Supporting Information

Daura et al. 10.1073/pnas.1619040114 SI The Gruta da Aroeira Site is not significant for samples with ages around 400 ka, so the Gruta da Aroeira (39° 30′ 20′′ N; 08° 36′ 57′′ W) is located in corrected and uncorrected results agree within uncertainties, and Torres Novas, central Portugal (Fig.1). The is part of the the results are robust. All the other samples have low detrital Almonda system, a labyrinthine network of passages con- components, and the correction is not significant. The un- taining a number of former entrances with Pleistocene infillings corrected and corrected ages are reported in Table S1 and are completely sealed by roof collapses. This system is formed of discussed in the next section. passages excavated at different elevations whose intersections with the 70-m-high escarpment rising above the extant spring of the SI Chronostratigraphy Almonda River, a tributary of the Tagus River, correspond to A minimum of five site formation phases can be differentiated fossil outlets of its subterranean course (Fig. S1). (Fig. 2 and Table S1). Episode 5 corresponds to the period Published reports of previous excavation work, carried out be- predating the opening of the cave and includes layer XII (unit 3) tween 1998 and 2002, have used the designation of “Galerias and the formation of a massive stalagmitic column that, based on Pesadas” for this site (6). However, this designation corresponds available cross-sectional views, would seem to have grown from to inner conduits of the karst system situated at this elevation that, its top. The column’s last layer of calcite, formed before the at present, remain unconnected to the exterior. In 1991, spe- accumulation of the deposit that eventually buried it, dates to leoarcheological surveys identified a cone of sediments in these 406 ± 30 ka (sample no. 8) and provides a minimum age for the conduits with Pleistocene fauna and stone tools (3) and eventually sequence of events subsumed under episode 5 and a maximum led to the location of a collapsed, sediment-sealed entrance—the age for the deposition of the Aroeira 3 cranium inside the cave. Gruta da Aroeira. The 1998–2002 excavations, as well as the new Episode 4 is represented by the accumulation of layer X (the phase of the project begun in 2013, were carried out here (Fig. 1 Acheulean deposit) and layer XI. The age of the cranium, found and Fig. S1). at the base of layer X, is bracketed by the age of the outermost layer of the stalagmitic column formed during the SI Stratigraphic Outline previous phase, against which it leaned and the basal age of the At the back of the cave, where bedrock probably has been reached, ARO2 flowstone (sample no. 2.1; 418/+37/−27 ka); therefore, the Aroeira stratigraphy spans a thickness of ∼4 m and comprises the age of the cranium must lie in the 390–436 ka interval. The three major stratigraphic units. Uppermost unit 1 is an infilling dating of calcite crystals sampled from the cranium’s breccia fill breccia capped by the ARO1 flowstone. Unit 2 is a 2.2-m-thick (samples no. 3 and 4) has provided minimum ages (390 ± 14 ka mud-supported breccia rich in angular and subangular clasts, and 408 ± 18 ka, respectively) consistent with this stratigraphic corresponding to the Acheulean layer X–Xb/c (upper and lower bracketing (Table S1). parts of a single layer, excavated in 1998–2002 and 2013–2015, Episode 3 corresponds to a sedimentation hiatus, probably be- respectively), and is capped by the ARO2 flowstone. The human cause of the infilling of the entrance to the cave. During the hiatus, cranium (inventory no. ARO-606, found in square H6, at a depth the ARO2 flowstone formed over the then-extant cave floor. In of 608 cm below datum) was recovered at the base of this unit (Fig. areas where the fill was not fully sealed, and as a result of sub- 1 and Fig. S4). Basal unit 3 is an endokarst fluvial deposit and sidence/suffosion, water percolating from above precipitated calcite comprises two layers: XI is a 0.4-m-thick layer of silty sand with in the voids and along cracks and fissures formed between sediment scattered gravel and faunal remains but no artifacts, and XII is a and cave wall or between sediment and the massive stalagmitic 0.5-m-thick, archeologically sterile, layer of slightly gravelly sand. column of episode 1. This water percolation would explain the results of 300 ± 3 ka for crystals formed in voids toward the top of SI U-Series Results layer X (samples no. 5 and 6) and of 236 ± 3 ka for calcite formed U concentrations of ARO1 and ARO2 are between 40 between the sediment column and the outer, sediment-dirty rim of and 90 ng/g (Table S1). The base of BL1 has a higher U the stalagmitic column (sample no. 7). Together with the dates concentration of 171 ng/g. The calcite crystals that formed in obtained for the upper part of the ARO2 flowstone (sample no. sediment voids (ARO14-03 and ARO14-04) have U concentra- 2.2.; 326.4 ± 13.4 ka) and for the BL1 stalagmite (sample BL1.1; tions of 191 and 129 ng/g, respectively; the outer layer of the 278.5 ± 12.7 ka) found exteriorly at the Brecha das Lascas locus stalagmite column (ARO14-H6-727) has a concentration of 105 ng/g; but in the same stratigraphic position, these results place the end of and a postunit 2 sedimentation crust on top of the column has much this episode after 239 ka. higher U concentration of 613 ng/g. The calcite crusts that formed Episode 2 corresponds to a collapse of the roof in the area of on the cranium (ARO-SK4 and ARO-SK6) have U concentrations rows 10–14 of the grid. A second round of infill deposits even- of 305 and 273 ng/g, respectively. tually filled the newly formed shaft. 232Th is a proxy for detrital components in the calcium car- Episode 1 corresponds to the formation of the ARO1 flowstone bonate, and the degree of contamination is assessed by today’s (sample no. 1). This flowstone is found at the top of the fill in the 230Th/232Th activity ratio. A correction for the detrital contribution interior space behind the new infill deposit, for which it provides a is done using a detrital 238U/232Th activity ratio of 0.8 with 50% minimum age of 44.8 ± 2.0 ka. uncertainty and also assuming secular equilibrium of the 238U Based on these results, we propose the following correlation of decay chain in the detritus. ARO1 has a high degree of detrital the sequence with the global record of MIS: episode 5, MIS 11c or components, and two of three dating results (ARO1-1 and earlier; episode 4, MIS 11c, ∼390–420 ka; episode 3, MIS 7a–11b, ARO1-3) are dominated by detritus, resulting in high uncer- ∼190–390 ka; episode 2, MIS 4–7a, ∼60–190 ka; and episode 1, tainties. ARO1-2 also has a high detrital contribution, but the MIS 3c ∼40–60 ka. corrected age is robust. The results for the column and the precipitates on the cranium also show a high detrital contribu- SI In Situ Extraction of the Fossil tion, with 230Th/232Th around 30 for ARO-SK-4 and ARO-SK-6 The cranium, heavily fossilized and well preserved, was contained and >10 for the column sample. However, the detrital correction within a cemented breccia (Fig. S4). Because of the extreme

