Brontornis Burmeisteri Moreno & Mercerat, Un Anseriformes (Aves

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Brontornis Burmeisteri Moreno & Mercerat, Un Anseriformes (Aves Rev. Mus. Argentino Cienc. Nat., n.s. 9(1): 15-25, 2007 Buenos Aires, ISSN 1514-5158 Brontornis burmeisteri Moreno & Mercerat, un Anseriformes (Aves) gigante del Mioceno Medio de Patagonia, Argentina Federico L. AGNOLIN Laboratorio de Anatomía Comparada y Evolución de los Vertebrados, Museo Argentino de Ciencias Naturales «Bernardino Rivadavia», Av. Angel Gallardo 470 (1405) Buenos Aires, Argentina. Área Paleontología, Fundación de Historia Natural «Félix de Azara», Departamento de Ciencias Naturales y Antropología. CEBBAD - Universidad Maimónides, Valentín Virasoro 732 (C1405BDB) Buenos Aires, Argentina, e-mail: [email protected]. Abstract: Brontornis burmeisteri Moreno & Mercerat, a giant Anseriformes (Aves) from the Middle Miocene of Patagonia, Argentina. Since its original description, Brontornis burmeisteri was considered as closely related to the family Phorusrhacidae (Gruiformes). Brontornis is known only by scarce postcranial remains, fragmentary mandibles and a quadrate bone. All the material comes from the Miocene of Santa Cruz Province, Argentina. A reanalysis of this material indicates that Brontornis may be more properly included within the Anseriformes as a basal member of this group, together with Diatryma and the extinct family Dromornithidae. Its inclusion within the Anseriformes suggests a more complex evolutionary history of the poorly known paleoavifaunas of South America. Key words: Brontornis, Phorusrhacidae, Anseriformes, Miocene, Argentina. ____________ Desde su descripción original, Brontornis dos con la graviportalidad (Tonni, 1977; Alva- burmeisteri Moreno & Mercerat, 1891 fue consi- renga & Hofling, 2003). derado como el representante más robusto y de Un análisis detallado del escaso material mayor tamaño de los Stereornithes (= Pho- craneano conocido, así como del material post- rusrhacidae Ameghino, 1889), criterio seguido craneano, indica una diferente posición filo- por la mayoría de los autores subsiguientes (e.g., genética para Brontornis. En este trabajo se pro- Patterson & Kraglievich, 1960; Alvarenga, 1993; pone que este género se encontraría más cer- Alvarenga & Hofling, 2003; Tambussi & Noriega, canamente relacionado a los Anseriformes que a 1996). Moreno & Mercerat (1891) remarcaron las los Phorusrhacidae (Gruiformes). Los Anse- notables semejanzas que presentaba Brontornis riformes vivientes se encuentran representados con el género Cygnus (Anseriformes), en espe- por el clado Anseres, el cual se compone de tres cial en el tibiotarso y fémur. Por otro lado, familias de aves acuáticas voladoras: los Ana- Dolgopol de Sáez (1927) indicó importantes dife- tidae, de distribución cosmopolita, los Anse- rencias (especialmente en el tarso y falanges) ranatidae y finalmente los Anhimidae, endémi- entre Brontornis y el resto de los Phorusrhacidae cos de Oceanía y Sudamérica, respectivamente. (=Stereornithes), y en consecunecia erigió, para Por otro lado, el registro fósil de Anseriformes el primero, un nuevo grupo de aves denominado basales se encuentra restringido principalmente a Brontornithes. Esta hipótesis ha sido criticada por formas de gran tamaño, de proporciones Kraglievich (1932) y la supresión definitiva de los graviportales y con capacidad nula de vuelo. Brontornithes, junto con su inclusión den-tro de Den-tro de estos taxones se presenta el género los Phororhacoidea, fue llevada a cabo pos- extin-to Diatryma Cope, 1876 del Eoceno de teriormente por Patterson & Kraglievich (1960). América del Norte, el cual ha sido considerado Dentro de los Phororhacoidea, Brontornis fue original-mente como un posible Palaeognathae considerado como cercanamente emparentado a (Cope, 1876) y posteriormente como un miembro los géneros Physornis Ameghino, 1895 y Para- del orden Gruiformes, probablemente relacionado physornis Alvarenga, 1993, conformando la fa- a los Cariamoidea (Mathew & Granger, 1917). Su milia (o subfamilia) Brontornithinae, la cual ha inclusión dentro de los Gruiformes ha sido pues- sido fundamentada principalmente porque todos ta en duda primeramente por Olson (1985). sus integrantes poseen extremidades posteriores Andors (1992) ha ubicado Diatryma dentro de los cortas y robustas y el tarsometatarso ancho y Galloanserae y lo incluyó acertadamente dentro anteroposteriormente aplanado (Alvarenga & de los Anseriformes. Consecuentemente, Diatry- Höfling, 2003), caracteres todos estos relaciona- ma ha pasado de ser una gigantesca grulla carní- 16 Revista del Museo Argentino de Ciencias Naturales, n. s. 9 (1), 2007 Fig. 1. Cladograma de consenso indicando la posición filogenética de Brontornis. Los números en ne-grita indican el número de carácter y el número entre paréntesis el estado del mismo. Los caracteres