Submitted : April 30 th , 2020 – Accepted : July 1 6th , 2020 – Posted online : July 2 3th , 2020
To link and cite this article:
doi: 10.5710/AMGH.16.07.2020.3356
1 NEW MIDDLE TRIASSIC POLLEN TAXA OF THE SAN RAFAEL BASIN,
2 MENDOZA PROVINCE, ARGENTINA
3 NUEVOS TAXONES DE POLEN TRIÁSICO MEDIO DE LA CUENCA SAN
4 RAFAEL, PROVINCIA DE MENDOZA, ARGENTINA
5
6 PEDRO R. GUTIÉRREZ1, ANA MARÍA ZAVATTIERI2
7 1Sección Paleopalinología, Área Paleontología, Museo Argentino de Ciencias Naturales
8 “B. Rivadavia”, CONICET, Av. Ángel Gallardo 470, C1405DJR Ciudad Autónoma de
9 Buenos Aires, Argentina. [email protected]
10 2Unidad de Paleopalinología, Departamento de Paleontología, Instituto Argentino de
11 Nivología, Glaciología y Ciencias Ambientales (IANIGLA), CCT-CONICET-Mendoza,
12 Av. A. Ruíz Leal s/n, Parque General San Martín, M5502IRA Mendoza, Argentina.
14
15 40 pag. (26 text + 14 references); 9 figs.; 2 tables
16
17 Proposed header: GUTIÉRREZ & ZAVATTIERI: PALYNOFLORAS FROM THE
18 LADINIAN, ARGENTINA.
19
20 Short Description: New pollen taxa are described from the Ladinian Quebrada de los
21 Fósiles Formation, Puesto Viejo Group, San Rafael Basin, Argentina.
22
23 Corresponding author: Pedro Raúl Gutiérrez. [email protected].
1
24
25 Abstract. New pollen grains are described from diverse palynological assemblages
26 recorded from the Quebrada de los Fósiles Formation, lower unit of the Puesto Viejo
27 Group, San Rafael Basin (Mendoza, Argentina), southwestern Gondwana. Two new
28 pollen genera, Mendozapollenites and Crackipollenites are proposed, and three new
29 species, Mendozapollenites variabilis, Angustisulcites hexagonalis and Crackipollenites
30 polygonata and Crackipollenites sp. A, as well as, a monosaccate pollen indeterminate
31 (Form A) are fully described and illustrated. The microflora contains high proportions of
32 sphenopsid and lycopsid spores together with true ferns and bryophyte spores as minor
33 components; non-taeniate pteridosperm bisaccate pollen grains, taeniate monosaccate and
34 bisaccate pollen together with scarce monosulcate and inaperturate grains, among other
35 less frequent gymnosperm pollen and the co-occurrence of chlorococcalean algae and
36 fungal spores. The sedimentation of the Quebrada de los Fósiles Formation took place
37 during Middle Triassic Ladinian age, confirmed by recent radiometric data carried out
38 from both base and top effusive levels (ignimbrites) of the main profile of the unit. Thus,
39 this is the first unequivocal and unique microflora recorded at present for the Ladinian in
40 South America.
41 Key words: Palynology. Pollen grains. Systematics. Ladinian. Argentina. Southwestern
42 Gondwana.
43 Resumen. NUEVOS TAXONES DE POLEN TRIÁSICO MEDIO DE LA CUENCA
44 SAN RAFAEL, PROVINCIA DE MENDOZA, ARGENTINA. Se describen nuevos
45 granos de polen de diversas asociaciones palinológicas registradas de la Formación
46 Quebrada de los Fósiles, unidad inferior del Grupo Puesto Viejo, cuenca de San Rafael
2
47 (Mendoza, Argentina), suroeste de Gondwana. Se proponen dos nuevos géneros,
48 Mendozapollenites y Crackipollenites, se describen e ilustran las nuevas especies,
49 Mendozapollenites variabilis, Angustisulcites hexagonalis, Crackipollenites polygonata y
50 C. sp. A, y también un grano de polen monosacado indeterminado (Forma A). La
51 microflora contiene altas proporciones de esporas de esfenópsidas y licópsidas, junto con
52 esporas de helechos verdaderos y de briófitas, como componentes menores; granos de
53 polen bisacados no taeniados de pteridospermas, y granos de polen monosacados y
54 bisaccados taeniados, junto con escasos granos monosulcados e inaperturados entre otros
55 granos de pollen de gimnospermas menos frecuentes, y la co-ocurrencia de algas
56 clorococales y esporas de hongos. La sedimentación de la Formación Quebrada de los
57 Fósiles tuvo lugar durante el Triásico Medio (Ladiniano), confirmado por dataciones
58 radiométricas recientes realizadas en los niveles de ignimbritas de la base y del techo del
59 perfil tipo de la unidad. Por lo tanto, esta es la primera e inequívoca palinoflora registrada
60 hasta ahora para el Ladiniano en América del Sur.
61 Palabras clave. Palinología. Granos de polen. Sistemática. Ladiniano. Argentina.
62 Suroeste de Gondwana.
3
63
64 THE TRIASSIC CONTINENTAL DEPOSITS IN THE SAN RAFAEL BASIN (also referred as San
65 Rafael depocenter by Ottone et al., 2018 or San Rafael Block by Cariglino et al., 2019
66 and Monti & Franzese, 2016), southwest of Mendoza Province, central-west Argentina
67 (Fig. 1), are represented by the Puesto Viejo Group (Stipanicic et al., 2007). It consists of
68 ca. 1000 m thick alluvial to fluvial sedimentary sequences intercalated with
69 volcanoclastic rocks, tuffs, basalts, andesite intrusives and rhyolitic ignimbrites
70 accumulated from the early Middle to early Late Triassic interval. The Puesto Viejo
71 Group includes the lower Quebrada de los Fósiles Formation and the upper Río Seco de
72 la Quebrada Formation (Stipanicic et al., 2007). The contact between the basal unit and
73 the volcanic basement of Middle Permian–Lower Triassic Choiyoi Group is an erosional
74 unconformity (González Díaz, 1972). The upper unit is overlain unconformably by the
75 Miocene Aisol Formation (González Díaz, 1972). The complete stratigraphy and origin
76 of the units of the San Rafael Basin have been summarized by Domeier et al. (2011),
77 Ottone et al. (2014) and Monti & Franzese (2016, 2019). The basal part of the Quebrada
78 de los Fósiles Formation consists of coarse epiclastic strata related to proximal alluvial
79 fan systems, sometimes covered by pyroclastic flows. Finer grained levels of the upper
80 section of the Quebrada de los Fósiles Formation represent meandering fluvial systems of
81 low to high sinuosity that were also interrupted by the emplacement of pyroclastic flows
82 (Spalletti, 1994; Monti & Franzese, 2016, 2019). The upper unit, the Río Seco de la
83 Quebrada Formation, is constituted by deposits of braided to meandering low-sinuosity
84 fluvial systems, and towards the top, by coarse-grained strata interpreted to represent the
85 distal sections of alluvial fans intercalated by lava flows (Monti & Franzese, 2016, 2019).
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86 At its type section, the Quebrada de los Fósiles Formation initiates and culminates
87 with pyroclastic flow (ignimbrite) deposits. The uppermost rhyolitic ignimbrite level
88 marks the boundary with the overlying Río Seco de la Quebrada Formation. Ottone et al.
89 (2014) performed SHRIMP U–Pb radiometric dating from juvenile magmatic zircons of
90 the upper ignimbrite deposit which provided an age constraint of 235.8±2 Ma. Domeier et
91 al. (2011) carried out 40Ar/39Ar isotopic dating on volcanoclastic and ignimbritic rocks
92 from the basal levels of the Quebrada de los Fósiles Formation which yielded an age of
93 ~245 Ma. Recently, the basal ignimbrite has been dated using LA-ICPMS U–Pb
94 indicating a late Anisian age (Monti in Sato et al., 2015). Thus, the Quebrada de los
95 Fósiles Formation as a whole, was deposited from the Middle (late Anisian) to early Late
96 (Carnian) Triassic, being mostly constrained to the Ladinian (Cohen et al., 2018) (Fig. 2).
