Accepted Manuscript

New Mesozoic and Cenozoic fossils from Ecuador: Invertebrates, vertebrates, plants,
and microfossils

Edwin A. Cadena, Alejandra Mejia-Molina, Carla M. Brito, Sofia Peñafiel, Kleber J.

Sanmartin, Luis B. Sarmiento

PII: S0895-9811(17)30528-X
DOI: 10.1016/j.jsames.2018.02.004
Reference: SAMES 1876

To appear in: Journal of South American Earth Sciences

Received Date: 19 December 2017

Revised Date: 7 February 2018
Accepted Date: 8 February 2018

Please cite this article as: Cadena, E.A., Mejia-Molina, A., Brito, C.M., Peñafiel, S., Sanmartin, K.J.,
Sarmiento, L.B., New Mesozoic and Cenozoic fossils from Ecuador: Invertebrates, vertebrates, plants,
and microfossils, Journal of South American Earth Sciences (2018), doi: 10.1016/j.jsames.2018.02.004.

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 ACCEPTED MANUSCRIPT

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1 New Mesozoic and Cenozoic fossils from Ecuador: invertebrates, vertebrates, plants, and

2 microfossils

Edwin A. Cadena1*, Alejandra Mejia-Molina1, Carla M. Brito1, Sofia Peñafiel1, Kleber J.

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5 Sanmartin1, Luis B. Sarmiento1

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7 School of Geological Sciences and Engineering, Yachay Tech, San Miguel de Urcuquí,

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8 Ecuador <cadenachelys@gmail.com>

9 * Corresponding author

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11 Abstract.—Ecuador is well known for its extensive extant biodiversity, however, its
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12 paleobiodiversity is still poorly explored. Here we report seven new Mesozoic and Cenozoic

13 fossil localities from the Pacific coast, inter-Andean depression and Napo basin of Ecuador,
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14 including vertebrates, invertebrates, plants, and microfossils. The first of these localities is
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15 called El Refugio, located near the small town of Chota, Imbabura Province, from where we

16 report several morphotypes of fossil leaves and a mycetopodid freshwater mussel of the
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17 Upper Miocene Chota Formation. A second site is also located near the town of Chota,

18 corresponding to potentially Pleistocene to Holocene lake deposits from which we report the
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19 occurrence of leaves and fossil diatoms. A third locality is at the Pacific coast of the country,
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20 near Rocafuerte, a town in Esmeraldas Province, from which we report a late Miocene palm

21 leaf. We also report the first partially articulated skull with teeth from a Miocene scombridid

22 (Mackerels) fish from El Cruce locality, and completely preserved seeds from La Pila

23 locality, both sites from Manabí Province. Two late Cretaceous fossil sites from the Napo

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24 Province, one near Puerto Napo showing a good record of fossil shrimps and a second near

25 the town of Loreto shows the occurrence of granular amber and small gymnosperms seeds

26 and cuticles. All these new sites and fossils show the high potential of the sedimentary

27 sequences and basins of Ecuador for paleontological studies and for a better understanding

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28 of the fossil record of the country and northern South America.

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29 Keywords: Paleobiodiversity, Neotropics, Northern South America, Paleontology

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31 1. Introduction

32 The fossil record of Ecuador is still poorly explored and documented as has been recently

33
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discussed in a review study on the paleontology of the country (Cadena and Román-Carrión, in
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34 press). This is due to several factors including for instance the lack of specific and permanent
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35 paleontological fieldwork. In the following paragraphs we briefly describe some of the most

36 famous localities of Ecuador where fossil vertebrates, invertebrates, plants, and microfossils have
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37 been reported and studied.

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38 In terms of fossil vertebrates, Ecuador has shown to be rich, particularly for Neogene faunas,

39 including several Pleistocene and Holocene localities from Santa Elena Province (southwestern
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40 region of the country) (Ruiz-Sánchez et al., 2014; Lindsey and Lopez, 2015; Lindsey and

41 Seymour, 2015; Tanaka et al., 2017, Cadena et al., 2017, references therein), an important
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42 number of localities from the inter-Andean basin, including one from the historic center of Quito
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43 (capital of the country) (Román-Carrión, 2012a, 2012b, 2012c, 2012d; Carlini et al., 2013), and

44 the localities near the small town of Bolivar, Carchi Province (northern of the country)

45 (Ficcarelli et al., 1992, 1995; Fejfar et al., 1993, 1996; Ferretti, 2010; Tomiati and Abbazzi,

46 2002; Ficcarelli et al., 2003).