Daura et al. www.pnas.org/cgi/content/short/1619040114 1of9 hardness of the sediments and the difficulty of excavation, several Subsequent to the acquisition of the Portuguese Heritage fragments of the cranium were separated from the main por- authority’s pertinent temporary export permit, the block con- tion at the moment of discovery by the impact of the heavy-duty taining the main portion of the cranium and the detached frag- demolition hammer being used at the time (hence, the circular ments were transported to the Conservation and Restoration hole apparent in Fig. 3 and Figs. S4–S7). The detached frag- Laboratory at the Centro de Investigación Universidad Com- ments and the contour and thickness of the sectioned cranial plutense de Madrid-Instituto de Salud Carlos III sobre la vault made it immediately apparent that a human fossil had Evolución y Comportamiento Humanos in Madrid for further been hit by the tool. Work was interrupted on the spot, all the preparation. The cranium was painstakingly extracted from the detached fragments were collected, and preparations for ex- breccia and restored over a period of two years (Figs. S5 and S6). traction of the remainder from the breccia were initiated. The The most recent criteria for conservation and restoration were visible sections of the main portion of the fossil were protected followed, including prioritizing the conservation of the fossil over with gauze coating impregnated with Paraloid B-72 (Rohm & its restoration and using products that are homogeneous and Haas) at 5–15% acetone (CH3COCH3) concentration (47–49). compatible with the bone as well as reversible to facilitate future Subsequently, the fossil was covered with a polyurethane resin interventions and avoid additional damage. A detailed diagnosis to protect it further during the rock-cutting, with appropriate of the specimen was carried out before each intervention, and all machinery, of a large block of the hard calcareous breccia that steps during the conservation and restoration process were contained it (Fig. S4). documented (50, 51).

