Liolaemus warjantay, Ubalde-Mamani & Gutiérrez & Chaparro & Aguilar-Kirigin & Cerdeña & Huanca-Mamani & Cárdenas-Ninasivincha & Lazo-Rivera & Abdala, 2021

Ubalde-Mamani, Misshell D., Gutiérrez, Roberto C., Chaparro, Juan C., Aguilar-Kirigin, Alvaro J., Cerdeña, José, Huanca-Mamani, Wilson, Cárdenas-Ninasivincha, Stefanny, Lazo-Rivera, Ana & Abdala, Cristian S., 2021, A new species of Liolaemus (Squamata: Liolaemidae) from the Reserva Paisajística Subcuenca del Cotahuasi, southwestern Peru, Amphibian & Reptile Conservation (e 287) 15 (2), pp. 172-197 : 175-181

Liolaemus warjantay sp. nov. ( Fig. 1 )

2008 Liolaemus signifer annectens, AEDES Guía de Anfibios y Reptiles. Reserva Paisajística Subcuenca del Cotahuasi

2020 Liolaemus aff. qalaywa, Huamaní-Valderrama et al. Amphibian & Reptile Conservation

2021 Liolaemus aff. qalaywa, Quiroz et al. Zoological Studies

urn:lsid:zoobank.org:act:33CAECD9-B6E0-49B7-8F39-18AEEEB547AB

Holotype. MUSA 5700 , an adult male ( Figs. 1 , 2C–E , and 4A, D, G, J, M, P ), from 6.4 km NE of Pampamarca, District of Pampamarca , Province of La Unión, Department of Arequipa, Peru (15°5’41.24”S 72°57’7.06”W) at 4,529 m asl, collected on 20 December 2019, by M. Ubalde and L. Arapa.

Paratypes. Thirteen specimens. All specimens belong to District of Pampamarca, Province of La Unión, Department of Arequipa, Peru. Six adult males: MUSA 5691–92 from 4.1 km NE of Pampamarca, (15°9’37.79”S, 72°55’30.10”W) at 4,500 m asl, collected on 6 August 2019, by M. Ubalde, J. Bedregal, J. Zegarra, L. Cáceres, and E. Guillén. MUSA 5695 from 0.06 km S of holotype (15°5’54.30”S, 72°57’5.49”W) at 4,510 m asl, collected on 19 December 2019, by M. Ubalde and L. Arapa. MUSA 5701–02 and MUBI 17684, same data as holotype. Five adult females: MUSA 5693–94 from 0.2 km S of holotype (15°5’47.80”S, 72°57’5.19”W) at 4,510 m asl, collected on 6 August 2019, by M. Ubalde, J. Bedregal, J. Zegarra, L. Cáceres, and E. Guillén. MUBI 17683, an adult female, from 6.3 km NE of Pampamarca, (15°5’44.14”S, 72°57’6.93”W) at 4,503 m asl, collected on 6 August 2019, by M. Ubalde, J. Bedregal, J. Zegarra, L. Cáceres, and E. Guillén. MUSA 5696 and 5699 from 6.2 km NE of Pampamarca (15°5’42.68”S, 72°57’8.28”W) at 4,504 m asl, collected on 19 December 2019, by M. Ubalde and L. Arapa. One subadult female: MUSA 5698 from 0.4 km S of holotype (15°5’55.81”S, 72°57’6.35”W) at 4,501 m asl, collected on 19 December 2019, by M. Ubalde and L. Arapa.

Diagnosis. We assign Liolaemus warjantay 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 ( Etheridge 1995; Abdala et al. 2020) and based on molecular ( Fig. 6 ) and morphological evidence ( Fig. 5 ). The species of the L. montanus group differ from those of the L. boulengeri group by the complete absence of patches of enlarged scales in the posterior part of the thigh ( Abdala 2007). Compared to the species of the L. montanus group, Liolaemus warjantay sp. nov. is a robust lizard differing by its larger size (max SVL = 89.56 mm) from Liolaemus andinus , L. anqapuka , L. audituvelatus , L. balagueri , L. cazianiae , L. chiribaya , L. duellmani , L. eleodori , L. erroneus , L. etheridgei , L. evaristoi , L. fabiani , L. famatinae , L. fittkaui , L. foxi , L. gracielae , L. griseus , L. hajeki , L. halonastes , L. huacahuasicus , L. insolitus , L. montanus , L. multicolor , L. nazca , L. omorfi , L. orko , L. ortizi , L. pantherinus , L. poconchilensis , L. poecilochromus , L. porosus , L. pulcherrimus , L. reichei , L. robertoi , L. rosenmanni , L. ruibali , L. schmidti , L. stolzmanni , L. tajzara , L. thomasi , L. torresi , L. vallecurensis , L. williamsi , and L. yarabamba (all with SVL between 50–80 mm). The presence of imbricate dorsal scales with keels differentiates L. warjantay sp. nov. from species with smooth juxtaposed or sub-imbricate scales, such as Liolaemus andinus , L. audituvelatus , L. balagueri , L. cazianiae , L. chiribaya , L. eleodori , L. fabiani , L. foxi , L. gracielae , L. halonastes , L. insolitus , L. jamesi , L. nigriceps , L. omorfi , L. patriciaiturrae , L. pleopholis , L. poconchilensis , L. poecilochromus , L. porosus , L. reichei , L. robertoi , L. robustus , L. rosenmanni , L. ruibali , L. schmidti , L. scrocchii , L. torresi , L. vallecurensis , L. victormoralesii , and L. vulcanus .

