**4. 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].

Despite the similarity of coloration patterns of *Cercosaura anomala* with species of *Pholidobolus*, and "*C. manicata boliviana"* with *C. manicata*, both species were not nested in their designated genera. This result shows that external morphological characters in gymnophthalmid lizards can converge in coloration, and pholidosis [10,12,20,52]. Examples of convergence are the body shape of six divergent lineages of semi-aquatic lizards of the genera *Centrosaura*, *Echinosaura*, *Gelanesaurus*, *Neusticurus*, *Potamites*, and *Rheosaurus*, which share similar body shape ("cocodrile like morphology"), presence of irregulars scales on the back, and a laterally flattened tail that aids in water locomotion [31,58]; body elongations in *Anotosaura*, *Bachia*, *Calyptommatus*, *Heterodactylus*, *Nothobachia*, and *Scriptosaura*, [32,59]; legs reduction in *Bachia*, *Colobosaura*, and *Scriptosaura* [9,32,60]; and external ear loss in *Antonosaura*, *Bachia, Heterodactylus*, *Nothobachia*, *Rachisaurus brachylepis*, and *Scriptosaura catimbau* [32,60]. In light of high frequency of evolutionary convergence, it is expected that lizards of the genera *Cercosaura*, *Pholidobolus*, *Macropholidus*, and *"Cerosaura manicata boliviana"* share similarities in their coloration patterns. Future evolutionary studies will further elucidate evolutionary convergence in these lizards.

**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.

**Author Contributions:** Conceptualization: L.M., J.C.C., C.C. and A.C.; methodology, L.M., C.C. and A.C.; software, L.M., C.C., C.A., and A.C.; validation, L.M., J.C.C., C.C., C.A., C.Y.S. and A.C.; formal analysis, L.M. and A.C.; investigation, L.M., J.C.C., C.C., C.A., C.Y.S., and A.C.; resources, A.C.; data curation, L.M., J.C.C., C.C., C.A., C.Y.S., and A.C.; writing—original draft preparation, L.M., C.C. and A.C.; writing—review and editing, L.M., J.C.C., C.C., C.A., C.Y.S. and A.C.; visualization, L.M., C.C. and A.C.; supervision, C.C. and A.C.; project administration, L.M. and A.C.; funding acquisition, L.M. and A.C. All authors have read and agreed to the published version of the manuscript.

**Funding:** The Asociación para la Conservación de la Cuenca Amazónica (ACCA) provided logistical support at Urusayhua and Tucantinas in Departament of Cusco, Peru.

**Acknowledgments:** We thank Evaristo <sup>L</sup>ópez (MUSA), Pablo Venegas (CORBIDI), and MUBI staff for allowing access to their respective herpetological collections during this study; Juan Carlos Chávez-Arribasplata (CORBIDI) for allowing us to use his photographs (Figure 3C,D); Tiffany M. Doan and an anonymous reviewer for their comments and suggestions that improved this manuscript.

**Conflicts of Interest:** The authors declare no conflict of interest.

## **Appendix A. Specimens Examined**

*Cercosaura anomala*—Peru: Department of Cusco: Province of Urubamba, District of Machupicchu, sector Puente Ruinas (MUBI 640, 641, 817, 5277); Province of La Convención, District of Santa Ana, sector Urusayhua (MUBI 13626), sector Tucantinas (MUBI 13328, 13529); District of Maranura (MUSA 4537); District of Quellouno (MUBI 16169).

*Cercosaura manicata*—Peru: Department of Cusco, Province of La Convención, District of Kimbiri, sector Pomoreni (MUBI 6789) and Pichari (MUBI 15734, 15735, 15736).

*Cercosaura pacha* sp. nov.—Peru: Department of Pasco, Province of Oxapampa, District of Huancabamba, sector Lanturachi (MUBI 14512, 14515).

*Cercosaura* sp.—Peru: Department of Cusco, Province of Quispicanchi, District of Camanti, sector Sirigua (MUBI 5881).

*"Cercosaura manicata boliviana"*—Peru: Department of Puno: Province of Paucartambo, District of Kosñipata, Parque Nacional del Manu, Trocha Unión (MUBI 5045), sector San Pedro (CORBIDI 16500); Departament of Puno, Province of Carabaya, sector Gallucunka (MUBI 4657), sector Ollachea (MUBI 11575), Province of Sandia, District of Limbani, Santo Domingo (CORBIDI 18716).
