Accepted Manuscript

A new and complete peirosaurid (Crocodyliformes, Notosuchia) from Sierra Barrosa

(Santonian, Upper Cretaceous) of the Neuquén Basin, Argentina

Rodolfo A. Coria, Francisco Ortega, Andrea B. Arcucci, Philip J. Currie

PII: S0195-6671(18)30083-1
DOI: https://doi.org/10.1016/j.cretres.2018.11.008
Reference: YCRES 4011

To appear in: Cretaceous Research

Received Date: 5 March 2018

Revised Date: 5 October 2018
Accepted Date: 12 November 2018

Please cite this article as: Coria, R.A., Ortega, F., Arcucci, A.B., Currie, P.J., A new and complete
peirosaurid (Crocodyliformes, Notosuchia) from Sierra Barrosa (Santonian, Upper Cretaceous) of the
Neuquén Basin, Argentina, Cretaceous Research, https://doi.org/10.1016/j.cretres.2018.11.008.

This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to
our customers we are providing this early version of the manuscript. The manuscript will undergo
copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please
note that during the production process errors may be discovered which could affect the content, and all
legal disclaimers that apply to the journal pertain.
 ACCEPTED MANUSCRIPT

PT
RI
U SC
AN
M
D
TE
EP
C
AC
 ACCEPTED MANUSCRIPT
1 A new and complete peirosaurid (Crocodyliformes, Notosuchia) from

2 Sierra Barrosa (Santonian, Upper Cretaceous) of the Neuquén Basin,

3 Argentina

PT
6 Rodolfo A. Coria1*, Francisco Ortega2, Andrea B. Arcucci3 and Philip J. Currie4

RI
7
1
8 CONICET – UNRN, Subsecretaría de Cultura de Neuquén - Museo Carmen

SC
9 Funes, Av. Córdoba 55, 8318 Plaza Huincul, Neuquén, Argentina.

10 rcoria@unrn.edu.ar

11 2

U
Grupo de Biología Evolutiva, Facultad de Ciencias, UNED, Paseo de la Senda
AN
12 del Rey 9, 28040. Madrid, Spain. fortega@ccia.uned.es
3
M

13 Universidad Nacional de San Luis, Área de Zoología, IMIBIO-CONICET, Av.

14 Ejército de los Ándes 950, 1st block, 2nd floor, 5700 San Luis, Argentina.
D

15 andrea.arcucci@gmail.com
TE

4
16 University of Alberta, CW405 Biological Sciences Building, Edmonton, Alberta,

17 Canada. pjcurrie@ualberta.ca
EP

18
C

19
AC

20

21 *Corresponding author

22

23

24

25
 ACCEPTED MANUSCRIPT
26 Abstract

27 A new peirosaurid crocodyliform from the Upper Cretaceous (Santonian) of

28 South America, Barrosasuchus neuquenianus gen. et sp. nov. is here

29 described. Barrosasuchus is distinguished by a combination of features that

PT
30 include: presence of a foramen at the mid-point of the dorsal surface of the

31 mandibular symphysis; quadratojugal dorsally broad, extensively contacting the

RI
32 postorbital articulation; absence of ventral exposure of splenials along

33 mandibular rami, posterior to the symphysis; mid to posterior teeth with roots

SC
34 and crowns highly compressed laterally; presence of longitudinal depressions

U
35 on palatal surface of maxillae; and anterior dentary alveoli strongly procumbent.
AN
36 The holotype specimen of Barrosasuchus neuquenianus, MCF-PVPH-413 is

37 represented by an almost complete skull and most of the articulated postcranial

M

38 skeleton. Thus, Barrosasuchus is currently the most complete peirosaurid taxon

39 known from Patagonia, and represents a key element for future reviews of the
D

40 phylogeny of the group. Peirosauridae represents a successful clade in

TE

41 Gondwana, and particularly in Patagonia, with at least six distinct genera

EP

42 currently recorded.

43
C

44 Key words
AC

45 Barrosasuchus; Peirosauridae; Crocodyliformes; Santonian; Upper Cretaceous;

46 Patagonia

47
 ACCEPTED MANUSCRIPT
48 1. Introduction

49 Notosuchian mesoeucrocodylians are relatively well documented in the

50 Cretaceous of Gondwana and comprise many species discovered in South

51 America (Argentina, Brazil, Uruguay), Africa, Madagascar, and the Indian

52 subcontinent (Pakistan) (Woodward, 1896; Rusconi, 1933; Price, 1945, 1950,

PT
53 1955, 1959; Kellner, 1987; Chiappe, 1988; Bonaparte, 1991; Gasparini et al.,

RI
54 1991; Carvalho, 1994; Gomani,1997; Carvalho and Bertini, 1999; Larsson and

55 Gado, 2000; Buckley et al., 2000; Wilson et al., 2001; Pol, 2003; Sereno et al.,

SC
56 2003, Turner and Calvo, 2005, Carvalho et al. 2011; Pol and Powell, 2011,

57 Martinelli et al., 2012). Over the past few decades, discoveries have extended

58
U
the paleobiogeographical distribution of basal mesoeucrocodylians to regions
AN
59 outside Gondwana, including Asia (China, Mongolia; Wu and Sues, 1996; Pol
M

60 and Norell, 2004), and Europe (Antunes, 1975; Ortega et al., 2000). Thus,

61 several systematic studies of crocodyliform taxa have been published (Clark,

D

62 1994; Ortega et al., 2000; Buckley et al., 2000; Sereno et al., 2001; Pol, 2003;
TE

63 Pol and Norell, 2004; Gasparini et al., 2006; Turner and Sertich, 2010; Pol et

64 al., 2014; Sertich and O’Connor, 2014).

EP

65 Peirosauridae, in particular, is considered a controversial group in

regards to both phylogenetic relationships and diversity. The first definition of

C

66

the group included Peirosaurus (Gasparini, 1982) and Lomasuchus (Gasparini

AC

67

68 et al., 1991). Later, it was expanded with the inclusion of an increasing number

69 of taxa with a broader geographical distribution in the Cretaceous of Gondwana

70 (Buckley and Brochu, 1999; Larsson and Gado, 2000; Sereno et al., 2001,

71 Carvalho et al., 2004; Krause et al., 2006; Larson and Sues, 2007; Leardi and

72 Pol, 2009; Sereno and Larson, 2009; O’Connor et al., 2010; Campos et al.,
 ACCEPTED MANUSCRIPT
73 2011; Martinelli et al., 2012; Sertich and O’Connor, 2014; Lio et al., 2016;

74 Barrios et al., 2016). Nonetheless, the phylogenetic relationships of peirosaurids

75 among Mesoeucrocodylia have not, at present, achieved a general consensus

76 and different phylogenetic hypotheses have been proposed to explain the

77 evolution of this clade (Gasparini et al., 1991; Larsson and Gado, 2000; Buckley

PT
78 et al., 2000; Ortega et al., 2000; Carvalho et al., 2004).

RI
79 The first peirosaurid described was Peirosaurus tormini Price, 1955 from

80 the Bauru Basin (Peirópolis, Uberaba County, Brazil). However, the clade was

SC
81 defined in later contributions by Gasparini (1982) and Gasparini et al. (1991).

82 Besides the African taxa Hamadasuchus (Buffetaut, 1994; Larson and Gado,

83
U
2000) and Stolokrosuchus (Larson and Sues, 2007), the highest diversity of
AN
84 peirosaurids is definitively recorded in South America. Among them, four genera
M

85 have been recognized from Brazil -- Montealtosuchus, Peirosaurus,

86 Pepesuchus and Uberabasuchus (Price, 1955; Carvalho et al., 2004, 2007;

D

87 Candeiro y Martinelli 2006; Candeiro et al., 2006; Campos et al., 2011).

TE

88 However, several authors (Larson and Sues, 2007; Martinelli et al., 2012) have

89 suggested Uberabasuchus is a probable junior synonym of Peirosaurus.

EP

90 At present, peirosaurid diversity from the Late Cretaceous of Patagonia

91 includes the Aptian-Albian Barcinosuchus (Leardi and Pol, 2009), the

C

Cenomanian-Turonian Bayomesasuchus (Barrios et al., 2016), the Turonian-

AC

92

93 Coniacian Lomasuchus (Gasparini et al., 1991; Calvo and Porfiri, 2010),

94 Patagosuchus (Lio et al., 2016), the Santonian-Campanian Gasparinisuchus

95 (Gasparini et al., 1991; Martinelli et al., 2012), and Kinesuchus (Filippi et al.,

96 2018).
 ACCEPTED MANUSCRIPT
97 In February 2001, a joint expedition of the Carmen Funes Museum

98 (Plaza Huincul, Neuquén, Argentina) and the Royal Tyrrell Museum of

99 Palaeontology (Alberta, Canada) conducted fieldwork at the locality of Sierra

100 Barrosa, 30 km NE from Plaza Huincul, Neuquén Province (Fig 1.).

101 The joint expedition collected numerous vertebrate fossils -- including the

PT
102 bones of dinosaurs and mammals, and bird footprints -- from different

RI
103 stratigraphic levels of Upper Cretaceous deposits (Coria et al., 2002; Coria and

104 Currie, 2016). Among the discoveries was an almost complete specimen of a

SC
105 peirosaurid crocodilyform from the Bajo de la Carpa Formation (Santonian,

106 Upper Cretaceous, Garrido, 2010). The specimen includes a complete cranium

107
U
articulated with most of its postcranial skeleton (Fig. 2).
AN
108 The specimen MCF-PVPH-413 represents a new peirosaurid taxon
M

109 (Barrosasuchus neuquenianus gen. et sp. nov.). The following description

110 focuses mostly on its cranial anatomy in order to assess its affinities and
D

111 phylogenetic relationships among Mesoeucrocodylia.

