southern south america

Submitted: August 28th, 2018 – Accepted: March 12th, 2018 – Published online: March 16th, 2018

To link and cite this article: doi: 10.5710/AMGH.12.03.2018.3152

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EMERGENCE OF GRASSY HABITATS DURING THE GREENHOUSE–

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ICEHOUSE SYSTEMS TRANSITION IN THE MIDDLE EOCENE OF

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SOUTHERN SOUTH AMERICA.

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APARICIÓN DE LOS ECOSISTEMAS DOMINADOS POR GRAMÍNEAS

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DURANTE LA TRANSICIÓN CLIMÁTICA GREENHOUSE–ICEHOUSE

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EN EL EOCENO MEDIO DEL SUDESTE DE AMÉRICA DEL SUR

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ALEJANDRO F. ZUCOL1, J. MARCELO KRAUSE2, 3, MARIANA BREA1, M. SOL

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RAIGEMBORN4 AND SERGIO D. MATHEOS4

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1

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Tecnología a la Producción, CICYTTP–CONICET, Dr. Materi y España S/N, E3105BWA

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Diamante, Entre Ríos, Argentina. [email protected]; [email protected]

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2

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Trelew, Chubut, Argentina. [email protected]

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3

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Provincial 1 s/n, (9000) Comodoro Rivadavia, Chubut, Argentina.

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4

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B1904DPK La Plata, Buenos Aires, Argentina. [email protected],

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[email protected]

Laboratorio de Paleobotánica, Centro de Investigaciones Científicas y Transferencia de

CONICET-Museo Paleontológico Egidio Feruglio, Avenida Fontana 140, U9100GYO

Departamento de Geología, Universidad Nacional de la Patagonia San Juan Bosco, Ruta

Centro de Investigaciones Geológicas (CONICET-UNLP), Diagonal 113 # 275,

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Total number of pages (text + references): 66

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Figures: 10; Supplementary material (Figures: 2, Tables: 2; and supplementary references)

1

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Proposed head: ZUCOL ET AL.: EOCENE PHYTOLITHS FROM PATAGONIA

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Corresponding author: Alejandro F. Zucol

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Abstract. Phytolith assemblages recovered from the middle Eocene Koluel-Kaike

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Formation in Central Patagonia, southern South America, confirm the presence of

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herbaceous components, mainly related to mixed ecosystems or savannahs, and

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subordinate forest communities. This new palaeobotanical information reveals a clear

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transition between previously described woodland-dominated communities from the

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early Eocene Las Flores Formation, and grass-rich ecosystems from the mid-Eocene

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Sarmiento Formation. The early appearance of grass-dominated ecosystems in southern

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South America, would be related to the worldwide greenhouse to icehouse systems

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transition initiated during the mid-Palaeogene.

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Key words. Phytoliths, palaeosols, Koluel-Kaike Formation, palaeoclimate, EECO,

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Central Patagonia.

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Resumen. APARICIÓN DE LOS ECOSISTEMAS DOMINADOS POR GRAMÍNEAS

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DURANTE LA TRANSICIÓN CLIMÁTICA GREENHOUSE–ICEHOUSE EN EL

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EOCENO MEDIO DE PATAGONIA CENTRAL. Asociaciones fitolíticas provenientes de

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la Formación Koluel-Kaike (Eoceno medio) en Patagonia Central, sur de América del Sur,

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confirman la presencia de componentes herbáceos, principalmente relacionados a

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ecosistemas mixto o sabanas, y comunidades arbóreas subordinadas. Esta nueva

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información paleobotánica revela una clara transición entre las comunidades previamente

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descriptas en las Formación Las Flores (Eoceno inferior), dominada por componentes

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leñosos, y la sección inferior de la Formación Sarmiento (Eoceno medio), caracterizada por

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una mayor abundancia de gramíneas. La aparición temprana de ecosistemas dominados por

2

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gramíneas en el sur de América del Sur, estaría relacionada a la transición climática global,

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desde sistema Greenhouse a Icehouse, iniciada a partir del Paleógeno medio.

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Palabras clave. Fitolitos, paleosuelos, Formación Koluel-Kaike, paleoclima, EECO,

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Patagonia Central.

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PREVIOUS studies on phytolith content from early Palaeogene successions in Central

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Patagonia have revealed the presence of forested communities during the early Eocene

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Las Flores Formation (Brea et al., 2008; Raigemborn et al., 2009), and grass-rich

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communities in the middle Eocene Gran Barranca Member (lower Sarmiento

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Formation, Mazzoni, 1979; Zucol et al., 1999, 2004; Andreis, 2008; Zucol et al.,

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2010a). Vegetation shift occurred in parallel sedimentary, environments and climatic

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conditions changes. For instance, lacustrine and fluvial sedimentation, under tropical

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conditions, occurred during the deposition of the Las Flores Formation (Raigemborn et

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al., 2009, 2010, 2014; Woodburne et al., 2014); and dominant loessic deposition, under

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semiarid-arid conditions, occurred during the deposition of the Gran Barranca Member,

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Sarmiento Formation (Mazzoni, 1979; Spalletti and Mazzoni, 1979; Bellosi et al., 2010;

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Bellosi and González, 2010; Bellosi and Krause, 2014).

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In order to better understand the palaeofloristical evolution in Central Patagonia, we

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document phytolith assemblages from the Koluel-Kaike Formation, a unit bracketed

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between the underlying Las Flores and the overlying Sarmiento formations. This new

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information fills the gap in the palaeofloristic record in the San Jorge Basin, Southern

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South America, between the mid-Ypresian woodland-dominated Las Flores

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communities, and the late Lutetian, grass-rich Sarmiento ones; and provides key

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knowledge for understanding the causes of the origin of grasslands in South America, a

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factor pertinent to regional palaeoecology and the evolution of mammals and feeding

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strategies (e.g., Strömberg, et al. 2013; Dunn et al., 2015). Our aims are to: 1) analyse

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the floristic composition and changes occurred during the deposition of the Koluel-

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Kaike Formation, and 2) consider these results in the context of the different hypotheses

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regarding the emergence of grasslands in South America and their relationship with the

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global palaeoclimate scenery.

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Figure 1

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GEOLOGICAL, SEDIMENTOLOGICAL AND

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PALAEONTOLOGICAL SETTING

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The Koluel-Kaike Formation is a continental unit cropping out in the San Jorge

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Basin, Central Patagonia, Argentina (Fig. 1.2). In Gran Barranca area the Koluel-Kaike

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Formation overlies, in probable paraconformity relationship, to the Las Flores

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Formation (Krause et al., 2017), and underlies in transition to the Gran Barranca

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Member (lower Sarmiento Formation) (Bellosi et al., 2010; Krause et al., 2010, 2017;

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Raigemborn et al., 2010). It composes of moderate to highly-silicified, silt-sized,

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pyroclastic deposits, bentonites, and tuffs, mostly modified to palaeosols (Krause and

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Bellosi, 2006; Krause et al., 2010; Raigemborn et al., 2010; Raigemborn et al., 2014).

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The fossils record of this unit currently restrict to silicified wood, referred to

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Taxaceoxylon katuatenkun Brea, Bellosi et Krause (Brea et al., 2009), and trace fossils

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of cicadas (Feoichnus challa Krause et al., 2008a). The temporal span of the middle Las

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Flores to uppermost Koluel-Kaike succession in the Gran Barranca area was

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constrained, on the base of magnetostratigraphic and geochronological data, to be ca.

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51.4–42.2 Ma (mid Ypresian–late Lutetian) in age, with the Koluel-Kaike being 46.7–

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42.2 Ma (Lutetian) in age (Krause et al., 2017) (Fig. 1.1). 4

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For this study, two localities were selected, Las Flores and Cañadón Lobo. The Las

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Flores locality (LF) (S 45° 43' 30''; W 68° 37' 22'') locates in the eastern edge of Gran

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Barranca, a classical Cenozoic land-mammal fossil locality (e.g., Ameghino, 1906;

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Simpson, 1948; Marshall et al., 1983; Madden et al., 2010), ~40 km to the east-southeast

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of Sarmiento city, south-central Chubut Province (Fig. 1.2). At LF the studied section is

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~42 m-thick (Fig. 2), and characterizes by tephric loessic beds, small alluvial channels,

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and shallow ponds (Krause et al., 2008b, 2010; Raigemborn et al., 2010, 2014). The

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sedimentation in the Koluel-Kaike Formation would have developed under different

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climates, with a clear temporal evolution from humid to subhumid-semiarid conditions,

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as evidenced by palaeosols and clay-mineral information (Krause et al., 2010; Krause,

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2012a, b; Raigemborn et al., 2014). Climofunctions applied on palaeo-Ultisols from the

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lower section, suggest values of Mean Annual Precipitation (MAP) from 1200 to 1300

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mm/yr, and Mean Annual Temperature (MAT) ~15° C (Krause et al., 2010), consistent

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with wet and megathermal conditions. Upward the section, climofunctions applied on

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weakly-developed palaeosols (e.g., palaeo-Andisols) give values of MAP ~1000 mm/yr

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and MAT ~12° C) (Krause et al., 2010), consistent with megathermal–mesic conditions.

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The change in clay mineralogy throughout the unit, from kaolinite-dominated

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assemblages, in the lower section, to smectite-dominated associations in the middle–

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upper ones, also indicates a climatic trend from humid to subhumid conditions, along

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with pulses of relative stronger seasonality or less intense precipitations periods (Krause

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et al., 2010; Raigemborn et al., 2014).

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The Cañadón Lobo (CL) locality (S 46° 58' 24"; W 66° 50' 13") places in northern

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Santa Cruz Province, ~15 km to the northwest of Punta Casamayor, in south border of the

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San Jorge Basin (Fig. 1.2). At CL the base of the Koluel-Kaike Formation is not

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exposed. The unit underlies to whitish, massive and tabular beds of the Sarmiento

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Formation assigned to the Gran Barranca Member, and bearing sparse, fragmentary

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vertebrate remains from the Casamayoran South American Land Mammal Age

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(SALMA), fossil dung-beetle brood balls (Coprinisphaera) and Feoichnus specimens

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(Pérez et al., 2012). The contact between the Koluel-Kaike and the Sarmiento

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formations looks conformable (Pérez et al., 2012), although the absence of around 10–

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15 m of whitish beds bearing weakly-developed palaeosols recognized in the upper

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Koluel-Kaike section at LF (Krause et al., 2010), suggests the presence of a

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paraconformity. The studied section is ca. 10 m-thick and mainly composes of massive,

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silt-sized, pyroclastic deposits contained within tabular and lenticular beds, with

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minority tabular, massive beds of mudstones (Figs. 2, 3.1–2). Lenticular beds display an

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erosive base, evidenced by the presence of muddy intraclasts; while tabular beds display

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sharp bases. Both types of beds lack in recognizable primary, sedimentary structures.

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Pedogenetic features, among them, iron-rich, reticulate mottles, iron nodules,

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slickensides, and rhizoliths are common within the tabular pyroclastic and muddy beds.

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Iron nodules are also present within the lenticular beds. Based on these features, Krause

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(2010) interpreted a palaeoenvironment characterized by small fluvial channels, cutting

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across pyroclastic and muddy floodplains (Fig. 3.2). The presence of iron nodules

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within the lenticular channel body indicates changes in water content in the original

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deposit, suggesting a probable ephemeral nature of the channel. On the other side,

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tabular beds composed of very fine sediments and bearing rhizoliths, slickensides, and

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iron mottling and concretions, support the presence of pedogenetically modified,

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floodplain deposits (Krause, 2010) (Fig. 3.3–4). Palaeosols at CL (Fig. 3.2–6)

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displaying reddish, reticulate mottles are strongly similar in morphology to those

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included in the lower Koluel-Kaike Formation at LF (e.g., Kápenk pedotype sensu

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Krause et al., 2010), which characterize ironstones levels (Krause et al., 2010) (Fig.

6

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3.5). Since ironstones of the Koluel-Kaike Formation occur typically in the lower

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section at Las Flores, and are absent in adjacent units (Krause et al., 2010), we correlate

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the CL section with the lower Koluel-Kaike Formation at LF.

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Figure 2

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THE EARLY PALAEOGENE OF CENTRAL PATAGONIA:

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PHYTOLITHS AND OTHERS PALAEOBOTANICAL RECORDS

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The early Palaeocene–middle Eocene span of time in Gran Barranca and surrounding

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areas is represented by five formations (Fig. 1.1): Salamanca (estuarine; Danian)

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(Martínez, 1992; Clyde et al., 2014; Comer et al., 2015), Peñas Coloradas (fluvial; latest

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Danian) (Raigemborn et al., 2010; Clyde et al., 2014), Las Flores (fluvial, lacustrine;

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Ypresian) (Raigemborn et al., 2010; Woodburne et al., 2014; Kohn et al., 2015; Krause et

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al., 2017), Koluel-Kaike (loessic, alluvial; Lutetian) (Krause et al., 2010, 2017;

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Raigemborn et al., 2010), and lower Sarmiento (Cañadón Vaca and Gran Barranca

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Members) (loessic, fluvial; late Lutetian–Bartonian) (Bellosi et al., 2010; Ré et al., 2010a,

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b; Dunn et al., 2013; Bellosi and Krause, 2014).

