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Nuevos Aportes a La Sistemática De Los “Plohophorini” De Uruguay (Mammalia, Cingulata, Glyptodontidae)

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Nuevos Aportes a La Sistemática De Los “Plohophorini” De Uruguay (Mammalia, Cingulata, Glyptodontidae) NUEVOS APORTES A LA SISTEMÁTICA DE LOS “PLOHOPHORINI” DE URUGUAY (MAMMALIA, CINGULATA, GLYPTODONTIDAE) Pablo Toriño Licenciatura en Ciencias Biológicas, profundización en Paleontología. Instituto de Ciencias Geológicas - Departamento de Paleontología. Facultad de Ciencias (UdelaR). Orientador: Dr. Daniel Perea Tribunal: Dr. Sergio Martínez, Dr. Martín Ubilla Montevideo, 2015 - 1 - Imagen de portada: Placa de coraza asignada al gliptodonte Pseudoplohophorus en el presente trabajo (FC-DPV 384), hallada en las barrancas de San Gregorio, departamento de San José (Uruguay). Procedencia estratigráfica: Formación Camacho (Mioceno Tardío). - 2 - A la memoria de Domingo Pablo Toriño (1944-2010), mi Padre, mi Héroe. Y de Maru (2004-2015), por tantos lindos recuerdos. - 3 - - 4 - Indice Resumen ………………………………………………………………………………………………………………………………………………………7 Abstract ………………………………………………………………………………………………………………………………………………………..8 1. Introducción …………………………………………………………………………………………………………………………………………….9 1.1. Antecedentes y fundamentación ………………………………………………………………………………………………………..9 1.2. Objetivos ……………………………………………………………………………………………………………………………………………21 2. Materiales y metodología …………………………………………………………………………………………………………………… 22 2.1. Relevamiento de colecciones ……………………………………………………………………………………………………………..22 2.2. Revisión taxonómica …………………………………………………………………………………………………………………………..24 2.3. Obtención de medidas ……………………………………………………………………………………………………………………….24 2.4. Análisis cuantitativos ………………………………………………………………………………………………………………………….34 3. Resultados ……………………………………………………………………………………………………………………………………………….36 3.1. Revisión y propuesta taxonómica ………………………………………………………………..……………………………………..36 Plohophorus ………………………………………………………………………………………………………………………………………..43 Plohophorus figuratus …………………………………………………………………………………………………………………….44 Parahoplophorus …………………………………………………………………………………………………………………………………47 Parahoplophorus paranensis ………………………………………………………………………………………………………….48 Stromaphoropsis ………………………………………………………………………………………………………………………………….50 Stromaphoropsis scavinoi ………………………………………………………………………………………………………………52 Pseudoplohophorus ……………………………………………………………………………………………………………………………..57 Nuevos sinónimos junior propuestos ……………………………………………………………………………………… 61 Berthawyleria ……………………………………………………………………………………………………………………… 61 Asymmetrura ………………………………………………………………………………………………………………………..63 Pseudoplohophorus orientalis ………………………………………………………………………………………………………..64 Pseudoplohophorus francisi ……………………………………………………………………………………………………………69 Pseudoplohophorus coloniensis ………………………………………………………………………………………………………70 Pseudoplohophorus berroi ……………………………………………………………………………………………………………..75 Pseudoplohophorus benvenutii ………………………………………………………………………………………………………78 Pseudoplohophorus rebuffoi …………………………………………………………………………………………………………..87 Pseudoplohophorus absolutus ………………………………………………………………………………………………………..93 Pseudoplohophorus compressus NOV. COMB. ……………………………………………………………………………………102 Pseudoplohophorus uncinatus NOV. COMB. ……………………………………………………………………………………. 106 Pseudoplohophorus gracilis NOV. COMB. ………..…………………………………………………………………………………109 - 5 - Pseudoplohophorus orientalis (non Ameghino) NOV. COMB. …………………………………………………………….114 Pseudoplohophorus sp. …………………………………………………………………………………………………………………..117 Ejemplares previamente asignados a otros taxones ………………………………………………………………….117 Nuevos materiales ……………………………………………………………………………………………………………………124 Status propuesto para otros materiales citados …………………………………………….………………………………….. 154 “Hoplophorinae” insertae sedis …………………………………………………………………………………………………….154 Zaphilus …………………………………………………………………………………………………………………………………. 154 Zaphilus larranagai …………………………………………………………..…………………………………………………155 “Hoplophorinae” gen. et sp. indet. …………………………………………………………………………………………………159 Cingulata gen. et sp. indet. …………………………………………………………………………………………………………….161 3.2. Análisis cuantitativos .…………………………………………………………………………………………………………………………164 4. Discusión ………………………………………………………………………………………………………………………………………………….171 4.1. Exoesqueleto y sistemática en “Plohophorini” …………………………………………………………………………………. 