The new species differs from Liolaemus chiribaya , L. evaristoi , L. etheridgei , L. insolitus , L. multicolor , L. omorfi , L. poconchilensis , L. pulcherrimus , L. robertoi , L. ruibali , and L. schmidti , by the absence of sky blue or light blue scales on the sides and dorsum of the body and tail. The number of scales around midbody in L. warjantay sp. nov. varies between 55–64 (mean = 60.3), which differentiates it from several species of the group with more than 65 scales, such as L. andinus , L. audituvelatus , L. cazianiae , L. duellmani , L. eleodori , L. erroneus , L. forsteri , L. foxi , L. gracielae , L. halonastes , L. inti , L. multicolor , L. nigriceps , L. patriciaiturrae , L. pleopholis , L. poecilochromus , L. porosus , L. pulcherrimus , L. robertoi , L. rosenmanni , L. ruibali , L. schmidti , L. multiformis , and L. vallecurensis . The number of ventral scales between the mental scale and the border of the vent in L. warjantay sp. nov. varies between 72–85 (mean = 78.2), and is lower than the number in the following species (with more than 90 ventral scales): L. andinus , L. cazianiae , L. erroneus , L. foxi , L. gracielae , L. halonastes , L. inti , L. multicolor , L. nigriceps , L. pachecoi , L. patriciaiturrae , L. pleopholis , L. poecilochromus , L. porosus , L. robertoi , L. rosenmanni , L. torresi , and L. vallecurensis ; and higher than the number in the following species (with less than 70 ventral scales): L. dorbignyi , L. fittkaui , L. melanogaster , L. polystictus , and L. thomasi . The number of dorsal scales of L. warjantay sp. nov. varies between 45–63 (mean = 52.8), while the species with more than 70 scales are L. andinus , L. audituvelatus , L. cazianiae , L. duellmani , L. eleodori , L. erroneous , L. fabiani , L. famatinae , L. forsteri , L. foxi , L. gracielae , L. halonastes , L. multicolor , L. nigriceps , L. orko , L. patriciaiturrae , L. pleophlolis , L. poecilochromus , L. porosus , L. pulcherrimus , L. robertoi , L. rosenmanni , L. ruibali , L. schmidti , L. torresi , and L. vallecurensis ; and the species with less than 45 dorsal scales are L. jamesi and L. pachecoi .

Only one female was found with two vestigial precloacal pores, that differentiates Liolaemus warjantay sp. nov. from species that do have pores in all females and in greater quantity as in L. aymararum , L. cazianiae , L. chiribaya , L. chlorostictus , L. dorbignyi , L. eleodori , L. erroneus , L. etheridgei , L. fabiani , L. famatinae , L. griseus , L. hajeki , L. huayra , L. huacahuasicus , L. inti , L. jamesi , L. montanus , L. nazca , L. nigriceps , L. orko , L. pachecoi , L. pantherinus , L. patriciaiturrae , L. porosus , L. pulcherrimus , L. qalaywa , L. scrocchii , L. multiformis , and L. vulcanus .

The color pattern clearly differentiates the new species from Liolaemus yauri , mainly the dorsal color of the head in both sexes of L. warjantay sp. nov. is dark grey and always darker than body, while in L. yauri the coloration is lighter and not in contrast with the body color; the color of the palpebral scales in females of L. warjantay sp. nov. is pale yellow, and in L. yauri is chestnut or grey; the shapes of the paravertebral spots in both sexes of L. warjantay sp. nov. are in the form of thin transversal lines and curved posteriorly (ocelli-shaped), while in L. yauri they are circular rhomboid or sub-quadrangular.