subrayados representan homoplasias. Galloanserae: 16(1), 21(1), 27(1); Anseriformes: 2(1), 6(1), 9(1), 12(1), 14(1), 17(1), 19(1), 20(1), 30(1), 36(1), 37(1), 38(1), 39(1), 40(1), 41(1), 43(1), 45(1), 47(0), 48(1); Nodo 1: 27(1), 25(1); Nodo 2: 18(1); Anseres: 3(1), 5(1), 15(1), 47(1), 48(0); Nodo 3: 4(1), 7(1), 10(1), 28(1), 29(1), 31(1), 32(1), 33(1), 34(1), 35(1). Abreviaturas: Phorus.: Phorusrhacidae. vora a un pacífico «pato gigante» (Murray & (altura hasta la cabeza) y su peso en más de 350 Megirian, 1998; véase Witmer & Rose, 1991). kg (Alvarenga & Hofling, 2003). Seguramente Adicionalmente, los géneros Gastornis Hébert, incapacitado de volar, como lo sugieren las pro- 1855 y Zhongyuanus Hou, 1980 del Eoceno de porciones de los huesos conservados (la longitud Europa y Asia, respectivamente han sido inclui- del tarsometatarso es aproximadamente un 50 dos junto con Diatryma en la familia Gastor- % de la del tibiotarso), y el peso estimado, nithidae (Martin, 1992; Andors, 1992). Recien- Brontornis con toda seguridad era un ave de con- temente, una familia de supuestos Palaeognathae dición graviportal (Tonni, 1977). La gran robus- graviportales de gran tamaño fue erigida para tez de la mandíbula de este taxón, sumada a sus abarcar materiales fósiles del Mioceno-Pleis- extremidades de proporciones indicativas de toceno de Oceanía. Esta agrupación fue denomi- graviportalidad, llevó a especular sobre los posi- nada Dromornithidae y originalmente fueron bles hábitos alimenticios de Brontornis, conside- considerados relacionados a los Casuariidae, prin- rándosele principalmente carroñeros (Tonni, cipalmente sobre la base de caracteres potcra- 1977; Alvarenga & Hofling, 2003). neanos (Rich, 1980; 1992). Sin embargo, Murray El objetivo del presente trabajo es reanalizar & Megirian (1998) después de un detallado estu- los materiales conocidos de Brontornis burmeis- dio del cráneo de los Dromornithidae concluye- teri, así como reconsiderar su posición filo- ron que esta familia debería ser incluida dentro genética. de los Anseriformes, como el posible grupo her- mano de los Anhimidae. De este modo, los gru- METODOS pos-hermanos de los Anseriformes vivientes los constituirían agrupaciones de aves con nula ca- Los caracteres utilizados en el análisis filo- pacidad de vuelo, graviportales y de gran tama- genético (Apéndice I) han sido tomados de ob- ño corporal. servaciones personales e información édita. La Brontornis burmeisteri es, sin duda alguna, matriz elaborada para analizar la posición el ave de mayor tamaño que habitó América, su- filogenética de Brontornis consiste en 51 carac- perando inclusive al género norteamericano teres y 12 taxones terminales. Se utilizan como Diatryma. Su tamaño ha sido calculado en 2,8 m. grupos externos las Familias Tinamidae y Ralli- Agnolin: Brontornis, Anseriformes basal del Mioceno de Argentina 17 dae. Se incluyen miembros bien conocidos de la Familia Phorusrhacidae (i.e., Paraphysornis, Tolmodus, Phorusrhacos), Galliformes, y varios Anseriformes basales (i.e., Diatryma, Dromor- nithidae, Anseranatidae, Anhimidae), así como los Anatoidea. No se ha incluido en el análisis cladístico al Phorusrhacidae Physornis (un gé- nero supuestamente cercano a Brontornis) debi- do a la naturaleza incompleta sobre la cual se ha basado este taxón. Del mismo modo, los Gas- tornithidae Gastornis y Zhongyuanus no son in- cluidos en el análisis cladístico debido a su natu- raleza fragmentaria (véase Martin, 1992). Se analizaron los datos bajo el programa NONA (Goloboff, 1993) mediante análisis heurístico con un máximo obtenible de árboles de 10000 y un número de replicaciones de 1000. La estrategia de búsqueda utilizada es TBR + TBR (mult*max*). Se obtuvieron tres clado-gramas, Fi. 2. Brontornis burmeisteri. Cuadrado izquierdo en vis- cuyo consenso estricto brindó un árbol de una tas: (A) anterior; (B) posterior; (C) distal. Escala 2 cm. longitud de 61 pasos, con un Indice de Consistencia de 81 y un Indice de Retención de 91 (Fig. 1). Todos los caracteres fueron corridos 1895; actualmente sinónimo junior de Brontornis con polaridad libre. platyonyx; según Alvarenga & Höfling, 2003). Se sigue la terminología osteológica de Diagnosis. Brontornis burmeisteri puede ser Howard (1926) y Baumel & Witmer (1993) con diagnosticado sobre la base de las siguientes modificaciones, y el esquema estratigráfico de autapomorfías: 1) tróclea III del tarsometatarso Legarreta & Uliana (1994). Con respecto a los con una fuerte proyección dorsomedial de la su- Phorusrhacidae se adopta la nomenclatura perficie articular (Alvarenga & Höfling, 2003), 2) taxonómica empleada por Tonni (1980), con mo- puente supratendinoso del extremo distal del dificaciones. tibiotarso ausente, 3) continuación dorsal del Abreviaturas. BMNH, British Museum of surco intercondilar del cuadrado. Natural
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