171 4.2. Caracterización del género Pseudoplohophorus ……………………………………………………………………………… 180 4.1. Consideraciones finales ………………………………………………………………………………………………………………………18 4 5. Referencias ………………………………………………………………………………………………………………………………………………18 6 Anexo I: Tablas de medidas ………………………………………………………………………………………………………………………20 1 Anexo II: Lista de materiales citados ………………………………………………………………………………………………20 7 Créditos de las imágenes ……………………………………………………………………………………………………………………………2 16 Agradecimientos …………………………………………………………………………………………………………………………………………2 18 - 6 - Resumen Los “Plohophorini” (Mioceno tardío–Pleistoceno de Argentina y Uruguay) constituyen uno de los grupos taxonómicamente más complejos dentro de los Glyptodontidae, con una monofilia cuestionada en recientes análisis filogenéticos. La tribu se caracteriza por una pobre representación de la mayoría de sus especies, varias de las cuales se basan en ejemplares tipo altamente fragmentarios, con escasos ejemplares referidos, cuando no inexistentes, a lo que se suma una notoria escasez de revisiones sistemáticas recientes. Además de algunos materiales fragmentarios dudosamente atribuidos a los géneros argentinos Plohophorus y Parahoplophorus, en Uruguay el registro de la tribu ofrece tradicionalmente un complejo de géneros endémicos mayormente conocidos por restos parciales de exoesqueletos (principalmente fragmentos de corazas y/o tubos caudales). Este complejo presenta esencialmente una distribución geográficamente acotada a la costa sur-oeste del país (departamentos de San José y Colonia), y estratigráficamente a la Formación Camacho (Piso/Edad Huayqueriense, Mioceno Tardío). A la luz de la constatación de la existencia de nuevos ejemplares uruguayos inéditos asignables a la tribu, se realizó un análisis comparativo de los mismos sobre la base de criterios cualitativos y cuantitativos (análisis multivariados de cráneos y tubos caudales), empleando a los “Plohophorini” de Uruguay y Argentina, y algunos “Hoplophorini” argentinos como grupo comparativo. Los resultados obtenidos señalan en primer lugar una mayor similitud de los “Plohophorini” uruguayos entre sí y con ciertos “Hoplophorini”, que con la mayoría de los “Plohophorini” argentinos, los cuales presentan una considerable variabilidad morfológica. En segundo lugar, el análisis de la totalidad de los ejemplares permite postular que varios de los taxones uruguayos involucrados fueron establecidos bajo una óptica netamente morfotípica y descriptiva, propia de una metodología ameghiniana de uso extendido al menos hasta la primera mitad del Siglo XX. Se propone reducir así la lista de géneros uruguayos a dos: Stromaphoropsis y Pseudoplohophorus (incluyendo en este a “Teisseiria”, Berthawyleria y Asymmetrura), manteniendo provisionalmente la identidad de las especies involucradas, hasta tanto nuevos hallazgos permitan comprender mejor la probable variabilidad intraespecífica presente en los taxones estudiados. - 7 - Abstract NEW CONTRIBUTIONS TO THE SYSTEMATICS OF THE “PLOHOPHORINI” OF URUGUAY (MAMMALIA, CINGULATA, GLYPTODONTIDAE). – The "Plohophorini" (Late Miocene – Pleistocene of Argentina and Uruguay) are one of the taxonomically more complex groups within Glyptodontidae, with a monophyly questioned in recent phylogenetic analysis. The tribe is characterized by a poor representation of the majority of its species, several of which are based on very fragmentary type specimens, with few referred specimens, even none, together with a noticeable shortage of recent systematic reviews. In addition to some fragmentary materials dubiously attributed to the argentinean genera Plohophorus and Parahoplophorus, in Uruguay the record of the tribe traditionally offers a complex of endemic genera mostly known by partial remains of exoskeletons (mainly fragments of carapaces and/or caudal tubes). This complex presents essentially a geographic distribution bounded to the south-west of the country (departments of San Jose and Colonia) and a stratigraphic distribution bounded to the Camacho Formation (Huayquerian SALMA, Late Miocene). In the light of the presence of new uruguayan, unpublished specimens referable to the tribe, a comparative analysis of them was made on the basis of qualitative and quantitative criteria (multivariate analysis of skulls and caudal tubes), using the "Plohophorini" from Uruguay and Argentina, and some argentinean “Hoplophorini " as a comparative group. The results obtained shows first a mayor similitude of the uruguayan "Plohophorini" among themselves and with certain "Hoplophorini", rather than with the most of the argentinean "Plohophorini", which shows a large morphological variability. Second, the analysis of all of the specimens allows to propose that several of the uruguayan taxa involved were established under a purely descriptive morphotypical criterion, own of an “ameghinian” methodology spread at least until the first half of the twentieth century. A reduction of the list of uruguayan genera up to two is proposed: Stromaphoropsis and Pseudoplohophorus (including in this one "Teisseiria", Berthawyleria and Asymmetrura), provisionally keeping the identity of the species involved, until new findings enable a better understanding about the probable intraspecific variability present in the studied taxa. - 8 - 1. Introducción 1.1. Antecedentes y fundamentación Desde un punto de vista taxonómico y nomenclatural, los gliptodontes constituyen uno de los grupos de mamíferos fósiles más complejos del Nuevo Mundo.
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