Accepted Manuscript

Inner skull cavities of the basal eusuchian Lohuecosuchus megadontos (Upper

Cretaceous, Spain) and neurosensorial implications

Alejandro Serrano-Martínez, Fabien Knoll, Iván Narváez, Stephan Lautenschlager,

Francisco Ortega

PII: S0195-6671(18)30123-X
DOI: 10.1016/j.cretres.2018.08.016
Reference: YCRES 3946

To appear in: Cretaceous Research

Received Date: 6 April 2018

Revised Date: 22 June 2018
Accepted Date: 25 August 2018

Please cite this article as: Serrano-Martínez, A., Knoll, F., Narváez, Ivá., Lautenschlager, S., Ortega, F.,
Inner skull cavities of the basal eusuchian Lohuecosuchus megadontos (Upper Cretaceous, Spain) and
neurosensorial implications, Cretaceous Research (2018), doi: 10.1016/j.cretres.2018.08.016.

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 ACCEPTED MANUSCRIPT
1 Inner skull cavities of the basal eusuchian Lohuecosuchus megadontos (Upper

2 Cretaceous, Spain) and neurosensorial implications

4 Alejandro Serrano-Martínez1*; Fabien Knoll2; Iván Narváez1; Stephan Lautenschlager3;

5 Francisco Ortega1

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6

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7 : Grupo de Biología Evolutiva. Facultad de Ciencias. Universidad Nacional de

8 Educación a Distancia. Paseo Senda del Rey 9. 28040, Madrid

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2
9 : ARAID-Fundación Conjunto Paleontológico Teruel-Dinópolis. Avda. Sagunto, S/N,

10 44002, Teruel. Spain

11 3

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: School of Geography, Earth and Environmental Sciences. University of Birmingham.
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12 B15 2TT, Birmingham. United Kingdom.
*
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13 Corresponding author

14
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15 1. Abstract
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16 The Late Cretaceous fossil site of Lo Hueco (Cuenca, Spain) has yielded a large

17 collection of tetrapod remains, in which crocodyliforms are one of the best represented
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18 groups. The crocodyliforms from Lo Hueco helped establish Allodaposuchidae as one

of the sister-groups of Crocodylia. A complete skull of the holotype of the

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19
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20 allodaposuchid Lohuecosuchus megadontos was CT scanned, and all its inner cavities,

21 including those of the brain, nerves, inner ear and blood vessels, as well as the

22 paratympanic sinus system and the paranasal sinuses, were three-dimensionally

23 reconstructed and compared to those of several extant and extinct crocodylians. The

24 endocranial anatomy of Lohuecosuchus megadontos is congruent with its phylogenetic

25 position as a basal eusuchian. Our work suggests that some of the neurosensorial
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26 capabilities found in the crown-group Crocodylia such as an acute sense of olfaction

27 and a low frequencies hearing are already present at the base of Eusuchia.

28

29 1.1 Institutional Abbreviations—CRARC: Centre de Recuperació d’Amfibis I Rèptils

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30 de Catalunya (Barcelona, Spain). FMNH: Field Museum of Natural History (Chicago,

31 Illinois). MCD: Museu de la Conca Dellà (Lleida, Spain). MLP: Museo de La Plata

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32 (Buenos Aires, Argentina). MZB: Museu Zoològic de Barcelona (Barcelona, Spain).

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33 OUVC: Ohio University Vertebrate Collections (Athens, Ohio). STUS: Sala de las

34 Tortugas “Emiliano Jiménez” de la Universidad de Salamanca (Salamanca, Spain).

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35 TMM: Texas Memorial Museum (Austin, Texas). UA: Université d’Antananarivo
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36 (Antananarivo, Madagascar). UF: University of Florida (Gainesville, Florida). UMZC:

37 University Museum of Zoology, Cambridge (Cambridge, UK).

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38
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39 1.2 Anatomical Abbreviations—am: auditory meatus. ant: anterior semicircular canal.

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40 bsd: basisphenoid diverticula. car: cerebral carotid artery. cer: cerebral hemispheres.

41 co: cochlear duct. dac: dorsal alveolar canal. ef: Eustachian foramen. en: external naris.
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42 fm: foramen magnum. ic: internal choana. ie: inner ear. itd: intertympanic diverticula.
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43 lat: lateral semicircular canal. mes: mesencephalon. mps: medial pharyngeal sinus. np:
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44 nasal passageway. npd: nasopharyngeal duct. ob: olfactory bulb. ol: optic lobe. ornc:

45 olfactory region of the nasal cavity. ot: olfactory tract. pd: parietal diverticulum. pfo:

46 pituitary (hypophyseal) fossa. pmmf: premaxillary-maxillary foramen. post: posterior

47 semicircular canal. pros: prosencephalon. pss: paranasal sinus system. pts:

48 pharyngotympanic sinus. pvvb: paravestibular vascular bundle. qs: quadrate sinus.

49 rhomb: rhombencephalon. II: optic nerve canal. III: oculomotor nerve canal. IV:
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50 trochlear nerve canal. V: trigeminal nerve canal. IX-XI: shared canal for

51 glossopharyngeal, vagus and accessory nerves. Xtymp: canal for tympanic branch of

52 vagus nerve. XII: hypoglossal nerve paired canals.

53

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54 2. Introduction

55 Extant crocodiles represent only a small fraction of Crocodylomorpha, whose

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56 evolutionary origins date back more than 230 million years. Crocodyliform remains,

57 including cranial elements, are very common in Mesozoic and Cenozoic fossil sites.

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58 They show a great diversity and abundance of this group in many past ecosystems.

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59 Despite the existence of a large number of complete and well-preserved skulls of fossil
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60 crocodiles, only a few have been studied using tomographic techniques. This is

61 particularly unfortunate as these methods allow non-invasive observation of the internal

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62 anatomy of the inner skull, including cavities housing soft-tissue structures .

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63 The first published CT-scan of a crocodyliform skull was that of the goniopholid
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64 Calsoyasuchus valliceps from the Early Jurassic of the USA (Tykoski et al., 2002).

