously confused with Liolaemus Wiegmann. Journal of Herpetology, 18, 357 - 373.","Cabrera, M. R. & Bee de Speroni, N. T. (1986) Composicion y distribucion de la lacertofauna de la provincia de Cordoba, Argentina. II. Amphisbaenidae, Anguidae e Iguanidae. Historia Natural en Corrientes, Argentina, 6, 1 - 12."]}

Phymaturus genus Most phylogenetic information available on Phymaturus species has commonly been provided in studies focusing on other lizard clades (often Liolaemus), in which a limited number of taxa belonging to this genus are used as outgroups (e.g. Schulte et al. 2000; Valladares et al. 2002; Díaz & Lobo 2006; Pincheira-Donoso et al. 2007a). The first phylogenetic hypothesis to include a proportionally large number of Phymaturus species was recently developed by Espinoza et al. (2004). In a comparative analysis exploring the evolution of trophic niche in Liolaemidae lizards, these authors reconstructed, on the basis of both morphological and molecular data, the phylogenetic relationships between seven Phymaturus taxa (when only 12 of the current 19 species were recognized; see below). A major achievement of Espinoza et al. ’s (2004) results was that they provided the first support to the hypothesis that these robust lizards are grouped into two main clades (Fig. 5), one primarily austral and Patagonian (patagonicus clade) and one essentially boreal and Andean (flagellifer clade = palluma clade; see Etheridge & Savage 2003 and Cei & Scolaro 2006, for nomenclatural details) as had previously been proposed by Etheridge (1995; see also Cei & Castro 1973; Cei 1986). More recently, Lobo & Quinteros (2005b) presented the only known explicit study focused on the phylogenetic relationships within the genus Phymaturus (Fig. 5). Lobo & Quinteros’s (2005b) phylogenetic hypothesis, entirely based on morphological data, incorporated the analysis of 15 Phymaturus species, when a total of 17 were known (in the same work, these authors provided the description of four new taxa; see below for details). Only P. calcogaster (Scolaro & Cei 2003), P. verdugo (Cei & Videla 2003b) and P. vociferator (Pincheira-Donoso 2004a) were excluded from that study (although P. dorsimaculatus – included in that study – and P. vociferator appear to be conspecific; see below). Even though Lobo & Quinteros’s (2005b) phylogeny differs importantly in some aspects from Espinoza et al. ’s (2004) tree (Fig. 5), it also supported substantially the hypothesis of two major clades forming the genus Phymaturus. In both phylogenetic hypotheses, the same Patagonian and Andean groups previously mentioned were found as independent evolutionary lines, consisting of the same subset of species (Espinoza et al. 2004; Lobo & Quinteros 2005b; Fig. 4). These recent attempts to clarify the phylogenetic relationships among Phymaturus species represent substantial progress for future phylogenetic-based systematic and evolutionary research (e.g. see Wiens 2004, for details on the value of morphological based phylogenies). Nevertheless, it is also worth noting that further studies are still needed to provide additional evidence on this matter. Indeed, different authors have recognized some of the limitations that morphological-based phylogenies may imply (Pimentel & Riggins 1987; Campbell & Frost 1993; Thiele 1993; Pleijel 1995; Wiens 