*3.2. Taxonomy*

*Cercosaura anomala* new. comb. (Müller, 1923) *Pholidobolus anomalus* Müller, 1923

Neotype: MUBI 5277 (Figure 3), an adult male from Puente Ruinas, Santuario Histórico de Machupicchu, District of Machupicchu, Province of Urubamba, Department of Cusco, Peru (13◦09-42" S, 72◦32-07" W, at 2060 m), collected by J.C. Chaparro on 20 April 1998.

**Figure 3.** Neotype of *Cercosaura anomala*, male MUBI 5277 (snout vent length (SVL) = 60.7 mm).

Referred specimens: An adult male (MUBI 641), an adult female (MUBI 640), and a subadult female (MUBI 819) from the same locality as the neotype; two adult females (MUBI 13328, 13626), and a subadult female (MUBI 13529) from Vilcabamba (Figure 4A,B); a subadult male (MUSA 4537) from Maranura; and an adult male (MUBI 16169) from Quellouno (Figure 4C,D). All sites are located in Department of Cusco.

**Figure 4.** Live specimens of*Cercosaura anomala*. (**A**,**B**) Adult females (Urusayhua, Vilcabamba, (**A**) MUBI 13626, SVL = 68.2 mm, (**B**) MUBI 13328, SVL = 72.1 mm); (**C**, **D**) Adult male (Quellouno, La Convención, MUBI 16169, SVL = 56.2 mm). Photo: (A, B) Luis Mamani, (C, D) Juan C. Chavez-Arribasplata.

Etymology: The specific epithet "*anomalus*" is a nominative adjective in ancient Latin meaning irregular (*anomalus*), which implicitly refers to the presence and irregular shape of the prefrontal scales; then *anomala* (feminine nominative) must follow the genus *Cercosaura* (feminine).

Diagnosis: (1) Body robust, maximum snout vent length (SVL) of females, 72.1 mm; males, 61.7 mm; (2) head flat, elongated, 1.5 times longer than wide; (3) ear opening distinct, slightly recessed; (4) nasals separated by frontonasal; (5) frontonasal undivided; (6) prefrontal, frontal, frontoparietals, parietals and interparietals present, prefrontal scales in contact, occasionally separate; (7) parietal longer than wide; (8) three supraoculars, three postoculars, three infraoculars; (9) three superciliars, complete series; (10) nasal suture present; (11) loreal present; (12) 7–8 supralabials, four supralabials anterior to the posteroventral angle of the subocular, occasionally five, 6–8 infralabials; (13) 5–6 genials in contact; (14) collar fold present; (15) 33–36 transverse rows of dorsal scales, hexagonal, slightly keeled, imbricate; (16) 19–22 transverse rows of ventral scales, quadrangular, smooth, juxtaposed; (17) 35–43 scales around midbody; (18) lateral reduced scales at midbody in 3–4 lines; (19) limbs pentadactyl, all digits clawed; (20) 12–14 subdigital lamellae under finger IV, 17–19 under toe IV; (21) 8–10 femoral pores in males, 4–6 in females; (22) 2 preanal scales, 3–4 postanal scales; (23) tail up to 1.9–2.6 times longer than body; (24) caudals subimbricate, keeled dorsally, smooth ventrally; (25) lower palpebral disc transparent and undivided; (26) in life the dorsum is light-brown with some black spots, flanks dark brown with diffuse ocelli; lips with a cream line that extends to the front of arm insertion; ventral surface cream-reddish with some small scattered black spots, gular region of head cream-reddish with small black spots (Figure 4).