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Phytolith analysis from the Salamanca Formation, proceeding from shallow estuarine and

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lagoon facies (Zucol et al., 2008) suggesting warm subtropical climate. This interpretation

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agrees with others palaeobotanical records from the Salamanca Formation, such pollen, leaf

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compression, permineralized woods and fruits (Berry, 1937; Romero, 1968; Archangelsky,

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1973, 1976; Archangelsky and Zamaloa, 1986; Somoza et al., 1995; Brea et al., 2005a, b,

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2008; Matheos et al., 2005; Iglesias et al., 2007, Jud et al., 2017; Ruiz et al., 2017, among

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others).

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Phytoliths have not been described for the Peñas Coloradas Formation, but

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complementary palynomorphs and permineralized woods remains indicate the presence of

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mixed subtropical to tropical and humid forests (Romero, 1973; Lema et al., 1999; Ruiz et

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al., 1999; Brea and Zucol, 2006; Raigemborn et al., 2009).

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The phytolith assemblages of the Las Flores Formation exhibits high abundance of dicot

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and palm phytoliths, in association with scarce grasses (Poaceae), sedges, and

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podostemoid elements, corresponding to subtropical to tropical lowland forests (Brea et

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al., 2008; Raigemborn et al., 2009). Woody and palm indicators constitute 12.27−39.61%

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and 8.05−37.19% respectively, with dicots (12.07–34.16%), diagnostic herbs (0–0.92%),

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non-diagnostic herb/bamboos (8.06–23.09%) and humid herbs (0−4.13%) as understory

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elements (Brea et al., 2008; Raigemborn et al., 2009).

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The phytolith content from the Gran Barranca Member (lower Sarmiento Formation) at

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Gran Barranca section reveals abundance of palms and up to 15% of herbaceous elements,

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and a lower proportion of woody plant elements (Zucol et al., 2010a; Strömberg et al.,

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2013; Dunn et al., 2015; Kohn et al., 2015). Zucol et al. (2010a), interpreted this

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association as the record of savannah palaeocommunities, composed of palms

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(9.19−38.61%), diagnostic herbs (1.40–18.48%), non-diagnostic herb/bamboo

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(10.90−35.28%), humid herbs (0−12.38%), dicots (3.57–19.33%), and woody elements

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(1.39−15.20%), and with a clear increase trend of megathermal components (from C4

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grass) upward the unit. Thus, the lower Gran Barranca Member characterizes by the

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highest abundances of palms (30.19−38.61%), which then decrease into the upper levels

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to the 12.44−22.50% range. In general, the woody elements have low abundances

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(1.39−9.74%) in the whole section, with the exception of one sample recorded in the

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middle-upper section (15.20% in the MMZ9504.0 sample, Zucol et al., 2010a).

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Diagnostic types of grass herbs and non-diagnostic herb/bamboo types respectively 8

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increase their abundance from 3.77–6.73% and 10.90–20.00%, in the lower section, to

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6.78−18.48% and 14.13–28.57%, in the upper one. This increase relates to the high

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abundance of panicoid phytoliths. Finally, upward the Gran Barranca Member, the

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floristic composition recordes an increase in humid herbs, which generally present in

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low percentages (0−3.43%) throughout the section, rising up to 12.38% in the upper

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section (sample MMZ9507.5; Zucol et al., 2010a).

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However, in spite of similar values of percentage among involved morphotypes

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obtained for the Gran Barranca Member at type locality obtained by Strömberg et al.

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(2013), Dunn et al. (2015), and Kohn et al. (2015), these authors do not share the

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interpretation of a savannah environment. For instance Strömberg et al. (2013), on the

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base of the abundance of palms, suggested an ecological scenery characterized by a

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closed–humid subtropical forest, an interpretation subsequently modified in Dunn et al.

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(2015). These authors, applying the leaf area index (LAI) proxy, concluded that the

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declination of the rLAI values, from the middle Eocene to the early Oligocene in the

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San Jorge Basin, indicating an opening of the landscape from dense vegetation through

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progressively more open vegetation, that is a nearly grass-free scrubland, an

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interpretation kept by Kohn et al. (2015).

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Figure 3

MATERIALS AND METHODS Twenty-four sedimentary samples (Fig. 2) were obtained from the Koluel-Kaike

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Formation for phytolith analysis: 19 from the LF locality (LF539–557), and 5 from the CL

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locality (CL1–CL5). Nine of these samples (LF541, LF545–546, LF548–549, LF552–555)

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provided an insufficient phytolith content to obtain a minimal representative sample and

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thus inadequate for their statistical analysis.

9

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Each sample consisted of 20 grams of sediment, which was processed following a

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protocol employing various chemical agents to remove soluble salts, organic matter and

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carbonates (Zucol et al., 2010b). Samples were washed and soluble salts dissolved with

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distilled water; carbonate material was removed with dilute HCl; organic material was

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removed by adding H2O2 followed by deflocculation by Sodium hexametaphosphate. The

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fractionation was quantified through sieving (coarse fraction) and sedimentation (fine

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fractions). Four fractions were obtained according to particle diameter: coarse (>250 µm),

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medium (5–53 µm and 53–250 µm) and fine (5 μm: “trapeziform/parallele pipedal”

Bilobates

Poaceae (GSSC)/PACMADclade/Danthonioideae? Bambusoideae? (C3)/PACMAD general/Polylobates

Bilobate with crenate central part (B/cr), (21). Bilobate with nodular/spiny shank (NB) (21). Trilobate (Tri) 3 ± regular lobes arranged in a line (19). Irregular 25. Polylobate complex bilobate 2 lobes with irregular lobe(s) on the shank (21). Regular complex bilobate > 3 ± regular lobes linearly arranged (21).

Polylobates (modified bilobates with one to several supplementary lobes on the shank), size: 18–25 μm

Polybates / Irregular, complex

2

Poaceae /PACMADclade/Danthonioideae? Agr (h) Bambusoideae? (C3)/PACMAD general

ID

Polylobate

3

Poaceae /PACMADclade/Chloridoideae (C4). Aristida-type

Agr (h)

ID

Thin, longshanked bilobate

4

Poaceae /PACMADclade/Other PACMAD (C3/C4)

Agr (h)

ID

5

Poaceae /PACMADclade/Other PACMAD (C3/C4)

Agr (h)

ID

6

Poaceae / PACMADclade/ Danthonioideae? Bambusoideae? (C3)/ PACMAD general

Agr (h)

ID

Poaceae (GSSC)/ PACMADclade/ Danthonioideae? Bambusoideae? (C3)/ PACMAD general/ Bilobates

7

Poaceae / basal P+BE clade/ Bambusoideae

Agr (hb)

ID

Poaceae (GSSC)/ Closedhabitat grasses (basal Poaceae+ BE clade)/ CH TOT/ Chusquea-type bilobate

8

Poaceae / Pooideae/ diagnostic/ Stipoideae

Agr (h)

ID

9

Poaceae / PACMADclade / Panicoideae (C3/C4)

Agr (h)

ID

10

Poaceae / PACMADclade / Other PACMAD Agr (h) (C3/C4)

ID

11

Poaceae / basal Poaceae+ BE clade/ Bambusoideae

ID

12

Poaceae / PACMADclade/ Chloridoideae (C4?)

Agr (h)

ID

13

Poaceae / PACMADclade/ Chloridoideae

Agr (h)

ID

Poaceae (GSSC)/PACMADclade/Other PACMAD (C3/C4)/Near-panicoid bilobate Poaceae (GSSC)/ PACMADclade/ Other PACMAD (C3/C4)/ Inverted bilobate/cross

Poaceae / basal P+BE clade

Poaceae (GSSC)/ Pooideae/ diagnostic/ POOID-D/ Stipatype bilobate.

Carinate

13. Carinate

Cross Thick Shank (QTS) Four lobes, thick shank (21). Cross Poaceae (GSSC)/ PACMADThin Shank (QIS) Four lobes, clade / Panicoideae (C3/C4)/ thin shank (21). Cross butterfly 26. Quadralobate/cross Various crosses. shape. Four lobes, transitional to short bilobate, margins crenate/wavy. Quadralobate/cr oss

Saddle (= Battle axe) Sa (3, and equivalent name reference therein).

Poaceae (GSSC)/ PACMADclade/ Chloridoideae (C4?)/ CHLOR/ True saddle.

Poaceae (GSSC)/ PACMADclade/ Chloridoideae (C4?)/ Pseudo-saddle (almost true saddle; end view not symmetric). Poaceae (GSSC)/ Closedhabitat grasses (basal Poaceae+ BE clade)/ CH TOT/ Collapsed saddle.

Saddles

Saddles ( Sa )

27. Saddle

Oxytenanthera saddle (15)

ID

30. Saddle ovate

17

Poaceae / PACMADclade/ Arundinoideae

Agr (h)

ID

31. Saddle plateau

18

Poaceae / PACMADclade/ Chloridoideae

Agr (h)

ID

32. Saddles quat

19

Poaceae /P+BE clade and PACMAD-clade

Truncated conical

ID

Mt21

21 22

23

24

Poaceae /P+BE clade and PACMAD-clade Poaceae /P+BE clade and PACMAD-clade Poaceae (GSSC)/P+BE clade and PACMADclade

MT20 Cónicos

Poaceae (GSSC)/ Pooideae/ Truncated cone (= Spool-shaped) Ct (3, non-diagnostic/ POOID-ND/ and equivalent name reference therein). Conical and keeled rondel.

Rondel Short cylinders or 42. Tower truncated/ beveled cones (22).

Truncated cones / Towers

Agr (h)

ID

43. Tower bevelled

Agr (h)

ID

44. Tower horned

Agr (h)

ID

45. Tower wide

Poaceae / PACMADclade/ Danthonioideae? Bambusoideae? (C3)/ PACMAD genera

Agr (hb)

ID

Poaceae / basal P+BE clade/ Bambusoideae

Agr (hb)

Rondell Mt30

ID

Saddles

Complex collapsed saddle

ID

20

Saddle

29. Saddle long, wavy side

Agr (h)

Agr (h)

Saddles (bodies with two opposite convex edges and two opposite concave edges in top view, “battles axes with double edges”), size: L=8–12 μm, h≤5 μm: “tabular” or h>5 μm: “trapeziform”

Collapsed saddle

Agr (hb)

Poaceae / basal P+BE clade Poaceae / PACMADclade/ Chloridoideae Arundinoideae

Crosses

28. Saddle long

ID

16

Cross body (all variants)

Poaceae (GSSC)/ PACMADclade / Other PACMAD (C3/C4)/ Near-panicoid cross.

Agr (hb)

15

Crosses (equidimensional bodies, 4- or 3-lobed), size: L=8–12 μm, h≤5 μm: “tabular” or h>5 μm: “trapeziform”

Poaceae (GSSC)/ Closedhabitat grasses (basal Poaceae+ BE clade)/ CH TOT/ Chusquoid body/cross with irregular/spiked top.

Mt34

14

12

Thin, longshanked bilobate

Other bilobate

Agr (hb)

11

Chusquea-type rondel

Poaceae (GSSC)/ PACMADclade/ Danthonioideae? Rondel (= Rondel) Rn (3, and equivalent Bambusoideae? (C3)/ name reference therein). PACMAD general/ Simple lobate.

Poaceae (GSSC)/ Closedhabitat grasses (basal Poaceae+ BE clade)/ CH TOT/ Chusquea-type rondel.

24. Ovate

Rondels (conical/cylindrical bodies, with Rondel/trapezoi Spooled/ round/oval base, size: d horned towers Ø 8–12 μm (except Ro8) and h=5–12 μm (except Ro-7))

Tall rondel

Rondels

Mt31

Square or rectangle (sensu 4)

25

Poaceae / Pooideae

Agr (h)

ID

Square

26

Poaceae / Pooideae/ Crenate “trapeziform polylobate”.

Agr (h)

ID

27

Poaceae / Pooideae/ Trapeziform short cells Agr (h) “pyramidal”.

ID

28

Poaceae / basal P+BE clade/ Ehrhartoideae

ID

29

Poaceae / basal Poaceae+ BE clade. Regular chusquoid body.

Agr (hb)

ID

Poaceae (GSSC)/ Closedhabitat grasses (basal Poaceae+ BE clade)/ CH TOT/ Regular chusquoid body.

30

Poaceae / basal Poaceae+ BE clade. Crescentic keeled rondel.