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47 Fossil invertebrates of Ecuador include freshwater, marine, and terrestrial forms. For

48 instance, Mesozoic mollusks reported and described from several localities of the eastern region

49 of the country (Amazon basin) (Aspden and Ivimey-Cook, 1992; Dommergues et al., 2004;

50 Bulot et al., 2005; Dhondt and Jaillard 2005). Cenozoic invertebrates, particularly mollusks,

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51 arthropods and ichnofossils have been reported and described from the inter-Andean depression,

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52 the Pacific coast and the Galápagos Islands (Walker, 2001; Di Celma et al., 2002; Landini et al.,

53 2002; Ragaini et al., 2002; Ragaini and Di Celma, 2009; Herrera and Román-Carrión, 2012;

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54 Sánchez et al., 2013; Zunino, 2013, references therein).

55 The fossil record of plants in Ecuador is restricted principally to the southern provinces in

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Mesozoic and Cenozoic sequences that include the famous petrified forest of Puyango
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57 (Shoemaker, 1982), and reports of leaves and seeds from different localities in Loja, El Oro, and

58 Azuay provinces (Berry, 1933; Burnham, 1995; Kowalski, 2001).

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59 The micropaleontology of Ecuador, as in many other countries has resulted highly

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60 beneficiated by the oil-exploration. However, as usual, much of this knowledge has not been
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61 published and makes part of internal reports or classified documents of the oil companies. In

62 spite of this, some studies describing microfossils and fossil palynomorphs of Ecuador include:
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63 Landini et al., 2002; Vallejo et al., 2002; Ordóñez et al., 2006; Niemann and Behling, 2008;

64 Collins et al., 2013; Bush et al., 2014, and references therein.

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65 Here we report and briefly describe seven new Mesozoic and Cenozoic localities and their
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66 fossil content from different regions of Ecuador, including vertebrates, invertebrates, plants, and

67 microfossils. We also discuss the relevance of these fossil findings for understanding the

68 biodiversity history of Ecuador and tropical South America, as well as their potential importance

69 for future paleobiological, paleoclimatic, and paleobiogeographical studies.

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71 2. New fossil localities and their geological setting

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73 2.1. El Refugio locality

74 Situated along the highway 35, which goes from Quito to Tulcán, approximately 800 m past the

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75 Ambuqui toll station, on the right side of the road (0º28’3.50”N, 78º2’1.17”W) (Fig. 1.1), town

76 of Chota, Ibarra Canton, Imbabura Province. The stratigraphic section (Fig. 2) makes part of the

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77 upper sequence (sequence S2) of the Chota Formation following Barragán et al., 1996, including

78 fine-grained lithic sandstones interbedded with siltstones, and layers of dark grey organic rich

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mudstones and laminated gypsum. Radiometric ages for the entire Chota Formation varies from
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80 28±2.0 to 1.1±0.6 Ma, with four of the five samples giving a range between 4.8±0.4 to 1.1±0.6
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81 Ma (Pliocene-Pleistocene), based on Winkler et al., (2005). In particular, radiometric ages

82 obtained from volcanic rocks overlying the Chota Formation along the Chota river by Barberi et
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83 al. (1988) indicate an age between 6.31±0.1 Ma and 6.30±0.06 Ma, suggesting that the fluvio-
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84 lacustrine deposits of the Chota Formation are the oldest of the stratigraphy sequence in this

85 region and should be at least Miocene in age (Barragán et al., 1996).

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86

87 2.2. Chota locality

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88 Located 200 m after the small town of Chota, crossing the Chota River bridge, on the left margin
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89 of the road (0º28’30.13”N, 78º3’57.37”W) (Fig. 1.2), Mira Canton, Carchi Province. This

90 outcrop is characterized by a thick bed of approximately 6 m of laminated white to pink

91 diatomite, pinched laterally and in contact with poorly consolidated fluvial deposits dominated

92 by conglomerates and conglomeratic sandstones. According to this stratigraphic configuration

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93 and the diatoms content (see microfossils section of this study) the age for this deposit is

94 hypothesized as Pleistocene to Holocene.