*Cercosaura anomala* is very similar to *C. hypnoides*, *C. manicata*, and *C. doanae*; all of which present a clear labial bar on both sides of the head and similar dorsal and lateral colorations. However, *C. anomala* differs from all other species of *Cercosaura* by the presence of smooth dorsal scales on the neck. In addition, *C. anomala* can be differentiated from *C. argulus* by the frontonasal scale being undivided (divided in *C. argulus*), 5–6 genial scales in contact (four genials in *C. argulus*); from *C. anodorsquama*, *C. bassleri*, *C. olivacea*, and *C. ocellata* because the keels of dorsal scales do not form lines arranged on the back (the keels of dorsal scales form continuous lines arranged on the back); from *C. hypnoides* by the presence of eight longitudinal ventral scales (six in *C. hypnoides*), absence of dorsolateral stripes (presence of continuous cream dorsolateral stripes in *C. hypnoides*); from *C. doanae* by the presence of dorsal scales with low keels (strongly keeled in *C. doanae*), maximum SVL of 72.1 mm in females (55.6 mm in *C. doanae*); from *C. eigenmanni* by the presence of 35–43 scales around the body (26–32 scales in *C. eigenmanni*), maximum SVL of 72.1 mm in females (47.0 mm in *C. eigenmanni*); from *C. manicata* by the presence of dorsal scales with low keels (strongly keeled in *C. manicata*), dorsal scales with the anterior and posterior edges almost blunt (anterior and posterior edges pointed in *C. manicata*); from *C. nigroventris* by the presence of 5–6 genials in contact (four genials in contact in *C. nigroventris*), maximum SVL of 61.7 mm in males (44.1 mm in *C. nigroventris*); from *C. oshaughnessyi* by the presence of the frontonasal scale not divided (divided in two in *C. oshaughnessyi*), maximum SVL of 72.1 mm in females (51 mm in *C. oshaughnessyi*); from *C. parkeri* by the presence of 35–43 scales around the body (24–30 in *C. parkeri*), 8–10 femoral pores per leg in males (3–5 per leg in *C. parkeri*); from *C. phelpsorum* by the presence of a white line on the upper lip (without a white line in C. *phelpsorum*), maximum SVL of 72.1 mm in females (55 mm in *C. phelpsorum*); from *C. quadrilineata* by the presence of 35–43 scales around the body (26 in *C. quadrilineata*), presence of 8–9 longitudinal ventral scales (four in *C. quadrilineata*); from *C. schreibersii* by the presence of three postoculars (two in *C. schreibersii*), 8–10 femoral pores per leg in males (3–6 in *C. schreibersii*); from *C. steyeri* by the presence of 8–9 longitudinal ventral scales (four in *C. steyeri*), dorsal scales not mucronate (dorsal scales mucronate in *C. steyeri*).

