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Journal of Human Evolution 63 (2012) 527e535

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Journal of Human Evolution

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First hominine remains from a w1.0 million year old bone bed atCornelia-Uitzoek, Free State Province, South Africa

James S. Brink a,b,*, Andy I.R. Herries c,d, Jacopo Moggi-Cecchi e, John A.J. Gowlett f, C. Britt Bousman g,h,John P. Hancox i, Rainer Grün j, Vera Eisenmann k, Justin W. Adams l, Lloyd Rossouwm

a Florisbad Quaternary Research, National Museum, P.O Box 266, Bloemfontein 9300, South AfricabCentre for Environmental Management, University of the Free State, Bloemfontein, South AfricacAustralian Archaeomagnetism Laboratory, Archaeology Program, School of Historical and European Studies, Faculty of Humanities and Social Sciences, La Trobe University,Melbourne 3086, VIC, AustraliadGeomagnetism Laboratory, School of Environmental Sciences, University of Liverpool, L69 7ZE, UKe Laboratori di Antropologia, Dipartimento di Biologia Evoluzionistica ’Leo Pardi’, Universita’ di Firenze, Firenze, ItalyfBritish Academy Centenary Project, Archaeology, Classics and Egyptology, University of Liverpool, L69 3GS Liverpool, UKgDepartment of Anthropology, Texas State University, San Marcos, TX 78666, USAh School of Geography, Archaeology and Environmental Studies, University of Witwatersrand, Johannesburg, South AfricaiBernard Price Institute for Palaeontology, University of the Witwatersrand, Johannesburg, South AfricajResearch School of Earth Sciences, The Australian National University, Canberra ACT 0200, AustraliakMNHN, Département Histoire de la Terre, CP 38, 8 rue Buffon, 75005 Paris, FrancelDepartment of Biomedical Sciences, Grand Valley State University Allendale, MI 49401, USAmDepartment of Archaeology, National Museum, P.O Box 266, Bloemfontein 9300, South Africa

a r t i c l e i n f o

Article history:Received 8 December 2011Accepted 22 June 2012Available online 26 July 2012

Keywords:Early HomoCornelian Land Mammal AgeAcheulianPalaeomagnetismSouthern Africa

* Corresponding author.E-mail addresses: jbrink@nasmus.co.za (J.S. Brin

(A.I.R. Herries), jacopo@unifi.it (J. Moggi-Cecch(J.A.J. Gowlett), bousman@txstate.edu (C.B. Bousm(J.P. Hancox), Rainer.Grun@anu.edu.au (R. Grün), veadamjust@gvsu.edu (J.W. Adams), lloyd@nasmus.co.z

0047-2484/$ e see front matter � 2012 Elsevier Ltd.http://dx.doi.org/10.1016/j.jhevol.2012.06.004

a b s t r a c t

We report here on evidence of early Homo around 1.0 Ma (millions of years ago) in the central plains ofsouthern Africa. The human material, a first upper molar, was discovered during the systematic exca-vation of a densely-packed bone bed in the basal part of the sedimentary sequence at the Cornelia-Uitzoek fossil vertebrate locality. We dated this sequence by palaeomagnetism and correlated thebone bed to the Jaramillo subchron, between 1.07 and 0.99 Ma. This makes the specimen the oldestsouthern African hominine remains outside the dolomitic karst landscapes of northern South Africa.Cornelia-Uitzoek is the type locality of the Cornelian Land Mammal Age. The fauna contains an archaiccomponent, reflecting previous biogeographic links with East Africa, and a derived component, sug-gesting incipient southern endemism. The bone bed is considered to be the result of the bone collectingbehaviour of a large predator, possibly spotted hyaenas. Acheulian artefacts are found in small numberswithin the bone bed among the fossil vertebrates, reflecting the penecontemporaneous presence ofpeople in the immediate vicinity of the occurrence. The hominine tooth was recovered from the central,deeper part of the bone bed. In size, it clusters with southern African early Homo and it is alsomorphologically similar. We propose that the early Homo specimen forms part of an archaic componentin the fauna, in parallel with the other archaic faunal elements at Uitzoek. This supports an emergentpattern of archaic survivors in the southern landscape at this time, but also demonstrates the presence ofearly Homo in the central plains of southern Africa, beyond the dolomitic karst areas.

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k), a.herries@latrobe.edu.aui), gowlett@liverpool.ac.uk

an), jhancox@cciconline.comra@mnhn.fr (V. Eisenmann),a (L. Rossouw).

All rights reserved.

Introduction

Cornelia-Uitzoek (or just Uitzoek, after the farm name) is therichest of a number of Pleistocene fossil localities near the town ofCornelia in the eastern Free State of South Africa. It is located115 km south of Johannesburg and about 160 km southeast of thedolomitic area containing the well-known hominine fossil-bearingsites, such as Swartkrans and Sterkfontein (Fig. 1). Uitzoek is thetype locality of the Cornelian Land Mammal Age (LMA) of southern

Figure 1. Locality map of Cornelia-Uitzoek. The shade coding of the site locations (B) conform to their chronological positions in the Land Mammal scheme (A) for southern Africa(after Brink (2005)). A north-looking panoramic view of the site (C) shows the Pleistocene valley-fill deposits in a basin of Permian-aged Ecca Group (Karoo Supergroup) and theposition of the excavation.

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Africa (Cooke, 1974; Hendey, 1974; Brink, 2005; Fig. 1) and hasproduced Acheulian artefacts (Clark, 1974). The site became knownin the 1920s and 1930s through the work of Van Hoepen (1930,1932a,b, 1947). Apart from a short field season in 1953 by Hoffmanand others (Clark, 1974), no further fieldwork was undertaken, untilthe early 1990s, when surface collecting was resumed (Bender andBrink,1992). In 1998, a small test cutting, aimed at investigating thesedimentary context and taphonomy of the mammalian fossils andtheir relationship with the Acheulian artefacts (Brink and Rossouw,2000; Brink, 2004), uncovered part of a bone bed, which subse-quently has been systematically excavated over several fieldseasons. These excavations produced the first hominine materialfrom the site, an upper first molar. Herewe report on this specimen,its context and preliminary geochronology.

Stratigraphic context

The Uitzoek site consists of a limited pocket of valley-fill, alluvialand colluvial fossil-rich gravels and clays that accumulated ina restricted basin carved into a basement of Permian-aged EccaGroup mudstones and siltstones (locally termed shales; Fig. 1;Butzer, 1974; Brink and Rossouw, 2000; Brink, 2004). Locally theEcca Group is extensively intruded by Mid-Jurassic-aged dolerites,which locally create aureoles of hornfels with the surrounding

mudrocks. The Uitzoek sedimentary fill has been cut through anderoded by a northward-flowing left bank tributary of the Vaal River,the Schoonspruit (Tooth et al., 2004). These exposures (locallytermed dongas) were originally described by Van Hoepen (1930)and Butzer (1974) who identified a series of stratigraphic hori-zons, Beds 1e6. The new work has identified the stratigraphicsequence shown in Fig. 2. The base of the succession is formed bya poorly sorted gravel layer termed the Banded Gravel Bed (BGB,Butzer’s Bed 1), which overlies the eroded Ecca Group bedrock(Fig. 2). Gravel clasts are predominantly locally sourced Ecca Groupderivatives, as well as other Karoo-aged lithologies, includingdolerite. Fossil bone is rarely preserved and where present it isusually rolled and abraded.

