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Home , Ein Gev, Erik Trinkaus, Jebel Sahaba, Late Stone Age, La Ferrassie, Taforalt

Late age and archaeological context for the hominin calvaria from GvJm-22 (Lukenya Hill, )

Christian A. Tryona,1, Isabelle Crevecoeurb, J. Tyler Faithc, Ekshtaina, Joelle Nivensd, David Pattersone, Emma N. Mbuaf, and Fred Spoorg,h aDepartment of , Harvard University, Cambridge, MA 02138; bUnité Mixte de Recherche 5199, de la Préhistoire à l’Actuel: Culture, Environnement, et Anthropologie, Centre National de la Recherche Scientifique, Université de Bordeaux, 33615 Talence, ; cArchaeology Program, School of Social Science, University of Queensland, Brisbane, QLD 4072, ; dDepartment of Anthropology, New York University, New York, NY 10003; eCenter for the Advanced Study of Hominid Paleobiology, Department of Anthropology, The George Washington University, Washington, DC 20052; fNational Museums of Kenya, Nairobi, Kenya 00100; gDepartment of , Max Planck Institute for Evolutionary Anthropology, D-04103, Leipzig, ; and hDepartment of Cell and Developmental , University College London, WC1E 6BT London,

Edited by Erik Trinkaus, Washington University, St. Louis, MO, and approved January 16, 2015 (received for review September 19, 2014)

Kenya National Museums Lukenya Hill Hominid 1 (KNM-LH 1) is Hominid 1 (KNM-LH 1) partial calvaria from site GvJm-22 at a Homo sapiens partial calvaria from site GvJm-22 at Lukenya Hill, Lukenya Hill, Kenya, the only eastern African fossil hominin Kenya, associated with Later (LSA) archaeological de- from a Last Glacial Maximum [LGM; 19–26.4 kya (19)] LSA posits. KNM-LH 1 is securely dated to the , and archaeological context. We construct a revised accelerator mass samples a time and region important for understanding the origins spectrometry (AMS) radiocarbon chronology built on 26 new of modern human diversity. A revised chronology based on 26 dates on OES fragments. The revised chronology confirms the accelerator mass spectrometry radiocarbon dates on ostrich egg- LGM age of KNM-LH 1 and expands the maximum age of the site shells indicates an age range of 23,576–22,887 y B.P. for KNM-LH to beyond the limits of the radiocarbon method. Increased ra- 1, confirming prior attribution to the Last Glacial Maximum. Addi- diometric age is consistent with new technological analyses that demonstrate previously unrecognized MSA elements that indicate tional dates extend the maximum age for archaeological deposits at > > assemblages spanning the MSA/LSA transition from deposits un- GvJm-22 to 46,000 y B.P. ( 46 kya). These dates are consistent derlying KNM-LH 1. Morphometric analyses using a robust com- with new analyses identifying both and LSA lithic parative dataset demonstrate the variability among African Late at the site, making GvJm-22 a rare eastern African re- Pleistocene hominins, including candidate populations for out- cord of major human behavioral shifts during the Late Pleistocene. of-Africa dispersals. They indicate that KNM-LH 1 is distinct Comparative morphometric analyses of the KNM-LH 1 cranium doc- from (i) modern Africans, (ii) H. sapiens from LSA sites, ument the temporal and spatial complexity of and (iii) European Upper modern , suggesting morphological variability. Features of cranial shape distinguish that it may sample a now extinct lineage. KNM-LH 1 and other Middle and Late Pleistocene African fossils from crania of recent Africans and samples from Holocene LSA and GvJm-22: Site Setting and Excavation History European sites. GvJm-22 is a one of a number of rock shelters on the ∼16-km2 inselberg of Precambrian gneiss known as Lukenya Hill (−1.48°, Middle Stone Age | Later Stone Age | ostrich eggshell | Homo sapiens | 37.0°) that rises ∼100–200 m above the grasslands of Kenya’s modern human origins Athi-Kapiti Plain (Fig. S1). The overlooks the historic migration path of large herds of blue wildebeest (Connochaetes or Late Pleistocene African populations of modern humans, Fthe constellation of behavioral changes encapsulated in the Significance transition from the Middle Stone Age (MSA) to the Later Stone Age (LSA) ∼70–20 kya represents a series of some of the most Modern human (Homo sapiens) fossils from eastern African pronounced changes in the archaeological record before the archaeological contexts from ∼70,000–20,000 years ago are rare, adoption of domesticated animals and plants and the use of limiting our ability to understand the relationship between bi- ceramics for and storage. It is among LSA sites that ological and behavioral change during a time and place charac- the strongest parallels with ethnographic and historic foragers terized by major human demographic shifts, including dispersals. are observed. Typical archaeological signatures include lithic tech- Our chronological, archaeological, and human paleontological nologies focused on the production of (small flakes, analyses of the GvJm-22 rock shelter and Kenya National Mu- blades, and bladelets with one edge blunted or “backed”)from seums Lukenya Hill Hominid 1 partial calvaria constrain the age bipolar, single-, and opposed-platform cores; an increased use of of major behavioral changes among African foragers (the shift ground-stone tools; and implements made of wood and bone. These to Later Stone Age technologies) and demonstrate the mor- new technologies occur with the appearance of material correlates phological distinctness of Late Pleistocene African hominins from of social identity and networks of long-distance exchange, including African Holocene or Late Pleistocene Eurasian hominins, com- ostrich eggshell (OES) beads, , and nonlocal stone raw ma- plicating the history of modern human diversity. terial, as as increased dietary breadth, all consistent with larger, more dense, or more interconnected populations (1–9). Author contributions: C.A.T., I.C., and J.T.F. designed research; C.A.T., I.C., J.T.F., R.E., J.N., This same interval of ∼70–20 kya witnessed a number of hu- D.P., E.N.M., and F.S. performed research; C.A.T., I.C., J.T.F., and R.E. analyzed data; and C.A.T. and I.C. wrote the paper. man dispersals across Africa, with eastern Africa host to one or more candidate populations for the expansion of Homo The authors declare no conflict of interest. sapiens out of Africa (10–15). However, the eastern African This article is a PNAS Direct Submission. hominin fossil record for this interval is extremely sparse and Freely available online through the PNAS open access option. poorly dated, and it consists largely of isolated teeth or highly 1To whom correspondence should be addressed. Email: [email protected]. fragmentary crania and postcrania (16–18). Here, we reassess the This article contains supporting information online at www.pnas.org/lookup/suppl/doi:10. age and context of the Kenya National Museums Lukenya Hill 1073/pnas.1417909112/-/DCSupplemental.

