Observations on Middle Stone Age human teeth from Klasies River Main Site, South Africa

Journal of Human Evolution 63 (2012) 750e758

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Observations on Middle Stone Age human teeth from Klasies River Main Site, South Africa Frederick E. Grine a, b a b

Department of Anthropology, Stony Brook University, Stony Brook, NY 11794-4364, USA Department of Anatomical Sciences, Stony Brook University, Stony Brook, NY 11794-8081, USA

a r t i c l e i n f o

a b s t r a c t

Article history: Received 21 June 2012 Accepted 15 August 2012 Available online 6 October 2012

The human fossils, artefacts and faunal remains from the Middle Stone Age (MSA) deposits of Klasies River Main Site have featured prominently in discussions of the evolution of modern human morphology and the emergence of human behavioral modernity. Nearly 40 human fossils were uncovered by John Wymer’s (1967e1968) excavations, and subsequent work by Hilary Deacon (1984e1995) has produced an additional dozen specimens. Many of the latter have been described, but most of the dental remains have been afforded only cursory mention and provisional identification. These specimens are documented here, and questions of individual association among some of the fossils from Wymer’s excavations are also addressed. Three teeth provide the first indisputable evidence for juvenile individuals in the deposit. The proportion of juvenile to adult remains in the MSA levels at Klasies is notably lower than in other penecontemporaneous South African coastal MSA sites such as Die Kelders Cave 1 and Blombos Cave, where the proportion of juveniles is seemingly in closer keeping with coastal, geographically proximate Later Stone Age sites such as Oakhurst Shelter and Matjes River Cave. The sizes of most of the recently identified human teeth from Klasies seem to affirm at least one arguable aspect of morphometric modernity in the MSA at this site in the form of a tendency for tooth size reduction. Ó 2012 Elsevier Ltd. All rights reserved.

Keywords: Dentition Crown size Deciduous Permanent Ontogenetic age profile

Introduction The ‘site’ of Klasies River comprises five caves or rock shelters located from 0.5 km to 2 km east of the mouth of the Klasies River on the Tsitsikamma Coast of the Eastern Cape Province of South Africa. Its significance to the study of human evolution derives from the abundant archaeological debris and the human fossils from the thick Middle Stone Age (MSA) deposits of the ‘Main Site.’ The artefacts and faunal remains have featured in arguments over the emergence of human behavioral modernity (e.g., Klein, 1975, 1976, 1989; Deacon, 1989, 1992, 2008; Ambrose and Lorenz, 1990; Klein and Cruz-Uribe, 1996; Milo, 1998; Bartram and Marean, 1999; Klein et al., 1999; Wurz, 1999, 2002, 2008, 2010; McBrearty and Brooks, 2000; Deacon and Wurz, 2001; McCall, 2006; Faith, 2008; Dusseldorp, 2010; d’Errico et al., 2012). The hominin fossils have played a prominent role in discussions of the evolution of modern human morphology (Singer and Wymer, 1982; Smith et al., 1989; Rightmire and Deacon, 1991; Bräuer et al., 1992; Smith, 1992; Frayer et al., 1993; Stringer and Bräuer, 1994; Bräuer and Singer, 1996; Churchill et al., 1996; Lam et al., 1996; Pearson and Grine, E-mail address: [email protected]. 0047-2484/$ e see front matter Ó 2012 Elsevier Ltd. All rights reserved. http://dx.doi.org/10.1016/j.jhevol.2012.08.005

1997; Groves, 1998; Pearson et al., 1998; Ahern and Smith, 2004; Royer et al., 2009). The Main Site consists of a ca. 21 m deep deposit of sediments spread across a series of interrelated recesses referred to as Caves 1, 1A, 1B, 1C and 2. Most of the work has focused on the Cave 1 and Cave 1A deposits. The first systematic excavation of the Main Site was undertaken by John Wymer, who made extensive inroads into the Cave 1, Cave 1A, Cave 1B and Cave 2 deposits in 1967e1968 (Singer and Wymer, 1982). These efforts yielded an abundance of stone tools and animal bones, as well as the bulk of the human fossils that have been recovered thus far (Singer and Wymer, 1982). Renewed excavations were initiated by Hilary Deacon in 1984, who continued this work until 1995. Deacon’s work provided an improved stratigraphic framework and a reasonable geochronology of the MSA deposits, as well as additional human fossils. A number of these specimens have been described by Rightmire and Deacon (1991, 2001), Bräuer et al. (1992), Churchill et al. (1996) and Rightmire et al. (2006). However, many of the dental remains uncovered by Deacon have been afforded only cursory mention and/or provisional identification. The purpose of the present contribution is to provide identification and descriptive documentation of these teeth.

F.E. Grine / Journal of Human Evolution 63 (2012) 750e758

Stratigraphy of the Main Site The stratigraphic framework provided by Deacon and Geleijnse (1988) combined layers with similar lithological characteristics into members, and placed most of the units and levels described by Singer and Wymer (1982) into this context. The lowermost sedimentary unit, which overlies bedrock, was referred to as the LBS (‘Light Brown Sand’) Member by Deacon and Geleijnse (1988). Levels 38 and 39 of Singer and Wymer (1982) fall within it. The sand, which is similar to that of the modern beach, was most likely deposited when sea level was very close to the present (i.e., following the last interglacial transgression of Marine Isotope Stage [MIS] 5e), at ca. 115e105 kya (thousands of years ago) (Shackleton, 1982; Deacon et al., 1988). A uranium-series date of ca. 108 kya obtained by Vogel (2001) for speleothem carbonates from layer 40 is consistent with this interpretation, as are optically stimulated luminescence dates of 115e110 kya recorded by Feathers (2002) for sand from Cave 1A (but see Millard, 2008). The lithic artefacts from the LBS Member have been categorized as conforming to MSA I by Singer and Wymer (1982), MSA 2b by Volman (1984), and the MSA I Klasies River sub-stage by Wurz (2002). The two maxillae (AA43 and Z44) recovered by Deacon from the LBS Member are the oldest hominin fossils from the site (Rightmire and Deacon, 1991; Bräuer et al., 1992). The LBS Member is overlain by the SAS (‘Shell and Sand’) Member.1 The SAS Member makes up the bulk of the thickness of the deposit in Caves 1 and 1A (Deacon and Geleijnse, 1988), and has been divided into several sub-members in these recesses to retain consistency with the units recognized by Singer and Wymer (1982). The RBS sub-member in Caves 1 and 1B appears to correspond to their level 37. Sub-members L (¼B),2 U, W and R of the SAS Member in Cave 1 correspond respectively to levels 17, 16, 15 and 14. The top of the SAS Member in Cave 1A is defined by Deacon and Geleijnse (1988) as being equivalent to layer 23 of Singer and Wymer (1982). The SAS deposits appear to have begun accumulation during MIS 5c (105e92 kya) (Shackleton, 1982; Deacon et al., 1988), and several dating methods have produced results consistent with an age of some 100e80 kya (Grün et al., 1990; Vogel, 2001; Feathers, 2002). The lithic artefacts from the SAS Member have been categorized as conforming to MSA II by Singer and Wymer (1982), MSA 2b by Volman (1984), and the MSA II Mossel Bay sub-stage by Wurz (2002). The vast majority of the human fossils recovered both by Singer and Wymer and by Deacon derive from the SAS Member, and most are from its lower units (Deacon, 2008). In Cave 1A, the relatively thin (ca. 0.5 m thick) RF (‘Rockfall’) Member, which is equivalent to layer 22 of Singer and Wymer (1982), overlies the SAS Member. The oxygen isotope profile of the RF Member correlates with MIS 5a (84e74 kya) (Deacon et al., 1988). Vogel (2001) and Feathers (2002) have reported concordant uranium-series and luminescence dates of some 77 kya and 80e 70 kya, respectively, for seemingly the lower part of this unit. The lithic artefacts from the RF Member have been categorized as conforming to MSA II by Singer and Wymer (1982), MSA 2b by Volman (1984), and the MSA II Mossel Bay sub-stage by Wurz (2002).