65 Digital explorations of fossil crocodilian cranial anatomy have significantly increased

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66 over the past few years. This holds true for both eusuchians Aegisuchus witmeri

67 (Holliday and Gardner, 2012), Mourasuchus nativus (Bona et al., 2013),

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68 ‘Allodaposuchus hulki’ (Blanco et al., 2015), Gryposuchus neogaeus (Bona et al.,

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69 2017), Diplocynodon tormis (Serrano-Martínez et al., unpublished work) and more

70 stemward forms (Brusatte et al., 2016; Fernández et al., 2011; Herrera et al., 2013; Kley

71 et al., 2010; Pierce et al., 2017). This has gone hand in hand with a similar improvement

72 of the knowledge of the cavities in extant crocodiles (Bona et al., 2017; Brusatte et al.,

73 2016; Dufeau and Witmer, 2015; Gold et al., 2014; Pierce et al., 2017; Witmer et al.,

74 2008; Witmer and Ridgely, 2008). The analysis of neuroanatomical structures in the
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75 Late Cretaceous taxa takes on particular interest for the study of early evolution of

76 Eusuchia. The radiation of eusuchians started sometime near the beginning of the

77 Cretaceous (Brochu, 2013; Buscalioni et al., 2011; Narváez et al., 2016; Turner, 2015).

78 The Late Cretaceous fossil record contains representatives of non-crocodylian Eusuchia

79 which may provide valuable information about the transition and development of brain

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80 structures at this stage. By the early Cenozoic, the three main lineages of Crocodylia

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81 were already geographically widespread.

82 Over the last years, several taxa have been described from the Campanian-Maastrichtian

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83 of Europe (Blanco et al., 2015, 2014; Buscalioni et al., 2001; Delfino and Smith, 2012;

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84 Martin, 2010; Martin and Buffetaut, 2008, 2005; Narváez et al., 2017, 2016, 2015;
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85 Puértolas-Pascual et al., 2014, 2011). Some of these taxa have been included in a

86 recently defined clade named Allodaposuchidae and considered as one of the closest
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87 outgroups to Crocodylia (Narváez et al., 2016). Within Allodaposuchidae, the

88 specimens described from the late Cretaceous site of Lo Hueco (Cuenca, Spain. Fig. 1)
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89 are noteworthy due to their completeness and are thus particularly interesting for the
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90 analysis of the brain and pneumatic structures.

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91 [Figure 1]
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92 Among the crocodiles from Lo Hueco, Lohuecosuchus megadontos was a medium size
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93 allodaposuchid with a broad and short rostrum and extremely hypertrophied teeth,

94 larger than those of any other related taxon (Narváez et al., 2015). The well-preserved

95 holotype specimen of Lohuecosuchus megadontos represents an unparalleled

96 opportunity to provide a detailed description of the complete inner skull cavities in basal

97 Eusuchia, an aspect of their anatomy on which information is scarcely available. These

98 cavities are further compared to those of the other described Crocodyliformes to identify
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99 possible differences and similarities in order to contribute to the development of a

100 neurosensorial evolutionary framework for this group of crocodiles.

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102 [Figure 2]

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103

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104 3. Material and Methods

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105 The specimen studied herein, HUE-04498 (Fig. 2), is a complete and fully articulated

106 skull, part of the holotype of Lohuecosuchus megadontos Narvaez et al. (2015), housed

107
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in the collection of the Museo de Paleontología de Castilla-La Mancha (MuPaCLM,
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108 Cuenca, Spain). The skull is well preserved, although it has suffered plastic deformation

109 in dorsoventral direction, and a shear deformation along its sagittal plane (~10-15º
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110 clockwise, in caudal view). It is also partially covered in a ferrous coating, which cannot
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111 be removed without damaging the fossil. For a detailed osteological description of the
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112 specimen, see Narváez et al. (2015).

113 In order to reconstruct digitally the endocast of the brain and the paratympanic and
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114 paranasal cavities, the specimen was scanned with a Toshiba Asteion Super 4 CT-
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115 scanner and with an Optima CT600, with a voltage of 135kV and a current of 75 mA.
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116 The inter-slice spacing was 0.33mm. The scan data were imported to ImageJ 1.49b

117 (National Institutes of Health, USA) for artefact removal and optimizing the contrast

118 and then into Avizo 7.1.0 (VSG, Burlington, MA, USA) for segmentation, visualization

119 and analysis.

120 The resulting reconstruction was retrodeformed to compensate for the compression

121 suffered by the fossil. With this end in view, the model was scaled in dorsoventral
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122 direction using the "transform editor" in Avizo. Based on specimens of related and

123 morphologically similar taxa, such as Allodaposuchus precedens from Oarda de Jos,

124 Romania (Delfino et al., 2008) and Agaresuchus subjuniperus from Beranuy, Spain

125 (Puértolas-Pascual et al., 2014), a scaling factor of 140% was applied to the Z-axis (Fig.

126 2C-E and 2F-H). Although the original condition is unknown, the applied

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127 retrodefromation method provides a “best-approach” hypothesis, approximating the

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128 alleged original skull morphology (Lautenschlager, 2016; Tschopp et al., 2013; Vidal

129 and Díez Díaz, 2017).

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130 The inner skull cavities of HUE-04498 were compared to those of all

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131 mesoeucrocodylians for which descriptions of the cranial cavities are currently
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132 available. The basalmost specimens are represented by the notosuchian Simosuchus

133 clarki (UA-8679, Kley et al., 2010) and the partially preserved allodaposuchid
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134 'Allodaposuchus hulki' (MCD-5139, Blanco et al., 2015). Within Crocodylia,

135 Alligatoroidea is represented by the basal alligatoroid Diplocynodon tormis (STUS-344,

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136 Serrano-Martínez Unpublished results), the alligatorine Alligator mississippiensis (MZB

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137 92-0231, pers. obs.; OUVC-9761, Witmer and Ridgely, 2008; Dufeau and Witmer,
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138 2015) and the caimanines Caiman crocodilus (caiman_crocodilus_CRARC, pers. obs.;

139 FMNH 73711, Brusatte et al., 2016) and Mourasuchus nativus (MLP73-IV-15-9, Bona
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140 et al., 2013). The crocodyloid sample comprises Crocodylus johnstoni (OUVC 10425,
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141 Witmer et al., 2008), Crocodylus niloticus (MZB 2003-1423, pers. obs.), Osteolaemus

142 tetraspis (MZB 2006-0039, pers. obs.) and Tomistoma schlegelii (TMM M-6342,

143 www.digimorph.org, pers. obs.). Digital reconstruction of the gavialoids Gavialis

144 gangeticus (UF118998, www.morphosource.org, pers. obs.; MLP 602, Bona et al.,

145 2017; UMZC R 5792, Pierce et al., 2017) and Gryposuchus neogaeus (MLP 68-IX-V-1,

146 Bona et al., 2017) complete the comparative sample. This taxon sample was divided
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147 into two size-based groups corresponding to the “yearling to subadult” and “adult”

148 categories based on the ontogeny of A. mississippiensis (Dufeau and Witmer, 2015).

