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
1
EMERGENCE OF GRASSY HABITATS DURING THE GREENHOUSE–
2
ICEHOUSE SYSTEMS TRANSITION IN THE MIDDLE EOCENE OF
3
SOUTHERN SOUTH AMERICA.
4
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
7 8
ALEJANDRO F. ZUCOL1, J. MARCELO KRAUSE2, 3, MARIANA BREA1, M. SOL
9
RAIGEMBORN4 AND SERGIO D. MATHEOS4
10 11
1
12
Tecnología a la Producción, CICYTTP–CONICET, Dr. Materi y España S/N, E3105BWA
13
Diamante, Entre Ríos, Argentina.
[email protected];
[email protected]
14
2
15
Trelew, Chubut, Argentina.
[email protected]
16
3
17
Provincial 1 s/n, (9000) Comodoro Rivadavia, Chubut, Argentina.
18
4
19
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,
21 22
Total number of pages (text + references): 66
23
Figures: 10; Supplementary material (Figures: 2, Tables: 2; and supplementary references)
1
24
Proposed head: ZUCOL ET AL.: EOCENE PHYTOLITHS FROM PATAGONIA
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Corresponding author: Alejandro F. Zucol
26 27
Abstract. Phytolith assemblages recovered from the middle Eocene Koluel-Kaike
28
Formation in Central Patagonia, southern South America, confirm the presence of
29
herbaceous components, mainly related to mixed ecosystems or savannahs, and
30
subordinate forest communities. This new palaeobotanical information reveals a clear
31
transition between previously described woodland-dominated communities from the
32
early Eocene Las Flores Formation, and grass-rich ecosystems from the mid-Eocene
33
Sarmiento Formation. The early appearance of grass-dominated ecosystems in southern
34
South America, would be related to the worldwide greenhouse to icehouse systems
35
transition initiated during the mid-Palaeogene.
36
Key words. Phytoliths, palaeosols, Koluel-Kaike Formation, palaeoclimate, EECO,
37
Central Patagonia.
38 39
Resumen. APARICIÓN DE LOS ECOSISTEMAS DOMINADOS POR GRAMÍNEAS
40
DURANTE LA TRANSICIÓN CLIMÁTICA GREENHOUSE–ICEHOUSE EN EL
41
EOCENO MEDIO DE PATAGONIA CENTRAL. Asociaciones fitolíticas provenientes de
42
la Formación Koluel-Kaike (Eoceno medio) en Patagonia Central, sur de América del Sur,
43
confirman la presencia de componentes herbáceos, principalmente relacionados a
44
ecosistemas mixto o sabanas, y comunidades arbóreas subordinadas. Esta nueva
45
información paleobotánica revela una clara transición entre las comunidades previamente
46
descriptas en las Formación Las Flores (Eoceno inferior), dominada por componentes
47
leñosos, y la sección inferior de la Formación Sarmiento (Eoceno medio), caracterizada por
48
una mayor abundancia de gramíneas. La aparición temprana de ecosistemas dominados por
2
49
gramíneas en el sur de América del Sur, estaría relacionada a la transición climática global,
50
desde sistema Greenhouse a Icehouse, iniciada a partir del Paleógeno medio.
51
Palabras clave. Fitolitos, paleosuelos, Formación Koluel-Kaike, paleoclima, EECO,
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Patagonia Central.
53 54
PREVIOUS studies on phytolith content from early Palaeogene successions in Central
55
Patagonia have revealed the presence of forested communities during the early Eocene
56
Las Flores Formation (Brea et al., 2008; Raigemborn et al., 2009), and grass-rich
57
communities in the middle Eocene Gran Barranca Member (lower Sarmiento
58
Formation, Mazzoni, 1979; Zucol et al., 1999, 2004; Andreis, 2008; Zucol et al.,
59
2010a). Vegetation shift occurred in parallel sedimentary, environments and climatic
60
conditions changes. For instance, lacustrine and fluvial sedimentation, under tropical
61
conditions, occurred during the deposition of the Las Flores Formation (Raigemborn et
62
al., 2009, 2010, 2014; Woodburne et al., 2014); and dominant loessic deposition, under
63
semiarid-arid conditions, occurred during the deposition of the Gran Barranca Member,
64
Sarmiento Formation (Mazzoni, 1979; Spalletti and Mazzoni, 1979; Bellosi et al., 2010;
65
Bellosi and González, 2010; Bellosi and Krause, 2014).