Daura et al. www.pnas.org/cgi/content/short/1619040114 2of9 Fig. S1. (A and E) Almonda escarpment with the position of Gruta da Aroeira and the Almonda River in the foreground. (B) General view of Gruta da Aroeira and the location of the main dated . (C) Estremadura Limestone Massif with the position of the Almonda spring, the Tagus River, and the Serra d’Aire. (D) Schematic cross-section of the Almonda karst system. (F) Acheulean biface (flint) from level Xb/c. (G) Gruta da Aroeira site plan.

Daura et al. www.pnas.org/cgi/content/short/1619040114 3of9 Fig. S2. Acheulean handaxes (bifaces) from level Xb/c. (A) Flint. (B–E) Quartzite.

Fig. S3. (A and B) The Aroeira 1 left mandibular canine (A) and the Aroeira 2 left maxillary third molar (B) in mesial (Mes), distal (Dist), buccal (Bucc), and lingual view (Ling). (C) Aroeira 3 maxilla in medial view. Arrows indicate the lateral nasal crest (1), turbinal crest (2); nasal floor (3), and root of the inferior nasal concha (4). (Scale bar, 2 cm.)

Daura et al. www.pnas.org/cgi/content/short/1619040114 4of9 Fig. S4. Various stages during the in situ extraction and restoration process of the fossil, and reconstruction of the Aroeira 3 cranium after cleaning. (A) Outline of the cranium embedded in the breccia. (B) Location of the cranium after its protection with gauze coating and polyurethane resin. (C) Cutting of the breccia with a circular saw to remove the cranium. (D) Protection of the fossil with wooden boards during the fracturing of the adhering to the wall with a pneumatic hammer. (E) Final stage of the extraction of the cranium. Wooden boards were placed along both sides of the wall while the breccia block containing the cranium was cut along the bottom. (F) Main portion of the cranium embedded within the breccia block. (G) Isolated cranial fragments before restoration. (H) Main portion of the cranium during the removal of the hardened sediments from the endocranial surface. (I) Main portion of the cranium during the cleaning process (Left) and reconstruction of some of the isolated cranial fragments (Right). (J) Detail of the cranial base and temporal bone during the cleaning process. The arrow indicates a thin layer of speleothem coating which remains on the superior portion of the petrous pyramid. (K) Fragments comprising the cranium before reconstruction. (L) Manually joining the fragments together with adhesive. (M) Endocranial view of the reconstructed cranium. (N) Lateral view of the reconstructed cranium. (Scale bars, 5 cm.) Images in K–N are from J. Trueba (photographer).

Daura et al. www.pnas.org/cgi/content/short/1619040114 5of9 Fig. S5. Virtual reconstruction of the Aroeira 3 cranium in frontal (A), posterior (B), superior (C), and endocranial (D) views. The frontal sinus in D is exposed in a parasagittal section located 4 mm to the right of the sagittal plane. (E) Virtual reconstruction of the Aroeira 3 cranium in a three-quarters view compared with Bilzingsleben B1 (cast).