TE

112

113 2. Institutional abbreviations

EP

114 MCF-PVPH, Museo Carmen Funes, Paleontología de Vertebrados de Plaza

Huincul (Plaza Huincul, Neuquén, Argentina).

C

115
AC

116

117 3. METHODS

118 The term Notosuchia (Gasparini, 1971) is defined as a stem group

119 composed of all crocodyliforms more closely related to Notosuchus terrestris

120 than to Crocodylus niloticus (Sereno et al., 2001). Carvalho et al. (2004)

121 explicitly proposed the inclusion of Peirosauridae in Notosuchia, a clade that

 ACCEPTED MANUSCRIPT
122 also includes Uruguaysuchidae and Ziphosuchia. The taxon Ziphosuchia was

123 proposed by Ortega et al. (2000) but was redefined by Carvalho et al. (2004),

124 and Turner and Sertich (2010) to encompass the most recent common ancestor

125 of Notosuchus terrestris, Libycosuchus brevirostris, and Baurusuchus pachecoi.

126 The taxonomic status of the new specimen was based on comparative

PT
127 studies by Price (1955), Gasparini (1982), Gasparini et al. (1991), and Carvalho

RI
128 et al. (2004).

129

SC
130 4. SYSTEMATIC PALEONTOLOGY

131 CROCODYLOMORPHA Hay, 1930 (sensu Walker, 1970)

132
U
CROCODYLIFORMES Hay, 1930 (sensu Clark, 1986)
AN
133 MESOEUCROCODYLIA Whetstone and Whybrow, 1983
M

134 NOTOSUCHIA Gasparini, 1971

135 PEIROSAURIDAE Gasparini, 1982

D

136
TE

137 Barrosasuchus neuquenianus

138 Gen. et sp. nov.

EP

139 Figs. 2-10

C

140
AC

141 Etymology: Barrosa, from the Sierra Barrosa, where the specimen was found,

142 and souchos, Greek for the Egyptian crocodile-headed deity; neuquenianus

143 refers to Neuquén Province.

144 Holotype material: MCF- PV 413, an articulated specimen including complete

145 skull with articulated jaws; presacral vertebral series articulated from the neck to

146 the proximal section of the tail; disarticulated hyoids at the posteroventral part of
 ACCEPTED MANUSCRIPT
147 the skull; right coracoid; humeri, ulnae and radii; both articulated hands

148 including carpals, metacarpals and phalanges; left tibia and fibula; articulated

149 pes with four incomplete digits; many relatively small, ornamented, dorsal,

150 ventral and appendicular osteoderms scattered across the ventral part of the

151 body.

PT
152 Locality: Sierra Barrosa , 30 Km NE to Plaza Huincul, Neuquén Province,

RI
153 Argentina (Fig. 1A).

154 Horizon: Bajo de la Carpa Formation, Rio Colorado Subgroup, Neuquén Group

SC
155 (Santonian, Upper Cretaceous) (Garrido, 2010) (Fig. 1B).

156 Diagnosis: Barrosasuchus is diagnosed by the unique combination of the

157
U
following characters (character numbers correspond to data matrix proposed by
AN
158 Pol et al., 2014 and modifications proposed by Barrios et al., 2016): presence of
M

159 a foramen at the mid-point of the dorsal surface of the mandibular symphysis

160 (autopomorphy); dorsal part of the quadratojugal broad, extensively contacting

D

161 the postorbital articulation (character 19, also present in some basal
TE

162 crocodyliforms); absence of ventral exposure of splenials along mandibular rami

163 posterior to the symphysis (character 125, reversal); mid to posterior teeth with
EP

164 roots and crowns highly compressed laterally (character 140, also present in

some sebecids); presence of longitudinal depressions on the palatal

C

165

surface of maxillae (character 253); and anterior dentary alveoli strongly

AC

166

167 procumbent (character 262).

168 Description

169 Skull: The skull of Barrosasuchus is almost complete and preserved in

170 articulation with the lower jaws and postcranial skeleton (Fig. 2). It is 315 mm
 ACCEPTED MANUSCRIPT
171 long, 170 mm wide across the squamosals, and 180 mm long from the anterior

172 limit of the orbits to the tip of the snout (See Table 1).

Skull length (distance between distal

level of the end of the quadrates to 315

PT
rostrum extremity)

RI
Skull posterior width (distance

between the quadrate external 156

SC
borders)

Skull anterior width (distance between

U
the borders of premaxilla at the level 67.5
AN
of the third tooth)
M

Interorbital width (distance between

the two orbits at the contact of anterior 113

D

and posterior palpebrals)

TE

Rostral width in lateral view (distance

between the tip of rostrum to the base 8,2

EP

of the premaxillary first alveolus)

Mandibular length (distance between

C
AC

the extremities of articular and 335

dentary)

Mandibular width (in the middle of the

170
mandibular fenestra)

Orbit anteroposterior length 53.3

Orbit lateromedial width 32

 ACCEPTED MANUSCRIPT
Infratemporal fenestra length 35

Infratemporal fenestra width 23.5

Antorbital fenestra length 13.2

Antorbital fenestra width 7.4

Mandibular fenestra length 22.5

PT
Mandibular fenestra height 7.2

RI
Supratemporal fenestra length 24.5

Supratemporal fenestra width 8.2

SC
External naris length 10.4

External naris width 8.8

U
Suborbital fenestra length 54
AN
Suborbital fenestra width 30
M

Angle between the longitudinal axis of

50°
the skull roof and the quadrate
D

173
TE

174 Table 1. Cranial measurements of Barrosasuchus neuquenianus, Holotype,

175 MCF-PVPH-413. Measurements in mm.

EP

176

Notwithstanding, most of the skull table and the dorsal part of the
C

177

occipital area (including the condylar ends of both quadrates) were eroded by
AC

178

179 weathering before collectin (Fig.3). The dorsal surface of the skull anterior to the

180 orbits was also affected by erosion. It is a slightly smaller specimen than those

181 of Gasparinisuchus and Lomasuchus (Gasparini et al., 1991; Martinelli et al.,

182 2012), which supposedly came from relatively close localities and same

183 stratigraphic levels.

 ACCEPTED MANUSCRIPT
184 In dorsal view (Fig. 3), the skull has a tubular (broad oreinirostral) and

185 moderately wide snout, although not as wide as that of Gasparinisuchus and

186 not as narrow as in either Lomasuchus or Uberabasuchus (Martinelli et al,

187 2012) (Table 1). The anterior surface is heavily ornamented with grooves and

188 pits that are distributed with differential density and arrangements in certain

PT
189 skull areas, including the jugals and the lateral borders of the cranial table. The

RI
190 preserved part of the cranial table, between the orbits and lateral to the right

191 supratemporal fenestra, is continuous with the dorsal border of the snout. In

SC
192 lateral view, the skull profile is flat dorsally and slightly festooned ventrally. The

193 external nares open anteriorly as in other peirosaurids (Price, 1955; Carvalho et

194
U
al., 2004; Candeiro and Martinelli, 2006; Campos et al., 2011). Only the right
AN
195 supratemporal fenestra is preserved, which is small, approximately half the
M

196 anteroposterior diameter of the orbit. The infratemporal fenestrae are slightly

197 smaller than the orbits and each is triangular, with the main axis oriented
D

198 anteroposteriorly.
TE

199 The orbits are subcircular, facing dorsolaterally and are smaller than in

200 other peirosaurids like Montealtosuchus or Uberabasuchus (Carvalho et al.,

EP

201 2004, 2007). Each orbit was covered by a heavily ornamented and triangular

anterior palpebral bone. There is a small and rounded antorbital fenestra, which
C

202

is approximately one third the size of the orbit. It is better preserved on the right
AC

203

204 side of the skull, and is positioned near the anterior border of the orbit.

205 In ventral view (Fig. 4), the palate is complete, and includes a long and

206 smooth secondary palate formed by the premaxilla, maxilla and palatine. The

207 secondary palate has no openings. The suborbital fenestrae are oval in general
 ACCEPTED MANUSCRIPT
208 shape, anteroposteriorly oriented, smaller than the orbits, and are positioned

209 posterior to the secondary palate.

210 The premaxillae are separated dorsally by the paired nasals at the

211 posterior border of the external nares (Fig.3). The premaxilla is

212 anteroposteriorly short and dorsoventrally high. There is a well-developed

PT
213 premaxillary-maxillary notch. The premaxillary-maxillary suture extends dorsally

RI
214 from the notch, in a posteromedially sinusoidal trajectory, dorsally to the contact

215 with the nasals. On the dorsal surface, parallel to the mid-line and posterior to

SC
216 the dorsal border of the external naris, there is a rounded notch between the

217 premaxilla laterally and the nasal medially. It faces dorsally as in other

218
U
peirosaurids. Each premaxillae bear four teeth. The palatal branches are
AN
219 anteroposteriorly short. The incisive foramen is small, circular, and located
M

220 close to the tooth row. Lingual to the tooth row, there is a series of small

221 occlusal pits for reception of the anterior mandibular teeth. The palatal
D

222 premaxillary-maxillary suture is anteromedially directed from the lateral

TE

223 premaxilla-maxilla notch to the rear border of the incisive foramen. The

224 presence of a septum that completely divides the external nares is uncertain.
EP

225 On the anterior contour of the premaxillary symphysis, there is a knob-like

structure as in other peirosaurids. The conjoined external nares face anteriorly,

C

226

have an elliptical outline, and are wider than high. There is an extensive
AC

227

228 perinarial region lateral to the external nares, which completely occupies the

229 anterior surface of the premaxillae. There is a foramen ventrolateral to each

230 side of the perinarial region. This region is not as well developed as in

231 Peirosaurus and Uberabasuchus.

 ACCEPTED MANUSCRIPT
232 The maxilla is mediolaterally wide and almost vertical at the lateral sides

233 of the snout, with a straight ventral edge. Festooning is present although is not

234 as pronounced as in other peirosaurids like Hamadasuchus and

235 Uberabasuchus. Posteriorly, the dorsal surface of the maxilla is not well

236 preserved. Therefore, it is not possible to determine its relationships with the

PT
237 prefrontal and lacrimal. The maxilla constitutes the anterior border of the

RI
238 antorbital fenestra. The maxilla-jugal suture is ventral to the antorbital fenestra.