2682–2687 | PNAS | March 3, 2015 | vol. 112 | no. 9 www.pnas.org/cgi/doi/10.1073/pnas.1417909112 taurinus), Burchell’s zebra (Equus quagga burchellii), hartebeest environment. They are dominated by the extinct alcelaphine bovid (Alcelaphus buselaphus), and impala (Aepyceros melampus) (20, 21). Damaliscus hypsodon, with arid-adapted taxa, such as Grevy’sze- R. M. Gramly discovered the site and, from 1970 to 1973, excavated bra (Equus grevyi) and oryx (Oryx beisa), found well outside their 21.75 m2 to ≤250 cm below surface (cmbs), uncovering >50,000 historic ranges (21, 27, 28). Our analysis of the identified micro- stone artifacts and >3,000 well-preserved identified faunal re- fauna (number of identified specimens = 9) revises Gramly’s mains, including a partial calvaria of H. sapiens (KNM-LH 1), the analysis (20) and indicates a semiarid savanna environment latter found at a depth of 138–140 cmbs in the initial test pit (Fig. (Gerbilliscus sp., Tachyoryctes cf. splendens,andPedetes sp., as well S2). Fossils and artifacts occur within variably CaCO3-cemented as Crocidura sp.), with the Vlei rat (Otomys sp.) and cane rat tan and gray sandy silts likely deposited by bedrock dissolution, (Thryonomys sp.) indicating wetter habitats likely at or near the slope wash, and aeolian processes; exfoliated slabs from the in- spring seeps that are still found near GvJm-22. selberg increase with depth (20, 22, 23). Artifact typology, the Gramly and Rightmire (29) report two radiocarbon dates on degree of fossilization of the fauna, and conventional radiocarbon bones bracketing KNM-LH 1 (from samples at 135–140 cmbs dates on charcoal and bone collagen from the 1970s (Table 1) and 140–145 cmbs), with fluorine and nitrogen values of KNM- indicate that four Pastoral and LSA archaeological LH 1 and the dated bones indicating a shared depositional his- assemblages from 0–110 cmbs date to the Holocene, separated by tory and an age between 17,670 ± 800 14C y B.P. and 17,700 ± an unconformity from two Pleistocene assemblages, termed oc- 760 14C y B.P. However, the ages of KNM-LH 1 and the GvJm- currence E (120–150 cmbs) and occurrence F (170–205 cmbs) (20, 22 sequence are usually considered unreliable because of the 25). The Pleistocene occurrence E and F lithic assemblages are stratigraphically inverted ages of samples underlying KNM-LH 1 characterized by the production of small (≤3 cm) flakes, blades, (Table 1) and skepticism about older radiocarbon dates on bone, and bladelets retouched primarily into scrapers and backed pieces, particularly from the tropics (30–32). Early collagen pretreat- and have been attributed to the LSA (20, 22, 23, 26). The faunal ment techniques yielded “collagen” where, in fact, none was assemblages from both occurrences indicate a dry, grassy, savanna present (as shown by later reanalysis with improved techniques)

Table 1. Radiocarbon dates from GvJm-22 † Date, 14C y B.P. Calibrated age,* y B.P. Material Trench Occurrence Depth, cmbs Laboratory no.

1,330 ± 100 1,234 ± 104 Charcoal (twig) Balk A 45–50 N-1076 1,510 ± 50 1,413 ± 61 Bone collagen Balk A 45–50 UCLA-1709D 2,250 ± 50 2,245 ± 62 Bone collagen Balk C 70–75 UCLA-1709C 17,670 ± 800 21,461 ± 977 Bone collagen Balk E 135–140 UCLA-1709A 17,700 ± 760 21,483 ± 926 Bone collagen Balk E 140–145 UCLA-1709B 3,526 ± 27 3,794 ± 49 OES A D/E 110–120 UBA-24889 13,008 ± 68 15,560 ± 137 OES A E 120–130 UBA-24884 12,752 ± 59 15,193 ± 100 OES A E 120–130 UBA-24885 12,854 ± 63 15,350 ± 122 OES A E 120–130 UBA-24886 12,953 ± 60 15,486 ± 124 OES A E 140–150 UBA-24881 15,391 ± 95 18,656 ± 101 OES A E 140–150 UBA-24883 18,056 ± 96 21,884 ± 152 OES A E 140–150 UBA-24882 ‡ 13,045 ± 67 15,615 ± 139 OES A E/F 160–170 UBA-24891 21,803 ± 142 26,048 ± 142 OES A F 190-breccia UBA-24890 4,116 ± 32 4,662 ± 87 OES B E 120–130 UBA-24877 3,2967 ± 518 37,213 ± 681 OES B E 130–140 UBA-24880 4,250 ± 32 4,814 ± 52 OES B E/F 150–160 UBA-24879 4,049 ± 34 4,533 ± 84 OES B E/F 160–170 UBA-24887 27,915 ± 287 31,824 ± 385 OES B F 170–180 UBA-24878 5,960 ± 35 6,793 ± 50 OES C D 105–110 UBA-24888 15,320 ± 450 18,609 ± 520 Bone collagen C + balk F 180–185 GX-3699 13,730 ± 430 16,628 ± 603 Bone collagen C + balk F 190–195 GX-3698 9,910 ± 300 11,487 ± 482 Bone collagen C + balk F 190–195 HEL-535 4,448 ± 32 5,103 ± 109 OES Test pit E 120–130 UBA-23927 12,884 ± 59 15,393 ± 120 OES Test pit E 130–140 UBA-23928 19,666 ± 120 23,691 ± 167 OES Test pit E 130–140 UBA-23929 19,594 ± 119 23,607 ± 176 OES Test pit E 130–140 UBA-23930 19,794 ± 127 23,826 ± 167 OES Test pit E 130–140 UBA-23931 19,568 ± 119 23,574 ± 179 OES Test pit E 140–150 UBA-23933 28,041 ± 320§ 31,985 ± 431 OES Test pit E 140–150 UBA-23934 28,230 ± 380§ 32,210 ± 502 OES Test pit E 140–150 AA102890 33,308 ± 612{ 37,571 ± 768 OES Test pit E 140–150 UBA-23935 ± { ± –