149 Medium-sized specimens of S. clarki, C. crocodilus, O. tetraspis and D. tormis have a

150 skull length (from the posterior edge of the supraoccipital to the anteriormost tip of the

151 premaxilla) between 100 and 300 mm. The large-sized sample includes A.

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152 mississippiensis, M. nativus, C. niloticus, C. johnstoni, T. schlegelii, G. neogaeus, G.

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153 gangeticus, 'A. hulki' and L. megadontos, with a skull length over than 300 mm.

154 For the neurosensorial analyses, only the generated reconstructions of D. tormis, A.

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155 mississippiensis, C. crocodilus, C. niloticus, O. tetraspis, T. schlegelii, G. gangeticus

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156 and HUE-04498 were taken into account. Further reconstructions of members of the
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157 outgroup were also included in the neurosensorial analyses: “Non archosauromorph

158 Sauropsida” is represented by the savannah monitor (Varanidae, Varanus

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159 exanthematicus FMNH 58299, Gauthier et al., 2012), the black-headed python

160 (Serpentes, Aspidites melanocephalus FMNH 97055, Gauthier et al., 2012) and the
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161 northern snapping turtle (Testudines, Elseya dentata TMM M-9315). We decided to add
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162 information from two extant birds as derived archosaurs: the ostrich (Struthio camelus
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163 TMM M-4826) and the black vulture (Coragyps atratus FMM uncatalogued). We

164 reconstructed all these specimens by CT slices obtained from the database of
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165 www.digimorph.org.
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166 We refer to the reconstructed bone-bounded spaces that house soft-tissue structures as if

167 they were the structures themselves (e.g., "facial nerve" instead of "endocast of facial

168 nerve canal").

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170 [Figure 3]
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171

172 4. Results

173 4.1 Neuroanatomy

174 The three-dimensional reconstruction obtained from the CT data provides a rendering of

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175 the inner skull cavities, including the brain cavity endocast, with the bases of the cranial

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176 nerves and the inner ear (Fig. 3), as well as the paratympanic sinus system and paranasal

177 sinuses (Fig. 4). However, the cavity boundaries are difficult to recognise in several

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178 regions, such as the hindbrain and the structures located around it, due to the similar

179 density of the sediment filling the cavities and the mineralised bone.

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180 4.1.1 Brain: The brain is the structure that has been affected the most by the taphonomic

181 dorsoventral compression. Despite the applied retrodeformation correction, the

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182 reconstructed anatomy does not fully represent the original morphology (Fig. 3B and

183 3C). However, the sigmoid morphology of the brain in a lateral view shown by all
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184 mesoeucrocodylians (Bona et al., 2017, 2013; Kley et al., 2010) is still distinguishable.
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185 The inner cavities also suffer from a slight shear deformation along the sagittal plane.
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186 The frontal forms the roof and lateral walls of the long olfactory tract. The cerebral

187 hemispheres are flattened, but can still be delimited rostrally by the position of the optic
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188 nerves and caudally by the pituitary fossa (Fig. 3B). The forebrain is covered dorsally
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189 by the frontal and the parietal, laterally by the laterosphenoids, and ventrally by the

190 basisphenoid.

191 The mesencephalon is an elongated region (Fig. 3B), delimited rostrally by the pituitary

192 fossa, and caudally by the concavities resulting from the otic capsules (Fig. 3C). The

193 hindbrain is limited dorsally by the parietal, dorsodistally by the otoccipital, distally by

194 the prootics and ventrally by the basioccipital. The hindbrain portion has two lateral
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195 concavities that made room for the otic capsules in life. The shear deformation of the

196 region resulted in the left concavity being larger than the right one (Fig. 3D). The

197 caudal region of the hindbrain, rostral to the foramen magnum, has also been

198 compressed, but to a lesser degree than the telencephalon. The pituitary extends from

199 the ventral surface of the brain, in the caudalmost area of the cerebral hemispheres. It is

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200 caudoventrally oriented and appears to be somewhat displaced due to the shear

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201 deformation. The cerebral carotid arteries emerge from the caudalmost part of the

202 pituitary. The left artery could be completely reconstructed, but the right one seemed to

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203 be only partially preserved (Fig. 3B and 3E). The arteries enter the foramen caroticum,

204 on the otoccipital, then course rostrally until reaching the pituitary space, in the

205 basisphenoid.
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206
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207 4.1.2 Cranial nerves: Right and left canals of the optic (II), oculomotor (III), trigeminal
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208 (V), glossopharyngeal (IX), vagus (X), accessory (XI) and hypoglossal (XII) nerves,
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209 and the right canal of the trochlear (IV) nerve are preserved (Fig. 3E). The abducens

210 nerves (VI) are usually very thin in eusuchian crocodiles and could not be recognized
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211 with the resolution of the CT data. The region surrounding the anterior part of the

212 hindbrain is obscured due to the density of the matrix being very similar to that of the
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213 bone, precluding the reconstruction of the facial (VII) and vestibulocochlear (VIII)
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214 nerves.

215 The optic nerves are located ventrally, close to the sagittal plane, in the rostralmost part

216 of the telencephalon. They are short but large nerves that exit the braincase rostrally

217 between the laterosphenoids and the basisphenoid. The oculomotor nerves are located

218 ventral to the flattened hemispheres, close to the rostral edge of the pituitary. They are
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219 caudodistally oriented. Their foramina are located on the laterosphenoids, although they

220 are not visible on the fossil as they are covered by the iron coating. The right trochlear

221 nerve is slender and long. It emerges from the mesencephalon and exits the braincase

222 ventrodistally through the laterosphenoid, rostral to the trigeminal foramen. The

223 trigeminal nerves are the largest, as in other archosaurs. They project laterally, being

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224 wider at their distal end than at their proximal base. Their distal branches cannot be

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225 distinguished. The glossopharyngeal, vagus and accessory nerves share a common

226 caudodistally oriented passage through the otoccipital, dorsolaterally from the foramen

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227 caroticum. The right nerve also preserves the tympanic branch of the vagus nerve

228 (Xtymp), which splits dorsally, close to the IX-XI foramen. The hypoglossal nerves find

229
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their way out the neurocranium via paired canals on each side. The rostral canals were
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230 fully recognized and reconstructed, but the caudal canals were only partially
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231 distinguishable, allowing only the reconstruction of the middle and distal parts. The

232 rostral pair direct laterally over its entire length. The caudal hypoglossal branches
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233 emerge from the hindbrain slightly more dorsally than the rostral ones and course
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234 dorsolaterally before deviating ventrally and exiting the braincase through the

235 otoccipital, close to the foramen magnum.