Daura et al. www.pnas.org/cgi/content/short/1619040114 6of9 Fig. S6. The Aroeira 3 cranium compared with Atapuerca SH Cranium 4 and Cranium 5. Atapuerca SH Cranium 5 is oriented in the Frankfurt horizontal orientation; this orientation is estimated for the Aroeira 3 cranium and Atapuerca SH Cranium 4. Aroeira 3 shows a slightly depressed nasion (n) with respect to the glabella (g), whereas in the Atapuerca SH Cranium 5 the nasion and glabella are located in the same vertical plane. The mastoid projection measured from the parietal incisure is shorter in the Aroeira 3 cranium than in the Atapuerca (SH) crania, but the projection is similar in all three crania when measured from the level of the porion. It is possible that the relatively high position of the transverse sulcus in the Aroeira 3 cranium (see Text) is related to the low position of the parietomastoid suture and asterion (a). A clear torus angularis (t.a.) is present is both the Aroeira 3 cranium and Atapuerca SH Cranium 4 but can only be insinuated in Atapuerca SH Cranium 5.

Fig. S7. The Aroeira 3 cranium compared with Atapuerca SH Cranium 5 and the Steinheim specimen. The Frankfurt horizontal orientation is estimated in both the Aroeira 3 cranuim and the Steinheim specimen. The nasion seems depressed relative to the glabella in the Steinheim cranium, but the strong deformation of this specimen precludes a conclusive assessment. The Aroeira 3 cranium is reminiscent of the Steinheim specimen in both the low position of the parie- tomastoid suture (compared with Atapuerca SH Cranium 4 and 5) and the corresponding short mastoid process when measured from the parietal incisure.

Daura et al. www.pnas.org/cgi/content/short/1619040114 7of9 ar tal. et Daura

www.pnas.org/cgi/content/short/1619040114 Table S1. U-Th results from Aroeira speleothems, calcite samples adhering to the cranium, and the stalagmite column Sample field-sub Sample 238 232 230 232 230 238 234 238 234 238 ID no. sample no. laboratory ID no U, ng/g Th, ng/g [ Th/ Th] [ Th/ U] [ U/ U] Uncorrected age, ka Corrected age, ka [ U/ U]initial Observations

1.3 ARO1-3 CENIEH-UT0276 49.4 ± 0.3 66.924 ± 0.428 1.22 ± 0.02 0.540 ± 0.010 1.073 ± 0.004 75.65 ± 2.00 32.5 ± 25.7 1.124 ± 0.037 Upper part of flowstone capping the Pleistocene fill 1.2 ARO1-2 CENIEH-UT0275 42.2 ± 0.3 5.578 ± 0.067 8.83 ± 0.16 0.382 ± 0.006 1.063 ± 0.004 48.37 ± 1.04 44.8 ± 2.0 1.074 ± 0.005 Middle of same flowstone 1.1 ARO1-1 CENIEH-UT0214 41.5 ± 0.3 28.202 ± 0.166 2.17 ± 0.02 0.483 ± 0.005 1.069 ± 0.004 65.05 ± 1.03 45.4 ± 10.4 1.096 ± 0.013 Basal part of the same flowstone; minimum age of the underlying deposit 2.2 ARO2-2 CENIEH-UT0212 70.3 ± 0.3 1.125 ± 0.010 203.84 ± 1.44 1.067 ± 0.006 1.093 ± 0.003 326.80 ± 13.40 326.4 ± 13.4 1.234 ± 0.008 Upper part of flowstone capping the Acheulian layer 2.1 ARO2-1 CENIEH-UT0211 88.6 ± 0.4 1.112 ± 0.013 261.05 ± 2.06 1.071 ± 0.006 1.069 ± 0.003 418.01 ± 31.50 417.7 +37.3/−27.5 1.226 ± 0.016 Basal part of the same flowstone; minimum age of the site’s Acheulian occupation BL1-1 CENIEH-UT0210 171.0 ± 1.1 1.184 ± 0.027 413.8 ± 7.7 0.938 ± 0.009 1.013 ± 0.003 278.7 ± 12.7 278.5 ± 12.7 1.0287 ± 0.006 Basal part of stalagmite growing on the flowstone that caps the Pleistocene fill at the site’s Brecha das Lascas locus 3 ARO-SK4-1 UEVA 495 304.7 ± 1.1 31.5 ± 0.1 30.71 ± 0.06 1.039 ± 0.003 1.050 ± 0.002 393 ± 13 390 ± 14 1.156 ± 0.005 Samples of syndepositionally formed calcite taken for U-series dating from the breccia adhering to inner side of the partial cranium 4 AR-SK6-1 UEVA 496 273.2 ± 1.1 29.0 ± 0.1 29.82 ± 0.06 1.037 ± 0.003 1.045 ± 0.002 410 ± 17 408 +19/-16 1.147 ± 0.006 Samples of syndepositionally formed calcite taken for U-series dating from the breccia adhering to inner side of the partial cranium 5 ARO 14-03-1 UEVA 137 191.3 ± 0.6 1.83 ± 0.01 318.2 ± 0.7 0.995 ± 0.003 1.048 ± 0.002 300 ± 6300± 6 1.112 ± 0.003 Calcite crystals precipitated in sediment voids 6 ARO 14-04-1 UEVA 138 129.1 ± 0.3 3.25 ± 0.01 120.8 ± 0.3 0.994 ± 0.003 1.044 ± 0.002 307 ± 6307± 6 1.104 ± 0.003 Calcite crystals precipitated in sediment voids 7 ARO 14-H6-727-2 UEVA 139 613.1 ± 2.7 19.36 ± 0.08 87.69 ± 0.17 0.906 ± 0.003 1.018 ± 0.002 237 ± 3236± 3 1.036 ± 0.003 The outermost flowstone overgrowth formed post-depositionally at the interface between the Acheulian layer and the stalagmitic column buried by that layer 8 ARO 14-H6-727-3 UEVA 493 105.0 ± 0.5 28.8 ± 0.1 11.79 ± 0.04 1.058 ± 0.005 1.061 ± 0.003 413 ± 24 406 +34/-26 1.206 ± 0.013 The formation of the stalagmitic column predates the accumulation of the sediments containing the cranium, and its outer layer provides a terminus post quem for the fossil

