*Liolaemus chiribaya*

2018. *Liolaemus* "Moquegua" Aguilar-Puntriano et al.

Holotype. MUSM 31547: adult male collected near "Cerros Los Calatos", 16.91892S, 70.89596W, Torata District, Mariscal Nieto Province, Moquegua Department, Per ú, 2615 m, December 19, 2014 by César Aguilar, Jessie Montalvo and Maribel Angeles.

Paratypes. MUSM 31553: adult male collected in Jaguay Chico, Torata District, Mariscal Nieto Province, Moquegua Department, Peru 16.94567 S, 70.88486 W, 2928 m, December 19, 2014; MUSM 31549–31550: one female and juvenile collected in the same location and date as previous specimen, 2942 and 2913 m respectively; MUSM 31548: a female collected near the Asirune Archaeological Zone, near Jaguay Chico, Torata District, Mariscal Nieto Province, Moquegua Department, Peru, 16.95213 S, 70.87854 W, 2990 m; all above paratypes collected by César Aguilar, Jessie Montalvo and Maribel Angeles; MUSM 31386–31388, 31390–31391: five males and MUSM 31389: one female collected in "Cerro Los Calatos", Torata District, Mariscal Nieto Province, Moquegua Department, Per ú, 2794–2988 m, 27–29 December 2012 by Juana Suárez.

Referred specimens. BYU 51568, BYU 51570: two males, collected in Jaguay Chico, Torata District, Mariscal Nieto Province, Moquegua Department, Per ú; MUSM 31546: male collected near the Asirune Archaeological Zone, near Jaguay Chico, Torata District, Mariscal Nieto Province, Moquegua Department, Per ú; BYU 51564, BYU 51566–51567: two females and one juvenile with same data as MUSM 31546.

**Figure 2.** Dorsal (**A**) and ventral (**B**) variation in males of *Liolaemus nazca.* From left to right: MUSM 31520 (SVL = 60.9 mm), BYU 50472 (SVL = 64 mm), MUSM 31523 (holotype, SVL = 64.5 mm), BYU 50471(SVL = 60.0 mm), MUSM 31526 (SVL = 53.1 mm).

Diagnosis. *Liolaemus chiribaya* is identified as a member of the *L. montanus* group by the absence of a patch of enlarged scales on posterior thighs. *Liolaemus chiribaya* forms a clade with *L. insolitus*, *L. poconchilensis* and *L. nazca* sp. nov. It differs from *L. poconchilensis* by having a fourth finger extending beyond the armpit when a hindlimb is brought forward (fourth finger does not exceed past the armpit in *L. poconchilensis*); male *L. chiribaya* further differs from *L. poconchilensis* by the presence of dorsal turquoise spots (absent in males of *L. poconchilensis*), and differs from *L. nazca* by having smooth dorsal body scales (dorsal body scales slightly keeled in *L. nazca*). In addition, *L. chiribaya* lacks an orange or yellow venter with dark spots. *Liolaemus chiribaya* differs from *L. insolitus* by having a greater number of scales around midbody (58–69 vs. 45–53). *Liolaemus chiribaya* differs from *L. nazca*, *L. insolitus* and other Peruvian species of the *L. montanus* group (except *L. poconchilensis*) by having a "phrynosauroid" head. *Liolaemus chiribaya* also differs from other Peruvian species of the *L. montanus* group by having dorsal turquoise scales and a maximum 68.8 mm SVL, being a smaller species than *L. aymararum*, *L. evaristoi*, *L. melanogaster*, *L. polystictus*, *L. robustus*, *L. thomasi* and *L. williamsi* (SVL 70.1–103.0 mm). *L. chiribaya* has fewer scales around midbody (54–66) than *L. signifer* (67–110), fewer maximum number of dorsal scales (between occiput and anterior level of hindlimb; 64) than *L. evaristoi* (75) and *L. signifer* (129), and more than *L. aymararum*, *L. ortizii* and *L. thomasi* (all ≤ 53). It also differs from *L. etheridgei*, *L. ortizii* and *L. thomasi* by lacking strongly keeled scales. Females of *L. chiribaya* also have vestigial precloacal pores, which are absent in females of *L. melanogaster*, *L. polystictus* and *L. thomasi*.

**Figure 3.** Dorsal (**A**) and ventral (**B**) variation in females of *Liolaemus nazca.* From left to right: MUSM 31525 (SVL = 61.4 mm), BYU 50506 (SVL = 66.1 mm), BYU 50507 (SVL = 54.9 mm), MUSM 31521 (SVL = 56.4 mm), BYU 50508 (SVL = 52.6 mm), BYU 50510 (SVL = 47.2 mm).