97 Monti (2015) and Monti & Franzese (2016, 2019) gave full details of the litho-
98 sedimentological description of both units integrating the Puesto Viejo Group and defined
99 five depositional systems: 1) debris-flow deposits, 2) channel-fill deposits, 3) floodplain
100 deposits, 4) pyroclastic deposits, and 5) volcanic intrusives and effusives. For detailed
101 lithofacies analysis of the Quebrada de los Fósiles Formation, we refer to these
102 mentioned contributions.
103 The Quebrada de los Fósiles flora recorded at the type locality of the unit was initially
104 referred as “Pleuromeia flora” composed only of scarce remains of lycophytes and
105 sphenophytes (Morel & Artabe, 1994; Coturel et al., 2016). Recently, Cariglino et al.
106 (2018) described a new, more diverse macrofloral assemblage of Ptilozamites,
107 Pleuromeia, Lepacyclotes, Equisetites and Neocalamites from the Quebrada de los
108 Fósiles Formation at its type locality. This unique and exceptional novel plant assemblage
5
109 of the lower unit of the Puesto Viejo Group includes typical Northern Hemisphere
110 Ptilozamites and Lepacyclotes genera—recorded for the first time in Gondwana—
111 together with Pleuromeia as a typical Middle Triassic assemblage recorded in the
112 Quebrada de los Fósiles Formation (Cariglino et al., 2018).
113 Ottone & García (1991) systematically described for the first time the palynoflora
114 from the type locality of Quebrada de los Fósiles Formation. Vázquez (2013) described a
115 microflora assemblage from the Quebrada de los Fósiles Formation at the Río Seco de la
116 Quebrada creek (see more details in Ottone et al., 2014: 193). During several field
117 seasons from 2010 to 2018, new palynological assemblages were recovered from the
118 Quebrada de los Fósiles Formation at both creeks which are being studied. Zavattieri et
119 al. (in press) have recently described and illustrated diverse freshwater green algal
120 microfossils and dispersed fungal spores among rich terrestrial palynofloras recovered in
121 the recent years from the upper part of the Quebrada de los Fósiles Formation, Puesto
122 Viejo Group.
123 The microfloras were recovered from three sectors of the type section of the Quebrada
124 de los Fósiles Formation, cropping out at the homonymous creek (Figs. 1, 2): a) thin
125 tabular tuffaceous shales with abundant organic matter (AMZ2 section of the integrated
126 type profile; Figs. 1, 2); b) tuffaceous shales and limestones with light gray to whitish
127 shales having fine and densely distributed root casts indicating development of incipient
128 paleosols (AMZ1 section of the integrated type profile; Figs. 1, 2), and c) tabular deposits
129 of laminated siltstones interbedded with light yellow-orange biolaminated calcareous
130 layers (GzD section of the integrated type profile, Figs. 1, 2). In the Quebrada del Río
131 Seco section (Fig. 3), the palynological samples were recovered from grey bentonitic
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132 clays, dark-brown to black carbonaceous shales (coal levels) with frequent organic plant
133 debris and from grey thin lamination siltstones with frequent organic plant debris.
134 Institutional abbreviations. IANIGLA, CCT-CONICET, Argentine Institute of
135 Nivology, Glaciology and Environmental Sciences, Technological Scientific Center-
136 National Scientific and Technical Research Council-Argentina, Mendoza, Argentina;
137 MACN, Argentine Museum of Natural Sciences “Bernardino Rivadavia”, Ciudad
138 Autónoma de Buenos Aires, Argentina; MPLP, Paleopalinoteca-Paleopalynological
139 Laboratory; BAPal, the Palynological Collection of the Argentine Museum of Natural
140 Sciences “Bernardino Rivadavia”.
141 MATERIALS AND METHODS
142 The palynological materials described herein have been recovered from 22 levels of
143 the type locality of the Quebrada de los Fósiles Formation exposed at the homonymous
144 creek and from 17 levels of the Río Seco de la Quebrada Creek. Most of the studied
145 levels in the two sections contain abundant and diverse spores, pollen grains, freshwater
146 algae and fungi.
147 Standard palynological extraction techniques were performed at the
148 Paleopalynological Laboratory of IANIGLA, CCT-CONICET and involved standard
149 treatments with HCl-HF acids. Some organic residues were subjected to further oxidation
150 using concentrated nitric acid (HNO3) (>2–10 minutes as required). Other oxidized
151 samples were stained with safranin for light microscopy observation and
152 microphotography. The organic residues were briefly washed (45 seconds) in 5%
153 ammonium hydroxide to remove the oxidation products, then sieved using a 10 µm mesh
154 and finally mounted with glycerine jelly (Volkheimer & Melendi, 1976). The specimens
7
155 were examined and photographed with a Nikon H550S transmitted light microscope
156 equipped with a DS-U2-Fil-U2 digital camera at the Paleopalynology Laboratory
157 (MACN), as well as by an Olympus BX 50 light microscope (LM), equipped with an
158 Olympus digital camera at the Paleopalinology Unit, IANIGLA, CCT-CONICET. The
159 studied slides with the prefix MPLP are housed at the Mendoza Paleopalinoteca-
160 Paleopalynological Laboratory (IANIGLA) and identified by the catalogue numbers
161 5877, 9076–9082, 9085, 10374–10377, 10379, 11051–11056, 11064. Other slides with
162 prefix BAPal are housed at the Palynological Collection of the MACN and identified by
163 6612–6614. Specimens are located on each slide with England Finder coordinates
164 (E.F.co.). Residues for Scanning Electron Microscope (SEM) were mounted on
165 aluminium stubs and coated with a gold-palladium alloy. A Phillips XL30 TMP electron
166 microscope at the MACN, Buenos Aires, was used for detailed examination and
167 microphotography of selected specimens, identified by the number of the stub (T) and the
168 photogram number (f). Dimensions for all specimens are expressed as “minimum (mean)
169 maximum”.
170 SYSTEMATIC PALEONTOLOGY
171 Anteturma POLLENITES Potonié, 1931
172 Turma SACCITES Erdtman, 1947
173 Subturma MONOSACCITES Chitaley, 1951 emend. R. Potonié & Kremp, 1954
174 Genus Mendozapollenites gen. nov.
175 Type species. Mendozapollenites variabilis sp. nov., Ladinian of San Rafael Basin,
176 Mendoza Province, Argentina.
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177 Derivation of name. The name alludes to the Mendoza Province (Argentina) where the
178 San Rafael Basin is located.
179 Diagnosis. Pollen grain monosaccate, taeniate, subcircular to oval in polar view, both in
180 transversal and longitudinal axis. Insertion of sacci sub-equatorial on both faces of the
181 central body. Proximal insertion of sacci slightly marked; on the distal face, a narrow
182 tapering cappula, whose edges converge towards the ends. Near the roots of the sacci,
183 striae are present parallel to the sides of the cappula and generally associated with
184 transverse folds. Saccus intrareticulate, with well-defined limb. Central body variable in
185 shape. Cappa exhibiting straight to curved taeniae, irregularly arranged, mainly in the
186 longitudinal direction of the grain, parallel to each other or even crossed between them.
187 Geographic occurrence. Quebrada de los Fósiles and Río Seco de la Quebrada creeks,
188 San Rafael Basin, Mendoza Province, Argentina.
189 Stratigraphic provenance. Upper section of the Quebrada de los Fósiles Formation,
190 Ladinian, Puesto Viejo Group.
191 Remarks and comparison. Mendozapollenites gen. nov. is characterized by the
192 arrangement of its structures: sub-equatorial saccus attachment on both sides, very
193 narrow cappula developed on distal face of the central body and in a transverse direction;
194 central body bearing straight, curved and irregularly arranged taeniae and an equatorial
195 limbus of the saccus.
