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TESIS SISTEMÁTICA MOLECULAR Y TAXONOMÍA DE LOS SAURIOS .Pdf Universidad de Concepción Facultad de Ciencias Naturales y Oceanográficas Programa de Magíster en Ciencias con mención en Zoología SISTEMÁTICA MOLECULAR Y TAXONOMÍA DE LOS SAURIOS ALTOANDINOS DEL GÉNERO PROCTOPORUS (SQUAMATA: GYMNOPHTHALMIDAE) Tesis presentada a la Facultad de Ciencias Naturales y Oceanográficas de la Universidad de Concepción para optar al grado académico de de Magíster en ciencias con mención en zoología POR: Luis Mamani Ccasa Profesor guía: Claudio Correa Quezada Profesor co-guía: Alessandro Catenazzi Giannoni Concepción, Chile 2020 Se autoriza la reproducción total o parcial, con fines académicos, por cualquier medio o procedimiento, incluyendo la cita bibliográfica del documento. ii Dedicatoria Esta tesis es dedicada con mucho cariño a Don Alejandro Mamani de la Cruz (Q.E.P.D.) y a mis padres, José Miguel Mamani Cruz y Victoria Ccasa Quispe. iii TABLA DE CONTENIDOS Dedicatoria .......................................................................................................... iii TABLA DE CONTENIDOS .................................................................................. iv ÍNDICE DE TABLAS ............................................................................................ v ÍNDICE DE FIGURAS .......................................................................................... v RESUMEN ......................................................................................................... vii ABSTRACT ......................................................................................................... ix INTRODUCCIÓN ................................................................................................ 1 PROBLEMA A ESTUDIAR ............................................................................ 10 OBJETIVOS ...................................................................................................... 11 OBJETIVOS ESPECÍFICOS ......................................................................... 11 MATERIALES Y MÉTODOS ............................................................................. 12 Material de estudio, extracción, amplificación y secuenciación de ADN........ 12 Reconstrucción filogenética ........................................................................... 16 Caracterización de la folidosis y medidas morfométricas .............................. 18 RESULTADOS .................................................................................................. 26 Análisis filogenético molecular ....................................................................... 26 Resolución de la polifilia de Proctoporus y sus especies ............................... 30 DISCUSIÓN ...................................................................................................... 35 CONCLUSIONES ............................................................................................. 42 REFERENCIAS ................................................................................................ 43 ANEXOS ........................................................................................................... 58 Anexo 1. Especies, vouchers y secuencias de ADN obtenidas de Genbank para este estudio. .......................................................................................... 58 Anexo 2. Revisión Taxonómica de Proctoporus y descripción de géneros nuevos. .......................................................................................................... 62 Género: Opipeuter Uzzell 1969 .................................................................. 92 Género: Perusaurus gen. nov. .................................................................... 96 Género: Alejandrosaurus gen. nov. ............................................................ 99 Género: Incasaurus gen. nov.................................................................... 104 iv Género: Sachasaura gen. nov. ................................................................. 109 Anexo 3. Clave de las especies de Proctoporus, Perusaurus, Incasaurus, Alejandrosaurus y Sachasaura. ................................................................... 115 Anexo 4. A new species of Andean gymnophthalmid lizard (Squamata: Gymnophthalmidae) from the Peruvian Andes, and resolution of some taxonomic problems ..................................................................................... 118 ÍNDICE DE TABLAS Tabla 1. Especímenes de los cuales se obtuvieron secuencias de ADN para este estudio. ............................................................................................... 14 Tabla 2. Primers y condiciones de la PCR usados en este estudio ........... 