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96 2.3. El Cruce locality

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97 Located 300 m before the round-point on road 15 that goes from Manta to Rocafuerte,

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98 approximately 3 km before this town, Rocafuerte Cantón, Manabí Province (00°54’47.32”S,

99 80°29’16.526”W) (Fig. 1.3). The outcrop is dominated by calcareous mudstones, mapped as

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100 belonging to the Dos Bocas Formation (Egüez et al., 2017), which is considered late Oligocene

101 to middle Miocene by Bristow and Hoffstetter, (1977). A sample for calcareous nannofossils

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(YT-Nan-0001) was studied from this locality and the results and implications for the age of the
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103 section are presented in the discussion section of this study.
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105 2.4. La Pila locality

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106 Located 80 m after La Pila town on the road that goes from Montecristi to Jipijapa, Montecristi
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107 Cantón, Manabí Province (01°61’41.84”S, 80°34’53.10”W) (Fig. 1.4). This outcrop is

108 represented by siliceous siltstones, mapped as belonging to the Dos Bocas Formation (Egüez et
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109 al., 2017), which is considered late Oligocene to middle Miocene by Bristow and Hoffstetter,

110 (1977).
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111
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112 2.5. Rocafuerte locality

113 Located 1.2 km west of Rio Fuerte town, along the beach, Pacific coast, Rio Verde Canton,

114 Esmeraldas Province (1º04’28.62”N, 79º23’17.95”W) (Fig. 1.5). The outcrop is dominated by

115 calcareous fine-grained sandstones with abundant hematite laminae and nodules, mapped as

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116 belonging to the Viche Formation (Egüez et al., 2017), which is considered as lower to lower

117 middle Miocene in age (Cantalamessa et al., 2007).

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119 2.5. Puerto Napo locality

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120 Located 200 m before the town of Puerto Napo on the right margin of road 45 that goes from

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121 Tena to Puyo, Napo Province (1º02’31.76S, 77º47’38.54”W) (Fig. 1.6). This section is

122 represented by an interbedded sequence of biomicrites and grey to black shales, belonging to the

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123 upper segment of Napo Formation, Coniacian age (Bristow and Hoffstetter, 1977).

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125 2.6. Pungarayacu quarry

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126 Located 7 km after the small town of Jondachi, along 20 road that goes from Tena to Loreto,

127 Loreto Canton, Napo Province (0º42’24.14S, 77º44’28.78”W) (Fig. 1.7). An old quarry in the
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128 middle of a dense tropical forest exposes a sequence of limestones, interbedded with grey to
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129 black-oily siltstones and claystones potentially belonging to the middle segment of Napo
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130 Formation, Turonian in age (Bristow and Hoffstetter, 1977).

131
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132 3. Materials and methods

133 All macro fossil specimens and nannofossil samples reported and figured herein were prepared,
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134 studied, and deposited at the Paleontology Lab collections of Yachay Tech, San Miguel de
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135 Urcuquí, Imabura Province, Ecuador. Specimens and micropaleontological slides receive a

136 collection identification number starting with the university initials (YT) followed by an

137 abbreviation of the group to which they belong e.g.,: Ver (vertebrates), and finally a consecutive

138 serial number of four digits e.g.,: 0001.

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139 Most of the specimens were studied and photographed using an Olympus SZX16

140 stereomicroscope. Additionally some specimens of diatoms and small seeds were observed and

141 studied using a Phenom ProX Scanning Electron Microscope under 5 and 10 kV conditions (YT

142 Paleontology Lab facility). Preliminary taxonomic identifications of the fossils described herein

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143 were consulted with experts in each of these groups, and are referred to as personal

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144 communications in each of the sections that we described the fossils.

145

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146 4. Fossil invertebrates

147 4.1. Bivalvia

148
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Large pearly freshwater mussels were found in a siltstone layer at the El Refugio locality (middle
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149 segment of the section, Fig. 2) (Fig. 3.1). At least six specimens (YT-Inv-0001-0006) were
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150 collected, being the largest one being 10.5 cm long. These bivalves belong to the

151 Mycetopodidiae family (F.P. Wesselingh, personal communication, 2017).

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152 4.2. Decapoda

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153 Shrimps preserving tails, arms, and claws with delicate cuticles were found in the Puerto Napo

154 locality, Napo Formation (Fig. 3.2–3.5) (YT-Inv-0007-0011). A total of seven specimens were
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155 found, and these seem to belong to the infraorder Thalassinidea (J. Luque, personal

156 communication, 2017).

