A New Species of Andean Gymnophthalmid Lizard (Squamata: Gymnophthalmidae) from the Peruvian Andes, and Resolution of Some Taxonomic Problems

: The family Gymnophthalmidae is one of the most speciose lineages of lizards in the Neotropical region. Despite recent phylogenetic studies, the species diversity of this family is unknown and thus, its phylogenetic relationships remain unclear and its taxonomy unstable. We analyzed four mitochondrial (12S, 16S, Cytb, ND4) and one nuclear (c-mos) DNA sequences of Pholidobolus anomalus , Cercosaura manicata boliviana and Cercosaura sp., using the maximum likelihood method to give insights into the phylogenetic relationships of these taxa within Cercosaurinae. Our results suggest that Pholidolus anomalus is nested within the clade of Cercosaura spp., that material we collected near Oxapampa belongs to a new species of Cercosaura , and that lizards identified as Cercosaura manicata boliviana belong to a separate lineage, possibly a new genus. We assign Pholidobolus anomalus to Cercosaura , redescribe the species, and designate a neotype to replace the lost holotype. In addition, we describe the new species of Cercosaura, and comment about the taxonomic status of “ Cercosaura manicata boliviana” incertae sedis . 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


Introduction
The eastern slopes of the Peruvian Andes are one of the regions with the greatest diversity of flora and fauna [1]. During the last few years, many species of plants and animals from the Peruvian Andes have been discovered (e.g., [2,3]). In particular, researchers have discovered many species of lizards of the family Gymnophthalmidae in poorly explored regions [4][5][6][7][8][9]. Recent phylogenetic inferences using molecular sequences have uncovered the phylogenetic relationships of many gymnophthalmid taxa [10][11][12][13]. However, there are still species whose phylogenetic position remains unclear, such as some populations currently assigned to the genus Cercosaura Wagler, 1830 and Pholidobolus anomalus Müller, 1923. Furthermore, the existence of poorly explored areas in the Peruvian Andes suggests that knowledge of the species richness of this group is still incomplete. Orange triangles = "Cercosaura manicata boliviana"; light-blue circle = C. manicata; purple circle = C. hypnoides; blue circle = C. doanae; green square = C. anomala; red star = C. pacha sp. nov.; yellow circle = Cercosaura sp. Data taken from the literature [16,17,30] and museum records. Orange triangles = "Cercosaura manicata boliviana"; light-blue circle = C. manicata; purple circle = C. hypnoides; blue circle = C. doanae; green square = C. anomala; red star = C. pacha sp. nov.; yellow circle = Cercosaura sp. Data taken from the literature [16,17,30] and museum records.

Designation of Neotype and Species Descriptions
In the original description of Pholidobolus anomalus, Müller [21] was not very precise about the type locality, and mentioned Cusco as the collecting locality. However, the most probable place where the holotype was collected is the Historical Sanctuary of Machupicchu (HSM), in the Cordillera de Vilcabamba. After its discovery by Hiram Bingham in 1911, the HSM became very popular with naturalists. The resulting scientific collections were deposited in different natural history museums [42,43].
We designed a neotype of Pholidobolus anomalus because the holotype was lost. This designation is covered by Article 75.3 of the International Code of Zoological Nomenclature (ICZN) [44]. We used the neotype to redescribe the species, and support the generic allocation. We followed Uzzell [45], Kizirian [46], and Doan and Cusi [24] for character definitions and measurements, and Chávez et al. [7] for description format. One of us (LM) observed morphological characters of species and took all measurements using a caliper with a precision of 0.1 mm. We referred to the literature for patterns of scalation and coloration of the following taxa: Cercosaura anordosquama, C. ocellata, C. bassleri and C. olivacea [13]; Cercosaura argulus, C. eigenmanni, and C. oshaughnessyi [47]; C. hypnoides [17]; C. quadrilineata, C. schreibersii, and C. phelpsorum [18]; C. doanae [16]; C. manicata [30]; C. nigroventris [48]; C. parkeri [49]; and C. steyeri [50]. We also examined specimens of Cercosaura deposited in the CORBIDI, MUBI, and MUSA collections (Appendix A). The electronic version of this article in portable document format will represent a published work according to the International Commission on Zoological Nomenclature, and hence the new names contained in the electronic version are effectively published under that Code from the electronic edition alone. This published work and the nomenclatural acts it contains have been registered in ZooBank, the online registration system for the ICZN. The Life Sciences Identifier (LSID) for this publication is: urn:lsid:zoobank.org:pub:FF0EC17F-965E-410D-AF0A-356B19BD4431.
Based on our molecular analyses ( Figure 2, Tables 3 and 4), we conclude (1) that P. anomalus needs to be redescribed and allocated to the genus Cercosaura, including designating a neotype; (2) Cercosaura sp. (MUBI 14515) is a new species; and (3) Cercosaura manicata boliviana is a valid species, the sister lineage of the semiaquatic lizards of the genus Potamites [20], but considered here as incertae sedis.
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 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)  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]. Table 5. Morphometric measurements and pholidosis of Cercosaura anomala and C. pacha sp. nov.