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236
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237 [Figure 4]
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238

239 4.1.3 Endosseous labyrinth and paratympanic sinus system: The endosseous labyrinths

240 are located in the otic capsules, formed by the prootics and the otoccipitals. Both

241 labyrinths are preserved in the specimen, but their morphology appears to have been

242 altered by the aforementioned shear deformation of the skull. In fact, the angle between
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243 the planes of the rostral and caudal semicircular canals is 120º in the left labyrinth (Fig.

244 3H) but in the right one it is spread almost 180º (Fig. 3I). The semicircular canals of

245 HUE-04498 do not reach the size expected in comparison to other crocodylians.

246 Furthermore, the rostral canal is the smallest (Fig. 3F and 3G), whereas it uses to be the

247 most developed one in archosaurs (Sobral and Müller, 2016). Whereas the left lateral

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248 semicircular canal is circumscribed at the ventral extremity of the rostral and caudal

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249 canals, the right lateral canal lies almost entirely rostral to the crus commune. The left

250 cochlear duct projects ventrally, although its tip is slightly curved rostrally. The right

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251 cochlea only preserves the dorsalmost part, which is ventrally oriented.

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252 The paratympanic and median pharyngeal sinus systems are housed in bony recesses
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253 around the mid/hindbrain (Fig. 4). The effects of the shear deformation are clearly

254 visible, resulting in these cavities having rotated clockwise (Fig. 4E) in caudal view of
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255 the skull. In this region of the braincase, the similar density between the matrix and the

256 bone made the distinction of these cavities difficult.

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257 The intertympanic diverticula are located dorsally to the endocast, housed by the

258 parietal, supraoccipital and both prootics. The lateral regions are wider (more
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259 prominently on the right side) than the central part (Fig. 4C), which appears reduced to

260 a transverse narrow canal in dorsal view. The parietal diverticulum forms a ring-shaped
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261 structure, rostrally oriented, which emerges from both lateral regions of the
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262 intertympanic diverticula and converge rostromedially

263 The pharyngotympanic sinus connects the intertympanic diverticula and the inner ear

264 with the external auditory meatus, which opens distally between the quadrate and the

265 squamosal. The other two passages of the pharyngotympanic sinus, the cranioquadrate

266 canal and the siphonial tube were not recognizable in this specimen, and could not be
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267 reconstructed. Ventrally, the pharyngotympanic sinuses are reduced to two relatively

268 short and narrow canals that course obliquely in caudal view (Fig. 4E). In eusuchians,

269 the pharyngotympanic sinuses connect with the pharynx by two narrow canals, but these

270 could not be distinguished in this specimen.

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271 The median pharyngeal sinus system is located ventral to the pharyngotympanic

272 sinuses, in bony recesses excavated in the basisphenoid.

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273 Both pharyngotympanic sinuses merge in the median pharyngeal sinus. From there, a

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274 long single canal passes caudoventrally to finally open between the basisphenoid and

275 the basioccipital (Fig. 4D). The basisphenoid diverticula only preserve the ventral

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276 connection with the median pharyngeal sinus, close to the merging point of the two
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277 ventral branches of the pharyngotympanic sinuses (Fig. 4D). The basisphenoid

278 diverticula form a single canal, rostrodorsally oriented.

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279
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280 4.1.4 Nasal cavity and associated structures: Archosaurs have a large system of nasal
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281 inner cavities, and those of Lohuecosuchus megadontos extend from the external nares,
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282 at the tip of the snout, to the internal choana, in the caudoventral region of the

283 basicranium. Insufficient resolution of the CT data makes it difficult to distinguish the
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284 limits of these structures. The external nares, formed by the premaxillae, are slightly
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285 wider than long (Fig. 4F). The nasal passageway proceeds from them ventrally, over a

286 short distance before turning at an almost right angle to extend caudally. It is ventrally

287 and laterally delimited by the maxillae, and dorsally by the nasals. Fairly ventrally to

288 the lacrimo-prefrontal area, the nasal passageway divides into two structures. Dorsally,

289 the canal widens, forming the olfactory region of the nasal cavity. Its rendering looks

290 like a large, bilaterally symmetrical bulb, with a sagittal groove on its dorsal surface
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291 (Fig. 4F). This chamber, caudally delimited by the prefrontal pillars and dorsally by the

292 frontals, continues caudally to the olfactory bulbs, which are connected with the brain

293 by long olfactory tract. On the ventral side of the olfactory region of the nasal cavity,

294 the nasal passageway continues caudally in the form of paired nasopharyngeal ducts.

295 These canals pass firstly through the palatine, and then caudally through the pterygoid.

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296 Their dorsal boundaries with the olfactory region are damaged (Fig. 4G), but the

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297 remains of the septum between the nasopharyngeal ducts is still discernible as a sagittal

298 groove on their ventral surface (Fig. 4H). Both ducts course ventrocaudally before

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299 merging in a single canal. Finally, the fused nasopharyngeal ducts open at the

300 caudoventral end of the cranium by the internal choanae, in the pterygoids (Fig. 4G).

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301 The paranasal sinus system is housed in the maxillae on both sides of the nasal

302 passageway (Fig. 4F and 4H). Although the cavities are recognizable, the boundaries
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303 with the nasal passageway were hardly distinguishable. The paranasal sinus system is

304 dorsoventrally compressed and has distally expanded projections. This system is
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305 composed of different cavities such as the antorbital sinus and the postvestibular sinus
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306 (Pierce et al., 2017; Witmer and Ridgely, 2008), but their limits cannot be distinguished
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307 in this specimen.

308 On each side, the dorsal alveolar canal extends lateral to the paranasal sinus systems. It
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309 housed the maxillary branches of the trigeminal nerve and the maxillary veins and
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310 arteries. The neurovascular canals pass rostrodorsally through the maxillae and

311 premaxillae. The dorsal alveolar canal expands in the rostral part of the course forming

312 the paravestibular vascular bundle. From it emerge a canal oriented medially that

313 connects with the nasal passageway, forming the medial subnarial anastomosis (Fig. 4F

314 and 4H). From this bundle also emerges a ventrally oriented canal that opens onto the

315 palate, by the premaxillary-maxillary foramen, located on the palatal premaxillary-

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316 maxillary suture (Fig. 4G). Caudally, the dorsal alveolar canal extends forming a large

317 cavity before opening into the olfactory region of the nasal cavity.