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157

158 5. Fossil vertebrates

159 5.1. Actinopterygii

160 A partially complete skull, considerably crushed, but preserving in place most of its teeth in the

161 premaxilla and dentary (Fig. 4.1–4.3) (YT-Ver-0001). The specimen was found at El Cruce

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162 locality, Manabí Province. This fish is potentially a representative of the Scombridae family,

163 resembling the Scomberomorus genus, which include also the extant mackerels (J. Carrillo,

164 personal communication, 2018).

165

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166 6. Fossil plants

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167 6.1. Angiosperms

168 Several indeterminate families of fossil leaves and seeds were found in four different horizons at

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169 the El Refugio locality, Chota Formation (Fig. 2). At least thirteen different morphotypes have

170 been identified, including large leaves with entire margin, some with dentate margin, two

171
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different type of grasses, and two potential small seeds (Fig. 5.1–5.8) (YT-Bot-0001-0022, 0028)
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172 (Supplementary data 1). Insect damage has also been recognized in some of these fossil leaves,
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173 as well as different levels of veins structures (Fig. 5.3).

174 Other locality from which we have found fossil leaves is the Rocafuerte locality, Viche
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175 Formation. Here we found only one small leaf that potentially represents a palm, with abundant
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176 oily microbubbles attached to the cuticle surface (Fig. 5.9) (YT-Bot-0025).

177 Fossil leaves have also been found in the paleolake deposits of the Chota basin particularly at
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178 the Chota locality. The fossils are found well-preserved in the diatomite layers; however we only

179 have identified one morphotype characterized by long leaves with entire margin and low degree
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180 of venation (Fig. 5.10) (YT-Bot-0023-0024).

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181 A large drift seed was found associated with abundant plant remains in La Pila locality, Dos

182 Bocas Formation (YT-Bot-0029) (Supplementary data 1). This fossil seed resembles in shape

183 and size the seeds of the extant Hura genus (S. Manchester, personal communication, 2017).

184 6.2. Gymnosperms

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185 Small seeds and cuticle remains well-preserved were recovered from grey siltstones rich in

186 organic matter and other plant remains (Fig. 6.1–6.5) (YT-Bot-0026), found at the Pungarayacu

187 quarry, Napo Province. These seeds potentially belong to the Gnetales order (F. Herrera,

188 personal communication, 2017).

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189

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190 6.3. Amber

191 Granular amber that varies from coarse grains to pebbles, are very abundant in the same

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192 siltstones that the Gnetales seeds and cuticles were found (YT-Bot-0027, Supplementary data 1).

193 At present, none of these amber granules have shown to preserve fossils, only abundant “dusty”

194 like inclusions and potentially microbubbles.

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195
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196 7. Microfossils

197 7.1. Diatoms

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198 At least twelve different taxa have been found in the diatomites from the paleolake deposits of
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199 the Chota basin, Chota locality. Some of the taxa identify include: Cocconeis placentula (Fig.

200 6.6) and Navicula cryptocephala (Fig. 6.7) (YT-Dia-0001-0002). (S. Fritz, personal
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201 communication, 2017), (Supplementary data 2).

202
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8. Discussion
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203

204 8.1. Relevance of the new fossils

205 The fossil fish reported here from El Cruce locality, Manabí Province represents the first skeletal

206 remain with in-place dentition reported for this region of Ecuador, and potentially the first fossil

207 record of scombridid from the Pacific margin of South America, from the late Miocene, based on

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208 the identification of the calcareous nannofossils NN8 zone, represented by Catinaster coalitus

209 (this study). Isolated marine fish remains principally teeth and otoliths assemblages have been

210 reported from the Pliocene Onzole Formation, Esmeraldas Province (Bianucci et al., 1993;

211 Carnevale et al., 2011), and nearly complete freshwater fish skeletons have been reported from

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212 the Loja Province (White, 1927; Costa, 2011).

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213 The marine fossil shrimps from Puerto Napo locality represent the first record of these

214 decapods in the Mesozoic of Ecuador. Fossils of freshwater shrimps have been reported from the

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215 Miocene of southern Ecuador (Herrera and Román-Carrión, 2012).

216 The freshwater mussels from the Chota Formation represent the first report of these bivalves

217
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in the Chota basin and at the same time represent the westernmost fossil record of the
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218 Mycetopodidiae family in South America. Thus, considering that the age of this sequence is

219 potentially late Miocene (Barragán et al., 1996), make these fossil bivalves have relevance to
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220 establish a possible west extension of the late Miocene Pebas wet-land system that existed
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221 previous to the origin of the current Amazon drainage system, a study that is currently being
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222 developed by the senior author. Fossil invertebrates particularly gastropods were only previously

223 mentioned in the general description of the Chota Formation (Barragán et al., 1996), however,
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224 we did not find evidence of these at the Chota Formation localities explored.