196 Misrapollenites Anand-Prakash, 1970 is the most morphologically similar genus
197 (attachment of the saccus and presence of the limbus) but the central body has no taeniae
198 on its distal surface, the cappula has a greater development and the saccus do not exhibit
199 the parallel taeniae next to the margin of the cappula. The same type of the attachment is
9
200 also presented by Korbapollenites Tiwari, 1964, Primuspollenites Tiwari, 1964,
201 Fimbriaesporites Leschik, 1959, Direticuloidospora Tiwari, 1964, Rhizomaspora
202 Wilson, 1962 and Ibisporites Tiwari, 1968, although they lack an equatorial limb on the
203 sacci. Korbapollenites, Primuspollenites, Fimbriaesporites and Direticuloidospora have
204 central bodies with reticulate striae with varied polygonal patterns, while Rhizomaspora
205 has ribs on the proximal face of the central body and lacks limbus. Otherwise, Ibisporites
206 has a smooth central body.
207 Mendozapollenites variabilis sp. nov.
208 Figures 4.1−4.6, 5.1−5.8, 6.1−6.7
209 Derivation of name. The name alludes to the variable shape of the central body of the
210 pollen grain.
211 Type material. Holotype. MPLP 5877(C) C44(1) (Fig. 4.1). Paratypes. MPLP 10348(G)
212 J44(0) (Fig. 4.2), MPLP 5877(C) H40(0) (Fig. 4.3), MPLP 5877(E) S52(4) (Fig. 4.4),
213 MPLP 5877(C) F53(1) (Fig. 4.5), BAPal 6612(7) D26(1) (Fig. 4.6), MPLP 5877(C)
214 U38(0) (Fig. 5.1), MPLP 5877(C) F51(0) (Fig. 5.2), MPLP 5877(E) G48(0) (Fig. 5.3),
215 BAPal 6612 (7) P45(4) (Fig. 5.4), MPLP 10352(E) K53(0) (Fig. 5.6), MPLP 10348(G)
216 W39(4) (Fig. 5.8), BAPal MEB 6612 (TM) f011 (Fig. 6.1), BAPal MEB 6612(TM) f029
217 (Fig. 6.2), BAPal MEB 6612(TM) f036 (Fig. 6.3), BAPal MEB 6612(TP) f050 (Fig. 6.4),
218 BAPal MEB 6612 (T1) f042 (Fig. 6.5), BAPal MEB 6612(TQ) f013 (Fig. 6.7), BAPal
219 MEB 10374(TJ) f011 (Fig. 6.6).
220 Referred material. MPLP 10348(G) R38(0) (Fig. 5.5), BAPal 6614(8) E43(0) (Fig. 5.7),
221 BAPal 6612(3) Z46(2), BAPal 6612(6) N52(0), BAPal 6612(7) Q51(3), BAPal MEB
222 6612(TP) f032, BAPal 6614(1) N60(0), BAPal 6614(3) B43(4), MPLP 5877(A) R43(0),
10
223 MPLP 5877(C) M39(3), MPLP 5877(C) P44(4), MPLP 5877(D) W50(2), MPLP 5877(E)
224 P34(3), MPLP 5877(F) C44(4), MPLP 10351(M) M54(2), MPLP 10362(I) L56(1),
225 MPLP 11054(B) V31(3), MPLP MEB 10374(TJ) f040.
226 Diagnosis. Pollen grain monosaccate, taeniate. In polar view it exhibits a subcircular to
227 oval shape. Insertion of sacci subequatorial on both faces of the central body; proximal
228 insertion slightly marked; on the distal face a narrow and tapering cappula is developed
229 along the corpus length, not reaching to the corpus margin. Near of the saccus, one or two
230 grooves parallel to the cappula borders are present, generally associated to the folds.
231 Intexine medium to thick. Central body of central to eccentrical position in polar view,
232 variable in shape and size and it has been associated to compressed folding of irregular
233 position. The shape of the central body varies from polygonal, sub-circular to oval, with
234 the major axis parallel or perpendicular to the cappula. Cappa taeniate with up to 10
235 straight to curved taeniae, irregularly disposed but mainly developed in the longitudinal
236 axis of the grain; they are partially parallel to each other or crossed between them in
237 variable angles.
238 Geographic occurrence. Quebrada de los Fósiles and Río Seco de la Quebrada creeks,
239 San Rafael Basin, Mendoza Province, Argentina.
240 Stratigraphic occurrence. Upper section of the Quebrada de los Fósiles Formation,
241 Ladinian, Puesto Viejo Group.
242 Description. The pollen grains are monosaccate, taeniate, subcircular to oval in shape,
243 both in transverse and longitudinal axis. The insertion of saccus is subequatorial, slightly
244 discernible on the proximal face, occasionally associated to exinal folds. On the distal
245 face, the insertion is bilaterally subequatorial, defining a narrow tapered cappula
11
246 (generally 3.2 µm wide), developed along the length of the corpus, but not reaching the
247 margin of the corpus, with exception of breaking. Between roots of the sacci and the
248 taeniae, the exoexine presents parallel folds of the cappula. The central body is polygonal
249 (pentagonal, hexagonal, rhomboidal, trapezoidal) to irregular in shape, irregular folds can
250 be present generally in central position of the body. On the cappa, one and ten straight to
251 curve taeniae are observed; they can cross in variable angles (from very sharp to straight),
252 but not forming reticulate pattern. Most of the taeniae are perpendicular to the cappula.
253 The limb is generally visible, 1 to 8 µm in width, conferring a rigid aspect to the grain.
254 Dimensions (390 specimens). In polar view: overall equatorial diameter, 70(116.4)211 x
255 63(111.8)175 µm; width of the corpus, 34(73.4)149 µm; corpus length, 30(79.6)144 µm;
256 ratio overall equatorial diameter major axis/minor axis, 0.6(1.1)1.8, ratio width of the
257 corpus/corpus length, 0.3(1.0)3.5; cappula width, 1(3.2)16 µm; width of limb: 1(3.8)8
258 µm. Number of taeniae, 1(4)10.
259 Remarks and comparisons. Mendozapollenites variabilis sp. nov. it is characterized by
260 its body of variable shape (pentagonal, hexagonal, rhomboidal, trapezoidal to irregular)
261 and the presence of 1 to 10 taeniae (straight to curve) on the proximal face, which cross
262 in various ways but without exhibiting a reticulated pattern. Densipollenites sp. A
263 described and illustrated by Jain (1968, p. 16, pl. 2, fig. 34) shows similar morphology to
264 M. variabilis sp. nov., due to the presence of two curved taeniae on the central body and
265 the presence of the cappula, both features were not included in the original description.
266 Cf. Parasaccites densicorpus Lele, 1975 described and illustrated by Nagamadhu &
267 Bilwa (2012: 92, fig. 22) presents a central body of similar morphology, dimensions and
268 general aspect of the grain allowing comparison with M. variabilis sp. nov. However, the
12
269 regular preservational state of cf. P. densicorpus and the poor quality of its illustration do
270 not permit more accurate comparison between both forms.
271 In the palynological literature of the Permian-Triassic of Gondwana we have found a
272 set of very similar monosaccate grains, which could be compared with M. variabilis sp.
273 nov. but may or may not be considered co-specific. This material has been illustrated
274 and/or described and referred, mainly to the genus Densipollenites Bharadwaj, 1962, and
275 seems to present, although not clearly visible, stretch marks on the proximal face of the
276 central body and/or an insertion bilateral-distal of the saccus (defining a narrow cappula).
