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ORIGINAL ARTICLE Rock Hyraxes (Procavia capensis) from Middle Stone Age Levels at Blombos Cave, South Africa Shaw Badenhorst & Karen L. van Niekerk & Christopher S. Henshilwood Published online: 28 March 2014 # Springer Science+Business Media New York 2014 Abstract The 100 ka Middle Stone Age levels at Blombos Cave, southern Cape, South Africa, contain numerous rock hyrax (Procavia capensis) remains. It is often ambiguous to interpret rock hyrax remains from archaeological deposits deriving from cave and shelter sites in southern Africa as the agent or agents of accumulation may be difficult to establish. In this paper, the different taphonomic signatures separating anthropogenic from natural accumulations at Blombos Cave are considered. The analysis indicates that although a few specimens show evidence for raptor and carni- vore accumulation, there is also substantial evidence that suggests humans preyed on these small mammals during different times of the year. Résumé Dans la grotte de Blombos (Cap-Occidental, Afrique du Sud), les niveaux Middle Stone Age datés denviron 100 ka se caractérisent notamment par labondance des restes de Daman du Cap (Procavia capensis). L interprétation de labondance du Daman dans les sites archéologiques sud-africains est souvent problématique, car il est Afr Archaeol Rev (2014) 31:2543 DOI 10.1007/s10437-014-9154-7 S. Badenhorst (*) Department of Archaeozoology, Ditsong National Museum of Natural History (former Transvaal Museum), 432 Paul Kruger St, Pretoria 0001, South Africa e-mail: [email protected] S. Badenhorst Department of Anthropology and Archaeology, University of South Africa, PO Box 392, UNISA 0003, South Africa K. L. van Niekerk : C. S. Henshilwood Institute for Archaeology, History, Culture and Religion, University of Bergen, Postbox 7805, 5020 Bergen, Norway K. L. van Niekerk e-mail: [email protected] C. S. Henshilwood e-mail: [email protected] C. S. Henshilwood Evolutionary Studies Institute, University of the Witwatersrand, Private Bag 3, Wits 2050, South Africa

Rock Hyraxes (Procavia capensis) from Middle Stone Age Levels at Blombos Cave, South Africa

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ORIGINAL ARTICLE

Rock Hyraxes (Procavia capensis) fromMiddle Stone AgeLevels at Blombos Cave, South Africa

Shaw Badenhorst & Karen L. van Niekerk &

Christopher S. Henshilwood

Published online: 28 March 2014# Springer Science+Business Media New York 2014

Abstract The ∼100 ka Middle Stone Age levels at Blombos Cave, southern Cape,South Africa, contain numerous rock hyrax (Procavia capensis) remains. It is oftenambiguous to interpret rock hyrax remains from archaeological deposits deriving fromcave and shelter sites in southern Africa as the agent or agents of accumulation may bedifficult to establish. In this paper, the different taphonomic signatures separatinganthropogenic from natural accumulations at Blombos Cave are considered. Theanalysis indicates that although a few specimens show evidence for raptor and carni-vore accumulation, there is also substantial evidence that suggests humans preyed onthese small mammals during different times of the year.

Résumé Dans la grotte de Blombos (Cap-Occidental, Afrique du Sud), les niveauxMiddle Stone Age datés d’environ 100 ka se caractérisent notamment par l’abondancedes restes de Daman du Cap (Procavia capensis). L’interprétation de l’abondance duDaman dans les sites archéologiques sud-africains est souvent problématique, car il est

Afr Archaeol Rev (2014) 31:25–43DOI 10.1007/s10437-014-9154-7

S. Badenhorst (*)Department of Archaeozoology, Ditsong National Museum of Natural History (former TransvaalMuseum), 432 Paul Kruger St, Pretoria 0001, South Africae-mail: [email protected]

S. BadenhorstDepartment of Anthropology and Archaeology, University of South Africa, PO Box 392, UNISA 0003,South Africa

K. L. van Niekerk : C. S. HenshilwoodInstitute for Archaeology, History, Culture and Religion, University of Bergen, Postbox 7805,5020 Bergen, Norway

