A New Species of Lizard Endemic to Sierra de Fiambalá, Northwestern Argentina (Iguania: Liolaemidae: Phymaturus). Integrated Taxonomy Using Morphology and DNA Sequences: Reporting Variation Within the antofagastensis Lineage

© 2019 Academia Sinica, Taiwan Open Access A New Species of Lizard Endemic to Sierra de Fiambalá, Northwestern Argentina (Iguania: Liolaemidae: Phymaturus). Integrated Taxonomy Using Morphology and DNA Sequences: Reporting Variation Within the antofagastensis Lineage Lobo Fernando1,2,*, Thomas Hibbard2, Matías Quipildor2, and Soledad Valdecantos2 1División Herpetología, Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, CONICET, Avenida Ángel Gallardo 470, C1405DJR Buenos Aires, Argentina. *Correspondence: E-mail: [email protected] (Fernando) 2IBIGEO. Instituto de Bio y Geociencias del NOA (CONICET-UNSa), 9 de Julio 14, Rosario de Lerma, Salta, Argentina Received 1 October 2018 / Accepted 28 June 2019 / Published 5 September 2019 Communicated by Benny K.K. Chan The northernmost distributed group of lizards belonging to Phymaturus occurs in rocky outcrops of the Puna region between 3600–4200 m in Argentina. In a recent phylogenetic study based on morphological and genetic information, the monophyly of this small lineage was corroborated. This group is formed by Phymaturus antofagastensis, P. laurenti, P. denotatus, P. mallimaccii and a population of uncertain taxonomic status until the present study. After obtaining new samples and observations, we described a new species belonging to this lineage that is known only from Sierra de Fiambalá, being the species of Phymaturus living at the highest elevation ever recorded (4500 m). Males have a homogeneous yellow dorsum and lack melanic coloration over their heads, a phenomenon found in males of most species of the palluma group. We provide a detailed diagnosis, including characters from the squamation, coloration and significant differences found among continuous characters (ANOVA). Furthermore, we present genetic distances among members of the mallimaccii subclade based on sequences of the cytb marker. We provide color photos showing pattern variation of males and females. We reanalyze the phylogenetic relationships within the entire palluma group and update info on all members of the antofagastensis lineage based on new samples and make a better supported hypothesis. We also evaluate the phylogenetic position of the new taxon. Key words: Phymaturus fiambala sp. nov., Taxonomy, Squamata, Liolaemidae, Argentina. Citation: Fernando L, Hibbard T, Quipildor M, Valdecantos S. 2019. A new species of lizard endemic of Sierra de Fiambalá, northwestern Argentina (Iguania: Liolaemidae: Phymaturus). Integrated taxonomy using morphology and DNA sequences: reporting variation within the antofagastensis lineage. Zool Stud 58:20. doi:10.6620/ZS.2019.58-20. BACKGROUND The genus Phymaturus is known for its extremely endemic species, often known only from their type locality, despite extensive sampling done over the years by different herpetologists. This pattern is likely caused by the genus habitat, which consists of rocky outcrops with crevices that these animals use as refuge from predators. Unlike its morphologically diverse sister genus, Liolaemus, Phymaturus’ morphology is highly conserved, being significantly dorso-ventrally flattened in order to make better use of the crevices (GonzálezZoological Studies 58: 20 (2019) doi:10.6620/ZS.2019.58-20 1 © 2019 Academia Sinica, Taiwan Marín et al. 2018, Troncoso-Palacios et al. 2018). They are also exclusively herbivorous and viviparous, with biennial reproduction. Due to this morphological conservatism, recognizing new species calls for in-depth knowledge of these animals’ systematic and diagnostic traits. Furthermore, given the extremely endemic nature of these species, their low population densities, and their biennial form of reproduction (Boretto and Ibargüengoytía 2006 2009), all Phymaturus were considered vulnerable in their latest categorization (Abdala et al. 2012). Therefore, recording the morphological diversity and delineating species within this clade is a primary goal for their conservation. Etheridge (1995) divided the genus Phymaturus in two species groups: the patagonicus and the palluma groups, based on a study of morphological characters. In his study, he proposed apomorphies, but did not present a formal phylogenetic analysis. Lobo and Quinteros (2005) performed an analysis using phylogenetic methods for the first time, confirming Etheridge’s division (1995), although the patagonicus group was less supported than the palluma group. Based mainly on morphological characters, Lobo and Quinteros (2005) recovered a clade within the palluma group (Node 12 fig. 8) formed at this time by P. antofagastensis Pereyra 1985, P. mallimaccii Cei 1980, P. punae Cei, Etheridge and Videla 1983, P. cf. punae and P. cf. antofagastensis. All members of this clade are distributed in the highland Andean areas