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The first dinosaur remains from the Cretaceous of Ecuador

Sebastián Apesteguía, John E. Soto Luzuriaga, Pablo A. Gallina, José Tamay

Granda, Galo A. Guamán Jaramillo

PII: S0195-6671(19)30302-7
DOI: https://doi.org/10.1016/j.cretres.2019.104345
Reference: YCRES 104345

To appear in: Cretaceous Research

Received Date: 23 July 2019

Revised Date: 25 October 2019
Accepted Date: 29 November 2019

Please cite this article as: Apesteguía, S., Soto Luzuriaga, J.E., Gallina, P.A., Granda, J.T., Guamán
Jaramillo, G.A., The first dinosaur remains from the Cretaceous of Ecuador, Cretaceous Research,
https://doi.org/10.1016/j.cretres.2019.104345.

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 1 The first dinosaur remains from the Cretaceous of Ecuador

3 Sebastián Apesteguía1, John E. Soto Luzuriaga2, Pablo A. Gallina1, José Tamay Granda2,

4 Galo A. Guamán Jaramillo2

5
1
6 Fundación de Historia Natural “Félix de Azara” – Univ. Maimónides. Hidalgo 755, 7º piso

7 (1405) Buenos Aires, Argentina. sebastian.apesteguia@fundacionazara.org.ar;

8 pablo.gallina@fundacionazara.org.ar
2
9 Departamento de Geología y Minas e Ingeniería Civil. Universidad Técnica Particular de

10 Loja. San Cayetano Alto, Apartado Postal 11-01-608, Ecuador. jesoto@utpl.edu.ec;

11 jvtamay@utpl.edu.ec; gaguaman2@utpl.edu.ec

12 Correspondence: gaguaman2@utpl.edu.ec; sebastian.apesteguia@fundacionazara.org.ar

13

14

15

16
17 ABSTRACT

18 Yamanasaurus lojaensis gen. et sp. nov. is a new titanosaur (Saurischia, Sauropoda) from the

19 Upper Cretaceous of the Alamor-Lancones Basin, southern Ecuador. The fossil remains were

20 found in rocks of the Río Playas Formation, which is regarded as Campanian-Maastrichtian in

21 age. Remains include a partial sacrum, a partial mid-caudal vertebra, and several associated

22 limb bones. Yamanasaurus is characterized by: (1) anterior to mid-caudal vertebrae with

23 a dorsoventrally compressed condyle, with the posterior tip elevated respect to the midline, no

24 longitudinal ventral ridge, and spongy inner structure with absence of internal cavities (i.e.,

25 camellate bone, shared with Neuquensaurus); (2) last sacral centrum as long as tall,

26 with small ovoid, shallow blind fossa on the lateral side; and (3) radius robust with flattened

27 diaphysis and a marked neck or cingulum right under the epiphysis, with an heptagonal

28 concave proximal surface. Morphology, size, and age suggest that Yamanasaurus is closely

29 related to Neuquensaurus, being the northernmost saltasaurine known by far.

30

31 Key words: Titanosauria; South America; Loja; Late Cretaceous; Saltasaurinae

32

33 1. INTRODUCTION

34 The finding of dinosaur remains in South America has depicted a different history from that

35 shown in other parts of the world. This is based both in the quantity of remains and the oddity

36 of their species. Since the beginning of twenty century, the main countries that provided those

37 remains were Argentina and Brazil, but Chile, Peru, and Bolivia provided also a good

38 quantity of bones and thousands of trackways (e.g., Apesteguía, 2002b, Bonaparte and Coria,

39 1993, Bonaparte and Gasparini, 1980, Campos and Kellner, 1999, Campos et al., 2005, Coria

40 et al., 2003, Kellner and De Azevedo, 1999, Leanza et al., 2004, Meyer et al., 2001, Novas,

41 2009, Powell, 2003). Andean region was probably a vast, coastal corridor for north-south
42 faunal movement (Meyer et al., 2001). However, although Chile, Bolivia, and Peru show

43 abundant ichnological evidence of dinosaur presence, and the same countries plus Colombia

44 (e.g., Carballido et al., 2015) provided body remains, no evidence was reported yet from

45 Ecuador.

46 A recent finding of bones in the Cretaceous rocks of the Loja Area, in the

47 southernmost basins of Ecuador, provided dinosaur bones for the first time in this country.