66
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
68
between the underlying Las Flores and the overlying Sarmiento formations. This new
69
information fills the gap in the palaeofloristic record in the San Jorge Basin, Southern
70
South America, between the mid-Ypresian woodland-dominated Las Flores
71
communities, and the late Lutetian, grass-rich Sarmiento ones; and provides key
72
knowledge for understanding the causes of the origin of grasslands in South America, a
73
factor pertinent to regional palaeoecology and the evolution of mammals and feeding
3
74
strategies (e.g., Strömberg, et al. 2013; Dunn et al., 2015). Our aims are to: 1) analyse
75
the floristic composition and changes occurred during the deposition of the Koluel-
76
Kaike Formation, and 2) consider these results in the context of the different hypotheses
77
regarding the emergence of grasslands in South America and their relationship with the
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global palaeoclimate scenery.
79 80
Figure 1
81 82
GEOLOGICAL, SEDIMENTOLOGICAL AND
83
PALAEONTOLOGICAL SETTING
84
The Koluel-Kaike Formation is a continental unit cropping out in the San Jorge
85
Basin, Central Patagonia, Argentina (Fig. 1.2). In Gran Barranca area the Koluel-Kaike
86
Formation overlies, in probable paraconformity relationship, to the Las Flores
87
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;
89
Raigemborn et al., 2010). It composes of moderate to highly-silicified, silt-sized,
90
pyroclastic deposits, bentonites, and tuffs, mostly modified to palaeosols (Krause and
91
Bellosi, 2006; Krause et al., 2010; Raigemborn et al., 2010; Raigemborn et al., 2014).
92
The fossils record of this unit currently restrict to silicified wood, referred to
93
Taxaceoxylon katuatenkun Brea, Bellosi et Krause (Brea et al., 2009), and trace fossils
94
of cicadas (Feoichnus challa Krause et al., 2008a). The temporal span of the middle Las
95
Flores to uppermost Koluel-Kaike succession in the Gran Barranca area was
96
constrained, on the base of magnetostratigraphic and geochronological data, to be ca.
97
51.4–42.2 Ma (mid Ypresian–late Lutetian) in age, with the Koluel-Kaike being 46.7–
98
42.2 Ma (Lutetian) in age (Krause et al., 2017) (Fig. 1.1). 4
99
For this study, two localities were selected, Las Flores and Cañadón Lobo. The Las
100
Flores locality (LF) (S 45° 43' 30''; W 68° 37' 22'') locates in the eastern edge of Gran
101
Barranca, a classical Cenozoic land-mammal fossil locality (e.g., Ameghino, 1906;
102
Simpson, 1948; Marshall et al., 1983; Madden et al., 2010), ~40 km to the east-southeast
103
of Sarmiento city, south-central Chubut Province (Fig. 1.2). At LF the studied section is
104
~42 m-thick (Fig. 2), and characterizes by tephric loessic beds, small alluvial channels,
105
and shallow ponds (Krause et al., 2008b, 2010; Raigemborn et al., 2010, 2014). The
106
sedimentation in the Koluel-Kaike Formation would have developed under different
107
climates, with a clear temporal evolution from humid to subhumid-semiarid conditions,
108
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
110
lower section, suggest values of Mean Annual Precipitation (MAP) from 1200 to 1300
111
mm/yr, and Mean Annual Temperature (MAT) ~15° C (Krause et al., 2010), consistent
112
with wet and megathermal conditions. Upward the section, climofunctions applied on
113
weakly-developed palaeosols (e.g., palaeo-Andisols) give values of MAP ~1000 mm/yr
114
and MAT ~12° C) (Krause et al., 2010), consistent with megathermal–mesic conditions.
115
The change in clay mineralogy throughout the unit, from kaolinite-dominated
116
assemblages, in the lower section, to smectite-dominated associations in the middle–
117
upper ones, also indicates a climatic trend from humid to subhumid conditions, along
118
with pulses of relative stronger seasonality or less intense precipitations periods (Krause
119
et al., 2010; Raigemborn et al., 2014).