239 Each maxilla has ten ziphodont teeth. On both sides, the premaxillary-

SC
240 maxillary notch region is slightly damaged, which makes it difficult to determine

241 the exact number of tooth positions. However, it is likely that there was one

242
U
more tooth at the anterior end, and two more tooth positions at the posterior end
AN
243 of the dental row, suggesting there were up to 13 maxillary teeth. The teeth are
M

244 implanted in separate alveoli with the apices oriented vertically, as in

245 Montealtosuchus and Uberabasuchus . In ventral view, the tooth row has a
D

246 gently sinusoidal curvature with maximum development of the lateral

TE

247 convexities level with the 3rd and 10th tooth positions. The palatal branches of

248 the maxillae are sutured along the entire midline of the secondary palate
EP

249 anterior to the contact with the palatines. The vomer is not exposed on the

palatal surface, and there are no foramina of significant size. Some shallow
C

250

depressed areas of the palatal surface medial to the tooth row are recognizable
AC

251

252 as in Lomasuchus. The posterior end of the palatal branch of the maxilla forms

253 the anterolateral border of the suborbital fenestra, and the posterior projection

254 contacts the ectopterygoid at the level of the lingual edge of the last alveolus.

255 Thus, the ectopterygoid is practically excluded from the contact with tooth row.
 ACCEPTED MANUSCRIPT
256 On the sagittal line, the maxillae have posteriorly short and blunt processes that

257 separate the anterior ends of the palatal branches of the palatines.

258 The nasal extends anteriorly between the premaxilla and maxilla and

259 forms the posterior border of the external naris. Due to preservational

260 constraints, it is unclear whether or not the nasal contacted the ascending

PT
261 process of the premaxilla to form an internarial septum. The nasals have a clear

RI
262 sagittal suture, and in dorsal view the edges of the maxillary sutures diverge

263 posteriorly.

SC
264 The external surface of the jugal is oriented dorsolaterally. In lateral view

265 the jugal almost reaches the level of the antorbital fenestra, beyond the most

266
U
anterior border of orbit. The jugal forms the ventral border of the orbit as a thick,
AN
267 lateral rim. The anterior region forming the ventral border of the orbit is slightly
M

268 deeper than the posterior region, but both regions are compressed transversely.

269 No large foramina can be seen near the postorbital bar. The jugal contacts the
D

270 quadratojugal near the posterior corner of the infratemporal fenestra and the
TE

271 suture extends posteroventrally. The medial surface of the jugal is overlapped

272 by an anterior projection of the quadratojugal. The ectopterygoid suture on the

EP

273 medial side of the jugal does not extend posteriorly to the base of the postorbital

bar.
C

274

The postorbital process projects dorsally from the main body of the jugal.
AC

275

276 On its lateral surface, it forms more than half of the postorbital bar, and

277 contributes to the posterior edge of the entire bar. The base of the postorbital

278 bar is flush with the lateral side of the main body of the jugal, and is not inset as

279 in most members of the Neosuchia. The postorbital bar is anteroposteriorly

280 narrow, and flat mediolaterally.

 ACCEPTED MANUSCRIPT
281 The quadratojugal forms the posterodorsal border of the infratemporal

282 fenestra and broadly contacts the postorbital on its dorsal end. The lateral

283 surface is smooth dorsally and bears dense sculpturing on the ventral side. The

284 most ventral portion consists of a buttress that contacts the lateral quadrate

285 condyle as in Hamadasuchus rebouli.

PT
286 The lacrimal forms part of the anterior border of the orbit and the dorsal

RI
287 and posterior borders of the antorbital fenestra as in most peirosaurids. The

288 dorsal surfaces of both lacrimals are heavily weathered and it is not possible to

SC
289 identify sutures with the surrounding bones. Only a marked depression at the

290 anteromedial end of the orbit is easily observed, and corresponds to the facet

291
U
where the anterior palpebral was positioned.
AN
292 The prefrontals are poorly preserved. The dorsal surfaces are
M

293 completely weathered, although the ventral regions provide some information.

294 The prefrontal pillar extends along the internal side of the lacrimal and reaches
D

295 the secondary palate, contacting the dorsal surface of the palatine. The dorsal
TE

296 part is anteroposteriorly compressed, but forms a high wall on the internal side

297 of the preorbital bar.

EP

298 The frontals are badly damaged by weathering, and their limits are

difficult to discern. The interorbital bar is wide and is not upturned at the orbital
C

299

borders. Most of the lateral border of the frontal shows an open suture for the
AC

300

301 supraorbitals. Posterolaterally, the frontal contacts the postorbital.

302 Only a small portion of the right postorbital represents what it is left of

303 the anterolateral corner of the skull table. It has a heavily sculptured dorsal

304 surface, and forms most of the anterior border of the supratemporal fenestra.

305 Along with the squamosal, the skull table diverges strongly posterolaterally. The
 ACCEPTED MANUSCRIPT
306 postorbital bar projects ventrally from the horizontal bar near the lateral border.

307 At the confluence of postorbital and horizontal lamina, there is a marked

308 depression, which could indicate the facet for the posterior palpebral. In this

309 area, the postorbital bar projects anteriorly as a short process. Beneath the

310 horizontal bar of the postorbital, the postorbital bar is a lateromedially

PT
311 compressed sheet that encloses anteriorly the infratemporal fenestra. Also, the

RI
312 postorbital extensively contacts the quadratojugal posteriorly.

313 The squamosal is sutured tightly to the posterior end of the posterior

SC
314 process of the postorbital. Only the anterolateral process of the squamosal is

315 preserved, which forms the posterolateral border of the supratemporal fenestra.

316
U
The squamosal extensively overlies the otic recess in a condition that is similar
AN
317 in all peirosaurids. The dorsal and lateral surfaces of the squamosal are heavily
M

318 ornamented.

319 Anterolaterally, the quadrate is tightly sutured to the quadratojugal. The

D

320 ascending process, the pterygoid process, and the proximal part of the distal
TE

321 quadrate body are preserved. Most of the quadrate condyles are missing from

322 both sides. In the area of the otic recess, the quadrate has a single fenestra
EP

323 anterior to the tympanic cavity, but does not show the arrangement of

324 pneumatic foramina characteristic of other Notosuchia (Baurusuchus,

C

Notosuchus). The tympanic foramen is limited by an anterolateral crest and

AC

325

326 projects laterally between the dorsal part of the quadrate and the ventral edge

327 of the squamosal. It is comparatively much smaller in Barrosasuchus than in

328 Hamadasuchus (Larson and Sues, 2007). The posterior branch of the quadrate

329 is oriented posteroventrally, forming an angle of 30° with the skull table. The

330 preserved part of the paraoccipital process would indicate that the
 ACCEPTED MANUSCRIPT
331 cranioquadrate canal is closed behind the tympanic cavity. The small preserved

332 part of the left quadrate condyle would indicate that the posterior branch of the

333 quadrate, which is posterior to the paraoccipital process, is short. In ventral

334 view, there is a narrow pterygoid branch. Its ventral surface has a deep groove

335 delimited by two sharp ridges. The sutural contact with the pterygoid is unclear.

PT
336 Two anterior palpebral bones were found disarticulated in the ventral

RI
337 region of the skull, near the posterior part of the palate. When cleaned, they fit

338 perfectly in their original positions over the orbits. Each is a thick, triangular

SC
339 bone that is heavily ornamented dorsally. Ventrally, the anterior part has a thick

340 boss that fits into a socket formed by the lacrimal and prefrontal bones, whereas

341
U
the posterior part is smooth and slightly concave. Two much smaller posterior
AN
342 supraorbitals are present anterior to the horizontal bars of the postorbitals
M

343 (Fig.3).

344 The palatines form the posterior end of the secondary palate (Fig.4).
D

345 Each extends a short distance between the posterior part of the maxilla and the
TE

346 choana, which is level with the middle of the suborbital fenestra. Anteriorly, the

347 palatal branches of the palatines form a blunt projection between the maxillae to
EP

348 form a W-shaped suture. The posterior borders of the palatines constitute the

anterior margin of the choana, which are separated by a septum. The palatine
C

349

projects posteriorly along the medial border of the suborbital fenestrae to its
AC

350

351 posterior corner.

352 The pterygoid bounds the posterior extension of the nasopharyngeal

353 canal. It forms a depression posterior to the choana, which projects laterally into

354 a thin pterygoid wing. The well developed pterygoid wing ends posterolaterally

355 in a thickening that constitutes a poorly developed pterygoid flange. The

 ACCEPTED MANUSCRIPT
356 pterygoid wing is shorter anteroposteriorly than in Hamadasuchus (Larson and

357 Sues, 2007). The posterior border of the two pterygoid wings consists of a

358 thickening interrupted at the midline by two poorly developed pterygoid

359 processes. The pterygoid closes the choana posteriorly. The anterior portion of

360 the distal end of the pterygoid wing is hidden from the ventral view by an

PT
361 overlapping process of the ectopterygoid that expands from the pterygoid

RI
362 flange. However, it is not possible to know if that pterygoid projection reaches

363 the posterior end of the pterygoid wing.