Description of the neotype (MUBI 5277): Adult male, SVL = 60.7 mm, tail length = 126.4 mm; head scales smooth, without striations, or rugosities; rostral scale wider (2.6 mm) than tall (1.5 mm), meeting supralabials on either side at above the height of supralabials, and becoming higher medially, in contact with frontonasal, nasal, and first supralabials; frontonasal pentagonal, wider than longer, widest in the mid, in contact with rostral, nasal, and prefrontals; prefrontals paired, pentagonal in contact with frontonasal, loreal, first superciliar, first supraocular, and frontal; frontal longer than wide, hexagonal, not in contact with superciliars, but in contact with first supraocular, and frontoparietals; frontoparietals polygonal, in contact with the frontal, all supraoculars, parietals, and interparietal; three supraoculars, all in contact with superciliaries, frontal, frontoparietals, parietal, and postocular; interparietal longer than wide, heptagonal, in contact with frontoparietals anteriorly, with parietal laterally, and with postparietals posteriorly; parietals polygonal, anteriorly in contact with frontoparietals, third supraocular, and postocular, laterally in contact with interparietals and supratemporals, and posteriorly with postparietals; three postparietals, smaller than parietals, the mid postparietal is smaller than lateral postparietals. Nasal divided, longer than high, in contact with first, and second supralabials; loreal present, in contact with second and third supralabials, in contact with nasal, first superciliar, and frenocular; four superciliars, first expanded onto surface of head; frenocular triangular in contact with third supralabials, first subocular, and loreal scales; palpebral disc made up of a single transparent scale; three suboculars; three postoculars; temporals smooth, glossy, and polygonal; four anterior supralabials to the posteroventral angle of the third subocular. Mental wider than long, in contact with first infralabial, and postmental; postmental single, polygonal, in contact with first and second infralabial, and first pairs of genials; three pairs of genials, five in contact, anterior pair in contact with second and third infralabials, middle pair in contact with third, fourth and fifth infralabials, posterior pair of genials in contact with fifth infralabial, and pregulars; two enlarged pregulars on left and right side, and 22 small pregulars irregularly distributed among enlarged pregulars; eight rows of gular scales including the collar, and the middle scales enlarged; collar fold distinct; lateral neck scales round, and smooth; dorsal neck scales smooth. Dorsal hexagonal, longer tan wide, juxtaposed, slightly keeled, in 36 transverse dorsal scale rows; 27 longitudinal dorsal scale rows at midbody; continuous lateral scale series, smaller than dorsals; reduced scales at limb insertion regions present; 22 transverse ventral scale rows; eight longitudinal ventral scale rows at midbody; a pair of anterior preanal plate scales; four posterior preanal plate scales; scales on tail rectangular, juxtaposed, and smooth. Limbs pentadactyl; digits clawed; dorsal brachial scales polygonal, imbricate, and smooth; ventral brachial scales small, rounded, and smooth; antebrachial scales polygonal, subequal in size, smooth, and imbricate; ventral antebrachial small, subimbricate, and rounded; dorsal manus scales polygonal, smooth, and subimbricate; palmar scales small, rounded, and domelike; dorsal scale on fingers smooth, quadrangular, imbricate, three on finger I, six on II, eight on III, nine on IV, and five on V; scales on anterodorsal surface of thigh large, polygonal, smooth, and subimbricate; scales on posterior surface of thigh small, rounded, juxtaposed, and keeled; scales on ventral surface of thigh large, roundish, flat, and smooth; nine femoral pores; preanal pores absent; scales on anterior surface of crus polygonal, keeled, and juxtaposed, decreasing in size distally; scales on posterior surface of crus small, roundish, keeled, and subimbricate; scales on dorsal surface of foot polygonal, smooth, and imbricate; scales on ventral surface of foot small, rounded, juxtaposed, and domelike; scales on dorsal surface of toes quadrangular, smooth, overhanging supradigital lamellae, four on toe I, seven on II, 10 on III, 12 on IV, and eight on V; fore and hind limbs overlapping when adpressed against the body.

Coloration in preservative: The dorsal surface of the head, neck, and body is brown with two clear dorsolateral lines on both sides of head that start from the supraoculars and disappear at the middle of body; the lateral surface of the head, neck, and body is dark brown; on both sides of the head and neck there is a cream labial line that extends from the tip of the head to the anterior part of the insertion of brachium; the ventral surface of the head, neck and body is dark gray with irregular cream spots. The dorsal surface of the limbs is brown, and ventral surface of limbs is similar to the ventral surface of the body (Figure 3).

Coloration in life: According to notes and photographs taken by LM of live specimens, the dorsal surface of the head and neck is brown with small black spots; the lateral sides of the head and neck are blackish brown with a cream labial line that extends from the tip of the head to the anterior part of the insertion of brachium; the ventral surface of the head is cream with small brown spots scattered; pregular and gular regions are similar to the ventral surface of head. The dorsal surface of body is brown with scattered black spots; lateral surface of body is blackish brown with black and cream spots that resemble ocelli; the ventral surface of body is reddish cream with scattered black spots. The dorsal surface of the limbs is brown with small black spots, the ventral surface is reddish cream with small black spots. The dorsal surface of the tail is similar to the dorsum, and the ventral surface of the limbs is similar to the ventral surface of the body (Figure 4).