120
The Cañadón Lobo (CL) locality (S 46° 58' 24"; W 66° 50' 13") places in northern
121
Santa Cruz Province, ~15 km to the northwest of Punta Casamayor, in south border of the
122
San Jorge Basin (Fig. 1.2). At CL the base of the Koluel-Kaike Formation is not
123
exposed. The unit underlies to whitish, massive and tabular beds of the Sarmiento
5
124
Formation assigned to the Gran Barranca Member, and bearing sparse, fragmentary
125
vertebrate remains from the Casamayoran South American Land Mammal Age
126
(SALMA), fossil dung-beetle brood balls (Coprinisphaera) and Feoichnus specimens
127
(Pérez et al., 2012). The contact between the Koluel-Kaike and the Sarmiento
128
formations looks conformable (Pérez et al., 2012), although the absence of around 10–
129
15 m of whitish beds bearing weakly-developed palaeosols recognized in the upper
130
Koluel-Kaike section at LF (Krause et al., 2010), suggests the presence of a
131
paraconformity. The studied section is ca. 10 m-thick and mainly composes of massive,
132
silt-sized, pyroclastic deposits contained within tabular and lenticular beds, with
133
minority tabular, massive beds of mudstones (Figs. 2, 3.1–2). Lenticular beds display an
134
erosive base, evidenced by the presence of muddy intraclasts; while tabular beds display
135
sharp bases. Both types of beds lack in recognizable primary, sedimentary structures.
136
Pedogenetic features, among them, iron-rich, reticulate mottles, iron nodules,
137
slickensides, and rhizoliths are common within the tabular pyroclastic and muddy beds.
138
Iron nodules are also present within the lenticular beds. Based on these features, Krause
139
(2010) interpreted a palaeoenvironment characterized by small fluvial channels, cutting
140
across pyroclastic and muddy floodplains (Fig. 3.2). The presence of iron nodules
141
within the lenticular channel body indicates changes in water content in the original
142
deposit, suggesting a probable ephemeral nature of the channel. On the other side,
143
tabular beds composed of very fine sediments and bearing rhizoliths, slickensides, and
144
iron mottling and concretions, support the presence of pedogenetically modified,
145
floodplain deposits (Krause, 2010) (Fig. 3.3–4). Palaeosols at CL (Fig. 3.2–6)
146
displaying reddish, reticulate mottles are strongly similar in morphology to those
147
included in the lower Koluel-Kaike Formation at LF (e.g., Kápenk pedotype sensu
148
Krause et al., 2010), which characterize ironstones levels (Krause et al., 2010) (Fig.
6
149
3.5). Since ironstones of the Koluel-Kaike Formation occur typically in the lower
150
section at Las Flores, and are absent in adjacent units (Krause et al., 2010), we correlate
151
the CL section with the lower Koluel-Kaike Formation at LF.
152 153
Figure 2
154 155
THE EARLY PALAEOGENE OF CENTRAL PATAGONIA:
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PHYTOLITHS AND OTHERS PALAEOBOTANICAL RECORDS
157
The early Palaeocene–middle Eocene span of time in Gran Barranca and surrounding
158
areas is represented by five formations (Fig. 1.1): Salamanca (estuarine; Danian)
159
(Martínez, 1992; Clyde et al., 2014; Comer et al., 2015), Peñas Coloradas (fluvial; latest
160
Danian) (Raigemborn et al., 2010; Clyde et al., 2014), Las Flores (fluvial, lacustrine;
161
Ypresian) (Raigemborn et al., 2010; Woodburne et al., 2014; Kohn et al., 2015; Krause et
162
al., 2017), Koluel-Kaike (loessic, alluvial; Lutetian) (Krause et al., 2010, 2017;
163
Raigemborn et al., 2010), and lower Sarmiento (Cañadón Vaca and Gran Barranca
164
Members) (loessic, fluvial; late Lutetian–Bartonian) (Bellosi et al., 2010; Ré et al., 2010a,
165
b; Dunn et al., 2013; Bellosi and Krause, 2014).
166
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
168
agrees with others palaeobotanical records from the Salamanca Formation, such pollen, leaf
169
compression, permineralized woods and fruits (Berry, 1937; Romero, 1968; Archangelsky,
170
1973, 1976; Archangelsky and Zamaloa, 1986; Somoza et al., 1995; Brea et al., 2005a, b,
171
2008; Matheos et al., 2005; Iglesias et al., 2007, Jud et al., 2017; Ruiz et al., 2017, among
172
others).
