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Fig 1.

Geological and geographical setting of La Retama fossil site within the Loranca Basin in the Iberian Peninsula.

Map illustration by IMS, modified from [41].

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Fig 2.

Cranial remains of Xenokeryx amidalae gen. et sp. nov. from La Retama.

A, MNCN-74448 (holotype), occipital appendage in anterior view; B, MNCN-74448 (holotype), occipital appendage in lateral view; C, MNCN-74448 (holotype), occipital appendage in posterior view; D, MNCN-74449, right ossicone and supra-orbital region of a skull in latero-distal view; E, MNCN-74449, right ossicone and supra-orbital region of the skull in medial view; F, MNCN-74446, left juvenile ossicone and supra-orbital region of the skull in lateral view; G, MNCN-74446, left juvenile ossicone and supra-orbital region of the skull in medial view; H, MNCN-74447, left juvenile ossicone in apical view, showing its transversal section; I, MNCN-74449, detail of the ossicone-frontal bone contact showing the suture line (not to scale). Abbreviations: Bp, bumps; Nf, nuchal fossa; Ow, ossicone ‘wing’; Sl, suture line between ossicone and frontal bone.

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Fig 3.

Upper dentition remains of Xenokeryx amidalae gen. et. sp. nov. from La Retama.

A, MNCN-74495, left maxillar with DP2-M1 in buccal view; B, MNCN-74495, left maxillar with DP2-M1 in occlusal view; C, MNCN-74452, right M3 in lingual view; D, MNCN-74452, right M3 in buccal view; E, MNCN-74452, right M3 in occlusal view; F, MNCN-74450, left P4-M1 in lingual view; G, MNCN-74450, left P4-M1 in buccal view; H, MNCN-74450, left P4-M1 in occlusal view; I, MNCN-74451, right P3 in lingual view; J, MNCN-74451, right P3 in buccal view; K, MNCN-74451, right P3 in occlusal view; L, MNCN-74453, right P2 in buccal view; M, MNCN-74453, right P2 in lingual view; N, MNCN-74453, right P2 in occlusal view.

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Fig 4.

Lower dentition remains of Xenokeryx amidalae gen. et sp. nov. from La Retama.

A, MNCN-74455, right hemimandibular fragment with m2-m3 in buccal view; B, MNCN-74455, right hemimandibular fragment with m2-m3 in lingual view; C, MNCN-74455, right hemimandibular fragment with m2-m3 in occlusal view; D, MNCN-74456, left hemimandibular fragment with m1-m3 in buccal view; E, MNCN-74456, left hemimandibular fragment with m1-m3 in lingual view; F, MNCN-74456, left hemimandibular fragment with m1-m3 in occlusal view; G, MNCN-74458, right hemimandible with p3-m3 in buccal view; H, MNCN-74458, right hemimandible with p3-m3 in lingual view; I, MNCN-74458, right hemimandible with p3-m3 in occlusal view; J, MNCN-74457, right m3 in buccal view; K, MNCN-74457, right m3 in lingual view; L, MNCN-74457, right m3 in occlusal view.

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Fig 5.

Fore limb remains and phalanges of Xenokeryx amidalae gen. et sp. nov. from La Retama.

A, MNCN-74460, proximal fragment of right radius in cranial view; B, MNCN-74460, proximal fragment of right radius in caudal view; C, MNCN-74459, distal fragment of right radius in cranial view; D, MNCN-74459, distal fragment of right radius in caudal view; E, MNCN-74472, proximal fragment of metacarpal III-IV in proximal view; F, MNCN-74473, proximal fragment of left radius in proximal view; G, MNCN74461, articular fragment of left scapula in lateral view; H, MNCN-74465, distal fragment of metacarpal III-IV in dorsal view; I, MNCN-74465, distal fragment of metacarpal III-IV in palmar view; J, MNCN-74486, first phalanx in external view; K, MNCN-74486, first phalanx in interdigital view; L, MNCN-74486, first phalanx in palmar/plantar view; M, MNCN-74487 in external view; N, MNCN-74487 in interdigital view; O, MNCN-74493, right pyramidal in lateral view; P, MNCN-74493, right pyramidal in medial view; Q, MNCN-74484, right semilunate in proximal view; R, MNCN-74484, right semilunate in distal view; S, MNCN-74483, left magnotrapezoid in proximal view; T, MNCN-74483, left magnotrapezoid in distal view.