277 However, the diagnostic characteristics of Mendozapollenites variabilis sp. nov. were not
278 originally described for Densipollenites and as a result, it is not possible to advance more
279 than in a superficial comparison. Among some of these forms we can mention some of
280 the specimens referred to D. indicus Bharadwaj (1962: 87, pl. 6, figs. 103–104; Tiwari,
281 1964: 187, pl. 5, fig. 95; Maheshwari, 1966: 270–271, pl. 5, fig. 41; Segroves, 1969: 187,
282 pl. 3, fig. B; Tiwari & Tripathi, 1992, pl. 4, fig. 2; Vijaya et al., 2012, pl. 1, figs. A–B;
283 Murthy & Sarate, 2016, pl. 1, fig. 6), D. invisus Bharadwaj & Salujha (1963: 198, pl. 5,
284 figs. 74–75; Tripathi, 2001, pl. I, fig. 6; Vijaya et al., 2012, fig. 5T; Murthy et al., 2013,
285 pl. I, fig. 22; Meena et al., 2013, pl. 1, fig. 1; Mishra & Joshi, 2015, fig. 3H; Mahesh et
286 al., 2016, fig. 7F; Singh et al., 2017, fig. 7.10); D. pullus Segroves (1969: 187–188, pl. k,
287 figs. A–B; pl. 5, figs. B–C), D. oviformis Shwartsman in Inossova et al., 1976 (Pittau et
288 al., 2002, fig. 7.11) and D. magnicorpus Bharadwaj, 1962 (Kumar et al., 2012, pl. 1, fig.
289 6).
290 INCERTAE SEDIS
291 Genus Indeterminate
13
292 Monosaccate indeterminate Form A
293 Figure 6.8–6.11
294 Referred material. BAPal 6612(3) O47(0) (Fig. 6.8), BAPal 6612(3) J32(1) (Fig. 6.9),
295 MPLP 10362(E) U34(2), (Fig. 6.10), BAPal 6612(3) B50(0) (Fig. 6.11).
296 Geographic occurrence. Quebrada de los Fósiles and Río Seco de la Quebrada creeks,
297 San Rafael Basin, Mendoza Province, Argentina.
298 Stratigraphic occurrence. Upper section of the Quebrada de los Fósiles Formation,
299 Ladinian, Puesto Viejo Group.
300 Description. Monosaccate pollen grain of bilateral symmetry, equatorial outline oval to
301 elliptical in polar and equatorial views. In polar view, the exine layers are separated in the
302 equatorial-proximal and distal zones; the endexine conforms to the central body of oval to
303 elliptical contour. The ektexine is fused to the endexine in the greater part of the proximal
304 face, having radial folds on the proximal surface of the central body. The sacci are
305 narrow, and present their greatest development at the ends of the grain. A monolete mark
306 is present in the proximal face, associated to fold that do not reach to the margin of the
307 body. It has no opening in the distal area. The lateral-distal tenuitas are not clearly
308 observed.
309 Dimensions (8 specimens). Overall length, 105(121.9)170 µm, overall width, 32(37.8)45
310 µm.
311 Remarks and comparisons. The pollen grains characterized as Form A include
312 monosacate specimens of bilateral symmetry, with a proximal monolete mark, the saccus
313 attached equatorially on the proximal face and totally encompassing the distal face of the
14
314 central body. No comparable genus has been found in the literature. The few specimens
315 recovered of this type do not allow to propose a new taxon.
316 Subturma DISACCITES Cookson, 1947
317 Infraturma STRIATITI Pant, 1954
318 Genus Angustisulcites Freudenthal, 1964
319 Type species. Angustisulcites klausii Freudenthal, 1964. Lower Triassic of Netherlands.
320 Angustisulcites hexagonalis sp. nov.
321 Figures 7.1–7.7
322 Derivation of name. The name alludes to the shape of the central body.
323 Type material. Holotype: MPLP 10348(G) Y33(0) (Fig. 7.5). Paratypes: MPLP
324 10348(D) X53(1) (Fig. 7.1), MPLP 10348(C) Y52(1) (Fig. 7.3), MPLP 10376 (E) B55(0)
325 (Fig. 7.7).
326 Referred material. MPLP 10348(G) P44(1) (Fig. 7.2), MPLP 10348(C) X37(2) (Fig.
327 7.4), MPLP 10348(D) J37(0) (Fig. 7.6).
328 Diagnosis. Pollen grain bisaccate, taeniate. Central body of thick exine, transversaly sub-
329 hexagonal to oval in shape. Asymmetric trilete mark, with two rays extending to the
330 equator and the third in transverse sense and less developed. Sacci larger than the central
331 body, slightly distally inclined, and laterally they are juxtaposed. Narrow cappula of
332 straight sides, concave or convex having sporadic nexinal associated folds.
333 Geographic occurrence. Quebrada de los Fósiles and Río Seco de la Quebrada creeks,
334 Mendoza Province, Argentina.
335 Stratigraphic occurrence. Upper section of the Quebrada de los Fósiles Formation,
336 Ladinian, Puesto Viejo Group.
15
337 Description. The pollen grains are bisaccate, taeniate. The central body has a thick wall,
338 and is transversely oval, rhomboidal to sub-hexagonal in shape (two lateral sides shorter
339 than the other four). The trilete mark is asymmetric, with two rays that extend until
340 reaching the proximities of the body’s equator, while the third one is developed in a
341 transverse direction and extends between 1/3 and one radius of the central body. The
342 sacci are larger than the central body, slightly distally inclined and laterally close to each
343 other. The cappula is narrow, biconvex to rectangular with exinal folds associated to the
344 margins.
345 Dimensions (125 specimens). Measures in polar view. Overall length, 50(81.4)120 µm,
346 width of the sacci, 40(64.4)92 µm; length of the sacci, 9(23.6)50 µm (proximal face),
347 21(38.8)75 µm (distal face); corpus length, 27(41.9)60 µm; corpus width, 36(56.0)74 µm;
348 ratio corpus length/width, 0.5(0.8)1.0; cappula width, 1(4.1)28 µm; width of the distal
349 folds, 2(9.3)23 µm.
350 Remarks and comparisons. Angustisulcites hexagonalis sp. nov. differs from the
351 remaining Angustisulcites species (see Visscher, 1966) by the transversely oval to
352 rhomboid shape of the central body, its thick wall and its haploxylonoid to slightly
353 diploxylonoid contour.
354 Turma HILATES Dettmann, 1963
355 Subturma AZONOHILATES Ramanujam & Varma, 1981
356 Genus Crackipollenites gen. nov.
357 Type species. Crackipollenites polygonalis sp. nov. Ladinian of San Rafael Basin,
358 Mendoza Province, Argentina.
16
359 Derivation of name. The name refers to the characteristic polygonal cracked surface of
360 the pollen grain (of the Latin “crack” = cracked).
361 Diagnosis. Pollen grain heteropolar with subcircular to elliptical outline in polar view;
362 irregular area of the opening (hilum) on one of the poles. Hilum not always distinctive, in
363 part covered by secondary folds and/or by breaking. Wall minutely and evenly punctate,
364 ca. 1−2 µm thick; negatively reticulate, divided into large irregular polygonal areas.
365 Comparisons. Crackipollenites gen. nov. is characterized by the presence of an irregular
366 hilum, an exine not divided into visible layers and by its surface with an irregular
367 negative reticulum (polygonal grooves that divide positive polygonal areas of different
368 sizes and shapes). The most similar genera are Sciadopityspollenites Raatz ex Potonié,
369 1958 and Haberkornia Scheuring, 1978. The first one is distinguished by its verrucose
370 ornamentation, while Haberkornia has short trilete mark instead of the hilum.
371 Balmeiopsis Archangelsky, 1979 has a superficial resemblance however, it has the exine
372 separated into two layers and a finely granulated surface.
373 Crackipollenites polygonalis sp. nov.
374 Figures 7.8–7.12, 8.1–8.8, 9.1–9.6
375 Synonymy.
376 1991. Inaperturopollenites sp. 1; Zavattieri: 220, pl. VI, figs. 10-11.
377 Derivation of name. The name alludes to the “polygonal cracked” appearance of the
378 surface.
379 Type material. Holotype: MPLP 5877(C) F49(0) (Fig. 8.1). Paratypes: BAPal 6612(8)
380 Y53(3) (Fig. 7.8), MPLP 5877(D) U47(4) (Fig. 8.2), MPLP 5877(F) M35(1) (Fig. 8.3),
17
381 MPLP 5877(F) W36(2) (Fig. 8.4), BAPal MEB 6612(TN) f025 (Fig. 9.3), BAPal MEB
382 6612(TN) 045 (Fig. 9.5).