7
173
Phytoliths have not been described for the Peñas Coloradas Formation, but
174
complementary palynomorphs and permineralized woods remains indicate the presence of
175
mixed subtropical to tropical and humid forests (Romero, 1973; Lema et al., 1999; Ruiz et
176
al., 1999; Brea and Zucol, 2006; Raigemborn et al., 2009).
177
The phytolith assemblages of the Las Flores Formation exhibits high abundance of dicot
178
and palm phytoliths, in association with scarce grasses (Poaceae), sedges, and
179
podostemoid elements, corresponding to subtropical to tropical lowland forests (Brea et
180
al., 2008; Raigemborn et al., 2009). Woody and palm indicators constitute 12.27−39.61%
181
and 8.05−37.19% respectively, with dicots (12.07–34.16%), diagnostic herbs (0–0.92%),
182
non-diagnostic herb/bamboos (8.06–23.09%) and humid herbs (0−4.13%) as understory
183
elements (Brea et al., 2008; Raigemborn et al., 2009).
184
The phytolith content from the Gran Barranca Member (lower Sarmiento Formation) at
185
Gran Barranca section reveals abundance of palms and up to 15% of herbaceous elements,
186
and a lower proportion of woody plant elements (Zucol et al., 2010a; Strömberg et al.,
187
2013; Dunn et al., 2015; Kohn et al., 2015). Zucol et al. (2010a), interpreted this
188
association as the record of savannah palaeocommunities, composed of palms
189
(9.19−38.61%), diagnostic herbs (1.40–18.48%), non-diagnostic herb/bamboo
190
(10.90−35.28%), humid herbs (0−12.38%), dicots (3.57–19.33%), and woody elements
191
(1.39−15.20%), and with a clear increase trend of megathermal components (from C4
192
grass) upward the unit. Thus, the lower Gran Barranca Member characterizes by the
193
highest abundances of palms (30.19−38.61%), which then decrease into the upper levels
194
to the 12.44−22.50% range. In general, the woody elements have low abundances
195
(1.39−9.74%) in the whole section, with the exception of one sample recorded in the
196
middle-upper section (15.20% in the MMZ9504.0 sample, Zucol et al., 2010a).
197
Diagnostic types of grass herbs and non-diagnostic herb/bamboo types respectively 8
198
increase their abundance from 3.77–6.73% and 10.90–20.00%, in the lower section, to
199
6.78−18.48% and 14.13–28.57%, in the upper one. This increase relates to the high
200
abundance of panicoid phytoliths. Finally, upward the Gran Barranca Member, the
201
floristic composition recordes an increase in humid herbs, which generally present in
202
low percentages (0−3.43%) throughout the section, rising up to 12.38% in the upper
203
section (sample MMZ9507.5; Zucol et al., 2010a).
204
However, in spite of similar values of percentage among involved morphotypes
205
obtained for the Gran Barranca Member at type locality obtained by Strömberg et al.
206
(2013), Dunn et al. (2015), and Kohn et al. (2015), these authors do not share the
207
interpretation of a savannah environment. For instance Strömberg et al. (2013), on the
208
base of the abundance of palms, suggested an ecological scenery characterized by a
209
closed–humid subtropical forest, an interpretation subsequently modified in Dunn et al.
210
(2015). These authors, applying the leaf area index (LAI) proxy, concluded that the
211
declination of the rLAI values, from the middle Eocene to the early Oligocene in the
212
San Jorge Basin, indicating an opening of the landscape from dense vegetation through
213
progressively more open vegetation, that is a nearly grass-free scrubland, an
214
interpretation kept by Kohn et al. (2015).
215 216 217
Figure 3
MATERIALS AND METHODS Twenty-four sedimentary samples (Fig. 2) were obtained from the Koluel-Kaike
218
Formation for phytolith analysis: 19 from the LF locality (LF539–557), and 5 from the CL
219
locality (CL1–CL5). Nine of these samples (LF541, LF545–546, LF548–549, LF552–555)
220
provided an insufficient phytolith content to obtain a minimal representative sample and
221
thus inadequate for their statistical analysis.