Variation: Morphometric characters and pholidosis are presented in Table 5; *Cercosaura anomala* apparently has sexual dimorphism in size, females (maximum SVL = 72.1 mm, n = 5) are larger than males (maximum SVL = 61.7 mm, n = 4). The condition of the prefrontal scales is variable, all specimens examined have joined prefrontal scales, and only one subadult female (MUBI 819) and the lost specimen (holotype) have small and separate prefrontal scales.

Distribution and natural history: *Cercosaura anomala* inhabits montane forests on the eastern slopes of Cordillera de los Andes, in Department of Cusco, between 1745–2218 m a.s.l. We have observed this lizard on litter and on rocks from 10:00 to 14:00 h on sunny days at five localities in the Cordillera de Vilcabamba: Urusayhua, Tucantinas, Historical Sanctuary of Machupicchu, Maranura, and Quellouno (Figures 1 and 5). Sympatric gymnophthalmid lizards include *Proctoporus machupicchu*, *P. guentheri, P. unsaacae,* and *Proctoporus* sp. [8].



**Figure 5.** Habitat of *Cercosaura anomala* in the montane forest of the Historical Sanctuary of Machupicchu, Department of Cusco.

## *Cercosaura pacha* sp. nov.

Zoobank LSID: urn:lsid:zoobank.org:act:6B3FCF87-82E4-4E2B-8C3B-99FD93E051CD

Holotype: MUBI 14515 (Figure 6), an adult female from Lanturachi, Fundo los Cocos, District of Huancabamba, Province of Oxapampa, Department of Pasco, Peru (10◦23-02" S, 75◦34-49" W, at 1986 m), collected by J.C. Chaparro and C. Alarcón on 21 September 2014.

**Figure 6.** Holotype of *Cercosaura pacha* sp. nov. female MUBI 14515 (SVL = 49.7 mm).

Paratype: MUBI 14512 (Figure 7), a subadult female from near the type locality (10◦23-29" S, 75◦34-12" W, 1845 m).

Etymology: The specific epithet "*pacha*" is a female noun in Quechua language that means Earth.

Diagnosis: (1) Body robust, SVL 49.7 mm in a single adult female; (2) head flat, elongated, 1.4 times longer than wide; (3) ear opening distinct, slightly recessed; (4) nasals separated by frontonasal; (5) frontonasal undivided; (6) prefrontals, frontal, frontoparietals, parietals and interparietal present; (7) parietals longer than wide; (8) three supraocular; (9) 3–5 superciliar series complete; (10) nasal suture absent; (11) loreal present, in contact with the second supralabial; (12) 7–8 supralabial, four supralabials anterior to the posteroventral angle of the subocular, four infralabials; (13) 4–5 genial, all in contact; (14) collar fold present; (15) 32–35 transverse rows of dorsal, hexagonal, keeled, imbricate; (16) 19–20 transverse ventral rows, quadrangular, smooth, juxtaposed; (17) 37–38 scales around midbody; (18) lateral reduced scales at midbody in three lines; (19) limbs pentadactyl, all digits clawed; (20) 12–13 subdigital lamellae under finger IV, 18 under toe IV; (21) 6–7 femoral pores in females; (22) two preanal scales, three postanal scales; (23) tail up to 2.0 times longer than body; (24) caudals subimbricate, keeled dorsally, smooth ventrally; (25) lower palpebral disc transparent and divided in two; (26) in life the dorsum is brown with two cream dorsolateral stripe that stars over the eyes and join in the middle of the body forming a vertebral dorsal stripe that extends to the tail; lips with a cream line that extend from the third supralabial to the front of back leg; a cream lateral line between arm and leg, below the lateral line; all cream lines are bordered by continuous black spots; the venter is cream-reddish with some small scattered black spots, the gular region of head is cream-reddish with small black spots; tail is orange, with small dark spots ventrally and dorsally, and a cream-orange line laterally that begins at the back of legs and continues to tip of the tail (Figures 6 and 7).