Sharply overlying the BGB is the Mottled Yellow Clay (MYC,Butzer’s Bed 2). This unit contains the richest fossil horizon,a densely-packed bone bed, from which the hominine tooth wasrecovered, along with Acheulian artefacts. This bone bed occurspart way through the MYC, which is erosively overlain by theLaminated Orange Clay (LOC, Butzer’s Bed 3). The LOC is in turnoverlain by the Orange Coarse Gravels (OCG) and the Dark Grey Clay(DGC). The stratigraphic relationship of the OCG and DGC is notclear yet and they are not dealt with here further. The top of thesuccession has been pedogenetically altered to form the Black TurfSoil (BTS). Younger Acheulian material occurs in the strata above

Table 1Taxonomic list of Cornelia-Uitzoek, comparing the material excavated from the MYCbone bed until 2010 with previously collected material (Old collection), according tothe number of identified specimens.

1998e2010 Old collection

PrimatesHomo sp. 1 e

CarnivoraPanthera leo 3 5Indet. 1 e

ProboscideaIndet. 5 2

PerissodactylaEyrygnathohippus cornelianusa 1 6Equus sp. cf. E. capensisa 25 69Equus sp. cf. E. quagga 25 125Rhinocerotidae indet. 1 2

ArtiodactylaHippopotamus gorgopsa 2 49Phacochoerus sp. 3 e

Metridiochoerus modestusa 1 11Metridiochoerus compactusa 3 2Kolpochoerus heselonia 1 4Syncerus antiquusa 11 54Damaliscus niroa 209 237Megalotragus eucornutusa 12 24Connochaetes gnou laticornutus 79 150Antidorcas reckia 4 e

Antidorcas bondia 197 257Aepyceros helmoedia 12 1Sylvicapra grimmia 8 2Raphicerus sp. 1 e

a Extinct species.

Figure 2. Stratigraphic column and magnetic polarity compared to the GPTS for Cor-nelia-Uitzoek.

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the LOC, within the OCG/DGC deposits. To the southeast of theexcavation site, Middle Stone Age occurrences are documented ina series of younger deposits that occur at the same elevation as andonlap the eroded older strata.

Initially, we attempted combined Electron Spin Resonance/U-series on teeth from the site, but the uranium concentrationswere too high to obtain sensible results. In 2006, a pilot study wasundertaken at the Geomagnetic Laboratory of the University ofTexas at Austin using the methods outlined in Gose (2000) on tenpaleomagnetic samples in standard palaeomagnetic sample cubes,from five superimposed positions within the MYC, LOC and DGC.These indicated a sequence of normal (N), reversed (R) and normal(N) polarity using alternating field (AF) demagnetization. In 2008,we sampled 23 levels from the Uitzoek type section at the site ofthe excavation, with three to six sub-samples per level, usingstandard sample cubes in less consolidated sediments, andoriented block samples where possible. The plastic cubes weresubject to AF demagnetization and the block samples to thermaldemagnetization at the University of Liverpool GeomagnetismLaboratory. Subsequent analysis has been conducted at theAustralian Archaeomagnetism Laboratory. Methods employed herefollow the protocols established in Braun et al. (2010), Dirks et al.(2010), and Herries and Shaw (2011).

The magnetic sequence established in the pilot study wasconfirmed, with changes in polarity occurring at the well-definederosional or non-depositional surfaces. A single reversed polaritydirection was identified from the BGB, so the polarity of this unit isnot conclusive andwork is continuingwith the excavation of a deeptest trench. The matrix of the MYC bone bed records mostly inter-mediate directions of polarity, which are argued to reflect mixing ofthe sediments within the bone bed. Samples from the edge of thebone occurrence record a stable normal polarity as do the

surrounding MYC samples, both at the same stratigraphic level andabove the bone accumulation. Again, further analysis will takeplace as the excavation proceeds. The overlying LOC recordedreversed polarity, and the capping beds all recorded normalpolarity (Fig. 2).

The vertebrate fauna, magnetobiostratigraphy andtaphonomy

The vertebrate assemblage from Uitzoek (Table 1) recordssimultaneously a diminishing biogeographic link with East Africaand evidence for incipient southern endemism. A number of taxa isshared with the upper part of the Olduvai sequence (Upper Bed II,Beds III and IV), including Eurygnathohippus cornelianus, Hippo-potamus gorgops,Metridiochoerus compactus, Kolpochoerus heseloni,Antidorcas recki and Syncerus antiquus (Cooke, 1974; Gentry andGentry, 1978; Bernor et al., 2010; Bishop, 2010). The fauna fromBeds III and IV at Olduvai Gorge may be slightly older than 1.07 Ma(millions of years ago), using the chronology of Tamrat et al. (1995).However, it should be noted that Tamrat et al. (1995) only identifiedone layer with a reversed magnetic polarity above Bed IV, in theNorkilili Formation of the Masek Beds. As such, the assignment ofthis level to the Matuyama is tenuous at present. It may instead, asTamrat et al. (1995) note, represent a short excursion event withinthe Brunhes Chron, of which many are documented (Roberts,2008). It may also be an artefact of the recording environment. Ifthis were the case, then Beds III and IV would date to slightly olderthan 780 ka (thousands of years ago), with the Jaramillo and othershort geomagnetic events such as Cobb Mountain absent due toerosional phases.

The above mentioned equid and suid taxa are typical of theCornelian LMA and they became extinct in the south before theFlorisian LMA (Brink, 1987), with the possible exception ofM. compactus, which may have survived into the Florisian LMA(Klein, 1984). The Florisian LMA is first recorded in the Gladysvale

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external deposits sometime between 780 and 560 ka (Lacruz et al.,2002, Fig. 1). These taxa can be seen to represent an archaiccomponent in the Uitzoek fauna. In contrast, southern endemics,reflecting a derived grazing component in the fauna, include anearly morphotype of the black wildebeest, Connochaetes gnou lat-icornutus, a derived temporal form of Damaliscus niro and Bond’sspringbok, Antidorcas bondi (Brink and Lee-Thorp, 1992; Thackerayand Brink, 2004; Brink, 2005). The evolution of the black wilde-beest in particular marks the appearance of southern endemism,reflecting the typical open, Highveld-type grasslands becominga permanent feature of the landscape of the central plains ofsouthern Africa (Brink, 2005). Thus, the dual character of the Uit-zoek fauna, i.e., the archaic component reflecting a previousbiogeographic connection with East Africa and the derivedcomponent suggesting southern endemism, points to an age equalto or younger than Beds III and IV of Olduvai, which may be olderthan 1.07 Ma based on Tamrat et al. (1995), as discussed above(Fig. 2).