9
222
Each sample consisted of 20 grams of sediment, which was processed following a
223
protocol employing various chemical agents to remove soluble salts, organic matter and
224
carbonates (Zucol et al., 2010b). Samples were washed and soluble salts dissolved with
225
distilled water; carbonate material was removed with dilute HCl; organic material was
226
removed by adding H2O2 followed by deflocculation by Sodium hexametaphosphate. The
227
fractionation was quantified through sieving (coarse fraction) and sedimentation (fine
228
fractions). Four fractions were obtained according to particle diameter: coarse (>250 µm),
229
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
Woody/herbaceous basal angiosperms and Ad (arh) eudicotyledons.
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
Woody/herbaceous basal angiosperms and Ad (arh) eudicotyledons.
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
Woody/herbaceous basal angiosperms and Ad (arh) eudicotyledons.
ND
97
Woody/herbaceous basal angiosperms and Ad (arh) eudicotyledons.
ND
98
Woody/herbaceous basal angiosperms and Ad (arh) eudicotyledons.
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
Woody/herbaceous basal angiosperms and Ad (arh) eudicotyledons.
D
111
Woody/herbaceous basal angiosperms and Ad (ar) eudicotyledons.
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).
Woody/herbaceous basal angiosperms and Ad (ar) eudicotyledons.
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.
Woody/herbaceous basal angiosperms and Ad (ar) eudicotyledons.
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)
A. Sclerenchyma / A.2. Sclereid / Sclereid pitted p.p.
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
Various plants and unknown.
ND (nd)
ND
cylindric scalariform or scrobiculate
118
Various plants and unknown.
ND (nd)
ND
Cylindroid bulbous
13. Cylindroid bulbous. Cylindroid/renate with a psilate texture and irregular bulbuous enlargements.
119
Various plants and unknown.
ND (nd)
ND
Cylindroid columellate
14. Cylindroid columellate. Very large cylindroid with surface covered by processes.
120
Woody/herbaceous basal angiosperms and Ad (arh) eudicotyledons.
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
Woody/herbaceous basal angiosperms and Ad (arh) eudicotyledons.
122
Woody/herbaceous basal angiosperms and Ad (ar) eudicotyledons.
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
Woody basal angiosperms and eudicotyledons.
Ad (ar)
ND
Sclereid pitted
A. Sclerenchyma / A.2. Sclereid / Sclereid pitted p.p.
125
Woody basal angiosperms and eudicotyledons.
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
Woody/herbaceous basal angiosperms and Ad (arh) eudicotyledons.
ND
130
Woody/herbaceous basal angiosperms and Ad (arh) eudicotyledons.
ND
131
Woody basal angiosperms and eudicotyledons.
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
Woody basal angiosperms and eudicotyledons.
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
Woody basal angiosperms and eudicotyledons.
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
Woody basal angiosperms and eudicotyledons.
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
Woody/herbaceous basal angiosperms and Ad (arh) eudicotyledons.
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
Woody/herbaceous basal angiosperms and Nd (nd) eudicotyledons, conifers, ferns, etc.
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
Woody/herbaceous basal angiosperms and Nd (nd) eudicotyledons, conifers, ferns, etc.
ND
140
Woody/herbaceous basal angiosperms and Nd (nd) eudicotyledons, conifers, ferns, etc.
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
Various plants and unknown.
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
Various plants and unknown.
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
Various plants and unknown.
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
Various plants and unknown.
Nd (nd)
ND
D. Elongate. Elongate vesicular. Elongate, thin, with large, spherical to elliptic concavities, only in wood of Capparis tomentosa.
147
Various plants and unknown.
Nd (nd)
ND
D. Elongate. Elongate with globular part(s). Cylindroid, with spherical to elliptic central or terminal swellings.
148
Various plants and unknown.
Nd (nd)
ND
D. Elongate. Elongate irregular. Elongate, high variability in shape and size.
149
Various plants and unknown.
Nd (nd)
ND
150
Various plants and unknown.
Nd (nd)
ND
151
Various plants and unknown.
Nd (nd)
ND
3. Blocky corniculate. Ovate, rectangular, polygonal or irregular block with corniculate projections. Variable textures.
152
Various plants and unknown.