of Puna (an expansive region of western South America which is a part of the Central Andes and forms the world’s second largest plateau). Puna is a desertic area situated at high elevation (above 3000 m), has a typical dominant vegetation of shrubs (there are no trees) (Martínez Carretero 1995) and its aridity and general landscape has existed at least since the Miocene (Strecker et al. 2007). In an updated analysis of 206 morphological characters (mainly based on squamation, colors and patterns, body proportions and skeletons), Lobo, Abdala and Valdecantos (2012) recovered this Puna clade (Node G, fig. 9) formed by P. mallimacci, P. punae, P. antofagastensis, P. laurenti (Lobo, Abdala and Valdecantos 2010), P. extrilidus (Lobo, Espinoza, Sanabria and Quiroga 2012b), and other five P. spp. (candidate species). At this time, two subclades were recognized within that node (a northern one with species inhabiting mountains of Catamarca and La Rioja provinces, and a southern one formed by species inhabiting La Rioja and San Juan provinces). After that revisionary contribution, three other species of this clade were formally described by Lobo et al. (2012c) from Laguna Blanca, Catamarca: P. denotatus (Lobo et al. 2013), and from San Juan province: P. aguanegra and P. williamsi. Using nuclear and mitochondrial sequences, Morando et al. (2013) recovered the same Puna-endemic clade including P. punae, P. extrilidus, P. mallimaccii and P. laurenti subdivided into the same two subclades previously recovered using morphology in Lobo et al. (2012a). Morando et al. (2013) named this clade the mallimaccii group. Recently, Lobo et al. (2016) sequenced fragments of cytochrome b (cytb), 12S, and ND4 for all terminals; described 45 new morphological characters; and incorporated all DNA sequences available from GenBank. Within the palluma group, two sister clades were recovered, the vociferator and bibroni clades, and two subclades within the latter: the roigorum and mallimaccii subclades (the latter equivalent to the mallimaccii group of Morando et al. 2013). The mallimaccii subclade consists of 13 terminal taxa, to which two Chilean species have been added in the last cladistic analyses: P. bibroni (Guichenot 1848) and P. aguedae Troncoso-Palacios and Esquerré 2014. Lobo et al. (2016) divided the mallimaccii subclade into two lineages: the antofagastensis lineage (P. mallimaccii, P. antofagastensis, P. laurenti, P. denotatus, sp. gua and P. sp. fia) and the punae lineage (P. punae, P. extrilidus, P. williamsi, P. aguanegra, P. bibroni and P. sp. lar). Grosso et al. (2017) studied the chromosome morphology of six species of the palluma group, including in their analysis three species of the mallimaccii subclade (P. laurenti, P. denotatus and P. williamsi). In this last study, interesting chromosome variation within the palluma group was described, including a multiple sex-chromosome system and several Robertsonian rearrangements. More recently, Troncoso et al. (2018) provided a multilocus phylogenetic analysis of the vociferator clade adding other species and previously unsampled populations from Chile to their data set. Troncoso et al. (2018) included most terminals of the mallimaccii subclade, recovering the two main lineages with a third one endemic to Chile. The present study refines the description of an unnamed population of Phymaturus that had previously been included in phylogenetic analyses. We reanalyze the phylogenetic relationships within the palluma group based on an updated data matrix after studying new samples of nine terminal taxa. We present here a description and diagnosis of other members of the lineage. MATERIALS AND METHODS We examined 203 specimens belonging to nine species of Phymaturus, including the type series of the new ones therein described (see Appendix 1). For this work, we collected sample data from Fiambalá mountains (WGS 27.25583 S 67.20980 W; altitude: page 2 of 18Zoological Studies 58: 20 (2019) © 2019 Academia Sinica, Taiwan 4533 m), Fiambalá Department, Catamarca Province, Argentina. Four adult males, two juvenile males and six females were sampled. We provide an original description of a new taxon of Phymaturus, with data on their variation and phylogenetic relationships. Appropriate actions were taken to minimize pain or discomfort of all lizards involved in this study, in accordance with international standards on animal welfare and national regulations of the “Comité Nacional de Ética en la Ciencia y la Tecnología” of Argentina (Expte. 