33,410 570 37,665 729 OES Test pit E 140 150 AA102889 ANTHROPOLOGY 46,710 ± 3,852 N.A. OES Test pit E 140–150 UBA-23932

Radiocarbon dates (14C y B.P.) are reported as mean and 1 standard deviation. Laboratory codes: AA, University of Arizona; GX, Geochron Laboratories; HEL, National Museum of (Helsinki); N, Nishina Memorial; UBA, Queen’s University Belfast; UCLA, University of California Los Angeles. N.A., not applicable. *Ages were calibrated using IntCal13 (24) and reported at the 95% confidence interval. † OES dates from GvJm-22 are reported here for the first time; others are from Gramly (20, 25). ‡ Gray-black color indicates that the sample is burnt. §Multiple analyses of the same specimen by different laboratories. {Multiple analyses of the same specimen by different laboratories.

Tryon et al. PNAS | March 3, 2015 | vol. 112 | no. 9 | 2683 or failed to remove contaminating modern carbon fully, strongly the KNM-LH 1 cranium. This age is very close to the ages of biasing dates older than ∼15 kya (33). The dates bracketing 21,461 ± 977 and 21,483 ± 926 cal y B.P. for the conventional KNM-LH 1 used the Longin technique of collagen extraction (34), radiocarbon dates on bone fragments from 135–140 cmbs and a modified form of which is still used today, whereas unknown 140–145 cmbs originally reported by Gramly and Rightmire (29). methods were used to generate the lower, stratigraphically in- verted dates. MSA Archaeology at GvJm-22 Our reanalysis of the archaeological deposits below occurrence E A Revised Radiocarbon Chronology and the KNM-LH 1 partial calvaria suggests assemblages that A revised chronology for the GvJm-22 sequence is provided by span the MSA/LSA transition at GvJm-22. The sequence is char- 26 AMS radiocarbon dates on the carbonate fraction of un- acterized by the progressive shift in emphasis from flake production modified OES fragments. OES is reliably dated by the AMS ra- by Levallois methods to and bladelet manufacture from single diocarbon method, and dating OES is appropriate for the 1970– and opposed platform cores and an increased use of backed pieces 1973 collections housed at the National Museums of Kenya (NMK) (microliths) at the expense of points and scrapers. The shift to LSA because the material is resistant to diagenetic contamination and is seen in other artifacts as well. Large grindstones that does not degrade with storage time like charcoal (2, 35–38). Sample show repeated, heavy use for grinding appear only in occurrence E, preparation was similar to the procedures of Janz et al. (37) and with the largest being fragmented but 154 mm in maximum di- incorporated a strong preanalysis acid etch designed to remove mension and weighing 1.1 kg, with 52-mm-wide and 73-mm-wide outer surfaces and any associated contamination. To measure depressions on either side, producing an hourglass-shaped profile reliability further, two OES samples were split and sent to dif- along the broken edge similar to ethnographic examples used for ferent laboratories (University of Arizona processing ochre, cartilage, seeds, fibrous tubers, or vegetables (40, and Queen’s University Belfast), yielding statistically indistinguish- 41). Ochre is present in small amounts (5.2 ± 4.4 g per 10-cm in- able results for both samples. All dates are calibrated using the terval) throughout the sequence (from 30–225 cmbs), with a single IntCal13 calibration curve (24) and are summarized in Table 1. piece with wear facets at 185–190 cmbs. Finished and unfinished The new dates confirm an unconformity of ∼9,000 y at ∼120 OES beads (n = 7 from 120–160 cmbs) appear at 150–160 cmbs, cmbs separating the Holocene from the underlying Pleistocene at and immediately below the occurrence E interface. deposits. This unconformity may be due to erosion or reduced We examined the cores found in strata below occurrence E, sedimentation with increased vegetation and greater moisture including occurrence F as defined by Gramly (20) at 170–205 availability during the Holocene. OES fragments from the cmbs, as well as an interval between occurrences E and F (160– Pleistocene strata were dated from across the site (Gramly’s test 170 cmbs) and the lowermost deposits (210–250 cmbs), which pit; trenches A, B, and C; and remnant sediment balks; Fig. S2). Gramly referred to as occurrence G in his unpublished notes These results (i) expand the age range of occurrence E to ∼15 (Table S1). Data from neither 160–170 cmbs nor occurrence G to >46 kya; (ii) confirm Gramly’s hypothesis (20) of postde- have been reported previously. Levallois cores form 10–33% of positional mixing in trench B; and (iii)suggestthatGramly’s5- the studied sample (n = 207) from below occurrence E. Our to 10-cm-thick horizontal excavation levels cross-cut sedimentary understanding of the Levallois concept follows the definition of deposits that dip ∼2° westward, with an age offset of ∼7,000 to Boëda (42). Most of the GvJm-22 Levallois cores were originally 10,000 y between layers at equivalent depths in trench A and the identified by Gramly (20) as “disk-like” cores, and many are adjacent test pit (Table 1 and Fig. S2). consistent with what Yellen et al. (43) term “micro-Levallois” and We developed a Bayesian age model (39) for the KNM-LH 1 “Aduma” cores. The GvJm-22 Levallois cores show primarily cranium (found at 138–140 cmbs) using only dates from the test centripetal and bidirectional patterns of flake removal from the pit (n = 11) in which it was originally found. OxCal 4.2 software upper surface and near-exclusive use of recurrent methods of flake was used to combine dates for samples with replicate analyses production immediately before discard (Fig. 1); éclats débordants from multiple laboratories and to develop a bounded sequence (n = 3) further indicate on-site knapping and Levallois core edge of dates and phases (groups of dates from the same 10-cm-thick maintenance during reduction. Levallois flakes are, on average, excavation level). The KNM-LH 1 hominin lies near the boundary small (n = 23, 29.8 ± 6.2 mm) but significantly larger (t = 2.72, of two excavated levels (130–140 cmbs and 140–150 cmbs). Both df = 43, P = 0.009) than Levallois cores (n = 54, 25.4 ± 6.8 mm), levels have dated samples that are substantially younger (∼9,000– although the scale of the difference (∼5 mm) is quite small in 15,000 y) than the other samples from the same 10-cm interval practical terms. The slight size difference between Levallois cores (UBA-23928 and UBA-23933; Table 1). We modeled these and Levallois flakes at discard suggests short operational chains, samples as younger than the remaining samples within their re- particularly for cores of chert (n = 31, 25.1 ± 3.9 mm) and ob- spective 10-cm-thick excavation level, assuming that their younger sidian (n = 12, 20.41 ± 3.9 mm), the most common raw materials age resulted from increased stratigraphic height and proximity to among Levallois cores. Both chert and obsidian are locally the overlying 10-cm-thick excavation unit. The modeled results available only as small (∼5 cm) clasts and lapilli (23). Quartz indicate an age range of 23,576–22,887 calibrated (cal) y B.P. for Levallois core size (n = 10, 32.5 ± 10.8 mm) is significantly larger