1 The SAS Member has also been referred to as the ‘Sands-Ash-Shell’ Member (Grün et al., 1990), although this nomenclature appears to have been abandoned by Deacon in favor of the original usage of ‘Shell and Sand.’ 2 Deacon and Geleijnse (1988) designated the stratigraphic unit immediately overlying the RBS sub-member of the SAS in Cave 1 as sub-member B. The name for this sub-member was altered to ‘L’ by Deacon (2001) in an unpublished ‘Guide to Klasies River 2001.’ Although ‘B’ continued to be used by him (e.g., Rightmire et al., 2006), ‘L’ was also employed (e.g., Deacon, 2008). Because ‘L’ has been employed consistently by Wurz (2002; Wurz et al., 2003) and appears to have been preferred by Deacon (2008), it is used here.

751

The stratigraphic sequence in Cave 1A is capped by the Upper Member. The majority of its thickness (i.e., Singer and Wymer layers 10e21) contains Howiesons Poort lithics. The lithic artefacts from the uppermost portion of the Upper Member have been categorized as conforming to a distinctive post-Howiesons Poort MSA III by Singer and Wymer (1982; see also Wurz, 2002; Villa et al., 2010). The oxygen isotope profile of the Upper Member is consistent with its deposition during MIS 4 at some 71 kya (Deacon et al., 1988); several methods have produced dates between 65 and 40 kya (Grün et al., 1990; Vogel, 2001; Feathers, 2002). The RF and Upper Member deposits have yielded two human parietal fragments (Singer and Wymer, 1982) and three isolated teeth (Rightmire and Deacon, 1991). In Cave 1, the WS (‘White Sand’) Member, which is equivalent to layer 13 of Singer and Wymer, caps the SAS (Deacon and Geleijnse, 1988). This unit is devoid of human fossils. Human fossils from the Main Site The SingereWymer excavations resulted in the recovery of nearly 40 human fossils, and almost all of these were documented in their 1982 monograph. Four others (a lumbar vertebra, a fragment of temporal bone, a partial atlas, and a left hallucial metatarsal) were identified subsequently by Richard Klein from among the faunal remains recovered from those excavations. These have been documented by Grine et al. (1998) and Rightmire et al. (2006). The vast majority of the human fossils derive from layers 14 to 17 of Cave 1 according to the field nomenclature of Singer and Wymer (1982), or the lower part of the SAS Member e sub-members L and U of Deacon and Geleijnse (1988) and Deacon (2008). Deacon’s excavations recovered a dozen human fossils. They derive from the LBS Member of Cave 1A, the SAS Member of Caves 1 and 1A, and the Upper Member of Cave 1A (Table 1). As noted above, a number of them have been described (Rightmire and Deacon, 1991, 2001; Bräuer et al., 1992; Churchill et al., 1996; Rightmire et al., 2006). Ontogenetic age profile of the Main Site assemblage With regard to the large collection that has been described from the SingereWymer and Deacon excavations, only one, the KRM 16425 frontal fragment, has been identified as possibly representing a subadult individual (Smith, 1992). The supraorbital morphology exhibited by this specimen is undeniably modern (Singer and Wymer, 1982; Rightmire and Deacon, 1991), and Smith (1992) has proposed that it may derive from an immature individual in explanation of its weak brow and glabellar prominence. Thus, Ahern and Smith (2004: 14) have argued that it might represent a “subadult version of a late archaic” African cranium. The evidence for the ontogenetic immaturity of KRM 16425, however, is neither overwhelming nor universally concordant. Thus, for example, Singer and Wymer (1982: 142) state that the frontonasal suture is “almost obliterated.” Any degree of fusion might be unexpected in a juvenile, but unfortunately there are no published data on age changes in this particular suture in recent humans. Nevertheless, its full fusion would seem to be very rare (ca. 20% of individuals over the age of 70 years; D. Ubelaker, Personal communication). A close examination of this feature would seem to be a reasonable topic of discovery since it might well be the best evidence available to address the question of the age of KRM 16425. Habgood (1989: 20) has stated that “the degree of frontal sinus development suggests that this fragment is from an adult individual,” although he provided no evidence in support of this claim. Indeed, Ahern and Smith (2004) observed that while the frontal sinus of KRM 16425 is ‘respectable,’ so too are those of Neandertal juveniles from Vindija and Le Moustier. In this regard, they also

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F.E. Grine / Journal of Human Evolution 63 (2012) 750e758

Table 1 Klasies River Main Site hominin specimens recovered during Deacon's (1984e1995) excavations. Specimen

Grid Coordinate

Member

Cave

Maxillary fragment Maxillary fragment Ldi1 RP4 Rdm1 Right proximal ulna Rdm2 I2 LM2 LM3 Left metatarsal II Right metatarsal V