5344/99 Res. 1047). All specimens were collected in the summer of 2017 by noose or by hand, and then fixed using 10% formalin and deposited in 70% ethanol. All herpetological collection data of the new species are recorded in collection databases in the Museo de Ciencias Naturales, Universidad Nacional de Salta (MCN-UNSa) and Instituto de Bio y Geociencias del Noa, Argentina (IBIGEO). Genetic data on all species in the group were extracted from GenBank, and accession numbers are indicated in table S1. GenBank accession numbers for sequences of the new species are KT203836 (12S), KT203831 (cytb), KT203850 (ND4) and KT203819 (Cmos), first published in Lobo et al. (2016). The genetic distances for cytb among members of the mallimaccii subclade are shown in table 1. All positions with less than 95% site coverage were eliminated. That is, fewer than 5% alignment gaps, missing data, and ambiguous bases were allowed at any position. Analyses were conducted in MEGA5 (Tamura et al. 2011). Phylogenetic relationships were analyzed updating a data matrix used in previous studies for the palluma group (Lobo et al. 2012 2016; Hibbard et al. 2019) and DNA sequences available in GenBank (including those used recently by Troncoso et al. 2018) using TNT, a parsimony software (Goloboff et al. 2008). Accession numbers for sequences from GenBank are reported in a table S1. In previous studies, this new taxon was mentioned as P. sp5 in Lobo et al. (2012) and as P. sp. fia in Lobo et al. (2016). At the time of those analyses, we had sequences of cytb, 12S and ND4 taken from a female individual (MCN-UNSa 2123) of the new species (see accession numbers above), but morphology data were taken only from two females and a juvenile. In this case we collected all information on males and obtained data from a total of twenty specimens of the new species. Coding procedures were described in detail in the studies above mentioned. We added eighteen new characters (254–267: scale organs; 268–270: color pattern; 271: integumentary glands), which are listed at the end of appendix 1. Also, we improved our samples for the whole morphology of P. mallimacci, P. antofagastensis and P. laurenti taking data from FML, MLP and DC-JMC collections. We updated morphological information on P. maulense, P. vociferator, P. damasense, and P. timi (see Appendix 1). Some of the terminals were assigned to species in accordance with the most recent literature. In previous Table 1. Estimates of divergence among cytb sequences of ten species of the Phymaturus mallimaccii subclade (shaded), plus three other representatives of other palluma group lineages and two species of the patagonicus group. Sequences of P. sp. gua, P. sp. lar, sensu Lobo et al. (2016) were not available; nor were those of P. aguanegra. The number of base differences per site from between sequences are shown. All positions with less than 95% site coverage were eliminated. That is, fewer than 5% alignment gaps, missing data, and ambiguous bases were allowed at any position. Analyses were conducted in MEGA5 (Tamura et al. 2011) 1234567891011 12 13 14 15 1) aguedae 2) antofagastensis 0.040 3) bibroni 0.037 0.017 4) denotatus 0.043 0.006 0.017 5) extrilidus 0.039 0.015 0.007 0.015 6) fiambala 0.044 0.014 0.018 0.011 0.017 7) laurenti 0.043 0.006 0.017 0.000 0.015 0.011 8) mallimaccii 0.040 0.015 0.018 0.012 0.019 0.017 0.012 9) punae 0.037 0.019 0.010 0.019 0.010 0.019 0.019 0.022 10) williamsi 0.041 0.018 0.012 0.015 0.011 0.017 0.015 0.017 0.014 11) dorsimaculatus 0.061 0.064 0.057 0.066 0.066 0.066 0.066 0.068 0.061 0.064 12) palluma 0.043 0.041 0.032 0.041 0.033 0.041 0.041 0.040 0.033 0.033 0.059 13) querque 0.055 0.052 0.044 0.052 0.048 0.057 0.052 0.055 0.046 0.046 0.054 0.046 14) indistinctus 0.146 0.140 0.137 0.145 0.144 0.148 0.145 0.144 0.146 0.149 0.134 0.149 0.145 15) somuncurensis 0.148 0.151 0.149 0.156 0.153 0.159 0.156 0.152 0.156 0.159 0.138 0.155 0.152 0.065 page 3 of 18Zoological Studies 58: 20 (2019) © 2019 Academia Sinica, Taiwan studies, we included Phymaturus from Termas de Chillán (P. cf. palluma CH, P. sp2 or P. sp. chi in Lobo and Quinteros 2005, Lobo et al. 2012 2016); now we assigned this sample to P. vociferator following Urra et al. (2017). We assigned other samples from El Planchón (P. cf. palluma EP, P. sp3 or P. sp. pla in Lobo and Quinteros 2005, Lobo et al. 2012 2016) to P. damasense following Ramírez-Alvarez et al. (2017). We analyzed our data matrix with TNT v. 1.5 applying strict parsimony (Goloboff et al. 2008). Support for individual nodes was assessed with jackknifing resampling (Siddall 1995) using 1000 replicates and a deletion value of 25%. Measurements were taken using digital calipers at 0.02 mm of precision; pictures of live specimens were taken in the field using a digital camera, and most character details were examined under a stereomicroscope. Most characters described in diagnoses and descriptions followed standards published in Smith (1946), Cei (1986 1993), Laurent (1984 1986), Etheridge (1995), Lobo and Quinteros (2005) and Lobo et al. (2010). Additionally, we chose 17 continuous characters of squamation plus snout-vent length (Table 2) to analyze if significant differences exist among species belonging to the antofagastensis lineage. These characters were: SVL (snout-vent length), number of scales around midbody, Hellmich’s index (scales counted along the mid-line of the head between the