Fig. 1. Artifacts from strata beneath occurrence E of the 1970–1973 excavations at GvJm-22. Levallois cores (A–G), Levallois flake (H), bladelet cores (I and J), bipolar core (K), “fan” (L), backed piece (M, ), and bifacial point (N). Artifacts E, F, J, and N are from the occurrence E/F interface (160– 170 cmbs), artifact A is from occurrence G, and the remainder are from occurrence F. All are made chert, except for G (quartz) and K and N (obsidian).

2684 | www.pnas.org/cgi/doi/10.1073/pnas.1417909112 Tryon et al. (F = 12.05, df = 2, P < 0.001) and more variable, likely reflecting the immediate availability of this material from veins within the Lukenya Hill inselberg. Levallois cores and flakes co-occur with a number of other approaches to the production of small blanks (Fig. 1). Non-Levallois cores are also small (n = 154, 26.5 ± 8.9 mm) and do not differ significantly in size from the Levallois cores (t = 1.01, P = 0.31) or in the relative abundance of the use of chert, quartz, or ob- sidian (χ2 = 4.068, df = 2, P = 0.13). These other flake, blade, and bladelet production methods form the technological sub- strate throughout the Pleistocene sequence at GvJm-22, and the difference between occurrence E and strata beneath it lies in the increased frequency of platform cores for blade and bladelet production, increased use of the bipolar method, and the aban- donment of Levallois and other centripetal approaches over time, differences that are significant (χ2 = 232.45, df = 6, P < 0.001). A similar gradation is found among the retouched tools, with differences between occurrence E and the strata below being primarily a difference of quantity rather than kind, with backed pieces and scrapers (Fig. 1) present at ≤195 cmbs. Based on counts from Gramly (20) and our metric data, backed pieces form the dominant retouched tool in occurrence E (measured sample: n = 128, 21.3 ± 4.7 mm) but do not differ in size from those backed pieces in occurrence F (n = 15, 23.8 ± 7 mm; t = 1.35, P = 0.20), where they are more rare. Scrapers are the most abundant retouched tool in occurrence F, and the difference between the archaeological occurrences of backed pieces and scrapers is significant (χ2 = 158.06, df = 1, P < 0.001). Points, although rare, are one of the few retouched tools restricted to strata below occurrence E. The sample includes a small (length = 42.3 mm), thin (6.6 mm) bifacial point of obsidian (Fig. 1) at 160–170 cmbs in trench A, with a second quartz biface tip from 180–185 cmbs from trench C + balk. The combination of artifact types suggests that strata below occurrence E span the MSA/LSA transition, although whether this combination reflects “transi- Fig. 2. Three-dimensional model of the frontal bone in anterior view, show- tional” assemblages or a postdepositional admixture of MSA and ing the healed frontal trauma and the inner structure of the frontal squama. LSA artifact-bearing strata is uncertain (SI Text). The zone of trauma is demarcated by the white rectangle. (A and B) Two The age of the deposits below occurrence E is constrained only sagittal sections intersect the healed trauma, as shown by white . (C) by the >46,710 ± 3,852 14C y B.P. date from 140 to 150 cmbs in Coronal section is given at the level of the postmortem perforation of the the test pit and the 25,868 ± 186 cal y B.P. date from 190 cmbs to right side of the frontal squama (left ). The right arrow outlines the bedrock from trench A (Table 1). This age range of ∼26 to >46 pathological inner structure changes of the diploe and outer table. kya for the lower strata at GvJm-22 is consistent with data from similarly aged industries from sites near GvJm-22, including the well represented in our Late Pleistocene North African (LPA) Nasera and Lemuta Industries at Nasera and perhaps Mumba = rock shelters in and the Sakutiek at Enkapune sample (Table S2) where preserved (25%, n 67). ya Muto in Kenya (SI Text and Fig. S1). All contain MSA/LSA The frontal bone exhibits two areas of antemortem and post- sequences and transitional artifact industries with small Levallois mortem damage (Fig. 2). One centimeter above the glabella, cores and flakes, retouched points, an abundance of small scrapers, a healed linear transverse trauma (10 mm in length) is present. ∼ backed pieces ∼24 mm in length, grindstones, ochre, and OES The surrounding area (over a radius of 15 mm) is depressed and beads (2, 4, 44–46). Pleistocene occurrences from GvJm-22, slightly remodeled, affecting the curvature of the frontal bone at Nasera, Mumba, and Enkapune ya Muto contain obsidian from this level. Sagittal CT sections of this portion reveal a healed sources near Lake Naivasha at distances 100–300 km from the complete fracture of the frontal bone without displacement, cor- source (47–49). Similar archaeological sequences and shared raw responding to the outer linear trauma (compare Fig. 2 A and B). material sources suggest population interconnections and extensive The second area of damage is located 6 cm from the right orbital Late Pleistocene social networks throughout southern Kenya and margin, along the broken and eroded right edge of the frontal northern Tanzania. squama. A circular perforation about 6 mm in diameter and 5 mm in depth is visible. It crosses the outer table and half of the diploe KNM-LH 1 in Context (compare Fig. 2C). As suggested by A. Walker (51), it might be KNM-LH 1 occurs within occurrence E at 138–140 cmbs, dates relatedtoatoothmarkfromacarnivore. to ∼23 kya, and is the only hominin cranium found with an LGM/ The cranium is thick [10.2-mm thickness at the (left) parietal LSA assemblage from eastern Africa. KNM-LH 1 thus plays an eminence (TPE), 10.0-mm thickness for the frontal squama]. CT important role in our understanding of the morphology of these images show a relatively thick vault compared with modern humans, ANTHROPOLOGY populations. Our formal description of KNM-LH 1 extends the with wide diploe and proportionate tables, with the exception of description of Gramly and Rightmire (29), augmented by com- the right part of the frontal squama. In this area, the outer table puted tomography (CT). KNM-LH 1 preserves two-thirds of the seems to have thinned, the diploe is wider, and slight porosities are frontal bone and most of the left parietal bone (Fig. S3). The visible on the external surface of the bone. Nonpathological high superciliary ridges are very pronounced and joined medially by cranial thickness is a common characteristic of Late Pleistocene a projecting glabella that is clearly separated from the rest of the H. sapiens that may reflect elevated activity levels compared with frontal bone. A sulcus separates the ridges from the robust lat- recent populations (52). However, the morphology of the right eral trigone [supraorbital torus grade 4 (50)]. This morphology is side of the frontal bone, with its expanded and slightly modified rare among recent modern humans (RMHs) from Africa but is diploe, suggests some caution in accepting this interpretation for