Agr (hb)

ID

Poaceae (GSSC)/ Closedhabitat grasses (basal Poaceae+ BE clade)/ CH TOT/ Crescentic keeled rondel.

Poaceae (GSSC)/ Pooideae/ diagnostic/ POOID-D/ Crenate (“trapeziform polylobate”).

Trapezoid crenate

31

32

33

34

Agr (h)

Poaceae / PACMADclade/ Danthonioideae? Bambusoideae? (C3)/ Agr (hb) Very tall with prominently faceted tops. Poaceae / PACMADclade / Panicoideae Agr (h) (C3/C4) Poaceae / PACMADclade/ Chloridoideae Agr (h) (C4?) Grass morphotypes no intrafamily diagnostic

Poaceae

Agr (hb)

Other “Pooide Class” (21).

46. Trapezoid crenate

Trapeziform bodies (sixTrapeziform sided), size: L=8–10 polybate μm.

Poaceae (GSSC)/ Pooideae/ non-diagnostic/ POOID-ND/ Trapeziform short cells (“pyramidal”). Scooped bilobate

Scooped bilobate

Other phytolith types (nonGSSCs)

Other phytolith types (non-GSSCs)

Poaceae (GSSC)/ PACMADclade/ Danthonioideae? Bambusoideae? (C3)/ PACMAD general/ Very tall with prominently faceted tops.

ID

Triangle cross (Tr/cr) 3 lobes arranged in a triangular shape.

ID

ID

14. Conical

D

Point-shaped

Mt12

Point-shaped (= Point-shaped, unciform MT26 Aculeolithum hair) Ps (3, and equivalent name sp.1 reference therein). 1–10 Ps1–Ps10 s.l.

Poaceae (GSSC)/ Unknown Poaceae/ NDG/ Short unciform prickle ("scutiform opal").

Acicular bodies Aci-1 Smooth (50–75 μm); Aci-2 Smooth, often infilled with black material (40–50 μm).

Pri/ Prickle, scutiform to triangular, oval to circular base with short pointed tips.

Point-shaped

35

Poaceae

Agr (hb)

D

36

Poaceae

Agr (hb)

D

Hks/ Unciform hair cell. Spo/ Sharp pointed hairs with no base. Mac/ Fairly long, unicellular hair. Mic/ Shorter than macrohairs,commonly two celled.

37

Poaceae

Agr (hb)

D

38

Poaceae

Agr (hb)

D

39

Poaceae

Agr (hb)

D

Mt28

Fan-shaped

Fan-shaped

40

Poaceae

Agr (hb)

D

Mt29

Polyhedral bulliforms

Polyhedrons MT29 Poliédricos

41

Poaceae

Agr (hb)

D

42

Poaceae

Agr (hb)

D

43

Poaceae

Agr (hb)

D

Blo/pol/ss / Polyhedral blocky bodies, straight margins and psilate surface.

44

Poaceae

Agr (hb)

D

Blo/pol/fac / Polyhedral blocky bodies, straight margins and faceted surface.

45

Poaceae

Agr (hb)

D

Mt24

Prismatic elongated with serrate contour

Prismatic serrate

Elongate (= Parallelepipedal or Prismatic) MT23 Macroprismatolithu El (3, and equivalent name reference therein). 64 El10. m sp.1

Pl/IBIC / Rectangle with discs evenly spaced along central axis.

46

Poaceae

Agr (hb)

D

Mt25

Prismatic elongated with denticulate contour

Prismatic denticulate

Grasses and other Elongate (= Parallelepipedal or Prismatic) MT24 monocotyledons, conifers / Macroprismatolithu El (3, and equivalent name reference NDG / Echinate, and therein). 57 El03. m denticulathum dendritic long cell.

El/sm/Sm / Elongate, straight ends/margins and psilate surface.

47

Poaceae

Agr (hb)

D

Mt27

Prismatic elongated with undulate contour

48

Poaceae

Agr (hb)

D

49

Poaceae

Agr (hb)

D

50

Poaceae

Agr (hb)

D

51

Poaceae

Agr (hb)

D

MT27 Flabelolithum sp1

Poaceae (GSSC)/ Unknown Poaceae/ NDG/ (Cunciform) bulliform cell.

Bul/cun/wv / Wedge-shape bulliform with wavy margins.

Parallepipedal bulliforms (sensu 45)

Poaceae (GSSC)/ Unknown Poaceae/ NDG/ (Cunciform) bulliform cell.

Bul/par/ss/str / Parallelepipedal bulliform, straight margins and psilate surface.

Blocky bodies / Blo-5 Parallelepipedal with constricted sides, approximately cuneiform (30–100 μm).

Bul/par/ss/wv / Parallelepipedal bulliform, wavy margins and psilate surface.

Blocky bodies / Blo-8 Parallelepipedal, smooth, crenate margins, often infilled with black material (40–100 μm).

Other parallelepipedal bulliform types

12. Bulliform

Bul/par/fac / Parallelepipedal bulliform, straight margins and faceted surface.

Elongate (= Parallelepipedal or Prismatic) Grasses and other El (3, and equivalent name reference monocotyledons, conifers / therein). 62 El08 El. NDG / Elongate sinuous.

El/sm/Sf / Elongate, straight ends/margins and faceted surface.

El/sm/ss / Elongate, straight ends/margins and granulate surface. El/sin/sw / Elongate, sinuate ends/margins and warty surface. El/sin/ss / Elongate, sinuate ends/margins and psilate surface. El/sin/sst / Elongate, sinuate ends/margins and granulate surface.

Other prismatic elongated

Blocky bodies / Blo-4 Perfectly cuneiform (bulliform cell) (30–100 μm).

Fan-shaped (= Cuneiform ?) Fs (3, and equivalent name reference therein).

Elongate bodies / El-3 Tabular/parallelepiped al bodies with smooth, sinuate or echinate margins (50–200 μm).

Bulliforms

52

Poaceae

Agr (hb)

D

53

Poaceae

Agr (hb)

D

54

Poaceae

Agr (hb)

D

55

Poaceae

Agr (hb)

D

56

Poaceae

Agr (hb)

D

Poaceae

Agr (hb)

D

57

El/sin/sfac / Elongate, sinuate ends/margins and faceted surface. Other prismatic elongated

El/wv/ss / Elongate, straight ends, wavy margins, and psilate surface. El/sp/ss / Elongate, straight ends, spiny margins and psilate surface. El/conc/ss / Elongate, concave ends, straight margins and psilate surface. El/conc/sfac / Elongate, concave ends, straight margins and faceted surface. El/tap/ss / Elongate, tapering ends, straight margins, psilate surface.

Other Grass morphotypes no intrafamily diagnostic (NID) 58

Poaceae (GSSC) / Unknown Poaceae / OTHG / Various GSSC. Grasses and other monocotyledons, conifers / NDG / Acicular hair cell.

Various GSSC Poaceae

Agr (hb)

D

59

Acicular hair cell

60

Poaceae

Agr (hb)

D

Circular, conical or inflated with domed to flat center

61 62 63 64 65 66

Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae

Agr (hb) Agr (hb) Agr (hb) Agr (hb) Agr (hb) Agr (hb)

D D D D D D

67

Poaceae

Agr (hb)

D

68

Poaceae

Agr (hb)

D

Elongate

20. Elongate orthogonal section.

69

Poaceae

Agr (hb)

D

Epidermal crenate

21. Epidermal crenate.

70

Poaceae

Agr (hb)

D

Epidermal polygonal

22. Epidermal polygonal.

71

Poaceae

Agr (hb)

D

Globular psilate

23. Globular psilate.

72

Poaceae

Agr (hb)

D

Scutiform

33. Scutiform.

73

Poaceae

Agr (hb)

D

Stomata/Hair

34. Stomata/Hair.

74 75 76 77 78 79 80

Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae

Agr (hb) Agr (hb) Agr (hb) Agr (hb) Agr (hb) Agr (hb) Agr (hb)

D D D D D D D

Tabular

35. Tabular crenate. 36. Tabular dendriform. 37. Tabular elongate. 38. Tabular facetate. 39. Tabular scrobiculate. 40. Tabular sinuate. 41. Tabular thin.

81

Poaceae

Agr (hb)

D

Vessel laminate

47. Vessel laminate.

Unciform, smooth

82

Poaceae

Agr (hb)

D

Poaceae

Agr (hb)

D

Pap / Circular, conical or inflated with domed to flat center. 10. Blocky crenate. 11. Blocky pilate. 15. Cylindroid. 16. Cylindroid bulbous. 17. Cylindroid pilate. 18. Cylindroid reticulate.

Blocky

Cylindroid

19. Cylindroid scrobiculate.

Polyhedral plates/ Pla9a Segmented, psilate (“vertebral column”, 23) (25–40 μm).

Segmented, psilate

84

Epidermal silicified structures / Sto-1 Silicified stomate (25–50 μm).

Elongate bodies / El-5a Unciform, smooth (“simple solid trichome”, 23). Epidermal silicified structures/ Trac-1 Silicified tracheid (50–150 μm).

Silicified tracheid ?

83

Epidermispolyhedral epidermis.

Epidermisanticlinal epidermis. Epidermispitted polyhedral epidermis. Epidermisdiagnostic polyhedral epidermal form.

85

86 Poaceae

Agr (hb)

Other epidermal forms

D

87

Globular types

88

Arecaceae, Dipterocarpaceae (13), Apa (p) / A (ar) / Mo Strelitziaceae (12, 14), (h) Bromeliaceae (14)

ND

Mt1

Globular spherical echinate

Globular type 1

MT1 Globulolithum Globular (= Spherical) Gl (3, and sphaeroechinulathu equivalent name reference therein). 44 Gl04. m

24. Globular echinate

Globular bodies / Glo-1 Large globular Echinate (Ø 8–17 μm). echinate

Globularechinate.

25. Globular echinate large

Globular bodies / Glo-6 Globular Micro-echinate (Ø echinate 15–20 μm).

31. Globular psilate

Globular bodies / Glo-5 GlobularGlobular psilate Psilate (Ø 15–20 μm). psilate.

Forest indicator taxa (FI) / Palm / PALM / Globular echinate.

89

Arecaceae, Dipterocarpaceae (13), Apa (p) / A (ar) / Mo Strelitziaceae (12, 14), (h) Bromeliaceae (14)

ND

90

Capparidaceae

D

Ellipsoid echinate Boscia type

91

Woody/herbaceous basal angiosperms and Ad (arh) eudicotyledons.

ND

Mt3

Globular Smooth spherical smooth spherical type 1 type 1

Globular (= Spherical) Gl (3, and MT3 Globulolithum equivalent name reference therein). 45 sphaeropsilathum Gl05 p.p.

92


ND

Mt5

Globular elipsoidal smooth

Globular (= Spherical) Gl (3, and MT5 Globulolithum equivalent name reference therein). 43 sp 3 Gl03.

36. Globulose, Ovate with psilate texture.

93


ND

Mt6

Globular Smooth spherical smooth spherical type 2 type 2

Macro-globular (= Macro-spherical) Mg (3, and equivalent name reference therein). 53 Mg01.

32. Globular psilate large.

Ad (ar)

Mt2

Globular elipsoidal echinate

Globular typo 2

Globular (= Spherical) Gl (3, and MT2 Globulolithum equivalent name reference therein). 42 sp 1 Gl02.

F. Globular s.l. / Ellipsoid echinate; Ellipsoid with a echinate surface, single or composed, only in bark of Boscia species.

Smooth elipsoidal

MT6 Globulares lisos de más de 12 μm de diámetro

Forest indicator taxa (FI) / Other FI: woody/herbaceous basal angiosperms and eudicotyledons, conifers, ferns, etc. / Other FI / Simple and compound globular psilate (3-15 µm).

F. Globular s.l. / Globular/ Subglobular psilate; Psilate sphere, single or composed, sometimes with dark core.

Globular bodies / Glo-7 Tuberculate with large smooth rounded projections (Ø 15–20 μm).

94

Marantaceae / Woody/herbaceous A (arh) basal angiosperms and eudicotyledons.

Globular (= Spherical) Gl (3, and equivalent name reference therein).

ND Mt4

Ad (ar)

Globular spherical rugulose

Rugulose spherical

MT4 Globulolithum sp 2

Forest indicator taxa (FI) / Other FI: woody/herbaceous basal angiosperms and eudicotyledons, conifers, ferns, etc. / Other FI / Globular verrucate (> 15 µm).

F. Globular s.l. / Globular/ Subglobular decorated; Sphere with various surface decorations, single or composed.

Globular bodies / Glo-7 Tuberculate with large smooth rounded projections (Ø 15–20 μm).

Acanthaceae (11)

96


ND

97


ND

98


ND

99

Thymelaceae

Ad (arh)

D

Globular facetate

100

Strelitziaceae (Zingiberales?)