All ratios are activity ratios. The degree of detrital 230 Th contamination is indicated by the measured 230 Th/232 Th activity ratio, and corrections were calculated using a 238 U/232 Th activity ratio of 0.8 ± 0.4 and assuming secular equilibrium of the 238 U decay chain in the detritus. Analytical errors are at the 95% confidence level. Activity ratios are calculated from isotope concentration ratios using decay constants according to refs. 52 (λ238 ), 53 (λ234 and λ230 ), and 54 (λ232 ). 8of9 Table S2. Principal measurements (in millimeters) in Aroeira 3 compared with the Atapuerca (SH) sample Howell’s Atapuerca SH Atapuerca SH Atapuerca SH Atapuerca (SH) adults‡, † Measurement Martin no.* abbreviation Aroeira 3 Cranium 4 Cranium 5 Cranium 6 mean ± SD (n)

Maximum cranial breadth M8 XCB 143–145 164.0 146.0 136.0 152.8 ± 11.85 138.9–175.7 (8) Minimum frontal breadth M9 <116.0 117.0 105.7 100.0 Maximum frontal breadth M10 XFB 122–124 126.0 118.5 117.1 ± 6.93 108.3–129.6 (9) Bistephanic breadth M10b STB 122–124 119.5 110.8 116 Biauricular breadth M11 AUB 136–138 155.5 139.0 122.0 Biasterionic breadth M12 ASB 121–125 132.0 116.5 117.6 Auriculo–bregmatic height M20 124–125 121.0 107.5 121.0 Frontal sagittal chord M29 FRC 113.9 115.0 106.0 98.0 109.0 ± 5.11 101–115 (6) Bregma–asterion chord M30c 136–137 143.5 129.4 137.5 Interorbital breadth M49(a) DKB 37.3 38.0 33.0 Anterior interorbital breadth M50 37.3 31.2 30.0 Bregma radius BRR 102–103 107.8 97.7 115.0

Bilateral measurements in Aroeira 3 are based on the mirror-imaged virtual reconstruction. *After ref. 55. † Ref. 56. ‡Includes the late adolescent/young adult individuals: crania 7, 10, and 16 (23).

Daura et al. www.pnas.org/cgi/content/short/1619040114 9of9