Preliminary taphonomic interpretation of the MYC bone bedsuggests that it was accumulated on a palaeosurface within anabandoned channel, and that the primary agent of accumulationwas a large bone-gathering predator, possibly spotted hyaenas(Brink, 2004). The bone accumulation has a NWeSE linear orien-tation andwas evidently deposited within a short tunnel within thedonga system. Many specimens show carnivore damage andhyaena coprolites are found in the bone bed. The bones do not showevidence of abrasion, as seen in fluvially transported bones. Exca-vations started at the northern-most extent of the tunnel where thebones were eroding from the section. At this point, stone toolsoccur only as occasional inclusions (Brink, 2004). The excavationshave now progressed in a southeasterly direction to where theoccurrence may have approached the ancient land surface oropened up into an unroofed section of the palaeo-donga channel.This is indicated by a rise towards the south-east of the floor of thebone occurrence and by evidence for less water logging in the fossilspecimens (Brink, 2004). At this interface zone, there is also a muchdenser accumulation of Acheulian artefacts. They are unabradedand have random in situ orientations, some being deposited on end,as if tumbled into the deposit through an open entrance or througha collapse of the roof of the tunnel. Although the Acheulian arte-facts and the vertebrate remains are preserved in the same sedi-mentary context, they may have different local origins. A directconnection between artefacts and some bones may exist, but in thecurrent absence of clear archaeological patterning and of cutmarks,no evidence yet points towards this explanation. The localconcentration of handaxes and flakes (see below) indicatesa substantial if intermittent hominine presence in the area. Thehominine tooth came from within the central, deeper part of thebone bed, which we interpret as probably reflecting the boneaccumulating behaviour of hyaenas, the primary taphonomic agent,and not a chance inclusion from the ancient land surface.

Hominine tooth

The hominine specimen (COR 2011) is identified as a right M1

(Fig. 3) because of its overall morphology, the lack of hypocone andmetacone reduction, and the straight roots (not tilted distally). Itcomprises a worn, but complete crownwith roots damaged duringexcavation. Most of the root system was recovered and glued backin their original position. The crown is well preserved, but thetexture and colour of the occlusal surface has been altered post-depositionally to the extent that some of the morphologicalfeatures are better visible on a cast. The mesio-buccal root ispresent, although a large fragment is missing on its mesial portion.Wear has reduced cusp heights, with the plane of occlusion on the

buccal cusps sloping lingually. Pits of dentine are exposed on theprotocone and the paracone. A tiny pit of dentine is also evident onthe metacone. A large, flat wear facet is present on the disto-lingualcorner of the hypocone. A large, flat interproximal wear facet(bucco-lingual 6.7 mm, occluso-cervical 2.4 mm) occupies themesial face. On the distal face, a large interproximal wear facet(bucco-lingual 7.5 mm, occluso-cervical 4.1 mm) is clearly visible. Itshows two distinct grooves emanating radially from the occlusaledge.

The occlusal outline of the crown is rhomboidal in shape, withthe buccal cusps placed mesially to the midline. All four cusps areequally well developed, with the hypocone bulging distally andlingually, and the metacone slightly larger than the paracone.Although wear has obscured some morphological details of theocclusal surface, a faint mesial fovea, buccally extended, is stillvisible (more so on a cast), bounded mesially by the mesialmarginal ridge. Most notably, a large ridge of enamel, a parastyle, isclearly evident, though worn, and is mesially placed to the para-cone. The central fovea is broad, with the mesial and buccal fissuresdeparting from it. Although worn, the trigon crest is still clearlyvisible. A distal fissure is evident, merging into the distal fovea thatis reduced to a slit by the thick distal marginal ridge. An additionalridge of enamel (metastyle) is evident also on the disto-buccalcorner of the metacone, protruding bucally. The parastyle and themetastyle give a characteristic outline to the occlusal surface. Thebuccal face is almost straight occluso-cervically, with a very slightbasal prominence. The buccal groove is moderately developed,bounded distally by an additional thin groove on the metacone.Faint mesio-buccal and disto-buccal grooves are evident in theposition of the parastyle and the metastyle, respectively. Thecervical enamel line is straight, with a slight cervical extension nearthe root bifurcation. The lingual face is slightly convex occluso-cervically, more so at the level of the hypocone. A faint lingualgroove is present. On the mesio-lingual corner, two subverticalfurrows suggest the presence of a Carabelli trait. The cervicalenamel line is almost horizontal. The preserved portions of thebuccal roots have an oval outline, mesio-distally compressed intheir central part. The remnants of the lingual root have a conicalshape.

The morphological features (a relative MD elongation, rathersteep buccal and lingual faces, lack of any bucco-lingual expansion)suggest attribution to the genus Homo. The mesio-distal (MD)length versus the bucco-lingual (BL) width dimensions fall withinthe range of the early Homo specimens (including Homo habilissensu lato, and Homo sp. from southern Africa and Ethiopia), veryclose to OH 24 and SK 27, and are larger than Homo ergaster, Homoantecessor and European Homo heidelbergensis specimens (Fig. 3).Its BL diameter falls in the range of the African Middle Pleistocenefossils, with its MD dimension surpassed only by the Kabwe spec-imen. However, the larger area of the metacone relative to that ofthe paracone has been described by Quam et al. (2009) as morefrequent in early Homo specimens, whereas the opposite trend canbe observed in later Homo. Although not strictly related to thepresent discussion, it is interesting to note that the African MiddlePleistocene fossils tend to have larger BL diameters than theEuropean Pleistocene specimens and, with the exception of Kabwe,smaller MD diameter than the majority of the early Homo group.

The two grooves evident on the distal interproximal attritionfacet extend radially from the occlusal edge. Similar grooves havebeen reported in several individuals of Middle and Late Pleistocenehuman populations from Europe (e.g., Villa and Giacobini, 1995;Pérez-Pérez et al., 2003; Rosas et al., 2006). It has been suggestedthat these grooves develop in life as a result of a heavy masticatorystress, and are usually located in correspondence with the Hunter-Schreger bands of the enamel. When the tooth is worn down in its

Figure 3. The early Homo tooth (RM1, COR 2011) from the Cornelia-Uitzoek MYC bone bed: occlusal view (A), occlusal view of a cast (B), buccal view (C), distal interproximal wearfacet (D), showing the grooves mentioned in the text, and a scatterplot (E) of the mesio-distal (MD) versus. bucco-lingual (BL) dimensions. Relevant measurements of COR 2011 are:MD, measured, 13.2 mm; MD, corrected, 13.8 mm; BL 13.4 mm; MD, cervical, 10.8 mm; BL, cervical, 13.2 mm; mesio-buccal root height (mesio-buccal corner) 10.1 mm; disto-buccalroot height (distal face) 7.5 mm; lingual root height (lingual face) 14.2 mm. Scale in cm.

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interproximal attrition facet, these grooves tend to become moreevident.

Table 2A provisional classification of artefacts from the Cornelia-Uitzoek MYCbone bed, reflecting the excavations up to 2010.

Artefact category Number

Hand-axes 7Cleavers 2Biface flakes 4Probable broken tips 3Cores/Polyhedrons 6Large (intermediate) flakes 10Flakes 65Biface trimming flakes 5Angular debitage 36Angular fragments (<20 mm) 16

Archaeology

The stone tool assemblage from the new excavations into theMYC bone bed includes as of 2010, 13 Acheulean bifaces and biface-flakes, and a number of flakes (Table 2; Fig. 4). They are madealmost entirely on hornfels, technically of poor quality, but allowingthe production of large flakes. Its exact source is uncertain,although hornfels similar to the raw material used for the MYCbifaces is present less than 1 km to the east of the site, close toa dolerite intrusion within the Ecca shales. Analyses show that thebifaces are highly distinctive in character, but compare closely withthe previous finds from the Uitzoek sediments described by Clark(1974). Several specimens stand out for their great size. Two largehandaxes made from flake blanks exceed 240mm in length (Fig. 4).Bifaces of this length are known from many sites in Africa (e.g.,Kalambo Falls, Buia, Kilombe, Peninj or Canteen Koppie: Roe, 2001;Martini et al., 2004; Gowlett, 2005; de la Torre et al., 2008; McNabb

and Beaumont, 2011, respectively) and elsewhere (e.g., Cuxton, UK:Wenban-Smith, 2004), but it is unusual for them to predominate inan assemblage. The Olorgesailie catwalk site offers a rare exampleof an assemblage where the mode is >200 mm (Isaac, 1977).

