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Sistemática De Fitolitos, Pautas Para Un Sistema Clasificatorio. Un Caso En Estudio En La Formación Alvear (Pleistoceno Inferior)

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Sistemática De Fitolitos, Pautas Para Un Sistema Clasificatorio. Un Caso En Estudio En La Formación Alvear (Pleistoceno Inferior) AMEGHINIANA (Rev. Asoc. Paleontol. Argent.) - 42 (4): 000-000. Buenos Aires, 30-12-2005 ISSN 0002-7014 Sistemática de fitolitos, pautas para un sistema clasificatorio. Un caso en estudio en la Formación Alvear (Pleistoceno inferior) Alejandro Fabián ZUCOL1 y Mariana BREA1 Abstract. PHYTOLITH SYSTEMATICS, GUIDELINE FOR A CLASSIFICATORY SYSTEM. A STUDY CASE IN THE ALVEAR FOR- MATION (LOWER PLEISTOCENE). The classification rules to establish a phytolith systematics are outlined, in such a way allowing a precise treatment and delimitation of their ranks according to the botanical no- menclatural code. The morphological terminology used is explained and these rules are applied to the predominant phytolith morphotypes of the Alvear Formation in their type area, Puerto General Alvear (Diamante department, Entre Ríos). Its sediments were deposited during an semi-arid interval in typical Lower Pleistocene pampean conditions. Globulolithum gen. nov. (G. sphaeroechinulathum sp. nov., G. spha- eropsilathum sp. nov.), Aculeolithum gen. nov. (A. rostrathum sp. nov., A. acuminathum sp. nov., A. ancistrat- hum sp. nov., A. aciculathum sp. nov.), Flabelolithum gen. nov. (F. euflabelathum sp. nov., F. complanathum sp. nov.) and Macroprismatolithum gen. nov. (M. psilaristathum sp. nov., M. denticulathum sp. nov., M. ondulat- hum sp. nov., M. excavathum sp. nov.) are described. This assemblage represents the first fossil record for this formation. Resumen. Se plantean las pautas clasificatorias para establecer una sistemática de fitolitos que permita un preciso tratamiento y delimitación de sus categorías, de acuerdo a las normativas de la nomenclatura bo- tánica. Conjuntamente se expone la terminología morfológica que se utiliza y se aplican estas pautas a la descripción de los morfotipos fitolíticos más abundantes de la asociación fitolítica hallada en la Formación Alvear, expuesta en su área tipo, la localidad de Puerto General Alvear (departamento Diamante, Entre Ríos), cuyos sedimentos han sido depositados durante un intervalo de clima semiárido de características típicamente pampeanas del Pleistoceno inferior. Se describen las características de los géneros Globulolithum gen. nov. (G. sphaeroechinulathum sp. nov., G. sphaeropsilathum sp. nov.), Aculeolithum gen. nov. (A. rostrathum sp. nov., A. acuminathum sp. nov., A. ancistrathum sp. nov., A. aciculathum sp. nov.), Flabelolithum gen. nov. (F. euflabelathum, sp. nov. F. complanathum sp. nov.) y Macroprismatolithum gen. nov. (M. psilaristathum sp. nov., M. denticulathum sp. nov., M. ondulathum sp. nov., M. excavathum sp. nov.). Esta asociación es el único registro fósil conocido para esta formación. Key words. Phytoliths. Classification rules. Alvear Formation. Lower Pleistocene. Palabras clave. Fitolitos. Pautas clasificatorias. Formación Alvear. Pleistoceno inferior. Introducción tos estableciendo una serie de morfogéneros y mor- foespecies; este sistema sería modificado más tarde Los estudios fitolíticos han tenido una evolución por Deflandre (1963). en estrecha vinculación con distintas disciplinas Bertoldi de Pomar (1971) propone un ensayo de científicas como consecuencia de sus aplicaciones y clasificación morfológica definiendo grupos y mor- de la potencialidad de estos estudios en Botánica, Pa- fotribus de fitolitos a los cuales da una denominación leobotánica, Pedología, Sedimentología y Arqueolo- particular. Ambos sistemas clasificatorios y nomen- gía (Zucol, 1992). Si bien en sus comienzos los fitoli- claturales, en un primer momento independientes, tos fueron estudiados como parte componente de los fueron integrados por Taugourdeau-Lantz et al., (1976) tejidos vegetales, pronto surgió la necesidad de esta- en una propuesta clasificatoria. Estos autores, por blecer su tratamiento en forma desarticulada del ve- principio de prioridad, toman las categorías de Eh- getal que los originó. Ehrenberg (1841, 1846 y 1854) renberg, y las recategorizan en grupo, género y espe- realiza la primera clasificación morfológica de fitoli- cies morfológicas, incluyendo las clases morfológicas propuestas por Bertoldi de Pomar. En forma coetánea surgen varias clasificaciones de fitolitos, las que se caracterizan por abarcar gru- pos de vegetales restringidos ya sea taxonómica o 1Laboratorio de Paleobotánica. Centro de Investigaciones Cientí- ficas, Diamante (CONICET), Matteri y España sn, 3105 Diamante, geográficamente (Zucol, 1995). Este enfoque se conti- Argentina. [email protected] nuó utilizando hasta la actualidad, y como conse- ©Asociación Paleontológica Argentina AMGHB2-0002-7014/05$00.00+.50 2 A.F. Zucol y M. Brea cuencia