Nd (nd)
ND
5. Blocky hairy. Ovate, rectangle, polygonal or irregular block with irregular/scrobiculate texture; long pointed projections.
153
Various plants and unknown.
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
Various plants and unknown.
Nd (nd)
ND
7. Blocky polygonal. Polygonal/irregular block with a psilate/scrobiculate texture.
155
Various plants and unknown.
Nd (nd)
ND
8. Blocky radiating. Orbicular to rectangular block with a surface covered by concentric layers.
156
Various plants and unknown.
Nd (nd)
ND
157
Various plants and unknown.
Nd (nd)
ND
158
Various plants and unknown.
Nd (nd)
ND
159
Various plants and unknown.
Nd (nd)
ND
160
Various plants and unknown.
Nd (nd)
ND
161
Various plants and unknown.
Nd (nd)
ND
162
Various plants and unknown.
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
Various plants and unknown.
Nd (nd)
ND
164
Various plants and unknown.
Nd (nd)
ND
165
Various plants and unknown.
Nd (nd)
ND
50. Tabular scrobiculate. Variably wide parallelepiped with an elongate outline and scrobiculate texture.
166
Various plants and unknown.
Nd (nd)
ND
51. Tabular thick contorted. Thick parallelepiped with a contorted profile and irregular to laminate texture.
167
Various plants and unknown.
Nd (nd)
ND
52. Tabular thick dendritic. Large elongate parallelepiped with psilate texture and sides covered by irregular dedritic processes.
168
Various plants and unknown.
Nd (nd)
ND
53. Tabular thick lacunate. Large thick parallelepiped with scrobiculate/lacunose texture.
169
Various plants and unknown.
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
Various plants and unknown.
Nd (nd)
ND
55. Tabular thin. Thin parallelepiped, square to rectangular shape with psilate texture.
171
Various plants and unknown.
Nd (nd)
ND
56. Tabular thin pilate. Thin parallelepiped, scrobiculate or lacunate texture and rodlike processes.
172
Various plants and unknown.
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
Woody basal angiosperms and eudicotyledons.
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
Woody basal angiosperms and eudicotyledons.
Ad (ar)
ND
Cork aerenchyma
C. Cork aerenchyma. Cork aerenchyma. Tabular silicified tissue, various shapes, with irregular perforations of various size.
204
Woody basal angiosperms and eudicotyledons.
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
Other parallelepipedal bulliform types
42
Poaceae
Agr (hb) Poaceae
D
Other parallelepipedal bulliform types
43
Poaceae
Agr (hb) Poaceae
D
Other parallelepipedal bulliform types
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
Woody basal angiosperms and eudicotyledons.
Ad (ar) Woody basal angiosperms and eudicotyledons
ND
132
Woody basal angiosperms and eudicotyledons.
Ad (ar) Woody basal angiosperms and eudicotyledons
ND
133
Woody basal angiosperms and eudicotyledons.
Ad (ar) Woody basal angiosperms and eudicotyledons
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
Woody basal angiosperms and eudicotyledons.
Ad (ar) Woody basal angiosperms and eudicotyledons
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
Woody / herbaceous basal angiosperms and eudicotyledons.
Ad (arh) Woody / herbaceous basal angiosperms and eudicotyledons
ND
Globular elipsoidal smooth
Mt5
93
Woody / herbaceous basal angiosperms and eudicotyledons.
Ad (arh) Woody / herbaceous basal angiosperms and eudicotyledons
ND
Globular spherical smooth type 2
Mt6
96
Woody / herbaceous basal angiosperms and eudicotyledons.
Ad (arh) Woody / herbaceous basal angiosperms and eudicotyledons
ND
Globular granulate
97
Woody / herbaceous basal angiosperms and eudicotyledons.
Ad (arh) Woody / herbaceous basal angiosperms and eudicotyledons
ND
Globular granulate
98
Woody / herbaceous basal angiosperms and eudicotyledons.
Ad (arh) Woody / herbaceous basal angiosperms and eudicotyledons
ND
Globular granulate
104
Woody / herbaceous basal angiosperms and eudicotyledons.
Ad (arh) Woody / herbaceous basal angiosperms and eudicotyledons
ND
Other Globular
109
Woody / herbaceous basal angiosperms and eudicotyledons.