SC
364 The ectopterygoid is gracile, and is narrower in Barrosasuchus than it is

365 in Hamadasuchus. The anterior ramus forms the posterolateral border of the

366
U
suborbital fenestra, and extends to the last dental alveolus, although it does not
AN
367 contribute to the alveolar margin. The dorsal ramus is sutured extensively with
M

368 the medial side of the anterior process of the jugal, but does not participate in

369 the postorbital bar. The pterygoid branch of the ectopterygoid covers a short
D

370 portion of the ventral surface of the pterygoid.

TE

371 The occipital view of the skull was badly damaged by weathering, which

372 has affected the occipital views of the parietals, the supraoccipital, the posterior
EP

373 ends of the squamosals, both dorsal regions of the exoccipitals, and the dorsal

part of the basioccipital. The occipital condyle is missing.

C

374

The lower part of the basioccipital is a triangular, posteroventrally

AC

375

376 oriented lamina that is wider than high. It can only be seen in ventral view

377 (Fig.4). The occipital plate is swollen with poorly developed lateral knobs where

378 it contacts the parabasisphenoid and quadrates. Below each of the knobs, and

379 along the quadrate-basioccipital suture, there is a small, slot-like, lateral

380 Eustachian foramen. The basioccipital tubera are poorly developed. Anterior to
 ACCEPTED MANUSCRIPT
381 the basioccipital on the mid-line, there is the opening of a large medial

382 Eustachian foramen, the anterior border of which is formed by the posterior end

383 of the parabasisphenoid.

384 The basisphenoid is exposed ventrally. It is triangular, and wider than

385 long. The basisphenoid is enclosed on either side by the pterygoid branch of the

PT
386 quadrate where it contacts the pterygoid process.

RI
387 Only the lower parts of the otoccipitals are preserved and each one is

388 strongly posteroventrally oriented. The vagus foramen is relatively large,

SC
389 oriented roughly dorsolaterally, and seems to be subdivided into two small

390 foramina, probably for the exits of cranial nerves IX and X. The exit for the

391
U
cranial nerve XII is positioned slightly dorsolaterally to the vagus foramen.
AN
392 Laterally, the exoccipital extends into the paraoccipital process, in which heavy
M

393 weathering has exposed large pneumatic cavities.

394 Mandible: Both mandibular rami are well preserved and complete. The
D

395 mandible in lateral view (Fig. 5) is more slender than in Uberabasuchus,

TE

396 particularly at the posterior corner where the angular projects less

397 dorsoventrally. The mandibular fenestra is smaller than that of Uberabasuchus.

EP

398 It lies in a shallow depressed area framed by the thickened borders of the

surangular and angular.

C

399

The mandibular symphysis is formed by both the dentaries and splenials,

AC

400

401 and extends posteriorly to the level of the 8th alveolus, as in Gasparinisuchus. In

402 contrast, the posterior end of the symphysis of Montealtosuchus reaches the

403 10th tooth position. In dorsal view (Fig. 6A,B), the mandibular rami diverge

404 posteriorly at a 35° angle from the mid-line. They both are rather straight at the

405 articular contact. The symphysial area is relatively short (compared with the
 ACCEPTED MANUSCRIPT
406 total lower jaw length) and wide as in other peirosaurids like Gasparinisuchus

407 (Martinelli et al., 2012) (Fig.6C), but unlike the extensive symphysis of

408 Kinesuchus (Filippi et al., 2018) (Fig.6D). The anterior part of the mandible is

409 dorsoventrally compressed, and in lateral view has a slightly concave dorsal

410 margin as in the peirosaurids Gasparinisuchus and Hamadasuchus. In dorsal

PT
411 view (Fig. 7A-B), the symphysial suture has a conspicuous foramen at the mid-

RI
412 point of the symphysial suture. Although the functionality of this feature remains

413 unclear for the moment, here it is considered an autapomorphic feature of

SC
414 Barrosasuchus neuquenianus.

415 The dentaries are dorsoventrally low anteriorly, and the rami are vertical

416
U
posterior to the symphysis. In dorsal view, the symphysial area has a convex
AN
417 anterior border. The suture with the splenials is concave, not showing the
M

418 posterior intersplenial wedging process present in Gasparinisuchus (Fig.

419 6B,C).The maximum width of the symphysis is at the level of the fourth tooth.
D

420 Posterior to the fourth tooth, there is a strong transverse constriction to the level
TE

421 of the eighth tooth.

422 The symphysis reaches the level of the sixth tooth on the dentary, but
EP

423 continues posteriorly on the splenial. The dental row is sigmoidal in dorsal view.

The teeth are tightly arranged with practically no interalveolar spacing. The first
C

424

three teeth are procumbent and project anterolaterally. The fourth tooth is the
AC

425

426 largest of the tooth row. Posteriorly, the sizes of the teeth decrease up to the

427 eighth tooth. From that position, the dental size increases up to the 13th dental

428 position. There are 16 teeth preserved in the left ramus, whereas there are 17 in

429 the right ramus. However, the total number of teeth was likely 18 in each. The

430 lateral sides of the dentaries are deeply sculptured by grooves and pits, mainly
 ACCEPTED MANUSCRIPT
431 towards the ventral edge. In lateral view, the dorsal border of the dentary is

432 sigmoidal, with two convexities that reach maximum heights at the level of the

433 fourth and 13th teeth. Throughout the entire length, there is a line of nutrient

434 foramina parallel to the dorsal border. Posterior to the symphysis, the lateral

435 sides of the mandibular rami slope dorsomedially. Thus, in dorsal view, the

PT
436 ventral border is lateral to the tooth-bearing edge. On the posterior end, the

RI
437 dentary projects below the surangular and defines the dorsal and anterior

438 borders of the external mandibular fenestra. The dentary-angular suture

SC
439 extends anterodorsally from the anterior end of the external mandibular fenestra

440 to the level of the last tooth position. In ventral view, the dentaries constitute

441
U
more than two thirds of the transverse width of the mandibular rami.
AN
442 The splenial is a thin lamina restricted to the medial side of the
M

443 mandibular ramus. It forms less than one third of the symphysial length. On the

444 medial side posterior to the symphysis, the splenial has a large, slot-like
D

445 intermandibular oralis foramen. The posterior lamina of the splenial, behind the
TE

446 ninth tooth position, constitutes the lingual border of the dentary alveoli. The

447 alveolar border of the splenial is higher than the labial alveolar border. In ventral
EP

448 view, the splenial overlaps the dentary at the level of the 13th tooth position.

More posteriorly, the splenial and dentary are separated by an anterior process
C

449

of the angular. Behind that position, the splenial-angular suture extends

AC

450

451 posteriorly on the medial side. The posterior end of the splenial forms the

452 anterior border of the abductor fossa.

453 The dorsal border of the surangular is bowed, and constitutes the

454 posterior segment of the mandibular ramus. In dorsal view, the surangular

455 exhibits a posterior flat, transversely wide surface. Anteriorly, it has a sharp
 ACCEPTED MANUSCRIPT
456 projection that wedges between the dentary and splenial and reaches the tooth

457 row. In lateral view, the surangular has a flat, sculptured surface above the

458 external mandibular fenestra, and forms the posterodorsal border of that

459 fenestra. In lateral view, the posterior end of the surangular overlaps all the

460 preserved portions of the retroarticular process. The surangular suture with the

PT
461 angular extends from the posterior end of the mandibular fenestra to the

RI
462 posterior end of the preserved portion of the retroarticular process. Posteriorly,

463 the surangular overlaps the lateral side of the articular, partially forming the

SC
464 lateral part of the glenoid cavity.

465 The angular extends on the ventral side of the mandible (Fig. 7) from the

466
U
level of the posterior end of the tooth row to the posterior border of the
AN
467 retroarticular process.
M

468 The lateral side is deeply sculptured and forms the ventral area of the

469 mandible posterior to the external mandibular fenestra. The ventral border of
D

470 the angular thickens into a ridge. In ventral view, the posterior end forms a flat,
TE

471 transversely wide surface, for the insertion of the m. pterygoideus. In medial

472 view, the angular encloses the lower part of the abductor fossa. There is no
EP

473 evidence of an intramandibular fenestra associated with the medial border of

the abductor fossa.

C

474

The articular has a triangular descending process, which is relatively

AC

475

476 short and is sutured to the medial surfaces of the angular and surangular. In

477 dorsal view, the glenoid surface is roughly rectangular, wider than

478 anteroposteriorly long. Posteriorly, there is a sharp transverse ridge. The

479 retroarticular process projects slightly posterodorsally. In dorsal view, the bone

480 has a triangular outline, and the posterior tip points posteroventrally. The medial
 ACCEPTED MANUSCRIPT
481 border has a well-developed, pendant and extensive lamina as in peirosaurids

482 like Montealtosuchus and other notosuchians like Baurusuchus.

483 Dentition: All teeth are ziphodont, have smooth enamel, and bear mesial and

484 distal carinae with discrete denticles. In general, the teeth are labiolingually

485 compressed, and symmetrical in lateral view. All the teeth are triangular and

PT
486 slightly expanded transversely.

RI
487 The premaxillary teeth are slightly more conical than the maxillary and

488 posterior dentary teeth. The first two are small and tightly spaced. The sizes of

SC
489 the premaxillary teeth increase posteriorly, the largest one being the fourth.