48 Although it is obvious that this land was part of the vast South American dinosaur territory,

49 no evidence was collected previously from its Mesozoic outcrops.

50 The material here described was found close to the locality of Yamana, cantón Paltas,

51 in the Loja Province, SW Ecuador (Fig. 1A). After its finding by a local farmer, the main

52 material was deposited in the Instituto Nacional de Patrimonio Cultural whereas an additional

53 element of the same finding, two fused, articulated sacral vertebrae, was deposited in the

54 collection of the Universidad Técnica Particular de Loja for its research and conservation. The

55 bearing rocks are the yellowish sandstones of the Río Playas Formation, associated to

56 turbidites and conglomerate lenses belonging to the Río Playas Formation (Fig. 1B),

57 considered by the authors as late Campanian to early Maastrichtian as was suggested by

58 Kennerley (1973) and Jaillard et al. (1996).

59 The small sizes of the specimen, the anatomy, plus the proposed age are consistent

60 with its belonging to the Saltasaurinae, a group restricted to littoral regions from the end of

61 the Cretaceous and associated to dwarfism and insularity (Apesteguía, 2002; Leanza et al.,

62 2004). The presence in Ecuador of this kind of dinosaurs represents a considerable extension

63 of its distribution.

64

65 Institutional abbreviations

66 AMNH: American Museum of Natural History, USA.

67 NHMUK: The Natural History Museum, London, UK.

68 CM: Carnegie Museum, USA.

69 INPC: Instituto Nacional de Patrimonio Cultural, Ecuador.

70 INIGEMM: Instituto Nacional de Investigaciones Geológicas Mineras y Metalúrgicas,

71 Ecuador.

72 MACN: Museo Argentino de Ciencias Naturales ‘Bernardino Rivadavia’, Buenos Aires,

73 Argentina.

74 MCF-PVPH: Museo ‘Carmen Funes’, Plaza Huincul, Neuquén, Argentina.

75 MCS: Museo de Cinco Saltos, Río Negro, Argentina.

76 MJG-R: Museo ‘Jorge Gerhold’, Ingeniero Jacobacci, Río Negro, Argentina.

77 MLP: Museo de La Plata, La Plata, Argentina.

78 MPCA: Museo Provincial ‘Carlos Ameghino’, Cipolletti, Argentina.

79 MUCPV: Museo de la Universidad del Comahue, Neuquén, Argentina.

80 MPCF: Museo Provincial ‘Carmen Funes’, Plaza Huincul, Neuquén, Argentina.

81 UNP: Universidad Nacional de la Patagonia ‘San Juan Bosco’, Comodoro Rivadavia, Chubut,

82 Argentina.

83 UTPL: Universidad Técnica Particular de Loja, Ecuador.

84 YM; Yamana locality collection, Ecuador.

85

86 2. Systematic Paleontology

87 Saurischia Seeley 1888

88 Sauropoda Marsh 1878

89 Titanosauriformes Salgado, Coria and Calvo 1997

90 Titanosauria Bonaparte and Coria 1993

91 Saltasaurinae Powell 1992

 92 Yamanasaurus gen. nov.

93

94 Type species: Yamanasaurus lojaensis described below.

95 Etymology: Genus name refers to the provenance locality, Yamana, in the Casanga Valley,

96 southwestern Ecuador.

97 Diagnosis: as for the species.

98 Yamanasaurus lojaensis sp. nov.

99

100 Holotype: The type material is composed by one specimen preserved as two conjoined sacral

101 vertebrae (YM-UTPL_002), a fragment comprising approximately the posterior half of a

102 caudal procoelous vertebra (YM-INPC-014), the proximal half of a left humerus (YM- INPC-

103 016), the proximal half of a radius (YM- INPC-015), and a very eroded fragment possibly the

104 proximal part of a tibia (YM- INPC-017) (Fig. 1G).

