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Figure 1.

Photographs of the holotype of Europejara olcadesorum gen. et sp. nov. (MCCM-LH 9413).

(A) Main slab under ultraviolet light. (B) Acid-prepared counterslab. Scale bar: 50 mm.

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Figure 2.

Line drawing of the holotype of Europejara olcadesorum gen. et sp. nov. and life restoration.

(A) Interpretative line drawing of the skull as observed on the acid-prepared counterslab. (B) Reconstruction of the skull (based in part on Tapejara) showing preserved parts in red. Life restoration of the head of Europejara in lateral (C) and frontal (D) views. apj, anterior process of the jugal; aprj, anterior process of the right jugal; d, dentary; ec, ectopterygoid; hy, hyoids; j, jugal; l, lacrimal; ld, left dentary; lj, left jugal; lm, left maxilla; lpo, left postorbital; lq, left quadrate; lsa, left surangular; lsq, left squamosal; ltf, lower temporal fenestra; m, maxilla; naof, nasoantorbital fenestra; o, orbit; pf, postfrontal; po, postorbital; pt, pterygoid; q, quadrate; rap, retroarticular process; rd, right dentary; rm, right maxilla; scd, sagittal crest of the dentary; scp, scleral plates; sq, squamosal. Scale bar: 50 mm.

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Figure 3.

Skull of Europejara olcadesorum gen. et sp. nov.

(A) Close-up of the crushed left post-orbital region of the skull (acid-prepared counterslab) in lateral view showing the narrowness of the lower temporal fenestra (ltf), the mandibular condyle of the quadrate (cq) and the distal extremities of the hyoid apparatus (hy). (B) Close-up of the posterior area of the palate (main slab under ultraviolet light) in dorsal view showing the thin, elongated vomer (v) septum separating the two choanae (ch). Note the medial surface of the left dentary (ld) and the hyoid apparatus (hy), adjacent to the ventral margin of the mandible. (C) Detail of the posterior area of the palate (acid-prepared counterslab) in dorsal view showing the pterygoid (pt), the ectopterygoid (ec), and the right maxilla (rm). Note the robustness of right mandibular ramus (in lateral view) and its thin, edentulous dorsal edge (arrow). ch, choanae; cq, mandibular condyle of the quadrate; ec, ectopterygoid; hy, hyoid apparatus; j, jugal; lbd, lingual bulge of the dentary; ld, left dentary; lft, lower temporal fenestra; po, postorbital; pt, pterygoid; q, quadrate; rd, right dentary; rm, right maxilla; sq, squamosal; v, vomer. Scale bars: 10 mm.

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Figure 4.

Lower jaw of Europejara olcadesorum gen. et sp. nov.

(A) Close-up of the symphyseal area (acid-prepared counterslab) in right lateral view showing the typical step-like dorsal margin of the dentary in tapejarines (arrow). Note the strong lateral compression of the mandible and the trabecular structure of the sagittal crest of the dentary (scd). (B) Close-up of the best preserved margin of the dentary crest (main slab). Note the concave posterior border (arrow) giving the peculiar recurved aspect of the dentary crest of Europejara. scd, sagittal crest of the dentary. Scale bars: 10 mm.

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Figure 5.

Comparisons of tapejarine lower jaws.

(A) Europejara olcadesorum gen. et sp. nov. (specimen MCCM-LH 9413): dentary crest height/mandibular ramus height (DCH/MRH) ratio = 4. (B) Tupandactylus imperator (specimen CPCA 3590): DCH/MRH ratio = 3. (C) Tapejara wellnhoferi (specimen AMNH 24440): DCH/MRH ratio = 2.5. (D) Sinopterus dongi (specimen GMN-03-11-001, holotype of Huaxiapterus jii): DCH/MRH ratio = 2.2. (E) Sinopterus dongi (specimen IVPP V 13363): DCH/MRH ratio = 1.5. (F) Kem Kem tapejarine (specimen BSP 1997 I 67): DCH/MRH ratio unknown. Gray areas indicate missing parts. Scale bars: 50 mm.

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Figure 6.

Strict consensus cladogram showing the relationships of Europejara olcadesorum gen. et sp. nov. to other Pterodactyloidea.

Europejara olcadesorum was found to be a member of the clade Tapejarinae (sensu [7]). Tree length = 205, consistency index = 0.7561, retention index = 0.8538, rescaled consistency index = 0.6456 (see Appendix S1 for character list and data matrix). Nodes: 1, Pterosauria; 2, Pterodactyloidea; 3, Azhdarchoidea; 4, Tapejarinae (sensu [7]). As indicated by its toothless condition, Europejara is included within the clade Azhdarchoidea (synapomorphy: character 57, state 3). Europejara shares with other tapejarines the apomorphic feature consisting of the presence of a step-like dorsal margin of the dentary in lateral view (synapomorphy: character 52, state 1).

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Figure 7.

Distribution through time and space of tapejarine pterosaurs and early angiosperms.

(A) Early Cretaceous (Aptian) paleogeographical map showing the three main areas where tapejarids co-occur with early angiosperms (black stars). (B) Stratigraphic distribution of the different tapejarid taxa (silhouettes above each taxon denote geographical occurrences); note that the diversification of tapejarids coincides with the Phase I of the early angiosperm diversification (EAD) [60][62]. (C) Leaves of an early angiosperm (cf. Jixia) from the late Barremian of Las Hoyas (MCCM-LH 30351), one of the oldest known macrofossil of a terrestrial flowering plant. Early angiosperm pollen grains Afropollis (D) and Stellatopollis (E) from the La Huérguina Formation, both worldwide distributed during the Barremian–Aptian interval. Ber, Berriasian; Val, Valanginian; Hau, Hauterivian; Bar, Barremian; Apt, Aptian; Alb, Albian; Cen, Cenomanian; Tur, Turonian; Con, Coniacian; San, Santonian; Cam, Campanian; Maa, Maastrichtian. Scale bars: 10 mm (C) and 10 µm (D,E).

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