AA43/SAS4 SHB Z44/SAS4 SHC E50/TSAS E50/AV H51/CP1 ? 01/C1/SMB A2/1/SMB A2/3/SMB A1/4/SMB A2/2/SMB2 B1/3/SMB

LBS LBS Upper Upper Upper SAS SAS U SAS U SAS U SAS U SAS U SAS U

1A 1A 1A 1A 1A 1A 1 1A 1A 1A 1A 1A

note that the sinus in the Klasies specimen does not extend superiorly into the frontal squama or laterally along the brow. They maintained that this is commensurate with an adolescent age for the specimen according to the Bolton Standards. However, it is uncertain that developmental standards established by a sample of 32 white North Americans aged three to 18 years from Cleveland, Ohio (Broadbend et al., 1975) necessarily apply to later Pleistocene African populations. Finally, with regard to KRM 16425, Ahern and Smith (2004: 14) noted that: “the browridge may thin laterally and exhibits a distinct supraorbital notch. Both of these features are characteristic of modern human browridges but are also found in adolescent Neandertals, as our observations on Vindija 224 and 279 show. They are also present on late archaic African specimens such as Florisbad.” However, what appears distinctly modern on the KRM 16425 frontal is the presence of separate medial and lateral components of the brow (i.e., a very faint, oblique depression separates a weak superciliary arch [¼eminence] medially from a flattened supraorbital trigone laterally [Cunningham, 1908]). The Florisbad brow lacks any such separation, and nor is it readily apparent in Le Moustier 1, Vindija 224 or Vindija 279 to judge from Ahern and Smith’s (2004) illustrations. Indeed, the characteristic cranial differences between Neandertals and modern humans, including those of the frontal, arise early in development and are maintained throughout ontogeny (Ponce de León and Zollikofer, 2001). Thus, although the ontogenetic age of the individual represented by the KRM 16425 frontal fragment is a matter of discussion, Smith’s (1992) suggestion is of interest not only because of its potential significance in relation to the morphology exhibited by the Klasies hominin fossils, but also from the perspective of the ontogenetic age profile of this assemblage. The potential presence of only a single subadult individual in the Klasies assemblage stands in marked contrast to those from other geographically proximate South African coastal MSA sites where more than a few human remains have been recovered (e.g., Die Kelders Cave 1 and Blombos Cave). Thus, the assemblage from Die Kelders Cave 1 comprises 27 specimens (24 isolated teeth, a mandibular fragment and two manual phalanges) that likely represent ten individuals; the vast majority are children, and all may be subadult (Grine et al., 1991; Avery et al., 1997; Grine, 2000). Similarly, of the nine isolated human teeth from Blombos Cave, five are deciduous (Grine et al., 2000; Grine and Henshilwood, 2002). Comparable situations pertain to the large skeletal collections from the geographically proximate Later Stone Age coastal rock shelters of Oakhurst and Matjes River. Excavations by Goodwin (1938) at the former recovered some 35 mid-Holocene human burials, of which 17 are children (Drennan, 1938a,b). Prolonged work at Matjes River

Reference Rightmire Rightmire Rightmire Rightmire Rightmire Rightmire Rightmire Rightmire Rightmire Rightmire Rightmire Rightmire

and Deacon (1991: 145); Brauer et al. (1992: 419e420) and Deacon (1991: 145); Brauer et al. (1992: 420) and Deacon (1991: 145, no identification) and Deacon (1991: 145, as a P3) and Deacon (1991: 145, no identification) and Deacon (1991: 146e147); Churchill et al. (1996: 214e220) and Deacon (2001: 538, possible dm2) and Deacon (2001: 538e539, anterior tooth) and Deacon (2001: 539, same individual as A1/4/SMB) and Deacon (2001: 540, same individual as A2/3/SMB) et al. (2006: 98) et al. (2006: 98e99)

(Dreyer, 1933; Hoffman, 1958; Louw, 1960) uncovered the remains of between 88 and 103 individuals, and over half are subadult (L’Abbé et al., 2008). Observations on the human teeth In 1996, the present author was afforded the opportunity to examine the human fossils that had been excavated by H.J. Deacon from the Main Site. In the course of this study, all of the teeth that had been only briefly mentioned and provisionally identified by Rightmire and Deacon (1991, 2001) were found to be identifiable to specific type. All of these specimens were re-examined by the present author in February, 2011, aided by a low-power binocular microscope, and the identifications were confirmed. Mesiodistal (MD) and buccolingual (BL) dimensions were measured according to the definitions of Tobias (1967), where MD represents the maximum distance between the mesial and distal surfaces determined by the longitudinal axis of the crown, and BL is the maximum distance between the buccal and lingual surfaces at a right angle to the longitudinal axis of the crown. One aspect of these identifications is that they serve for the first time to unquestionably document juvenile individuals at Klasies. Singer and Wymer recovered a partial mandibular corpus (KRM 13400) containing the RP4-RM2 and four isolated permanent teeth (three molars and a premolar) in close proximity to one another in Layer 14 (i.e., sub-member R of the SAS Member of Deacon and Geleijnse, 1988) of Cave 1. There has been disagreement between Singer and Wymer (1982) and Rightmire and Deacon (1991) as to their association with one another, and particularly over whether the isolated premolar belongs with the others or represents a second individual. These fossils were re-examined with an eye to addressing this issue. The Deacon specimens Because Deacon did not provide specimen numbers for the human remains and they have yet to be accessioned by the Iziko South African Museum, their excavation grid coordinates are employed here in lieu of specimen numbers. Rightmire and Deacon (1991: 145) noted that, in addition to the two maxillae recovered from the LBS Member, “several isolated teeth have been collected. Some are too incomplete to be studied, but one lower premolar has an intact root. Enamel has chipped away from all of the buccal half of the crown, but the small lingual cusp is only slightly worn. Proportions of the part of the crown remaining are consistent with identification of this tooth as a P3. Mesiodistal length can be estimated as 6.7 mm”

F.E. Grine / Journal of Human Evolution 63 (2012) 750e758

Although the stratigraphic derivation of these isolated teeth was not indicated, and Rightmire and Deacon’s (1991) text might suggest that they are from the LBS Member, they come from the Upper Member of Cave 1A (Sarah Wurz, Personal communication). They are associated with the Howiesons Poort and MSA III levels. The three specimens in question are from square E50, units AV and TSAS, and from square H51, unit CP1 (Table 1). The E50/AV premolar is considered here to be a RP4 rather than a P3, and the other two isolated teeth referred to by Rightmire and Deacon (1991) are identified here as a Ldi1 (E50/TSAS) and a fragment of a Rdm1 (H51/CP1). All three derive from Cave 1A. Rightmire and Deacon (2001) also mentioned an ‘anterior tooth,’ tentatively regarded by them as a permanent upper lateral incisor. This specimen derives from square A2/1, unit SMB. They posited that it possibly derived from the same individual as two isolated molars (LM2 and LM3) described by them from the same unit (¼layer) in subjacent squares A2/3 and A1/4. Their identification of the tooth as an I2 is almost certainly correct. Moreover, its state of wear, color and patina are entirely consistent with its belonging to the same individual as the molars. This tooth was provided neither description nor illustration. Rightmire and Deacon (2001: 538) also made mention of a “fragment of tooth crown,” which they considered to be “probably a lower molar,” and “likely on size considerations to be deciduous.” This fragment is the mesial end of a Rdm2; it was neither described nor illustrated by them. It was recovered from Square 01/C1, unit SMB of the ‘witness baulk’ in Cave 1. Ldi1 E50/TSAS This specimen comprises a very heavily worn crown with a short (4.3 mm long) segment of root (Fig. 1). The root is broken obliquely, with a longer mesial edge. The crown is worn such that the labial face retains only 2.8 mm of enamel height; the labial margin is slightly beveled. Incisal wear has produced a strong distolingual slope, with only a small remnant of the distolingual enamel margin preserved. The incisal outline of the crown is testament to the presence of a distolingually disposed basal tubercle. There is no evidence of enamel hypoplasia on the remnant of the labial face. The root measures 6.0 mm MD and ca. 4.0 mm BL at the cervical margin. The labial aspect of the root is crossed by a very shallow, 0.6 mm broad furrow immediately above the cervical margin. The crown, as preserved, measures 6.5 mm MD and 5.0 mm BL; the degree of crown loss through wear precludes estimation of the original MD diameter. The BL diameter is compared with other South African MSA homologs in Table 2. It is somewhat smaller in this dimension than incisors from Die Kelders Cave 1 and Blombos,

Figure 1. The E50/TSAS Ldi1 in labial (left) and incisal (right) views. Mesial to the left in both views; labial to the top in (b).