Two cleavers made on flakes are also massive, withmeasurements of 185 � 104 � 60 mm and 188 � 104 � 68 mm,

Figure 4. Plan and side views of two hand-axes from the MYC bone bed (COR MYC) assemblage (A) and discriminant analysis (B) with the Uitzoek OCG and DGC specimens, andother southern African, East African and North African/Middle East assemblages. In A, the scalloped edge appearance is clearly visible. These and other specimens also show similarcurved long sections, a result of the strongly convex ventral surfaces of the large primary flakes from which they were made. The discriminant analysis is based on 10 measuredvariables: T, Thickness; L, Length; B, Breadth; BM, Breadth at 0.5 L; BA, Breadth at 0.8 L (from butt); BB, Breadth at 0.2 L; PMB1, Point of maximum breadth (first side); PMB2, Point ofmaximum breadth (second side); TM, Thickness at 0.5; LTA, Thickness at 0.8 L (see Lycett and Gowlett (2008) for further details, and Gowlett et al. (2001) for full measurementscheme). The diagram shows mean assemblage scores. The sites are: COR MYC ¼ Cornelia-Uitzoek MYC bone bed; COR OCG and DGC ¼ Cornelia-Uitzoek OCG and DGC; K FallsA ¼ Kalambo Falls A6; K Falls B ¼ Kalambo Falls B4 Sangoan; V Douglas ¼ Vaal River Gravels, Douglas West; V Hols ¼ Vaal River Gravels, Holsdam (Van Riet Lowe, 1952); KarL ¼ Kariandusi lava; Kar U ¼ Kariandusi Upper site, obsidian; Stic ¼ Sidi Abderrahman (Biberson, 1961); Umm Q ¼ Umm Qatafa (Neuville, 1951). Umm Qatafa is included as anoutgroup.

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respectively. Even the shortest hand-axe flake is ca. 140 mm long,and the mean length for the Uitzoek bone bed series is 202 mm(standard deviation ¼ 34, N ¼ 12). In addition to finished hand-axes and cleavers, the presence of cleaver flakes and other largeblanks suggests that bifaces were made or shaped close to thefindspot.

The bifaces show a particular competence in the manufactureof long flakes as blanks, reflected in the large size and also in therelatively low breadth/length ratio mean of 0.54, and thickness/breadth ratio mean of 0.55. The specimens give an impression ofbold flaking, both in the striking of blanks and in the secondaryworking (Fig. 4). The large size of the flake scars and lack of finetrimming give the pieces a scalloped appearance around themargin, a feature also seen in the specimens illustrated by Clark(1974). Our experiments suggest that this appearance is a resultof the flaking properties of the local hornfels raw material, whichtends to resist fracture with normal weight blows, then to flakesomewhat explosively when hit harder. The finds from the earlierexcavations include a greater variety of biface forms and aresomewhat smaller, like those recently found in the youngerUitzoek LOC and DGC levels. In contrast to the purely hornfelsmaterial of the MYC stone tools, those from LOC and DGC aremade on a wider variety of raw material including quartziticsandstone. These include specimens as small as 130 mm in length(e.g., COR 3380).

The remaining artefacts in the Uitzoek MYC bone bed compriseflakes and angular debitage, a number of large flakes that includepossible reject biface blanks, and possible rare scrapers, but nolarge cores from which biface blanks were struck (‘giant cores’ cf.Sharon, 2009). These last are the only part of the operational chainthat is missing. Small flakes including biface trimming flakes arescarce but present. The pattern may suggest a site area wherebifaces were completed and used, following the import of blanks.The transport of large irregular flakes is an interesting feature. de laTorre et al. (2008) discuss ‘intermediate’ flakes at Peninj that are

large enough tomake bifaces, but appear to be by-products of blankproduction from large cores. At Uitzoek, such flakes may occurbecause they had sufficient utility to be transported along with thebiface blanks.

We restrict comments of comparison chiefly with the distinctivelower assemblage from the MYC bone bed, as further finds from thehigher levels have been made recently, and a more comprehensivepaper on the archaeology is in preparation. Few sites within SouthAfrica are directly comparable with the lower Cornelia assemblage,but there are three assemblages that are seemingly older than Uit-zoek. TheWonderwerk CaveAcheulian assemblage fromStratum11includes some large-sized bifaces and is dated to sometimebetweenc. 1.8 and 1.1 Ma (Chazan et al., 2008; Matmon et al., 2012). TheRietputs Formation sequence has material that is perhaps as old as1.6e1.4 Ma (Gibbon et al., 2009; Herries, 2011). However, theAcheulian assemblage described byGibbon et al. (2009) comes fromundated gravels and the entire sequence of gravels spans a timeperiod between 2.1 Ma and 1.1 Ma. The Sterkfontein Member 5cAcheulian assemblage (Kuman and Clarke, 2000) is dated to some-time between 1.4 and 1.1 Ma (Herries and Shaw, 2011) and may besimilar in age to the small SwartkransMember 2 probable Acheulianassemblage (Clark, 1993; Kuman, 2007; Balter et al., 2008).

There are numerous other Acheulean assemblages across SouthAfrica, but mainly younger. The Cave of Hearths Bed 1e3 assem-blages and the single hand-axe from Gladysvale are younger than780 ka (Hall et al., 2006; Herries and Latham, 2009). The smallSwartkrans Member 3 assemblage is potentially dated to between1.0 Ma and 600 ka (Balter et al., 2008; Herries et al., 2009), whileElandsfontein has been suggested to date to sometime betweenw1.0 Ma and 600 ka (Klein et al., 2007). The Elandsfontein faunahas a Cornelian character, but it is probably marginally youngerthan Uitzoek (Cooke, 1974; Brink, 2005). Canteen Kopje has largehand-axes (McNabb and Beaumont, 2011) and preliminary datesfor this site place it somewhere between 1.6 Ma and 800 ka(Gibbon, 2009).

J.S. Brink et al. / Journal of Human Evolution 63 (2012) 527e535 533

In East Africa, where early Acheulean was recently dated to ca.1.75 Ma (Lepre et al., 2011) the closest comparisons may be withmiddle/upper Olduvai Bed II (ca. 1.5e1.2 Ma) and IV (>w0.78 or>w1.07 Ma; Leakey and Roe, 1995; Tamrat et al., 1995), althoughthe Uitzoek bifaces are generally far larger. The bifaces from theMYC appearmore refined than the very early Acheulian from Peninjin Tanzania (in the range 1.5e1.1 Ma; Isaac, 1967; Isaac and Curtis,1974; de la Torre et al., 2008) in their blank striking and shaping,and less so than the well-dated late Lower Pleistocene group ofKilombe, Kariandusi and Olorgesailie (w1.1e0.8 Ma; Isaac, 1977;Gowlett and Crompton, 1994; Potts et al., 1999; Noll and Petraglia,2003; Gowlett, 2005, 2011; Herries et al., 2011). This appearancemay, however, result largely from the nature of the local hornfelsraw material. In long section, the large Cornelia hand-axes havea distinct curved profile arising from the shape of the blank flakes(Fig. 4).