225 Fossil leaves from the El Refugio locality (Chota Formation) represent the first occurrence of
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226 this type of fossils in northern Ecuador and could contribute to the understanding of
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227 paleovegetation changes of this region during the onset of the uplifting of this part of South

228 American Andes. As well as, represent a good site for comparisons with the floristic composition

229 and paleobiogeography distribution between this and the Miocene south localities of the country,

230 particularly with the Nabon basin fossils (Kowalski, 2001).

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231 The small seeds and cuticles potentially from gymnosperms (Gnetales), as well as the

232 granular amber from Pungarayacu locality offer a new opportunity for understanding aspects of

233 the vegetation and possibly climatic conditions of this part of South America during the late

234 Cretaceous.

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235 Finally, the fossil diatoms from the lake deposits of the Chota basin, particularly from the

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236 Chota locality constitute the first fossil record of this type of microfossils in northern Ecuador

237 and could shed light on paleoenvironmental conditions of this region of the country during the

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238 Neogene and the Quaternary, as well as for correlation with other paleolake deposits along the

239 South American Andes.

240
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241 9. Conclusions
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242 The fossil sites and fossils reported and briefly described herein are evidence of the high

243 potential that the sedimentary rocks of Ecuador have to improve our knowledge on the
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244 paleobiodiversity not only of the country but also of northern South America. At the same time,
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245 with this work we show the poorly explored state in which many of the basins and sequences of

246 Ecuador are in terms of their fossil content. We hope this study will trigger other near future
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247 contributions describing and studying in detail the taxonomic, systematic paleontology, as well

248 as to establish the paleoenvironmental, paleobiogeographical and paleoclimatic implications of

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249 the fossils described herein. Also with this type of contributions we hope to promote and support
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250 the paleontological activity and research in Ecuador.

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252 Acknowledgments

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253 We thank to E. Mariño, D. Rojas, R. Madden, J. Román, K. Choez for assisting during the

254 fieldwork. We also thank to F. Mejia from the Instituto Nacional de Patrimonio Cultural (INPC)

255 of Ecuador for helping us to obtain permits to work at the Chota basin. Special thanks to J.

256 Carrillo, G. Ballen, J. Luque, S. Fritz, F. Herrera, S. Manchester, and F. Wesselingh for their

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257 personal comments on the taxonomy of the fossils reported here. Thanks to the reviewers J.

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258 Carrillo and J. Abella for their comments and suggestions to improve this manuscript.

259

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260 Funding

261 Funding for this study was provided by the School of Geological Sciences and Engineering of

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Yachay Tech, and the internal grants program of this univeristy, grants 27 and 09.
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263 Accessibility of supplemental data
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264 Supplementary data 1. Complete catalogue of the leaves morphotypes from El Refugio locality,

265 Chota basin and drift seed from La Pila locality.

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266 Supplementary data 2. Complete catalogue of the fossil diatoms from Chota locality, Chota
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267 basin.

268 References
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269 Aspden, J.A. and Ivimey-Cook, H.C., 1992. Nuevos datos paleontológicos del Centro y Sureste
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329 First record a copemyine-peromyscine cricetid (Rodentia, Mammalia) in South America:

330 hypotheses regarding its ancestry in the Palaeartic. Acta Zoologica Cracoviensia v. 39, 137–
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331 145.
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332 Ferretti, M.P., 2010. Anatomy of Haplomastodon chimborazi (Mammalia, Proboscidea) from the
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338 on the South American mastodons referred to Haplomastodon and Cuvieronius. Geobios v.

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340 Ficcarelli, G., Coltorti, M., Moreno-Espinosa, M., Pieruccini, P.L., Rook, L. and Torre, D., 2003.

341 A model for the Holocene extinction of the mammal megafauna in Ecuador. Journal of

342 South American Earth Sciences , v. 15, 835–845.

343 Herrera, M.M., and Román-Carrión, J.L. 2012. Registro de camarones de río en el Mioceno

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345 Kowalski, E.A., 2001. Middle to Late Miocene environment of Southern Ecuador: Temperature,

346 elevation, and fossil plants of the Nabón basin [Ph.D. thesis]. The University of Michigan,

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347 Ann Arbor, 451 p.