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319 [Table 1]

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320

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321 4.2 Estimation of neurosensorial capabilities

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322 Although we are conscious that assessing neurosensorial capabilities in a fossil taxon

323 such as Lohuecosuchus megadontos is fraught with difficulties, we seized the

324
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opportunity offered by our digital reconstruction of the inner cranial cavities to attempt
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325 to provide a general picture of the sensorial and cognitive acuities of this taxon. Indeed,

326 the endocranial components described above allow calculating linear and volumetric
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327 measurements that have bearing on the sensory abilities and cognitive capabilities of L.
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328 megadontos.
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329 Olfactory acuity depends on the number and size of mitral cells and the number of smell

330 receptors and olfactory receptor genes, and these factors seem to be related to the
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331 absolute and relative size of the olfactory bulb (Zelenitsky et al., 2011, 2009 and
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332 references therein). The olfactory ratio was calculated by comparing the largest
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333 diameter of the olfactory bulb to that of the cerebral hemispheres for each taxon. The

334 results were subsequently log transformed. The obtained results agree with those

335 reported by Zelenitsky et al. (2009): In this study, the olfactory ratio of A.

336 mississippiensis (1.76) was found close the range assigned to this taxon (1,70 - 1,74) by

337 Zelenitsky et al. (2009). The other crocodylians of the sample were also recovered close

338 to the Alligator range proposed by Zelenitsky et al. (2009). The olfactory ratios of the
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339 extant taxa range between 1.70 and 1.82. The olfactory ratio of L. megadontos falls

340 within this range at 1.79 (Table 1).

341 The bony labyrinth houses the neurosensory organs related to the hearing and balance.

342 The cochlear region (Endosseous Cochlear Duct or ECD sensu Witmer et al., 2008), in

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343 the ventral part of the inner ear, houses the hearing organs. The hearing acuity in

344 reptiles and birds seems to be related to the size of the ECD (Walsh et al., 2009). The

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345 ECD of the specimens were measured, scaled to the basicranium length and log

346 transformed (Table 2). The extant crocodyliformes of the study have a scaled ECD that

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347 range between -0.73 and -1.09, close to the values given by Walsh et al. (2009) (range

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348 between -0.62 and -1). L. megadontos has a scaled ECD of -1.13, lower, but similar to
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349 those of A.mississippiensis and C. niloticus.

350 The visual acuity in vertebrates is often related to the size of the eyeball (Hall and Ross,
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351 2007; Schmitz, 2009). The relationship between the size of the eyeball and the ocular
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352 orbit has been studied in lepidosaurs (Hall, 2009) and birds (Hall, 2008), but no such
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353 work is available for crocodiles. However, crocodilian endocasts have an easily

354 distinguishable optic lobe (Fig. 3B; Jirak and Janacek, 2017), which can be used as a
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355 proxy for optic capabilities. The relative volume of the mesencephalon with respect to

356 the whole brain in the extant specimens of our sample ranges between 10 and 15% in
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357 the large-sized specimens, and between 19 and 22% in the medium-sized ones. The
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358 mesencephalon of HUE-04498 has a relative size of 13%, similar to those of other

359 large-sized specimens.

360 The volume of the endocast of the brain can be used to estimate cognitive capabilities

361 using an equation adjusted to accommodate reptilian brain relative proportion, the

362 Reptile Encephalization Quotient (REQ: Hurlburt, 1999). Volume measurements of the
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363 brain endocast were obtained using the measurement tool of AVIZO and converted into

364 the mass assuming a density of 1g/cm3 (Franzosa, 2004). The body mass was estimated

365 based on skull measurements (Dodson, 1975; Platt et al., 2011; Webb and Messel,

366 1978). The endocast volume of L. megadontos is 13.6 cm3 and its body mass was

367 estimated to be around 89 kg (Webb and Messel, 1978). The actual brain volume of L.

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368 megadontos was calculated using Jirak and Janacek's (2017) data on extant crocodiles,

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369 using a regression formula that compares the endocast volume with the relative volume

370 of the brain (Suppl). As a result of this adjustment, the REQs were reduced to 0.9-1.2 in

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371 the large-sized specimens and about 1.7 in the medium-sized ones. The REQ of L.

372 megadontos is 0.82, a bit low compared with those of the large-sized specimens. This is

373
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probably due the crushing suffered by the brain cavity.
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374
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375 [Table2]
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376
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377 5. Discussion
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378 5.1 Morphological implications

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379 Braincases are usually considered conservative structures (Holloway, 2011; Jirak and
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380 Janacek, 2017; Paulina-Carabajal and Filippi, 2017; Witmer and Ridgely, 2009), and the

381 differences found in these structures are potentially useful in phylogenetic analyses

382 (Knoll et al., 2012; Paulina-Carabajal et al., 2017). The compression and shear forces

383 suffered by the specimen have substantially modified the shape of the inner skull

384 cavities of Lohuecosuchus megadontos. However, the different regions that compose

385 the brain and pneumatic cavities can still be distinguished. The reconstructed
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386 endocranial components of L. megadontos were compared to those of all

387 mesoeucrocodylians previously described.

388 Although the morphology of the inner cavities is similar to the rest of the sample overall

389 (Fig. 5), L. megadontos shows particular character states that are consistent with its

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390 phylogenetic position as a basal eusuchian (Narváez et al., 2015), shows distinct

391 character states and polarization proposed in Serrano-Martínez (Unpublished results).

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392 The endocranial cavities of HUE-04498 show all character states that can be considered

393 eusuchian plesiomorphies, such as a flat caudodorsal outline of the brain hemispheres in

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394 lateral view, an intertympanic diverticula with distal expansions more developed than

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395 the central part , and a median pharyngeal sinus that is long and circular in cross-section
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396 along most of its length.

397 In all studied crocodylians, the nasopharyngeal ducts turn at a right angle to open
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398 ventrally through the internal choana. In HUE-04498, however, the caudalmost part of
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399 the nasopharyngeal ducts does not show the straight angle turn. Instead, these ducts
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400 show a gradual caudoventral curvature (Fig. 5), due to the ventral displacement of the

401 internal choana (process explained by Tarsitano (1985)).

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402 Regarding the dorsal alveolar canal, a notable feature is observed. In some alligatoroids,
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403 such as C. crocodilus, the caudal end of the canal opens into the olfactory region of the
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404 nasal cavity chamber, keeping its tubular structure all along its length. However, in

405 crocodyloids, this neurovascular canal extends distally and then expands into a large

406 cavity just rostral to the point where the septum between it and the nasal cavity chamber

407 disappears (Fig. 5). In G. gangeticus, the dorsal alveolar canal opens into the olfactory

408 region cavity, as in C. crocodilus. HUE-04498 shows a configuration recalling that seen

409 in crocodyloids: the dorsal alveolar canal widens and opens separately from the
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410 olfactory region of the nasal cavity. The primitive condition may be the presence of a

411 large cavity in the caudalmost region of the dorsal alveolar canal. Crocodyloidea retain

412 this character state, as do some alligatoroids such as A. mississippiensis, while in

413 Gavialoidea and C. crocodilus this enlargement of the dorsal alveolar canal has been

414 lost. Brochu (1999) also noticed this character, suggesting it to be a Crocodyloidea

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415 apomorphy.