occiput and rostrum), number of scales contacting interparietal, number of infralabial scales, number of subocular scales, scales contacting nasal, lorilabial scales, temporal scales, superciliary scales, gular scales and the number of precloacal pores in males (in this lineage there are no pores in females), scales contacting mental, ventral scales, scales projecting over auditory meatus, number of dorsal scales (counted at middle of the trunk in a head-length), scales between frontalrostral and scale organ on postrostrals. The data met assumptions of normality and homogeneity of variance. We performed an ANOVA (analysis of variance) using a test of multiple comparisons LSD of Fisher running the statistical package INFOSTAT (Di Rienzo et al. 2016). RESULTS TAXONOMY Family Liolaemidae Frost and Etheridge, 1989 Genus Phymaturus Gravenhorst, 1838 Phymaturus fiambala sp. nov. Lobo, Hibbard, Quipildor and Valdecantos (Figs. 1, 3–6) urn:lsid:zoobank.org:act:04F90EF0-A70A-49C4-A91FA1FDFCB7466 Table 2. ANOVA results obtained after comparisons across species of the antofagastensis lineage of Phymaturus for eighteen continuous characters. Different capital letters following mean ± standard error between species indicate a significant difference. Sample size indicated between parentheses P. antofagastensis (n = 19) P. denotatus (n = 15) fiambala (n = 18) P. laurenti (n = 23) P. mallimaccii (n = 16) Prueba p SVL 90.02 ± 2.08 B98.37 ± 2.15 A97.73 ± 1.96 A91.43 ± 1.74 B86.44 ± 2.08 BF = 6.26 0.0002 Scales around midbody 198.11 ± 2.84 B207.80 ± 3.19 C193.17 ± 2.91 A B 193.87 ± 2.58 B185.25 ± 3.09 AF = 6.93 0.0001 Hellmich index 21.63 ± 0.42 B22.60 ± 0.47 B C 18.78 ± 0.43 A23.26 ± 0.38 C21.56 ± 0.46 BF = 16.70 < 0.0001 Contacting interparietal 8.63 ± 0.26 A8.60 ± 0.29 A8.28 ± 0.27 A8.70 ± 0.24 A9.69 ± 0.28 BF = 3.67 0.0083 Infralabial scales 9.47 ± 0.29 9.80 ± 0.32 9.67 ± 0.29 10.17 ± 0.26 9.56 ± 0.31 F = 1.01 0.4077 Subocular scales 2.16 ± 0.22 B C 2.80 ± 0.24 C D 1.89 ± 0.22 A B 2.91 ± 0.20 D 1.31 ± 0.23 AF = 8.84 < 0.0001 Contacting nasal 9.00 ± 0.21 A B 9.33± 0.24 B8.39 ± 0.22 A9.22 ± 0.20 B8.56 ± 0.23 AF = 3.37 0.0130 Lorilabial scales 11.68 ± 0.28 A14.33 ± 0.31 C14.06 ± 0.29 C12.74 ± 0.25 B12.19 ± 0.30 A B F = 15.45 < 0.0001 Temporal scales 10.21 ± 0.25 A11.73 ± 0.28 C11.44 ± 0.26 B C 10.70 ± 0.23 A10.75 ± 0.28 A B F = 5.37 0.0007 Superciliary scales 10.16 ± 0.28 A11.20 ± 0.31 B11.28 ± 0.28 B10.35 ± 0.25 A10.81 ± 0.30 A B F = 3.18 0.0173 Gular scales 83.58 ± 1.70 A B 80.27 ± 1.91 A89.67 ± 1.7 C85.78 ± 1.55 B C 83.25 ± 1.85 A B F = 10.86 < 0.0001 Precloacal pores 9.22 ± 0.40 B C 8.00 ± 0.69 A B 10.43 ± 0.45 C7.40 ± 0.31 A7.22 ± 0.40 AF = 3.75 0.0074 Contacting mental scales 6.37 ± 0.16 A B 6.80 ± 0.22 B C 6.11 ± 0.08 A6.04 ± 0.04 A7.00 ± 0.20 CH = 18.55 < 0.0001 Ventral scales 182.11 ± 2.30 189.93 ± 2.81 183.44 ± 2.28 181.87 ± 1.93 181.13 ± 2.87 F = 1.95 0.1093 Scales projected over auditory meatus 4.05 ± 0.18 C2.00 ± 0.49 B4.06 ± 0.39 C0.83 ± 0.29 A5.19 ± 0.54 D F = 23.33 < 0.0001 Dorsal scales 34.89 ± 1.23 36.73 ± 1.55 35.67 ± 0.74 35.61 ± 0.90 37.06 ± 0.65 F = 0.68 0.6052 Scales between frontalrostral Scale 9.05 ± 0.30 B10.07 ± 0.25 C8.22 ± 0.27 A10.39 ± 0.29 C10.50 ± 0.29 CF = 11.76 < 0.0001 organs on postrostrals 1.33 ± 0.15 1.72 ± 0.18 1.18 ± 0.11 1.43 ± 0.24 1.77 ± 0.19 F = 1.65 0.1698 page 4 of 18Zoological Studies 58: 20 (2019) © 2019 Academia Sinica, Taiwan Synonymy: Phymaturus sp5: Lobo et al. 2012: 21. Phymaturus sp. fia: Lobo et al. 2016: 650. Deposition of types: Holotype: IBIGEO 5756. Male (Fig. 1). Paratypes: IBIGEO 5757–59, 5765, 5774 (4 adult males); 5769–70 (2 juvenile males). 5760–61, 63–64, 5766, 68 (6 females) deposited at the Reptiles collection of the Instituto de Bio y Geociencias del Noa (IBIGEO), Salta, Argentina. Site: 27.25583 S 67.20980 W; altitude: 4533 m. Locality: Cerca del Puesto de la lagunilla, Fiambalá Department, Catamarca Province, Argentina. Dates: 6 December 2017. Collectors: Thomas Hibbard and Matías Quipildor. MACN 51034035 (ex IBIGEO 5762, 5767). Same data, deposited at the Herpetological collection of the Museo Argentino de Ciencias Naturales, Buenos Aires, Argentina. MCNUNSa 2122, 2123 (2 females, MCN-UNSa 2123 is voucher of DNA sequences), deposited at Museo de Ciencias Naturales, Universidad Nacional de Salta, Salta, Argentina. Locality: Puesto la Lagunita, 35–38 km NE of Medanitos, climbing from Medanitos, Fiambalá Department, Catamarca Province, Argentina. Dates: 23 March 2006. Collectors: Sebastián Barrionuevo, Juan Manuel Díaz Gómez and Sebastián Quinteros. MCNUNSa 2125 juvenile. Same data, deposited at Museo de Ciencias Naturales, Universidad Nacional de Salta, Salta, Argentina. Diagnosis: Phymaturus fiambala sp. nov. Doral pattern with very thin spray, throats and chests light Fig. 1. (A) Dorsal view of the holotype of Phymaturus fiambala sp. nov. IBIGEO 5756. (B) Ventral view of the same specimen (Photos: M. Quipildor). (A) (B) page 5 of 18Zoological Studies 58: 20 (2019) © 2019 Academia Sinica, Taiwan gray, rostral scales always undivided. Males with enlarged postcloacal scales, females with slender white transversal lines over trunk, enlarged scales on posterior gular fold, a patch of enlarged scales between gular folds evident, vertebral stripe absent. Deposition of types: Holotype: IBIGEO 5756. Male (Fig. 1). Paratypes: IBIGEO 5757–59, 5765, 5774 (4 adult males); 5769–70 (2 juvenile males). 