Liolaemus warjantay sp. nov. can be distinguished from L. annectens and L. qalaywa (two geographically close species) by a combination of the following characters: trifid scales from the plantar surface, absence of pores in the base of the tail, presence of gular fold, and the presence of ocelli in males ( Table 2 , Fig. 4 ). The analysis of DNA sequences of L. warjantay sp. nov. reveals differences of 2.4–5.1% with L. qalaywa , and 9.2–9.5% with L. annectens .

Description of the holotype ( Fig. 1 ). Adult male (MUSA 5700), SVL 89.56 mm. Head greater in length (22.45 mm) than width (17.02 mm). Head height 12.51 mm. Neck width 21.43 mm. Eye diameter 3.83 mm. Interorbital distance 13.33 mm. Auditory meatus elliptical 5.97 mm high, 1.98 mm wide. Internasal width 3.52 mm. Subocular scale length 6.18 mm. Trunk length 34.53 mm, width 28.64 mm. Tail length 122.3 mm. Tail width 15.08 mm (cloaca level). Femur length 20.53 mm, tibia 18.61 mm, and foot 26.01 mm. Humerus length 12.03 mm, width 8.43 mm. Forearm length 8.43 mm, width 17.54 mm. Hand length 15.84 mm. Fourth finger length of the foot 15.52 mm. Pygal region length 8.89 mm, and cloacal region width 12.31 mm. Nasal separated from rostral by one scale. Two internasals slightly longer than wide. Nasal surrounded by seven scales, separated from canthal by one scale. Six scales between frontal and rostral. Frontals divided into four scales. Interparietal surrounded by eight scales. Preocular separated from lorilabials by one scale. Five superciliaries and 15 upper ciliary scales, lower ciliaries are neither projected nor open between them. Three differential scales at anterior margin of auditory meatus and a large diagonal auricular scale. Nine temporary granular scales. Five lorilabial scales, in contact with subocular.Eight supralabials, which are not in contact with subocular. Five supraocular. Ten lorilabials, five scales are in contact with the subocular scale and separated by a scale from the preocular. Six infralabials. Four scales around the mental scale. Four scales in contact with the second infralabial scale, and six scales separate the fourth shields. Dorsal head with 15 scales, 42 scales up to the neck, 24 up to the antehumeral fold (following the longitudinal fold), 59 scales around the body, 51 dorsal from the occiput to the hind limbs. The dorsal scales are triangular, with a pronounced keel, mucron, and imbricate. With 79 ventral scales, eight pygals, and seven precloacal pores.

Four chin shields, 4 th pair separated by six scales. Seventy scales around half a body. Fifty-nine triangular dorsal body scales, imbricated, and with a keel and mucron; laminar anterior on members, imbricate and slightly keeled; laminar on hind limbs, imbricate and slightly keeled; tail with dorsal scales juxtaposed. Seventy-nine ventral scales, from the mental to the cloacal region, following the ventral midline of the body, laminar, imbricated. Thirty-seven imbricate gulars, smooth. Neck with longitudinal fold with 42 granular, not-keeled scales, ear fold and antehumeral fold present. Forelimbs ventrally laminar, subimbricate to imbricate, with keeled scales; hind legs laminar, imbricate, with keeled scales. Twenty subdigital lamellae on the 4 th finger of the hand. Twenty-two subdigital lamellae of the 4 th toe, with three keels, plantar trifid scales with keels and mucrons. Lamellar ventral scales on tail, imbricate, not keeled. Seven precloacal pores. Supernumerary pores absent.

Color of holotype in life ( Figs. 2D–F ). Head completely melanic. Temporal region with clear edges, supralabial, infralabial, and lorilabial scales are gray with black spots. Palpebral scales pale yellow. Neck dorsally black and yellow on the sides with some black or dark hues. Body uniform brownish-yellow color, vertebral field not defined, vertebral line and dorsolateral bands absent. Paravertebral spots diffuse, imperceptible, black in color, and in the form of thin transversal lines, curved posteriorly. These spots project to the sides of the body, which are lighter yellow in color with no obvious spots. Front and rear legs brown with yellow hues and dark scales. Fingers light gray. Tail of the same color as the body, a little lighter on the sides and at its distal end. Venter light gray or whitish throughout the body, with some dark shades in the center of the abdomen and yellow on the sides of the abdomen.

