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Higher TaxaLiolaemidae, Iguania, Sauria, Squamata (lizards)
Common Names 
Ctenoblepharis adspersus — BOULENGER 1885: 136
Ctenoblepharus sp. — PÉFAUR et al. 1978
Liolaemus insolitus — CEI & PÉFAUR 1982
Ctenoblepharys adspersa — ETHERIDGE 1995
Phrynosaura sp. — NUÑEZ 2004
Liolaemus cf. insolitus — GUTIÉRREZ & QUIRÓZ, 2010
Liolaemus species 2 — LANGSROTH 2011: 32
Liolaemus aff. insolitus7 — ABDALA et al. 2020 
DistributionPeru (Arequipa)

Type locality: between Quebrada San Jose and Quebrada Tinajones, District of Uchumayo, Province of Arequipa, Department of Arequipa, Peru (16°31’47”S, 71°39’04”W) at 2,460 m asl  
TypesHolotype. MUSA 5573, an adult male (Figs. 2–3), collected on 10 November 2013, by C.S. Abdala, R. Gutiérrez, A. Quiroz, L. Huamani, and J. Cerdeña.
Paratypes. MUSA 5574–75, six adult females, same data as holotype. MUSA 1766, from Quebrada Tinajones, 300 m southeast of holotype (16°31’54.29”S, 71°38’57.547”W) at 2,492 m asl, collected on 9 October 2010, by A. Quiroz and J. Cerdeña. MUBI 13522, MUSA 1767, from Quebrada Tinajones, 600 m southeast of holotype (16°31’54.207”S, 71°38’46.187”W) at 2,528 m asl, collected on 9 October 2010, by A. Quiroz and J. Cerdeña. MUBI 14680, from Quebrada Tinajones (16°31’22.705”S, 71°37’35.666”W) at 2,561 m asl, collected on 27 July 2007, by R. Gutiérrez and A. Quiroz. 
DiagnosisDiagnosis: We assign Liolaemus anqapuka sp. nov. to the L. montanus group because it presents a blade-like process on the tibia, associated with the hypertrophy of the tibial muscle tibialis anterior (Abdala et al. 2020; Etheridge 1995) and its placement in the morphological and molecular phylogenies (Fig. 11). Within the L. montanus group, Liolaemus anqapuka sp. nov. differs from L. andinus, L. annectens, L. aymararum, L. cazianiae, L. chlorostictus, L. dorbignyi, L. fabiani, L, forsteri, L. foxi, L. gracielae, L. huayra, L. inti, L. jamesi, L. melanogaster, L. multicolor, L. nigriceps, L. orientalis, L. pachecoi, L. pantherinus, L. patriciaiturrae, L. pleopholis, L. polystictus, L. puritamensis, L. qalaywa, L. robustus, L. scrocchii, L. signifer, L. vallecurensis, L. victormoralesii, L. vulcanus, and L. williamsi, for being species of larger size (SVL greater than 75 mm) unlike L. anqapuka sp. nov., which has a maximum SVL of 73.5 mm. Liolaemus anqapuka sp. nov., has between 58 and 72 (mean = 64.8) scales around the body, which differentiates it from species of the group with more than 80 scales, such as L. cazianiae, L. duellmani, L. eleodori, L. erguetae, L. forsteri, L. gracielae, L. molinai, L. multicolor, L. nigriceps, L. patriciaiturrae, L. pleopholis, L. poecilochromus, L. porosus, L. pulcherrimus, L. robertoi, L. rosenmanni, L. ruibali, and L. vallecurensis; and also from species with less than 55 scales, like L. aymararum, L. jamesi, L. pachecoi, and L. thomasi. Liolaemus anqapuka sp. nov. have 60–72 dorsal scales (mean = 65.5), and differs from L. andinus, L. cazianiae, L. eleodori, L. erguetae, L. forsteri, L. foxi, L. gracielae, L. halonastes, L. molinai, L. multicolor, L. nigriceps, L. patriciaiturrae, L. pleophlolis, L. poecilochromus, L. porosus, L. pulcherrimus, L. robertoi, L. rosenmanni, L. ruibali, L. schmidti, and L. vallecurensis, which have between 75–102 dorsal scales. The number of ventral scales between 73–87 (mean = 81.3) differentiates it from species with more than 90 ventral scales, such as L. andinus, L. cazianiae, L. erguetae, L. eleodori, L. foxi, L. gracielae, L. halonastes, L. hajeki, L. molinai, L. nigriceps, L. patriciaiturrae, L. pleopholis, L. poecilochromus, L. porosus, L. robertoi, L. rosenmanni, and L. vallecurensis. Liolaemus anqapuka sp. nov. has juxtaposed or subimbricate dorsal scales, without keel or mucron, this differentiates it from species with conspicuous keel and mucron, as L. aymararum, L. etheridgei, L. famatinae, L. fittkaui, L. griseus, L. huacahuasicus, L. montanus, L. orko, L. ortizi, L. polystictus, L. pulcherrimus, L. qalaywa, L. signifer, L. tajzara, L. thomasi, L. victormoralesii, and L. williamsi. Females of L. anqapuka sp. nov. present 1–4 (mean = 2.6) precloacal pores, this character differentiates it from species like L. andinus, L. balagueri, L. fittkaui, L. multicolor, L. ortizi, L. polystictus, L. puritamensis, L. robertoi, L. robustus, L. rosenmanni, L. ruibali, L. thomasi, and L. vallecurensis, because they do not present precloacal pores in females.