348 Landini, W., Bianucci, G., Carnevale, G., Ragaini, L., Sorbini, C., Valleri, G., Bisconti, M.,

349
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350 the development of the Panamic bioprovince after the rising of Central American Isthmus.

351 Canadian Journal of Earth Science, v. 39, 27–41.

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352 Lindsey, E.L. and Lopez R., 2015. Tanque Loma, a new late-Pleistocene megafaunal tar seep
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353 locality from southwest Ecuador. Journal of South American Earth Sciences v. 57, p. 61–82.
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354 Lindsey, E.L. and Seymour K.L., 2015. “Tar pits” of the western neotropics: Paleoecology,

355 taphonomy, and mammalian biogeography. In: J.M. Harris (Ed.), La Brea and Beyond: The
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356 Paleontology of Asphalt-preserved Biotas. Natural History Museum of Los Angeles County

357 Science Series nº 42, p. 111–124.

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358 Niemann, H. and Behling, H., 2007. Late Quaternary vegetation, climate and fire dynamics
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359 inferred from the El Tiro record in the southeastern Ecuadorian Andes. Journal of

360 Quaternary Science, v. 23, 203–212.

361 Ordóñez, M., Jiménez, N. and Suárez, J., 2006. Micropaleontología ecuatoriana. Centro de

362 Investigaciones Geológicas, Guayaquil, 634 p. [in Spanish]

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363 Ragaini, L., Bianucci, G., Cantalamessa, G., Valleri, G., and Landini, W., 2002. Paleoecology

364 and paleobiogeography of fossil mollusks from Isla Isabela (Galápagos Ecuador). Journal of

365 South American Earth Sciences 13:381–389.

366 Ragaini, L. and Di Celma, C., 2009. Shell structure, taphonomy and mode of life of a Pleistocene

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367 ostreid from Ecuador. Bolletino della Societa Paleontologica Italiana, v. 48, 79–87.

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369 Quito. Revista Politécnica v. 30, 136–146. [in Spanish]

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376 Ruiz-Sánchez, F.J., Abella, J., Román-Carrión, J.L., Lindsey, E., Santana, J., López, E.,
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377 Marquina, R., Crespo, V.D., Mansino, S., Díez, N., Ramírez, M., Vera, D., Escalante, K.M.,

378 Flores, F.F., Molina, J.O., Ramos, M.B., Ronquillo, I.L. and Cornejo, M.H., 2014. Nuevos
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379 datos sobre las faunas fósiles de vertebrados de la zona de Quebrada Seca (Santa Elena,

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381 215–218. [in Spanish]

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383 beetle brood balls from Pleistocene highland palaeosols of Andean Ecuador: A reassessment

384 of Sauer's Coprinisphaera and their palaeoenvironments. Palaeogeography,

385 Palaeoclimatology, Palaeoecology, v. 386, 257–274.

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386 Shoemaker, R.E., 1982. Fossil leaves from the lower Cretaceous Ciano Formation southwestern

387 Ecuador. Palaeontographica B, v. 180, 120–132.

388 Tanaka, Y., Abella, J., Aguirre-Fernández, G., Gregori, M., Fordyce, R.E. 2017. A new tropical

389 Oligocene dolphin from Montañita/Olón, Santa Elena, Ecuador. PLoS ONE 12:e0188380.

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390 Tomiati, C. and Abbazzi, L., 2002. Deer fauna from Pleistocene and Holocene localities of

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391 Ecuador (South America). Geobios v. 35, 631–645.

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394 Ecuador. Cretaceous Research, v. 23, 845–859.

395
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396 Upper Pliocene of Ecuador. Palaeogeography, Palaeoclimatology, Palaeoecology, v. 166,

397 141–163.
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398 White, E.I., 1927. On a Fossil Cyprinodont from Ecuador. Annals and magazine of natural
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399 history, v. 20, 519–522.

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400 Winkler, W., Villagómez, D., Spikings, R., Abegglen, P., Tobler, St., and Egüez, A., 2005. The

401 Chota basin and its significance for the inception and tectonic setting of the inter-Andean
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402 depression in Ecuador. Journal of South American Earth Sciences, v. 19, 5–19.

403 Zunino, M., 2013. The first dung beetle retrieved from Coprinisphaeridae ichnofossils: Phanaeus
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405 Mexicana, v. 29, 219–226.