105 Etymology: Named after the Province of Loja, where the material was found.

106 Locality: Yamana region, Paltas Cantón, Loja Province, southwestern Ecuador.

107 Age and horizon: Late Cretaceous, 66.9 My, late Maastrichtian, Río Playas Formation (=

108 Casanga Formation, Jaillard et al. 1999), Alamor-Lancones Basin.

109 Diagnosis: Yamanasaurus lojaensis is a saltasaurine titanosaur diagnosed by the following

110 autapomorphies (marked by an asterisk), as well as a unique combination of character states:

111 anterior to mid-caudal vertebrae with a dorsoventrally compressed condyle, with the posterior

112 tip elevated respect to the midline, no longitudinal ventral ridge, and spongy inner structure

113 with absence of internal cavities (i.e., camellate bone, shared with Neuquensaurus); last sacral

114 centrum as long as tall, with small ovoid, shallow blind fossa on the lateral side (*); radius

115 robust with flattened diaphysis and a marked neck or cingulum right under the epiphysis, with

116 an heptagonal proximal articular surface (*).

117

118 3. Material

119 All the studied material comes from the same area and was collected by one single

120 person, Mr. Víctor Francisco Celi Ríos, but the material was later housed in two different

121 institutions. Mr. Marco Antonio Paladines Balcázar donated the main group of bones to the

122 INPC collection whereas an additional material is preserved at the UTPL. Considering area,

123 size, and preservational features they probably belong to a single specimen. The collected

124 materials include one partial sacrum composed by two conjoined vertebral centra (YM-

125 UTPL_002), a fragment comprising approximately the posterior half of a caudal procoelous

126 vertebra (YM-INPC-014), the proximal half of a left ulna (YM- INPC-016), the proximal half

127 of a radius (YM- INPC-015), and a very eroded fragment possibly part of a tibia or an ulna

128 (YM- INPC-017).

129

130 4. Locality and horizon

131 The Alamor-Lancones Basin is located in SW Ecuador, as a branch of the northern

132 Perú Lancones Basin (Fig. 1). In Ecuador it includes the Western area of the Loja Province.

133 The stratigraphy, geochemical and geochronological aspects of this region were first studied

134 by Kennerley (1973), and later by Jaillard et al. (2004), Hungerbühler (1997, 2002), among

135 other. The “Celica-Lancones Basin s.s.” includes different tectonic unities with variegated

136 Lower Cretaceous stratigraphical series, separated by major faults. The units can be grouped

137 in a SE paleogeographical Province, mainly characterized by volcanoclastic deposits, and a

138 NW Dominion characterized by clastic deposits rich in detritic quartz (Jaillard et al., 1999).

139 The Casanga Valley is located in the western part of the Alamor-Lancones Basin, 18.8

140 Km from the Catacocha Town in the Paltas Cantón (Fig. 1A). The valley shows a rhomboidal
141 shape with its major axis oriented NE-SW and with levels reaching 730 to 1,250 m.o.s.l. what

142 makes its weather dry and hot. The Casanga Valley forms the sedimentary basin of the Río

143 Playas, which is at the same time the NE part of the Alamor-Lancones Basin, where outcrop

144 sedimentary Late Cretaceous to Paleogene rocks (Jaillard, et al., 1999). The local stratigraphy

145 remains poorly studied and despite INIGEMM 2013 proposed new units, there are still several

146 stratigraphic inconsistences in the basin. We describe here these sequences based on the

147 INIGEMM (2013) and other authors.

148 The basement of the basin is formed by the gabbro and basalts of the Punta de Piedra

149 Formation, a unit transitionally covered by the Celica Unit, composed by breccia, basalt–

150 andesitic lavas and hyaloclastites intruded by the Tangula Batolite. After stratigraphic

151 correlations with rocks of the Punta de Piedras Formation (Alamor Basin), it would