16 Tabla 3. Características morfométricas y folidosis de los géneros tratados en este estudio ........................................................................................... 34 ÍNDICE DE FIGURAS Figura 1. Distribución actual del género Proctoporus. .................................. 5 Figura 2. Caracterización de la folidosis en Proctoporus caideni ............... 25 Figura 3. Relaciones filogenéticas de Proctoporus y especies afines. ....... 29 Figura 4. Distribución geográfica de los géneros tratados en este estudio. 33 Figura 5. Distribución geográfica del género Proctoporus.. ........................ 64 Figura 6. Vista dorsolateral y ventral de Proctoporus caideni. .................... 67 Figura 7. Vista dorsolateral y ventral de machos de Proctoporus lacertus. 73 Figura 8. Vista dorsolateral y ventral de Proctoporus machupicchu. .......... 77 Figura 9. Vista lateral y ventral de Proctoporus oreades. ........................... 80 Figura 10. Vista dorsolateral y ventral de Proctoporus pachyurus. ............ 83 Figura 11. Vista dorsolateral y ventral de Proctoporus rahmi.. ................... 88 Figura 12. Vista dorsal y ventral de Proctoporus spinalis. .......................... 91 Figura 13. Vista dorsal y ventral de Opipueter xestus. ............................... 93 Figura 14. Distribución de género Opipeuter. ............................................. 95 v Figura 15. Vista dorsolateral y ventral de Perusaurus labioocularis. .......... 97 Figura 16. Distribución del género Perusaurus.. ........................................ 98 Figura 17. Vista dorsolateral y ventral de Alejandrosaurus bolivianus. .... 102 Figura 18. Distribución del género Alejandrosaurus. ................................ 104 Figura 19. Vista dorsolateral y ventral de Incasaurus CA6. ...................... 107 Figura 20. Distribución del género Incasaurus. ........................................ 108 Figura 21. Vista dorsolateral y ventral de Sachasaura. ............................ 110 Figura 22. Distribución del género Sachasaura. ....................................... 112 vi RESUMEN Proctoporus es un género de lagartos de hábitos principalmente semifosoriales, que habitan los bosques montanos, pastizales húmedos y valles interandinos de la vertiente oriental de la Cordillera de los Andes. Los primeros estudios filogenéticos moleculares resolvieron las relaciones filogenéticas y propusieron rearreglos taxonómicos. Sin embargo, estudios filogenéticos moleculares más recientes, basados en muestreos más amplios de especies, mostraron indicios de la polifilia de Proctoporus, lo que concuerda con la divergencia morfológica existente entre sus especies. En este estudio, se obtuvieron 69 terminales adicionales de Proctoporus y especies afines, de las cuales se obtuvieron cuatro marcadores mitocondriales (12S, 16S, Cytb, ND4) y un marcador nuclear (c- mos). Se realizó un análisis filogenético de inferencia Bayesiana (IB) y se realizó una revisión de especímenes de Proctoporus, depositados en colecciones científicas de Perú y Bolivia. Los resultados obtenidos en este estudio confirman la polifilia de Proctoporus; se demuestra que este género está conformado por cinco linajes altamente soportados, algunos de los cuales están más relacionados con otros géneros de gimnoftálmidos, tres especies (Proctoporus lacertus, P. laudahnae, P. guentheri) fueron recuperados en diferentes linajes, Euspondylus caideni fue anidado con Proctoporus chasqui y P. sucullucu fue anidado con P. lacertus. Con base en la filogenia y la revisión morfologica, se proponen los siguientes rearreglos taxonómicos: se resucita al género Opipeuter, vii se erigen cuatro géneros nuevos (Alejandrosaurus sp. nov., Incasaurus gen. nov., Perusaurus gen. nov., Sachasaura gen. nov.), se resuelve la polifilia de P. lacertus, P. laudahnae y P. guentheri, se sinonimiza a Proctoporus chasqui con P. caideni y a P. sucullucu con P. lacertus, se transfiere a Pholidobolus anomalus al género Cercosaura y Cercosaura manicata boliviana es validada como buena especie y nombrada como “Cercosaura manicata boliviana” incertae sedis. PALABRAS CLAVE: Cercosaurinae, Cordillera de los Andes, taxonomía, lagartos semifosoriales, Opipeuter, Alejandrosaurus gen. nov., Incasaurus gen. nov., Perusaurus gen. nov., Sachasaura gen. nov. viii ABSTRACT The species of the genus Proctoporus are mainly semifossorial, inhabiting montane forests, humid grasslands, and inter-Andean valleys on the eastern slopes of the Cordillera de los Andes. The first molecular phylogenetic studies resolved phylogenetic relationships and proposed taxonomic rearrangements of Proctoporus. However, recent molecular phylogenetic studies, based on a broader sampling of species, suggested the polyphyly of Proctoporus, which is consistent with the morphological divergence among its species. In this
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