Mo (h)

D

Druse-like bodies

D

Forest indicator taxa (FI) / Other FI: woody/herbaceous basal angiosperms and eudicotyledons, conifers, ferns, etc. / Other FI / Globular granulate (3-10 µm).

Globular granulate

G. Aggregates / Nodular / Aggregation of small, smooth nodules with various shapes: globular, elongate or irregular.

28. Globular granulate, Spheroid with granular texture.

G. Aggregates / Granular / Irregular aggregation of small, smooth granules.

Globular bodies / Glo-3 Granulate/tuberculate 29. Globular granulate large / with small smooth Large globular Large spheroid with granular rounded projections granulate texture. and often dark core (Ø 8–15 μm).

Druse-like bodies

Mo (h)

D

102

Woody/herbaceous basal angiosperms and Ad (ar) eudicotyledons.

D

103

Fabaceae / Bombacaceae

D

27. Globular folded, Subspheroid with extended, thickened rim.

ND

33. Globular scrobiculate, Spheroid with pitted texture.

ND

34. Globular tuberculate, Large spheroid beset with nodular processes.

105

Globulargranulate.

Forest indicator taxa (FI) / Zingiberales / ZINGI / Costustype body.

Zingiberales

Woody/herbaceous basal angiosperms and Ad (arh) eudicotyledons. Marantaceae / Woody/herbaceous A (arh) basal angiosperms and eudicotyledons.

Globular bodies / Glo-2 Globular Granulate (Ø 12–17 granulate μm).

Globular bodies / Glo-4 30. Globular granulate Granulate / oblong / Ovate with granular tuberculate with a texture. central vacuum (Ø 8–15 μm). 26. Globular facetate / Facetate spheroid/hemisphere.

101

104

Globularnodular.

Large globular with rugulose hemisphere

95

Ad (ar)

Nodulose or crushed decoration globular

Zingiber type

F. Globular s.l. / Ellipsoid/Lobate; Elliptic to irregular (with two or more lobes), psilate.

106

Ebenaceae

Ad (ar)

D

35. Globular verrucate oblong, Large elongated subspheroid with irregular texture and processes.

107

Solanaceae

Ad (arh)

D

37. Globulose bisected, Cubic body, orbicular to square in outline, with adaxial bisection and psilate texture.

108

Marantaceae

Mo (h)

D

Globularirregularly angled/folded

ND

Cystoliths

Other Globular

109

Woody/herbaceous basal angiosperms and Ad (arh) eudicotyledons. Large faceted elements

110


D

111


D

112

Various plants and unknown. Elongate facetate.

ND

113

Various plants and unknown. Elongate facetate.

Ad (arh)

Mt7

Faceted isodiametric

Faceted spherical

Multicavate (= Faceted) Mc (3, and MT7 Isodiamétricos equivalent name reference therein). 50 facetados Mc02.

Forest indicator taxa (FI) / Other FI: woody/herbaceous basal angiosperms and eudicotyledons, conifers, ferns, etc. / Other FI / Blocky polyhedron” (equidimensional irregular or facetate body).

Mt8

Faceted elongate

Faceted elongate

MT8 Elongados facetados

Multicavate (= Faceted) Mc (3, and equivalent name reference therein). 52 Mc04.

Forest indicator taxa (FI) / Other FI: woody/herbaceous basal angiosperms and eudicotyledons, conifers, ferns, etc. / Other FI / Faceted elongate or blocky bodies with tubercles.

A. Sclerenchyma / A.1. Fibre / Fibre facetate / Long and thin, sometimes with pointed end(s), psilate, with large concave facets.

45. Tabular facetate. Facetate with partly psilate and partly corniculate textures.

Multi-faceted bodies

Faceted elongate

D. Elongate / Elongate facetate / Elongate, with concave facets Various plants and unknown 4. Blocky facetate. Faceted, either in side or surface view, / NDO / Elongate facetate. psilate block. representing impressions of adjacent cells.

Other faceted

Ad (arh)

Irregular faceted

A. Sclerenchyma / A.2. Sclereid / Sclereid pitted / (iii) elongate columellate facetate.

ND

Tracheid, Sclereid and vascular elements

ND

Mt13

Tracheids with helical thickkenings

Helical MT11 Traqueidas thickkenings

Straited oblong form from xylem cell (16); Tracheid p.p. (17), Tracheid (sensu 4).


ND

Mt14

Tracheids with irregular body and striate ornamentation

MT13 Cuerpos Irregular type irregulares con 1 ornamentaciones estriadas.

Short tracheid (= Irregular body with striate ornamentation) Ta (3, and equivalent name reference therein).

A. Sclerenchyma / A.2. Sclereid / Sclereid pitted p.p.


ND

Mt15

Tracheids with irregular body Irregular type and widely 2 spaced verrucate nodes

Tracheid p.p. (17); Irregular body with widely spaced verrucate nodes (18)


114

Various plants and unknown.

115

116

ND (nd)

MT14 Cuerpos irregulares con ornamentaciones verrucosas espaciadas

Irregular stipulate tracheid

Tracheary elementstracheid.

Grasses and other monocotyledons, conifers / NDG / Monocotyledon tracheary element (cylindric scalariform or scrobiculate), etc.

117


ND (nd)

ND

cylindric scalariform or scrobiculate

118


ND (nd)

ND

Cylindroid bulbous

13. Cylindroid bulbous. Cylindroid/renate with a psilate texture and irregular bulbuous enlargements.

119


ND (nd)

ND

Cylindroid columellate

14. Cylindroid columellate. Very large cylindroid with surface covered by processes.

120


ND

Irregular sulcate or facetate vascular cell

Forest indicator taxa (FI) / Other FI: woody/herbaceous basal angiosperms and eudicotyledons, conifers, ferns, etc. / Other FI / Irregular sulcate or facetate vascular cell (incluiding Magnolia-type “terminal tracheid” and tuberculated rounded sclereid/tracheid).

Favose aggregate sclerenchyma and similar

Forest indicator taxa (FI) / Other FI: woody/herbaceous basal angiosperms and eudicotyledons, conifers, ferns, etc. / Other FI / Favose aggregate sclerenchyma and similar (irregular clavate body, often branched, sometimes facetate or papillate surface).

121


122


ND

Fibre pitted

A. Sclerenchyma / A.1. Fibre / Fibre pitted / Long and thin, sometimes with pointed end(s), psilate surface, with elongate, crest-like knobs in oblique position to the long axe of the phytolith (¼ silica filling of fibre pits).

123

Woody basal angiosperms and eudicotyledons.

Ad (ar)

ND

Fibre psilate

A. Sclerenchyma / A.1. Fibre / Fibre psilate / Long and thin, with pointed end(s), sometimes bent, psilate.

124


Ad (ar)

ND

Sclereid pitted


125


Ad (ar)

ND

Sclereid psilate

A. Sclerenchyma / A.2. Sclereid / Sclereid psilate / i) elongate or irregular with thick silicified walls, non-silicified lumen and pit channels.

126

127

128

Woody/herbaceous basal angiosperms and Ad (arh) eudicotyledons. Woody/herbaceous basal angiosperms and Ad (arh) eudicotyledons. Woody/herbaceous basal angiosperms and Ad (arh) eudicotyledons.

ND

Echinate tracheid

Blocky bodies. Blo-10 Angular, attenuate ends (“sclereid”, 24; 23) (45–50 μm).

15. Cylindroid crenate. Cylindroid with scalloped edges. 16. Cylindroid lacunate. Cylindroid with lacunose texture. 17. Cylindroid large. Elongate bodies. El-1a Cylindroid of large size with Cylindric, smooth psilate texture. (100–150 μm). 18. Cylindroid reticulate. Subcylindroid psilate body partly covered by a net-like pattern.

ND

ND

ND Cylindroid

129


ND

130


ND

131


Ad (ar)

ND

Mt16

Vascular Vascular element element with with simple simple perforation perforate plates plates

MT15 Elementos vasculares con Vascular cell or tissue p.p. (4). placa de perforación simple

132


Ad (ar)

ND

Mt17

Vascular Vascular element element with with scaleriform scaleriform perforation perforate plates plates

MT16 Elementos vasculares con placa de perforación escalariforme .

Vascular cell or tissue p.p. (4).

133


Ad (ar)

ND

Mt18

Vascular Vascular element element with with intervessel intervessel pits bordered pits alternate

MT17 Elementos vasculares con punteaduras areoladas

Vascular cell or tissue p.p. ( 4).

134


Ad (ar)

ND

Mt19

Vascular Vascular element element with with intervessel intervessel pits scaleriform pits scaleriform

MT18 Elementos vasculares con punteaduras escalariformes

Vascular cell or tissue p.p. (4).

MT9 Lobulados

Lobular (= Jigzaw) Lb (3, and equivalent name reference therein). 48 Lb01.

19. Cylindroid scrobiculate. Large cylindroid with scrobiculate texture.

Epidermal polyhedral or puzzles / hairs / hair bases

135


ND

Mt9

Laminar puzzle forms

136

Woody/herbaceous basal angiosperms and Ad (nd) eudicotyledons.

ND

Mt10

Structured puzzle forms

137

Woody/herbaceous basal angiosperms and Nd (nd) eudicotyledons, conifers, ferns, etc.

138


Puzzles

20. Epidermal jig-saw. Thin epidermal cells with puzzlelike outline. Variable textures.

Structured lobular (= Jigzaw ?) sLb (3, and equivalent name reference therein). 94 sLb02.

ND

Anticlinal epidermis silicified nongrass (reniform)

Forest indicator taxa (FI) / Other FI: woody/herbaceous basal angiosperms and eudicotyledons, conifers, ferns, etc. Other FI / Anticlinal epidermis silicified non-grass (reniform).

ND

Thin epidermal polygonal cells

Forest indicator taxa (FI) / Other FI: woody/herbaceous basal angiosperms and eudicotyledons, conifers, ferns, etc. Other FI / Polyhedral epidermis (various types).

22. Epidermal polygonal. Thin epidermal polygonal cells. Variable textures.

Polyhedral plates

Sclereid

Tracheary elements

Point-shaped (= Point-shaped, unciform hair) Ps Zucol et al. 2010a and equivalent name reference therein. 1–10 Ps1–Ps10 s.l.

139


ND

140


ND

Multi-layered trichome

141

Woody/herbaceous basal angiosperms and Ad (nd) eudicotyledons, conifers, ferns, etc.

ND

Guard and subsidiary cells

Forest indicator taxa (FI) / Other FI: woody/herbaceous basal angiosperms and eudicotyledons, conifers, ferns, etc. Other FI / Guard and subsidiary cells.

142

Grasses and other monocotyledons, conifers.

ND

Stomatal complex

Grasses and other monocotyledons, conifers. / NDG / Silicified monocotyledon stomatal complex.

Mo (nd)

Mt11

Hair bases

Hair bases

MT10 Base de elementos aguzados

Hairs

Acicular bodies. Aci-3a Smooth, compartimented (“multi-layered trichome” , 23) (30–100 μm).

41. Stomata/hair/base. Stomatal complex, hairs and hair bases.

Other prismatic elements

143


Nd (nd)

ND

Mt22

Brief elongate (= Brief prismatic) Be, and Short prismatic Prismatic MT21 Prismáticos Micro–elongate (= Micro-elongate) Me Z elongated with smooth type de menos de 30 μm (3, and equivalent name reference smooth contour 1 de largo therein).

144


Nd (nd)

ND

Mt23

Elongate (= Parallelepipedal or Prismatic) Prismatic Prismatic MT22 Various plants and unknown elongated with smooth type Macroprismatolithu El (3, and equivalent name reference / NDO / Elongate psilate. therein). 63 El09. smooth contour 2 m psilaristathum

145


Nd (nd)

ND

D. Elongate. Elongate scalariform. Elongate, with unsilicified elliptic cavities (pits?) on the surface arranged perpendicular to the length of the body, reminding of a staircase.

146


Nd (nd)

ND

D. Elongate. Elongate vesicular. Elongate, thin, with large, spherical to elliptic concavities, only in wood of Capparis tomentosa.

147


Nd (nd)

ND

D. Elongate. Elongate with globular part(s). Cylindroid, with spherical to elliptic central or terminal swellings.

148


Nd (nd)

ND

D. Elongate. Elongate irregular. Elongate, high variability in shape and size.

149


Nd (nd)

ND

150


Nd (nd)

ND

151


Nd (nd)

ND

3. Blocky corniculate. Ovate, rectangular, polygonal or irregular block with corniculate projections. Variable textures.

152


Nd (nd)

ND

5. Blocky hairy. Ovate, rectangle, polygonal or irregular block with irregular/scrobiculate texture; long pointed projections.

153


Nd (nd)

ND

6. Blocky pilate. Orbicular, oblong, rectangle, polygonal or irregular block with scrobiculate texture and projections that have straight or slightly curved ends.