Morphological variation ( Table 1 ). Thirteen specimens including MUSA 5691–96 , MUSA 5698–5702 , and MUBI 17683–84 (seven males and six females). Considering both sexes, individuals of L. warjantay sp. nov. reach a maximum snout-vent length of 95.12 mm, with males tending to be larger than females ( SVL male mean: 85.98 mm; SVL female mean: 78.39 mm) ( Table 1 ). Other characters: A line of lorilabial scales. Dorsal scales juxtaposed, triangulars the majority and with keeled scales between occiput and hindlimb insertion. Parietals slightly smaller than interparietals. Occiput scales granular or conical in males and granular in females. Nasal and canthal separated by two scales. Upper ear border with enlarged anterior diagonal scale. Temporary scales granular and without keel. Subocular in contact with three to five lorilabials. Second right infralabial scale in contact with four or five scales. Dorsal body scales, subimbricated to imbricate, males with dark, light and triangular scales, females with dark scales and rounded or triangular posterior border. Ventral scales, imbricated in the gular, pectoral, abdominal, and pygal region. Precloacal pores evident in males, only one female with two small pores. Dorsal scales at the base of the tail are imbricated. Dorsal scales on forelimbs imbricate without keel or slight keel. Dorsal scales on hind limbs imbricate and with a slight to strong keel, only one female with subimbricate. Heteronotes in the region where the femoral patch would be present. Palmar scales, imbricated and with a triangular posterior border. Plantar scales, with slight or strong keel and triangular rear edge, one male rounded and without keel and one female without keel. Subdigital lamellae of the fourth toe with three keels.

Color variation in life ( Figs. 2–3 ). Liolaemus warjantay sp. nov. shows evident sexual dichromatism. In males, head and temporal region are gray or dark brown, always darker than body. Lorilabial, supralabial, and infralabial scales are always lighter in color than the rest of the head. Color of body is highly variable, varying from brown to dark gray. Paravertebral spots are dark and can vary in shape and intensity. Most of the specimens have the shape of light ocelli with two edges, one light internal and the other dark external, which can vary in intensity and thickness. These edges can project to the sides of the body. Paravertebral spots are more evident in juvenile specimens, and in larger males they can be very diffuse or imperceptible. No vertebral line, dorsolateral bands, antehumeral arch, or scapular spots. The sides of the body are lighter than the back, and can vary from yellow to orange. No lateral spots, with small circular spots, with yellow orange scales. No blue scales anywhere on the body. The forelimbs and hindquarters are generally lighter in color than the body and with yellow or black dots or scales. Tail with spots and lines of the body change merging or fading until they are completely lost by approximately the first fifth of the tail, then they become lighter or darker in color than the rest of the body ( Fig. 2 ). Ventrally, the color is variable, some males are completely yellow, orange, or white; some with dark spots or scales on different parts of the body. Females have a similar design as males, but with less dramatic colors ( Fig. 3 ). Head color also varies from chestnut to dark gray. Supralabial, infralabial, and lorilabial scales are lighter in color than the back of the head, with gray being the most common color. Dorsal body is light to dark brown. Paravertebral spots are also ocelli-shaped, but they are black or dark brown with a white border. These ocelli are more evident in smaller females and may be absent in larger ones. Some females with edges of the paravertebral spots projecting to the sides of the body in a “zigzag” line shape, with no obvious vertebral line or dorsolateral bands. Lateral body is generally the same color as dorsum. No lateral spots. Tail and hind limbs have the same design and color as the body, however, they are lighter in color after the first third. Immaculate white underneath. Some have dark spots or scales on different parts of the body.

Distribution and natural history ( Figs. 7–8 ). Liolaemus warjantay sp. nov. is restricted to the type locality, Pampamarca ( Fig. 8 ), in the RPSCC, Department of Arequipa, Peru, at elevations between 4,500 –4,529 m asl ( Fig. 7 ). This species inhabits high Andean dry puna ( Fig. 8 ), where the climate has hostile conditions due to the high elevations, with a wide range of temperatures (-8.9–14.6 ºC), and an annual average of 4 ºC (WorldClim database, based on collected information on environmental variables for 30 years) ( Fick and Hijmans 2017). Individuals were registered and collected during the dry and wet seasons (July–December), in natural rocky areas with rocks which varied in size ranging between 30–200 cm, small bushes (Parasthrephia sp.), grassland ( Stipa sp. and Festuca sp. ), and to a lesser extent an area of yareta ( Azorella sp. ). The peak of lizard activity was during 1100–1300 h. In the month of July, 100% of the adult females observed were pregnant; while in December, no pregnant females were observed, but there were juvenile individuals. In most cases, the juveniles were associated with shelters that were channels under the rocks (approximately 40 cm long by 20 cm high) and were observed together with adult females in some cases, and the number of juveniles observed in each refuge was between 3–6 individuals. The potential predators of L. warjantay sp. nov. are: American Kestrel (Falco sparverius), Andean Fox ( Lycalopex culpaeus ), and snakes ( Tachymenys peruviana ), which were observed during field work in the type locality of L warjantay sp. nov., and these species are known to include a percentage of lizards in their diets ( Jaksić et al. 1981, 1982; Guzmán-Sandoval et al. 2007; Walker et al. 2007; Santillán et al. 2009; Miranda et al. 2015; Pozo-Zamora et al. 2017).