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416 On the other hand, results are not conclusive on some controversial issues within

417 crown-group Crocodylia. As aforementioned, T. schlegelii has a caudal end of the

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418 dorsal alveolar canal similar to those of crocodyloids, but a short median pharyngeal

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419 sinus, as in G. gangeticus and the alligatoroids. The first character is in agreement with
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420 the morphological phylogenetical hypotheses that include T. schlegelii within

421 Brevirostres (Fig. 5A. Brochu, 2011, 1999; Piras et al., 2010), whereas the second one it
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422 is in line with molecular phylogenetical hypotheses (Fig. 5B) in which T. schlegelii and

423 G. gangeticus are closely related (Densmore and Owen, 1989; Meganathan et al., 2010).
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424 [Figure 5]

425
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426 5.2 Neurosensorial implications

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427 Crocodylians are characterized by big olfactory bulbs and a sharp sense of smell
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428 (Zelenitsky et al., 2011, 2009 and references therein). Our results suggest that olfactory

429 acuity of crocodiles is beyond that of birds and most dinosaurs (Table 1; Fig. 2 from

430 Zelenitsky et al., 2009) (as well as beyond the outgroup taxa Elseya dentata and

431 Varanus exanthematicus), but at the same level as the snake Aspidites melanocephalus.

432 When crocodyloids and alligatoroids are considered separately, it is notable that the

433 olfactory ratio remains relatively constant, regardless of skull-size, with crocodyloids
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434 having a slightly higher olfactory ratio than alligatoroids. With an olfactory ratio of

435 1.79, HUE-04498 falls between crocodyloids (average olfactory ratio = 1.82) and

436 alligatoroids (average olfactory ratio = 1.76).

437 With regards to hearing, crocodiles seem to have an acoustic acuity specialized towards

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438 low frequencies (Brusatte et al., 2016; Pierce et al., 2017; Vergne et al., 2009; Walsh et

439 al., 2009). The maximum scaled cochlear length of the studied sample is similar to that

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440 found by Walsh et al. (2009) for these taxa (-1 to -0.62). Furthermore, the minimum

441 scaled cochlear length established for the biggest samples (-1.09 to -0.95; Table 2), are

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442 close to the average position for their hearing capability according to the regression line

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443 proposed by Walsh et al. (2009): around 700Hz (Vergne et al., 2009; Walsh et al.,
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444 2009). HUE-04498 has a small scaled EDC (-1.13), even smaller, but still similar to A.

445 mississippiensis and C. niloticus. The hearing range of L. megadontos would therefore
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446 be lower than those of its extant relatives. Vergne et al. (2009) noticed that larger

447 specimens are found to show a trend towards lower frequencies hearing. Since L.
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448 megadontos has a remarkable short snout (Narváez et al., 2015), the measurements
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449 based on the rostrocaudal axis may be underestimated. However, L. megadontos has the
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450 largest skull width between the samples here studied, so it may be the largest specimen

451 of this study. Therefore should not be surprising it has the lower hearing frequency. On
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452 the other hand, short cochlear size in HUE-04498 may also be explained by the
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453 compression suffered by the specimen.

454 The deformation of most of the brain cavity and the inner ear of HUE-04498 makes the

455 results of some analyses, such as the encephalization quotient, the visual acuity or the

456 balance capability less conclusive, and less distorted specimens are necessary to clarify

457 which of this anomalous morphology is due to distortion.

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458

459 6. Conclusion

460 The digital reconstruction of the holotype skull of Lohuecosuchus megadontos reveals

461 an endocranial anatomy as would be expected for a taxon of its size and phylogenetical

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462 position as a basal eusuchian. A flat caudodorsal outline of the cerebral hemispheres in

463 lateral view, large distal expansions of the intertympanic diverticula compared with the

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464 central region, and a long and circular cross-sectional median pharyngeal sinus are the

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465 most notable plesiomorphies of the endocranial cavities of L. megadontos. A broad

466 caudal end of the dorsal alveolar canal is shared with crocodyloids, whereas the gradual

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467 ventral turn of the nasopharyngeal ducts becomes a right angle in more derived
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468 eusuchians.
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469 The neurosensorial analyses are consistent with eusuchians having an acute olfactory

470 capability and are adapted to low frequency hearing. The infered olfactory acuity of L.
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471 megadontos falls between that of the two major crocodylian groups: Alligatoroidea and
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472 Crocodyloidea. The inner ear of L. megadontos seems to be adapted to even lower

473 frequences than extant alligatoroids and crocodyloids. The sensorial capabilities
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474 deduced from the morphology of the semicircular canals and the general shape of the
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475 brain must be considered with caution, as these structures might have been considerably
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476 deformed by diagenetic processes. Further investigations based on undeformed basal

477 eusuchian specimens will allow to confirm the results of this study and to provide

478 further understanding of the neurological and sensorineural evolution of the early

479 evolution of modern crocodiles.

480

481 7. Acknowledgements
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482 We are grateful to the staff of the Centro Médico de Diagnóstico (Talavera de la Reina)

483 and the Hospital Quirón Ruber (Madrid) for CT scanning many of the specimens

484 analysed in this work. This article greatly benefited from comments by D. Vidal that

485 undoubtedly improve this article. The authors want to acknowledge the valuable

486 suggestions of two anonymous referees.

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487 This is a contribution to a research project from the Ministerio de Economía, Industria y

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488 Competitividad, Spain [CGL2015-68363-P] and from the Junta de Comunidades de

489 Castilla-La Mancha, Spain [DGCLM/140565]. ASM is an FPU grantee [FPU13/03362]

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490 of the Ministerio de Educación, Cultura y Deporte (Spain). FK is an ARAID Senior

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491 Researcher. AN
492
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493 References

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650 Jahrb. für Geol. und Paläontologie Abhandlungen 170, 27–44.

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651 Tschopp, E., Russo, J., Dzemski, G., 2013. Retrodeformation as a test for the validity of

652 phylogenetic characters: an example from diplodocid sauropod vertebrae.

653 Paleontol. Electron. 16, 1–23.

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654 Turner, A.H., 2015. A review of Shamosuchus and Paralligator (Crocodyliformes,
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655 Neosuchia) from the Cretaceous of Asia. PLoS One 10, e0118116.