154


Nd (nd)

ND

7. Blocky polygonal. Polygonal/irregular block with a psilate/scrobiculate texture.

155


Nd (nd)

ND

8. Blocky radiating. Orbicular to rectangular block with a surface covered by concentric layers.

156


Nd (nd)

ND

157


Nd (nd)

ND

158


Nd (nd)

ND

159


Nd (nd)

ND

160


Nd (nd)

ND

161


Nd (nd)

ND

162


Nd (nd)

ND

1. Blocky. Block with lacunose/slightly scrobiculate texture. 2. Blocky cavate. Block with psilate texture and centric depression.

Other prismatic (elongate, blocky and tabular

9. Blocky trapezoid. Trapezoid block with lacunose texture. 10. Blocky tuberculate. Irregular block with lacunose/scrobiculate/tuber culate texture. 42. Tabular corniculate. Elongate parallelepiped with corniculate texture. 43. Tabular crenate. Tabular body with psilate texture and scalloped edges. 44.Tabular elongate. Elongate. 46. Tabular lanceolate. Large tabular, psilate body shaped like a spear. 47. Tabular oblong. Large tabular body with orbicular, ovate or oblong outline and psilate texture.

Hair bases

Stomata

48. Tabular psilate. Elongate to oblong body with psilate texture. 49. Tabular ridged. Thin parallelepiped, orbicular to square, with surface covered by ridges.

163


Nd (nd)

ND

164


Nd (nd)

ND

165


Nd (nd)

ND

50. Tabular scrobiculate. Variably wide parallelepiped with an elongate outline and scrobiculate texture.

166


Nd (nd)

ND

51. Tabular thick contorted. Thick parallelepiped with a contorted profile and irregular to laminate texture.

167


Nd (nd)

ND

52. Tabular thick dendritic. Large elongate parallelepiped with psilate texture and sides covered by irregular dedritic processes.

168


Nd (nd)

ND

53. Tabular thick lacunate. Large thick parallelepiped with scrobiculate/lacunose texture.

169


Nd (nd)

ND

54. Tabular thick sinuate. Thick parallelepiped/subcylindroid with psilate texture. One or two sides are sinuate; sometimes, one side is sinuate and the other one is columellate.

170


Nd (nd)

ND

55. Tabular thin. Thin parallelepiped, square to rectangular shape with psilate texture.

171


Nd (nd)

ND

56. Tabular thin pilate. Thin parallelepiped, scrobiculate or lacunate texture and rodlike processes.

172


Nd (nd)

ND

57. Tabular trapezoid. Elongate body with a trapezoidal to triangular cross-section and psilate texture.

Cyperaceae cones and Wetland plant forms

173

Sedges, unknown Acy (h) aquatic monocotyledon

D

174

Sedges, unknown Acy (h) aquatic monocotyledon

D

175

Cyperaceae

Acy (h)

D

176

Cyperaceae

Acy (h)

D

177

Cyperaceae

Acy (h)

D

178

Cyperaceae

Acy (h)

D

Mt32

Laminar with pentagonal or hexagonal contour

Polygonal plate (= Polygonal plate) Pg (3, and equivalent name reference therein).

Cyperoid type

Wetland plants / Sedges, Equisetum, unknown aquatic monocotyledon / AQ / Epidermal plate with coneshapes ("papilae").

Blocky bodies

Sinuous suborbicular with central protuberance

Mt26

Prismatic elongated with irregular simetric or asimetric contour

Sinuous suborbicular with central protuberance Polyhedral plates / Elongate bodies. El-6 Reflexed parallelepipedal, smooth (30–75 μm).

Elongate (= Parallelepipedal or Prismatic) MT25 Macroprismatolithu El (3, and equivalent name reference therein). 55 El01 and 60 El06. m sp. 2 Wetland plants / Sedges, Equisetum, unknown aquatic monocotyledon / AQ / Silicified stellate facetate parenchyma.

Silicified stellate facetate parenchyma

Pla-3a. Stellate, psilate (25–30 μm).

Blocky bodies. Blo-1 Long acicular to oblong, sometimes pinched-point, often infilled with black material (50–75 μm). Blocky bodies. Blo-2 Irregular with non welldefined shape (30–100 μm).

Blocky bodies

179

Cyperaceae

Acy (h)

D

Blocky bodies. Blo-7 Parallelepipedal, smooth, often infilled with black material (25–100 μm).

180

Cyperaceae

Acy (h)

D

Blocky bodies. Blo-9 Orbicular to ovate, smooth, slightly curved (10–15 μm).

181

Cyperaceae

Acy (h)

D

Polyhedral plates. Pla4 Planar, hexagonal, scorbiculate, cavate in the center (30–35 μm).

182

Cyperaceae

Acy (h)

D

Polyhedral plates. Pla5 “Sinuous shape, knobby apex, psilate sculpturing, satellites absent = NKXWI” (from 25) (15–20 μm).

D

Polyhedral plates. Pla6 “Angular to subrounded shape, pointed apex, psilate sculpturing, satellites present = A(R)KXWI” (from 25); papillae cell or “hat-shaped” (26) (10–20 μm).

D

Polyhedral plates. Pla7a “Rounded shape, knobby apex, psilate sculpturing, satellites absent = RPXWI” (from 25); papillae cell or “hat-shaped” (26) (10–15 μm). Epidermal silicified structures. Str-1a Articulated epidermal structure of Cyperaceae-papillae (more than three articulated elements)

Polyhedral plates 183

184

Cyperaceae

Cyperaceae

Acy (h)

Acy (h)

185

Cyperaceae

Acy (h)

D

186

Podostemaceae

Apo (h)

D

Mt36

Irregular elongated

Granulate polygonal platelet with central Cyperaceae protuberance / cones Smooth conical body with satellites.

Podostemac MT28 Fusiformes eae type

Fusiform Fu (3, and equivalent name reference therein).

Podostemaceae (15)

Others forms

187

Diagnostic forms Marantaceae

Mo (h)

D

Troughed

Troughed 39. Shield. Tabular to trapezoidal body with one side curved and the other one wavy. Psilate to scrobiculate texture. Rim is bevelled.

188

Solanaceae / Sterculiaceae

Ad (arh)

D

Shield

189

Commelinaceae

Mo (h)

D

Anisopolar polygonal top

Anisopolar polygonal top, decorated cylindrical bottom

190

Zingiberales

Mo (h)

D

Trough body

Trough body

191

Marantaceae

Mo (h)

D

Cylindrical with protuberances or decoration

Cylindrical with protuberances or decoration and central protuberance

192

Amaranthaceae

Ad (arh)

D

Lenticular concave/convex

193

Asteraceae

Ad (h)

D

Opaque perforated plates B.Parenchyma/Cork (= P/C) / B.1. Blocky / P/C Blocky with irregular projections / Polyhedral to paralellepipedal, psilate, with irregular projections at the edges, only in bark of Stereospermum kunthianum, single and multicell.

38. Lenticular concave/convex.Hemispheri c body with rugose to sulcate texture. Opaque perforated plates

194

Bignoniaceae

Ad (ar)

D

Blocky with irregular projections

195

Capparidaceae

Ad (ar)

D

Blocky multilayered

B.Parenchyma/Cork (= P/C). B.1. Blocky. P/C Blocky multilayered. ±Paralellepipedal blockies consisting of numerous thin tabular layers, not seen as multicell, only in bark of Maerua crassifolia. B.Parenchyma/Cork (= P/C). B.1. Blocky. P/C Blocky cavate fimbriate. Polyhedral to paralellepipedal with a central cavity and fimbriate edges, psilate, only in bark of Celtis mildbraedii, single and multicell.

196

Cannabinaceae

Ad (arh)

D

Blocky cavate fimbriate

197

Cannabinaceae

Ad (arh)

D

Echinate irregular platelet

198

Dipterocarpaceae

Ad (ar)

D

Club-shaped body

199

Fabaceae

Ad (ar)

D

Tabular, subpolygonal body

200

Proteaceae

Ad (ar)

D

Spooliform.

201

Arecaceae

Apa (p)

D

Conical echinate

Echinate irregular platelet 11. Clavate granulate. Clubshaped body with granular texture. 21. Epidermal laminate. Tabular, sub-polygonal body covered with layers. 40. Spooliform. Cylindroid body with mesial strangulation. Conical echinate (hatshaped)

Non diagnostic forms

202


Ad (ar)

ND

Parenchyma strand

B.Parenchyma/Cork (= P/C). B.2.Parenchmyma strand. Parenchmyma strand. Multicell strand of elongate cells, rarely paralellepipedal blockies, with ± straight margins, psilate; also single elongate cells.

203


Ad (ar)

ND

Cork aerenchyma

C. Cork aerenchyma. Cork aerenchyma. Tabular silicified tissue, various shapes, with irregular perforations of various size.

204


Ad (ar)

ND

Blocky polyhedral or tabular, with penta- to hexagonal faces.

B.Parenchyma/Cork (= P/C) / B.1. Blocky / P/C Blocky polyhedral / (i) isodiametric with penta- to hexagonal faces, psilate, single and multicell. / (ii) tabular, penta- to hexagonal, psilate, single and multicell.

205

ND

ND (nd)

ND

Blocky irregula

E. Blocky unspecific. Blocky irregular. Variable in shape and size.

206

ND

ND (nd)

ND

Tabular irregular

E. Blocky unspecific. Tabular irregular. Tabular, thin, variable in shape and size.

207

ND

ND (nd)

ND

Aggregated

H. Silica particles/ Accumulations / Small irregular silica particles sometimes occurring aggregated, amorphous silica aggregations.

208

ND

ND (nd)

ND

Mt33

209

ND

ND (nd)

ND

Mt35

210

ND

ND (nd)

ND

Mt37

Laminar with different contour Echinate and verrugate or plates echinate surface Triangular Irregular isodiametric

MT12 Cuerpos prismáticos laminares de superficie columelada o equinada

Plate papillose (= Laminar papillae plate) Lp (3, and equivalent name reference therein). Triangular (= Triangular) Tr (3, and Proteiform (= Irregular ovoid) Pr and Scrobiluar (= Nuxolita) Nx (3, and

Conical/ hatshaped

B.Parenchyma/Cork (= P/C) / B.1. Blocky / P/C Blocky paralellepipedal / Rectangular (2D)/paralellepipedal (3D), L/W 2, psilate, single and multicell.

211

ND

ND (nd)

ND

Blocky paralellepipedal

212

ND

ND (nd)

ND

Single-cell, fluidconducting tissue

213

ND

ND (nd)

ND

Parallelepipedal to cubic, granulate

Blocky bodies. Blo-3 Parallelepipedal to cubic, granulate (25–100 μm).

214

ND

ND (nd)

ND

Cubic, smooth

Blocky bodies. Blo-6 Cubic, smooth, often infilled with black material (25–75 μm).

215

ND

ND (nd)

ND

parallelepipedal, pinched-point

Elongate bodies. El-2 Tabular / parallelepipedal bodies, pinched-point (50–75 μm).

216

ND

ND (nd)

ND

Cylindric, laminate

Elongate bodies. El-4 Cylindric, laminate, with a median swelling (75–150 μm).

217

ND

ND (nd)

ND

Polyhedral plates

218

ND

ND (nd)

ND

Orbicular, stellate inside

219

ND

ND (nd)

ND

Stalked granulate body

220

ND

ND (nd)

ND

Orbicular planoconvex verrucose

23. Epidermal tracheid. Single-cell, fluid-conducting tissue.

Polyhedral plates. Pla2a Rectangular, lacunate (30–40 μm). Polyhedral plates. Pla8a Orbicular, stellate inside (hair base) (Ø 40–60 μm). Stalked granulate body (Type I) Orbicular planoconvex verrucose (Type II)

Table 3. Phytolith morphotypes sketch.