5760–61, 63–64, 5766, 68 (6 females) deposited at the Reptiles collection of the Instituto de Bio y Geociencias del Noa (IBIGEO), Salta, Argentina. Site: 27.25583 S 67.20980 W; altitude: 4533 m. Locality: Cerca del Puesto de la lagunilla, Fiambalá Department, Catamarca Province, Argentina. Dates: 6 December 2017. Collectors: Thomas Hibbard and Matías Quipildor. MACN 51034– 035 (ex IBIGEO 5762, 5767). Same data, deposited at the Herpetological collection of the Museo Argentino de Ciencias Naturales, Buenos Aires, Argentina. MCNUNSa 2122, 2123 (2 females), deposited at Museo de Ciencias Naturales, Universidad Nacional de Salta, Salta, Argentina. Locality: Puesto la Lagunita, 35–38 km NE of Medanitos, climbing from Medanitos, Fiambalá Department, Catamarca Province, Argentina. Dates: 23 March 2006. Collectors: Sebastián Barrionuevo, Juan Manuel Díaz Gómez and Sebastián Quinteros. MCNUNSa 2125 juvenile. Same data, deposited at Museo de Ciencias Naturales, Universidad Nacional de Salta, Salta, Argentina. Description of holotype (Fig. 1): Male. SVL 98.4 mm. Head length: 18.1 mm. Head width: 18.7 mm. Head height (at parietal): 8.2 mm. Axilla-groin length: 50.1 mm (50.9% of SVL). Tail length (complete, not regenerated): 71.3 mm to the point of regeneration. Body moderately wide, trunk width: 36.5 mm (37.1% of SVL). Twenty smooth dorsal head scales. Three scale organs in three postrostrals. Nasal bordered by ten scales, not in contact with rostral. Canthal separated from nasal by two scales. Loreal region flat. Twelve enlarged supralabial scales, none contacting subocular. Ten enlarged infralabials. Auditory meatus oval shaped (height: 3.9 mm; width: 2.1 mm) with four enlarged, flat and keeled backwardly projecting scales on the anterior margin. Auricular scale absent. Twelve convex, juxtaposed temporals. Auditory meatusciliary scales posterior commissure distance: 6.3 mm. Rostral undivided. Mental scale sub-pentagonal, in contact with six scales. Interparietal scale bordered by eight scales, being of larger size than postparietals. Frontal region without an azygous scale. Supraorbital semicircles inconspicuous. No distinctly enlarged supraoculars. Eleven juxtaposed superciliaries, seventeen upper ciliaries and sixteen lower ciliaries. Subocular fragmented in two scales. Fifteen lorilabials, without contacting subocular. Preocular larger than canthal, separated by one scale. Preocular separated from lorilabial row by three scales. Scales of throat round, small, and juxtaposed. Eighty-eight gulars between auditory meata. Lateral nuchal folds well developed, with granular scales over longitudinal fold. Antehumeral pocket well developed. Sixty-four scales between auditory meatus and shoulder. Fortyone scales between antehumeral fold and shoulder. In ventral view, anterior and posterior gular folds present, their anterior margins with two to three enlarged scales on their borders. Dorsal scales round, smooth and juxtaposed. Thirty-six dorsal scales along midline of the trunk in a length equivalent to head length. Scales around midbody: 178. Ventral scales larger than dorsals. Ventral scales between mental and precloacal pores: 187. Ten precloacal pores in an undivided row with two supernumerary pores. Two slightly enlarged postcloacal scales. Brachial and antebrachial scales smooth, with round posterior margins. Supracarpals laminar, round and smooth. Subdigital lamellae of fingers have three keels. Subdigital lamellae of finger (left manus) IV: 21. Claws moderately long (fourth toe’s claw: 2.6 mm). Supradigital lamellae convex, imbricate. Infracarpals and infratarsals have round margins and 2–3 keels. Supracarpals and supratarsals smooth, with rounded posterior margins. Subdigital lamellae of toe (left pes) IV: 25. Coloration of holotype: the holotype exhibits a homogeneous yellow dorsal background, with small light brown scales scattered irregularly over all its body. Dorsum of tail of the same yellow coloration as trunk (no ringed or variegated pattern). Head uniformly light brown, with this coloration extended over the lateral neck folds. Throat immaculate light gray with no variegation. It has almost inconspicuous, very small and disperse spots, slightly darker than the background. Immaculate chest and belly entirely yellow from the anterior gular fold to the cloacal opening, extended over fore and hindlimbs and ventral surfaces of thighs and tail. Ventral surface of tail does not have any pattern. Color of a female (Fig. 2): background dorsal coloration light brown all over head, trunk, tail