753

Table 2 BL diameter of the Klasies E50/TSAS di1 compared with values recorded for recent African, MSA African and MP Eurasian samples. Sample/specimen

n

Mean

Klasies E50/TSAS Die Kelders AP 6249 Die Kelders AP 6278 Blombos AP 6293

1 1 1 1

5.0 5.3 6.4 5.7

Blombos AP 8973

1

5.2

59 22 21 15

5.1 5.0 5.0 6.1

KhoeSan _ þ \ South African _ þ \ Liberian _ þ \ Neandertal

s.d.

0.3 0.4 0.3 0.4

Obs. range

4.5e5.8 4.2e5.9 5.4e6.9

Reference This study Grine (1998) Grine (1998) Grine and Henshilwood (2002) Grine and Henshilwood (2002) Grine (1998) Grine (1986) Moss and Chase (1966) Grine et al. (1991),a

a

The MP Eurasian Neandertal sample includes specimens from Chateauneuf, Engis, Kebara, Tabun, La Ferrassie, Hortus, Krapina, Lazaret, Neussing, Pech de l’Aze, Roc de Marsal, Shanidar, Staroselje and Subalyuk. See Grine et al. (1991: table 6) for the sources from which the measurements for these specimens were taken.

and smaller also than penecontemporaneous Middle Palaeolithic (i.e., Neandertal) di1s from Europe. The BL diameter of the Klasies di1 corresponds to the values recorded for recent sub-Saharan African population samples. RP4 E50/AV This specimen consists of the damaged crown and complete root of an isolated RP4 (Fig. 2). It was identified as a P3 by Rightmire and Deacon (1991). Enamel has been lost from the buccal half of the protoconid. Occlusal wear is very slight on the metaconid; no dentine is exposed. There are small (1.9 mm BL) mesial and larger (minimum of 2.8 mm BL) distal interproximal contact facets. The distal facet presents several shallow, vertical furrows. The protoconid was substantially larger than the metaconid, which is set distal to the midcrown transverse axis that bisects the protoconid. The protoconid and metaconid are separated by a deep longitudinal fissure that extends buccally at its ends to delineate short mesial and distal fovea. The mesial and distal marginal ridges are low, but complete. There is no indication of enamel hypoplasia on preserved crown surfaces. The single slender root measures 5.2 mm MD and 6.7 mm BL at the cervix. It is long (17.6 mm), and vertically disposed except at the tip, which has a slight mesial curve. The root tapers evenly both MD and BL to a sharp apex.

Figure 2. The E50/AV RP4 in occlusal view. Mesial is to the left and buccal to the top.

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F.E. Grine / Journal of Human Evolution 63 (2012) 750e758

Table 3 MD diameter of the Klasies E50/AV P4 compared with values recorded for recent African, MSA African and MP Eurasian samples. Sample/specimen Klasies E50/AV KRM 13400/14692 Die Kelders AP6279 KhoeSan _ þ \ South African _ South African \ Atapuerca Neandertals

n

Mean

1 2 1 13 293 92 10

6.9 8.1 8.3 7.3 7.4 7.3 7.2

37

7.8

s.d.

Obs. range

Reference

0.6 0.5 0.5 0.5

6.5e8.3 6.2e9.0 6.2e8.9 6.6e7.7

0.6

6.9e9.4

This study This study Grine (1998) Grine (1998) Jacobson (1982) Jacobson (1982) Bermúdez de Castro (1993) Bermúdez de Castro (1986)

The crown measures 6.7 mm MD, as noted by Rightmire and Deacon (1991). Only an additional 0.2 mm is necessary to compensate for interproximal wear. It is not possible to determine the BL diameter of the crown. The MD diameter is compared with other South African MSA homologs in Table 3. The E50/AV P4 is notably smaller than the other Klasies (KRM 13400) homolog. It is also smaller than the Die Kelders Cave 1 and the majority of European Middle Palaeolithic premolars. While the E50/AV P4 diameter falls within the fiducial limits of recent southern African population samples, it is below the mean for each. Rdm1 H51/CP1 This specimen consists of an incomplete (distobuccal two-thirds) crown and a small root fragment of a Rdm1 (Fig. 3). The metacone, distal marginal ridge, the buccal half of the hypocone, a small part of the paracone, and the distobuccal corner of the protocone are preserved. The crown is moderately worn. An appreciable dentine island is exposed on the protocone, and this extends along the distal trigon ridge. The paracone and hypocone preserve remnants of moderate dentine islands; there is a very small exposure on the metacone. The distal crown face preserves part of a BL broad, deeply concave interproximal contact facet. This appears to have been artificially deepened by dentine loss. The distal marginal ridge is thick and complete. The talon basin is represented by a narrow, 2.6 mm long BL fissure that extends mesially around the base of the hypocone. The buccal crown face presents a broad, V-shaped depression between the paracone and metacone and a second, smaller depression between the metacone protuberance and the distal margin of the crown. The latter partially delineates a small distoconule at the buccal end of the distal marginal ridge. The cervical enamel prominence above the metacone shows evidence of mesial expansion into a tuberculum molare.

Figure 3. The H51/CP1 Rdm1 in occlusal (left) and buccal (right) views. Mesial is to the right in the occlusal view, and to the left in the buccal view.

A short segment of the buccal side of the root neck is preserved, which appears to have been very low (ca. 1 mm from the cervical margin to the bifurcation of the buccal roots). The distobuccal root appears to have flared to a considerable degree. A strip of dental calculus is preserved along the margin of the buccal enamel cervix. The crown fragment measures 7.5 mm MD and 7.2 mm BL. These cannot be taken as even approximating the original crown dimensions. It is clear that the original crown dimensions would have been much larger, and that this was a comparatively large tooth. I2 A2/1/SMB This is the very heavily worn crown and complete root of a maxillary permanent lateral incisor (Fig. 4). The side from which it derives cannot be determined; it is treated here as a LI2 only for descriptive purposes. The crown is worn to the level of the mesial and distal cervical margins. A short segment of the labial enamel face is preserved, together with a narrow rim of the lingual cervical enamel margin. Wear on the incisal surface has exposed a secondary dentine infilling of the radicular canal. The crown, as preserved, measures 5.5 mm MD. This, of course, is of no meaning beyond establishing a minimum MD diameter. The remnant of lingual enamel permits the BL crown diameter to be determined at 6.2 mm. The root surface is smooth save for a short, shallow furrow on its mesial face. The apical two-thirds of the root is set at a slight lingual tilt to the cervical third, and this change in orientation is most notable across the labial surface. The apical portion of the root is slightly tapered BL. The root is 13.5 mm long (measured labially), and 4.4 mm MD and 6.0 mm BL at the cervix. The BL diameter of this crown is compared with European Middle Palaeolithic and recent South African homologs in Table 4. The Klasies value is well below the observed ranges for European MP samples, and while it is somewhat lower than the means for recent South African Bantu-speaking males and females, it falls comfortably within the observed ranges and fiducial limits for both samples.

Figure 4. The A2/1/SMB I2 in ?mesial (left) and incisal (right) views. Labial is to the top in the incisal view.