**Figure 4.** Habitat of *Liolaemus nazca.*

Description of the holotype. Adult male, SVL 52.6 mm, head length 14.2 mm, head width 11.7 mm, head height 7.6 mm, groin armpit distance 20.3 mm (38.6% SVL), foot length 14.3 mm (27.2 % SVL), tail length 54.2 mm (103% SVL). 21 dorsal scales on head; smooth dorsal head scales, occipital, parietal and frontonasal area with convex scales, parietal scales polygonal; scale organs numerous in prefrontal, lorilabial and loreal scales; supralabial scales with few scales organs; five organs in the left postrostral and four in the right. Nasal scale separated from rostral, separated by a scale from first supralabial, right nasal limited by eight scales; canthal separated from nasal by two scales. Seven supralabials. Eight lorilabial scales, three in contact with the subocular. Seven infralabials. Oval auditory meatus (height 1.8 mm; width 1.2 mm), with two small protruding scales on anterior margin. Eight smooth and convex temporal scales. Distance between orbit and auditory meatus 4.1 mm. Rostral three times wider than high (width 2.4 mm; height 0.8 mm). Mental subpentagonal, almost twice as wide as high (width 2.4 mm; height 1.3 mm). Interparietal pentagonal, with elongated posterior apex, bordered by five scales, parietals larger than interparietal. Frontal scales polygonal. Complete supraorbital semicircles on both sides. Semicircles formed by 13 scales. Three enlarged supraoculars. Eight superciliares. 14 upper and 12 lower ciliaries. Subocular divided into three, longer than eye diameter, posterior subocular larger and separated from supralabials by a single row of lorilabials. Supralabials of similar size. Lorilabial eighth and seventh in contact with subocular. Preocular of medium size, separated from lorilabial row by a scale. Postoculars of similar size to preocular. Mental scale in contact with four scales: two infralabials (on each side) and two enlarged chin scales. Chin scales forming a longitudinal row of four enlarged scales separated from each other by 12 scales. Gular scales rounded, flat and overlapped with very few scale organs. 25 gular scales between auditory meatus. Longitudinal neck fold without keeled scales, almost half in size of dorsal scales. Antehumeral pocket and well developed antehumeral neck folds. 30 scales between auditory meatus and shoulder (counting along post-auricular and longitudinal neck fold), 25 scales between the auditory meatus and the neck fold. Gular fold absent. Dorsal scales juxtaposed or poorly imbricated. 60 dorsal scales between occiput and groin level. 64 scales around midbody. Dorsal scales smooth on flanks and belly. Ventral scales slightly wider than dorsal scales. 74 ventral scales between mental and cloaca; four precloacal pores present. Smooth laminar supracarpals, with oval or rounded margins. Subdigital lamellae of fingers with three keels, and with formula I:7; II:11; III:15; IV:17; V:8 (right hand). Smooth and imbricated supradigital lamellae. Smooth infratarsals and keeled infracarpals, both strongly imbricated. Supratarsal smooth, oval or rounded. Subdigital lamellae of toes with formula I:9; II:11; III:16; IV:18; V:13 (right foot).

Color pattern in life (Figure 5). Dorsal paravertebral orange and turquoise scales from neck region to tail base. Dorsal head with pale orange and brown scales on occipital and frontal region. Dark brown scales on supraocular, parietal region and snout. Dorsal limbs pale orange, with some orange scales and dark brown scales forming zigzag spots up to base of digits (hand) or on digits (foot). First third of tail with turquoise and orange scales, with dark scales forming transverse bands or spots. Ventral scales on head, limbs and tail white smoke, belly with two orange lateral stripes separated by a central area of pale yellowish scales. Ventral tail white smoke.

Color pattern in preservative. Dorsal light gray scales on vertebral region and dark brown scales on paravertebral region, from neck to tail base. Dorsal head with light gray scales on occipital, parietal, frontal and prefrontal regions. Dark olive scales on supraocular, and some on parietal and temporal region, creamy scales on snout. Dorsal limbs pale gray, with gray scales forming zigzag marks up to base of digits (hand) or on digits (foot). Tail pale gray with dark scales forming transverse bands or spots. Ventral scales white smoke on head, limbs, tail and belly.