and limbs. Light brown coloration speckled with darker brown scales, which become confluent on the sides of neck and shoulders. Scapular spot conspicuous. Flanks with yellow coloration that extends to the belly as symmetrical patches. Most of ventral surfaces immaculate, light gray to white. Dorsal pattern of tail ringed. Ventral surface of tail lacks any kind of pattern. Etymology: The species inhabits Sierra de Fiambalá (Fiambalá mountains). The toponym Fiambalá comes from an ancient language (Cacán) of natives who lived in northwestern Argentina before Quechua (Inca) and Spanish became dominant. “Cacán” voice: page 6 of 18Zoological Studies 58: 20 (2019) © 2019 Academia Sinica, Taiwan fiambalao (fiambal = wind; ao = house, place), meaning “house of winds”. Variation: based on 16 adult specimens (7 males and 9 females). SVL 90.2–102.3 mm (x = 97.5; SD = 3.2) (two juveniles not included to avoid including ontogenetic variation). Head length 16.7–18.9% (x = 17.7%; SD = 0.7) of SVL. Tail length 0.80–1.08 (x = 0.96; SD = 0.08) times SVL. Scales around midbody 178–212 (x = 192.4; SD = 9.6). Dorsal head scales 15–22 (x = 18.6; SD = 1.9). Ventral scales 168–203 (x = 184.4; SD = 8.3). Scales surrounding interparietal 7–10 (x = 8.3; SD = 0.9). Scales surrounding nasal 7–10 (x = 8.4; SD = 0.9). Number of scale organs on postrostrals 1–3 (x = 1.1; SD = 0.5). Superciliaries 10–13 (x = 11.2; SD = 0.9). Subocular fragmented in half of the sample (ten specimens). Mental scale in contact with 6–7 (x = 6.1; SD = 0.3). Number of chinshields 2–7 (x = 4.5; SD = 1.8). All specimens exhibit enlarged scales on the border of the posterior gular fold (varying in number). Lorilabials 12–16 (x = 13.9; SD = 1.1). Enlarged scales on the anterior border of the auditory meatus 3–7 (x = 4.5; SD = 1.3) (Fig. 3A). Scales of neck along longitudinal fold from posterior border of auditory meatus to shoulder 60–76 (x = 68.4; SD = 4.6). Gulars 77–100 (x = 89.2; SD = 6.2). Scales between rostral and frontal 6–10 (x= 8.2; SD = 1.2). Subdigital lamellae on fourth finger 18–21 (x = 19.4; SD = 1.1). Subdigital lamellae on fourth toe 22–28 (x = 24.1; SD = 1.6). Males with 9–12 precloacal pores (x = 10.3; SD = 1.0). No females show precloacal pores. A small, newborn-sized individual was collected with 52.1 mm SVL (IBIGEO 5770). It shows two conspicuous enlarged postcloacal scales and a row of differentiated scales that will house later (at its maturity) precloacal pores (Fig. 3B). A juvenile male (IBIGEO 5769) with 78.2 mm SVL shows a row of differentiated scales but without pit or any signal of secretion. It has slightly conspicuous enlarged postcloacal scales and it exhibits a Fig. 2. (A) Dorsal view of a female of Phymaturus fiambala sp. nov. IBIGEO 5763. (B) Ventral view of the same specimen (Photos: M. Quipildor). (A) (B) page 7 of 18Zoological Studies 58: 20 (2019) © 2019 Academia Sinica, Taiwan Fig. 3. (A) Details of the head in a female of Phymaturus fiambala sp. nov. (MCN-UNSa 2123); (B) newborn P. fiambala sp. nov. (IBIGEO 5770); (C) ventral view of females of P. laurenti; (D) females of P. denotatus. Character 35 (number of precloacal pores in males) 112 (row of precloacal pores); character 138 (1) presence of enlarged postcloacal scales in males; character 156 (1) preocular scale small separated from canthal by another scale; character 165 (1) three to seven enlarged scales on the anterior border of the auditory meatus; character 166 (0) enlarged scales on the anterior border of auditory meatus projected posteriorly over the ear opening (Photos: M. Quipildor). (A) (B) (D) (C) page 8 of 18Zoological Studies 58: 20 (2019) © 2019 Academia Sinica, Taiwan typical ringed tail as the smallest juvenile. Males (Fig. 4) exhibit a yellow coloration all around their trunks. This color can be extended over tails, and in lesser degree over fore and hindlimbs. Dorsum and flanks speckled of dark brown/black scales that become more densely distributed in the neck and shoulders. Dorsal melanism of neck incomplete over the mid vertebral line (character 172). All males show a scapular yellow spot (character 139). Chest and abdomen immaculate yellow. Heads light brown, not a single specimen exhibits melanization common in the palluma group (character 124). Females (Fig. 5) homogeneous brown (head, body, limbs and tail), their trunks speckled with black to dark brown scales. Lateral sides of neck, in several cases melanic, can be extended over the shoulders and the axilla. Scapular spot conspicuous in most females (character 140). Most females exhibit white slender transverse stripes across their backs (character 180). Ventral surfaces light gray almost white with a pair of yellow patches on the sides extended over flanks (character 183). Tails patterned (irregularly distributed darker spots) like in males but more conspicuous. Distribution: At present, only known from its type locality. Detailed comparisons to other members of the antofagastensis