F.E. Grine / Journal of Human Evolution 63 (2012) 750e758 Table 4 BL diameter of the Klasies River A2/1/SMB I2 compared with values recorded for recent African and MP Eurasian samples. Sample/specimen Klasies A2/1/SMB Atapuerca Neandertals South African _ South African \

n

Mean

s.d.

Obs. range

1 4

6.2 7.9

0.3

7.6e8.3

24

8.6

0.6

7.5e9.9

195 71

6.7 6.4

0.5 0.5

4.9e8.2 5.0e7.4

Reference This study Bermúdez de Castro (1993) Bermúdez de Castro (1986) Jacobson (1982) Jacobson (1982)

Rdm2 O1/C1/SMB This specimen comprises the mesial end of a moderately worn Rdm2 (Fig. 5). The crown preserves part (approximately half) of the metaconid, the protoconid and the mesial half of the hypoconid. The metaconid has a small dentine exposure at its apex. The protoconid and hypoconid have been reduced to a flat plane with confluent lakes of dentine. The mesial face exhibits three broad vertical furrows, and a ‘doubled’ interproximal contact facet with superior and inferior beveled surfaces. This suggests that the dm1 had been shed and the P3 had erupted into functional occlusion prior to the death of this individual (or to the dm2 having been shed). The mesial root is preserved for only a short distance (ca. 2.8 mm) beyond the cervical margin. The cervical margin of the hypoconid preserves a thin strip of calculus. The trigonid is 8.3 mm BL. It is not possible to estimate the BL diameter of the talonid. Because the trigonid of the dm2 is most commonly narrower than the talonid (Grine, 1984), almost all studies that record the maximum BL diameter of this tooth relate to the talonid. The only samples for which separate trigonid and talonid dimensions have been recorded are recent KhoeSan and South African Bantu-speaking populations. The BL diameter of the Klasies dm2 trigonid falls between the means for these recent samples (Table 5).

Figure 5. The O1/C1/SMB Rdm2 in occlusal view. Mesial is to the left.

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Table 5 BL diameter of the Klasies River O1/C1/SMB dm2 trigonid compared with values recorded for recent African samples. Sample/specimen Klasies O1C/1 SMB KhoeSan _ þ \ South African _ þ \

n

Mean

1 101 35

8.3 8.1 8.4

s.d. 0.4 0.4

Obs. range

Reference

6.8e9.0 7.5e9.3

This study Grine (1984) Grine (1984)

The Singer and Wymer specimens Singer and Wymer recovered five fossils in close proximity to one another in the SAS Member (i.e., Layer 14) of Cave 1. The largest is a partial mandibular corpus containing the RP4-RM2. The other four are isolated permanent teeth. They are cataloged as:

KRM KRM KRM KRM KRM

13400 14692 14691 14693 14694

(SAM-AP (SAM-AP (SAM-AP (SAM-AP (SAM-AP

6223) 6227) 6228) 6229) 6230)

Mandible with RP4-RM2 LP LM1 LM2 LM3

The mandible consists of a robust right corpus preserving the RP4-M2, as well as the sockets of the L&R I1-P3 and part of the LP4 alveolar margin. It was illustrated and briefly described by Singer and Wymer (1982). Rightmire and Deacon (1991) provided additional descriptive detail. The isolated teeth were illustrated and briefly described by Singer and Wymer (1982). These specimens are shown here together in Fig. 6. Singer and Wymer (1982) associated all four isolated teeth with the KRM 13400 mandible. Rightmire and Deacon (1991) agreed with the association of the LM1, LM2 and LM3 with the mandibular corpus, but argued that the premolar belonged to a different individual. Singer and Wymer (1982) regarded the KRM 14692 premolar as a LP3, and stated that it fits into the appropriate socket of KRM 13400. Rightmire and Deacon (1991) accepted the identification of this tooth as a LP3, but observed that when the KRM 14692 root is aligned with the empty RP3 socket in KRM 13400, the crown projects well above the level of the arcade that is preserved on the right side of the mandible. They concluded, therefore, that despite its archaeological provenance, KRM 14692 “should not be grouped with the lower jaw” (Rightmire and Deacon, 1991: 139). With regard to the KRM 14692 premolar, both Singer and Wymer (1982) and Rightmire and Deacon (1991) are correct on one count, but mistaken on others. Contra both Singer and Wymer (1982) and Rightmire and Deacon (1991), this isolated premolar is not a LP3. Rather, KRM 14692 is a LP4. KRM 14692 is clearly the antimere of the KRM 13400 RP4 in terms of crown morphology, crown size, wear and root length. The right (KRM 13400) P4 crown is complete; the left (KRM 14692) is missing a small enamel chip from the lingual half of the mesial surface. Nevertheless, in both, the crown has a nearly square occlusal outline, and the protoconid is larger than the metaconid, which is set mesial of the midcrown transverse axis that bisects the protoconid. In both, the longitudinal fissure is continuous distally with an elongate fovea posterior, which has subequal buccal and lingual limbs. The small fovea anterior is deflected buccally and opens mesially, incising the narrow, low mesial marginal ridge. The distal marginal ridge is thick and continuous, slightly swelling the distolingual corner of the crown. The buccal and lingual surfaces are unremarkable, and neither crown evinces any evidence of enamel hypoplasia.

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Figure 6. The KRM 13400 (SAM-AP 6223) mandible with RP4-RM2, KRM 14692 (SAM-AP 6227) LP4, KRM 14691 (SAM-AP 6228) LM1, KRM 14693 (SAM-AP 6229) LM2, and KRM 14694 (SAM-AP 6230) LM3. Left: occlusal view in articulation. Scale bar ¼ 1 cm. Right: lateral flat plane radiographs of the KRM 13400 mandibular corpus (below), and the KRM 14692 (SAM-AP 6227) LP4, KRM 14691 (SAM-AP 6228) LM1, KRM 14693 (SAM-AP 6229) LM2, and KRM 14694 (SAM-AP 6230) LM3 (above) in articulation. The radiographs reveal the similarities in root length and configuration between the left and right antimeres. The radiographs were recorded in 1996 using an Elma Orbix with a 120 cm source-image distance and 0.3 mm2 focal spot size. The mandibular corpus was recorded at 60 kV and 22 MAS; the isolated teeth at 55 kV and 20 MAS.

Occlusal wear has produced enamel facets on the buccal and lingual cusps; there is a small dentine exposure in the center of the protoconid on the right. The distal interproximal contact facet of KRM 14692 matches the mesial facet of the KRM 14691 LM1. It is also evident that despite being incomplete, the mesial interproximal contact facet of KRM 14692 was broad, being comparable in size to that on the KRM 13400 RP4. The principal dimensions of the crowns are identical, or virtually so (Table 6). The LP4 root is single and long, measuring 9.1 mm in length. It is moderately inclined distally, and the apical half has a slight distal curvature. The tip exhibits slight hypercementosis. These features and dimensions accord with the radiographic appearance of its antimere in the KRM 13400 mandible. Thus, the KRM 14692 P4 is associated with the isolated molars and the KRM 13400 mandible. This is in accord with the individual associations proposed by Singer and Wymer (1982), who appear to have been right for the wrong reason. Discussion Five human teeth recovered by Deacon from the Upper and SAS Members of the Klasies River Main Site are here identified and described. Three of these provide the first indisputable documentation of juvenile remains from the site. The degree of wear of the Ldi1 (E50/TSAS) and Rdm1 (H51/CP1) from the Upper Member is compatible with both having derived from a single individual. However, these two teeth are separated by over a meter of sediment (Villa et al., 2010; S. Wurz, Personal Table 6 Principal crown dimensions of the premolars and molars in the KRM 13400 mandible and the isolated teeth associated with it. Specimen KRM KRM KRM KRM KRM KRM KRM

14692 13400 14691 13400 14693 13400 14694

LP4 RP4 LM1 RM1 LM2 RM2 LM3

MD meas.