490 The anterior maxillary teeth are subconical. The posterior maxillary teeth

491
U
are apically-basally shorter, and their crowns are rather pointed in lateral view
AN
492 (Fig.8), which is similar to other peirosaurids such as Hamadasuchus and
M

493 Lomasuchus.

494 As preserved, the largest tooth in the maxilla is the second from the front,
D

495 although it probably represents the third position of the maxillary series, as is
TE

496 common among peirosaurids. Tooth size decreases posteriorly from the biggest

497 (third) of the series.

EP

498 The dentary teeth are, in general, smaller than maxillary teeth, and the

most posterior ones are blunt. The first dentary tooth of each ramus is well
C

499

developed and directed straight forward. The second and third teeth are smaller
AC

500

501 than the first, and the fourth is the largest one. The 13th mandibular tooth is

502 broader transversally than the rest. More posterior dentary teeth are short and

503 globoid.
 ACCEPTED MANUSCRIPT
504 Ceratobranchialia: Both horns of the ceratobranchialia were preserved near

505 the posterior part of the palate. They are very thin, bowed and dorsoventrally

506 compressed bones (Fig.9).

507 The posterior ends that connected with the basihyoids are thicker,

508 whereas the opposite ones are slender and strongly curved inward. Although

PT
509 the available record of preserved hyobranchial cornua in mesoeucrocodylian is

RI
510 very scarce (ceratobranchialia is known in Simosuchus, Kley et al., 2010),

511 hyoids of Barrosasuchus are similar in both shape and form to the hyoids of

SC
512 extinct and extant crocodiles.

513

514 Postcrania
U
AN
515 The holotype specimen of Barrosasuchus neuquenianus was found lying
M

516 on its ventral side, its skull articulated with most of the postcranial skeleton (Fig.

517 2). It includes most of the presacral vertebral column, forelimbs, one hind limb
D

518 and several semi-articulated osteoderms. However, in ventral view, the ventral
TE

519 armor partially obscures several skeletal elements.

520 The postcranial skeleton was prepared in ventral view, keeping all the
EP

521 elements in position. Although some descriptions for certain elements are

provided, detailed postcranial descriptions will be presented elsewhere.

C

522

Axial skeleton: In the postcranial jacket, the last cervical vertebrae can be
AC

523

524 recognized, which are articulated with their corresponding cervical ribs (Fig.

525 2.3). The current preservation of the specimen prevents a more precise

526 identification in the vertebral column. The centra are amphycoelous,

527 anteroposteriorly shorter than wide. Each of the visible centra has a notch on

528 the ventral border of the anterior articular surface, and a distinctive longitudinal
 ACCEPTED MANUSCRIPT
529 ventral keel. Laterally, the centra expand anteriorly, which is related to the

530 presence of a well-developed lamina that connects the anterior end of the

531 centrum with each parapophysis.

532 The dorsal vertebrae that are not obscured correspond to anterior

533 elements, although the exact sequence remains unclear. The centra are

PT
534 amphycoelous, anteroposteriorly short, and have no ventral keels. The anterior

RI
535 border of the longitudinal midline is slightly swollen to form an incipient

536 hypapophysis. Two posterior dorsal vertebrae are amphycoelous,

SC
537 anteroposteriorly short, and lack both ventral keels and swollen anterior

538 processes. The transverse processes are laminar.

539
U
The shaft of each cervical rib is relatively short, and overlaps distally with
AN
540 the anterior projection of the next vertebra. At least ten dorsal ribs are
M

541 preserved almost in anatomical position on the left side. They are ventrally

542 grooved, with no recognizable uncinate processes.

D

543 Appendicular skeleton: Only a right coracoid is preserved from the pectoral
TE

544 girdle (Fig.2.1). It is exposed in lateral view and almost complete. It is an

545 elongate, plate-like bone. The scapular contact is anteroposteriorly short and
EP

546 straight. Dorsally, there is a round coracoid foramen. The glenoid cavity is

rather wide and is defined by a lip-like process. The coracoid shaft is elongate,
C

547

slightly constricted at mid-shaft, and has expanded proximal and distal ends.
AC

548

549 Both fore-arms are preserved, and the one from the left side is almost

550 complete. The left humerus is partially exposed in anterior view (Fig. 2.2). The

551 proximal end is obscured by some manual elements, and only the medial

552 humeral process is visible. The deltopectoral crest is straight, occupies a third of

553 the total humeral length, and is poorly developed. Distal to the deltopectoral
 ACCEPTED MANUSCRIPT
554 crest, there is a well-developed deltopectoral tubercle. The shaft is slender at its

555 mid-point, and has a square cross-section. Both distal condyles are well

556 developed, but the ulnar condyle expands more anteriorly.

557 The left ulna is almost complete and is exposed in medial view (Fig. 2.2).

558 The proximal end is badly weathered. However, a relatively well-developed

PT
559 olecranon can be observed. The ulnar shaft is slightly bowed and has a

RI
560 transversely compressed cross-section. It is articulated with the left radius,

561 which cannot be described given the current stage of preparation.

SC
562 The right proximal carpals are exposed in anterior view, and in

563 anatomical position (Fig. 2.4). The radiale and ulnare are elongate. In anterior

564
U
view, the radiale is axe-shaped. The ulnare is approximately two thirds the
AN
565 length of the radiale. The preserved portion of the pisciform is rather spherical in
M

566 shape.

567 The proximal ends of the five metacarpals are preserved. Metacarpal I is
D

568 robust, and expands proximally. Digit I has two phalanges. The first phalanx is
TE

569 half the length of the metacarpal. The ungual phalanx is more robust than the

570 penultimate phalanx.

EP

571 Metacarpal II is slightly longer and more slender than Mtc I. Digit II has

three phalanges. The first phalanx is twice the length of the second. The ungual
C

572

phalanx is transversely compressed and is smaller than the first ungual. Only
AC

573

574 the proximal portion of the metacarpal and the most distal phalanges are

575 preserved for the third digit. The third ungual is smaller than that of Digit II. Only

576 the proximal sections of metacarpals IV and V are preserved.

577 The left hindlimb is missing the femur, but the tibia and fibula are

578 articulated with the pes (Fig.2.5). The tibia is exposed in lateral view. The
 ACCEPTED MANUSCRIPT
579 proximal end expands anteroposteriorly, and the shaft is compressed

580 transversely. Only the proximal ends of Metatarsals I to IV are preserved.

581 Osteoderms: The osteoderms preserved are of at least three different

582 types: 1- rectangular, 2- square and 3- smaller, round forms (Fig. 10). Most of

583 them are disarticulated and scattered on the ventral side of the skeleton. A

PT
584 small area in the posterior part of the skeleton has preserved an articulated

RI
585 portion of the ventral armor.

586 The rectangular osteoderms (Fig. 10A) are interpreted as elements of the

SC
587 dorsal armor that were arranged in a pair of parasagittal rows. Some narrower,

588 rectangular osteoderms are associated with the cervical vertebrae, suggesting

589
U
they were part of the nuchal armor. They are dorsoventrally thin, heavily
AN
590 ornamented with pits on the dorsal surfaces, and very smooth on the ventral
M

591 sides. Each of these osteoderms is two times wider than long, and exhibits a

592 dorsal keel that differentiates the element into two parts. The medial one
D

593 occupies two thirds of the element and faces more dorsally, whereas the lateral,
TE

594 smaller part slopes ventrally at an angle of approximately 45°. The entire

595 external surface is sculptured with no indication of any kind of feature that would
EP

596 suggest there were overlapping articulations with other osteoderms.

597 The osteoderms are similar to those referred to Itasuchus, Peirosaurus

C

(Price, 1955) and Uberabasuchus (Marinho et al., 2006). Among the Brazilian
AC

598

599 material, Peirosaurus shows a dorsal keel similar to that in an osteoderm of

600 Barrosasuchus.

601 The square (Fig. 10B) and rounded (Fig.10C) osteoderms contact each

602 other to build a ventral armor composed of multiple rows. The ventral surface of

603 a square osteoderm is flat and evenly ornamented. Small, roundish osteoderms
 ACCEPTED MANUSCRIPT
604 appear occasionally, which could be associated with the lateral shield of the

605 appendicular skeleton.

606

607 5. PHYLOGENETIC DISCUSSION

608 In order to establish the phylogenetic relationships of Barrosasuchus, the

PT
609 new Patagonian form was included in the data set proposed by Pol et al. (2014)

RI
610 with the modifications proposed by Barrios et al. (2016) (Appendix 1), which is

611 based in several previous contributions (Clark, 1994; Turner and Sertich, 2010;

SC
612 Pol et al., 2012) and includes 412 cranial and postcranial characters distributed

613 among 111 crocodyliform taxa. The analysis was conducted by using TNT

614
U
software (Goloboff et al., 2008), 1.5 version (Goloboff and Catalano, 2016) with
AN
615 the application of Traditional Search analysis, replicating the settings proposed
M

616 by Pol et al. (2014, Supp. information).

617 The Traditional Search option provided 1050 most parsimonious trees
D

618 (MPTs) of 1623 steps with a Consistency Index of 0.290 and Retention Index of
TE

619 0.678. The strict consensus showed Barrosasuchus as the sister taxon of

620 Montealtosuchus, Uberabasuchus, Gasparinisuchus and Lomasuchus, within

EP

621 an unsolved politomy with other peirosaurids (Appendix 1.B). The Pruned Trees

622 option shows Lomasuchus and Bayomesasuchus as “wild cards”. When those
C

taxa and that character were inactivated, the new analysis resulted in 1140
AC

623

624 MPTs of 1618 steps, with CI= 0.299 and RI= 0.692 (Appendix 1.C). However,

625 the phyletic position of Barrosasuchus remained unaltered, as the sister taxon

626 of Montealtosuchus and Gasparinisuchus + Uberabasuchus, and the

627 Peirosauridae was recovered in a resolved monophyletic group and sister taxa

628 of the Mahajangasuchidae (Fig. 11).