656 https://doi.org/10.1371/journal.pone.0118116
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657 Tykoski, R.S., Rowe, T.B., Ketcham, R.A., Colbert, M.W., 2002. Calsoyasuchus

658 valliceps, a new crocodyliform from the Early Jurassic Kayenta Formation of
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659 Arizona. J. Vertebr. Paleontol. 22, 593–611.

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660 Vergne, A.L., Pritz, M.B., Mathevon, N., 2009. Acoustic communication in
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661 crocodilians: From behaviour to brain. Biol. Rev. 84, 391–411.

662 https://doi.org/10.1111/j.1469-185X.2009.00079.x

663 Vidal, D., Díez Díaz, V., 2017. Reconstructing hypothetical sauropod tails by means of

664 3D digitization: Lirainosaurus astibiae as case study. J. Iber. Geol. 43, 293–305.

665 https://doi.org/10.1007/s41513-017-0022-6
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666 Walsh, S. a, Barrett, P.M., Milner, A.C., Manley, G.A., Witmer, L.M., 2009. Inner ear

667 anatomy is a proxy for deducing auditory capability and behaviour in reptiles and

668 birds. Proc. R. Soc. 276, 1355–1360. https://doi.org/10.1098/rspb.2008.1390

669 Webb, G.J.W., Messel, H., 1978. Morphometric analysis of Crocodylus porosus from

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670 the north coast of Arnhem Land, Northern Australia. Aust. J. Zool. 26, 1–27.

671 https://doi.org/10.1071/ZO9780001

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672 Witmer, L.M., Ridgely, R.C., 2009. New insights into the brain, braincase, and ear

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673 region of tyrannosaurs (Dinosauria, Theropoda), with implications for sensory

674 organization and behavior. Anat. Rec. 292, 1266–1296.

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675 https://doi.org/10.1002/ar.20983
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676 Witmer, L.M., Ridgely, R.C., 2008. The paranasal air sinuses of predatory and armored
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677 dinosaurs (Archosauria: Theropoda and Ankylosauria) and their contribution to

678 cephalic structure. Anat. Rec. 291, 1362–1388. https://doi.org/10.1002/ar.20794

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679 Witmer, L.M., Ridgely, R.C., Dufeau, D.L., Semones, M.C., 2008. Using CT to peer
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680 into the past : 3D visualization of the brain and ear regions of birds, crocodiles, and
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681 nonavian dinosaurs, in: Endo, H., Frey, R. (Eds.), Anatomical Imaging towards a

682 New Morphology. Springer, Tokyo, pp. 67–87. https://doi.org/10.1007/978-4-431-

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683 76933-0
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684 Zelenitsky, D.K., Therrien, F., Kobayashi, Y., 2009. Olfactory acuity in theropods:

685 palaeobiological and evolutionary implications. Proc. R. Soc. B Biol. Sci. 276,

686 667–673. https://doi.org/10.1098/rspb.2008.1075

687 Zelenitsky, D.K., Therrien, F., Ridgely, R.C., McGee, A.R., Witmer, L.M., 2011.

688 Evolution of olfaction in non-avian theropod dinosaurs and birds. Proc. R. Soc. B
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689 Biol. Sci. 278, 3625–3634. https://doi.org/10.1098/rspb.2011.0238

690

691 Figure captions

692 Fig. 1: Geographic location of the Spanish fossil site of Lo Hueco (Cuenca, Spain).

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693 Modified from Narváez et al. (2015). [planned for 2 column size]

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694

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695 Fig. 2: Skull of Lohuecosuchus megadontos (HUE-04498). A: dorsal view of the skull.

696 B: dorsal view of the unmodified digital model. C: left view of the skull. D: left view of

697
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the unmodified digital model. E: left view of the retrodeformed digital model. F: caudal
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698 view of the skull. G: caudal view of the unmodified digital model. H: caudal view of

699 the retrodeformed digital model. I: ventral view of the skull. J: ventral view of the
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700 unmodified digital model. Abbreviations: am: auditory meatus; ef: Eustachian
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701 foramen; en: external naris; fm: foramen magnum; ic: internal choana; pmmf:
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702 premaxillary-maxillary foramen. Scale bar equals 5cm. [planned for 2 column size]

703
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704 Fig. 3: Three-dimensional reconstruction of the endocranial components of

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705 Lohuecosuchus megadontos (HUE-04498). A: in-situ position of the brain endocast

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706 with the skull rendered semi-transparent, in left lateral view. B-E: HUE-04498 brain,

707 nerves, endosseous labyrinth and arteries in B, unmodified, in left lateral view; C, left

708 lateral view after retrodeformation; D, dorsal and E, ventral views. Scale bar equal to

709 1cm. F-I: HUE-04498 endosseous labyrinths in: F, left endosseous labyrinthin left

710 lateral view; G, right endosseous labyrinth in right lateral view; H, left endosseous

711 labyrinth in dorsal view and I, right endosseous labyrinth in dorsal view. Scale bar
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712 equals 1mm. Abbreviations: ant, anterior semicircular canal; car, cerebral carotid

713 artery; cer, cerebral hemisphere; co, cochlear duct; ie, inner ear; lat, lateral semicircular

714 canal; mes, mesencephalon; ob, olfactory bulb; ol, optic lobe; ot, olfactory tract; pfo,

715 pituitary (hypophyseal) fossa; post, posterior semicircular canal; pros, prosencephalon;

716 rhomb, rhombencephalon; II, optic nerve; III, oculomotor nerve; IV, trochlear nerve; V,

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717 trigeminal nerve; IX-XI, glossopharyngeal, vagus and accessory nerves; Xtymp,

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718 tympanic branch of vagus nerve; XII, hypoglossal nerve. [planned for 2 column size]

719

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720 Fig. 4: Three-dimensional reconstruction of the paratympanicsinus system and

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721 paranasal structures within the braincase of Lohuecosuchus megadontos (HUE-04498).
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722 A, B: in-situ position of the paratympanic sinus system and paranasal structures with the

723 skull rendered semi-transparent, in A, dorsal; B, left lateral views. C-E: HUE-04498 the
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724 median pharyngeal and pharyngotympanic sinus systems with the brain rendered
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725 transparent, in C, dorsal; D, left lateral; E, caudal views. F-H: HUE-04498 the
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726 paranasal system with the brain rendered transparent, in F, dorsal; G, left lateral; H,

727 ventral views. Abbreviations: bsd, basisphenoid diverticula; dac, dorsal alveolar canal;
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728 en, external naris; ic, internal choana; itd, intertympanic diverticula; mps, medial