MD est.

BL meas.

BL est.

7.6 7.6 12.9 12.8 12.7 12.5 12.2

8.1 8.1 13.2 13.2 13.5 13.3 12.4

9.2 9.3 11.2 11.1 10.5 10.8 9.9

9.2 9.3 11.2 11.1 10.5 10.8 9.9

communication), indicating that it is most unlikely that they represent a single individual. Thus, it would appear that they attest to the presence of two juvenile individuals within the Upper Member strata of Cave 1A. While the state of preservation of the Rdm1 (H51/CP1) cannot preclude the possibility that this tooth had simply been shed in the deposit, there is no evidence of resorption of the root of the Ldi1 (E50/TSAS). At the same time, the E50/AV RP4 from the Upper Member may represent a subadult individual given its state of wear, but it is extremely unlikely that it could have derived from the same individual as the E50/TSAS Ldi1. Stratigraphic as well as developmental considerations also likely preclude the premolar being associated with the Rdm1 (H51/CP1) from the Upper Member. The Rdm2 (O1/C1/SMB) derives from the SAS Member (submember U) of Cave 1, which would almost certainly preclude its association with either of the other two deciduous teeth discussed above. Moreover, there is evidence that the individual from which the dm2 derived had already erupted the P3 into occlusion, and under normal circumstances this would preclude the di1 (E50/ TSAS) being associated with the dm2 (O1/C1/SMB). Although the degree of wear, preservation of the enamel, and dentine coloration are strikingly similar on the Rdm1 (H51/CP1) and Rdm2 (O1/C1/ SMB), which might be taken as being consistent with their having derived from a single individual, their different stratigraphic derivations certainly speak against this association. On the other hand, the level from which the Rdm2 (O1/C1/SMB) derives e the SAS Member (sub-member U) of Cave 1 e is equivalent to level 16 of Singer and Wymer (1982). Interestingly, this is the level from which the KRM 16425 (¼SAM-AP 6103) frontonasal fragment suspected by Smith (1992), Ahern and Smith (2004) as belonging to a juvenile individual also derives. The presence of a deciduous molar provides incontrovertible evidence of a juvenile individual from this particular stratum. While the possibility that the molar and frontal fragment may derive from a single individual should be entertained, it is not possible at present to provide any evidence for this association. Indeed, there are other adult hominin fossils from this particular sub-member (Singer and Wymer, 1982) with which KRM 16425 could be associated. Even though juvenile hominin remains have now been indisputably documented from the MSA layers at Klasies River Main Site,

F.E. Grine / Journal of Human Evolution 63 (2012) 750e758

and even if the three deciduous teeth derive from separate individuals and the KRM 16425 frontal fragment does represent a subadult individual (Smith, 1992), the proportion of juvenile to adult remains in this repository remains notably lower than in other penecontemporaneous South African coastal MSA sites. Thus, at Die Kelders Cave 1 and Blombos Cave between 50% and 100% of individuals are juvenile, a proportion that is seemingly in closer keeping with coastal, geographically proximate LSA sites such as Oakhurst Shelter and Matjes River Cave. The possibility that the preponderance of adult human remains at Klasies River might in some way be related to the practice of cannibalism at the site (White, 1987; Deacon, 1992, 2008) warrants further investigation. With regard to individual associations at Klasies Main Site, that between the KRM 13400 partial mandible and the four isolated teeth (KRM 14691e14694) suggested by Singer and Wymer (1982) (contra Rightmire and Deacon, 1991) is confirmed. Although no meaningful dimension could be recorded for the H51/CP1 dm1 fragment, it appears to have derived from a comparatively large tooth. On the other hand, the other two deciduous teeth for which crown dimensions could be recorded (the E50/TSAS di1 and 01/C1/SMB dm2) are comparatively small. In this regard, the di1 is somewhat smaller BL than South African MSA homologs from Die Kelders Cave 1 and Blombos, corresponding to the values recorded for recent sub-Saharan African population samples. Similarly, the BL diameter of the dm2 trigonid falls between the means recorded for two recent southern African human populations. The permanent teeth recovered by Deacon are also comparatively small. Thus, in its MD diameter, the E50/AV P4 is notably smaller than other Klasies River (KRM 13400/14692) and Die Kelders Cave 1 homologs, falling below the means for recent South African population samples. Similarly, the BL diameter of the Klasies A2/1 SMB I2 is not only smaller than European Middle Palaeolithic homologs, it is also somewhat lower that the means for recent South African samples. As such, the comparatively small sizes of most of the recently identified human teeth from Klasies River appear to affirm at least one arguable aspect of morphometric modernity at this MSA site in the form of a tendency for tooth size reduction. While a number of Klasies River fossils exhibit arguably archaic traits, and the degree of size dimorphism seems unmatched among recent people, the limited evidence provided by the teeth is consistent with other MSA cranial remains from this site in suggesting a pattern of general, albeit incomplete morphological modernity. As aptly observed by Smith (1992: 148), the “somewhat primitive aspects of certain features in some specimens” are to be expected in an otherwise morphologically modern population of this geological antiquity. The mosaic nature of evolution is manifest in these Late Pleistocene representatives of the human lineage. Acknowledgments I thank the late Hilary Deacon for the invitation in 1996 to his home in Stellenbosch to study the human remains that he had excavated so carefully from Klasies River, and I thank Sarah Wurz for permission to re-examine them in 2011. Sarah also graciously provided confirmation of their stratigraphic derivation, and cogent comments on the manuscript. I am grateful to Wilhelmina Seconna for her gracious hospitality and support during my work on this material at the Iziko South African Museum, Cape Town in 2011. I thank Luci Betti-Nash for her masterful illustrations, and Gillan Bowie, Department of Radiology, Groote Schuur Hospital, for the radiographs. In addition to Sarah Wurz, I thank Fred Smith and Philip Rightmire as well as the Associate Editor and three anonymous reviewers for their comments and suggestions, which improved the manuscript.