383 Referred material. BAPal 6612(8) F27(4) (Fig. 7.9), BAPal 6612(3) K48(2) (Fig. 7.10),
384 BAPal 6613(2) D37(2) (Fig. 7.11), BAPal 6614(1) U48(0) (Fig. 7.12), MPLP 10379(E)
385 G35(4) (Fig. 8.5), MPLP 5877(C) R39(1) (Fig. 8.6), MPLP 11056(G) N43(0) (Fig. 8.7),
386 BAPal 6614(1) R56(3) (Fig. 8.8), BAPal 6614(2) V50(4) (Fig. 9.1), MPLP 10348(D)
387 K48(0) (Fig. 9.2), BAPal 6613(2) Y56(2) (Fig. 9.4), BAPal 6612(3) M37(1) (Fig. 9.6),
388 BAPal 6612(7) Y32(1), BAPal 6612(8) X39(3), BAPal 6614(2) W43(1), MPLP 5877(C)
389 L45(0), MPLP 5877(D) G39(3), MPLP 9085(F) J38(1), MPLP 9085(F) W45(0), MPLP
390 10362(A) W43(3), MPLP 10375(1) Y42(4), MPLP 10376(E) W52(3), MPLP 10376(E)
391 Q52(1).
392 Diagnosis. Pollen grain originally large, spheroidal, with wall divided into variable shape
393 and dimensions of polygonal sections, producing a negative reticulate pattern on the
394 surface of the grain. Flat-topped polygonal sections separated by interconnected shallow
395 “cracks” that have a “v” shape in cross-section.
396 Geographic occurrence. Quebrada de los Fósiles and Río Seco de la Quebrada creeks,
397 Mendoza Province, Argentina.
398 Stratigraphic occurrence. Upper section of the Quebrada de los Fósiles Formation,
399 Ladinian, Puesto Viejo Group.
400 Description. The pollen grains are hilate, spheroidal in shape, circular to subcircular in
401 equatorial contour (relationship between the major and the minor diameters, usually 1.3).
402 They present frequent folding parallel to the equator. On one face, a hilum of a size
403 equivalent to 2/3 of the grain diameter is observed, bordered by irregular folds. The exine
18
404 is 1 µm thick, covered by elements of polygonal contour and flat tops (from 3.3 to 4.8 µm
405 in average diameter, high 0.7 µm), separated from each other by narrow “grooves” of 0.5
406 µm in approximate similar width and forming a polygonal design that give the
407 appearance of negative reticulum. The elements of the sculpture have irregular sizes and
408 shapes, and appearing less marked until they disappear towards the center of the hilum.
409 By the projection of the sculptural elements, the equatorial contour is slightly dentate.
410 Dimensions (136 specimens). Larger equatorial diameter, 55(79.7)130 µm; minor
411 equatorial diameter, 39(61.3)100 µm; ratio equatorial diameter larger:minor 1(1.3)2.2;
412 polygonal section: larger diameter 2(4.8)9 µm, smaller diameter 1(3.3)6 µm; height,
413 0.5(0.7)1.0 µm.
414 Remarks. Due to the large dimensions and probably the weak construction of the wall by
415 the polygonal cracks, the specimens are frequently broken or folded.
416 Comparisons. Inaperturopollenites sp. 1 recorded in the Cerro de Las Cabras Formation
417 (Zavattieri, 1991, p. 220, pl. VI, figs. 10–11) is herein assigned to Crackipollenites
418 polygonalis gen et sp. nov. because it presents the diagnostic characteristic features, as
419 the presence of the hilum, shape and size of the specimens and the same type and size of
420 the exine ornamentation.
421 Crackipollenites sp. A
422 Figure 9.7–9.11
423 Referred material. MPLP 10348(D) V52(0) (Fig. 9.7), BAPal MEB 6612(TQ) f16 (Fig.
424 9.8), MPLP 11054(D) B42(3) (Fig. 9.9), MPLP 10379(E) V36(0) (Fig. 9.10), MPLP
425 10348(C) X33(3) (Fig. 9.11), MPLP 10348(G) 33,3/103,0, MPLP 10350(G) M37(1),
19
426 MPLP 10352(B) C36(4), MPLP 10374(C) M40(0), MPLP 10376(A) G41(4), MPLP
427 11052(C) O46(0), MPLP 11071(A) S35(2), BAPal 6613(4) F29(4).
428 Geographic occurrence. Quebrada de los Fósiles and Río Seco de la Quebrada creeks,
429 San Rafael Basin, Mendoza Province, Argentina.
430 Stratigraphic occurrence. Upper section of the Quebrada de los Fósiles Formation,
431 Ladinian, Puesto Viejo Group.
432 Description. The pollen grains are hilate, spheroidal in shape, circular to subcircular
433 equatorial contour (ratio between major and minor diameters generally 2.9), with frequent
434 folds disposed parallel to the contour. A hilum is present on one face of the grain; it has a
435 size equivalent to the 2/3 of the grain diameter and bordered by irregular folding. The
436 exine is 2 µm thick, covered by elements of polygonal contour and flat tops (8 to 13 µm
437 of larger diameter by 2 to 5 µm of smaller diameter, and height of 0.5 to 1.5 µm),
438 separated from each other by narrow grooves (0.5 µm wide approximate) of irregular and
439 broken pattern that give the appearance of negative reticulum. The elements of the
440 sculpture have variable sizes and predominantly rectangular shapes; they appear less
441 marked until they disappear towards the center of the hilum. Due to the projection of the
442 sculptural elements, the margin of the grain appears slightly dentate.
443 Dimensions (8 specimens). Larger equatorial diameter, 50(65.5)80 µm; minor equatorial
444 diameter, 39(52.7)60 µm; ratio equatorial diameter larger:minor 1(1.2)1.5; polygonal
445 section: larger diameter 3(8)13 µm; smaller diameter, 2(3)5 µm; height, 0.5–1,5 µm.
446 Remarks and comparison. Crackipollenites sp. A differs from C. polygonata sp. nov. in
447 the rectangular shape of the ornamentation, which gives the appearance of aligned
448 polygonal plates, and by smaller size.
20
449 COMPOSITION OF THE QUEBRADA DE LOS FÓSILES MICROFLORA
450 The palynological assemblage from the type locality of the Quebrada de Los Fósiles
451 Formation were recovered from the floodplain facies of a meandering fluvial system that
452 laterally intergrades with episodic shallow lacustrine and palustrine deposits associated to
453 incipient development of paleosols and carbonate levels. On the other hand, in the
454 Quebrada del Río Seco section, the palynological assemblages were recovered from the
455 subaqueous subenvironment facies, such as swamp, marginal lacustrine bodies, stagnant
456 pools, moist soils and peats (Spalletti, 1994; Zavattieri et al., 2003; Monti & Franzese
457 2016) (Figs. 2–3).
458 The vegetation, as reflected by the palynoflora recorded in the Quebrada de los Fósiles
459 Formation, is dominated by monolete and trilete spores, taeniate and non-taeniate pollen
460 grains, algae and fungi (Tabs. 1–2). The spores are dominant components in AMZ1 and
461 AMZ2 sections of the Quebrada de los Fósiles Creek and in the section of the Río Seco
462 de la Quebrada Creek (Tabs. 1–2). The dispersed spores include representatives of
463 lycophytes (mainly Aratrisporites Leschik, 1955; Densoisporites Weyland & Krieger,
464 1953 emend. Dettmann, 1963; Lundbladispora Balme, 1963), sphenophytes
465 (Calamospora Schopf et al., 1944), true ferns (mainly Apiculatisporis Potonié & Kremp,
466 1956; Verrucosisporites Ibrahim, 1933; Leschikisporis Potonié, 1958), bryophyte
467 (Retusotriletes Naumova, 1953; Stereisporites Pflug in Thomson & Pflug, 1953)
468 (Zavattieri et al., in press). AMZ1 and GzD levels report Pleuromeiaceae and
469 equisetalean plant remains, and high representations of their dispersed microspores
470 reflecting local vegetation of lowland environments (floodplains, riparian lakes and/or
471 ponds).