 ACCEPTED MANUSCRIPT
629 Barrosasuchus is clearly nested among Peirosauridae by the presence of the

630 following synapomorphies: the retention of nearly parallel nasal lateral edges

631 along the suture with the maxilla (character 128, most crocodyliforms have

632 oblique nasal lateral edges, either converging of diverging anteriorly);

633 Postorbital process of jugal posteriorly positioned (character 243, unknown in

PT
634 Gasparinisuchus and Uberabasuchus); sinusoidal ventral edge of maxilla in

RI
635 lateral view (except in Hamadasuchus, which shows the primitive condition);

636 and the presence of a prominent depression on the palate near the alveolar

SC
637 margin at level of the sixth or seventh alveolus (character 396, unknown in

638 Uberabasuchus).

639
U
Barrosasuchus neuquenianus was collected from the Bajo de la Carpa
AN
640 Formation, and the detailed stratigraphical section of the holotype specimen’s
M

641 provenance is provided in the present contribution (Fig. 1). Other peirosaurid

642 taxa, like Gasparinisuchus peirosauroides (Martinelli et al., 2012), Kinesuchus

D

643 overoi (Filippi et al., 2018), and Lomasuchus palpebrosus (Gasparini et al.,
TE

644 1991), are also claimed to have been collected from comparable strata,

645 although no detailed stratigraphic information is provided with the descriptions

EP

646 of these forms. Nonetheless, putting aside the uncertainties of the geographical

and stratigraphical data for those remains, anatomical comparisons between

C

647

Barrosasuchus and the other Bajo de la Carpa taxa show sufficient differences
AC

648

649 to distinguish the new taxon described here. First, the current phylogenetic

650 analysis carried out, nested all these taxa in different phyletic positions (Fig. 10

651 and App. 1 B-C). Also, in regard to direct comparisons, a number of features

652 support the distinction of Barrosasuchus as a new taxon. Kinesuchus overoi

653 (Filippi et al., 2018) shows a long and narrow symphysial mandibular suture,
 ACCEPTED MANUSCRIPT
654 unlike the rather short and broad symphysial suture of Barrosasuchus,

655 Gasparinisuchus, Lomasuchus and most peirosaurids. On the other hand, there

656 are several differences between Barrosasuchus and Lomasuchus (i.e., general

657 geometry of the skull) to distinguish them as two different taxa. Major similarities

658 exist between Barrosasuchus and Gasparinisuchus. However, detailed

PT
659 observation allows detection of several differences that justify their distinction as

RI
660 two different taxa (see Table 2 for a summary of those differences).

661

SC
CHARACTER Barrosasuchus Gasparinisuchus
Compared snout width narrower wider
Dentary-splenial suture in splenials wedge into

U
concave
mandibular symphysis dentaries
similar side than anterior
AN
Posterior premaxillary teeth hipertrophied
ones
Number of premaxillary
4 5
teeth
Unsculpted region in dentary
M

present absent
below tooth row
Dentary with anterior lateral
present absent
concavity
D

Lateral contour of snout in

sinusoidal straight
dorsal view
TE

Wedge-like process of the

maxilla in lateral surface of absent present
premaxilla-maxilla suture
Longitudinal depression in
EP

present absent
palatal surface of maxilla
662
C

663 Table 2. Comparison between Barrosasuchus and Gasparinisuchus.

AC

664

665 Currently, there is a remarkable peirosaurid diversity in the Late Cretaceous

666 of Patagonia. Whether or not such diversity is enhanced by over-assigment of

667 new taxa upon very fragmentary material (i.e. Bayomesasuchus, Patagosuchus,

668 Kinesuchus) is something that exceeds the goals of the present contribution.

669
 ACCEPTED MANUSCRIPT
670 6. CONCLUSIONS

671 Barrosasuchus is a new peirosaurid crocodile taxon from the Upper

672 Cretaceous of South America. The current diversity of the clade Peirosauridae

673 shows its highest peak in Patagonia, with at least six different genera. Although

674 some of these taxa could prove to be invalid in the future due to their

PT
675 incompleteness (i.e. Patagosuchus, Bayomesasuchus and Kinesuchus; Barrios

RI
676 et al., 2016; Lio et al., 2015, Fillippi et al., 2017), it is clear that the

677 Peirosauridae represents a successful clade in Gondwana, and particularly in

SC
678 Patagonia.

679 Barrosasuchus is by now the most complete peirosaurid known from

680
U
Patagonia, and represents a key element for future reviews of the phylogeny of
AN
681 the group.
M

682

683 Acknowledgments
D

684 The skeleton was found and collected by Mike Getty (Royal Tyrrell Museum of
TE

685 Palaeontology) and lab-prepared by M. Alegría (MCF). For access to

686 specimens we thank J. Powell (Colección de Paleontología del Instituto Miguell

EP

687 Lillo, Tucumán, Argentina) and A. Garrido (Museo Olsacher de Zapala,

Neuquén). The authors also thank to the many people involved in 2001
C

688

fieldwork in Sierra Barrosa (L. Filippi, A. Garrido, M. Getty, D. Hernández, E.

AC

689

690 Koppelhus, S. Saldivia and W. Sloboda) that participated in collecting MCF-

691 PVPH-413. Thanks are extended to NSERC grant (Grant #203091–06),

692 Troodon Resources (Calgary, Canada), the Dinosaur Research Institute

693 (Calgary, Canada) for funding support to P.J. Currie, for travel and fieldwork

694 assistance. E. Koppelhus (University of Alberta, Canada) for photography,

 ACCEPTED MANUSCRIPT
695 picture setup, field assistance and logistics. K. Kamra assisted in English

696 editing. We are grateful to D. Pol and another anonymous reviewer for greatly

697 improving the manuscript, and to E. Koutsoukos (CR) for editorial assistance.

698 We also thank the teachers and students of School number 291 of Sierra

699 Barrosa, for their hospitality during our field season, and the Municipalidad de

PT
700 Plaza Huincul and Subsecretaría de Cultura de Neuquén for institutional

RI
701 support.

702

SC
703 References

704 Antunes, M.T., 1975. Iberosuchus, crocodile sebecosuchien nouveau,

705
U
l’Eocene iberique au Nord de la Chaine Centrale, et l’origine du Canyon
AN
706 de Nazare. Comunicações dos Serviços Geológicos de Portugal, 59:
M

707 285-330.

708 Barrios, F., Paulina-Carabajal, A. and Bona, P., 2016. A New Peirosaurid
D

709 (Crocodyliformes, Mesoeucrocodylia) from the Upper Cretaceous of

TE

710 Patagonia, Argentina. Ameghiniana, 53 (1): 14-25.

711 Bonaparte, J.F., 1991. Los vertebrados fósiles de la Formación Rio

EP

712 Colorado, de la ciudad de Neuquén y sus cercanías, Cretácico Superior,

Argentina. Revista Museo Arg. de Cs. Naturales “Bernardino Rivadavia”,

C

713

Paleontología, 4: 17-123.
AC

714

715 Buckley, G.A. and Brochu, C.A., 1999. An enigmatic new crocodile from the

716 Upper Cretaceous of Madagascar. Special Papers in Palaeontology

717 60:149-175.

718 Buckley, G.A., Brochu, C.A., Krause, D.W. and Pol, D., 2000. A pug-nosed
 ACCEPTED MANUSCRIPT
719 crocodyliform from the Late Cretacous of Madagascar. Nature 405

720 (6789): 941-944.

721 Buffetaut, E., 1994. A new crocodilian from the Cretaceous of southern

722 Morocco. Comptes Rendus de l’Académie des Sciences 319(2): 1563-

723 1568.

PT
724 Calvo, J.O. and Porfiri, J.D., 2010. New material of Peirosaurids in Neuquén,

RI
725 Patagonia: its age. Brazilian Geographical Journal: Geosciences and

726 Humanities research medium, 1 (1): 50-64.

SC
727 Campos, D. A., Oliveira, G.R., Figueiredo, R.G., Riff, D., Azevedo, S.A.K.,

728 Carvalho, L.B. and Kellner, A.W.A., 2011. On a new peirosaurid

729
U
crocodyliform from the Upper Cretaceous, Bauru Group, southeastern
AN
730 Brazil. Anais da Academia Brasileira de Ciências, 83 (1): 317-327.
M

731 Candeiro, C.R.A. and Martinelli, A.G., 2006. A review of paleogeographical

732 and chronoestratigraphical distribution of mesoeucrocodylian species

D

733 from the upper Cretaceous beds from the Bauru (Brazil) and Neuquén
TE

734 (Argentina) groups, Southern South America. Jour. South Am. Earth Sci.

735 22(1-2): 116-129.

EP

736 Candeiro, C.R.A., Martinelli, A.G., Avilla, L. and Rich, T., 2006. Tetrapods

from the Upper Cretaceous (Turonian- Maastrichtian) Bauru Group of

C

737

Brazil: a reapraisal. Cretaceous Research 27(6): 923-946.

AC

738

739 Carvalho, I.S., 1994. Candidodon: um crocodilo com heterodontia

740 (Notosuchia, Cretáceo Inferior - Brasil). Anais da Academia Brasileira de

741 Ciências, 66(3): 331-346.

742 Carvalho, I.S. and Bertini, R.J., 1999. Mariliasuchus: um novo

 ACCEPTED MANUSCRIPT
743 Crocodylomorpha (Notosuchia) do Cretáceo da Bacia Bauru, Brasil.