Ad (arh) Woody / herbaceous basal angiosperms and eudicotyledons
ND
Other Globular
110
Woody / herbaceous basal angiosperms and eudicotyledons.
Ad (arh) Woody / herbaceous basal angiosperms and eudicotyledons
ND
Faceted isodiametric
Woody / herbaceous basal angiosperms and eudicotyledons.
Ad (arh) Woody / herbaceous basal angiosperms and eudicotyledons
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
Woody / herbaceous basal angiosperms and eudicotyledons.
Ad (arh) Woody / herbaceous basal angiosperms and eudicotyledons
ND
Other faceted
120
Woody / herbaceous basal angiosperms and eudicotyledons.
Ad (arh) Woody / herbaceous basal angiosperms and eudicotyledons
ND
121
Woody / herbaceous basal angiosperms and eudicotyledons.
Ad (arh) Woody / herbaceous basal angiosperms and eudicotyledons
ND
Irregular sulcate or facetate vascular cell Favose aggregate sclerenchyma and similar
126
Woody / herbaceous basal angiosperms and eudicotyledons.
Ad (arh) Woody / herbaceous basal angiosperms and eudicotyledons
ND
Cylindroid
127
Woody / herbaceous basal angiosperms and eudicotyledons.
Ad (arh) Woody / herbaceous basal angiosperms and eudicotyledons
ND
Cylindroid
128
Woody / herbaceous basal angiosperms and eudicotyledons.
Ad (arh) Woody / herbaceous basal angiosperms and eudicotyledons
ND
Cylindroid
129
Woody / herbaceous basal angiosperms and eudicotyledons.
Ad (arh) Woody / herbaceous basal angiosperms and eudicotyledons
ND
Cylindroid
130
Woody / herbaceous basal angiosperms and eudicotyledons.
Ad (arh) Woody / herbaceous basal angiosperms and eudicotyledons
ND
Cylindroid
135
Woody / herbaceous basal angiosperms and eudicotyledons.
Ad (arh) Woody / herbaceous basal angiosperms and eudicotyledons
ND
Laminar puzzle forms
Mt9
136
Woody / herbaceous basal angiosperms and eudicotyledons.
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
Various plants and unknown.
ND (nd) Various plants and unknown
ND
144
Various plants and unknown.
ND (nd) Various plants and unknown
ND
145
Various plants and unknown.
ND (nd) Various plants and unknown
ND
146
Various plants and unknown.
ND (nd) Various plants and unknown
ND
147
Various plants and unknown.
ND (nd) Various plants and unknown
ND
148
Various plants and unknown.
ND (nd) Various plants and unknown
ND
149
Various plants and unknown.
ND (nd) Various plants and unknown
ND
150
Various plants and unknown.
ND (nd) Various plants and unknown
ND
151
Various plants and unknown.
ND (nd) Various plants and unknown
ND
152
Various plants and unknown.
ND (nd) Various plants and unknown
ND
153
Various plants and unknown.
ND (nd) Various plants and unknown
ND
154
Various plants and unknown.
ND (nd) Various plants and unknown
ND
155
Various plants and unknown.
ND (nd) Various plants and unknown
ND
156
Various plants and unknown.
ND (nd) Various plants and unknown
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
Various plants and unknown.
ND (nd) Various plants and unknown
ND
158
Various plants and unknown.
ND (nd) Various plants and unknown
ND
159
Various plants and unknown.
ND (nd) Various plants and unknown
ND
160
Various plants and unknown.
ND (nd) Various plants and unknown
ND
161
Various plants and unknown.
ND (nd) Various plants and unknown
ND
162
Various plants and unknown.
ND (nd) Various plants and unknown
ND
163
Various plants and unknown.
ND (nd) Various plants and unknown
ND
164
Various plants and unknown.
ND (nd) Various plants and unknown
ND
165
Various plants and unknown.
ND (nd) Various plants and unknown
ND
166
Various plants and unknown.
ND (nd) Various plants and unknown
ND
167
Various plants and unknown.
ND (nd) Various plants and unknown
ND
168
Various plants and unknown.
ND (nd) Various plants and unknown
ND
169
Various plants and unknown.
ND (nd) Various plants and unknown
ND
170
Various plants and unknown.