21
472 The shallow ephemeral lacustrine systems and small water bodies developed in the
473 stratotype of the Quebrada de los Fósiles Formation as well as in the Quebrada del Río
474 Seco sections are evidenced by variable proportions of aquatic (fresh and/or brackish)
475 algae (0–97.8%, see Zavattieri et al., in press). Green algae are mainly represented by the
476 Zygnematales (Lecaniella Cookson & Eisenack, 1962; Maculatasporites Tiwari, 1964,
477 Mehlisphaeridium Segroves, 1967; and Ovoidites Potonié ex Thomson & Pflug, 1953),
478 scarce colonies of chlorophyte algae (Botryococcus Kützing, 1849), and other organic-
479 walled incertae sedis microfossils (Leiosphaeridia Eisenack, 1958) (Zavattieri et al., in
480 press).
481 Fungal spores are quite rare in the palynological spectrum of the Quebrada de los
482 Fósiles Formation represented mainly by the genera Portalites Hemer & Nygreen, 1967
483 and Microsporonites Jain, 1968. Their higher proportions in the studied sections are
484 recorded within peats and/or coal levels of the Río Seco de la Quebrada section
485 (Zavattieri et al., in press).
486 The pollen grain assemblages of the Quebrada de los Fósiles Formation in the studied
487 sections include conifers, and pteridosperms, particularly peltasperms and corystosperms,
488 and rare monosulcate pollen of ginkgoaleans, cycadaleans and/or bennettitaleans. The
489 conifers and pteridosperms are dominated by non-taeniate bisaccate pollen (partly
490 Alisporites Daugherty, 1941/Falcisporites Leschik, 1956; Platysaccus Naumova ex
491 Ishchenko, 1952; Minutosaccus Mädler, 1964/Protodiploxypinus Samoilovich, 1953; and
492 Klausipollenites Jansonius, 1962) and they are well represented in the Río Seco de la
493 Quebrada Creek section (Tab. 2). Pollen grains of the mentioned groups might have
22
494 derived from upland vegetation (drier parts of the floodplains, and surrounding areas) that
495 supplied allochthonous elements to the sites of deposition.
496 The pollen grains described here belong to miospore groups with low representation in
497 the type locality of the Quebrada de los Fósiles Formation (see Tabs. 1–2). However, they
498 are forms that are easily identifiable by their morphology and constitute distinctive
499 components of the Quebrada de los Fósiles microflora. In this sense, among the taeniate
500 monosaccate pollen grains of this palynoflora (Tabs. 1–2), Mendozapollenites gen. nov.
501 constitutes the dominant component, accompanied by few grains of the genera
502 Striomonosaccites Bharadwaj, 1962 and Crustaesporites Leschik, 1956. On the other
503 hand, among the non-taeniate pollen grains, several species of the genera Accinctisporites
504 Leschik, 1955, Cladaitina Maheshwari & Meyen, 1975 and Sulcosaccispora Klaus, 1964
505 were identified.
506 The taeniate bisaccate pollen grains also appear in low proportions within the
507 composition of the palynofloras of the Quebrada de los Fósiles Formation (see Tabs. 1–
508 2). The genus appearing in quantities which are included in the counts are:
509 Protohaploxypinus Samoilovich, 1953; Striatopodocarpites Sedova, 1956;
510 Angustisulcites Freudenthal, 1964; Striatoabieites Sedova, 1956; Hamiapollenites
511 Wilson, 1962; and Lunatisporites Leschik, 1956. Angustisulcites is mainly represented by
512 A. hexagonalis. sp. nov.; A. gorpii Visscher, 1966; A. grandis (Freudenthal, 1964)
513 Visscher, 1966; and A. klausii (Freudenthal, 1964) Visscher, 1966.
514 Finally, hilate pollen grains appear in even smaller proportions (Tabs. 1–2) and are
515 represented by Crackipollenites polygonalis gen. et sp. nov. This is the only species of
516 those described in this work that has a previous record in the Triassic of Argentina,
23
517 referred previously as Inaperturopollenites sp. 1 by Zavattieri (1991) from the Cerro de
518 Las Cabras Formation, Cuyana Basin (Mendoza, Argentina).
519 FINAL COMMENTS
520 This work presents new palynological components of the microfloral assemblages
521 recovered from two sections of the Quebrada de los Fósiles Formation, lower unit of the
522 Puesto Viejo Group, San Rafael Basin, central-western Argentina. The microflora of the
523 unit is referred to Ladinian age based on recent radiometric dating and stratigraphical
524 interpretations.
525 The composition of the Quebrada de los Fósiles microflora can be clearly
526 differentiated from all other known late Middle to Upper Triassic palynological
527 assemblages of Argentina and Gondwanan strata which are characteristically dominated
528 by non-taeniate corystospermous and peltaspermous bisaccate pollen mainly assigned to
529 Falcisporites/Alisporites, Pteruchipollenites, Platysaccus, Minutosaccus,
530 Klausipollenites, Vitreisporites, among other gymnosperm pollen grains as well as
531 abundant and diverse dispersed spores of true ferns and sphenopsids. It also differs from
532 those of the uppermost Permian (Lopingian) microflora recorded in Argentina dominated
533 by glossopterids with sub-dominant conifers (Voltziales), Cordaitales, peltasperms and
534 corystosperm pollen grains, and which also have low proportion of spores related to true
535 ferns and shenophytes (Gutiérrez et al., 2017, 2018; Zavattieri et al., 2017, 2018).
536 Similarly, the flora from the Quebrada de los Fósiles Formation is unique when
537 compared to other Middle and Late Triassic assemblages in Argentina, in that it includes
538 Ptilozamites (commonly registered in the Northern Hemisphere), and abundant
539 pleuromeids and sphenophytes (Cariglino et al., 2018). Other typical plant elements of
24
540 the well-known Dicroidium flora (such as corystosperms and peltasperms) common to
541 the various Triassic floras from Argentina and other Gondwanan regions are absent
542 (Cariglino et al., 2018).
543 Among the pollen grains constituting the composition of the Quebrada de los Fósiles
544 microflora, a small set of forms hitherto unknown to the Triassic of Argentina are
545 described herein assigned to Mendozapollenites variabilis gen. nov. et sp. nov.,
546 Crackipollenites gen. nov. et sp. nov., monosacate indeteterminate Form A and
547 Angustisulcites hexagonalis sp. nov., associated with well-known species of the Triassic
548 of Gondwana such as Alisporites/Falcisporites, Platysaccus, Pteruchipollenites,
549 Minutosaccus/Protodiploxypinus, Klausipollenites, Lunatisporites, Accinctisporites,
550 Sulcosaccispora, Striomonosaccites and Crustaesporites. Unlike the palynofloras of the
551 late Middle and Late Triassic of Argentina (Zavattieri & Batten, 1996), the inaperturate,
552 polyplicate and monsulcate pollen grains appear in very low proportions in the microflora
553 of the Quebrada de los Fósiles Formation.
554 This unique and exceptional novel microfloral assemblages of the lower unit of the
555 Puesto Viejo Group are integrated by diverse terrestrial pollen grains and spores among
556 rich freshwater green algal microfossils and dispersed fungal spores (Zavattieri et al.,
557 2020). Most of these algae, fungi and terrestrial components constitute the first Triassic
558 records for the entire Gondwanan continent. The terrestrial and aquatic vegetation grew
559 associated with shallow water bodies in floodplain environments developed under warm
560 temperate, strongly seasonal climatic conditions when profuse volcanic activity
561 introduced large volumes of ash and greenhouse gases into the atmosphere. The volcanic
562 events developed during deposition of the Quebrada de los Fósiles Formation promoted
25
563 changes in the environmental conditions that influenced the development of the peculiar
564 composition of the vegetation and the algae community of the water bodies, creating
565 stressful conditions and the development of the plant communities within the ecosystem
566 (Spalletti et al., 2003; Cariglino et al., 2018).