de ello es frecuente encontrar en la literatura fitolítica que un mismo tipo morfológico es denomi- nado de diferentes modos, o que en las distintas cla- sificaciones un mismo tipo morfológico posee rangos de variabilidad disímiles. En la presente contribución se plantean las pautas clasificatorias para establecer una sistemática de fito- litos que permita un preciso tratamiento y delimita- ción de las categorías, de acuerdo a las normativas de la nomenclatura botánica. Conjuntamente se expone la terminología morfológica que se utiliza y se apli- can estas pautas en la descripción de los morfotipos fitolíticos más abundantes en la asociación fitolítica hallada en la Formación Alvear, expuesta en la loca- lidad de Puerto General Alvear, departamento Diamante, provincia de Entre Ríos (figura 1). La Formación Alvear se presenta en forma casi continua en la margen derecha del río Paraná desde el arroyo Nogoyá hasta la ciudad de Paraná. Esta unidad de origen palustre esta integrada por sedi- mentos carbonáticos y fue asignada al Pleistoceno in- ferior, de acuerdo a su posición estratigráfica. Este paquete sedimentario fue depositado durante un in- tervalo de clima semiárido de características típica- mente pampeanas (Iriondo, 1980). La presente comu- nicación forma parte de la descripción detallada de la asociación fitolítica de esta Formación (Zucol y Brea, 2000 b, en prensa), la cual, hasta la actualidad consti- Figura 1. Ubicación geográfica y sedimentológica de la Formación tuye el único tipo de registro fósil hallado. Alvear en la localidad de Puerto General Alvear / Sedimentological and geographical setting of Alvear Formation in Puerto General Alvear locality. Pautas para el tratamiento morfológico de los fitolitos tativamente superior, que el espectro fitolítico se en- cuentra dominado por los fitolitos de origen silíceo o Para la cabal comprensión de la morfología de los silicofitolitos. fitolitos se debe tener en cuenta un conjunto de ca- Los fitolitos están presentes en todo el reino vege- racterísticas, que Bertoldi de Pomar (1975) denominó tal, pero es en los vegetales superiores donde su “caracteres biogenéticos”; tales como, plantas prove- identificación y asignación sistemática se torna más edoras, concentración de fitolitos en las plantas y en controvertida. En lo que respecta a su producción, los las distintas etapas fenológicas, tipos de tejidos y cé- fitolitos se encuentran en representantes de pteridó- lulas en que se originan, mecanismos de deposita- fitas, gimnospermas y angiospermas (Piperno, 1988; ción, composición química y propiedades físicas de Pearsall, 2000), siendo ciertas familias de las mono- los fitolitos, para luego establecer los caracteres mor- cotiledóneas (como Poaceae, Cyperaceae, Arecaceae) fométricos de cada tipo morfológico. las principales proveedoras. En estos vegetales, la Como resultado del metabolismo, en los vegetales presencia de sílice se encuentra asociada no sólo a se producen distintos tipos de depositaciones mine- procesos metabólicos intrínsecos, sino también al cre- rales; muchas de las cuales, como consecuencia de cimiento y productividad del vegetal, y a fenómenos sus propiedades físico químicas, no llegan a perdurar de resistencia a enfermedades, toxicidades, ataques como fitolitos en el amplio sentido de la palabra. de insectos y efectos producidos por la herbivoría Dentro del espectro de sustancias minerales biogéni- (Rovner, 1988). cas, en los vegetales son predominantes -ya sea por De acuerdo a ello, la concentración de los fitolitos sus características físico químicas como por su abun- en la planta variará con la etapa fenológica del vege- dancia- los depósitos de minerales cálcicos y silíceos. tal, como así también, en relación con las diferentes Químicamente, los minerales cálcicos (tales como los partes de la planta (Bertoldi de Pomar, 1975; Zucol, oxalatos de calcio) son más fácilmente alterables que 2001). La concentración de sílice será variable de los silíceos. Es por ello, y por una producción cuanti- acuerdo a los procesos metabólicos que acompañan AMEGHINIANA 42 (4), 2005 Sistemática de fitolitos 3 la vida de la planta, y a la localización de muchos de prismaticos con predominio de uno de sus ejes se en- estos procesos en ciertos órganos. Parry y Smithson cuentran orientados en sentido paralelo a la superfi- (1958 a) describen el proceso de silicificación de célu- cie de los órganos en donde se originan (con excep- las buliformes foliares de gramíneas, y observan que ción de por ejemplo las células buliformes silicifica- ésta mineralización se incrementa en etapas próxi- das), es que sus caras se denominan: periclinales, a mas a la desecación de la hoja, que es cuando éstas las caras dispuestas en forma paralela a la superficie células pierden su importancia funcional en la aper- del órgano vegetal y anticlinales a las que se dispo- tura y cierre higroscópico de la hoja, como así tam- nen en
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