**Figure 7.** Live specimen of *Cercosaura pacha* sp. nov., subadult female MUBI 14512 (paratype, SVL = 32.4 mm), from Lanturachi, near Oxapampa, Department of Pasco, Peru. Photo: Consuelo Alarcón.

*Cercosaura pacha* sp. nov. is similar to *C. anomala*, *C. doanae*, *C. hypnoides*, and *C. manicata*. However, *C. pacha* sp. nov. differs from *C. anomala* by having dorsal surface of neck keeled (smooth in *C. anomala*), six genials (4–5); from *C. doanae* by having dorsal scale of neck polygonal, keeled, and the distal edges of scales are blunt (strongly keeled, and the distal edges are pointed in *C. doanae*), dorsolateral stripes forming a vertebral dorsal stripe (not forming a vertebral stripe); from *C. hypnoides* by having loreal scales in contact with supralabials (not in contact with supralabias), eight longitudinal ventral scales (six), dorsal scales of neck polygonals (rounded); from *C. manicata* by having three postoculars (four in *C. manicata*), three suboculars (4–5), eight longitudinal ventral scales (six). Furthermore, *C. pacha* sp. nov. differs from *C. anordosquama*, *C. argulus*, *C. bassleri*, *C. eigenmanni*, *C. nigroventris*, *C. ocellata*, *C. olivacea*, *C. oshaughnessyi*, *C. parkeri*, *C. phelpsorum*, *C. quadrilineata*, *C. schreibersii*, and *C. steyeri* in having a clear labial bar that extends from the third supralabial to the point of insertion of the posterior limbs, and cream dorsolateral stripes that extends over the eyes and join in the middle of the body forming a single vertebral dorsal stripe that reaches the tail. Additionally, *C. pacha* sp. nov. can be distinguished from *C. argulus* and *C. oshaughnessyi* by having an undivided frontonasal (divided in *C. argulus* and *C. oshaughnessyi*); from *C. anordosquama*, *C. bassleri*, *C. ocellata* and *C. olivacea* by having the keels of the dorsal scales not organized in longitudinal rows, and eight longitudinal ventral rows (organized in longitudinal rows, and six in *C. anordosquama*, *C. bassleri*, *C. ocellata* and *C. olivacea*); from *C. eigenmanni* by having 37–38 scales around midbody (26–32 in *C. eigenmanni*); from *C*. *nigroventris* by having 37–38 scales around the midbody (40–44 in *C. nigroventris*), dorsal scales strongly keeled (weakly keeled in *C*. *nigroventris*); from *C. parkeri* by having 37–38 scales around the mid-body (24–30 in *C. parkeri*); from *C. phelpsorum* by having dorsal scales strongly keeled (weakly keeled in *C. phelpsorum*); from *C. quadrilineata* by having eight longitudinal ventral scales (four in *C. quadrilineata*); from *C. schreibersii* by having eight longitudinal ventral scales (six in *C. schreibersii*); from *C. steyeri* by having eight longitudinal rows of ventral scales (four in *C. steyeri*) and 37–38 scales around midbody (17).