567 ACKNOWLEDGMENTS
568 This research has been partially funded by the National Agency for Scientific and
569 Technological Promotion, Argentina (research grants ANPCYT-PICT 2016-0637 and
570 ANPCYT-PICT 2016-0431). Parts of the research that led to the revision of this material
571 were financially supported by the Consejo Nacional de Investigaciones Científicas y
572 Técnicas (CONICET), Argentina (PIP Nº 0705 PRG and PIP Nº 11220090100605
573 AMZ). We are deeply indebted to Dr. Mariana Monti for her assistance in the field
574 works, interpretation and information about the geology of the Puesto Viejo Group, as
575 well as to Dr. Bárbara Cariglino for her invaluable assistance in the field. Thanks to A.
576 Moschetti (IANIGLA-CCT-CONICET-Mendoza) for palynological laboratory
577 preparations of samples for light optical microscopy studies, as well as to F. Tricárico
578 who assisted with the scanning electron microscopy.
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37
819
820 Figure captions
821 Figure 1. Location maps (taken from Zavattieri et al., in press). A, Distribution of the
822 Triassic basins in central-western of Argentina, showing the location of the San Rafael
823 Basin. B, Outcrops of the Puesto Viejo Group in the San Rafael Basin. C, Geological
824 map of the Puesto Viejo Group in the study area. Locations of the palynological sampling
825 sections in the Quebrada de los Fósiles Creek: 1, GzD; 2, AMZ1; 3, AMZ2 (see Fig. 2);
826 4, location of the sampling in the section Río Seco de la Quebrada Creek; Qda. Quebrada;
827 Fm: Formation.
828 Figure 2. Integrated sedimentological section of the Quebrada de los Fósiles Formation
829 at the type locality (Agua de los Burros Creek; taken from Zavattieri et al., in press).
830 Characteristics of the sections studied from the type locality and the locations of the
831 studied palynological samples (see Fig. 1, Tab. 1). Stars show sample dating levels of the
832 basal and top effusive levels of the type section.
833 Figure 3. Sedimentological section of the Quebrada de los Fósiles Formation cropping
834 out at the Río Seco de la Quebrada Creek succession, indicating the characteristics of the
835 sequence and the location of the studied palynological samples (Fig. 1, Tab. 2; taken from
836 Zavattieri et al., in press).
837 Figure 4. 1−6, Mendozapollenites variabilis gen. et sp. nov. 1, MPLP 5877(C) C44(1),
838 holotype; 2, MPLP 10348(G) J44(0), paratype; 3, MPLP 5877(C) H40(0), paratype; 4,
839 MPLP 5877(E) S52(4), paratype; 5, MPLP 5877(C) F53(1), paratype; 6, BAPal 6612(7)
840 D26(1), paratype. Scale bar, 20 µm.
38
841 Figure 5. 1-8, Mendozapollenites variabilis gen. et sp. nov. 1, MPLP 5877(C) U38(0),
842 paratype; 2, MPLP 5877(C) F51(0), paratype; 3, MPLP 5877(E) G48(0), paratype; 4,
843 BAPal 6612(7) P45(4), paratype; 5, MPLP 10348(G) R38(0); 6, MPLP 10352(E) K53(0);
844 7, BAPal 6614(8) E43(0); 8, MPLP 10348(G) W39(4), paratype. Scale bar, 20 µm.
845 Figure 6. 1-7, Mendozapollenites variabilis gen. et sp. nov. 1, BAPal MEB 6612(TM)
846 f011, distal face; 2, BAPal MEB 6612(TM) f029, paratype, proximal face; 3, BAPal
847 MEB 6612(TM) f036, paratype, distal face; 4, BAPal MEB 6612(TP) f050, paratype,
848 distal face; 5, BAPal MEB 6612(T1) f042, paratype, distal face; 6, MPLP MEB
849 10374(TJ) f011, paratype, proximal face; 7, BAPal MEB 6612(TQ) f013, paratype,
850 proximal face. 8-11, Monosaccate indeterminate Form A. 8, BAPal 6612(3) O47(0); 9,
851 BAPal 6612(3) J32(1); 10, MPLP 10362(E) U34(2); 11, MPLP 6612(3) B50(0); Scale
852 bar, 20 µm.
853 Figure 7. 1-7, Angustisulcites hexagonalis sp. nov., 1, MPLP 10348(D) X53(1), paratype,
854 2, MPLP 10348(G) P44(1); 3, MPLP 10348(C) Y52(1) paratype, 4, MPLP 10348(C)
855 X37(2); 5, MPLP 10348(G) Y33(0), holotype; 6, MPLP 10348(D) J37(0); 7, MPLP
856 10376 (E) B55(0), paratype. 8-12. Crackipollenites polygonalis gen. et sp. nov., 8, BAPal
857 6612(8) Y53(3), paratype; 9, BAPal 6612(8) F27(4); 10, BAPal 6612(3) K48(2); 11,
858 BAPal 6613(2) D37(2); 12, BAPal 6614(1) U48(0). Scale bar, 20 µm.
859 Figure 8. 1-8. Crackipollenites polygonalis gen. et sp. nov., 1, MPLP 5877(C) F49(0),
860 holotype; 2, MPLP 5877(D) U47(4), paratype; 3, MPLP 5877(F) M35(1), paratype; 4,
861 MPLP 5877(F) W36(2), paratype; 5, MPLP 10379(E) G35(4); 6, MPLP 5877(C) R39(1);
862 7, MPLP 11056(G) N43(0); 8, BAPal 6614(1) R56(3). Scale bar X 20 µm.
39
863 Figure 9. 1-10. Crackipollenites polygonalis gen. et sp. nov., 1, BAPal 6614(2) V50(4);
864 2, MPLP 10348(D) K48(0); 3, BAPal MEB 6612(TN) f025, paratype; 4, BAPal 6613(2)
865 Y56(2); 5, BAPal MEB 6612(TN) 045, paratype; 6, BAPal 6612(3) M37(1). 7-11.
866 Crackipollenites sp. A, 7, MPLP 10348(D) V52(0); 8, BAPal MEB 6612(TQ) f016; 9,
867 MPLP 11054(D) B42(3); 10, MPLP 10379(E) V36(0); 11, MPLP 10348(C) X33(3).
868 Scale bar, 20 µm.
869 Table 1. Composition and distribution (in %) of miospore groups and the main saccate
870 pollen genera recorded in the Quebrada de los Fósiles Formation, Puesto Viejo Group in
871 the sections integrating the type section, at the Quebrada de los Fósiles Creek.
872 Table 2. Composition and distribution (in %) of miospore groups and the main saccate
873 pollen genera recorded in the Quebrada de los Fósiles Formation, Puesto Viejo Group at
874 Río Seco de la Quebrada Creek section.
40
TABLE 1. Composition and distribution (in %) of miospore groups and the main saccate pollen genera recorded in the Quebrada de Los Fósiles Formation, Puesto Viejo Group in the sections integrating the type section, at the Quebrada de los Fósiles Creek.