K. L. van Niekerke-mail: [email protected]

C. S. Henshilwoode-mail: [email protected]

C. S. HenshilwoodEvolutionary Studies Institute, University of the Witwatersrand, Private Bag 3, Wits 2050, South Africa

particulièrement difficile en contexte de grotte ou d’abri de déterminer le ou les agent(s)responsable(s) de l’accumulation de ses restes. Dans cette contribution, les différentessignatures taphonomiques utilisables pour distinguer accumulations naturelles etanthropiques sont considérées. Nos analyses indiquent que, même si quelques restesprésentent des traces évidentes d’action par un rapace ou un carnivore, de nombreuxarguments suggèrent que ce petit mammifère représentait une proie de choix pour lesgroupes humains, et ce à différentes périodes de l’année.

Keywords Middle StoneAge . Blombos Cave . Southern Cape . Rock hyrax . Procaviacapensis

Introduction

The rock hyrax (Procavia capensis) occurs throughout most of Africa. It is a small, rabbit-like mammal with an average weight of 3.6 kg for females and 4 kg for males. Rockhyraxes feed on a variety of herbaceous plants (Olds and Shoshani 1982). They havemany predators (Burton and Burton 2002: 1271). They live in rocky areas, where cavesand shelters are common (cf. Ditlhogo and Setshogo 2001). Due to the variety ofpredators, their agents of accumulation in archaeological and paleontological depositsfrom cave and shelter sites are often difficult to determine because raptors, carnivores andhominins make use of shelters (Plug et al. 2003). In this paper, we review themain lines ofevidence used to infer different agents of accumulation of rock hyrax remains in archae-ological and paleontological deposits. We test these arguments against the sample fromthe ∼100,000-year-old Middle Stone Age levels at Blombos Cave (BBC) in the southernCape of South Africa, where large numbers of rock hyrax remains have been excavated.

Natural Accumulations of Rock Hyrax Remains

Here, we review evidence from previous studies for the agents that may have beenresponsible for natural accumulations of rock hyraxes in shelters and cave sites.

Raptors

Remains of rock hyraxes can accumulate in caves or shelters due to raptor predation.Several species of eagle, such as black eagles (Aquila verreauxi), crowned eagles(Stephanoaetus coronatus), and martial eagles (Polemaetus bellicosus) prey on rockhyraxes (Fourie 1983). However, eagles do not enter caves to nest. Black eaglesconstruct large roosts, about 1.5–2 m in diameter, usually on a ledge of a cliff (Brain1981; Watson 2004: 65), and sometimes in trees (Maclean 1985: 117). Crowned andmartial eagles roost in trees (Maclean 1985). Bones that have been accumulated byblack eagles are usually found in direct association with nests (cf. De Ruiter et al. 2010:132). Eagles nesting on a ledge or tree above a cave could result in the bones from theroost becoming incorporated in the cave deposit. According to Cruz-Uribe and Klein(1998: 143), rock hyraxes are too large to be introduced at cave and shelter sites by anyraptor smaller than a Cape eagle owl (Bubo capensis). In southern Africa, the barn owl(Tyto alba) is one of the most common owls that inhabits caves and shelters, but usually

26 Afr Archaeol Rev (2014) 31:25–43

preys on smaller rodents and is unlikely the contributor of hyrax remains (Maclean1985). Some species of eagle owls such as the spotted eagle owl (Bubo africanus)frequent caves, but their prey size seldom exceeds 140 g (Andrews 1990). The gianteagle owl (Bubo lacteus) preys on hyraxes but rarely inhabits caves. The Cape eagleowl preys on hyraxes, frequents caves and, of the owl species, is the most likelycontributor of hyrax remains (Brain 1981: 123–124).

Raptor-introduced bones may be identified by modifications such as beak damage(Brain 1981), regurgitated hair, ‘fresh’ bone (Shaffer and Neely 1992; Plug 1993;Badenhorst 2003), dried flesh, and partially articulating elements (Plug 1992). Hyraxremains left by the Cape eagle owl typically consists of crania discarded with the pellets(Brain 1981: 123–124). Hare crania show characteristic damage to the calvariae andnasals when consumed by Cape eagle owls, and similar damage could be expected oncrania from hyraxes, comparable in size to hares (Brain 1981: 123–124).