744 Geología Colombiana, 24: 83-105.

745 Carvalho, I.S., Ribeiro, L.C.B. and Avilla, L. de S., 2004. Uberabasuchus

746 terrificus sp.nov. A new Crocodylomorpha from the Bauru Basin (Upper

747 Cretaceous), Brazil. Gondwana Research 7 (4): 975-1002.

PT
748 Carvalho, I.S., Teixeira, V.P.A., Ferraz, M.L.F., Ribeiro, L.C.B., Martinelli,

RI
749 A.G., Neto, F.M., Sertich, J.J.W., Cunha, G.C., Cunha, I.C. and Ferraz,

750 P.F., 2011. Campinasuchus dinizi gen. et sp. nov., a new Late

SC
751 Cretaceous baurusuchid (Crocodyliformes) from the Bauru Basin,

752 Brazil. Zootaxa, 2871: 19-42.

753
U
Carvalho, I.S., Vasconcellos, F.M. and Tavares, S.A.S., 2007.
AN
754 Montealtosuchus arrudacamposi, a new peirosaurid crocodile
M

755 (Mesoeucrocodylia) from the Late Cretaceous Adamantina Formation of

756 Brazil. Zootaxa 1607: 35-46.

D

757 Chiappe, L.M., 1988. A new trematochampsid crocodile from the Early
TE

758 Cretaceous of north-western Patagonia, Argentina and its

759 palaeobiogeographical and phylogenetic implications. Cretaceous

EP

760 Research, 9(4), 379-389.

Clark, J., 1994. Patterns of evolution in Mesozoic Crocodyliformes; Chapter

C

761

5: 84-97 in Fraser, N.C. and Sues, H-D. (eds.), In the Shadows of the
AC

762

763 Dinosaurs, Early Mesozoic Tetrapods, Cambridge Press.

764 Coria, R.A., and Currie, P.J., 2016. A new megaraptoran dinosaur

765 (Dinosauria, Theropoda, Megaraptoridae) from the Late Cretaceous of

766 Patagonia. PloS one, 11 (7), e0157973.

767 Coria, R.A., Currie, P.J., Eberth, D., and Garrido, A., 2002. Bird Footprints
 ACCEPTED MANUSCRIPT
768 from the Anacleto Formation (Lower Campanian, Upper Cretaceous).

769 Ameghiniana 39 (4): 453-463.

770 Filippi, L.S., Barrios, F., and Garrido, A.C., 2018. A new peirosaurid from the

771 Bajo de la Carpa Formation (Upper Cretaceous, Santonian) of Cerro

772 Overo, Neuquén, Argentina. Cretaceous Research 83: 75-83.

PT
773 https://doi.org/10.1016/j.cretres.2017.10.021

RI
774 Garrido, A.C., 2010. Estratigrafía del Grupo Neuquén, Cretácico Superior de

775 la Cuenca Neuquina (Argentina): nueva propuesta de ordenamiento

SC
776 litoestratigráfico. Revista del Museo Argentino de Ciencias Naturales

777 12(2): 121-177.

778
U
Gasparini, Z.B., 1982. Una nueva familia de cocodrilos zifodontes cretácicos
AN
779 de América del Sur. Actas del V Congreso Latinoamericano de Geología,
M

780 Buenos Aires, 317-329.

781 Gasparini, Z., Chiappe, L.M. and Fernández, M., 1991. A new senonian
D

782 peirosaurid (Crocodylomorpha) from Argentina and a synopsis of the

TE

783 South American Cretaceous crocodilians. Journal of Vertebrate

784 Paleontology 11(3): 316-333.

EP

785 Gasparini, Z., Pol, D. and Spalletti, L.A., 2006. An unusual marine

crocodyliform from the Jurassic- Cretaceous boundary of Patagonia.

C

786

Science 311(5757): 70-73.

AC

787

788 Goloboff, P.A., Farris, J.S. and Nixon, K.C., 2008. TNT, a free program for

789 phylogenetic analysis. Cladistics, 24(5): 774–786. doi:10.1111/j.1096-

790 0031.2008.00217.x

791 Goloboff, P.A., and Catalano, S., 2016. TNT, version 1.5, with a full
 ACCEPTED MANUSCRIPT
792 implementation of phylogenetic morphometrics. Cladistics DOI

793 10.1111/cla.12160.

794 Gomani, E.M., 1997. A crocodyliform from the Early Cretaceous dinosaur

795 beds, Northern Malawi. Journal of Vertebrate Paleontology, 17(2): 280-

796 294.

PT
797 Kellner, A.W., 1987. Ocurrencia de um novo Crocodiliano no Cretaceo

RI
798 Inferior da Bacia do Araripe, Nordeste do Brasil. Anais da Academia

799 Brasileira de Ciencias, 59(3): 219-232.

SC
800 Kley, N.J., Sertich, J.J., Turner, A.H., Krause, D.W., O'Connor, P.M., and

801 Georgi, J.A., 2010. Craniofacial morphology of Simosuchus clarki

802
U
(Crocodyliformes: Notosuchia) from the Late Cretaceous of
AN
803 Madagascar. Journal of Vertebrate Paleontology, 30 (sup. to 6):13-98.
M

804 Krause, D.W., O'Connor, P.M., Curry-Rogers, K., Sampson, S.D., Buckley,

805 G.A. and Rogers, R.R., 2006. Late Cretaceous terrestrial vertebrates
D

806 from Madagascar: implications for Latin American biogeography. Annals

TE

807 of the Missouri Botanical Garden 93(2):178-208.

808 Larsson, H.C.E. and Gado, B., 2000. A new Early Cretaceous crocodyliform
EP

809 from Niger. Neues Jharbuch Geol Palont. Abh. Vol 217(1): 131-141.

Larsson, H.C.E. and Sues, H-D., 2007. Cranial osteology and phylogenetic
C

810

relationships of Hamadasuchus rebouli (Crocodyliformes:

AC

811

812 Mesoeucrocodylia) from the Cretaceous of Morocco. Zoological Journal

813 of the Linnean Society, 149)4): 533-567.

814 Leardi, J.M. and Pol, D., 2009. The first crocodyliform from the Chubut

815 Group (Chubut Province, Argentina) and its phylogenetic position within

816 basal Mesoeucrocodylia. Cretaceous Research, 30 (6): 1376-1386.

 ACCEPTED MANUSCRIPT
817 Lio, G., Agnolín, F.L., Valieri, R.J., Filippi, L., and Rosales, D., 2016. A new

818 peirosaurid (Crocodilyformes) from the Late Cretaceous (Turonian–

819 Coniacian) of Patagonia, Argentina. Historical Biology, 28 (6): 835-841.

820 Marinho, T.S., Ribeiro, L.C.B., and Carvalho, I.S, 2006. Morfologia de

821 osteodermos de Crocodilomorfos do sítio paleontológico de Peirópolis

PT
822 (Bacia Bauru, Cretáceo Superior). Anuário do Instituto de

RI
823 Geociências, 29(2): 44-53.

824 Martinelli, A.G., Sertich, J.J., Garrido, A.C. and Praderio, Á.M., 2012. A new

SC
825 peirosaurid from the Upper Cretaceous of Argentina: Implications for

826 specimens referred to Peirosaurus torminni Price (Crocodyliformes:

827
U
Peirosauridae). Cretaceous Research, 37: 191-200.
AN
828 O'Connor, P.M., Sertich, J.W., Stevens, N.J., Roberts, E M., Gottfried, M.D.,
M

829 Hieronymus, T.L., Jinnah, Z.A., Ridgely, R., Ngasala, S.E. and Temba,

830 J., 2010. The evolution of mammal-like crocodyliforms in the Cretaceous

D

831 period of Gondwana. Nature 466: 748-751

TE

832 Ortega, F., Gasparini, Z., Buscalioni, A.D. and Calvo, J.O., 2000. A new

833 species of Araripesuchus (Crocodylomorpha, Mesoeucrocodylia) from

EP

834 the Lower Cretaceous of Patagonia (Argentina). Journal of Vertebrate

Paleontology 20(1): 57-76.

C

835

Pol, D., 2003. New remains of Sphagesaurus huenei (Crocodylomorpha,

AC

836

837 Mesoeucrocodylia) from the Late Cretaceous of Brazil. Journal of

838 Vertebrate Paleontology 23(4): 817-831.

839 Pol, D. and Norell, M.A., 2004. A new crocodyliform from Zos Canyon,

840 Mongolia. American Museum Novitates, 3445: 1-36.

841 Pol, D. and Powell, J.E., 2011. A new sebecid mesoeucrocodylian from the
 ACCEPTED MANUSCRIPT
842 Rio Loro Formation (Palaeocene) of north-western Argentina. Zoological

843 Journal of the Linnean Society, 163 (suppl.1): S7-S36.

844 Pol, D., Leardi, J.M., Lecuona, A. and Krause, M., 2012. Postcranial

845 anatomy of Sebecus icaeorhinus (Crocodyliformes, Sebecidae) from the

846 ocene of Patagonia, Journal of Vertebrate Paleontology 32 (2): 328-354.

PT
847 Pol, D. Nascimento, P.M., Carvalho, A.B., Riccomini, C., Pires-Domingues,

RI
848 R.A., and Zaher, H., 2014. A new Notosuchian from the Late Cretaceous

849 of Brazil and the Phylogeny of Advanced Notosuchians. PLoS ONE 9(4):

SC
850 e93105. doi:10.1371/journal.pone.0093105.