1995, 1998, 2001; Strong & Lipscomb 1999; Scotland et al. 2003). For example, whereas delimitation and definition of morphological character states are difficult and sometimes arbitrary, this is not a problem for molecular variables, whose delimitation and definition is practically automatic (Wiens 2001). Consequently, the elaboration of Phymaturus phylogenetic hypotheses based on molecular dataset are desirable. In this work, we recognize within the Liolaemidae family the main clades and groups detailed in figs. 2, 3, 4 and 5, and as follows Family Liolaemidae Frost & Etheridge 1989 (family status according to Frost et al. 2001) Genus Ctenoblepharys Tschudi Genus Liolaemus Wiegmann Subgenus Liolaemus Wiegmann (clade chiliensis) Series signifer (see Pincheira-Donoso & Núñez 2005; also Etheridge 1995; Schulte et al. 2000; Espinoza et al. 2004) Subgenus Donosolaemus Pincheira-Donoso & Núñez (2005) (clade archeforus-kingii) Line magellanicus Subgenus Vilcunia Donoso-Barros & Cei (1971) (clade lineomaculatus) Subgenus Eulaemus Girard (Schulte et al. 2000; Pincheira-Donoso & Núñez 2005) Clade montanus (Etheridge 1995; Schulte et al. 2000) Group reichei (Núñez et al. 1998, 2003) Clade anomalus (see Espinoza et al. 2004; Abdala 2007) Complex boulengeri (Etheridge 1995, 2000) Clade fitzingerii (Etheridge 1995; Espinoza et al. 2004; Pincheira-Donoso & Núñez 2005) Clade wiegmannii (Etheridge 1995, 2000; Cruz et al. 2005) Genus Phymaturus Gravenhorst Clade flagellifer (Etheridge 1995; Lobo & Quinteros 2005b; Cei & Scolaro 2006) Clade patagonicus (Etheridge 1995; Lobo & Quinteros 2005b), Published as part of Pincheira-Donoso, Daniel, Scolaro, J. Alejandro & Sura, Piotr, 2008, A monographic catalogue on the systematics and phylogeny of the South American iguanian lizard family Liolaemidae (Squamata, Iguania), pp. 1-85 in Zootaxa 1800 on pages 17-18, DOI: 10.5281/zenodo.6789337, {"references":["Schulte II, J. A., Macey, J. R., Espinoza, R. E. & Larson, A. (2000) Phylogenetic relationships in the iguanid lizard genus Liolaemus: multiple origins of viviparous reproduction and evidence for recurring Andean vicariance and dispersal. Biological Journal of the Linnean Society, 69, 75 - 102.","Valladares, J. P., Etheridge, R., Schulte II, J. A., Manriquez, G. & Spotorno, A. (2002). Nueva especie de lagartija del norte de Chile, Liolaemus molinai (Reptilia: Liolaeminae). Revista Chilena de Historia Natural, 75, 473 - 489.","Diaz, J. M. & Lobo, F. (2006) Historical biogeography of a clade of Liolaemus (Iguania: Liolaemidae) based on ancestral areas and dispersal-vicariance analysis (DIVA). Papeis Avulsos de Zoologia, 46, 261 - 274.","Pincheira-Donoso, D., Scolaro, J. A. & Schulte II, J. A. (2007 a) The limits of polymorphism in Liolaemus rothi: molecular and phenotypic evidence for a new species of the Liolaemus boulengeri clade (Iguanidae, Liolaemini) from boreal Patagonia of Chile. Zootaxa, 1452, 25 - 42.","Espinoza, R. E., Wiens, J. J. & Tracy, C. R. (2004) Recurrent evolution of herbivory in small, cold-climate lizards: Breaking the ecophysiological rules or reptilian herbivory. Proceedings of the National Academy of Sciences, USA, 101, 16819 - 16824.","Etheridge, R. & Savage, J. M. (2003) Phymaturus Gravenhorst, 1837 and Lacerta palluma Molina, 1782 (currently Phymaturus palluma; Reptilia, Sauria): proposed