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References Ahern, J.C.M., Smith, F.H., 2004. Adolescent archaic or adult moderns? Le Moustier 1 as a model for estimating age at death of fragmentary supraorbital fossils in the modern human origins debate. Homo 55, 1e19. Ambrose, S.H., Lorenz, K.G., 1990. Social and ecological models for the Middle Stone Age in southern Africa. In: Mellars, P.A. (Ed.), The Emergence of Modern Humans. Edinburgh University Press, Edinburgh, pp. 3e33. Avery, G., Cruz-Uribe, K., Goldberg, P., Grine, F.E., Klein, R.G., Lenardi, M.J., Marean, C.W., Rink, W.J., Schwarcz, H.P., Thackeray, A.I., Wilson, M.L., 1997. The 1992e93 excavations at the Die Kelders Middle and Later Stone Age cave site, South Africa. J. Field Archaeol. 24, 263e291. Bartram, L.E., Marean, C.W., 1999. Explaining the ‘Klasies pattern:’ Kua ethnoarchaeology, the Die Kelders Middle Stone Age archaeofauna, long bone fragmentation and carnivore ravaging. J. Archaeol. Sci. 26, 9e29. Bermúdez de Castro, J.M., 1986. Dental remains from Atapuerca (Spain) I. Metrics. J. Hum. Evol. 15, 265e287. Bermúdez de Castro, J.M., 1993. The Atapuerca dental remains. New evidence (1987e1991 excavations) and interpretations. J. Hum. Evol. 24, 339e371. Bräuer, G., Deacon, H.J., Zipfel, F., 1992. Comment on the new maxillary finds from Klasies River, South Africa. J. Hum. Evol. 23, 419e422. Bräuer, G., Singer, R., 1996. The Klasies zygomatic bone: archaic or modern? J. Hum. Evol. 30, 161e165. Broadbend, B.H., Broadbend Jr., B.H., Golden, W.H., 1975. Bolton Standards of Dentofacial Developmental Growth. C.V. Mosby, St. Louis. Churchill, S.E., Pearson, O.M., Grine, F.E., Trinkaus, E., Holliday, T.W., 1996. Morphological affinities of the proximal ulna from Klasies River Main Site: archaic or modern? J. Hum. Evol. 31, 213e237. Cunningham, D., 1908. The evolution of the eyebrow region of the forehead, with reference to the excessive supraorbital development in the Neanderthal race. Trans. R. Soc. Edinb. 46, 243e310. Deacon, H.J., 1989. Late Pleistocene palaeoecology and archaeology in the southern Cape, South Africa. In: Mellars, P.A., Stringer, C.B. (Eds.), The Human Revolution: Behavioural and Biological Perspectives on the Origins of Modern Humans. Edinburgh University Press, Edinburgh, pp. 547e564. Deacon, H.J., 1992. Southern Africa and modern human origins. Philos. Trans. R. Soc. Lond. B 337, 177e183. Deacon, H.J., 2001. Guide to Klasies River 2001. http://academic.sun.ac.za/ archaeology/krguide2001.pdf, pp. 1e12. Deacon, H.J., 2008. The context of the 1967e8 sample of human remains from Cave 1 Klasies River Main Site. S. Afr. Archaeol. Soc. Goodwin Ser. 10, 143e149. Deacon, H.J., Geleijnse, V.B., 1988. The stratigraphy and sedimentology of the Main Site sequence, Klasies River, South Africa. S. Afr. Archaeol. Bull. 43, 5e14. Deacon, H.J., Wurz, S., 2001. Middle Pleistocene populations and the emergence of modern behaviour. In: Barham, L., Robson Brown, K. (Eds.), Human Roots e Africa and Asia in the Middle Pleistocene. Western Academic and Specialist Press, Bristol, pp. 55e63. Deacon, H.J., Talma, A.S., Vogel, J.C., 1988. Biological and cultural development of Pleistocene people in an Old World southern continent. In: Prescott, J.R. (Ed.), Early Man in the Southern Hemisphere. Department of Physics and Mathematical Physics, University of Adelaide, Adelaide, pp. S23eS31. d’Errico, F., Moreno, R.G., Rifkin, R.F., 2012. Technological, elemental and colorimetric analysis of an engraved ochre fragment from the Middle Stone Age levels of Klasies River Cave 1, South Africa. J. Archaeol. Sci. 39, 942e952. Drennan, M.R., 1938a. Archaeology of the Oakhurst Shelter, George. Part III. The cave-dwellers. Trans. R. Soc. S. Afr. 25, 259e280. Drennan, M.R., 1938b. Archaeology of the Oakhurst Shelter, George. Part IV. The children of the cave-dwellers. Trans. R. Soc. S. Afr. 25, 281e293. Dreyer, T.F., 1933. The archaeology of the Matjes River Rock Shelter. Trans. R. Soc. S. Afr. 21, 187e209. Dusseldorp, G.L., 2010. Prey choice during the South African Middle Stone Age: avoiding dangerous prey or maximizing returns? Afr. Archaeol. Rev. 27, 107e133. Faith, J.T., 2008. Eland, buffalo, and wild pigs: were Middle Stone Age humans ineffective hunters? J. Hum. Evol. 55, 24e36. Feathers, J.K., 2002. Luminescence dating in less than ideal conditions: case studies from Klasies River Main Site and Duinefontein, South Africa. J. Archaeol. Sci. 29, 177e194. Frayer, D.W., Wolpoff, M.H., Thorne, A.G., Smith, F.H., Pope, G.G., 1993. Theories of modern human origins: the paleontological test. Am. Anthropol. 95, 14e50. Goodwin, A.J.H., 1938. The archaeology of the Oakhurst Shelter, George. Trans. R. Soc. S. Afr. 25, 229e324. Grine, F.E., 1984. The deciduous dentition of the Kalahari San, the South African Negro and the South African Plio-Pleistocene hominids. Ph.D. Dissertation, University of the Witwatersrand. Grine, F.E., 1986. Anthropological aspects of the deciduous teeth of South African Blacks. In: Singer, R., Lundy, J.K. (Eds.), Variation, Culture and Evolution in African Populations. University of the Witwatersrand Press, Johannesburg, pp. 47e83. Grine, F.E., 1998. Additional human fossils from the Middle Stone Age of Die Kelders Cave 1, South Africa: 1995 excavation. S. Afr. J. Sci. 94, 229e235. Grine, F.E., 2000. Middle Stone Age human fossils from Die Kelders Cave 1, Western Cape Province, South Africa. J. Hum. Evol. 38, 129e145.