Description of the holotype (MUBI 14515): Adult female, SVL = 49.7 mm, tail length = 98.1 mm; head scales with some rugosities; rostral scale wider (2.3 mm) than tall (1.2 mm), meeting supralabials on either side at above the height of supralabials, and becoming higher medially, in contact with frontonasal, nasal, and first supralabials; frontonasal polygonal with blunt edges, wider than longer, widest at the back, in contact with rostral, nasals, loreals, and prefrontals; prefrontals paired, polygonal, in contact with frontonasal, loreal, first superciliar, first supraocular, and frontal; frontal longer than wide, polygonal, in contact with first and second supraocular, and frontoparietals; frontoparietals polygonal, in contact with the frontal, second and third supraoculars, parietals, and interparietal; three supraoculars, all in contact with superciliaries, frontal, frontoparietals, parietal, and postocular; interparietal longer than wide, heptagonal, in contact with frontoparietals anteriorly, with parietal laterally, and with postparietals posteriorly; parietals polygonal, anteriorly in contact with frontoparietals, third supraocular, and postocular, laterally in contact with interparietals and temporals, and posteriorly postparietals; three postparietals, smaller than parietals, the mid postparietal is smaller than laterals postparietals. Nasal undivided, longer than high, in contact with first and second supralabials; loreal present, in contact with second supralabials, nasal, first superciliar, and frenocular; four superciliars, the first expanded onto surface of head; frenocular trapezoidal, in contact with second and third supralabials, infraocular, subocular, and loreal scales; palpebral disc divided in two semitransparent scales; three suboculars; three postoculars; temporals with keeled and smooth scales (the big scales smooth and the small scales keeled), and polygonal; four supralabials anterior to the posteroventral angle of the subocular. Mental wider (2.3 mm) than long (1.5 mm), in contact with first infralabial and postmental posteriorly; postmental single, polygonal, in contact with first and second infralabial, and first pairs of genials; five genials, all in contact, on the left side two and on the right three, in contact with second, third, and fourth infralabials; 35 pregulars irregularly distributed, and small in the mid; seven rows of gular scales including the collar, the middle scales enlarged; collar fold distinct, formed by large scales; lateral neck scales round, upper scales keeled, and lower scales smooth; dorsal neck scales polygonal and keeled. Dorsal hexagonal, longer tan wide, juxtaposed, strongly keeled, in 35 transverse rows; 30 longitudinal dorsal scale rows at midbody; lateral scale series slightly smaller than dorsal; reduced scales at limb insertion regions; 20 transverse ventral scale rows; eight longitudinal ventral scale rows at midbody; four anterior preanal plate scales (the lateral scales are smaller), three posterior preanal plate scales; scales on tail rectangular, juxtaposed, and smooth. Limbs pentadactyl; digits clawed; dorsal brachial scales polygonal, imbricate, and slightly keeled; ventral brachial scales small, rounded, and smooth; dorsal antebrachial scales polygonal, subequal in size, smooth, and imbricate; ventral antebrachial small, subimbricate, and rounded; dorsal manus scales polygonal, smooth, and subimbricate; palmar scales small, rounded, and domelike; dorsal scale on fingers smooth, quadrangular, imbricate, four on finger I, six on II, eight on III, nine on IV, and five on V; scales on anterodorsal surface of thigh large, polygonal, smooth, and sub-imbricate; scales on dorsal surface of thigh large, keeled; scales on posterior surface of thigh small, rounded, juxtaposed, and keeled; scales on ventral surface of thigh large, roundish, flat, and smooth; seven femoral pores; preanal pores absent; scales on anterior surface of crus small, polygonal, keeled, juxtaposed, and decreasing in size distally; scales on posterior surface of crus small, roundish, keeled, and sub-imbricate; scales on ventral surface of crus large, roundish, flat, and smooth; scales on dorsal of foot roundish, smooth, and imbricate; scales on ventral of foot small, rounded, juxtaposed, and domelike; scales on dorsal surface of toes quadrangular, smooth, overhanging supradigital lamellae, three on toe I, six on II, 10 on III, 12 on IV, and eight on V; fore and hind limbs overlapping when adpressed against the body.

Coloration: In preservative, the dorsal surface of the head, neck, and back is dark-brown, the dorsolateral lines are cream grayish and join at midbody to form a vertebral stripe that extends to the tail; the dorsal surface of the tail is dark brown with a dorsal stripe in the anterior part of the tail, and pale orange with some gray spots in the distal part; the lateral sides of the head and neck are blackish brown with a cream labial line that extends from the third supralabial to the anterior part of the insertion of posterior limbs; the ventral surface of the head is gray with small, irregular, brown spots; gular and ventral surfaces of the body are dark gray with cream spots around some scales; the ventral surface of the limbs and tail, are cream with some irregular, dark gray spots (Figure 6). In life, the dorsal surface of the body is brown with scattered black spots; the lateral surface of the

body is blackish brown with black and cream spots that resemble ocelli; the ventral surface of the body is reddish cream with scattered black spots. The dorsal surface of the limbs is brown with small black spots, the ventral surface is reddish cream with small black spots. The dorsal and ventral surfaces of the tail are orange, and the ventral surfaces of the limbs are similar to the ventral surface of the body (Figures 6 and 7).

