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7. Phytolith Paleosols Analysis

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UNIVERSIDAD COMPLUTENSE DE MADRID FACULTAD DE GEOGRAFÍA E HISTORIA TESIS DOCTORAL Reconstrucción de la paleovegetación y uso de útiles líticos en el yacimiento de homínidos de la garganta de Olduvai (Tanzania) mediante el uso de microfósiles vegetales Paleovegetation and stone tool use at a selection of hominin sites and their associated landscapes from Olduvai Gorge (Tanzania): a study of plant microfossils MEMORIA PARA OPTAR AL GRADO DE DOCTOR PRESENTADA POR Héctor Arráiz Rodríguez Directores Manuel Domínguez-Rodrigo Doris Barboni Madrid, 2017 © Héctor Arráiz Rodríguez, 2016 Tesis Doctoral Reconstrucción de la paleovegetación y uso de útiles líticos en el yacimiento de homínidos de la garganta de Olduvai (Tanzania) mediante el uso de microfósiles vegetales Paleovegetation and stone tool use at a selection of hominin sites and their associated landscapes from Olduvai Gorge (Tanzania). A study of plant microfossils Autor: HÉCTOR ARRÁIZ RODRÍGUEZ Directores: MANUEL DOMÍNGUEZ-RODRIGO DORIS BARBONI FACULTAD DE GEOGRAFÍA E HISTORIA Tesis Doctoral Reconstrucción de la paleovegetación y uso de útiles líticos en el yacimiento de homínidos de la garganta de Olduvai (Tanzania) mediante el uso de microfósiles vegetales Paleovegetation and stone tool use at a selection of hominin sites and their associated landscapes from Olduvai Gorge (Tanzania). A study of plant microfossils. Autor: HÉCTOR ARRÁIZ RODRÍGUEZ Directores: MANUEL DOMÍNGUEZ-RODRIGO DORIS BARBONI FACULTAD DE GEOGRAFÍA E HISTORIA Agradecimientos Conviene empezar por los formalismos, pero es justo agradecer al Ministerio de Economía y Competitividad (otrora Ministerio de Ciencia e Innovación) el haberme concedido una beca FPI y posibilitar el desarrollo de este trabajo. Gracias también a la Universidad Complutense de Madrid por haber sido la sede académica de esta tesis y del que esto escribe. Mil gracias, infinitas gracias, a Doris Barboni y a Manuel Domínguez-Rodrigo, codirectores de esta tesis, sin los que este trabajo hubiera sido imposible. Gracias por acoger a quien desconocía casi todo de la materia a la que se enfrentaba y ahora sólo desconoce gran parte. Doris, merci pour ton soutien, ton patience et ton amitié. Sans toi, l’atterrissage à Aix aurait été impossible. Gracias a Enrique Baquedano por abrirme las puertas del Museo Arqueológico Regional de la Comunidad de Madrid para poder desarrollar mi tesis doctoral. Qué decir de Belén, César y Mari, quienes han sido mi pequeña familia en esa casa, gracias por vuestros consejos. Gracias a Blanca Ruiz y a María José Gil por prestarme su laboratorio en la Universidad de Alcalá de Henares, a Tomás Martín por su ayuda en ese laboratorio y a los tres por los cafés en el patio. Agradezco a Santos Ballesteros (técnico de la UAH) la ayuda prestada a un extraño a esa institución. Gracias a todos los que, de alguna u otra manera, habéis colaborado en el trabajo de esta tesis doctoral (junto a los ya mencionados): Fernando Díez-Martín, David Uribelarrea, José Yravedra, Marine Pasturel, Nicolas Barbarin, Luc-Beaufort, Alice Novello, Jean Charles-Mazur… Agradezco, también, la labor de los evaluadores y miembros del tribunal que juzgan esta tesis. Aixoises, ¡qué momentos! Lo mejor que me llevo de esta tesis es un puñado de amigos y amigas, ¡la familia que hizo de Aix mi segunda casa!: Anita, Meritxell, Fatiha, Olivier Dahan, Vladi, Ghislain, Gaby, Fabrice, Greg, Barbara, Yolaina, Olivier, Paco, Elena… (el orden es aleatorio, no os pongáis celosos). Cito aparte a Adelia y Dave, 3 porque aparte de brindarme su amistad, he de agradecer su inestimable ayuda con la corrección del inglés de esta tesis. Nos volveremos a juntar muy pronto en el Hublot (gracias también a la “fauna” que allí habitamos y a Fethi por