Tryon et al. PNAS | March 3, 2015 | vol. 112 | no. 9 | 2685 5 of KNM-LH 1 were made on the 3D reconstruction of the calvaria withAvizo7andaregiveninTable S3 with the comparative ranges of variation of the early modern human (EMH), Late Pleistocene 4 modern human (LPMH), and RMH samples. KNM-LH 1 differs from the LPMH sample in relation to its vault thickness and the morphology of the frontal bone [frontotransverse index (IFT)]. 3 KNM-LH 1 possesses a significantly less divergent frontal squama (IFT = 92.79) than the LPMH group, lying in the EMH range of variation (IFT = 87.9 ± 3.5, n = 10; Table S3). Compared with 2 KNM-LH 1 the RMH group, KNM-LH 1 falls outside the upper limit of the RMH variation regarding its anterior frontal breadths (bifrontal Principal component 1 (42.2%) breadth and interorbital breadth) and the frontal length [frontal 1 chord (FRC)]. Focusing on calvarial shape, we performed a OA2 principal component analysis on five size-adjusted measurements

0 (55, 56). The results are shown in Fig. 3. In terms of the shape of the calvaria, KNM-LH 1 is distinct from recent Africans and Holocene LSA samples. It is also distinct from most of the com- -1 parably aged European Upper Paleolithic (EUP) samples, except, interestingly, Pes¸tera cu Oase, the oldest EUP cranium. Instead, KNM-LH 1 and several Late Pleistocene African crania (e.g., -2 Nazlet Khater 2, Iwo Eleru, the LPA group) fall outside of the RMH variation along the first axis. The separation along the first axis is related to the combination of large anterior frontal breadths -3 in relation to the maximal frontal breadth and the FRC. The separation of KNM-LH 1 along the third axis is related to its relatively great frontal length. DKB -4 Unlike face or basicranial shape, neurocranial shape is gen- FMB KNM-LH 1 LPA erally considered a useful measure of population history (57, 58). NK2 LPS Our results emphasize distinct temporal and spatial differences HOF LSA IWO NEOAF -5 in cranial shape among modern humans (H. sapiens). KNM-LH 1 MPH RMH and other Pleistocene African specimens, all of which are po- PAC FRC EMH NEAND XFB EUP tentially sampling candidate populations for dispersals across -6 and out of Africa during the Late Pleistocene (12–15, 50, 59), 4 3 2 1 0 -1 -2 -3 -4 Principal component 3 (17.5%) differ substantially not only from recent Africans but also from individuals drawn from Holocene LSA archaeological sites. Fig. 3. Scatter plot of the first and third principal components of the principal KNM-LH 1 and other Pleistocene African specimens (found component analysis computed on size-adjusted cranial dimensions. The ellipse with MSA and LSA artifacts) are also distinct from most EUP represents 95% of the RMH variation. (Inset) Correlation circle in the lower left individuals. The limited similarity between the Late Pleistocene corner of the projection shows the position of the craniometric variables in African and European samples indicates that KNM-LH 1 sam- relation to the two plotted principal components. DKB, interorbital breadth; ples elements of modern human variability not found in current FMB, bifrontal breadth; FRC, frontal chord; PAC, parietal chord; XFB, maximum lineages and attests to the still poorly understood interactions frontal breadth. Metric data for HOF, IWO, MPH, and NEAND come from between demography, dispersals, behavior, and climate that have published measurements (61–66). Data for NK2, La Chapelle-aux-Saints 1, La shaped human cranial variation over the past >70,000 y (60). Ferrassie 1, and Spy 1 were taken on the originals. NK2, Nazlet Khater 2; HOF, Hofmeyr; IWO, Iwo Eleru; OA2, Pes¸tera cu Oase 2; MPH, Middle Pleistocene Conclusions Homo fossils from Africa; EMH, early modern human fossils from Africa and Renewed chronometric, lithic technological, and morphological Southwest Asia; EUP, European Upper Paleolithic modern human fossils; LPA, Late Pleistocene fossils or samples from ; LPS, Late Pleistocene analyses of the hominin partial cranium (KNM-LH 1) from site fossils or samples