conservation of usage of the names by designation of a neotype for Lacerta palluma Molina, 1782. Bulletin of Zoological Nomenclature, 60, 38 - 41.","Scolaro, J. A. & Cei, J. M. (2006) A new species of Liolaemus from central steppes of Chubut, Patagonia Argentina (Reptilia: Iguania: Iguanidae). Zootaxa, 1133, 61 - 68.","Etheridge, R. (1995) Redescription of Ctenoblepharys adspersa Tschudi, 1845, and the taxonomy of Liolaeminae (Reptilia: Squamata: Tropiduridae). American Museum Novitates, 3142, 1 - 34.","Cei, J. M. & Castro, L. P. (1973) Taxonomic and serological researches on the Phymaturus patagonicus complex. Journal of Herpetology, 7, 237 - 247.","Cei, J. M. (1986) Reptiles del centro, centro-oeste y sur de la Argentina. Herpetofauna de las zonas aridas y semiaridas. Museo Regionale di Scienze Naturali di Torino, Monografie 4, 528 pp.","Lobo, F. & Quinteros, S. (2005 b) A morphology-based phylogeny of Phymaturus (Iguania: Liolaemidae) with the description of four new species from Argentina. Papeis Avulsos de Zoologia, 45, 143 - 177.","Scolaro, J. A. & Cei, J. M. (2003) Una excepcional nueva especie de Phymaturus de la precordillera de Chubut, Argentina (Liolaemidae, Iguania, Lacertilia, Reptilia). Facena, 19, 107 - 112.","Cei, J. M. & Videla, F. (2003 b) A new Phymaturus species from volcanic cordilleran mountains of the southwestern Mendoza Province, Argentina (Liolaemidae, Iguania, Lacertilia, Reptilia). Bolletino Museo Regionale di Scienze Naturali di Torino, 20, 291 - 314.","Pincheira-Donoso, D. (2004 a) Una nueva especie del genero Phymaturus (Iguania: Tropiduridae: Liolaemini) del centrosur de Chile. Multequina, Latin American Journal of Natural Resources, 13, 57 - 70.","Wiens, J. J. (2004) The role of morphological data in phylogeny reconstruction. Systematic Biology, 53, 653 - 661.","Pimentel, R. & Riggins, R. (1987) The nature of cladistic data. Cladistics, 3, 201 - 209.","Campbell, J. A. & Frost, D. R. (1993) Anguid lizards of the genus Abronia: revisionary notes, descriptions of four new species, phylogenetic analysis, and key. Bulletin of the American Museum of Natural History, 216, 1 - 121.","Thiele, K. (1993) The Holy Grail of the perfect character: the cladistic treatment of morphometric data. Cladistics, 9, 275 - 304.","Pleijel, F. (1995) On character coding for phylogeny reconstruction. Cladistics, 11, 309 - 315.","Wiens, J. J. (1995) Polymorphic characters in phylogenetic systematics. Systematic Biology, 44, 482 - 500.","Wiens, J. J. (1998) Testing phylogenetic methods with tree-congruence: phylogenetic analysis of polymorphic morphological characters in phrynosomatid lizards. Systematic Biology, 47, 411 - 428.","Wiens, J. J. (2001). Character analysis in morphological phylogenetics: problems and solutions. Systematic Biology, 50, 689 - 699.","Strong, E. E. & Lipscomb, D. (1999) Charactercoding and inapplicable data. Cladistics, 15, 363 - 371.","Scotland, R. W., Olmstead, R. G. & Bennett, J. R. (2003) Phylogeny reconstruction: the role of morphology. Systematic Biology, 52, 539 - 548.","Frost, D. R. & Etheridge, R. (1989) A phylogenetic analysis and taxonomy of iguanian lizards (Reptilia: Squamata). University of Kansas Museum Natural History, Miscellaneous Publications, 81, 1 - 65.","Frost, D. R., Etheridge, R., Janies, D. & Titus, T. A. (2001) Total evidence, sequence