tolerarnos). Je vous aime! Je ne sais pas pourquoi, mais je vous aime! A los de aquí (o que aquí viven)… Diego, gracias por todo, piso, amistad y gin- tonics intempestivos. Gracias Pablo y Ana Belén por los cafés en la complu y la charleta sobre las penas del doctorando, a Edgar y Rubén por las cervezas por tierra de gatos buscando tascas. Leoneses, cántabros, burgaleses, pucelanos, etc… los de siempre, ya sabéis quienes sois y sois tanto… colegio, instituto, los de la universidad (gracias Félix y Carmen por guiarme en los primeros pasos que di en la ciencia y a Estrella y Alicia por compartirlos). Son muchos años que forjaron mi carácter, por los bares compartidos, por los campings, por las lluvias, el teatro, los jueves santos, por el fútbol (¡aúpa Sporting de Pinilla!), por las tardes perdidas en vuestras casas (o no, según se mire)… A Ruth por todo. Y a mi familia, claro, por su cariño y ayuda. Sin vosotros esto hubiera sido imposible. 4 Resumen Esta tesis doctoral presenta los resultados de una serie de análisis multidisciplinares que tienen como objeto los microfósiles de plantas. Los objetivos de la tesis fueron mejorar las identificaciones gránulos de almidón mediante el uso de un sistema automatizado, evaluar las muestras modernas como método de inferencia a través del estudio de fitolitos de suelos y plantas relacionadas con áreas húmedas en África, y la reconstrucción del paleoambiente/paleopaisaje de dos yacimientos de la garganta de Olduvai para analizar su influencia en el comportamiento de los homínidos. Para mejorar la identificación de gránulos de almidón se utilizó un sistema de análisis de imagen que midió 123 caracteres ópticos y morfológicos de aproximadamente 5000 gránulos de 20 especies de plantas comestibles del este de África. Los datos obtenidos se analizaron mediante un sistema estadístico de aprendizaje automático (Random Forest). Los resultados muestran que, a pesar de que el sistema desarrollado no es perfecto, es más eficaz que las identificaciones de visu, cuyo acierto medio fue del 25% frente al 53% del sistema automatizado. Se observó que el sistema es sensible al número de especies analizadas y, en un menor grado, al número de caracteres utilizados, por lo que consideramos que reducir las potenciales especies a identificar es crucial para obtener identificaciones precisas y para ello es necesario combinar diversos análisis. De no ser así la identificación automática de gránulos de almidón no será lo suficientemente precisa para ser utilizada en arqueología. Para mejorar el conocimiento de los conjuntos de fitolitos que permitan obtener inferencias precisas de las muestras fósiles se analizaron los fitolitos de 22 suelos modernos provenientes del entorno de Olduvai y de 14 especies relacionadas con ambientes húmedos. Los resultados del análisis de suelos muestran que los fitolitos reflejan, parcialmente, la estructura vegetación, pero no el tipo de vegetación que los produjo y que las diferencias entre diferentes tipos de formaciones son sutiles. Sin embargo, se observó que dichas diferencias son apreciables cuando se aplican herramientas estadísticas. Los resultados muestran que los fitolitos reflejan mejor que el polen la estructura de la vegetación, pero, como era esperable, el polen traza mejor la 5 diversidad de especies. El estudio preliminar de 14 especies relacionadas con los ambientes húmedos mostró que los helechos (Pteridophyta) producen fitolitos capaces de marcar su presencia de manera inequívoca en el ambiente. Por otro lado, las otras especies no mostraron fitolitos característicos que permitiesen mejorar la identificación de alguna de las especies analizadas. El análisis de fitolitos de 27 trincheras del paleopaisaje que incluye los yacimientos del “complejo Zinj” (FLK Zinj, AMK, PTK y DS) reveló una gran área cubierta por vegetación boscosa en los tiempos de deposición de la toba IC (1,832 millones de años). La presencia de palmeras en el área viene a refutar hipótesis previas que sugerían la presencia del cauce de un río de a 50-200 metros al sureste de FLK Zinj. La aparición de fitolitos de helechos indica presencia de hábitats sombríos y húmedos, lo que sugiere presencia de agua dulce (ríos o surgencias) que pudo atraer a homínidos y otros animales. La disponibilidad de agua dulce y los restos animales y útiles líticos encontrados en los yacimientos llevan a pensar que los homínidos no sólo usaron estos lugares como refugio, si no como áreas de procesado de restos animales y otras actividades vinculadas con un modelo de forrajeo central. El análisis de los fitolitos de 24 muestras del yacimiento BK, muestreados en sentido vertical al de la estratigrafía, permitieron reconstruir la evolución de la paleovegetación que acompañó a la evolución de un sistema fluvial. Los resultados muestran una vegetación que encaja bien con la dinámica fluvial que formó BK. En las zonas donde aparece la vegetación esta estaba dominada por plantas leñosas y se mantuvo estable durante todo el marco de tiempo analizado. Por otro lado, la presencia de muestras estériles coincide con los lugares y momentos donde los suelos se renovaban continuamente debido al arrastre fluvial. El análisis de fitolitos de 42 útiles líticos del yacimiento FLK West, junto con los paleosuelos que sirvieron de control, mostró que algunas de las piezas recuperadas pudieron ser utilizadas para procesar palmeras y tejidos duros de dicotiledóneas. Los análisis de gránulos almidón efectuados en esas mismas piezas no produjeron resultados concluyentes. Un análisis de arqueología experimental demostró que la distribución de 6 los fitolitos no se ve afectada por el azar durante la deposición de las piezas líticas o por el manejo de las mismas en el laboratorio. En conclusión, los resultados de esta tesis muestran que el análisis
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  • Poaceae (Gramineae)

    Poaceae (Gramineae)

    FLORA OF SINGAPORE, Vol. 7: 219–501 (2019) POACEAE (GRAMINEAE) J.F. Veldkamp1, H. Duistermaat1, K.M. Wong2 & D.J. Middleton2 Barnhart, Bull. Torrey Bot. Club 22 (1895) 22; Monod de Froideville in Backer & Bakhuizen van den Brink, Fl. Java (Spermatoph.) 3 (1968) 495; Watson & Dallwitz, Grass Gen. World (1992) 1; Chen et al., Fl. China 22 (2006) 1; Chen et al., Fl. China Illus. 22 (2007) 1; Kellogg in Kubitzki (ed.), Fam. Gen. Vasc. Pl. 13 (2015) 3. Type: Poa L. Gramineae Juss., Gen. Pl. (1789) 28, nom. alt.; Ridley, Fl. Malay Penins. 5 (1925) 186; Schmid, Agron. Trop. (Nogent-sur-Marne) 13 (1958) 7; Bor, Grasses Burma, Ceylon, India & Pakistan (1960) 1; Gilliland, Rev. Fl. Malaya 3 (1971) 1; Clayton & Renvoize, Gen. Gram. [Kew Bull., Addit. Ser. 13] (1986) 1. Type: Poa L. Annual or perennial plants. Stems herbaceous or (in most bamboos and rarely in other grasses) heavily lignified and hard-tissued (‘woody’), prostrate (stoloniferous: creeping and rooting at nodes), or with distinct subterranean portions (rhizomes: mostly sympodial, in some bamboos monopodial) and above-ground portions (culms: that may be somewhat erect or, in some bamboos, scrambling, clambering or even twining); terete, hollow or solid, with transverse septs; branching at culm base intra- or extravaginal, producing equivalent culms, that at more distal nodes of the culm irregular and simple (especially in stoloniferous taxa) or (in most bamboos) more consistently present at most nodes and typically with few to many orders of branching (‘complex branching’); nodes when branch-bearing with the axillary bud initially enclosed within a prophyll (its bract-like first leaf whose back is addorsed to the main axis), bearing verticils of short roots (in stoloniferous grasses and the culm bases of some bamboos) or not, such roots rarely indurated (‘root-thorns’) in some bamboos.