from Southwest Asia; LSA, fossils or samples GvJm-22 at Lukenya Hill demonstrate (i) a Last Glacial Maxi- from Sub-Saharan Africa; NEOAF, Neolithic fossils or samples from North mum age for KNM-LH 1, (ii) an extended age range for > ± 14 Africa; RMH, recent modern human samples; NEAND, Neandertals. archaeological deposits at the site to 46,710 3,852 C y B.P., (iii) the presence of lithic artifacts typically associated with MSA or transitional MSA/LSA industries in deposits underlying KNM-LH 1 (compare Fig. 2C). Although located in a small area KNM-LH 1, and (iv) the substantial and still poorly documented of the calvaria and associated with no other criteria (e.g., absence variability among aspects of cranial shape among Late Pleisto- of cribra orbitalia, presence of frontal sinus), this thickening, to- cene H. sapiens across Africa and Eurasia. Although the record gether with the external porosity, could be related to healed ex- of MSA and LSA sites and hominin remains in eastern Africa is sparse and discovery of additional sites through further fieldwork pression of light porotic hyperostosis (53). The alternate hypothesis – for the localized porotic cranial changes on the right side of the is badly needed (46), our reexamination of the 1970 1973 frontal bone is that they are related to periosteal reactions resulting collections from GvJm-22 at Lukenya Hill show the value of from trauma infection (54). In either case, the TPE does not seem to a critical analysis of museum collections for understanding the be associated with bone remodeling. behavior and biology of Late Pleistocene H. sapiens. In their formal description of the specimen, Gramly and Materials and Methods Rightmire (29) emphasized the low and receding morphology of the frontal region as being distinct from the morphology All archaeological materials (including OES fragments) were studied and seen in modern African populations. Here, we expand the com- sampled from collections curated at the NMK in Nairobi. Comparisons of artifact dimensions used an unequal variance t test, unless otherwise spec- parison of the KNM-LH 1 partial cranium to a geographically and ified. Analyses of the KNM-LH 1 hominin cranial fragments (also stored at temporally diverse sample through univariate and multivariate the NMK) were augmented by CT scans made with a Siemens Sensation 16 analyses. The comparative sample groups are listed in Table S2 scanner. The three fragments were oriented perpendicular to the scan plane and include Middle Pleistocene to Holocene individuals from to give optimal visualization of the tables and diploe, using a slice spacing of Europe, Southwest Asia, and across Africa. The measurements 0.3 mm and a pixel size of 0.23 mm.

2686 | www.pnas.org/cgi/doi/10.1073/pnas.1417909112 Tryon et al. ACKNOWLEDGMENTS. We thank Drs. Greg Hodgins, Pierre Guyomarc’h, Laténium (M. Honegger), Muséum national d’Histoire naturelle (Ph. Mennecier), Alex Tokovinine, Christine Lane, Warren Sharp, and Linda Reynard for advice NMK, National Museum of Natural History (Smithsonian Institution; D. Hunt), and assistance, and particularly R. M. Gramly for his continued support Royal Belgian Institute of Natural Sciences (P. Semal), and Staatssammlung throughout this project. Sheila Nightingale provided the excellent artifact für Anthropologie und Paläoanatomie (G. McGlynn). This research was con- illustrations. We thank the following institutions and persons for providing ducted under Research Permit NCST/5/002/R/576 issued to C.A.T. by the Gov- permission to study the comparative material and the CT scan data: The ernment of the Republic of Kenya while an affiliate of the NMK and was (N. Spencer, D. Antoine), University of Colorado Boulder (D. Van supported by funds from the American School for Prehistoric Research and Gerven), Institut de Paléontologie Humaine (H. de Lumley, D. Grimaud-Hervé), Harvard University.

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Tryon et al. PNAS | March 3, 2015 | vol. 112 | no. 9 | 2687 Supporting Information