Cercosaura anomala (all adults)
Cercosaura 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].     (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). Paratype: MUBI 14512 (Figure 7), a subadult female from near the type locality (10°23′29″ S, 75°34′12″ W, 1845 m).  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).
Diversity 2020, 12, x FOR PEER REVIEW 18 of 24 a.s.l (Figure 1). We captured two specimens using pitfall traps set up for 10 days at the type locality ( Figure 8).
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.

Discussion
Our molecular and morphological evidence solves the taxonomy of Cercosaura anomala, reveals "Cercosaura manicata boliviana" as incertae sedis, and supports the description of a new species of Cercosaura from the Andes of Peru. Despite a complex taxonomic history, genetic data have supported recent changes in the systematics and taxonomy of cercosaurine lizards, increasing our understanding of their evolutionary history (e.g., [6,10,12,19,27,53]). However, genetic studies are still incomplete, and many genera and species are pending review and broader sampling of genetic sequences [6,10,12]. 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.

Discussion
Our molecular and morphological evidence solves the taxonomy of Cercosaura anomala, reveals "Cercosaura manicata boliviana" as incertae sedis, and supports the description of a new species of Cercosaura from the Andes of Peru. Despite a complex taxonomic history, genetic data have supported recent changes in the systematics and taxonomy of cercosaurine lizards, increasing our understanding of their evolutionary history (e.g., [6,10,12,19,27,53]). However, genetic studies are still incomplete, and many genera and species are pending review and broader sampling of genetic sequences [6,10,12].
The ML topology obtained in this study using concatenated sequences of mitochondrial and nuclear genes recovered the monophyly of Proctoporus and included the genus Wilsonosaura within Proctoporus. This topology contrasts with previous studies that did not support the monophyly of Proctoporus, suggesting additional studies are needed to solve the taxonomy and phylogenetic position of Proctoporus [6,11,12,51,52]. Moreover, our study considered 129 terminals and addressed the taxonomy and phylogenetic relationships of the three species of Cercosaura (Cercosaura anomala, "Cercosaura manicata boliviana", and Cercosaura pacha sp. nov.).
We designated a neotype for Cercosaura anomala, a designation carried out in accordance with article 75.3 of the ICZN, based on a specimen collected in Puente Ruinas, inside the Historical Sanctuary of Machupicchu, Department of Cusco, Peru. The designation of HSM as the type locality of C. anomala, and associated genetic data we provided in this work, are important because they will facilitate future taxonomic, phylogenetic, ecological, and evolutionary studies. Moreover, with the generic allocation of C. anomala and the description of a new species, we increase the diversity of the genus Cercosaura, which now contains 18 species.
In the original description of Cercosaura anomala, Müller [21] observed the small size of the prefrontal scales, and the separation between them, stating that these could be rudimentary. Among the material examined in this study, all specimens have large and attached prefrontals, except a subadult female (MUBI 819) with separate prefrontal scales. Variation in the form of prefrontals, and other characters, occurs in different species of gymnophthalmid lizards such as Pholidobolus vertebralis [24], Proctoporus spinalis [23], P. machupicchu [56], and P. laudhanae [57]. The high cryptic diversity, and the variation observed in the characters used in taxonomy of these lizards warn us that generic assignments and the description of new species should be undertaken with caution, and if possible, supported by genetic evidence [10,12,20,29].

Supplementary Materials:
The following are available online at http://www.mdpi.com/1424-2818/12/9/361/s1: Table S1: Vouchers and accession number codes of taxa included in this study available from GenBank. Figures S1 and S2: Mitochondrial and nuclear maximum likelihood trees respectively showing the phylogenetic relationships of Cercosaura, and other gymnophthalmid lizards.