152 correspond to the Early Cretaceous (Egüez and Poma, 2001). The Bramaderos Unit

153 transitionally covers the Celica Unit. It is constituted by agglomerates, tuffs, grauwaques and,

154 sparsely, basalt-andesitic lavas which form the base of the Naranjo and Casanga Units, which

155 also are overlaying them unconformably. Since the Bramaderos Unit is covered by

156 Cenomanian-Turonian sedimentary rocks (Jaillard et al,. 1999), it is considered as Albian-

157 Cenomanian in age.

158 The La Ramada Unit constitutes a sequence of grey, agglomerate tuffs, interbedded

159 with light grey to dark brown sandstones and brownish grauwacques. It covers in concordance

160 the volcanoclastic sequence of the Bramaderos Unit and is unconformably covered by the

161 Naranjo Unit. After its stratigraphic position they were considered as Turonian-Coniacian in

162 age. The Naranjo Unit is formed by fine, golden sandstones, limolites, and carbonate

163 sandstones covered by the Casanga Unit rocks, composed by conglomerates, coarse turbiditic

164 sandstones, and fewer dark brown limolites. Among conglomerates, lithic fragments of

165 marine basalts are recognizable, as well as silicified volcanoclastic rocks. Jaillard et al. (1996)
166 calculated the Casanga Unit thickness as between 200 and 400 m. Hungerbuhler et al. (2001)

167 considered it as late Campanian to early Maastrichtian in age. New correlations indicate that

168 parts of this unit are the conglomerates and sandstones outcropping in the Yamana and La

169 Cordillera areas, where they are known as Río Playas Formation. The datation of these rocks

170 was sampled by Hungerbuhler et al. (2001) about 800 m NE from the Playas bridge, in rocks

171 of the Bramaderos Unit, indicating an age of 66.9 My, corresponding to the late

172 Maastrichtian. The overlying unit, the grainy rocks of the Río Playas Formation could attain a

173 similar age. Covering this unit in the Cerro de Puerna area and the high part of the La Merced

174 area there are rhyolitic basalts dated as Oligocene in age (Hungerbuhler et al., 2001) close to

175 Barrial Blanco, probably corresponding to the Loma Blanca Unit. They lay over dacitic-

176 rhyolitic tuffs perhaps belonging to the Paleocene Sacapalca Formation.

177 Fossilized bones described here were found in yellowish sandstones of the Río Playas

178 Formation (Hungerbuhler et al., 2001; INIGEMM 2013) in a creek 50 meters under the top of

179 the surrounding hills. Considering the lithology, the position overlying Cretaceous volcanic

180 rocks exposed at Puente de Playas, the possible coeval age with the Bramaderos Unit

181 (INIGEMM, 2013), and the opinion of Jaillard et al. (1999), we consider the Río Playas

182 Formation as Upper Cretaceous in age (contra Hungerbuhler et al., 2001).

183

184 5. Results

185 5.1. Description

186 5.1.1. Sacrum (YM-UTPL_002, Fig. 2)

187 The partial recovered sacrum is composed by two conjoined vertebral centra,

188 represented by a small portion of the penultimate vertebral body and the nearly complete last

189 sacral centrum. The material was naturally prepared. The centra of both vertebrae are solidly

190 co-ossified showing a markedly dorsoventral angulation in the longitudinal axis of the sacrum
191 in lateral view (Fig. 2A) close to 45º (Powell, 2003, p.68). This condition is present in the

192 holotype of Neuquensaurus australis (Lydekker, 1893; Bonaparte and Gasparini, 1980;

193 D´Emic and Wilson, 2011) and presumably represents both the effect of an extra caudal

194 addition to the posterior sacrum in this taxon (Fig. 2E) plus a compensation of a relatively

195 horizontal tail with an obliquely oriented sacrum and illia as end of an arched back (Powell,

196 2003).