Tryon et al. 10.1073/pnas.1417909112 SI Text Therefore, we refrain from a formal archaeological taxonomic attribution for the occurrence F and occurrence G material from Archaeological Taxonomic Status of the Occurrence F and Gramly’s 1971–1973 excavation but, instead, emphasize the pres- Occurrence G Assemblages ence of MSA technology (Levallois flakes and cores as well as We reanalyzed archaeological material from the 1971–1973 Gramly bifacial points), where none was previously recognized. Thus, the excavations of GvJm-22 stored at the NMK. We focused on cores GvJm-22 artifact assemblages span the MSA/LSA transition but because, from a technological standpoint, they are among the do not necessarily represent one or more transitional industries. most informative aspects of any lithic assemblage. The results The situation at GvJm-22 is remarkably similar to the situation at show that Levallois cores, one of the defining features of MSA the Magosi rock shelter in , where the co-occurrence technology (1–3), comprise 10–33% of the archaeological material of MSA and LSA artifacts (Levallois technology, retouched attributed by Gramly to occurrences F and G. The Levallois cores points, and backed microliths) from excavations in 1926 de- (and flakes) from occurrence F at GvJm-22 co-occur in strata with fined the Magosian Industrial Complex, once considered to be other typically MSA artifact forms (i.e., bifacially flaked points), the clearest example of an industry of the “Second Intermediate as well as with those artifact forms considered diagnostic of LSA Period” between the MSA and LSA (9–11). However, re- sites, particularly microlithic backed pieces and small “fan” scra- excavation of Magosi in 1963 demonstrated that this combina- pers. Our examination of the formal tools from strata beneath tion of MSA and LSA artifacts was artificial and resulted from occurrence E simply confirmed previous typological attributions the removal of sediment in 61-cm-thick horizontal increments (4, 5). Although we collected metric data on artifact dimensions, across steeply dipping (∼45°) strata, a recognition that contributed we did not attempt a reanalysis of the abundance of different to the abandonment of both the Magosian Industrial Complex and artifact types because it was clear that a number of formal tools Second Intermediate Period concepts (11, 12). Because of have been separated from the remainder of the GvJm-22 col- similar stratigraphic problems at GvJm-22, a more precise lection at the NMK and could not be located. The sample size of definition of the assemblages below occurrence E and the na- retouched tools from previously unreported occurrence G is very ture of the MSA/LSA transition at the site are problems that can low (n = 2) and does not include retouched points, microlithic only be addressed with renewed excavations at the site using backed pieces, or fan scrapers. finer scale artifact provenience recording and microstratigraphic The co-occurrence of MSA and LSA lithic artifacts in occur- approaches. rence F might indicate that this archaeological aggregate rep- “ ” resents a transitional assemblage, formed by a gradual process Hominin Comparative Data of shifts in over time, as appears to be the case Metric comparisons of the KNM-LH 1 partial calvaria, sum- at sites in Tanzania, such as the Mumba and Nasera rock shelters that sample the same time interval (6–8). However, the strati- marized in Table S3, are based on the following measurements: graphic resolution of the material excavated in 1970–1973 is too maximum frontal breadth (XFB), bifrontal breadth, interorbital coarse to understand the nature of technological change between breadth, FRC, parietal chord (PAC), and frontal angle following MSA and LSA technologies at GvJm-22. The alternative hy- Howells (13); minimal frontal breadth [BFT; M9 (14)]; frontal = pothesis of formerly discrete assemblages with exclusively MSA arc (FAR; M26); parietal arc (PAR; M27); TPE (15); IFT (I.12 = and LSA artifacts that have been mixed postdepositionally or BFT/XFB); frontal curvature index (I.22 FRC/FAR); and pa- = during excavation cannot be rejected. Our radiocarbon dating rietal curvature index (I.24 PAC/PAR). Published measure- program focused on providing age sequences from multiple areas ments for RMHs are taken from Tweisselmann (15) and Howells of the 1970–1973 excavation area. When these results are inte- (16). The published measurements for EMHs and LPMHs used grated with site stratigraphy, they demonstrate that Gramly’sex- in Table S3 are taken from other publications (16–49). Mea- cavation of arbitrarily 5- to 10-cm-thick horizontal spits intersected surements in Table S3 of El Barga, El Fakhuri, Ishango, Jebel natural and cultural deposits that dip ∼2°, resulting in the ad- Sahaba, Lothagam, Olduvai Hominid 1, , and mixture of formerly discrete strata, a problem recognized by were taken from the originals. Measurements in Table S3 of Wadi Gramly (4) in his definitions of occurrences E and F, but one Kubbaniya were taken from a cast. The Ishango material was too that achieves greater significance with the identification of both fragmentary to be included in the principal component analysis. MSA and LSA technological elements in occurrence F. Shown in Table S3 are sample means, SD, and sample size.

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Tryon et al. www.pnas.org/cgi/content/short/1417909112 2of7 Lake Turkana Magosi

100 km

Mt. Elgon Mt. Kenya

Lake Enkapune Lake Naivasha Victoria ya Muto

Lukenya Hill

Nasera Mt. Kilimanjaro N

Elevation 8752 m Mumba

-407 m

Fig. S1. Topographic sketch map of eastern Africa showing archaeological sites and major geographic features mentioned in this study.

Tryon et al. www.pnas.org/cgi/content/short/1417909112 3of7 Fig. S2. R. M. Gramly’s 1970–1973 excavations at GvJm-22: the Lukenya Hill inselberg (A); the view of the Athi-Kapiti Plains from Lukenya Hill (B); GvJm-22 at the beginning of the excavation (C); plan view of the GvJm-22 excavations, showing excavations units A–E, the test pit (T.P.), and the sediment balk, with circles denoting units sampled in this study (D); and profile view of the south wall of the test pit and trench A, with stippled areas showing the location of archae- ological strata occurrence E and occurrence F (E). Modified with permission from ref. 4, © 1976 Royal Anthropological Institute of Great Britain and Ireland.

Fig. S3. Virtual reconstruction of the KNM-LH 1 calvaria (clockwise from upper left) in anterior, left lateral, superior, and inferior views.

Tryon et al. www.pnas.org/cgi/content/short/1417909112 4of7 Table S1. Frequency of cores from GvJm-22 Occurrence E, 160–170 Occurrence, F Occurrence G, Core type 120–150 cmbs* cmbs 170–205 cmbs 210–225 cmbs

Levallois preferential 0 2 3 2 Levallois recurrent 1 13 32 0 Levallois (method indeterminate) 0 0 2 0 Single platform 198 9 34 6 Double platform (opposed)† 63 0 (2) 2 (11) 3 (1) Multiple platform 17 0 8 2 ‡ Bipolar 145 9171 Centripetal 0 5 25 1 Irregular/other 0 4 8 1 Core fragment 36 1 1 2 Total 349 45 143 19

Cores are defined as pieces with three or more removals; core types follow Mehlman (1), Boëda (2), and Kuhn (3). *Data are from Barut (4). †Opposed platform cores are shown in parentheses. ‡Includes split pebbles (n = 3).

1. Mehlman MJ (1989) Late Quaternary archaeological sequences in northern Tanzania. PhD dissertation (University of Illinois at Urbana–Champaign, Champaign, IL). 2. Boëda É (1994) Le concept Levallois: Variabilité des méthodes (Centre National de la Recherche Scientifique Éditions, Paris). French. 3. Kuhn SL (1995) Lithic Technology: An Ecological Perspective (Princeton Univ Press, Princeton). 4. Barut S (1997) Later Stone Age lithic raw material use at Lukenya Hill, Kenya. PhD dissertation (University of Illinois at Urbana–Champaign, Champaign, IL).