Liolaemus anqapuka sp. nov. belongs to the clade of Liolaemus reichei sensu Abdala et al. (2020). The color pattern of Liolaemus anqapuka sp. nov. has a combination of characteristics in males and females that distinguish it from the rest of the Liolaemus of the group. The number of scales around the body is between 58–72 (mean = 64.8), which differentiates it from L. audituvelatus, L. balagueri, L. insolitus, and L. reichei (Table 3). The number of dorsal scales varies between 60–72 (mean = 65.5), which is lower than the number in L. audituvelatus, higher than in L. nazca, and has a variation in range of scales different than L. chiribaya, L. reichei, and L. torresi (Table 3). The numbers of ventral scales of Liolaemus anqapuka sp. nov. vary between 73–87 (mean = 81) which are different from L. audituvelatus, L. nazca, and L. torresi (Table 3). The presence of precloacal pores in females 1–4 (mean = 2.6), is different from L. audituvelatus, L. balagueri, and L. reichei, whose females do not have precloacal pores (Table 3). Coloration patterns on lateral sides have light blue scales, which are different from L. audituvelatus, L. balagueri, L. nazca, L. torresi, and L. reichei (Table 3). The existence of dorsal body scales with a keel differentiate it from L. nazca which have dorsal body scales without keel. Ventral thigh scales with keel are present in 100% of individuals of L. anqapuka sp. nov. but they are less evident than those present in L. chiribaya, where only 35% of individuals present this character (Table 3). The maximum SVL is greater than in L. audituvelatus, L. poconchilensis, L. reichei, L. stolzmanni, and L. torresi (Table 3 in HUAMANI-VALDERRAMA et al. 2020).

Color variation in life (Figs. 4–5). Liolaemus anqapuka sp. nov. shows evident sexual dichromatism. In males, head is darker than the gray body. In some specimens, supralabial and infralabial scales are generally lighter gray than the rest of the head. The subocular is generally white with irregular dark spots. The dorsal color of the neck is gray, varying in its hue, and may be dotted with some light blue scales and orange spots. The body color is always gray. The vertebral region in most males is well delimited with some light blue scales. No vertebral line, dorsolateral bands, antehumeral arch, or scapular spots. Few specimens have diffuse gray paravertebral spots, and rounded shape. As in the holotype, in the paravertebral, dorsolateral, and lateral regions of the body, irregular orange spots stand out, surrounded and dotted with celestial scales. Orange spots can vary in intensity and size, as light blue scales that can form thin irregular lines or clump together to form more conspicuous spots. In some specimens the amount of light blue scales is so remarkable that they cover the orange spots. Orange spots and light blue scales are distributed on the sides of the tail. In some individuals, the celestial scales reach the distal end of the tail. In some specimens, light blue scales are replaced by dark, bluish-green scales. In some, irregularly shaped white spots are distributed among the orange spots. The fore and hind limbs, as well as the tail, have the same design as the body. In the tail, incomplete rings of dark spots with light edges are formed. Ventrally, the majority of males are similar. The predominant color is white, some have faint yellow and a yellow hue that can vary in intensity, highlighted in the gular region and the hind limbs. On the sides of the belly, a thin orange longitudinal line protrudes from the armpit to the groin (Fig. 4).
Females have a totally different coloring pattern than males (Fig. 5). The color of the head varies from brown to gray, with some dark red spots and scales. The supralabial, infralabial, and lorilabial scales are lighter in color than the dorsal surface of the head. The back of the body can be light gray or brown; with small paravertebral spots, gray or dark brown, and circular or sub-quadrangular; with a small white spot on the back which can be the same size as the paravertebral; and with meager orange spots between the paravertebrals. A few females have light blue scales on paravertebral spots. On the sides of the body, there may be lateral spots of the same design as the paravertebral ones. The tail and hind limbs have the same design and color as the body, without dorsolateral bands. Ventrally they are white or faint yellow immaculate throughout the body. In some females, the tail has more intense yellow throughout its extension (Fig. 5 in HUAMANI-VALDERRAMA et al. 2020).