Variation. (Figures 5 and 6). Variation in selected characters is summarized in Table 2. Sexual dichromatism present. Females and juveniles on dorsal body have 6 to 8 triangular or quadrangular marks, sometimes bordered by lighter scales; without turquoise blue scales; males with or without triangular or quadrangular marks, but if present marks are usually covered by turquoise blue scales; venter whitish without orange lateral stripes.

**Figure 5.** Dorsal (**A**) and ventral (**B**) variation in males of *Liolaemus chiribaya.* From left to right: MUSM 31547 (Holotype, SVL = 52.6 mm), BYU 51568 (SVL unknown), BYU 51570 (SVL unknown), MUSM 31553 (SVL = 52.7 mm).



Vestigial precloacal pores.

\*

**Figure 6.** Dorsal (**A**) and ventral (**B**) variation in *Liolaemus chiribaya.* From left to right: MUSM 31546 (male, SVL = 52.5 mm), BYU 51564 (female, SVL unknown), MUSM 31548 (juvenile, SVL = 49.6 mm), BYU 51566 (juvenile, SVL unknown), BYU 51567 (juvenile, SVL unknown).

Distribution and natural history (Figure 7). *Liolaemus chiribaya* is only known in the District of Torata, Province of Mariscal Nieto, Department of Moquegua, at elevations of 2615–3005 m; they were found under rocks or on ground in desert areas with cacti and low shrubs; they were active between 10:00 and 14:00. Other lizard species present in the area were *Phyllodactylus gerrophygus* and *L. tacnae*.

*Etymology*. The specific epithet *chiribaya* is a noun used in apposition and honors the Chiribaya culture (900–1350 A.D.). Chiribayans were settled in the basin of the Ilo River, and expanded north to the Tambo valley (Arequipa) and the south to the Azapa valley (Chile), including the high altitude regions, up to nearly 3000 m of elevation.

*Liolaemus victormoralesii*

2017. *Liolaemus* "Abra Toccto" Aguilar et al. 2018.*Liolaemus*"AbraToccto"Aguilar-Puntrianoet

 al.

Holotype. MUSM 31461: adult male collected at Abra Toccto, Huamanga Province, Ayacucho Department, Perú, 13.346 S, 74.184 W, elevation 4222 m, on 01 June 2012 by César Aguilar, Víctor J. Vargas, Frank Huari and Elver Coronado.

Paratypes. MUSM 31371–31372, 31460: three adult males collected at Abra Toccto, Huamanga Province, Ayacucho Department, Perú, 13.298 S, 74.091 W, elevation 4193–4215 m on 3 December 2012 by Alfredo Guzmán and Víctor J. Vargas; MUSM 31460, 31468: two adult females, same data as holotype; MUSM 31463: juvenile collected at Abra Toccto, Huamanga Province, Ayacucho Department, Perú, 13.35 S, 74.187 W, elevation 4182m, on 04 June 2012 by César Aguilar, Víctor J. Vargas, Frank Huari and Elver Coronado. MUSM 25700, adult male collected at Chiara, Huamanga Province, Ayacucho Department, Perú, 13.341 S, 74.216 W, elevation 4145m, on 30 November 2006 by Margarita Medina.

**Figure 7.** Habitat of *Liolaemus chiribaya*.

Referred specimens. BYU 50431, BYU 50427, MUSM 31462: three males, same data as holotype; BYU 50433, BYU 50428: two females, same data as holotype.

