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La Familia Carditidae (Mollusca- Bivalvia) En El Cenozoico Del Atlántico Sudoccidental: Sistemática Y Filogenia

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La Familia Carditidae (Mollusca- Bivalvia) En El Cenozoico Del Atlántico Sudoccidental: Sistemática Y Filogenia Tesis Doctoral La familia Carditidae (Mollusca- Bivalvia) en el Cenozoico del Atlántico Sudoccidental: sistemática y filogenia Pérez, Damián Eduardo 2016-03-11 Este documento forma parte de la colección de tesis doctorales y de maestría de la Biblioteca Central Dr. Luis Federico Leloir, disponible en digital.bl.fcen.uba.ar. Su utilización debe ser acompañada por la cita bibliográfica con reconocimiento de la fuente. This document is part of the doctoral theses collection of the Central Library Dr. Luis Federico Leloir, available in digital.bl.fcen.uba.ar. It should be used accompanied by the corresponding citation acknowledging the source. Cita tipo APA: Pérez, Damián Eduardo. (2016-03-11). La familia Carditidae (Mollusca-Bivalvia) en el Cenozoico del Atlántico Sudoccidental: sistemática y filogenia. Facultad de Ciencias Exactas y Naturales. Universidad de Buenos Aires. Cita tipo Chicago: Pérez, Damián Eduardo. "La familia Carditidae (Mollusca-Bivalvia) en el Cenozoico del Atlántico Sudoccidental: sistemática y filogenia". Facultad de Ciencias Exactas y Naturales. Universidad de Buenos Aires. 2016-03-11. Dirección: Biblioteca Central Dr. Luis F. Leloir, Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires. Contacto: [email protected] Intendente Güiraldes 2160 - C1428EGA - Tel. (++54 +11) 4789-9293 UNIVERSIDAD DE BUENOS AIRES Facultad de Ciencias Exactas y Naturales Departamento de Ciencias Geológicas La familia Carditidae (Mollusca-Bivalvia) en el Cenozoico del Atlántico Sudoccidental: sistemática y filogenia Tesis presentada para optar al título de Doctor de la Universidad de Buenos Aires en el área de Ciencias Geológicas Damián Eduardo Pérez Director de Tesis: Dra. Claudia J. del Río Consejero de Estudios: Dra. G. Andrea Concheyro División Paleontología de Invertebrados Museo Argentino de Ciencias Naturales “Bernardino Rivadavia” (MACN), CONICET Buenos Aires, 2015 Fecha de defensa: 11/3/2016 Tesis Doctoral Universidad de Buenos Aires Damián Eduardo Pérez, 2015 ÍNDICE Resumen Abstract Agradecimientos I. Introducción 7 II. Objetivos e Hipótesis de trabajo 10 III. Antecedentes 13 IV. Materiales y Métodos 17 A. Materiales 18 B. Metodología 19 a. Morfometría geométrica 19 b. Filogenia 21 c. Sistemática 22 V. Marco geológico 24 VI. Morfometría geométrica 61 A. Variación ontogenética en Claibornicardia paleopatagonica 62 B. Variación sexual en Nov. Gen. C patagonica 68 C. Variación morfológica en Neovenericor austroplata 71 D. Discusión 79 E. Otros casos de variación intraespecífica 83 VII. Relaciones filogenéticas 84 A. Antecedentes 85 B. Análisis de caracteres 90 a. Costillas radiales 91 b. Tipos de costillas 95 c. Charnela 96 C. Grupo interno 97 D. Grupo externo 98 E. Búsqueda 99 F. Resultados 100 G. Discusión y Conclusiones 107 VIII. Paleontología sistemática 113 IX. Bioestratigrafía 201 X. Conclusiones 206 XI. Bibliografía 209 Láminas 246 Apéndices 265 A. Listado de materiales estudiados B. Listado de materiales comparativos C. Morfometría geométrica D. Listado de caracteres E. Matriz de caracteres F. Tabla de datos para GLM 1 Tesis Doctoral Universidad de Buenos Aires Damián Eduardo Pérez, 2015 La familia Carditidae (Mollusca-Bivalvia) en el Cenozoico del Atlántico Sudoccidental: Sistemática y Filogenia La familia Carditidae comprende a un grupo de bivalvos con un importante desarrollo durante el Cenozoico mundial, cuyas relaciones filogenéticas se encuentran pobremente dilucidadas. Si bien se encuentran presentes en casi todas las asociaciones cenozoicas de moluscos en Argentina, su conocimiento sistemático sigue siendo muy reducido. Este trabajo constituye en el primer estudio conjunto de la sistemática de los cardítidos fósiles patagónicos y del análisis filogenético cuantitativo de la familia, teniendo en cuenta las frecuentes variaciones morfológicas, de sus representantes. Para evaluar los casos de variación intraespecífica se utilizaron metodologías morfogeométricas, tanto contornos elípticos de Fourier como landmarks. En el caso de las relaciones filogenéticas se utilizó la metodología cladística, considerando 103 caracteres morfológicos de la conchilla y 61 terminales. Se identificaron los géneros Neovenericor Rossi de García, Levy y Caminos, 1980 (N. austroplata (Gardner y Bowles, 1939), N. paranensis (Borchert, 1801) y N. carrerensis (Griffin, 1991)), Cyclocardia Conrad, 1867 (“C.” cannada (Ihering, 1907), “C.” n. sp., “C.” nortensis (del Río, 1986), C.mesembria (Stilwell y Zinsmeister, 1992) y C. compressa (Reeve, 1843)), Pleuromeris Conrad, 1867 (P. sulcolunularis (Ihering, 1907) y P. fueguina (Steinmann y Wilckens, 19087)), Scalaricardita Sacco, 