851 Price, L.I., 1945. A new reptile from the Cretaceous of Brazil. Notas

852
U
Preliminares e Estudos, Serviço Geológico Mineralógico do Brasil, 25: 1-
AN
853 8.
M

854 Price, L.I., 1950. On a new crocodilian, Sphagesaurus, from the Cretaceous

855 of the State of São Paulo, Brazil. Anais da Academia Brasileira de

D

856 Ciências, 22(1): 77-83.

TE

857 Price, L.I., 1955. Novos crocodilídeos dos arenitos da Série Baurú, Cretáceo

858 do Estado de Minas Gerais. Anais da Academia Brasileira de Ciências,

EP

859 27(4): 487-498.

Price, L.I., 1959. Sôbre um crocodilídeo notosúquio do Cretácico Brasileiro.

C

860

Boletim do Departamento Nacional da Produção Mineral, Divisão de

AC

861

862 Geologia e Mineralogia, 188: 7-55.

863 Rusconi, C., 1933. Sobre reptiles cretáceos del Uruguay (Uruguaysuchus

864 aznarezi, n.g.n.sp.) y sus relaciones con los notosúquidos de Patagonia.

865 Boletín del Instituto de Geología y Perforaciones, 19: 1-64.

866 Sereno, P.C. and Larsson, H.C.E., 2009. Cretaceous Crocodyliforms from
 ACCEPTED MANUSCRIPT
867 the Sahara. Zookeys 28: 1-143.

868 Sereno, P.C., Larsson, H.C.E., Sidor, C.A. and Gado, B., 2001. The giant

869 crocodyliform Sarcosuchus from the Cretaceous of Africa. Science,

870 294(5546): 1516-1519.

871 Sereno, P.C., Sidor, C.A., Larsson, H.C.E. and Gado, B., 2003. A new

PT
872 notosuchian from the Early Cretaceous of Niger. Journal of Vertebrate

RI
873 Paleontology 23(2): 477-482.

874 Sertich, J., and O’Connor, P.M., 2014. A new crocodyliform from the middle

SC
875 Cretaceous Galula Formation, southwestern Tanzania, Journal of

876 Vertebrate Paleontology, 34:3, 576-596, DOI:

877 10.1080/02724634.2013.819808
U
AN
878 Turner, A.H. and Calvo, J.O., 2005. A new sebecosuchian crocodyliform
M

879 from the Late Cretaceous of Patagonia. Journal of Vertebrate

880 Paleontology, 25(1): 87-98.

D

881 Turner, A.H. and Sertich, J.W. 2010. Phylogenetic history of Simosuchus
TE

882 clarki (Crocodyliformes:Notosuchia) from the Late Cretaceous of

883 Madagascar. Journal of Vertebrate Paleontology 30, Supp.6: 177-236.

EP

884 Wilson, J.A., Malkani, M.S. and Gingerich, P.D., 2001. New crocodyliform

(Reptilia, Mesoeucrocodylia) from the Upper Cretaceous Pab Formation

C

885

of Vitakri, Balochistan (Pakistan). Contributions from the Museum of

AC

886

887 Paleontololy. The University of Michigan 30(12): 321-336.

888 Woodward, A.S., 1896. On two Mesozoic crocodilians Notosuchus (genus

889 novum) and Cynodontosuchus (genus novum) from the Red Sandstones

890 of the Territory of Neuquén (Argentina Republic). Anales del Museo de

891 La Plata, Paleontología, 4:1-20.

 ACCEPTED MANUSCRIPT
892 FIGURE CAPTIONS

893 Figure 1. Location map and geological section.A) Map of the locality with the

894 Barrosasuchus site indicated by the star, B) geological section measured at the

895 site (modified from Coria et al., 2002, 2016).

896 Figure 2. Barrosasuchus neuquenianus, MCF-PVPH-413, Holotype. Top of the

PT
897 figure: Complete skeleton with skull and postcranium articulated as collected:

RI
898 2.1: close-up of left coracoid; 2.2: close-up of right radius and humerus; 2.3:

899 close-up of 2nd,3rd and 4th cervical vertebrae; 2.4: close-up of elongated

SC
900 carpals and manus with 1st , 2nd , 3rd , 4th and 5th metacarpals and ungual

901 phalanges; 2.5: close-up of tibia and left pes with distal tarsals and incomplete

902
U
metatarsals I, II, III, IV and V. Abbreviations: ca, carpus; co, coracoid; cv2, cv3,
AN
903 cv4, 2nd, 3rd and 4th cervical vertebrae; mcI, mcII, mcIII, mcIV, mcV,
M

904 metacarpal I, II, III, IV and V; mtI, mtII, mtIII, mtIV, mtV, metatarsal I, II, III, IV

905 and V; r, radius; ra, radiale; ta, tarsals; ti, tibia; un, ungual phalanx. Scale bars:
D

906 5 cm.
TE

907 Figure 3. Barrosasuchus neuquenianus, MCF-PVPH-413, Holotype. Skull in

908 dorsal view. Abbreviations: j, jugal; m, maxilla; n, nasal; pf, prefrontal; pm,
EP

909 premaxilla; po, postorbital; qj, quadratojugal; soa, anterior palpebral; sop,

posterior palpebral. Scale bar: 10 cm.

C

910

Figure 4. Barrosasuchus neuquenianus, MCF-PVPH-413, Holotype. Skull in

AC

911

912 ventral view. Abbreviations: bo, basioccipital; bs, basisphenoid; ept,

913 ectopterygoid; f, frontal; j, jugal; l, lacrimal; m, maxilla; pl, palatine; pm,

914 premaxilla; pt, pterygoid; qj, quadrato-jugal; sp, squamosal. Scale bar: 10 cm.
 ACCEPTED MANUSCRIPT
915 Figure 5. Barrosasuchus neuquenianus, MCF-PVPH-413, Holotype. Lower jaw

916 in left lateral view. Abbreviations: a, angular; ar, articular; d, dentary; sa,

917 surangular; sp, splenial. Scale bar: 10 cm.

918 Figure 6. Barrosasuchus neuquenianus, MCF-PVPH-413, Holotype and other

919 peirosaurids. Lower jaws in dorsal view. A) photograph of Barrosasuchus

PT
920 neuquenianus, B) interpretative line drawing, C) Gasparinisuchus

RI
921 peirosauroides, D) Kinesuchus overoi. Abbreviations: a, articular; d, dentary; sa,

922 surangular; sp, splenial. Scale bar: 10 cm (C, after Martinelli et al., 2012; D,

SC
923 after Filippi et al., 2018).

924 Figure 7. Barrosasuchus neuquenianus, MCF-PVPH-413, Holotype. Lower jaws

925
U
in ventral view. Abbreviations: a, angular; ar, articular; d, dentary; sp, splenial.
AN
926 Scale bar: 10 cm.

Figure 8. Barrosasuchus neuquenianus, MCF-PVPH-413, Holotype.12th

M

927

928 maxillary tooth in lingual view. Scale bar: 5 mm.

D

929 Figure 9. Barrosasuchus neuquenianus, MCF-PVPH-413, Holotype.

TE

930 Ceratobranchialia . A) right and B) left hyoid cornua in ventral view. Scale bar: 5

931 cm.
EP

932 Figure 10. Barrosasuchus neuquenianus, MCF-PVPH-413, Holotype.

Osteoderms. A) rectangular, B) squared and C) rounded osteoderms in external

C

933

views. Abbreviations: r, rib; ti, tibia. Scale bar: 5 cm.

AC

934

935 Figure 11. Simplified cladogram A showing the phylogenetic position of

936 Barrosasuchus among other peirosaurids after using the character matrix

937 proposed by Pol et al. (2014) (see Appendix 1.C and D for detailed strict and

938 reduced consensus after TNT analysis).

939
 ACCEPTED MANUSCRIPT
940 Appendix 1

941 A. Character scoring of Barrosasuchus in Pol et al. (2014) after the

942 modifications presented by Barrios et al. (2016).

943 201?0001[1]2??001110101??111?0111?????211010001???2011?1010???

944 ????[23]?1?211?010011111??????????00?10?0?101001100?1?100??111

PT
945 001100010010000?0001200101??00????111?1300100??1?0??1?1?1?100

RI
946 ?1000000000?010??01000?1110000010000?0110?10?20?000001001?000

947 ?2000?0???11??00????1?1??1??1???10?0110??1?1?00??00000???????

SC
948 ????00?0000????01?0?11???????????????????????00?0????21?00000

949 ?01?11?1?01??1?10?10000?0??00?0?1?000?00?0????0??

950
U
AN
951
M
D
TE
C EP
AC
 ACCEPTED MANUSCRIPT
952

953 B. Strict consensus cladogram of Barrosasuchus neuquenianus

954 relationships as obtained from TNT after comparing 412 cranial and

955 postcranial characters distributed among 111 crocodile taxa using the

956 character matrix proposed by Pol et al. (2015) after the modifications

PT
957 done by Barrios et al. (2017).

RI
U SC
AN
M
D
TE
C EP
AC

958
 ACCEPTED MANUSCRIPT
959

960 C. Reduced consensus cladogram showing the relationships of Barrosasuchus

961 neuquenianus after excluding six crocodile unstable taxa (see Discussion in

962 text).

PT
RI
U SC
AN
M
D
TE
C EP
AC

963
 ACCEPTED MANUSCRIPT
Barrosasuchus neuquenianus is a new South American Upper Cretaceous
peirosaurid crocodile
Barrosasuchus is the most complete peirosaurid crocodile recorded in
Patagonia at present.
Barrosasuchus is a member of the major Upper Cretaceous diversity of
peirosaurids.

PT
RI
U SC
AN
M
D
TE
C EP
AC