A discriminant analysis is helpful for demonstrating the generalcharacter of the bifaces, based on 10 variables (Crompton andGowlett, 1993; Lycett and Gowlett, 2008, Fig. 4). In relation toother sites, it confirms the highly distinctive nature of the bifacesfrom the MYC bone bed, but also suggests some regional continuitythrough putting the two Uitzoek sets fairly close together (Fig. 4).Although the older Uitzoek assemblage stands apart, the LOC andDGC bifaces from Uitzoek also cluster with other southern Africanassemblages, including two from the Vaal River gravels measuredby us e Vaal Douglas, and Holsdam (Van Riet Lowe, 1952).

Conclusions

Given the sequential succession of magnetic polarity changesidentified from base to surface (Reversed, Normal, Reversed,Normal), the presence of a Cornelian LMA fauna and the characterof the Acheulian assemblage, the normal polarity of MYC and thebone bed, including the hominine tooth, is correlated to the Jar-amillo Normal polarity Subchron between 1.07 and 0.99Ma (Fig. 2).

The importance of the Cornelia-Uitzoek hominine specimen isemphasised by its age of between 1.07 and 0.99 Ma, a time forwhich the human fossil record in southern Africa is virtually terraincognita. It is also the oldest hominine specimen outside of theMalmani dolomite karst landscapes of northern South Africa(Taung on the edge of the Ghaap Plateau, Makapansgat in LimpopoProvince and the numerous fossil sites of the Gauteng area). It ismorphologically similar to southern African early Homo specimens,with the area of the metacone being relatively larger than that ofthe paracone, and it clusters in size with early Homo specimensfrom southern Africa and with the Kabwe specimen, rather thanwith African Middle Pleistocene Homo. The position of the Kabwespecimen is anomalous in this respect, since, although presentlyundated, it is generally considered to be in the range of 0.5 Ma oryounger (Klein, 1999).

Appendix

Hominine M1 specimens and their MD and BL measurements (mm), as plotted in Fig. 3.

Category Specimen M

COR 2011 13South African early Homo SK 27 13South African early Homo SKX 268 13South African early Homo SKW 3114 13South African early Homo SE 255 13South African early Homo STW 151 12South African early Homo DNH 70 12Homo habilis s.l. KNM-ER 1805 13Homo habilis s.l. KNM-ER 1813 12

While the occurrence of early Homo at ca. 1.0 Ma might seemunusual in an East African context, where fossils such as those fromOlorgesailie, Daka, and Buia have affinities with mid-Pleistoceneforms of Homo (Potts et al., 2004), this seems not to be the case insouthernAfrica.Australopithecushas been shown to occur at least halfa million years later in southern Africa than East Africa (Herries et al.,2010), if you take Australopithecus garhi as the last of its genus in EastAfrica. Kuman and Clarke (2000) suggest that the StW 80 mandiblefrom the Acheulian infill of Sterkfontein Member 5 representsH. ergaster. This deposit has been dated to sometime between 1.4 and1.1Ma (Herries and Shaw, 2011) and is therefore notmuch older thanthe Uitzoek hominine tooth. Another archaic hominine taxon, Para-nthropus, has also been recovered from Swartkrans Member 3 andthis has a current best age estimate of less than 1 Ma (Balter et al.,2008; Herries et al., 2009). As such, the occurrence of early Homo atUitzoek seems less unusual than it may at first appear.

In addition, the Uitzoek hominine is likely to be part of anarchaic component in the fauna, in parallel with the other archaictaxa in the fossil assemblage, probably reflecting an archaicelement in the palaeo-landscape. Thus, the late occurrence of earlyHomo supports an emergent biogeographic pattern of late survivingarchaic faunal elements in the southern African subregion at thistime. It also further emphasises the importance of the Uitzoeklocality, suggesting a wider occurrence of early Homo in the centralplains of southern Africa, distant from the hominine-bearingdolomitic caves further to the north.

Acknowledgements

Excavations at Cornelia-Uitzoek were funded initially by a grantfrom the French Department of Foreign Affairs, through a FrencheSouth African palaeontological agreement, to JSB and by theNational Museum, Bloemfontein, but later through a grant of theNational Research Foundation, African Origins Platform, User ID65245, to JSB. Palaeomagnetic sampling undertaken by BB in 2006was supported by the Leakey Foundation and Texas State University-SanMarcos. The 2006 samplesweremeasured and analyzed byHollyMeier, Deidra Aery and BB at the Geomagnetic Laboratory of theUniversity of Texas at Austin in collaboration with Wulf Gose. Allsubsequent palaeomagnetic analysis was conducted by AIRH at theUniversity of LiverpoolGeomagnetismLaboratorywith the supportofMimi Hill and Florian Stark, and the Australian ArchaeomagnetismLaboratory. This analysis was supported by an Australian ResearchFellowship as part of ARC Discovery Grant DP0877603. Without thehelpof the Florisbad team this researchwouldnot havebeenpossible.We acknowledge in particular the assistance of Mr. Abel Dichakane,Ms. Sharon Holt, Mr. Peter Mdala, who uncovered the homininemolar, Mr. Bonny Nduma, Mr. Isaac Thapo, Mr. Adam Thibeletsa andMr. Gary Trower.We thankWillem Carel Brink, Daryl Codron, SharonHolt and Liora Horwitz for reading drafts of this manuscript.

D BL Reference

.8 13.4 This paper

.7 13.3 Personal data

.1 12.7 Personal data

.6 12.2 Personal data

.3 13.4 Personal data

.9 12.9 Personal data

.7 13.1 Personal data

.2 13.4 Wood, 1991

.0 13.0 Wood, 1991

(continued on next page)

(continued )

Category Specimen MD BL Reference

Homo habilis s.l. KNM-ER 42703 12.6 13.3 Spoor et al., 2007Homo habilis s.l. OH 6 13.3 12.4 Wood, 1991Homo habilis s.l. OH 13 12.5 12.8 Wood, 1991Homo habilis s.l. OH 16 14.7 13.7 Wood, 1991Homo habilis s.l. OH 21 12.2 11.9 Wood, 1991Homo habilis s.l. OH 24 13.7 13.2 Wood, 1991Homo habilis s.l. OH 39 11.2 12.0 Tobias, 1991Homo habilis s.l. OH 41 13.9 14.1 Tobias, 1991Homo habilis s.l. OH 44 12.9 12.6 Tobias, 1991Homo habilis s.l. OH 45 13.5 13.1 Tobias, 1991Homo habilis s.l. OH 65 13.0 13.6 Blumenschine et al., 2003Homo habilis s.l. Omo L894-1 12.8 12.9 Wood, 1991Homo habilis s.l. A.L. 666-1 12.9 12.4 Kimbel et al., 1997Homo ergaster(African Homo erectus)