Grouping according to its botanical affinity and diagnosis

Nomenclator

Arundinoideae

BEP clade

Pooideae Ehrhartoideae Bambusoideae

Poaceae

D Poaceae morphotype

Arecaceae (in echinate Ø 8–40 μm)

Dipterocarpaceae (Ø 4–10 μm)

Zingiberales

Marantaceae Commelinaceae

Cyperaceae

ID ID ID

Polylobate Other bilobate Other bilobate

Agr (h) PACMAD general Agr (h) PACMAD-clade

ID ID

Other bilobate Quadralobate/cross

Agr (hb) PACMAD-clade

ID

Rondell

Agr (hb) PACMAD-clade Agr (h) Panicoideae Agr (h) Panicoideae Agr (h) Panicoideae Agr (h) Chloridoideae Agr (h) Chloridoideae Agr (h) Chloridoideae Agr (h) Chloridoideae Agr (h) Chloridoideae Agr (h) Chloridoideae - Arundinoideae Agr (h) Arundinoideae Agr (hb) basal BEP clade Agr (hb) basal BEP clade Agr (hb) basal BEP clade Agr (hb) basal BEP clade Agr (h) Pooideae Agr (h) Pooideae Agr (h) Pooideae Agr (h) Pooideae Agr (h) Ehrhartoideae Agr (hb) Bambusoideae Agr (hb) Bambusoideae Agr (hb) Bambusoideae

ID ID ID ID ID ID ID ID ID ID ID ID ID ID ID ID ID ID ID ID ID ID ID

Other phytolith types (non-GSSCs) Dumbbell Quadralobate/cross Other phytolith types (non-GSSCs) Thin, long-shanked bilobate Saddles Saddles Saddles Saddles Saddles Saddles Saddles Other phytolith types Other phytolith types Carinate Square Trapezoid crenate Trapezoid crenate Scooped bilobate Other bilobate Quadralobate/cross Chusquea-type rondel

Mt34 Mt34 Mt34 Mt34

Mt21 Mt21 Mt21 Mt21 Mt12 Mt12 Mt12 Mt12 Mt12 Mt28 Mt29

19 20 21 22 34 35 36 37 38 39 40

Poaceae /P+BE clade and PACMAD-clade Poaceae /P+BE clade and PACMAD-clade Poaceae /P+BE clade and PACMAD-clade Poaceae (GSSC)/P+BE clade and PACMAD-clade Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae

Agr (h) BEP and PACMAD clades Agr (h) BEP and PACMAD clades Agr (h) BEP and PACMAD clades Agr (h) BEP and PACMAD clades Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae

ID D D D D D D D D D D

Truncated conical / Towers Truncated conical / Towers Truncated conical / Towers Truncated conical / Towers Point-shaped Point-shaped Point-shaped Point-shaped Point-shaped Fan-shaped Polyhedral bulliforms

41

Poaceae

Agr (hb) Poaceae

D


42

Poaceae

Agr (hb) Poaceae

D


43

Poaceae

Agr (hb) Poaceae

D


44

Poaceae

Agr (hb) Poaceae

D

45

Poaceae

Agr (hb) Poaceae

D

46

Poaceae

Agr (hb) Poaceae

D

47 48 49 50 51 52 53 54 55 56 57 58 59

Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae

Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae

D D D D D D D D D D D D D

60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87

Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae

Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae Agr (hb) Poaceae

D D D D D D D D D D D D D D D D D D D D D D D D D D D D

Other parallelepipedal bulliform types Prismatic elongated with serrate contour Prismatic elongated with denticulate contour Prismatic elongated with undulate contour Other prismatic elongated Other prismatic elongated Other prismatic elongated Other prismatic elongated Other prismatic elongated Other prismatic elongated Other prismatic elongated Other prismatic elongated Other prismatic elongated Other prismatic elongated Various Acicular hair cell Circular, conical or inflated with domed to flat center Blocky Blocky Cylindroid Cylindroid Cylindroid Cylindroid Cylindroid Elongate Epidermal crenate Epidermal polygonal Globular psilate Scutiform Stomata/Hair Tabular Tabular Tabular Tabular Tabular Tabular Tabular Vessel laminate Unciform, smooth Silicified tracheid ? Segmented, psilate Other epidermal forms Other epidermal forms Other epidermal forms

ND

Globular spherical echinate

Mt1

ND D D D D D

Globular elipsoidal echinate Conical echinate Druse-like bodies Zingiber type Trough body Other Globular Cylindrical with protuberances or decoration Troughed Anisopolar polygonal top

Mt2

23

Chloridoideae

Agr (h) PACMAD general Agr (h) PACMAD-clade Agr (h) PACMAD-clade

31 1 9 32 3 12 13 18 33 16 17 14 15 29 30 8 25 26 27 28 7 11 24

6 10

Panicoideae

Botanical affinity/ Life forms

Poaceae / PACMAD-clade/ Danthonioideae? Bambusoideae? (C3) / PACMAD general Poaceae / PACMAD-clade / Other PACMAD (C3/C4) Poaceae / PACMAD-clade / Other PACMAD (C3/C4) Poaceae / PACMAD-clade / Danthonioideae? Bambusoideae? (C3) / PACMAD general Poaceae / PACMAD-clade / Other PACMAD (C3/C4) Poaceae / PACMAD-clade / Danthonioideae? Bambusoideae? (C3) / PACMAD genera Poaceae / PACMAD-clade / Danthonioideae? Bambusoideae? (C3) / Very tall with prominently faceted tops. Poaceae / PACMAD-clad e/Panicoideae (C3/C4) Poaceae / PACMAD-clade / Panicoideae (C3/C4) Poaceae / PACMAD-clade / Panicoideae (C3/C4) Poaceae /PACMAD-clade/Chloridoideae (C4). Aristida-type Poaceae / PACMAD-clade/ Chloridoideae (C4?) Poaceae / PACMAD-clade / Chloridoideae Poaceae / PACMAD-clade/ Chloridoideae Poaceae / PACMAD-clade/ Chloridoideae (C4?) Poaceae / PACMAD-clade/ Chloridoideae - Arundinoideae Poaceae / PACMAD-clade/ Arundinoideae Poaceae / basal P+BE clade Poaceae / basal P+BE clade Poaceae / basal Poaceae+ BE clade. Regular chusquoid body. Poaceae / basal Poaceae+ BE clade. Crescentic keeled rondel. Poaceae / Pooideae / diagnostic / Stipoideae Poaceae / Pooideae Poaceae / Pooideae/ Crenate “trapeziform polylobate”. Poaceae / Pooideae/ Trapeziform short cells “pyramidal”. Poaceae / basal P+BE clade/ Ehrhartoideae Poaceae / basal P+BE clade/ Bambusoideae Poaceae / basal Poaceae+ BE clade/ Bambusoideae Poaceae / basal P+BE clade/ Bambusoideae

2 4 5

PACMAD-clade

Botanical affinity

diagnostic (D)/ intrafamily diagnostic (ID)/ non diagnostic (ND) Acronym and name, this study

Arecaceae, Dipterocarpaceae (13), Strelitziaceae (12, 14), Bromeliaceae (14) Arecaceae Strelitziaceae (Zingiberales) Zingiberales Zingiberales Marantaceae

(Ø 8–40 μm) Apa (p) Arecaceae / (Ø 4–10 μm) A (ar) Dipterocarpaceae / Mo (h) Strelitziaceae, Bromeliaceae (Ø 8–40 μm) Apa (p) Arecaceae / (Ø 4–10 μm) A (ar) Dipterocarpaceae / Mo (h) Strelitziaceae, Bromeliaceae Apa (p) Arecaceae Mo (h) Zingiberales Mo (h) Zingiberales Mo (h) Zingiberales Mo (h) Marantaceae

191 187 189

Marantaceae Marantaceae Commelinaceae

Mo (h) Marantaceae Mo (h) Marantaceae Mo (h) Commelinaceae

D D D

173

Sedges, unknown aquatic monocotyledon

Acy (h) principally Cyperaceae

D

174

Sedges, unknown aquatic monocotyledon

Acy (h) principally Cyperaceae

D

175

Cyperaceae

Acy (h) Cyperaceae

D

Laminar with pentagonal or hexagonal contour Sinuous suborbicular with central protuberance Prismatic elongated with irregular simetric or asimetric contour

176 177 178 179 180 181 182 183

Cyperaceae Cyperaceae Cyperaceae Cyperaceae Cyperaceae Cyperaceae Cyperaceae Cyperaceae

Acy (h) Cyperaceae Acy (h) Cyperaceae Acy (h) Cyperaceae Acy (h) Cyperaceae Acy (h) Cyperaceae Acy (h) Cyperaceae Acy (h) Cyperaceae Acy (h) Cyperaceae

D D D D D D D D

Silicified stellate facetate parenchyma Blocky bodies Blocky bodies Blocky bodies Blocky bodies Polyhedral plates Polyhedral plates Polyhedral plates

or Strelitziaceae 88 and/or Bromeliaceae (Ø 4–10 μm) 89 201 100 101 190 108

Arecaceae, Dipterocarpaceae (13), Strelitziaceae (12, 14), Bromeliaceae (14)

Mt30

Mt20

Mt34 Mt34 Mt34

Mt31

Mt30

Mt24 Mt25 Mt27

Mt32

Mt26

Cyperaceae

Podostemaceae Capparidaceae Capparidaceae Acanthaceae Ebenaceae Fabaceae / Bombacaceae Fabaceae Bignoniaceae Dipterocarpaceae

Woody basal angiosperms and eudicotyledons Woody basal angiosperms and eudicotyledons

Herbs basal angiosperms and eudicotyledons

Cyperaceae Cyperaceae Podostemaceae

Acy (h) Cyperaceae Acy (h) Cyperaceae Apo (h) Podostemaceae

D D D

Polyhedral plates

90 195 95 106

Capparidaceae Capparidaceae Acanthaceae (11) Ebenaceae

Ad (ar) Capparidaceae Ad (ar) Capparidaceae Ad (ar) Acanthaceae Ad (ar) Ebenaceae

D D D D

Ellipsoid echinate Boscia type Blocky multilayered Globular spherical rugulose Other Globular

103 199 194 198 200 102 111

Fabaceae / Bombacaceae Fabaceae Bignoniaceae Dipterocarpaceae Proteaceae Woody/herbaceous basal angiosperms and eudicotyledons. Woody/herbaceous basal angiosperms and eudicotyledons.

Ad (ar) Fabaceae / Bombacaceae Ad (ar) Fabaceae Ad (ar) Bignoniaceae Ad (ar) Dipterocarpaceae Ad (ar) Proteaceae Ad (ar) Woody basal angiosperms and eudicotyledons Ad (ar) Woody basal angiosperms and eudicotyledons

D D D D D ND ND

115

Woody/herbaceous basal angiosperms and eudicotyledons.

Ad (ar) Woody basal angiosperms and eudicotyledons

ND

116 122 123 124 125

Woody/herbaceous basal angiosperms and eudicotyledons. Woody/herbaceous basal angiosperms and eudicotyledons. Woody basal angiosperms and eudicotyledons. Woody basal angiosperms and eudicotyledons. Woody basal angiosperms and eudicotyledons.

Ad (ar) Woody basal angiosperms and eudicotyledons Ad (ar) Woody basal angiosperms and eudicotyledons Ad (ar) Woody basal angiosperms and eudicotyledons Ad (ar) Woody basal angiosperms and eudicotyledons Ad (ar) Woody basal angiosperms and eudicotyledons

ND ND ND ND ND

131



ND

132



ND

133



ND

134 202 203

Woody basal angiosperms and eudicotyledons. Woody basal angiosperms and eudicotyledons. Woody basal angiosperms and eudicotyledons.

Ad (ar) Woody basal angiosperms and eudicotyledons Ad (ar) Woody basal angiosperms and eudicotyledons Ad (ar) Woody basal angiosperms and eudicotyledons

ND ND ND

204



ND

Other Globular Tabular, sub-polygonal body Blocky with irregular projections Club-shaped body Spooliform. Other Globular Faceted elongate Tracheids with irregular body and striate ornamentation Tracheids with irregular body and widely spaced verrucate nodes Fibre pitted Fibre psilate Sclereid pitted Sclereid psilate Vascular element with simple perforation plates Vascular element with scaleriform perforation plates Vascular element with intervessel pits bordered Vascular element with intervessel pits scaleriform Parenchyma strand Cork aerenchyma Blocky polyhedral or tabular, with pentato hexagonal faces.

193

Asteraceae

Ad (h) Asteraceae

D

Opaque perforated plates

192 196 197 107

Amaranthaceae Cannabinaceae Cannabinaceae Solanaceae

Ad (arh) Amaranthaceae Ad (arh) Cannabinaceae Ad (arh) Cannabinaceae Ad (arh) Solanaceae

D D D D

Lenticular concave/convex Blocky cavate fimbriate Echinate irregular platelet Other Globular

188 99

Solanaceae / Sterculiaceae Thymelaceae

Ad (arh) Solanaceae / Sterculiaceae Ad (arh) Thymelaceae

D D

Shield Globular facetate

91

Woody / herbaceous basal angiosperms and eudicotyledons.