lineage Phymaturus fiambala sp. nov. belongs to the antofagastensis lineage because it shares synapomorphies with all other members of the lineage: four discrete and three continuous characters (presence of flank color in females, loss of scale organ in mental, dark sides of neck speckled with small white spots among them) plus eight DNA changes (see below “Phylogenetic relationships”). Because of this, comparisons are restricted to all members of the lineage. Phymaturus fiambala sp. nov. males exhibit yellow tails continuing the same color of trunks, different from all other members of the mallimaccii subclade with males that exhibit brown tails (yellow tails are found in the vociferator clade, and the roigorum subclade). Phymaturus fiambala sp. nov. differs from P. antofagastensis in that has a pattern of very thin spray, while P. antofagastensis exhibits a typical aggregated pattern (Lobo and Quinteros 2005, fig. 12D) formed by larger brown spots irregularly distributed over its body. Males exhibit a yellow coloration covering head, trunk, limbs and tail (Fig. 1) while in P. antofagastensis, yellow is more restricted to flanks, shoulders and neck, never shown in tails (Fig. 6). Throats and chests in P. fiambala sp. nov. are light gray, being dark, almost completely melanic in P. antofagastensis. In P. fiambala sp. nov., granular scales among dorsal tibial scales are absent, while they are present in P. antofagastensis. In P. fiambala sp. nov. the rostral scale is always undivided, while 63% of studied individuals of P. antofagastensis show a divided rostral scale. In P. fiambala sp. nov., all males exhibit enlarged poscloacal scales (like in P. laurenti see Lobo et al. 2012c, fig. 3D) while only 37.5% of males of P. antofagastensis do. White transversal stripes are quite evident and wide in females of P. antofagastensis, but slender and almost inconspicuous in P. fiambala sp. nov. Also, P. fiambala sp. nov. shows significant differences in other five continuous characters (Table 2): P. fiambala sp. nov. shows a larger SVL than P. antofagastensis, more lorilabials, superciliaries and gular scales, fewer scales between rostral and frontal, and fewer scales along midline of head (Hellmich’s index). Phymaturus fiambala sp. nov. differs from P. mallimaccii in that males of the second species exhibit melanic throats, and several a very dark pattern formed by a dense distribution of small dark spots over dorsum. Also, in P. mallimaccii, a vertebral stripe is conspicuous, i.e., a vertebral stripe of a lighter coloration similar to the one shown by species of the punae lineage but absent in P. fiambala sp. nov. In P. mallimaccii, lateral neck folding is dark and speckled with small white spots even in males (Fig. 6D) but in certain individuals it is not so evident while in P. fiambala sp. nov. this character is absent. In P. fiambala sp. nov., enlarged scales on posterior gular fold and a patch of enlarged scales between gular folds are evident, while in P. mallimaccii they are inconspicuous or absent. Flank coloration in females of P. fiambala sp. nov. is yellow but orange in females of P. mallimaccii. According to statistical tests, P. fiambala sp. nov. have significantly larger SVL than P. mallimaccii, lower Hellmich’s index, fewer scales contacting interparietal, scales contacting mental (Lobo and Quinteros 2005, fig. 9C & D), scales projecting on the anterior margin of the auditory meatus (Fig. 3A), scales between rostral and frontal but more lorilabial scales, gular scales and precloacal pores. Phymaturus fiambala sp. nov. is different from P. laurenti in that the scapular spot is absent in P. laurenti, present in P. fiambala sp. nov. Flank coloration of females is orange in P. laurenti (Fig. 3C) but yellow in P. fiambala sp. nov. Tarsal scales in P. fiambala sp. nov. are strongly keeled but slightly keeled in P. laurenti. Enlarged poscloacal scales in males are larger in P. laurenti. Phymaturus fiambala sp. nov. lacks granular scales among dorsal tibial scales that are present in P. laurenti (see this character in Lobo et al. 2016, fig. 8F). Also, there are eight continuous characters that exhibit significant differences: P. fiambala sp. nov. has a larger SVL, lower Hellmich’s index, fewer subocular scales (Lobo and Quinteros 2005, fig. 9A & B), scales page 9 of 18Zoological Studies 58: 20 (2019) © 2019 Academia Sinica, Taiwan is the most basal taxon). In the combined analysis of Lobo et al. (2016, fig. 2), we recovered the same topology recovered here, with the exception of P. sp. gua, which now is not included in the lineage. In the present analysis with only the molecular information, we found with parsimony P. fiambala sp. nov. as the most basal species of the antofagastensis lineage but lack of support for P. mallimaccii as sister taxon of the rest of members of that lineage (Fig. 7A). Although the molecular analysis was made for comparison, we believe that total evidence analysis is more accurate, as