ND (nd) Various plants and unknown
ND
171
Various plants and unknown.
ND (nd) Various plants and unknown
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
Mercader, J., Bennett, T., Esselmont, C., Simpson, S. and Walde, D., 2009. Phytoliths in woody plants from the Miombo woodlands of Mozambique. Annals of Botany 104: 91-113. Mercader, J., Astudillo, F., Barkworth, M., Bennett, T., Esselmont, C., Kinyanjui, R., Grossman, D.L., Simpson, S., and Walde, D., 2010. Poaceae and Cyperaceae phytoliths from the woodlands of Niassa Rift, Mozambique. Journal of Archaeological Science 37: 1953–1967. Novello, A., D. Barboni, L. Berti-Equille, J.-C. Mazur, P. Poilecot, P. Vignaud. 2012. Phytolith signal of aquatic plants and soils in Chad, Central Africa. Review of Palaeobotany and Palynology 178 (2012) 43–58. Dickau, R., B.S. Whitney, J. Iriarte, F.E. Mayle, J. D. Soto, P. Metcalfe, F. A. Street-Perrott, N. J. Loader, K. J. Ficken, T.J. Killeen. 2013. Differentiation of neotropical ecosystems by modern soil phytolith assemblages and its implications for palaeoenvironmental and archaeological reconstructions. Review of Palaeobotany and Palynology 193 (2013) 15–37. Watling J., J. Iriarte. 2013. Phytoliths from the coastal savannas of French Guiana. Quaternary International 287 (2013) 162-180. Kealhofer, L., and D. R. Piperno. 1998. Opal Phytoliths in Southeast Asian Flora. Smithsonian Contributions to Botany, number 88,39 pages, 49 figures, 5 tables. Benvenuto M.L., Fernández Honaine M., Osterrieth M.L. and E. Morel. 2015. Differentiation of globular phytoliths in Arecaceae and other monocotyledons: morphological description for paleobotanical application. Turk J Bot (2015) 39: © TÜBİTAK doi:10.3906/bot-1312-72 . Garnier, A., Neumann, K., Eichhorn B. and L. Lespez. 2013. Phytolith taphonomy in the middle- to late-Holocene fluvial sediments of Ounjougou (Mali, West Africa). The Holocene 23(3) 416–431. Kondo, R. 1977. Opal phytoliths: inorganic, biogenic particles in plants ans soils. Japan Agricultural Research Quaterly 11: 198-203. Kondo, R., Childs, C. & Atkinson, I., 1994. Opal phytoliths of New Zealand. Manaaki Whenua Press. Canterbury. Wallis, L., 2003. An overview of leaf phytolith production patterns in selected northwest Australian flora. Review of Palaeobotany and Palynology 125: 201–248. Fahmy, A.G., 2008. Diversity of lobate phytoliths in grass leaves from the Sahel region, West Tropical Africa: tribe Paniceae. Plant Systematics and Evolution 270, 1-23. Piperno, D.R., 2006. Phytoliths. A Comprehensive Guide for Archaeologists and Palaeoecologists. AltaMira Press, Lanham. Gallego, L., Distel, R.A., 2004. Phytolith assemblages in grasses native to central Argentina. Annals of Botany 94, 865-874. Mulholland, S.C., 1989. Phytolith shape frequencies in North Dakota grasses: a comparison to general patterns. Journal of Archaeological Science 16, 489-511. Strömberg, C.A.E., 2003. The Origin and Spread of Grass-dominated Ecosytems during the Tertiary of North America and How it Relates to the Evolution of Hypsodonty in Equids. Ph.D. thesis. University of California, Berkeley. Neumann, K., Fahmy, A., Lespez, L., Balloche, A., Huysecom, E., 2009. The early Holocene palaeoenvironment of Ounjougou (Mali): phytoliths in a multiproxy context. Palaeogeography, Palaeoclimatology, Palaeoecology 276, 87-106. Ollendorf, A.L., 1992. Toward a classification scheme of sedge (Cyperaceae) phytoliths. In: Rapp Jr., G., Mulholland, S.C. (Eds.), Phytolith Systematics. Plenum Press, New York. Piperno, D.R., 1988. Phytolith Analysis: an Archaeological and Geological Perspective. Academic Press, San Diego.