197 The posterior or better preserved centrum of Yamanasaurus is short and tall, biconvex,

198 with rounded to hexagonal articular facets (Fig. 2C). It shows a marked constriction at mid-

199 length, depicting a narrowed shape with expanded articular facets in ventral view, in a similar

200 way to the saltasaurine Neuquensaurus australis (Apesteguía and Salgado, 2003; Salgado et

201 al., 2005; D´Emic and Wilson, 2011).

202 The ventral narrow condition seen in the material here described (Fig. 2H) clearly

203 differs from those more stout last sacral vertebrae present in other titanosaurs such as

204 Trigonosaurus pricei (Campos et al., 2005), Isisaurus colberti (Jain and Bandyopadhyay,

205 1997), the innominate MCT 1536-R (Campos and Kellner, 1999), and the giants

206 Futalognkosaurus dukei (Calvo et al., 2007, MUCPv-323) and Argentinosaurus huinculensis

207 (Bonaparte, 1999; Bonaparte y Coria, 1993).

208 The presence of a narrow last sacral vertebra is only described for the aeolosaurini

209 Overosaurus (Coria et al., 2013), but the preserved sacral ribs shows a difficult correlation

210 count with other titanosaurs. In most titanosaurs, there are two large and squared proximal

211 vertebrae followed by a third reductionary 3rd centrum which begins the stenosacrum, the

212 narrow part of the sacrum, commonly from 3rd to 6th centra, being the latter expansive. It is

213 possible that the last dorsal of Overosaurus could correspond to the first sacral of many other

214 titanosaurs. Furthermore, in Overosaurus the last sacral elements are aligned with the rest of

215 the sacrum, lacking the change of angulation described for YM-UTPL-002 in lateral view.
216 The left lateral side preserves a small ovoid, shallow blind fossa. Although no internal

217 cavities (i.e. camellate bone) are present in most of the centrum, small irregular pneumatic

218 cavities are recognized in the dorsal region and in the base of the sacral ribs.

219 Both sacral ribs were preserved in the same vertebra. Since in neosauropods the 4th,

220 5th, and 6th ribs are tightly articulated to the acetabulum, and fuse in a sacricostal yolk

221 (Wilson and Sereno, 1998, Wilson, 2002), this is expected in the specimen from Ecuador.

222 Differing from the lateroposteriorly directed 4th sacral rib, here is laterally directed, as in the

223 5th and 6th of titanosaurs. Also the 7th rib in Neuquensaurus australis (MCS-5/16) is laterally

224 directed, but distally expands to articulate to the postacetabular region of the ilia.

225

226 5.1.2. Anterior to mid-caudal vertebra (YM-014, Fig. 3)

227 It consists on a fragment comprising approximately the posterior half of a mid-caudal

228 vertebra. Despite cotyle is not preserved, the condyle is globous and massive respect to the

229 substantial dorsoventral reduction in the middle of the centrum. As preserved, the vertebral

230 centrum seems strongly procoelous. The condyle is mostly spherical, dorsoventrally

231 compressed, with the posterior tip elevated respect to the midline. Besides, a ventral concavity

232 or slight notch is present on the articular surface. The dorsoventral compression is not as

233 pronounced as in Saltasaurus and Rocasaurus.

234 Ventrally, the centrum is markedly concave with a pair of processes posteriorly

235 located, corresponding to the chevron facets. As seen in Neuquensaurus, there is no evidence

236 of a longitudinal ventral ridge that characterize the saltasaurines Rocasaurus and Saltasaurus.

237 No part of the neural arch was preserved but the absence of their bases shows that the position

238 of the arch was located in the anterior half of the centrum. There is no mark of the transverse

239 processes either.

240 The natural breakage evidences no conspicuous inner cavities in a “honeycomb”

241 camellate way, but instead spongy (Fig. 3F). The spongy bone present in the vertebral body

242 with irregular internal pneumatic cavities in the dorsolateral region of the vertebral centrum is

243 commonly view in anterior and mid-caudal vertebrae of Neuquensaurus (Cerda et al., 2012;

244 Zurriaguz and Cerda, 2017). This condition strongly contrasts with the observed in the other

245 saltasaurine taxa such as Saltasaurus and Rocasaurus which have typical camellate internal

246 texture in their anterior and middle caudal centra (Cerda et al., 2012; Zurriaguz and Cerda,

247 2017).

