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Los Panochthini (Xenarthra, Glyptodontidae): Sistemática Y Evolución

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Los Panochthini (Xenarthra, Glyptodontidae): Sistemática Y Evolución Los Panochthini (Xenarthra, Glyptodontidae): Sistemática y Evolución Tesis para optar al título de Doctor en Ciencias Naturales Licenciado Martín Zamorano Director: Gustavo Juan Scillato-Yané Codirector: Alfredo Eduardo Zurita Facultad de Ciencias Naturales y Museo Universidad Nacional de La Plata La Plata, 2012 A MI PAPÁ Y MI MAMÁ POR MÁS DE 33 AÑOS DE GENUINO CARIÑO ÍNDICE Agradecimi entos...........................................................................................................................I Resumen.......................................................................................................................................V Abstract..................................................................................................................................... VII 1. INTRODUCCIÓN 1. 1. Glyptodontidae: un peculiar grupo de Xenarthra............................................................ 1 1. 2. Características anatómicas de los Glyptodontidae......................................................... 4 1. 3. Ubicación de la familia Glyptodontidae (según distintos esquemas de consenso entre los propuestos por distintos autores).....................................................7 1. 4. Los Panochthini dentro de los Glyptodontidae: distintas propuestas...........................8 1. 5. Estado del conocimiento de los géneros y especies de la Tribu Panochthini en el momento de iniciar esta Tesis (2007)................................................................. 13 1. 6. Objetivos............................................................................................................................ 18 1. 7. Hipótesis.............................................................................................................................19 2. MATERIALES Y MÉTODOS 2. 1. Terminología anatómica utilizada y aspectos nomenclaturales y bioestratigráficos............................................................................................................ 20 2. 2. Repositorios y abreviaturas..............................................................................................21 2. 3. Ejemplares estudiados: número de repositorio, restos, procedencia geográfica y estratigráfica................................................................................................................... 25 2. 4. Metodología aplicada....................................................................................................... 35 2. 5. Medidas calculadas........................................................................................................... 38 Figuras I-XIV................................................................................................... 39 3. SISTEMÁTICA 3. 1. Aspectos históricos y nomenclaturales de los géneros de Panochthini.................... 48 3. 2. Revisión sistemática y contexto estratigráfico de los géneros y las especies de Panochthini considerados válidos en esta Tesis.................................................... 52 3. 2. 1. Nopachtus Ameghino, 1888.......................................................................... 52 3. 2. 2. Nopachtus coagmentatus Ameghino, 1888.................................................53 Figuras 1-2...........................................................................................58 3. 2. 3. Nopachtus cabrerai Zamorano et al., 2011.................................................60 Figura 3................................................................................................62 3. 2. 4. Propanochthus Castellanos, 1925.................................................................63 3. 2. 5. Propanochthus bullifer (Burmeister, 1874).................................................63 Figuras 4-5...........................................................................................67 3. 2. 6. Panochthus Burmeister, 1866....................................................................... 69 3. 2. 7. Panochthus tuberculatus (Owen, 1845)...................................................... 70 Figuras 6-9...........................................................................................83 3. 2. 8. Panochthus frenzelianus Ameghino, 1889..................................................87 Figuras 10-13.......................................................................................94 3. 2. 9. Panochtus intermedius Lydekker, 1895.......................................................97 Figuras 14-16.................................................................................... 106 3. 2. 10. Panochthus subintermedius Castellanos, 1937.......................................109 Figuras 17-20.................................................................................... 115 3. 2. 11. Panochthus greslebini Castellanos, 1942................................................119 Figuras 21-22.....................................................................................123 3. 2. 12. Panochthus jaguaribensis Moreira, 1965................................................125 Figuras 23-24.................................................................................... 128 3. 2. 13. Panochtus cf. subintermedius.................................................................. 130 Figuras 25-27.................................................................................... 134 3. 3. Especies nominadas de panoctinos consideradas no válidas o mal asignadas genéricamente................................................................................................................137 3. 3. 1. Panochthus giganteus Serres, 1866 (in Pouchet, 1866)..........................137 3. 3. 2. Panochthus bullifer Burmeister, 1874.......................................................138 3. 3. 3. Panochthus lundii Burmeister, 1874......................................................... 138 3. 3. 4. Panochthus morenoi Ameghino, 1881......................................................139 3. 3. 5. Panochthus eocenus Scalabrini, 1887......................................................... 141 3. 3. 6. Panochthus beyrichi Roth, 1888..................................................................142 3. 3. 7. Panochthus brocherii Moreno, 1888.......................................................... 142 3. 3. 8. Panochthus trouessarti Moreno, 1888........................................................143 3. 3. 9. Panochthus vogti Roth, 1888........................................................................143 3. 3. 10. Panochthus (?) nodotianus Ameghino, 18089........................................ 144 3. 3. 11. Panochthus voghti Ameghino, 1889......................................................... 146 3. 3. 12. Panochthus interundatus (Ameghino, 1885) Lydekker 1895.................147 3. 3. 13. Nopachtus trouessarti (Moreno, 1888) Castellanos, 1925..................... 147 3. 3. 14. Panochthus oliveiraroxoi Castellanos, 1942............................................148 3. 3. 15. Panochthus rusconii Castellanos, 1942.................................................... 148 3. 3. 16. Panochthus rochai Paula Couto, 1954......................................................149 3. 3. 17. Parapanochthus jaguaribensis (Moreira, 1965) Moreira, 1971........... 149 4. FILOGENIA 4. 1. Análisis filogenéticos de los Glyptodontidae.............................................................. 151 4. 2. Descripción, codificación e ilustración de los caracteres empleados en el análisis filogenético..............................................................................................153 4. 3. Descripción de la elaboración del análisis filogenético............................................. 183 4. 4. Resultados del análisis filogenético............................................................................. 186 4. 5. Discusiones sobre los resultados del análisis filogenético........................................ 186 5. DISTRIBUCIÓN CRONOLÓGICA Y GEOGRÁFICA DE LOS PANOCTINOS 5. 1. Distribución cronológica de los panoctinos.................................................................198 5. 2. Distribución geográfica de los panoctinos.................................................................. 208 6. DISCUSIÓN..........................................................................................................228 7. CONCLUSIONES............................................................................................... 231 8. APENDICE. Matriz e indices calculados en los análisis filogenéticos................................................................................................................235 9. BIBLIOGRAFÍA................................................................................................. 237 Se hace notar que el texto de esta Tesis Doctoral se atiene a las normas de la “Ortografía de la lengua española” (2011), la cual ha innovado sobre todo en lo atinente a la acentuación de numerosos vocablos. “Hay hombres que luchan un día y son buenos. Hay otros que luchan un año y son mejores. Hay quienes luchan muchos años, y son muy buenos. Pero hay los que luchan toda la vida, esos son los imprescindibles”. Bertolt Brecht (1898-1956). AGRADECIMIENTOS Procurando nombrar a todas las personas a las cuales deseo agradecer el inconmensurable apoyo y la vital colaboración que me brindaron durante la elaboración de esta Tesis, arrancaré a escribir estos renglones. El orden de aparición no será riguroso, me resultaría imposible encontrarle
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