Quebrada de los Fósiles Creek section
GzD section AMZ1 section AMZ2 section
Samples (prefixes MPLP/*BAPal) 9077 9077 9078 9079 9080 9081 9082 9085 5877 11064 10362 *6612 *6613 *6614 11068 10363 11071 10366 11072 10372 11073 10373 9076 9076 FUNGI 0.2 0 0 0 1.0 0 0 0.5 1.2 1.0 0 0 0.2 0.2 0.2 0 0 0 0 0 0 0 ALGAE 96.4 83.3 94.6 93.0 97.9 96.3 93.5 41.8 1.2 5.6 2.5 0.5 0 0.2 1.3 77.0 2.5 2.2 27.1 1.0 4.9 10.3 INCERTAE SEDIS 0 0 0 0 0 0 0 0 0 0 0.5 0 0 0 0 0.2 0 0 0.2 0 0 0 MONOLETE SPORES 0 4.5 1.0 0.8 0 0 1.3 23.8 2.1 85.4 58.6 59.1 69.9 64.4 97.7 17.5 91.0 81.5 29.6 89.3 61.0 73.2 TRILETE SPORES 2.0 3.8 2.2 1.5 0.5 1.5 1.7 14.8 12.0 5.2 28.4 31.4 21.6 25.3 0.6 3.2 3.3 15.3 2.8 9.2 24.7 13.2 MONOSACCATE NON-TAENIATE PG 0 1.5 0 0 0 0 0.2 3.3 35.9 0.2 2.2 3.6 2.0 2.5 0 0.2 0.2 0 2.2 0 0.8 0 Accinctisporites 0 1.5 0 0 0 0 0 0.5 3.5 0.2 0.5 0.8 0.2 0 0 0 0.2 0 2.0 0 0.8 0 Cladaitina 0 0 0 0 0 0 0.2 2.8 32.4 0 1.5 2.8 1.8 2.5 0 0 0 0 0.2 0 0 0 Sulcosaccispora 0 0 0 0 0 0 0 0 0 0 0.2 0 0 0 0 0 0 0 0 0 0 0 MONOSACCATE TAENIATE PG 0 1.5 0 0 0 0 0.2 6.0 10.3 0 2.2 0.5 0.8 1.5 0 0.2 0.2 0.2 0.8 0 0 0 Mendozapollenites 0 0 0 0 0 0 0 6.0 10.1 0 0.8 0.5 0.8 1.5 0 0.2 0.2 0.2 0.8 0 0 0 Crustaesporites 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Striomonosaccites 0 0 0 0 0 0 0 0 0.2 0 0 0 0 0 0 0 0 0 0 0 0 0 BISACCATE NON-TAENIATE PG 1.0 4.7 1.8 1.2 0.2 0.7 0 6.9 28.6 1.7 4.3 3.5 5.1 5.0 0.2 1.9 1.4 0.4 33.3 0.5 5.2 1.4 BISACCATE TAENIATE PG 0.4 2.0 0.2 3.5 0.4 1.3 0.5 1.5 7.0 0.5 0.2 1.2 0.4 0.2 0 0 0.4 0.2 2.7 0 0.9 0.4 Protohaploxypinus 0.2 1.5 0 3.0 0.2 1.3 0.5 1.5 4.8 0 0.2 0.5 0 0 0 0 0.2 0 2.5 0 0.5 0.2 Angustisulcites 0 0 0 0 0.2 0 0 0 0.2 0 0 0.2 0.2 0.2 0 0 0.2 0 0 0 0 0 Striatoabieites 0 0.5 0 0 0 0 0 0 1.0 0.5 0 0 0.2 0 0 0 0 0 0.2 0 0.2 0 Striatopodocarpites 0.2 0 0.2 0.5 0 0 0 0 1.0 0 0 0.5 0 0 0 0 0 0.2 0 0 0.2 0.2 Hamiapollenites 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Lunatisporites 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 INAPERTURATE PG 0 0 0 0 0 0 2.8 0.9 0.2 0.2 0.8 0 0 0.5 0 0 1.0 0.2 1.3 0 2.5 0 HILATE PG 0 0 0.2 0 0 0 0 0 1.3 0.2 1.0 0 0 0.2 0 0 0 0 0 0 0 1.5 Crackipollenites 0 0 0 0 0 0 0 0 1.3 0.2 1.0 0 0 0.2 0 0 0 0 0 0 0 1.5 SULCATE PG 0 0.2 0 0 0 0 0 0.5 0 0 0.5 0 0 0 0 0 0 0 0 0 0 0 POLYPLICATE PG 0 0 0 0 0 0.2 0 0 0 0 0.2 0.2 0 0 0 0 0 0 0 0 0 0 TOTAL 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100
TABLE 2. Composition and distribution (in %) of miospore groups and the main saccate pollen genera recorded in the Quebrada de Los Fósiles Formation, Puesto Viejo Group at Río Seco de la Quebrada Creek section.
Río Seco de La Quebrada Creek section
Samples (prefix MPLP) 10348 10374 11051 10375 11052 10376 11053 10377 10350 6543 6544 11054 10351 10379 11055 10352 11056 FUNGI 0.3 0.8 4.3 0 0 0 1.0 0.5 0.3 0 0.3 0.3 1.3 0.5 0.5 0.3 1.3 ALGAE 1.5 7.8 26.0 1.3 8.8 2.5 1.0 1.0 5.8 4.5 65.3 0.8 3.5 4.3 9.0 6.3 3.3 INCERTAE SEDIS 0 0 3.3 0 0.3 0 0.3 0.3 0 0 1.0 0 0 0 1.5 0 0.3 MONOLETE SPORES 0 0.8 18.0 82.0 11.8 43.0 74.0 88.5 35.3 0.3 3.5 0.5 0.5 8.3 1.3 28.8 1.0 TRILETE SPORES 12.8 40.0 42.5 14.0 44.3 46.5 21.5 8.8 33.5 95.0 25.8 63.0 11.5 15.5 83.5 47.0 19.6 MONOSACCATE NON-TAENIATE PG 4,3 9,3 1.0 0,2 6,7 1,0 0 0.5 2,5 0,0 0 2,5 4,8 4,0 0.5 1,3 5,3 Accinctisporites 2.5 2.3 0 0 1.2 0.5 0 0 1.8 0 0 2.3 2.5 1,5 0 0.5 2.0 Cladaitina 0.3 0 0.3 0 0 0 0 0 0 0 0 0.2 0 0.3 0.3 0 0.3 Sulcosaccispora 1.5 7.0 0.7 0.2 5.5 0.5 0 0.5 0.7 0 0 0 2.3 2.2 0.2 0.8 3.0 MONOSACCATE TAENIATE PG 2,2 1,7 0 0,3 0,3 1,0 0 0 3,3 0,3 0 4,0 2,2 1,3 0,0 1,0 1,0 Mendozapollenites 1.5 1.7 0 0.3 0.3 1.0 0 0 3.0 0.3 0 3.3 1.5 1.0 0 1.0 0.5 Crusteasporites 0 0 0 0 0 0 0 0 0 0 0 0.2 0 0 0 0 0 Striomonosaccites 0.7 0 0 0 0 0 0 0 0.3 0 0 0.5 0.7 0.3 0 0 0.5 BISACCATE NON-TAENIATE PG 66.0 34.0 0.5 0.5 22.3 3.5 1.2 0.3 15.5 0 0 15.5 39.8 46.3 1.3 14.5 49.9 BISACCATE TAENIATE PG 8.8 4.0 0 0.3 0.8 1.5 0.6 0 2.0 0 0 10.6 10.5 5.0 0 0 7.8 Protohaploxypinus 3.7 3.5 0 0.3 0.8 1.0 0.3 0 1.1 0 0 3.5 8.5 4.0 0 0 5.5 Striatopodocarpites 2.5 0 0 0 0 0.2 0 0 0.3 0 0 2.8 1.3 0 0 0 0.7 Angustisulcites 0.3 0.5 0 0 0 0 0.3 0 0 0 0 3.3 0 0.5 0 0 0.3 Striatoabieites 2.0 0 0 0 0 0.3 0 0 0.3 0 0 1.0 0.7 0.5 0 0 1.0 Hamiapollenites 0 0 0 0 0 0 0 0 0.3 0 0 0 0 0 0 0 0 Lunatisporites 0.3 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.3 INAPERTURATE PG 2.5 0.5 4.5 0.8 4.3 0 0.3 0.3 0.5 0 4.3 2.5 23.8 10.8 2.0 1.0 9.4 HILATE PG 0.3 0.3 0 0 0 0 0.2 0 0.5 0 0 0.3 0 0.5 0.3 0 0.3 Crackipollenites 0.3 0.3 0 0 0 0 0.2 0 0.5 0 0 0.3 0 0.5 0.3 0 0.3 SULCATE PG 1.5 0.8 0 0.8 0.8 0.3 0 0 1.0 0 0 0.3 2.0 3.8 0.3 0 1.0 POLYPLICATE PG 0 0.3 0 0 0 0.8 0 0 0 0 0 0 0.3 0 0 0 0 TOTAL 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100