Etymology. The specific name in the Quechua language ( “warjantay ”) refers to the local name in RPSCC and its surroundings assigned to the high Andean lizards of the genus Liolaemus .

Phylogeny. The morphology-based phylogeny presented here ( Fig. 5 ), performed with all the values of the concavity constant (K), indicates that Liolaemus warjantay sp. nov. belongs to the L. montanus group, and within this to the L. ortizi clade ( Abdala et al. 2020), and it is recovered as sister taxon of L. qalaywa . The molecular analysis ( Fig. 6 ) shows that the terminals of L. warjantay sp. nov. (MUBI 17683, MUSA 5692, MUSA 5685, VOI 009, and VOI 006) form a monophyletic subclade. This was also recovered as a sister of L. qalaywa in a clade with a terminal identified as L. signifer sensu lato (MUSM 29110), from Desaguadero, in the Department of Puno, near the Bolivian border, which was mentioned by Chaparro et al. (2020) and might represent a potential new species of the L. montanus group from the Titicaca Andean plateau in southern Peru. The topology of the molecular tree is consistent with previous results using either the same molecular markers ( Chaparro et al. 2020) or only cyt- b ( Huamaní-Valderrama et al. 2020; Quiroz et al. 2021).

Figures

Fig. 1. Holotype of Liolaemus warjantay sp. nov. MUSA 5700 (SVL = 89.56 mm, Tail = 122.3 mm): (A) dorsal and (B) ventral views of body; (C) dorsal, (D) ventral, and (E) lateral views of head; (F) ventral view of precloacal pores; (G) ventral body scales; (H) keeled dorsal body scales. Scale = 5 mm.

Fig. 2. Adult males of Liolaemus warjantay sp. nov. in dorsal, lateral, and ventral views: (A–C) MUSA 5695 (SVL = 88.18 mm); (D–F) MUSA 5700 (SVL = 89.56 mm, Tail = 123.3 mm); (G–I) MUSA 5702 (SVL = 86.21 mm, Tail = 123.56 mm); (J–L) MUBI 17684 (SVL = 86.21 mm, Tail = 123.56 mm); (M–O) MUSA 5702 (SVL = 84.89 mm, Tail = 117.47 mm).

Fig. 4. Comparisonsof distinctive characters between the new species described herein and the phylogenetically and morphologically closest species. Adult males of, Liolaemus warjantay sp. nov. (MUSA 5700, holotype): (A) Infralabials with absent keel, (D) Presence of enlarged scales on sides of gular fold, (G) Palmar scales trifid, (J) Plantar scales trifid, (M) Absence of pores at the base of the tail, (P) Dorsolateral fold present; L. qalaywa (MUBI 13286, holotype): (B) Infralabials barely keeled, (E) Absence of enlarged scales on sides of gular fold, (H) Palmar scales triangular, (K) Plantar scales rounded, (N) Presence of pores at the base of the tail, (Q) Dorsolateral fold absent; L. annectens (LECG 102): (C) Infralabials with absent keel, (F) Absence of enlarged scales on sides of gular fold, (I) Palmar scales trifid, (L) Plantar scales conical, (O) Absence of pores at the base of the tail, (R) Dorsolateral fold present.

Fig. 6. Phylogenetic tree showing the relationships between Liolaemus warjantay sp. nov. and species within the L. montanus group by molecular phylogenetic analysis.

Fig. 5. Phylogenetic tree showing the relationships between Liolaemus warjantay sp. nov. and species within the L. montanus group by morphological phylogenetic analysis.

Fig. 3. Adult females of Liolaemus warjantay sp. nov. in dorsal, lateral, and ventral views: (A–C) MUSA 5699 (SVL = 70.63 mm, Tail = 99.61 mm); (D–F) MUSA 5696 (SVL = 88.14 mm); (G–I) MUSA 5694 (SVL = 85.84 mm, Tail = 118.33 mm); (J–L) MUBI 17683 (SVL = 92.31 mm, Tail = 107.14 mm).

Fig. 7. Geographic distribution showing the type localities of species included in the Liolaemus montanus group in Peru.

Fig. 8. Habitat of the type locality of Liolaemus warjantay sp. nov. in the Department of Arequipa, Peru.