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Fig 6.

Hind limb remains of Xenokeryx amidalae gen. et sp. nov. from La Retama.

A, MNCN-74471, proximal fragment of left femur in cranial view; B, MNCN-74471, proximal fragment of left femur in caudal view; C, MNCN-74475, distal fragment of metatarsal III-IV in dorsal view; D, MNCN-74464, proximal fragment and diaphysis of left metatarsal III-IV in dorsal view; E, MNCN-74464, proximal fragment and diaphysis of left metatarsal III-IV in plantar view; F, MNCN-74466, distal fragment of left tibia in cranial view; G, MNCN-74466, distal fragment of left tibia in caudal view; H, MNCN-74466, distal fragment of left tibia in distal view; I, MNCN-74469, right calcaneus in medial view; J, MNCN-74469, right calcaneus in lateral view; K, MNCN-74470, right calcaneus in medial view; L, MNCN-74470, right calcaneus in lateral view; M, MNCN-74477, right navicular-cuboid in proximal view; N, MNCN-74479, left navicular-cuboid in proximal view; O, MNCN-74477, right navicular-cuboid in distal view; P, MNCN-74479, left navicular-cuboid in distal view; Q, MNCN-74480, right navicular-cuboid in plantar view; R, MNCN-74502, right ectomesocuneiform in proximal view; S, MNCN-74501, right ectomesocuneiform in proximal view; MNCN-74500, right ectomesocuneiform in proximal view; U, MNCN-74476, left astragalus in dorsal view; V, MNCN-74476, left astragalus in plantar view; X, MNCN-74482, right malleolar in medial view. Abbreviations: Cr, crest.

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Fig 7.

Phylogenetic relationships among palaeomerycids.

Bootstrapped MPT of 45 steps (CI = 0,889; RI = 0,861). Bootstrap values (1000 replicates) over each node.

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Table 1.

Xenokeryx and palaeomerycids, distribution of autapomorphic character states for the internal nodes and Xenokeryx.

Ambiguous synapomorphies in italics.

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Fig 8.

Phylogenetic relationships of palaeomerycids within the Pecora.

Bootstrapped MPT of 205 steps (CI = 0,541; RI = 0,700). Bootstrap values (1000 replicates) over each node. Dromomerycid and palaeomerycid terminals are highlighted.

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Fig 9.

Phylogenetic relationships of palaeomerycids within the Pecora.

Maximum clade credibility time-tree of major ruminant clades obtained from our ‘tip-dating’ Bayesian analysis. Uncertainty in divergence ages is shown with horizontal grey bars representing their 95% highest posterior densities (HPDs). Numbers above each branch show the posterior probability (PP) of the respective clade.

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Table 2.

Palaeomerycids within the Pecora, distribution of character states for the discussed internal nodes of the MPT.

Ambiguous synapomorphies in italics.

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Table 3.

Palaeomerycids within the Pecora, distribution of the reconstructed morphological character states for the discussed internal nodes of the Bayesian tree.

Ambiguous synapomorphies in italics.

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Fig 10.

Life reconstruction of the head of Xenokeryx amidalae gen. et sp. nov.

Adult male based on the fossils from La Retama. Illustration by IMS.

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Fig 11.

Evolution of palaeomerycids.

Summary scheme showing a calibrated phylogeny of palaeomerycids (based on the corresponding MPT) with special emphasis on the main morphological traits of each basal clade (Ampelomeryx-clade and Triceromeryx-clade represented by reconstructions of Ampelomeryx and Xenokeryx) including their biogeographic distribution. Illustrations by IMS.

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