KNM-WT15000 12.2 11.8 Wood, 1991

African Homo erectus KGA 11-350 12.0 12.8 Suwa et al., 2007Homo rudolfensis KNM-ER 1590 14.1 15.0 Wood, 1991Homo sp. KNM-ER 807 13.0 13.6 Wood, 1991Homo sp. KNM-ER 808 13.0 13.0 Wood, 1991Homo sp. OMO SH1-17 12.9 13.5 Wood, 1991Homo sp. OMO P933-1 14.5 14.0 Wood, 1991Dmanisi D-2282 12.7 12.9 Macaluso, 2010Dmanisi D-2700 13.1 13.0 Macaluso, 2010African Mid Pleistocene Tighenif 5A 12.0 14.4 Wolpoff, 1971African Mid Pleistocene Kabwe 14.0 13.5 Wolpoff, 1971African Mid Pleistocene Kebibat 12.0 12.0 Wolpoff, 1971African Mid Pleistocene Thomas III 12.5 14.0 Bermúdez de Castro, 1986African Mid Pleistocene Salè 11.8 13.8 Day, 1986African Mid Pleistocene Lainyamok 11.2 12.6 Shipman et al., 1983African Mid Pleistocene Wadi Dagadlé 10.4 13.0 de Bonis et al., 1984African Mid Pleistocene LH 18 10.8 12.5 Day et al., 1980African Mid Pleistocene Eyasi 11.5 13.0 Wolpoff, 1971Homo heidelbergensis AT-16 11.8 12.3 Bermúdez de Castro, 1986Homo heidelbergensis AT-196 11.0 10.9 Bermúdez de Castro, 1986Homo heidelbergensis AT-20 10.7 11.9 Bermúdez de Castro, 1986Homo heidelbergensis AT-26 11.2 11.4 Bermúdez de Castro, 1986Homo heidelbergensis AT-406 12.2 12.3 Bermúdez de Castro, 1986Homo heidelbergensis Arago XXI 12.0 12.8 Bermúdez de Castro, 1986Homo heidelbergensis Petralona 12.0 13.0 Bermúdez de Castro, 1986Homo heidelbergensis Visogliano 6 12.0 12.9 Abbazzi et al., 2000Homo heidelbergensis Steinheim 11.8 12.5 Wolpoff, 1971Homo heidelbergensis Pontnewydd 4 12.3 12.4 Day, 1986Homo antecessor ATD6-11 12.1 12.9 Bermúdez de Castro et al., 1999Homo antecessor ATD6-69 12.0 12.0 Bermúdez de Castro et al., 1999

J.S. Brink et al. / Journal of Human Evolution 63 (2012) 527e535534

References

Abbazzi, L., Fanfani, F., Ferretti, M.P., Rook, L., Cattani, L., Masini, F., Mallegni, F.,Negrino, F., Tozzi, C., 2000. New human remains of archaic Homo sapiens andLower Palaeolithic industries from Visogliano (Duino Aurisina, Trieste, Italy).J. Archaeol. Sci. 27, 1173e1186.

Balter, V., Blichert-Toft, J., Braga, J., Telouk, P., Thackeray, F., Albarède, F., 2008. UePbdating of fossil enamel from the Swartkrans Pleistocene hominid site, SouthAfrica. Earth Planet Sci. Lett. 267, 236e246.

Bender, P.A., Brink, J.S., 1992. A preliminary report on new large mammal fossil findsfrom the Cornelia-Uitzoek site. S. Afr. J. Sci. 88, 512e515.

Bermúdez de Castro, J.M., 1986. Dental remains from Atapuerca (Spain). I. Metrics.J. Hum. Evol. 15, 265e287.

Bermúdez de Castro, J.M., Rosas, A., Nicolás, M.E., 1999. Dental remains fromAtapuerca-TD6 (Gran Dolina site, Burgos, Spain). J. Hum. Evol. 37, 523e566.

Bernor, R.L., Armour-Chelu, M.J., Gilbert, H., Kaiser, T.M., Schulz, E., 2010. Equidae.In: Werdelin, L., Sanders, W. (Eds.), Cenozoic Mammals of Africa. The Universityof California Press, Berkeley, pp. 685e721.

Biberson, P., 1961. Le Paleolithique Inferieur du Maroc Atlantique. Publications duService des Antiquites du Maroc, Rabat. Fascicule 17.

Bishop, L.C., 2010. Suoidea. In: Werdelin, L., Sanders, W. (Eds.), Cenozoic Mammalsof Africa. The University of California Press, Berkeley, pp. 821e842.

Blumenschine, R.J., Peters, C.R., Masao, F.T., Clarke, R.J., Deino, A.L., Hay, R.L.,Swisher, C.C., Stanistreet, I.G., Ashley, G.M., McHenry, L.J., Sikes, N., van derMerwe, N.J., Tactikos, J.C., Cushing, A.E., Deocampo, D.M., Njau, J.K., Ebert, J.I.,2003. Late Pliocene Homo and hominid land use from western Olduvai Gorge,Tanzania. Science 299, 1217e1221.

Braun, D.R., Harris, J.W.K., Levin, N.E., McCoy, J.T., Herries, A.I.R., Bamford, M.,Bishop, L., Richmond, B.R., Kibunjia, M., 2010. Early hominin diet includeddiverse terrestrial and aquatic animals 1.95 Ma ago in East Turkana, Kenya. Proc.Natl. Acad. Sci. 107, 10002e10007.

Brink, J.S., 1987. The archaeozoology of Florisbad, Orange Free State. Mem. Nas.Mus., Bloemfontein 24, 1e151.

Brink, J.S., 2004. The taphonomy of an Early/Middle Pleistocene hyaena burrow atCornelia-Uitzoek, South Africa. Rev. Paléobiol. 23, 731e740.

Brink, J.S., 2005. The evolution of the black wildebeest, (Connochaetes gnou), andmodern large mammal faunas of central southern Africa. Ph.D. Dissertation,University of Stellenbosch.

Brink, J.S., Lee-Thorp, J.A., 1992. The feeding niche of an extinct springbok, Anti-dorcas bondi (Antilopini, Bovidae), and its palaeoenvironmental meaning. S. Afr.J. Sci. 88, 227e229.

Brink, J.S., Rossouw, L., 2000. New trial excavations at the Cornelia-Uitzoek typelocality. Navors. Nas. Mus. Bloemfontein 16, 141e156.

Butzer, K.W., 1974. Geology of the Cornelia Beds. Mem. Nas. Mus., Bloemfontein 9,7e32.

Chazan, M., Ron, H., Matmon, A., Porat, N., Goldberg, P., Yates, R., Avery, M.,Horwitz, L.K., 2008. Radiometric dating of the earlier Stone Age sequence inexcavation I at Wonderwerk Cave, South Africa: preliminary results. J. Hum.Evol. 55, 1e11.

Clark, J.D., 1974. The stone artefacts from Cornelia, O.F.S., South Africa. Mem. Nas.Mus., Bloemfontein 9, 33e62.

Clark, J.D., 1993. Stone artefact assemblages from Members 1e3, Swartkrans Cave.In: Brain, C. (Ed.), Swartkrans: A Cave’s Chronicle of Early Man. TransvaalMuseum, Pretoria, pp. 167e194.

Cooke, H.B.S., 1974. The fossil mammals of Cornelia, O.F.S., South Africa. Mem. Nas.Mus., Bloemfontein 9, 63e84.

Crompton, R.H., Gowlett, J.A.J., 1993. Allometry and multidimensional form inAcheulean bifaces from Kilombe, Kenya. J. Hum. Evol. 25, 175e199.

Day, M.H., 1986. Guide to Fossil Man, fourth ed. The University of Chicago Press,Chicago.

Day, M.H., Leakey, M.D., Magori, C., 1980. A new hominid fossil skull from theNgaloba Beds, Laetoli, northern Tanzania. Nature 284, 55e56.