Diagnosis. *Liolaemus victormoralesii* is identified as a member of the *L. montanus* group by the absence of a patch of enlarged scales on posterior thighs. *Liolaemus victormoralesii* forms a clade with *L. evaristoi*, *L. melanogaster*, *L. polystictus*, *L. robustus* and *L. williamsi*. It differs from closely related *L. evaristoi* by lacking blue scales on the dorsum and flanks, having a larger size (maximum SVL 88.9 mm in *L. victormoralesii* and 70.1 mm in *L. evaristoi*) and by lacking vestigial precloacal pores in females. *Liolaemus victormoralesii* differs from *L. melanogaster* by lacking black belly scales (gray scales in adult *L. victormoralesii*). Adult females of *L. victormoralesii* differ from *L. polystictus* and *L. williamsi* females by having a darker dorsal background coloration and few large contrasting marks dorsally (*L. polystictus* and *L. williamsi* have a lighter dorsal background coloration and large number of small contrasting marks dorsally). *Liolaemus victormoralesii* further differs from *L. williamsi* by having a larger size (maximum SVL 74.9 mm in *L. williamsi*) and by lacking vestigial precloacal pores in females. *Liolaemus victormoralesii* differs from *L. robustus* by lacking dorsal yellow greenish scales. Adult males of *Liolaemus victormoralesii* differs from *L. etheridgei* by lacking light blue dorsolateral scales, from *L. annectens* by having a darker dorsum, and from *L. signifer* by lacking bright yellow and fewer maximum number of dorsal scales (57 vs. 129). *Liolaemus victormoralesii* differs from *L. ortizi* and *L. thomasi* by lacking strongly keeled and by having smaller dorsal scales. *Liolaemus victormoralesii* differs from *L. aymararum* by having more scales around midbody (51–64 vs. 48–52) and smaller dorsal scales. It differs from *L. nazca* sp. nov. by lacking emerald green spots surrounded by black and yellow scales laterally on body. It differs from *L. chiribaya* and *L. poconchilensis* by lacking a "phrynosauroid" head, and from *L. poconchilensis* by lacking well-developed ciliary scales (serrate "combs") surrounding the eyes. It di ffers from *L. insolitus* by lacking dorsal and lateral blue spots. *L. victormoralesii* further di ffers from *L. insolitus* and *L*. *poconchilensis* by lacking dorsal smooth scales. *Liolaemus victormoralesii* also di ffers from *L*. *chiribaya* sp. nov., *L*. *etheridgei*, *L*. *nazca* sp. nov., *L*. *ortizii* and *L*. *poconchilensis* in having a larger SVL (88.9 mm versus 56–77 mm). *Liolaemus victormoralesii* females also di ffers from *L*. *annectens*, *L*. *aymararum*, *L*. *chiribaya*, *L*. *etheridgei*, *L*. *insolitus* and *L*. *nazca* sp. nov by lacking vestigial precloacal pores.

Description of the holotype. Adul male, SVL 83.8 mm, head length 19.2 mm, head width 18.2 mm, head height 10.8 mm, groin armpit distance 29 mm (34.6% SVL), foot length 19.5 mm (23.4 % SVL), tail length 107 mm (128.5% SVL). 19 dorsal scales on head; smooth dorsal head scales, occipital, parietal and frontonasal area with slightly convex scales, parietal scales polygonal, similar in size to interparietal; numerous scale organs in prefrontal and lorilabial, and few in loreal and supralabial scales; six and four scale organs on left and right postrostral respectively. Nasal scale separated from rostral, separated by a scale from first supralabial, right nasal limited by six scales; canthal separated from nasal by one scale. Nine supralabials. Seven lorilabials, six in contact with subocular. Seven infralabials. Oval auditory meatus (height 3.5 mm; width 2.7 mm). Eight smooth temporal scales. Distance between orbit and auditory meatus 9.4 mm. Rostral two times wider than tall (width 3.2 mm; height 1.5 mm). Mental trapezoidal, almost twice as wide as high (width 3.8 mm; height 2.1 mm). Interparietal hexagonal, with elongated posterior apex bordered by six scales, interparietal similar in size as parietals. Frontal divided into two scales. Complete supraorbital semicircles on both sides. Semicircles formed by 16 scales. Seven enlarged supraoculars. Seven superciliares. 14 upper and 12 lower ciliaries. Subocular not divided, longer than eye diameter. Supralabials of similar size. Some lorilabiales with rows of scale organs. Five lorilabials in contact with subocular. Preocular separated from lorilabial row by two scales. Postoculars similar in size as preocular. Mental scale in contact with four scales: one infralabial (on each side) and two enlarged chin scales (one on each side). Six chinshields (three on each side), second pair separated by four scales. Gular scales rounded, flat and without scale organs. 23 gular scales between auditory meatus. Longitudinal neck fold without keeled, but granular scales. Without antehumeral pocket, well developed antehumeral neck folds. 44 scales between auditory meatus and shoulder (counting along post-auricular and longitudinal neck fold). Gular fold absent. Dorsal scales rhomboid, slightly imbricate or juxtaposed, and slightly keeled. 57 scales between occiput and groin level. 58 scales around midbody. Scales of flanks rhomboidal and smooth. Ventrals slightly larger than dorsals, flat, and imbricated. 74 ventral scales between mental and cloaca; five precloacal pores present. Smooth laminar supracarpals, with oval or angular margins. Subdigital lamellae of fingers with three keels, and with formula I:8; II:13; III:16; IV:17; V:12 (right hand). Smooth and imbricated supradigital lamellae. Infratarsals and infracarpals keeled, both strongly imbricated. Supratarsals keeled, triangular or angular in shape. Subdigital lamellae of toes with formula I:8; II:14; III:18; IV:20; V:14 (right foot).