Tryon et al. www.pnas.org/cgi/content/short/1417909112 5of7 Table S2. Hominin comparative samples Groups Specimens Country

KNM-LH 1 Lukenya Hill Kenya NK2 Nazlet Khater 2 HOF Hofmeyr South Africa IWO Iwo Eleru MPH Middle Pleistocene Homo fossils from Africa Florisbad South Africa Kabwe 1 EMH Early modern human fossils from Africa and Southwest Asia Border Cave 1 South Africa Dar-es-Soltan H5 Herto 1 (BOU-VP-16/1) Ethiopia 1 and 2 Morocco Ngaloba LH18 Tanzania Omo Kibish 2 Ethiopia Qafzeh 3, 6, and 9 Israel Skhul IV, V, VI, and IX Israel LPMH: Late Pleistocene modern humans EUP European Upper Paleolithic modern human fossils France Barma Grande Brno 2 Czech Republic Cioclovina Romania Cro-Magnon 1, 2, and 3 France Dolní Vestonice 3, 13, 14, 15, and 16 Czech Republic Grotte des Enfants 4, 5, and 6 Italy Khvalynsk Russia La Crouzade V France Mladecˇ 1, 2, 5, and 6 Czech Republic Pavlov 1 Czech Republic Pes¸tera Muierii 1 Romania Pes¸tera cu Oase 2 Romania Podkumok Russia Predmostí 3, 4, 9, and 10 Czech Republic Satanay Russia Skhodnya Russia Sunghir 1 Russia LPA Late Pleistocene fossils or samples from North Africa Afalou-Bou-Rhummel (n = 37) Algeria El Barga Mesolithic (n = 26) Esna (El Fakhuri) Egypt Jebel Sahaba (n = 22) Sudan Taforalt (n = 30) Morocco Wadi Halfa (n = 15) Sudan Wadi Kubbaniya Egypt LPS Late Pleistocene fossils or samples from Southwest Asia Ein Gev 1 Israel El Wad (n = 16) Israel Erq El-Ahmar (n = 2) Israel Eynan Mallaha (n = 19) Israel Hayonim (n = 7) Israel Kebara (n = 5) Israel Nahal Ein Gev Israel Nahal Oren (n = 15) Israel Ohalo H2 Israel Rakefet Israel Shukbah (n = 5) Israel Wadi Mataha F-81 Israel

Tryon et al. www.pnas.org/cgi/content/short/1417909112 6of7 Table S2. Cont. Groups Specimens Country

LSA Late Stone Age fossils or samples from Sub-Saharan Africa Elmentieta A, B, D, E, F, and F1 Kenya Fish Hoek South Africa Gamble’s Cave II (4 and 5) Kenya Ishango NFPr (n = 3) Democratic Republic of Congo Lothagam 2, 4b, 7, and 10 Kenya Makalia 1 Kenya Matjes River Kenya Mumba X Tanzania Naivasha Kenya Nakuru IX Kenya Olduvai Hominid 1 Tanzania Tuinplaas 1 (Springbok Flats) South Africa Willey’s Kopje 1, 2, and 3 Kenya NEOAF Neolithic fossils or samples from North Africa Ain Dokkara Algeria Asselar Mali Columnata (n = 13) Algeria El Barga Neolithic (n = 25) Sudan Gambetta 1 Algeria Hassi-el-Habiod (n = 21) Mali Mechta el Arbi 1–3 and 5 Algeria Medjez II (n = 4) Algeria RMH Recent modern human samples Bushman (n = 82) South Africa Zulu (n = 101) South Africa Dogon (n = 99) Mali Teita (n = 83) Kenya Egypt (n = 111) Egypt NEAND Neandertal fossils Israel La Chapelle-aux-Saints 1 France 1 France Shanidar 1 Spy 1 Belgium Steinheim Germany

Table S3. Comparative craniometric descriptive statistics BFT XFB FMB DKB FRC PAC FRA FAR PAR TPE IFT ICF ICP

KNM-LH 1 103.00 111.00 107.40 30.20 120.00 114.00 128.41 140.00 125.00 10.20 92.79 85.71 91.20 EMH M 105.25 120.30 116.79 27.33 116.11 118.10 132.70 129.89 128.75 8.83 87.88 87.89 92.94 S 5.73 5.12 8.13 2.96 5.52 9.48 4.86 6.51 8.48 2.21 3.46 2.53 2.39 N 13 10 7 7 12 10 9 9 8 6 10 10 8 LPMH M 97.61 118.15 104.96 26.06 112.16 117.29 128.47 129.19 130.41 7.21 82.75 87.50 90.00 S 5.99 6.48 6.63 2.80 6.20 6.50 6.76 7.49 8.07 1.58 4.63 2.67 2.99 N 248 221 142 113 224 216 167 266 263 161 196 241 202 RMH M 111.72 96.69 22.34 108.65 111.31 126.66 7.70 S 5.69 4.46 2.54 5.24 6.40 4.36 0.98 N 476 476 476 476 476 476 64 Adjusted Z-scores* KNM-LH 1 vs. EMH −0.1740 −0.7653 −0.4411 0.3707 0.3075 −0.1823 −0.3634 0.6391 −0.1763 0.2231 0.5981 −0.3630 −0.2905 KNM-LH 1 vs. LPMH 0.4559 −0.5580 0.1854 0.7433 0.6398 −0.2560 −0.0045 0.7315 −0.3395 0.9360 1.0958 −0.3396 0.2031 KNM-LH 1 vs. RMH −0.0644 1.2218 1.5710 1.1008 0.2138 0.2043 1.2819

BFT, minimal frontal breadth; XFB, maximum frontal breadth; FMB, bifrontal breadth; DKB, interorbital breadth; FRC, frontal chord; PAC, parietal chord; FRA, frontal angle; FAR, frontal arc; PAR, parietal arc; TPE, vault thickness at parietal eminence; IFT, frontotransverse index; ICF, frontal curvature index; ICP, parietal curvature index; M, mean; S, standard deviation; N, number. *Adjusted Z-scores of KNM-LH 1 relative to both the Late Pleistocene modern human and recent modern human samples; the values in bold and italic (<−1or >1) are outside 95% of the comparative sample range of variation (49).

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