J.S. Brink et al. / Journal of Human Evolution 63 (2012) 527e535 535

de Bonis, L., Geraads, D., Guerin, G., Haga, A., Jaeger, J.J., Sen, S., 1984. Decouverted’un Hominide fossile dans le Pleistocene de la Republique de Djibouti. C.R.Acad. Sci., Paris 299, 1097e1100.

Dirks, P.H.D.M., Kibii, J.N., Kuhn, B.F., Steininger, C., Churchill, S.E., Kramers, J.D.,Pickering, R., Farber, D.L., Mériaux, S.-A., Herries, A.I.R., King, G.C.P., Berger, L.R.,2010. Geological setting and age of Australopithecus sediba from southern Africa.Science 328, 205e208.

de la Torre, I., Mora, R., Martinez-Moreno, J., 2008. The early Acheulean in Peninj(Lake Natron, Tanzania). J. Anthropol. Archaeol. 27, 244e264.

Gentry, A.W., Gentry, A., 1978. Fossil Bovidae (Mammalia) of Olduvai Gorge,Tanzania. Part I. Bull. Br. Mus. Nat. Hist. 29, 290e446.

Gibbon, R.J., 2009. The fluvial history of the lower Vaal River catchment. Ph.D.Dissertation, University of the Witwatersrand.

Gibbon, R.J., Granger, D.E., Kuman, K., Partridge, T.C., 2009. Early Acheulean tech-nology in the Rietputs Formation, South Africa, dated with cosmogenicnuclides. J. Hum. Evol. 56, 152e160.

Gose, W.A., 2000. Palaeomagnetic studies of burned rocks. J. Archaeol. Sci. 27,409e421.

Gowlett, J.A.J., 2005. Seeking the Palaeolithic individual in East Africa and Europeduring the Lower-Middle Pleistocene. In: Gamble, C.S., Porr, M. (Eds.), TheHominid Individual in Context: Archaeological Investigations of Lower andMiddle Palaeolithic Landscapes, Locales and Artefacts. Routledge, London,pp. 50e67.

Gowlett, J.A.J., 2011. The vital sense of proportion: transformation, Golden Section,and 1:2 preference in Acheulean bifaces. Paleoanthropology, 175e188.

Gowlett, J.A.J., Crompton, R.H., 1994. Kariandusi: Acheulean morphology and thequestion of allometry. Afr. Archaeol. Rev. 12, 3e42.

Gowlett, J.A.J., Crompton, R.H., Li, Yu, 2001. Allometric comparisons betweenAcheulean and Sangoan large cutting tools at Kalambo Falls. In: Clark, J.D. (Ed.),Kalambo Falls Prehistoric Site. The Earlier Cultures: Middle and Earlier StoneAge, vol 3. C.U.P, Cambridge, pp. 612e619.

Hall, G., Pickering, R., Lacruz, R., Hancox, J., Berger, L.R., Schmid, P., 2006. AnAcheulean handaxe from Gladysvale Cave site, Gauteng, South Africa. S. Afr. J.Sci. 102, 103e105.

Hendey, Q.B., 1974. The late Cenozoic Carnivora of the south-western Cape Province.Annals. S. Afr. Mus. 63, 1e369.

Herries, A.I.R., 2011. A chronological perspective on the Acheulian and its transitionto the Middle Stone Age in southern Africa: the question of the Fauresmith. Int.J. Evol. Biol.. http://dx.doi.org/10.4061/2011/961401

Herries, A.I.R., Curnoe, D., Adams, J.W., 2009. A multi-disciplinary seriation of earlyHomo and Paranthropus bearing palaeocaves in southern Africa. Quatern. Int.202, 14e28.

Herries, A.I.R., Latham, A.G., 2009. Archaeomagnetic studies at the cave of Hearths.In: McNabb, J., Sinclair, A.G.M. (Eds.), The Cave of Hearths: Makapan MiddlePleistocene Research Project. Southampton Monographs in Archaeology 1.Archaeopress, Oxford, pp. 59e64.

Herries, A.I.R., Hopley, P., Adams, J., Curnoe, D., Maslin, M., 2010. Geochronology andpalaeoenvironments of the South African early hominin bearing sites: a reply to‘Wrangham et al., 2009: shallow-water habitats as sources of fallback foods forhominins’. Am. J. Phys. Anthropol. 143, 640e646.

Herries, A.I.R., Davies, S., Brink, J.S., Curnoe, D., Warr, G., Hill, M., Rucina, S., Onjala, I.,Gowlett, J.A.J., 2011. New explorations and preliminary magnetobiostrati-graphical analysis of the Kilombe Acheulian locality, Central Rift, Kenya.Abstracts of the Paleoanthropology Society 2011 Meeting. Paleoanthropology2011, A16.

Herries, A.I.R., Shaw, J., 2011. Palaeomagnetic analysis of the Sterkfontein palaeocavedeposits; age implications for the hominin fossils and stone tool industries.J. Hum. Evol. 60, 523e539.

Isaac, G.L., 1967. The stratigraphy of the Peninj Group - early Middle Pleistoceneformations west of Lake Natron, Tanzania. In: Bishop, W.W., Clark, J.D. (Eds.), Back-ground to Evolution in Africa. University of Chicago Press, Chicago, pp. 229e258.

Isaac, G.L., 1977. Olorgesailie: Archaeological Studies of a Middle Pleistocene LakeBasin. University of Chicago Press, Chicago.

Isaac, G.L., Curtis, G.H., 1974. Age of early Acheulian industries from the PeninjGroup, Tanzania. Nature 249, 624e6627.

Kimbel, W.H., Johanson, D.C., Rak, Y., 1997. Systematic assessment of a maxilla ofHomo from Hadar Ethiopia. Am. J. Phys. Anthropol. 103, 235e262.

Klein, R.G., 1984. The large mammals of southern Africa: Late Pliocene to Recent. In:Klein, R.G. (Ed.), Southern African Prehistory and Palaeoenvironments. A.A.Balkema, Rotterdam, pp. 107e146.

Klein, R.G., 1999. The Human Career: Human Biological and Cultural Origins, seconded. University of Chicago Press, Chicago.

Klein, R.G., Avery, G., Cruz-Uribe, K., Steele, T., 2007. The mammalian fauna asso-ciated with an archaic hominin skullcap and later Acheulean artifacts atElandsfontein, Western Cape Province, South Africa. J. Hum. Evol. 52, 164e186.

Kuman, K., 2007. The Earlier Stone Age in South Africa: site context and the influ-ence of cave studies. In: Pickering, T.R., Schick, K., Toth, N. (Eds.), Breathing Lifeinto Fossils: Taphonomic Studies in Honor of C.K. (Bob) Brain. Stone AgeInstitute Press, Bloomington, pp. 181e198.

Kuman, K., Clarke, R.J., 2000. Stratigraphy, artefact industries and hominid associ-ations for Sterkfontein, M5. J. Hum. Evol. 38, 827e848.

Lacruz, R., Brink, J.S., Hancox, P.J., Skinner, A.R., Herries, A.I.R., Schmid, P., Berger, L.R.,2002. Palaeontology and geological context of a Middle Pleistocene faunalassemblage from the Gladysvale Cave, South Africa. Palaeont. Afr. 38, 99e114.

Leakey, M.D., Roe, D.A., 1995. Olduvai Gorge, vol. 5. C.U.P, Cambridge.