alignment, evolution of Polychrotid lizards, and a reclassification of the Iguania (Squamata: Iguania). American Museum Novitates, 3343, 1 - 38.","Pincheira-Donoso, D. & Nunez, H. (2005) Las especies chilenas del genero Liolaemus (Iguania, Tropiduridae, Liolaeminae). Taxonomia, sistematica y evolucion, Publicacion Ocasional del Museo Nacional de Historia Natural de Chile, Santiago, 486 pp.","Donoso-Barros, R. & Cei, J. M. (1971) New lizards from the volcanic Patagonian plateau of Argentina. Journal of Herpetology, 5, 89 - 95.","Nunez, H., Navarro, J., Sufan, J. & Galaz, J. L. (1998) Distribucion geografica de Phrynosaura (Reptilia, Tropiduridae). Noticiario Mensual Museo Nacional de Historia Natural, 334, 11 - 14.","Nunez, H., Navarro, J., Garin, C., Pincheira-Donoso, D. & V. Meriggio, D. (2003) Phrynosaura manueli y Phrynosaura torresi, nuevas especies de lagartijas para el norte de Chile (Squamata: Sauria). Boletin del Museo Nacional de Historia Natural de Chile, 52, 67 - 88.","Abdala, C. S. (2007) Phylogeny of the boulengeri group (Iguania: Liolaemidae, Liolaemus) based on morphological and molecular characters. Zootaxa, 1538, 1 - 84.","Etheridge, R. (2000) A review of lizards of the Liolaemus wiegmannii group (Squamata, Iguania, Tropiduridae), and a history of morphological change in the sand-dwelling species. Herpetological Monographs, 14, 293 - 352.","Cruz, F. B., Fitzgerald, L. A., Espinoza, R. E. & Schulte II, J. A. (2005). The importance of phylogenetic scale in tests of Bergmann's and Rapoport's rules: lessons from a clade of South American lizards. Journal of Evolutionary Biology, 18, 1559 - 1574."]}

Liolaemus tehuelche Abdala Liolaemus tehuelche Abdala 2003: 5 (type locality: Cerro Alto, on National Road 40, 54 km NW Pilcaniyeu, Pilcaniyeu Department, [40°44.074’S, 70°34.696’W], 1014 m, Río Negro Province, Argentina); 2007: 44; Scolaro 2006: 49; Pincheira-Donoso et al. 2007a: 30., Published as part of Pincheira-Donoso, Daniel, Scolaro, J. Alejandro & Sura, Piotr, 2008, A monographic catalogue on the systematics and phylogeny of the South American iguanian lizard family Liolaemidae (Squamata, Iguania), pp. 1-85 in Zootaxa 1800 on page 53, DOI: 10.5281/zenodo.6789337, {"references":["Abdala, C. S. (2003) Cuatro nuevas especies del genero Liolaemus (Iguania: Liolaemidae), pertenecientes al grupo boulengeri, de la Patagonia, Argentina. Cuadernos de Herpetologia, 17, 3 - 32.","Scolaro, J. A. (2006) Reptiles patagonicos: Norte. Una guia de campo, Ediciones Universidad Nacional de la Patagonia San Juan Bosco, Comodoro Rivadavia, 112 p.","Pincheira-Donoso, D., Scolaro, J. A. & Schulte II, J. A. (2007 a) The limits of polymorphism in Liolaemus rothi: molecular and phenotypic evidence for a new species of the Liolaemus boulengeri clade (Iguanidae, Liolaemini) from boreal Patagonia of Chile. Zootaxa, 1452, 25 - 42."]}