information given by the more than 260 morphological characters has been exhaustively researched for the last 10 years. The position of P. mallimaccii as sister to P. fiambala sp. nov. is therefore preferred. In any case, we find that independent lines of evidence both indicate that P. fiambala sp. nov. is indeed a member of the P. antofagastensis clade with good support, which indicates that at least the composition of this previously named group undoubtedly stays firm. Finally, it should be mentioned that a character that was recovered as an apomorphy of the antofagastensis lineage in Lobo et al. (2016)—brown-pigmented dorsal fascia of longissimus dorsi and transverso spinalis without pigmentation (character 216)—is not recovered now. The present study it is recovered it as an apomorphy that links both terminal lineages, with a secondary loss in the group formed by P. aguanegra and P. williamsi. Species of the mallimaccii subclade exhibit phenotypic characters that allow for a clear differentiation among them, but low cytb distances in comparison to other clades. Species of the antofagastensis lineage are an example of this phenomenon. This highlights the importance of including phenotypic characters in phylogenetic analyses. CONCLUSIONS Phymaturus fiambala sp. nov. belongs to the antofagastensis lineage because it shares synapomorphies with all other members of the lineage: four discrete and three continuous characters (presence of flank color in females, loss of scale organ in mental, dark sides of neck speckled with small white spots among them) plus eight DNA changes. Phymaturus fiambala sp. nov. males exhibit yellow tails continuing the same color of trunks, different from all other members of the mallimaccii subclade, which is composed of males that exhibit brown tails (yellow tails are found in the vociferator clade, and the roigorum subclade). Furthermore, we found morphological and molecular evidence that allows us to differentiate P. fiambala sp. nov. from other species within antofagastensis lineage. Our present phylogenetic analysis provides a new hypothesis of relationships within the mallimaccii subclade that allow for more confident studies on evolutionary comparisons and the biogeography of these Andean lizards in the future. Acknowledgments: This work and the new species name have been registered with ZooBank under urn:lsid:zoobank.org:pub:95B79C5B-823B-4A10AC8B-7049FB384E09. We thank S. Quinteros, S. Barrionuevo, J. M. Díaz Gómez and C. Abdala for their guidance on our Medanitos and Sierra de Fiambala expedition after their 2006 trip. To D. Barrasso for his critical review of a first version and S. Wenner (CSUN) for her invaluable help with the language. D. Slodki, S. Ruiz, L. Díaz Fernández and A. Paz for their continuous help and support in lab work at IBIGEO. The senior author acknowledges the valuable support of colleagues and friends at the Herpetology Division of MACN. This study was supported by grants (FL) from CONICET Consejo Nacional de Investigaciones Científicas y Técnicas of Argentina (PIP 0871) and CIUNSA Consejo de Investigaciones de la Universidad Nacional de Salta, Argentina (CIUNSA 2342). We thank the following colleagues (and museums) for allowing F. L. to study specimens under their care recently and over the last decade: B. Espeche (Unidad de Herpetología - Facultad de Química, Bioquímica y Farmacia - Universidad Nacional de San Luis, curator of the Diagnostic Collection José Miguel Cei), R. Espinoza (CSUN Herpetological Collection), E. Pereyra (Instituto de Biología Animal, Universidad Nacional de Cuyo, Mendoza), F. Videla (IADIZA, Mendoza), E. Lavilla and S. Kretzschmar (Instituto de Herpetología, Fundación Miguel Lillo, Tucumán), J. Faivovich and S. Nenda (Museo Argentino de Ciencias Naturales, Buenos Aires), J. Williams and L. Alcalde (Museo de La Plata), A. Scolaro (CENPAT, Pto. Madryn). Authors’ contributions: All the authors contributed in the same way. Competing interests: FL, TH, MQ, and SV declare that they have no conflict of interest. Availability of data and materials: The data and materials are specified in the materials and methods section and in appendix 1. Consent for publication: Not applicable. 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An updated phylogeny and morphological study of the Phymaturus vociferator clade (Iguania: Liolaemidae). Zootaxa 4441:447– 466. doi:10.11646/zootaxa.4441.3.2. Urra FA, Díaz HA, Werning H, Eisenberg T, Troncoso-Palacios J. 2017. Phymaturus vociferator Pincheira-Donoso, 2004 (Squamata: Liolaemidae): new records and updated geographic distribution. Check List 13:1–4. doi:10.15560/13.3.2137. Supplementary materials Appendix 1. List of individuals studied for each species and list of morphological characters added to the matrix. (download) Table S1. List of all the species, voucher numbers and Genbank accession numbers of the sequences employed in this study. (download) page 18 of 18Zoological Studies 58: 20 (2019)