758

F.E. Grine / Journal of Human Evolution 63 (2012) 750e758

Grine, F.E., Henshilwood, C.S., 2002. Additional human remains from Blombos Cave, South Africa: (1999e2000 excavations). J. Hum. Evol. 42, 293e302. Grine, F.E., Henshilwood, C.S., Sealy, J.C., 2000. Human remains from Blombos Cave, South Africa: (1997e1998 excavations). J. Hum. Evol. 38, 755e765. Grine, F.E., Klein, R.G., Volman, T.P., 1991. Dating, archeology and human fossils from the Middle Stone Age levels of Die Kelders, South Africa. J. Hum. Evol. 21, 363e395. Grine, F.E., Pearson, O.M., Klein, R.G., Rightmire, G.P., 1998. Additional human fossils from Klasies River Mouth, South Africa. J. Hum. Evol. 35, 95e107. Groves, C.P., 1998. The proximal ulna from Klasies River. J. Hum. Evol. 34, 119e121. Grün, R., Shackleton, N.J., Deacon, H.J., 1990. Electron-spin-resonance dating of tooth enamel from Klasies River Mouth. Curr. Anthropol. 31, 427e432. Habgood, P.J., 1989. The examination of regional features on middle and early Late Pleistocene sub-Saharan African hominids. S. Afr. Archaeol. Bull. 44, 17e22. Hoffman, A., 1958. New excavations in the Matjes River Rock Shelter. S. Afr. Mus. Assoc. Bull. 6, 342e348. Jacobson, A., 1982. The Dentition of the South African Negro. Higginbotham, Anniston. Klein, R.G., 1975. Middle Stone Age maneanimal relationships in southern Africa: evidence from Die Kelders and Klasies River Mouth. Science 190, 265e267. Klein, R.G., 1976. The mammalian fauna of the Klasies River Mouth sites, southern Cape Province, South Africa. S. Afr. Archaeol. Bull. 21, 75e98. Klein, R.G., 1989. Why does skeletal part representation differ between smaller and larger bovids at Klasies River Mouth and other archeological sites? J. Archaeol. Sci. 16, 363e381. Klein, R.G., Cruz-Uribe, K., 1996. Exploitation of large bovids and seals and Middle and Later Stone Age sites in South Africa. J. Hum. Evol. 31, 315e334. Klein, R.G., Cruz-Uribe, K., Milo, R.G., 1999. Skeletal part representation in archaeofaunas: comments on “Explaining the ‘Klasies Pattern’: Kua ethnoarchaeology, the Die Kelders Middle Stone Age archaeofauna, long bone fragmentation and carnivore ravaging” by Bartram and Marean. J. Archaeol. Sci. 26, 1225e1234. L’Abbé, E.N., Loots, M., Keough, N., 2008. The Matjes River Rock Shelter: a description of the skeletal assemblage. S. Afr. Archaeol. Bull. 63, 61e68. Lam, Y.M., Pearson, O.M., Smith, C.M., 1996. Chin morphology and sexual dimorphism in the fossil hominid mandible sample from Klasies River Mouth. Am. J. Phys. Anthropol. 100, 545e557. Louw, J.T., 1960. Prehistory of the Matjes River Rock Shelter. Memoir No. 1. National Museum, Bloemfontein. McBrearty, S., Brooks, A.S., 2000. The revolution that wasn’t: a new interpretation of the origin of modern human behavior. J. Hum. Evol. 39, 453e563. McCall, G.S., 2006. Multivariate perspectives on change and continuity in the Middle Stone Age lithics from Klasies River Mouth, South Africa. J. Hum. Evol. 51, 429e439. Millard, A.R., 2008. A critique of the chronometric evidence for hominid fossils. I. Africa and the Near East 500e50 ka. J. Hum. Evol. 54, 848e874. Milo, R.G., 1998. Evidence for hominid predation at Klasies River Mouth, South Africa, and its implications for the behaviour of early modern humans. J. Arcahaeol. Sci. 25, 99e133. Moss, M.L., Chase, P.S., 1966. Morphology of the Liberian Negro deciduous teeth. I. Odontometry. Am. J. Phys. Anthropol. 24, 215e230.

Pearson, O.M., Churchill, S.E., Grine, F.E., Trinkaus, E., Holliday, T.W., 1998. Multivariate analyses of the hominid ulna from Klasies River Mouth. J. Hum. Evol. 34, 653e656. Pearson, O.M., Grine, F.E., 1997. Re-analysis of the hominid radii from Cave of Hearths and Klasies River Mouth, South Africa. J. Hum. Evol. 32, 577e592. Ponce de León, M.S., Zollikofer, C.P.E., 2001. Neanderthal cranial ontogeny and its implications for late hominid diversity. Nature 412, 534e538. Rightmire, G.P., Deacon, H.J., 1991. Comparative studies of Late Pleistocene human remains from Klasies River Mouth, South Africa. J. Hum. Evol. 20, 131e156. Rightmire, G.P., Deacon, H.J., 2001. New human teeth from the Middle Stone Age deposits at Klasies River, South Africa. J. Hum. Evol. 41, 535e544. Rightmire, G.P., Deacon, H.J., Schwartz, J.H., Tattersall, I., 2006. Human foot bones from Klasies River Main Site, South Africa. J. Hum. Evol. 50, 96e103. Royer, D.F., Lockwood, C.A., Scott, J.E., Grine, F.E., 2009. Size variation in early human mandibles and molars from Klasies River, South Africa: comparison with other Middle and Late Pleistocene assemblages and with modern humans. Am. J. Phys. Anthropol. 140, 312e323. Shackleton, N.J., 1982. Stratigraphy and chronology of the Klasies River Mouth deposits: oxygen isotope evidence. In: Singer, R., Wymer, J. (Eds.), The Middle Stone Age at Klasies River Mouth in South Africa. University of Chicago Press, Chicago, pp. 194e199. Singer, R., Wymer, J.J., 1982. The Middle Stone Age at Klasies River Mouth in South Africa. University of Chicago Press, Chicago. Smith, F.H., 1992. Models and realities in modern human origins: the African fossil evidence. Philos. Trans. R. Soc. B 337, 243e250. Smith, F.H., Falsetti, A.B., Donnelly, S.M., 1989. Modern human origins. Yearb. Phys. Anthropol. 32, 35e68. Stringer, C.B., Bräuer, G., 1994. Methods, misreading, and bias. Am. Anthropol. 96, 416e424. Tobias, P.V., 1967. The Cranium and Maxillary Dentition of Australopithecus (Zinjanthropus) boisei. Olduvai Gorge, vol. 2. Cambridge University Press, Cambridge. Villa, P., Soriano, S., Teyssandier, N., Wurz, S., 2010. The Howiesons Poort and MSA III at Klasies River main site, Cave 1A. J. Archaeol. Sci. 37, 630e655. Vogel, J.C., 2001. Radiometric dates for the Middle Stone Age in South Africa. In: Tobias, P.V., Raath, M.A., Moggi-Cecchi, J., Doyle, G.A. (Eds.), Humanity from African Naissance to Coming Millennia e Colloquia in Human Biology and Palaeoanthropology. Firenze University Press, Firenza, pp. 261e268. Volman, T.P., 1984. Early prehistory of southern Africa. In: Klein, R.G. (Ed.), Southern African Prehistory and Palaeoenvironments. Balkema, Rotterdam, pp. 169e220. White, T.D., 1987. Cannibals at Klasies? Sagittarius 2, 7e9. Wurz, S., 1999. The Howiesons Poort backed artefacts from Klasies River: an argument for symbolic behavior. S. Afr. Archaeol. Bull. 54, 38e50. Wurz, S., 2002. Variability in the Middle Stone Age lithic sequence, 115,000e60,000 years ago at Klasies River, South Africa. J. Archaeol. Sci. 29, 1001e1015. Wurz, S., 2008. Modern behavior at Klasies River. S. Afr. Archaeol. Soc. Goodwin Ser. 10, 150e156. Wurz, S., 2010. Middle Stone Age tools from Klasies River main site, conventions and symbolic cognition. In: Nowell, A., Davidson, I. (Eds.), The Cutting Edge: Stone Tools and the Evolution of Cognition. Colorado University Press, Boulder, pp. 135e158. Wurz, S., le Roux, N.J., Gardner, S., Deacon, H.J., 2003. Discriminating between the end products of the earlier Middle Stone Age sub-stages at Klasies River using biplot methodology. J. Archaeol. Sci. 30, 1107e1126.