Lepre, L.J., Roche, H., Kent, D.V., Harmand, S., Quinn, R.L., Brugal, J.-P., Texier, P.-J.,Lenoble, A., Feibel, C.S., 2011. An earlier origin for the Acheulian. Nature 477, 82e85.

Lycett, S.J., Gowlett, J.A.J., 2008. On questions surrounding the Acheulean ‘tradition’.World Archaeol. 40, 295e315.

Macaluso, P.J., 2010. Variation in dental remains from Dmanisi, Georgia. Anthropol.Sci. 118, 31e40.

Martini, F., Yosief, L., Filippi, O., Asmeret, G., Habtom, K., Asmeret, K., Martino, G.,Dawit, O., Amaha, S., Tekeste, S., Zelalem, T., Dessale, Y., Samuel, Y., 2004.Characterization of lithic complexes from Buia (Dandiero Basin, DanakilDepression, Eritrea. In: Abbate, E., Beraki, W., Yosief, L., Tewelde, M.T., Rook, L.(Eds.), 2004. A Step Towards Human Origins: The Buia Homo One-Million YearsAgo in the Eritrean Danakil Depression (East Africa), vol. 110, pp. 99e132.Rivista Italiana di Paleontologia e Stratigrafia.

Matmon, A., Ron, H., Chazan, M., Porat, N., Horwitz, L.K., 2012. Reconstructing thehistory of sediment deposition in caves: a case study from Wonderwerk Cave.Geol. Soc. Am. Bull. 124, 611e625.

McNabb, J., Beaumont, P., 2011. A Report on the Archaeological Assemblages fromExcavations by Peter Beaumont at Canteen Koppie, Northern Cape, South Africa.University of Southampton Series in Archaeology 4. Archaeopress, Oxford.

Neuville, R., 1951. Le Paléolithique et Mésolithique du Désert du Judée. Masson, Paris.Noll, M., Petraglia, M.P., 2003. Acheulean bifaces and early human behavioural

patterns in East Africa and South India. In: Soressi, M., Dibble, H. (Eds.), MultipleApproaches to the Study of Bifacial Technologies. Museum of Archaeology andAnthropology, Philadelphia, pp. 31e53.

Pérez-Pérez, A., Espurz, V., Bermúdez de Castro, J.M., de Lumley, M.A., Turbón, D., 2003.Non-occlusal dental microwear variability in a sample of Middle and Late Pleisto-cenehumanpopulations fromEurope and theNear East. J. Hum. Evol. 44, 497e513.

Potts, R., Behrensmeyer, A.K., Deino, A., Ditchfield, P., Clark, J., 2004. Small mid-Pleistocene hominin associated with East African Acheulean technology.Science 305, 75e78.

Potts, R., Behrensmeyer, A.K., Ditchfield, P., 1999. Paleolandscape variation and earlyPleistocene hominid activities: members 1 and 7, Olorgesailie formation, Kenya.J. Hum. Evol. 37, 747e788.

Quam, R., Bailey, S.E., Wood, B.A., 2009. Evolution of M1 crown size and cuspproportions in the genus Homo. J. Anat. 214, 655e670.

Roberts, A.P., 2008. Geomagnetic excursions: knowns and unknowns. Geophys. Res.Lett. 35, L17307.

Roe, D.A., 2001. The Kalambo Falls large cutting tools: a comparative metrical andstatistical analysis. In: Clark, J.D. (Ed.), Kalambo Falls Prehistoric Site. The EarlierCultures: Middle and Earlier Stone Age, vol 3. C.U.P, Cambridge, pp. 492e599.

Rosas, A., Martínez-Maza, C., Bastir, M., García-Tobernero, A., Lalueza-Fox, C.,Huguet, R., Ortiz, J.E., Julia, R., Soler, V., de Torres, T., Martínez, E., Cañaveras, J.C.,Sánchez-Moral, S., Cuezva, S., Lario, J., Santamaria, D., de la Rosilla, M., Fortea, J.,2006. Paleobiology and comparative morphology of a late Neandertal samplefrom El Sidron, Asturias, Spain. Proc. Natl. Acad. Sci. 103, 19266e19271.

Sharon, G., 2009. Acheulian giant core technology: a worldwide perspective. Curr.Anthropol. 50, 335e367.

Shipman, P., Potts, R., Pickford, M., 1983. Lainyamok, a new middle Pleistocenehominid site. Nature 306, 365e368.

Spoor, F., Leakey, M.G., Gathogo, P.N., Brown, F.H., Antòn, S.C., McDougall, I.,Kiarie, C., Manthi, F.K., Leakey, L.N., 2007. Implications of new early Homo fossilsfrom Ileret, east of Lake Turkana, Kenya. Nature 448, 688e691.

Suwa, G., Asfaw, B., Haile-Selassie, Y., White, T., Katoh, S., Woldegabriel, G.,Hart, W.K., Nakaya, H., Beyene, Y., 2007. Early Pleistocene Homo erectus fossilsfrom Konso, southern Ethiopia. Anthropol. Sci. 115, 133e151.

Tamrat, E., Thouveny, N., Taieb, M., Opdyke, N.D., 1995. Revised magneto-stratigraphy of the Plio-Pleistocene sedimentary sequence of the OlduvaiFormation (Tanzania). Palaeogeogr. Palaeoclimatol. Palaeoecol. 114, 273e283.

Thackeray, J.F., Brink, J.S., 2004. Damaliscus niro horns from Wonderwerk Cave andother Pleistocene sites: morphological and chronological considerations.Palaeont. Afr. 40, 89e93.

Tobias, P.V., 1991. Olduvai Gorge. In: The Skulls, Endocasts and Teeth of Homohabilis, vol. IV. C.U.P, Cambridge.

Tooth, S., Brandt, D., Hancox, P.J., McCarthy, T.S., 2004. Geological controls onalluvial river behaviour: a comparative study of three rivers on the SouthAfrican Highveld. J. Afr. Earth Sci. 38, 79e97.

Van Hoepen, E.C.N., 1930. Fossiele perde van Cornelia, O.V.S. Paleont. Navors. Nas.Mus., Bloemfontein 2, 1e24.

Van Hoepen, E.C.N., 1932a. Die stamlyn van die sebras. Paleont. Navors. Nas. Mus.,Bloemfontein 2, 25e37.

Van Hoepen, E.C.N., 1932b. Voorlopige beskrywing van Vrystaatse soogdiere.Paleont. Navors. Nas. Mus., Bloemfontein 2, 63e65.

Van Hoepen, E.C.N., 1947. A preliminary description of new Pleistocene mammals ofSouth Africa. Paleont. Navors. Nas. Mus., Bloemfontein 2, 103e106.

Van Riet Lowe, C., 1952. The Vaal River chronology: an up-to-date summary. S. Afr.Archaeol. Bull. 7, 135e149.

Villa, G., Giacobini, G., 1995. Subvertical grooves of interproximal facets in Nean-dertal posterior teeth. Am. J. Phys. Anthropol. 96, 51e62.

Wenban-Smith, F., 2004. Handaxe typology and Lower Palaeolithic cultural devel-opment: ficrons, cleavers and two giant handaxes fromCuxton. Lithics 25,11e21.

Wolpoff, M.H., 1971. Metric Trends in Hominid Dental Evolution. Studies inAnthropology. Case Western Reserve University. The Press of Case WesternUniversity, Cleveland.

Wood, B.A., 1991. Koobi Fora Research Project IV: Hominid Cranial Remains fromKoobi Fora. Clarendon Press, Oxford.