Morphological variation. Twenty-two specimens (six males and 16 females). Dorsal surface of head rough with 14–21 scales (mean = 16.82; STD = 1.71). Nasal surrounded by 6–9 scales (mean = 7.41; STD = 0.73). Supralabials 7–10 scales (mean = 8.18; STD = 0.8), lorilabials 8–11 scales (mean = 9.32; STD = 0.89). A line of lorilabial scales. Supraoculars 4–6 (mean = 5.45; STD = 0.6). Interparietals smaller than parietals, surrounded by 4–8 scales (mean = 6.32; STD = 1.09). Infralabials 6–9 (mean = 7.14; STD = 0.77). Gulars 28–39 (mean = 33.41; STD = 2.99). Temporals smooth, 7–10 scales (mean = 9.09; STD = 0.97). Meatus auditory higher 1.37–2.47 mm (mean = 2.05; STD = 0.26), than wide 0.20–1.20 (mean = 0.81; STD = 0.25). Head longer 12.32–17.20 (mean = 14.91; STD = 1.31) than wide 9.15–15.92 (mean = 12.77; STD = 2.03). Head height 6.84–10.48 (mean = 8.38; STD = 0.87). Underarm to groin length 21.61–32.8 (mean = 28.58; STD = 2.76). SVL males 56.23–73.53 mm (mean = 65.05 mm; STD = 7.08) and females 52.15–71.10 mm (mean = 62.9 mm; STD = 4.61). Femur length 10.11–14.65 mm (mean = 12.31 mm; STD = 1.06). Humerus length 7.56–11.01 mm (mean = 8.86 mm; STD = 0.99). Forearm length 7.65–11.56 mm (mean = 9.59 mm; STD = 1.06). Hand length 8.03–11.25 (mean = 10.25; STD = 0.86). Scales around midbody 58–72 (mean = 65.09; STD = 3.7). Dorsal 60–72 (mean = 65.59; STD = 3.5), juxtaposed to sub-juxtaposed, and smooth scales. Infradigital lamellae of the 4th finger of the hand 15–21 (mean = 17.73; STD = 1.45) and of the 4th toe 20–26 (mean = 21.67; STD = 1.5). Ventral 73–87 (mean = 81.32; STD = 3.37) larger than dorsal scales. Tail length 46.77–67.16 mm (n = 17, mean = 56.83 mm; STD = 5.91). Males with 4–6 (mean = 4.67; STD = 0.82) precloacal pores, and females with 3–5 (mean = 4.22; STD = 0.83) precloacal pores. Body measurements, males (mean = 66.62 mm) slightly larger than females (mean = 62.90 mm), tail length in males slightly larger (mean = 61.74 mm) than females (mean = 54.80 mm) [Table 4 in HUAMANI-VALDERRAMA et al. 2020]. 
CommentSynonymy after HUAMANI-VALDERRAMA et al. 2020. 
EtymologyThe specific name refers to the coloration patterns of males. The word “anqapuka” is an original word in the Quechua language (spoken currently in the Peruvian Andes), corresponding to a complex word between “anqa” assigned to the blue color, and “puka” which means orange or red color. 
  • Abdala, Cristian Simón; Andrés Sebastián Quinteros, Romina Valeria Semhan, Ana Lucia Bulacios Arroyo, James Schulte, Marcos Maximiliano Paz, Mario Ricardo Ruiz-Monachesi, Alejandro Laspiur, Alvaro Juan Aguilar-Kirigin, Roberto Gutiérrez Poblete, Pabl 2020. Unravelling interspecific relationships among highland lizards: first phylogenetic hypothesis using total evidence of the Liolaemus montanus group (Iguania: Liolaemidae). Zoological Journal of the Linnean Society 189 (1): 349–377 - get paper here
  • Boulenger, G.A. 1885. Catalogue of the lizards in the British Museum (Natural History). Vol. 2, Second edition. London, xiii+497 pp. - get paper here
  • Cei & PÉFAUR 1982. Una nueva especie de Liolaemus (Iguanidae:Squamata): su sistemática, ecología y distribución. Act. 8th Congr. Latinoam. Zool. 2: 573-586
  • Etheridge,R. 1995. Redescription of Ctenoblepharys adspersa Tschudi, 1845, and the Taxonomy of Liolaeminae (Reptilia: Squamata: Tropiduridae). American Museum Novitates 3142: 1-34 - get paper here
  • Gutiérrez RC, Quiroz A. 2010. Herpetofauna del sur del Perú. apparently online only
  • Huamaní-Valderrama L, Quiroz AJ, Gutiérrez RC, Aguilar-Kirigin A, Huanca-Mamani W, Valladares-Faúndez P, Cerdeña J, Chaparro JC, Santa Cruz R, Abdala CS. 2020. Some color in the desert: description of a new species of Liolaemus (Iguania: Liolaemidae) from southern Peru, and its conservation status. Amphibian & Reptile Conservation 14(3) [Taxonomy Section]: 1–30 (e250) - get paper here
  • 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 - get paper here
  • Nuñez H. 2004. Cambios taxonómicos para la herpetofauna de Argentina, Bolivia y Chile. Noticiario Mensual Nacional de Historia Natural de Chile 353: 28–34
  • Pefaur, J.E., Davila, J., Lopez, E., & Nunez, A. 1978. Distribucion y clasificacion de los reptiles del Departamento de Arequipa [Chile]. Instit. Francais d'Etudes Andines, Lima, Bull. 7(1-2): 129-139.
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