248

249 5.1.3. Humerus (YM-016, Fig. 4A-G)

250 The material includes the eroded proximal half of a left humerus. This fragment

251 evidences a stout contexture as seen in the humeri of saltasaurine titanosaurs such as

252 Saltasaurus (Powell, 2003) and Neuquensaurus (Otero, 2010), being mediolaterally expanded

253 in both proximal and distal portions. This morphology depicts a short limb bone, with

254 diaphysis reaching half of the lateromedially proximal width.

255 The preserved portion of the bone has a triangular, concave anterior surface, while the

256 posterior face is convex. Although damaged, the humeral head can be located at the mid-

257 width of the proximal end. Its dorsal edge was not preserved but it was probably quite straight

258 as in Saltasaurus (Powell, 2003), Neuquensaurus (Otero, 2010), and Opisthocoelicaudia

259 (Borsuk-Białynicka, 1977). The external squared corner was not preserved either. The base of

260 a robust deltopectoral crest is present on the lateral edge of the bone. The extense and

261 thickened crest is concordant with that of Saltasaurus and Neuquensaurus, and unlike the

262 narrower ones present in non-saltasaurine titanosaurs, but it differs from the former in

263 reaching a more proximal point.

264 The bone is crescently-shaped in proximal view. The mid-shaft is anteroposteriorly

265 compressed, with an oval perimeter in cross section.

266

267 5.1.4. Radius (YM-015, Fig. 4H-M)

268 A proximal half of a right radius is present among the material. The diaphysis is

269 thicker than what is common in other titanosaurs and is some flattened. The proximal articular

270 surface is flat to slightly concave, with a peculiar heptagonal outline. In posterior view, the

271 proximal epiphysis widens both laterally and medially, leaving a neck in the diaphysis. This

272 peculiar shape of the radius could be considered an autapomorphy. It is no possible to

273 recognize longitudinal ridges over the diaphysis as expected for the posterior (ulnar) surface.

274 The broken diaphysis shows a quite peculiar subhexagonal outline.

275

276 5.1.5. Tibia (YM-017, Fig. 4N-S)

277 An extremely damaged proximal end of a right tibia was preserved too. It is a stout

278 bone, with an anteroposteriorly expanded proximal end. The shaft tapers towards the distal

279 part of the preserved diaphysis, as in other titanosaurs, but no details are visible. Laterally, the

280 proximal end is planar along its articulation with the femur and curves distally along the

281 cnemial crest, which is poorly preserved.

282 The minimum width of the shaft is located at the distal preserved end, approximately

283 its mid-length. There is a slightly concave surface which could be considered as the lateral

284 surface of the tibia behind the cnemial crest, as occurs in Saltasaurus and Gondwanatitan

285 faustoi (Kellner and Azevedo, 1999; fig. 21).

286

287 6. Discussion

288 6.1. Phylogenetic inferences

289 Although the incompleteness of Yamanasaurus holotype prevents against plotting it in

290 a data matrix to get reliable phylogenetic results, the relatively small size of the remains and

291 the evidence of proportionally short limb elements relates the Yamanasaurus with saltasaurine

292 titanosaurs; the smallest short-limbed South American sauropods known. The presence of

293 isolated features like the narrow last sacral vertebrae, the marked dorsoventral angulation

294 beetwen the last two sacral elements, the dorsoventrally compressed condyle in the anterior to

295 mid-caudal vertebra with posterior tip elevated respect to the midline, plus the absence of a

296 longitudinal ventral ridge and spongy inner structure with absence of internal cavities or

297 camellate bone in this element, approaches the position of Yamanasaurus to the well-known

298 Argentinian saltasaurine Neuquensaurus.

299

300 6.2. Temporal adjustments

301 Saltasaurine titanosaurs are a group of small and derived sauropods. They were

302 already discovered and described in Argentina since the XIX Century (e.g., Lydekker, 1893).