729 pharyngeal sinus; np, nasal passageway; npd, nasopharyngeal duct; ob, olfactory bulb;
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730 ornc, olfactory region of the nasal cavity; pd, parietal diverticulum; pts,
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731 pharyngotympanic sinus; pss, paranasal sinus system; pvvb, paravestibular vascular

732 bundle. Scale bar equals 1cm. [planned for 2 column size]

733

734 Fig. 5: Evolutionary pattern of the eusuchian brain. A, Phylogeny of Crocodylia based

735 on morphological characters, modified from Brochu, (1999, 2011). B, Phylogeny of

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736 Crocodylia based on morphological and molecular characters, modified from Densmore

737 and Owen, (1989) and Meganathan et al., (2010). It includes Lohuecosuchus

738 megadontos (HUE-04498) and the reconstructed specimens discussed in the text. a,

739 Alligatoroidea; b, Crocodyloidea; c, Gavialoidea. The images are posterior views of the

740 brain with cranial nerves, blood vessels and inner ear, and the paratympanic sinus

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741 system (left); and right lateral (middle) and dorsal (right) views of the brain with cranial

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742 nerves, blood vessels and inner ear, the paratympanic sinus system, and the paranasal

743 sinuses. Color key. Endocast colors were based on Witmer et al., (2008): blue, brain;

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744 yellow, cranial nerves; pink, inner ear; red, arteries. Paratympanic sinus system colors

745 were based on Dufeau and Witmer, (2015): intertympanic diverticula, blue;

746
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pharyngotympanic sinus, green; median pharyngeal recess, purple; basisphenoid
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747 diverticula, violet. Paranasal sinuses colors were based on Witmer and Ridgely, (2008):
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748 nasal cavity, yellow; dorsal alveolar duct, red; postvestibular sinus, blue; antorbital

749 sinus, green; nasolacrimal duct, orange. Scale bars equal 1cm. [planned for 2 column
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750 size]
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751
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752 Table1: Measurements for the olfactory capability calculations, based on Zelenitsky et

753 al. (2011, 2009). These data were plotted for a better visualization, in Suppl. Data
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755 Table 2: Measurements for the auditory capability calculations, based on Walsh et al.

756 (2009). Abbreviations: ECD, endosseous cochlear duct.

757

758 Appendix Fig. 1: Plot of the maximum endocast diameter versus the maximum

759 olfactory bulb diameter in the studied eusuchians.

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760

761 Appendix tables: Measurements for the semicircular canals, optic capability and REQ.

762

763 Appendix Fig. 2: Plot and regression of the relative volume of the encephalon versus

764 the endocast total volume during the ontogeny of crocodiles, basing on the data of Jirak

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765 and Janacek, (2017).

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766

767 Appendix skull.pdf: Interactive 3D PDFs of the inner skull cavities reconstructions are

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768 provided as supplementary figures to this paper.

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Semicircular canal maximum diameter (mm) Semicircular canal circumference (mm)

Taxon
lateral canal anterior canal posterior canal lateral canal anterior canal posterior canal

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A. mississippiensis 6,54 9 6,22 14,7 21,41 14
C. crocodilus 4,39 6,71 4,23 11,01 17,11 11,15
C. niloticus 6,5 10,39 6,75 15,04 24,79 15,78

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O. tetraspis 7,07 9,43 6,4 16,32 21,63 14,44
T. schlegelii 7,09 9,45 5,72 16,9 23,66 14,8

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G. gangeticus 7,85 10,56 7,62 18,18 26,22 18,41
L. megadontos 3,81 2,89 3,71 9,99 7,78 9,86
E. dentata 2,14 3,07 2,28 5,15 6,66 4,95

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A. melanocephalus 2,768 2,965 2,805 6,059 6,467 6,035

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V. exanthematicus 3,73 3,91 3,8 8,08 8,69 8,42
S. camelus 3,79 7,13 4,27 9,81 18,64 11,06
C. atratus 4,901 8,183 - 12,917 20,219 -

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Endocast Optic lobe Relative

Taxon
volume (mm3) volume (mm3) Volume
A. mississippiensis 18719,9 2467,94 0,13

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C. crocodilus 4941,74 936,482 0,19
C. niloticus 24764,8 3789,03 0,15
O. tetraspis 11593,1 2577,36 0,22

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T. schlegelii 19390,5 2105,72 0,11
G. gangeticus 38309,9 6996,52 0,18

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L. megadontos 13604,5 1885,87 0,14
Measurements for the relative size of the optic region, based on Jirak and Janacek, (2017).

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Endocast Brain volume*

Taxon Body mass (g) REQ
volume (cm3) (cm3)
A. mississippiensis 18,72 6,91 104328,281 0,94

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C. crocodilus 4,94 2,75 5887,67 1,74
C. niloticus 24,76 8,38 151389,302 0,94
O. tetraspis 11,59 4,96 20228,392 1,62

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T. schlegelii 19,39 7,08 93619,603 1,02
G. gangeticus 38,31 11,32 170000** 1,19

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L. megadontos 13,60 5,54 88719,462 0,82
Measurements for the Reptile Encephalization Quotient calculations, based on (Hurlburt et al., 2003). *The brain volume was estimated introducing the data
of Jirak and Janacek, (2017) in a regression plot (Suppl Fig. 2). The body mass was estimated using skull measurements, basing in 1Dodson, (1975); 2Webb

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Olfactory bulb Cerebral hemispheres Log olfactory
Taxon
(mm) (mm) ratio
A. mississippiensis 17,3 29,8 1,76
C. crocodilus 10,92 19,62 1,75
C. niloticus 20,37 30,5 1,82
O. tetraspis 15,43 24,9 1,79
T. schlegelii 19,13 28,89 1,82
G. gangeticus 16,65 33,29 1,70
L. megadontos 20,8 33,52 1,79

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E. dentata 4,56 9,26 1,69
A. melanocephalus 6,6 10,71 1,79
V. exanthematicus 7,21 15,56 1,67

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Taxon ECD length Basicranial Scaled ECD Scaled/transformed
(mm) axis (mm) length ECD length
A. mississippiensis 7,6 92,38 0,08 -1,08
C. crocodilus 6,6 35,28 0,19 -0,73
C. niloticus 8,55 104,33 0,08 -1,09
O. tetraspis 8 70,77 0,11 -0,95
T. schlegelii 8,21 73,88 0,11 -0,95
G. gangeticus 8,8 86,33 0,10 -0,99
L. megadontos 6,55 87,76 0,07 -1,13

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