Color pattern in life (Figure 8). Dorsal head between snout and anterior level of eyes brown with black on scale margins, and between anterior level of eyes and occipital region mainly black. Anterior third of dorsal body with a central band of reddish cream scales and posterior two thirds mostly black with reddish cream scale tips; dorsal flanks mostly black with reddish cream scale tips. Lateral body mostly reddish cream but some with scales half black. Dorsal region of tail, anterior and posterior limbs, similar to dorsal body but also with pale blue scales. No vertebral line or scapular spots. Ventral body and tail gray with pale yellow scales, throat region with gray longitudinal stripes, ventral thighs orange.

Color in preservative. Head darker than body, dorsal head mostly dark beige but some scales completely black, parietal zone dark beige. Trunk with scales having black anteriorly and beige posteriorly, scales close to neck with one third black and two thirds beige, scales close to middle and posterior body two thirds black and one third beige. Fore and hind limbs same color as trunk, but with lighter scales close to hands and foot. Without vertebral line, scapular or paravertebral spots nor dorsolateral stripes. Lateral body with scales having one half black anteriorly and other half beige. Dorsal tail similar to trunk, but with lighter scales close to scale tips. Ventrally, from mental scale to tail tip, mostly light gray but some scales cream.

**Figure 8.** Holotype (male, SVL = 83.3 mm, MUSM 31461) of *Liolaemus victormoralesii* in dorsal (**A**) and ventral (**B**) views.

Variation (Figures 9 and 10). Variation in selected characters is summarized in Table 3. Sexual dichromatism present in ventral coloration, with males having yellow scales on thighs. Juveniles exhibit gray dorsal background coloration. Juveniles with black spots on paravertebral region, sometimes enclosing pale orange scales. Females and juveniles ventrally from mental region to tail tip white or white with gray or black scales, some specimens with reddish or orange scales on body venter.

Distribution and natural history (Figure 11). *Liolaemus victormoralesii* is only known in Huamanga Province, Department of Ayacucho, at elevations of 4175-4252 m; *L. victormoralesii* was found under rocks or on the ground in grassland areas. It was found together with *Liolaemus wari* and the snake *Tachymenis peruviana*. Díaz [9] recorded body temperatures of 104 specimens of *Liolaemus victormoralesii* (as *L. <sup>a</sup>*ff*. melanogaster*; mean ± standard deviation): 21.3 ± 6.4 ◦C (substrate temperature: 14.5 ± 4.1 ◦C; air temperature: 13.2 ± 3.4 ◦C). However, lower body temperatures were recorded in the summer: 18.9 ± 6.8 ◦C (substrate temperature: 14.1 ± 3.1 ◦C; air temperature: 12.1 ± 2.7 ◦C); and higher body temperatures were recorded in the fall: 29.9 ± 0.1 ◦C (substrate temperature: 19.5 ± 3.4 ◦C; air temperature: 15.7 ± 2.2 ◦C). *Liolaemus victormoralesii* feeds on Araneae, Acari, Collembola, Scorpiones, Coleoptera, Diptera, Hymenoptera, Orthoptera, Lepidoptera, insect larvae and pupae, and vegetal matter [9]. *Liolaemus victormoralesii* feeds more on vegetal matter during summer and winter, and more on arthropods during spring (n = 56; [9]). This species is viviparous, one female had three embryos on each side at an advanced stage of development.

**Figure 9.** Dorsal and ventral (**B**) variation in *Liolaemus victormoralesii.*(**A**): BYU 50431 (male, SVL=87.8 mm), (**B**): MUSM 31462 (juvenile, SVL = 60.9 mm), (**C**): BYU 50427 (juvenile, SVL = 52.2 mm.

**Figure 10.** Dorsal and ventral variation in females of *Liolaemus victormoralesii.* (**A**): BYU 50433 (SVL = 79.7 mm), (**B**): BYU 50428 (SVL = 69 mm), (**C**): MUSM 31468 (SVL = 75.4 mm).


**Table 3.** Variation in selected characters among type specimens of *Liolaemus victormoralesii* sp. nov. All specimens from Museo de Historia Natural Universidad Mayor de San Marcos.

**Figure 11.** Habitat of *Liolaemus victormoralesii*.

*Etymology*. The specific term *victormoralesii* is a noun in apposition and is given to honor our friend and colleague Víctor Morales for his contributions to herpetology. Víctor Morales passed away in December 2015, but his publications and memory live on with us.