Ad (arh) Woody / herbaceous basal angiosperms and eudicotyledons

ND

Globular spherical smooth type 1

Mt3

92



ND

Globular elipsoidal smooth

Mt5

93



ND

Globular spherical smooth type 2

Mt6

96



ND

Globular granulate

97



ND

Globular granulate

98



ND

Globular granulate

104



ND

Other Globular

109



ND

Other Globular

110



ND

Faceted isodiametric



ND

Other faceted

Irregular elongated

Mt36

Mt4

Mt8 Mt14 Mt15

Mt16 Mt17 Mt18 Mt19

Asteraceae Amaranthaceae Cannabinaceae Cannabinaceae Solanaceae Solanaceae / Sterculiaceae Thymelaceae

Basal angiosperms and eudicotyledons

184 185 186

Woody / herbaceous basal angiosperms and eudicotyledons

Woody / herbaceous 112 basal angiosperms and eudicotyledons. 113

Angiosperms

Angiosperms and eudicotyledons, conifers, ferns, etc. Grasses and other monocotyledons, conifers



ND

Other faceted

120



ND

121



ND

Irregular sulcate or facetate vascular cell Favose aggregate sclerenchyma and similar

126



ND

Cylindroid

127



ND

Cylindroid

128



ND

Cylindroid

129



ND

Cylindroid

130



ND

Cylindroid

135



ND

Laminar puzzle forms

Mt9

136


Ad (nd) Woody/herbaceous basal angiosperms and eudicotyledons

ND

Structured puzzle forms

Mt10

ND

Globular spherical rugulose

Mt4

ND

Other Globular

ND ND

Guard and subsidiary cells Stomatal complex

ND

Anticlinal epidermis silicified non-grass (reniform)

ND

Thin epidermal polygonal cells

ND

Hair bases

ND ND ND ND ND

Multi-layered trichome Tracheids with helical thickkenings cylindric scalariform or scrobiculate Cylindroid bulbous Cylindroid columellate Short prismatic elongated with smooth contour Prismatic elongated with smooth contour Other prismatic (elongate, blocky and tabular Other prismatic (elongate, blocky and tabular Other prismatic (elongate, blocky and tabular Other prismatic (elongate, blocky and tabular Other prismatic (elongate, blocky and tabular Other prismatic (elongate, blocky and tabular Other prismatic (elongate, blocky and tabular Other prismatic (elongate, blocky and tabular Other prismatic (elongate, blocky and tabular Other prismatic (elongate, blocky and tabular Other prismatic (elongate, blocky and tabular Other prismatic (elongate, blocky and tabular

94

Marantaceae / Woody/herbaceous basal angiosperms and eudicotyledons.

105

Marantaceae / Woody/herbaceous basal angiosperms and eudicotyledons. Woody/herbaceous basal angiosperms and eudicotyledons, conifers, ferns, etc. Grasses and other monocotyledons, conifers.

A (arh) Marantaceae / Woody/herbaceous basal angiosperms and eudicotyledons A (arh) Marantaceae / Woody/herbaceous basal angiosperms and eudicotyledons Ad (nd) Woody/herbaceous basal angiosperms and eudicotyledons, conifers, ferns, etc. Mo (nd) Grasses and other monocotyledons

140 114 117 118 119

Woody/herbaceous basal angiosperms and eudicotyledons, conifers, ferns, etc. Woody/herbaceous basal angiosperms and eudicotyledons, conifers, ferns, etc. Woody/herbaceous basal angiosperms and eudicotyledons, conifers, ferns, etc. Woody/herbaceous basal angiosperms and eudicotyledons, conifers, ferns, etc. Various plants and unknown. Various plants and unknown. Various plants and unknown. Various plants and unknown.

Nd (nd) Woody/herbaceous basal angiosperms and eudicotyledons, conifers, ferns, etc. Nd (nd) Woody/herbaceous basal angiosperms and eudicotyledons, conifers, ferns, etc. Nd (nd) Woody/herbaceous basal angiosperms and eudicotyledons, conifers, ferns, etc. Nd (nd) Woody/herbaceous basal angiosperms and eudicotyledons, conifers, ferns, etc. ND (nd) Various plants and unknown ND (nd) Various plants and unknown ND (nd) Various plants and unknown ND (nd) Various plants and unknown

143


ND (nd) Various plants and unknown

ND

144



ND

145



ND

146



ND

147



ND

148



ND

149



ND

150



ND

151



ND

152



ND

153



ND

154



ND

155



ND

156



ND

141 142 137

Basal angiosperms and eudicotyledons, conifers, ferns, etc.

138 139

Various plants and unknown

Mt7

Mt11

Mt13

Mt22 Mt23

Various plants and unknown

ND (nd)

157



ND

158



ND

159



ND

160



ND

161



ND

162



ND

163



ND

164



ND

165



ND

166



ND

167



ND

168



ND

169



ND

170



ND

171



ND

172 205 206 207

Various plants and unknown. ND ND ND

ND (nd) Various plants and unknown ND (nd) ND (nd) ND (nd)

ND ND ND ND

208 209 210 211 212 213 214 215 216 217 218 219 220

ND ND ND ND ND ND ND ND ND ND ND ND ND

ND (nd) ND (nd) ND (nd) ND (nd) ND (nd) ND (nd) ND (nd) ND (nd) ND (nd) ND (nd) ND (nd) ND (nd) ND (nd)

ND ND ND ND ND ND ND ND ND ND ND ND ND

Other prismatic (elongate, blocky and tabular Other prismatic (elongate, blocky and tabular Other prismatic (elongate, blocky and tabular Other prismatic (elongate, blocky and tabular Other prismatic (elongate, blocky and tabular Other prismatic (elongate, blocky and tabular Other prismatic (elongate, blocky and tabular Other prismatic (elongate, blocky and tabular Other prismatic (elongate, blocky and tabular Other prismatic (elongate, blocky and tabular Other prismatic (elongate, blocky and tabular Other prismatic (elongate, blocky and tabular Other prismatic (elongate, blocky and tabular Other prismatic (elongate, blocky and tabular Other prismatic (elongate, blocky and tabular Other prismatic (elongate, blocky and tabular Blocky irregula Tabular irregular Aggregated Laminar with different contour and verrugate or echinate surface Triangular Irregular isodiametric Blocky paralellepipedal Single-cell, fluid-conducting tissue Parallelepipedal to cubic, granulate Cubic, smooth parallelepipedal, pinched-point Cylindric, laminate Polyhedral plates Orbicular, stellate inside Stalked granulate body Orbicular plano-convex verrucose

Mt33 Mt35 Mt37

Sample Morphotypes Mt1 Mt2 Mt3 Mt4 Mt5 Mt6 Mt7 Mt8 Mt9 Mt10 Mt11 Mt12 Mt13 Mt14 Mt15 Mt16 Mt17 Mt18 Mt19 Mt20 Mt21 Mt22 Mt23 Mt24 Mt25 Mt26 Mt27 Mt28 Mt29 Mt30 Mt31 Mt32 Mt33 Mt34 Mt35 Mt36 Mt37 Others Non Id.

BCH531 34,92 2,27 1,65 12,81 0,41 0,00 3,72 0,00 5,99 0,00 0,00 0,41 1,86 3,93 0,00 1,03 0,00 0,00 0,00 0,21 0,62 2,69 10,74 1,45 2,48 2,07 0,00 0,62 3,10 0,00 0,00 0,00 1,65 0,00 0,00 2,07 0,00 2,48 0,83

BCH532 23,62 4,60 1,23 3,68 1,23 0,00 9,20 0,00 12,27 0,00 0,00 1,53 3,68 2,15 0,00 0,31 3,07 1,53 1,23 0,00 0,92 0,00 10,74 2,45 2,76 0,00 0,00 2,15 7,67 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 2,45 1,53

BCH533 4,17 3,89 0,28 11,11 4,17 0,56 16,67 0,00 11,94 0,00 5,56 2,22 1,94 0,00 0,00 5,83 1,39 2,50 1,39 0,00 0,00 5,56 5,28 0,83 2,78 0,56 0,00 2,78 2,78 0,00 0,00 0,00 1,11 0,00 0,00 1,11 0,00 1,39 2,22

Las Flores Formation BCH534 BCH535 6,32 5,26 0,00 8,42 0,42 3,79 14,32 0,00 10,74 0,00 1,26 2,53 1,26 0,00 0,00 1,68 3,58 1,89 1,68 0,00 0,00 0,00 10,53 6,95 0,00 0,63 0,00 4,42 9,05 0,00 0,00 0,00 0,42 0,00 0,00 0,00 0,00 2,95 1,89

8,67 2,31 0,00 5,20 1,16 0,00 16,76 0,00 5,78 0,00 0,00 0,00 1,73 0,00 0,00 0,58 8,67 4,62 1,16 0,00 0,00 0,00 16,18 6,36 0,00 1,73 0,00 4,05 11,56 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 2,60 0,87

BCH536 22,64 7,55 0,00 5,28 0,00 0,00 6,04 0,00 6,04 0,00 0,00 3,77 0,00 0,00 0,00 0,75 5,66 3,02 3,02 0,00 0,00 0,00 10,57 3,77 0,00 0,00 0,00 7,55 6,04 0,00 0,00 0,00 3,02 0,00 0,00 0,00 0,00 3,77 1,51

n BCH537 12,68 2,82 0,00 6,76 2,25 0,00 9,86 0,00 8,45 0,00 0,00 2,82 2,82 0,00 0,00 1,41 2,82 3,38 0,00 0,00 0,00 0,00 16,90 2,82 2,25 0,00 0,00 5,35 9,86 0,00 0,00 0,00 1,97 0,00 0,00 0,00 0,00 3,10 1,69

BCH538 10,47 4,36 0,00 10,47 1,22 3,84 3,14 0,00 9,77 0,00 0,00 3,14 0,00 0,00 0,00 1,40 6,98 8,73 6,98 0,35 0,00 0,00 13,96 0,00 0,00 0,00 0,00 1,40 10,12 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 2,44 1,22

LF-539

LF-540 18,72 8,51 4,26 0,00 2,55 1,70 2,55 0,00 5,11 0,00 2,55 0,00 0,00 0,00 0,00 4,26 0,00 0,00 0,00 0,00 1,70 4,26 15,32 0,00 2,55 0,00 0,00 4,26 10,21 0,00 0,00 4,26 0,00 0,00 0,00 0,00 0,00 5,11 2,13

LF-541 7,63 3,82 2,29 0,00 0,00 0,00 3,82 0,00 6,11 0,00 0,00 1,53 1,53 0,00 0,00 0,00 0,00 2,29 0,00 1,53 0,00 9,16 38,17 0,00 6,11 0,00 0,00 2,29 7,63 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 3,05 3,05

LF-542 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00

LF-543 19,72 8,45 11,27 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 14,08 5,63 0,00 0,00 0,00 0,00 0,00 14,08 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 19,72 7,04

10,45 1,49 7,46 0,00 0,00 0,00 11,94 0,00 7,46 0,00 0,00 2,99 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 1,49 8,96 10,45 0,00 7,46 0,00 0,00 5,97 0,00 0,00 0,00 2,99 0,00 0,00 7,46 0,00 0,00 8,96 4,48

LF-544

LF-545 14,49 4,83 3,86 3,86 0,00 0,00 10,63 0,00 8,70 0,00 0,00 1,93 1,93 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 10,63 13,53 0,00 5,80 0,00 0,00 4,83 3,86 0,00 0,00 0,00 0,00 0,00 2,90 0,00 0,00 4,35 3,86

Koluel Kaike Formation, Las Flores profile LF-547 LF-548 LF-549 LF-550

LF-546 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00

0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00

14,93 6,97 2,99 1,99 0,00 0,00 6,97 0,00 8,96 0,00 0,00 1,99 0,00 0,00 0,00 0,00 4,98 0,00 0,00 4,98 4,98 2,99 11,94 0,00 0,00 0,00 0,00 5,97 5,97 0,00 0,00 0,00 0,00 0,00 4,98 0,00 0,00 5,47 2,99

0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00

0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00

9,09 0,00 3,90 3,90 0,00 0,00 9,09 0,00 12,99 0,00 3,90 3,90 0,00 0,00 0,00 0,00 19,48 0,00 0,00 0,00 0,00 0,00 5,19 2,60 0,00 0,00 0,00 12,99 6,49 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 4,55 1,95

ofile LF-551

LF-552 12,50 5,00 0,00 0,00 0,00 0,00 7,50 0,00 12,50 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 5,00 0,00 25,00 0,00 0,00 0,00 0,00 0,00 5,00 0,00 0,00 0,00 0,00 0,00 12,50 0,00 0,00 10,00 5,00

LF-553 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00

LF-554 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00

LF-555 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00

LF-556 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00

LF-557 13,16 2,63 0,00 0,00 0,00 0,00 22,37 0,00 6,58 0,00 0,00 3,95 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 6,58 3,95 19,74 0,00 1,32 0,00 0,00 10,53 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 5,26 3,95

18,32 4,58 0,00 0,00 0,00 0,00 15,27 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 10,69 4,58 12,21 0,00 6,11 0,00 0,00 7,63 7,63 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 6,87 6,11

CL-1

Koluel Kaike Formation, Cañadón Lobo profile CL-2 CL-3 CL-4 CL-5 12,11 4,15 2,08 0,00 1,38 0,00 2,08 0,00 4,84 0,00 0,69 3,46 2,08 0,00 0,00 1,38 0,00 0,00 0,00 0,69 0,69 10,38 17,65 0,69 0,00 0,00 0,00 6,23 17,65 0,00 0,00 0,69 0,00 0,00 0,00 0,00 0,00 6,23 4,84