Variation: Table 5 summarizes morphometric characters and pholidosis.

Distribution and natural history: *Cercosaura pacha* sp. nov. inhabits montane forests on the eastern slopes of Cordillera de los Andes, Department of Pasco, central Peru, between 1845–1986 m a.s.l (Figure 1). We captured two specimens using pitfall traps set up for 10 days at the type locality (Figure 8).

**Figure 8.** Type locality of *Cercosaura pacha* sp. nov, montane forest of Lanturachi, near Oxapampa, Department of Pasco, Peru. Photo: Consuelo Alarcón.

Taxonomic status of *Cercosaura manicata boliviana*

*Cercosaura manicata boliviana* is considered a subspecies of *Cercosaura manicata*, which is distributed from southeastern Peru to central Bolivia [30]. Werner [54] described *C. manicata boliviana* as *Prionodactylus bolivianus* based on a specimen (Muséum national d'Histoire naturelle, MNHN 00.4) collected in the montane forests of Bolivia (Chacó). Subsequently, *P. bolivianus* and *P. ockendeni* were considered synonyms of *Prionodactylus manicatus*, but both with subspecies status [30]. Finally, according to a phylogenetic study based on morphological data, *P. manicatus bolivianus* was transferred to the genus *Cercosaura* [18]. Echevarría et al. [16] and Uzzell [30] observed clear differences between both subspecies; however, only Echevarría et al. [16] considered this taxon as a putative separate species, and highlighted the need for genetic evidence.

The genus *Prionodactylus* was erected by O'Shaughnessy [55], and the type species was *Prionodactylus manicatus*. However, *P. manicatus* was transferred to the genus *Cercosaura*, and *Prionodactylus* was invalidated and considered as a synonym of *Cercosaura* [18].

According to the molecular evidence obtained in this study using two specimens from southern Peru (Figure 9), *Cercosaura manicata boliviana* is the sister lineage of the genus *Potamites*; therefore,

it should be excluded from *Cercosaura*. However, the taxonomic assignation of *Cercosaura manicata boliviana* remains uncertain, because the genus *Prionodactylus* (original genus) is no longer valid. We could assign this species to *Potamites*, but external morphological characters and ecological traits do not support this taxonomic change*. Potamites* is a genus of lizards strongly associated with aquatic ecosystems [20], whereas the individuals of *"Cercosaura manicata boliviana*" have semi-arboreal habits. Thus, we propose to maintain the name "*Cercosaura manicata boliviana*" *incertae sedis* until a dedicated study can ascertain its phylogenetic relationships.

**Figure 9.** Live specimen of "*Cercosaura manicata boliviana*"; (**A**) adult male CORBIDI 16500 (SVL = 42.5 mm), from near San Pedro, Kosñipata Valley, Department of Cusco, Peru; (**B**) CORBIDI 18716 (SVL = 44.1 mm), from Santo Domingo, District of Limbani, Province of Sandia, Department of Puno, Peru. Photo: Alessandro Catenazzi.

In conclusion, the molecular, ecological, and morphological evidence support the hypothesis that *"Cercosaura manicata boliviana* is a separate species and a new lineage, which is sister to lizards of the genus *Potamites*. Future studies should ascertain the relationship of this *incertae sedis* with *Potamites*, and determine whether *P. bolivianus* and *P. ockendeni* are conspecifics or separate species.