Liolaemus mapuche Abdala Liolaemus mapuche Abdala 2002: 5 (type locality: 15 km S Paso de Indios, Provincial Road 10, Zapala Department, Neuquen Province, Argentina), 2007: 44; Chebez et al. 2005: 33; Sura 2005: 429; Scolaro 2006: 40., Published as part of Pincheira-Donoso, Daniel, Scolaro, J. Alejandro & Sura, Piotr, 2008, A monographic catalogue on the systematics and phylogeny of the South American iguanian lizard family Liolaemidae (Squamata, Iguania), pp. 1-85 in Zootaxa 1800 on page 51, DOI: 10.5281/zenodo.6789337, {"references":["Abdala, C. S. (2002) Nuevo Liolaemus (Iguania: Liolaemidae) perteneciente al grupo boulengeri de la Provincia de Neuquen, Argentina. Cuadernos de Herpetologia, 16, 3 - 13.","Chebez, J. C., Rey, N. R. & Williams, J. D. (2005) Reptiles de los parques nacionales de la Argentina. Literatura of Latin America, Buenos Aires, 37 pp.","Sura, P. 2005. Encyclopedia of extant amphibians and reptiles, Wydawnictwo Fundacja, Nowy Sacz, 544 pp.","Scolaro, J. A. (2006) Reptiles patagonicos: Norte. Una guia de campo, Ediciones Universidad Nacional de la Patagonia San Juan Bosco, Comodoro Rivadavia, 112 p."]}

Liolaemus ditadai Cei Liolaemus anomalus ditadai Cei 1983: 193 (type locality: Salinas Grandes, 4 km from the limit between Córdoba and Catamarca, Argentina), 1986: 202, 1993: 293; Etheridge 1995: 34; Etheridge & Espinoza 2000: 4. Ceiolaemus anomalus ditadai Cabrera & Bee de Speroni 1986: 5; Cabrera 1996: 219. Liolaemus ditadai Abdala 2007: 39. Observations: After morphological comparisons, Abdala (2007: 39) suggested that L. anomalus ditadai should be given full species status (see above comments for L. anomalus)., Published as part of Pincheira-Donoso, Daniel, Scolaro, J. Alejandro & Sura, Piotr, 2008, A monographic catalogue on the systematics and phylogeny of the South American iguanian lizard family Liolaemidae (Squamata, Iguania), pp. 1-85 in Zootaxa 1800 on page 47, DOI: 10.5281/zenodo.6789337, {"references":["Cei, J. M. (1983) Una nueva subespecie de Liolaemus anomalus de la region arida halofila de Salinas Grandes (Provincia de Cordoba, Argentina). Deserta, 7, 172 - 178.","Etheridge, R. (1995) Redescription of Ctenoblepharys adspersa Tschudi, 1845, and the taxonomy of Liolaeminae (Reptilia: Squamata: Tropiduridae). American Museum Novitates, 3142, 1 - 34.","Etheridge, R & Espinoza, R. E. (2000) Taxonomy of the Liolaeminae (Squamata: Iguania: Tropiduridae) and a semiannotated bibliography. Smithsonian Herpetological Information Service, 126, 1 - 64.","Cabrera, M. R. & Bee de Speroni, N. T. (1986) Composicion y distribucion de la lacertofauna de la provincia de Cordoba, Argentina. II. Amphisbaenidae, Anguidae e Iguanidae. Historia Natural en Corrientes, Argentina, 6, 1 - 12.","Cabrera, M. R. (1996) Lista y distribucion geografica de saurios, anfisbenas y tortugas (Amniota, Reptilia) de la provincia de Cordoba. In: di Tada, I. E. & Bucher, E. H. (Ed.), Biodiversidad de Provincia Cordoba. I. Fauna. Universidad Nacional de Rio Cuarto, Rio Cuarto, Cordoba, 215 - 238.","Abdala, C. S. (2007) Phylogeny of the boulengeri group (Iguania: Liolaemidae, Liolaemus) based on morphological and molecular characters. Zootaxa, 1538, 1 - 84."]}

Liolaemus quilmes Etheridge Liolaemus darwinii (sensu lato) Cei 1993: 258. Liolaemus quilmes Etheridge 1993: 152 (terra typica: 3,2 km S Animaná, 1695 m, National Road 40, Cafayate Department, Salta Province, Argentina), 1995: 34, 2000: 331; Cei 1993: 734; Etheridge & Espinoza 2000: 8; Chebez et al. 2005: 34; Sura 2005: 429; Abdala 2007: 44; Díaz 2007: 115., Published as part of Pincheira-Donoso, Daniel, Scolaro, J. Alejandro & Sura, Piotr, 2008, A monographic catalogue on the systematics and phylogeny of the South American iguanian lizard family Liolaemidae (Squamata, Iguania), pp. 1-85 in