1899 (S. camaronesia (Ihering, 1907) y S. laciarina (Feruglio, 1954)), Cardites Link, 1807 (C. feruglioi (Petersen, 1946)), Claibornicardia Stenzel y Krause, 1957 (C. paleopatagonica (Ihering, 1907)), Rotundicardia Heaslip, 1968 (R. n. sp.) y Purpurocardia Maxwell, 1969 (P. elegantoides (Ortmann, 1899) y P. leonensis (del Río, 1986)). Se describieron tres nuevos géneros para incluir a las especies “Venericardia” patagonica (Sowerby, 1846), “V.” burmeisteri (Böhm, 1903) y “V.” tehuelchana (Ihering, 1907), respectivamente. El análisis filogenético permitió descartar el esquema subfamiliar propuesto por Chavan (1969) para la familia y se confirmó la separación de los linajes de cardítidos planicostados y alticostados. Neovenericor se ubicó en el primer grupo, pero separado de los planicostados del Hemisferio Norte. Los alticostados conforman un grupo monofilético e incluyen al nuevo género integrado por la especie “V.” burmeisteri. Se recuperó un linaje novedoso, que engloba a los géneros Cyclocardia, Pleuromeris y Scalaricardita. La ubicación en el análisis filogenético de las especies argentinas permitió corroborar lo propuesto en el análisis sistemático. “V.” iheringi (Bóhm, 1903) es el único taxón que atraviesa el límite K/P, mientras que el resto de las especies paleocenas se restringen al Daniano. Cyclocardia compressa es el único taxón fósil que sobrevivió hasta la actualidad. Palabras Clave: Patagonia - Argentina - Análisis filogenético - Bioestratigrafía - Paleógeno - Neógeno 2 Tesis Doctoral Universidad de Buenos Aires Damián Eduardo Pérez, 2015 The family Carditidae (Mollusca-Bivalvia) in the Cenozoic of the Southwestern Atlantic: Systematic and Phylogeny The family Carditidae comprises a group of bivalves with an important development during Cenozoic times. Despite this, their phylogenetic relationships are poorly elucidated. The carditids are recorded in almost all Argentinian Cenozoic fossil molluscan assemblages, but their systematic knowledge remains reduced. This work constitutes the first integrated approach that includes the systematic study of the Patagonian fossil carditidsand a quantative phylogenetic study of the family, taking into account the very frequent morphological variations among its representatives. To evaluate the intraspecific variations, morphometric geometric tools were applied, using both elliptic Fourier outlines and landmarks. In the case of the phylogenetic relationships, cladistic methodology was used, including 103 shell morphological characters, and 61 terminals. It was identified the presence of the genera Neovenericor Rossi de García, Levy and Caminos, 1980 (N. austroplata (Gardner and Bowles, 1939), N. paranensis (Borchert, 1801) and N. carrerensis (Griffin, 1991)), Cyclocardia Conrad, 1867 (“C.” cannada (Ihering, 1907), “C.” n. sp., “C.” nortensis (del Río, 1986), C.mesembria (Stilwell and Zinsmeister, 1992) andC. compressa (Reeve, 1843)), Pleuromeris Conrad, 1867 (P. sulcolunularis (Ihering, 1907) andP. fueguina (Steinmann y Wilckens, 19087)), Scalaricardita Sacco, 1899 (S. camaronesia (Ihering, 1907) and S. laciarina (Feruglio, 1954)), Cardites Link, 1807 (C. feruglioi (Petersen, 1946)), Claibornicardia Stenzel and Krause, 1957 (C. paleopatagonica (Ihering, 1907)), Rotundicardia Heaslip, 1968 (R. n. sp.) and Purpurocardia Maxwell, 1969 (P. elegantoides (Ortmann, 1899) and P. leonensis (del Río, 1986)). Three new genera were described where the species “Venericardia” patagonica (Sowerby, 1846), “V.” burmeisteri (Böhm, 1903) and “V.” tehuelchana (Ihering, 1907) were respective included. The phylogenetic analysis allowed to reject the subfamiliar scheme proposed by Chavan (1969), and confirmed the separation between planicostate and alticostate carditids. Neovenericor was placed in the first group, but separated from the Northern Hemisphere planicostates. The alticostate carditids are a monophyletic group, and comprises the new genus that included to “V.” burmeisteri. A novel lineage was recovered that embraces the genera Cyclocardia, Pleuromeris, and Scalaricardita. The systematic placement of the Argentinian species was corroborated by the phylogenetic analysis. Only “V.” iheringi (Böhm, 1903) goes through the K/P boundary, while remaining Paleocene carditids are restricted to Danian times. Cyclocardia compressa is the only fossil taxon that survived into Recent times. Keywords: Patagonia - Argentina - Phylogenetics Analysis - Biostratigraphy -Paleogene - Neogene 3 Tesis Doctoral Universidad de Buenos Aires Damián Eduardo Pérez, 2015 AGRADECIMIENTOS Esto es solo otro paso en el camino, otro punto para seguir andando. Cuando de chico le decía a mis padres que quería ser paleontólogo no tenía idea de como se iban a ir dando, y siento que estoy aportando un ladrillo más para cumplirle el sueño a ese niño. Nunca hubiera podido realizar esta tesis doctoral sino hubiese sido por el apoyo de mi familia,
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