303 Saltasaurine were considered as restricted for both Patagonia (Neuquensaurus, Rocasaurus,

304 Salgado and Azpilicueta, 2000) and northwestern Argentina (Saltasaurus, cf. Neuquensaurus,

305 El Brete quarry, Salta; Bonaparte and Powell, 1980; Powell, 1992) to the interval Campanian-

306 Maastrichtian (Heredia and Salgado, 1999; Leanza et al., 2004), for which the taxon seems to

307 be quite reliable for temporal inferences. Up to now, only Neuquensaurus was found to have

308 seven sacral vertebrae, the last of which is biconvex.

309

310 6.3. Size and environment

311 Although the history of sauropods shows several examples marked by growth as a

312 main survival strategy since their very beginnings at Late Triassic times (Apaldetti et al.,

313 2018), the presence of a split in the diversity in colossal and moderate lineages was present in
314 most clades (Carballido et al., 2017). For example, when Brachiosaurus was one of the most

315 impressive macronarian with 38 tons of body mass (Gunga et al., 2008), Europasaurus barely

316 reached 1 ton (Sander et al., 2006). Among titanosaurs, examples are even more striking. The

317 group successfully occupied diverse environments during Cretaceous times reaching the very

318 end of this period, probably owing to their intrinsic diversity and adaptability.

319 From 110 to 90 Mya they were part of the Age of Giants in southern continents (e.g.,

320 colossosaurian titanosaurs (González Riga et al., 2019) like Patagotitan, Argentinosaurus,

321 Mendozasaurus, Futalognkosaurus, in South America), with species reaching 70 tones

322 (Carballido et al., 2017 and cites therein).

323 However, after Santonian times, the largest forms were vastly replaced by more

324 average sized species, sometimes their own sister-groups, like the 6-10 tones rinconsaurian

325 titanosaurs (e.g., Rinconsaurus, Muyelensaurus) and the Saltasaurinae (Salgado, 2000,

326 Apesteguía 2002a, Leanza et al., 2004). Saltasaurine titanosaurs were extreme in this aspect.

327 Neuquensaurus australis, with a body length of about 7–9 m, attained a body mass of 3.5

328 tones (Carballido et al., 2017), and the same is true for both Saltasaurus and Rocasaurus.

329 We have no way to know if saltasaurine titanosaurs were actual island dwarfs

330 (Apesteguía, 2002a,b) or just their small size, uneven pneumaticity (Cerda et al., 2012), and

331 wider body shape (Powell, 2003) conferred them advantages to occupy the new and widely

332 distributed coastal and insular environments (Apesteguía, 2002b). They were result of the

333 Campanian-Maastrichtian marine transgressive events along most South America. Whereas in

334 northern Patagonia the main event is the Kawas Sea (associated to the saltasaurines

335 Neuquensaurus and Rocasaurus) (e.g., Leanza et al., 2004), NW Argentina and Southern

336 Bolivia were flooded by the Pacha Sea (associated to the saltasaurine Saltasaurus) (Salfity

337 and Marquillas, 1994, Apesteguía and Ares, 2010). Northern Perú and Ecuador were flooded

338 during the early Campanian, mid Campanian–early late Campanian, early Maastrichtian, and
339 terminal early Maastrichtian (Jaillard et al., 2005), where is associated now the saltasaurine

340 Yamanasaurus. Dwarfism was already proposed for titanosaurs from island environments of

341 Europe (e.g., Ampelosaurus, Lirainosaurus, Magyarosaurus) by the time when titanic

342 sauropods roamed continental environments in other places of the world (Nopcsa, 1915,

343 Weishampel et al., 1991, Jianu and Weishampel, 1999, Stein et al., 2010, Company, 2010,

344 Klein et al., 2012).

345

346 7. Conclusions

347 Yamanasaurus, a small to medium-sized titanosaur sauropod, represents the first

348 dinosaur known for Ecuador and the northernmost saltasaurine titanosaur recorded. The

349 presence of saltasaurine sauropods in this country does not result unexpected taking into

350 account the proposed coastal environment adaptations of these small sauropods (Apesteguía,

351 2002b), in the context of the important sea transgressions that occupied the northwest of

352 South America during Campanian-Maastrichtian times (Jaillard et al., 2005). Additionally,

353 since saltasaurines are only known for the interval Campanian-Maastrichtian, this fits quite

354 well with previous age assumptions for the Río Playas Formation.