Zootaxa 1800 on page 52, DOI: 10.5281/zenodo.6789337, {"references":["Cei, J. M. (1993) Reptiles del noroeste, nordeste y este de la Argentina. Herpetofauna de las selvas subtropicales, Puna y Pampas. Museo Regionale di Scienze Naturali di Torino, Monografie 14, 949 pp.","Etheridge, R. (1993) Lizards of the Liolaemus darwinii Complex (Squamata: Iguania: Tropiduridae) in Northern Argentina. Bollettino del Museo Regionale di Scienze Naturali di Torino, 11, 137 - 199.","Etheridge, R & Espinoza, R. E. (2000) Taxonomy of the Liolaeminae (Squamata: Iguania: Tropiduridae) and a semiannotated bibliography. Smithsonian Herpetological Information Service, 126, 1 - 64.","Chebez, J. C., Rey, N. R. & Williams, J. D. (2005) Reptiles de los parques nacionales de la Argentina. Literatura of Latin America, Buenos Aires, 37 pp.","Sura, P. 2005. Encyclopedia of extant amphibians and reptiles, Wydawnictwo Fundacja, Nowy Sacz, 544 pp.","Abdala, C. S. (2007) Phylogeny of the boulengeri group (Iguania: Liolaemidae, Liolaemus) based on morphological and molecular characters. Zootaxa, 1538, 1 - 84.","Diaz, J. M. (2007) Reptilia, Iguania, Liolaeminae, Liolaemus, Puna, Prepuna, and mountain ranges, northwestern Argentina. Check List, 3, 105 - 118."]}

Liolaemus laurenti Etheridge Liolaemus darwini Cei 1986: 218 (in part). Liolaemus laurenti Etheridge 1992: 2 (type locality: 12 km E Plaza Central de Pituíl, Provincial Road 11, Famatina Department, Catamarca Province, Argentina), 1993: 179, 1995: 34, 2000: 331; Cei 1993: 262; Avila et al. 1998: 11; Etheridge & Espinoza 2000: 6; Chebez et al. 2005: 32; Sura 2005: 429; Abdala 2007: 44., Published as part of Pincheira-Donoso, Daniel, Scolaro, J. Alejandro & Sura, Piotr, 2008, A monographic catalogue on the systematics and phylogeny of the South American iguanian lizard family Liolaemidae (Squamata, Iguania), pp. 1-85 in Zootaxa 1800 on page 51, DOI: 10.5281/zenodo.6789337, {"references":["Cei, J. M. (1986) Reptiles del centro, centro-oeste y sur de la Argentina. Herpetofauna de las zonas aridas y semiaridas. Museo Regionale di Scienze Naturali di Torino, Monografie 4, 528 pp.","Etheridge, R. (1992) A new psammophilus lizard of the genus Liolaemus (Squamata: Tropiduridae) from northwestern Argentina. Bollettino del Museo Regionale di Scienze Naturali di Torino, 10, 1 - 19.","Cei, J. M. (1993) Reptiles del noroeste, nordeste y este de la Argentina. Herpetofauna de las selvas subtropicales, Puna y Pampas. Museo Regionale di Scienze Naturali di Torino, Monografie 14, 949 pp.","Avila, L. J., Acosta, J. C. & Murua, A. F. (1998) Herpetofauna de la provincia de San Juan, Argentina: lista comentada y distribucion geografica. Cuadernos de Herpetologia, 12, 11 - 29.","Etheridge, R & Espinoza, R. E. (2000) Taxonomy of the Liolaeminae (Squamata: Iguania: Tropiduridae) and a semiannotated bibliography. Smithsonian Herpetological Information Service, 126, 1 - 64.","Chebez, J. C., Rey, N. R. & Williams, J. D. (2005) Reptiles de los parques nacionales de la Argentina. Literatura of Latin America, Buenos Aires, 37 pp.","Sura, P. 2005. Encyclopedia of extant amphibians and reptiles, Wydawnictwo Fundacja, Nowy Sacz, 544 pp.","Abdala, C. S. (2007) Phylogeny of the boulengeri group (Iguania: Liolaemidae, Liolaemus) based on morphological and molecular characters. Zootaxa, 1538, 1 - 84."]}