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409

410

411 Figure Captions

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412 Figure 1. Map of Ecuador showing the new six fossil localities describe herein: (1) El Refugio

413 locality, Chota Formation, Imbabura Province; (2) Chota locality, paleolake deposits, Carchi

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414 Province; (3) El Cruce locality, Dos Bocas Formation, Manabí Province; (4) La Pila locality,

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415 Dos Bocas Formation, Manabí Province; (5) Rocafuerte locality, Viche Formation, Esmeraldas

416 Province; (6) Puerto Napo locality, Napo Formation, Napo Province; (7) Pungarayacu quarry,

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417 Napo Formation, Napo Province.
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419 Figure 2. Stratigraphic column of the Chota Formation, El Refugio locality, showing the fossil
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420 leaves and freshwater bivalves horizons.

421
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422 Figure 3. Fossil invertebrates from Ecuador reported herein: (1) Freshwater mussel (YT-Inv-
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423 0005) belonging to the Mycetopodidiae family (F.P. Wesselingh, personal communication, 2017)
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424 from El Refugio locality; (2) Posterior portion of a fossil shrimp with the tail (YT-Inv-0009),

425 Puerto Napo locality; (3) Partial arm with claw of a fossil shrimp (YT-Inv-0007), Puerto Napo
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426 locality; (4) Proximal segment of an arm of a fossil shrimp (YT-Inv-0010), Puerto Napo locality;
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427 (5) Close-up of margin of YT-Inv-0010 (red rectangle in (4)). Scale bars are (1) 2 cm; (2) 3 cm;

428 (3, 4) 10 mm; (5) 1 mm.

429

430 Figure 4. Scombridid (J. Carrillo, personal communication, 2017) partial fossil fish skull, from

431 El Cruce locality (YT-Ver-0001): (1) Dorsolateral view showing the articulated (crushed) right

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432 maxilla, premaxilla and dentary; the print and a fragment of the right dentary also can be

433 observed (2) Close-up of the premaxilla and dentary showing teeth in-place; (3) Close-up of the

434 teeth (red rectangle in (2)). Scale bars are (1) 3 cm; (2) 5 mm; (3) 1 mm. Abbreviations: lde, left

435 dentary; rde, right dentary; rmx, right maxilla; rpm, right premaxilla.

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436

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437 Figure 5. Fossil leaves and seeds from El Refugio (1-7), Rocafuerte (8), and Chota (9) localities:

438 (1) Leaf with dentate margin (YT-Bot-0001); (2) Leaf with entire margin (YT-Bot-0014); (3)

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439 Detail of the venation pattern of YT-Bot-0012; (4) Leaf elongated with entire margin (YT-Bot-

440 0019); (5) Close-up of red rectangle area in (4), showing microverrucate texture; (6) Fern leaf

441
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(YT-Bot-0028); (7) Potential fossil seed (YT-Bot-0005); (8) Grass leaf (YT-Bot-0022); (9) Palm
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442 leaf fragment (YT-Bot-0025), and one of its oily microbubbles shown at the upper right corner

443 (scale bar 5 µm); (10) Leaf with entire margin (YT-Bot-0024), from the paleolake deposits of
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444 Chota locality. Scale bars are (1, 6, 8, 10) 5 mm; (2, 4, 9) 10 mm; (3, 7) 2 mm; (5) 1 mm.
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445
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446 Figure 6. Small seeds and diatoms from the Pungarayacu and Chota localities respectively: (1)

447 Small seed from Gnetales (F. Herrea, personal communication, 2017) (YT-Bot-0026); (2) SEM-
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448 photograph of the proximal region of YT-Bot-0026; (3) YT-Bot-0026 after being released from

449 the rock; (4) Fragment of a cuticle (YT-Bot-0026); (5) SEM-photograph of the cuticle shown in
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450 (4); (6) Cocconeis placentula diatom from Chota locality (YT-Dia-0001); (7) SEM-photograph
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451 of Navicula cryptocephala diatom (YT-Dia-0001), from Chota locality. Scale bars are (1, 4) 1

452 mm; (3) 0.5 mm; (6) 10 µm. SEM-photographs have the scale at the bottom black stripe, as well

453 as the magnification and voltage information.

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Research highlights

• The first skeletal fossil record of sksphyraenids from the Pacific margin of South
America
• The westernmost fossil record of the Mycetopodidiae family in South America
• Contribution to the knowledge of the paleobiodiversity of Ecuador

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