10,79 1,75 2,33 0,00 0,00 2,04 2,33 0,00 5,83 0,00 0,87 2,33 1,75 0,00 0,00 8,75 0,00 0,00 0,00 1,17 1,17 9,33 13,41 0,00 1,75 0,00 0,00 0,58 20,12 0,58 0,00 0,00 0,00 0,58 0,00 0,00 0,00 9,04 3,50

18,61 4,17 2,22 0,00 0,00 0,56 0,00 0,00 3,06 0,00 1,11 2,22 1,67 0,56 0,00 2,50 0,00 0,00 0,00 1,67 2,22 12,22 2,22 9,72 0,00 8,33 0,00 0,56 16,67 2,22 0,00 0,56 0,00 0,00 0,00 0,00 0,00 5,00 1,94

22,73 2,60 3,90 0,00 0,00 0,00 0,00 0,00 0,00 0,00 4,55 5,19 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 2,60 14,29 9,09 0,00 1,30 1,30 0,00 2,60 23,38 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 1,30 5,19

MMZ-1.0 29,72 0,00 1,75 0,00 1,40 0,35 0,35 0,00 0,00 0,00 2,80 2,10 1,75 0,00 0,00 0,00 0,00 0,00 0,00 3,85 1,75 9,44 9,44 0,00 0,35 1,40 0,00 1,05 15,38 1,40 0,00 0,00 0,00 0,35 0,00 4,55 0,00 5,94 4,90

13,77 1,27 1,27 0,00 2,54 0,00 0,00 0,00 9,32 0,00 0,00 2,33 0,00 0,00 0,00 2,33 0,00 0,00 0,00 0,85 0,85 16,74 20,34 0,21 1,06 0,64 0,00 1,69 11,02 0,00 0,00 0,00 0,00 0,21 0,00 0,00 0,00 7,63 5,93

MMZ-1.5 7,64 1,55 1,55 0,00 1,43 0,48 0,00 0,00 19,33 0,00 0,00 2,63 0,12 0,00 0,00 5,37 0,00 0,00 0,00 0,36 2,63 11,69 17,66 0,84 1,67 0,72 0,00 0,72 8,71 0,48 0,00 0,00 0,00 0,36 0,00 0,00 0,00 5,61 8,47

MMZ-2.0 22,85 7,34 2,31 0,00 1,47 0,00 5,45 0,00 11,53 0,00 0,00 2,94 1,05 0,00 0,00 3,77 0,00 0,00 0,00 0,63 2,94 5,87 10,48 0,42 0,63 0,00 0,00 0,21 6,71 0,00 0,00 0,00 8,60 0,21 0,00 0,00 0,00 4,19 0,42

MMZ-2.5 27,13 11,49 1,78 0,00 0,00 0,59 0,99 0,00 3,56 0,00 0,00 3,17 0,40 0,00 0,00 0,79 0,00 0,00 0,00 1,98 3,56 4,95 11,49 0,99 1,19 0,79 0,00 0,40 14,26 0,79 0,00 0,40 2,77 0,00 0,40 0,00 0,00 4,95 1,19

Sarmiento Formation (Gran Barranca Member), Gran Barranca profile MMZ-3.0 MMZ-3.3 MMZ-3.5 MMZ-4.0 MMZ-5.0 22,66 9,56 1,25 0,00 0,83 0,00 1,46 0,00 2,29 0,00 0,00 1,04 0,00 0,21 0,00 1,46 0,00 0,00 0,00 0,21 2,29 7,07 8,11 0,42 0,62 0,42 0,00 0,83 18,92 0,62 0,00 0,83 7,69 0,21 0,83 0,00 3,33 6,03 0,83

14,09 8,05 2,01 0,00 0,00 1,34 8,05 0,00 4,70 0,00 0,00 5,70 0,00 0,00 0,00 2,01 0,34 0,00 0,00 2,01 4,03 4,36 11,07 0,00 1,01 1,68 0,00 1,01 12,42 0,67 0,00 1,68 3,36 1,34 0,34 0,00 0,00 8,05 0,67

15,73 10,49 0,70 0,00 0,35 0,00 12,24 0,00 1,05 0,00 0,00 2,80 0,00 0,70 0,00 0,35 0,00 0,00 0,00 4,55 5,94 5,59 8,39 1,75 1,05 0,70 0,00 0,70 10,84 1,75 0,00 0,00 4,55 1,40 0,00 0,70 0,35 5,59 1,75

11,40 6,18 1,43 0,00 0,71 0,00 13,78 0,00 2,85 0,00 0,00 2,14 0,00 0,00 0,00 7,13 5,94 0,00 0,00 1,19 1,43 7,13 8,79 0,24 0,00 0,00 0,00 0,24 14,49 0,48 0,00 0,48 4,04 0,48 0,24 0,00 0,00 4,99 4,28

14,39 8,12 1,86 0,00 3,25 0,23 8,12 0,00 2,09 0,00 0,00 1,86 0,00 0,00 0,00 1,62 2,78 0,00 0,00 3,25 3,71 7,89 8,12 0,23 0,00 0,00 0,00 1,62 17,40 0,00 0,00 0,93 4,18 0,00 1,16 0,00 0,00 5,10 2,09

er), Gran Barranca profile MMZ-6.0 11,68 6,79 0,00 0,00 0,54 0,00 1,63 0,00 2,17 0,00 0,00 2,72 0,00 0,00 0,00 5,16 0,54 0,00 0,00 10,05 6,52 4,62 7,61 0,82 0,82 0,00 0,00 0,82 8,97 0,82 0,00 0,54 3,26 0,82 0,82 0,00 4,08 11,96 6,25

MMZ-6.5 14,29 6,07 0,36 0,00 0,36 0,00 1,79 0,00 1,79 0,00 0,00 2,14 0,00 0,00 0,00 2,86 0,00 0,00 0,00 3,57 1,79 7,50 12,14 0,71 0,71 0,36 0,00 1,43 23,57 1,07 0,00 0,00 3,57 0,36 0,36 0,00 0,00 8,93 4,29

MMZ-7.0 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00

MMZ-7.5 18,93 0,00 0,00 0,00 1,87 1,40 0,70 0,00 3,50 0,00 0,00 0,47 0,23 0,00 0,00 1,17 0,23 0,00 0,00 1,17 0,23 6,78 9,11 0,00 0,23 1,17 0,00 0,47 34,11 0,00 0,00 2,34 0,00 0,00 0,00 0,70 0,93 12,38 1,87

MMZ-8.0 8,58 3,86 0,00 0,00 0,00 5,15 14,16 0,00 1,72 0,00 0,00 1,29 0,00 0,00 0,00 1,29 0,00 0,00 0,00 0,86 0,00 4,29 13,30 0,00 0,00 0,43 0,00 0,00 27,90 0,86 0,00 0,86 2,58 0,00 0,00 0,00 0,00 11,16 1,72

Figure 2. Detailed box-plot of abundance of life forms in Koluel-Kaike Formation LF (PSG-LF1, PSG-LF2 and PSG-LF3) and CL (PSGCL) profiles.

For determining the vegetation palaeocommunity type, Palms phytoliths have been counted as a particular life-form, and then the relative abundance of palm phytoliths were compared to other plant indicators, basically with grasses or trees and shrubs morphotypes presence (Patterer, 2012). This method follows the methodological patterns of Couvreur et al. ( 2011), since palms are one of the most characteristic and ecologically important components of tropical rain forest worldwide; but they are also present in a broad range of environments, during mid-Cretaceous. These authors observed the differentiation of Nypoideae clade (mangrove swamp palms), while and during the Cretaceous/Paleocene, recorded the first appearance of the Coryphoideae clade (subtropical or non-rain-forest palms). In America, its extant species (Uhl et al., 1987; Henderson et al., 1999) grow in different environments such as deserts (Washingtonia ssp. for the Sonoran Desert), xeric environments (Tritrinax ssp. and Copernicia ssp. in the Chaco region), swamps (Mauritia ssp. northern South America), forests (with the abundance of species in the Amazon region), savannahs (Butia ssp. in southern Brazil, Uruguay and Argentina) and temperate forests (Jubaea chilensis (Molina) Baill. in Central Chile or Maule Forest). There are certain groups that may indicate the environment in which they develop (Tomlinson et al., 2011) and thus they can be applied in palaeopalynological and biogeographical studies (Rull, 1998, 1999, 2001). Phytolith morphotypes do not allow us to establish even this distinction. Couvreur, T.L.P., Forest, F. and Baker. W.J. 2011. Origin and global diversification patterns of tropical rain forests: inferences from a complete genus-level phylogeny of palms. BMC Biology 2011, 9:44. http://www.biomedcentral.com/1741-7007/9/44 Henderson, A., Galeno, G., and Bernal, R. 1999. Field guide to the palms of the Americas. Ed. Princeton, 352 p. Patterer, N.I. 2012. [Análisis fitolíticos de la Formación El Palmar (Pleistoceno tardío) en la Cuenca del río Uruguay (Este de Argentina). PhD Thesis, Universidad Nacional de Córdoba, Córdoba 307 p. Unpublished]. Rull, V. 1998. Biogeographical and evolutionary considerations of Mauritia (Arecaceae), based on palynological evidence. Review of Palaeobotany and Palynology 100: 109–122. Rull, V. 1999. A palynological record of a secondary succession after fire in the Gran Sabana, Venezuela. Journal of Quaternary Science 14: 137–152. Rull, V. 2001. A morphometric study of early Miocene Mauritiidites from Northern South America. Palaeoecological and evolutionary implications. Grana 40: 163–167. Tomlinson, P.B., Horn, J.W., and Fisher, J.B. 2011. The Anatomy of Palms. Arecaceae – Palmae. Oxford University Press, 251 p. Uhl, N.W., and Dransfield, J. 1987. Genera Palmarum: a classification of palms based on the work of Harold E. Moore. Allen Press, The L.H. Bailey

Diagnostic types:

diagnostic (D)

intrafamily diagnostic (ID)

non diagnostic (ND)

Botanical affinity:

A: Angiosperm

Apa: Arecaceae

Ad: Dicotyledonous

Mo: Monocotyledonous

Agr: Poaceae

Acy: Cyperaceae

Apo: Podostemaceae

ND: non-diagnostic

Life forms: (p) palms (ar) arborescent (arh) arborescent and/or herbs (hb) herbs and/or bamboos (h) herbs (nd) non-diagnostic Classifications synthesis of phytolith morphotypes in woody and herbaceous used for the Table 2. References: 1 Raigemborn, M., Brea, M., Zucol, A., and Matheos, S., 2009. Early Paleogene climatic conditions at mid latitude Southern Hemisphere: Mineralogical and paleobotanical proxies from continental sequences in Golfo San Jorge basin (Chubut, Patagonia, Argentina). Geologica Acta, Climate and Biota of the Early Paleogene, Special Volume 7: 125–145. 2 Brea, M., Zucol, A.F., Raigemborn, M.S., and Matheos, S., 2008. Reconstrucción de paleocomunidades arbóreas mediante análisis fitolíticos en sedimentos del Paleoceno superior –Eoceno? (Formación Las Flores), Chubut, Argentina. In: A. Korstanje and Babot, P. (Eds.), Matices Interdisciplinarios en Estudios Fitolíticos y de otros Microfósiles/ Interdisciplinary Nuances in Phytolith and other Microfossil Studies., BAR (British Archaeological Reports) International Series S1870, Chapter 9.Oxford, p. 91–108. 3 Zucol, A.F., Brea, M., and Bellosi, E.S., 2010. Phytolith studies in Gran Barranca (Central Patagonia, Argentina) focused in the Middle Eocene. In: R. Madden, A. Carlini, G. Vucetich and R. Kay (Eds.), The Paleontology of Gran Barranca: Evolution and Environmental Change through the Middle Cenozoic of Patagonia. Cambridge University Press, p. 313–336. 4 ICPNWG. Madella M, Alexandre A, Ball T. 2005. International Code for Phytolith Nomenclature 1.0. Annals of Botany 96: 253–260. 5 Stromberg, C.A.E., Dunn, R.E., Madden, R.H., Kohn, M.J., and Carlini, A.A., 2013. Decoupling the spread of grasslands from the evolution of grazer–type herbivores in South America. Nature Communications 4: 1478. doi:10.1038/ncomms2508. 6 Collura, L.V. and Neumann, K., 2016. Wood and bark phytoliths of West African woody plants. Quaternary International in press 1-18. http://dx.doi.org/10.1016/j.quaint.2015.12.070. 7 Neumann, K., Fahmy, A.G., Müller-Scheeßel, N., and Schmidt, M., 2016. Taxonomic, ecological and palaeoecological significance of leaf phytoliths in West African grasses. Quaternary International in press 1-18. http://dx.doi.org/10.1016/j.quaint.2015.11.039. 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26

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