355

356 Acknowledgements

357 The authors are grateful to Mr. Víctor Francisco Celi Ríos, collector of the studied

358 materials and to Mr. Marco Antonio Paladines Balcázar, who preserved them and donated to

359 the UTPL for its research. Also to Ms. Iovana Jaramillo from the Instituto Nacional de

360 Patrimonio Cultural (INPC-Loja) for allowing us the access to the material preserved at the

361 INPC. We are also grateful to the UTPL for supporting a visit to the field and to Jean-Noel

362 Martínez from the University of Piura, Perú for establishing the work contacts. To Hospital

363 IESS Phillip Brillance for Electronic microscope and CT scan images of the bones. Jorge
364 Antonio González performed the illustrations of the skeleton and bones. Two anonymous

365 reviewers contributed to present a better result from our work.

366

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531
532 FIGURE CAPTIONS

533

534 Fig. 1. Location of the Yamana fossiliferous site in the continental and country context, as

535 well as its location respect to the Lithotectonic Dominion of the Alamor-Lancones

536 Basin in southern Ecuador (A) (source: INIGEMM, 2013); stratigraphic column of the

537 Río Playas Formation with the fossiliferous level (B); different photographs of the

538 Cretaceous outcrops with authors exploring (C-E); fragment of bone outcropping in

539 the quarry (F); scheme of a Neuquensaurus australis skeleton in order to show the

540 relative size of the specimen and the bones found (in red) (G).

541

542

543 Fig. 2. Last two sacral centra in photograph (up) and line drawing (down) of the left (A), right

544 (B), caudal (C), ventral (H), dorsal (I), and cranial (J) views, in life position. SEM

545 image of the ventral view (D). CT scan to show the biconvex nature of the last sacral

546 centrum (G). Penultimate sacral and last biconvex sacral centrum in the holotype of

547 Neuquensaurus australis (MLP Ly 1 and 7) in right lateral (E articulated, reversed, F

548 isolated), ventral (K), and dorsal (L) views. Left half of the complete sacrum of

549 Neuquensaurus australis MCS 5/16 in ventral view (M). Scale bar: 10 cm (A-C, E-F,

550 H-J, K, L); 50 cm (M). E, F, K, L modified from D’Emic and Wilson (2011). M

551 modified from Salgado et al. (2005).

552

553

554 Fig. 3. Caudal half of a mid-caudal vertebra in left (A), right (B), ventral (C), dorsal (D),

555 cranial (E, broken with inner structure detail, F), and caudal (G) views. SEM image of

556 the dorsal view (H). CT scan at mid of the dorsoventral (I) and craniocaudal pathways
557 (J, K) to show the inner structure of the centrum. Scale bar: 10 cm (A-H).

558 Abbreviations: hf, haemal facet; nc, neural canal.

559

560

561 Fig. 4. Preserved appendicular elements. Left humerus in anterior (A), posterior (B), proximal

562 (C), distal (E, broken), lateral (F), and medial (G) views. Left radius in posterior (H),

563 anterior (I), medial (J), lateral (K), proximal (L), and distal (M) views. Right tibia in

564 medial (N), anterior (O), anteromedial (P), posterior (Q), proximal (R), and distal (S,

565 broken) views. Scale bar: 10 cm. A, B, I, P, line drawings to help visual identification.

566 D, line drawing of Neuquensaurus humerus modified from Otero (2010).

Authors’ Statement

All authors have contributed to the

Conceptualization, Methodology, Investigation,
Visualization, and Writing of the manuscript,
and approved the final version.
We have no conlict of interests regarding the material here presented

Sebastian Apesteguia