Black eagles feeding on hyraxes leave distinct taphonomic signatures, as they do notconsume the entire skull or large bones such as pelvises and limb bones (Gargett 1972).Bone samples from eagle roosts have a striking abundance of mandibles and maxillaecompared to post-cranial remains. Hind limb elements (pelvis, femur, tibia) outnumberthose from the forelimb (scapula, humerus, radius, ulna) in such collections (Cruz-Uribeand Klein 1998: 139–140). Eagles typically open the braincase of hyraxes from the backor side to remove the brain, and sometimes the entire calvaria (skullcap or upper part ofthe cranium) is consumed, leaving a distinct damage pattern on the skull (Brain 1981).Some damage on skulls, such as detached occipital bones, could be a post-depositionalphenomenon (Cruz-Uribe and Klein 1998: 137). Eagle-accumulated faunal assemblagestend to have a composition of species which falls within a very narrow weight factor (cf.De Ruiteret al. 2010: 133), and rarely contain bones from very young (neonate) hyraxes(Cruz-Uribe and Klein 1998). In a recent study, De Ruiter et al. (2010), based on thecriteria of Cruz-Uribe and Klein (1998), suggested that the rock hyrax remains fromKromdraai A, a Pleistocene site in Gauteng, South Africa, may have been accumulatedby eagles. This sample consists of 51 cranio-dental fragments and nine post-crania(eight of which are articulating vertebrae) (Brain 1981: 254).

Owls swallow bones, and subsequently regurgitate these as pellets, whereas eaglesmainly dismember their prey. Swallowed bones are partially digested by strong stom-ach acids before they are regurgitated in pellets (Cruz-Uribe and Klein 1998: 137). Theregurgitated pellets accumulate on the floor of the cave or shelter, and the bones thereincan become incorporated into archaeological deposits, either in distinct pockets orscattered due to post depositional processes (Brain 1981: 118; Levinson 1982).

Carnivores

Carnivores, in particular caracals and leopards, prey on rock hyraxes (Bothma and Du1971; Brain 1981; Fourie 1983; Norton et al. 1986; Palmer and Fairall 1988). Leopards(Panthera pardus) consume the entire post-cranial skeleton, leaving crania that showdistinctive modifications. Leopards shear off the braincase and posterior part of themandible to remove the brain and tongue, leaving ragged, tooth-marked edges oncrania and mandibulae. The remaining maxilla and mandible are rejected (Brain 1981;Plug and Keyser 1994). While leopards also swallow hyrax bones (e.g., Rautenbach2010), scats are usually located at strategically selected trees (Rautenbach 1982: 148)

Afr Archaeol Rev (2014) 31:25–43 27

and not in caves or shelters. Caracal scats are usually dropped in open terrain (e.g.,Braczkowski et al. 2012) and they seldom use caves or shelters; hence, they areunlikely to have been significant contributors of hyrax remains in these sites.

Plio-Pleistocene hyrax remains have been recovered from a number of sites insouthern Africa: (a) At Swartkrans, leopards were likely the main agent of accumula-tion, with some contribution by owls and eagles, of the P. capensis, Procaviatransvaalensis and Procavia antique bones (but see McMahon and Thackeray 1994),which are dominated by cranio-dental remains (Watson 2004); (b) At Haasgat Cave,only hyrax crania are present and the chew marks on these suggest that carnivores werethe predators (Plug and Keyser 1994).

Natural Deaths

Hyraxes die naturally in caves and shelters from starvation, diseases and injury. Theseremains can become incorporated into archaeological deposits andmay contain articulatedelements and complete long bones and a ‘fresh’ appearance (Badenhorst 2003). Studies ofnatural hyrax death assemblages have shown that these tend to be dominated by youngindividuals (<2 years of age) (Fourie 1983). The relatively high mortality of young rockhyraxes is an indication of the large proportion of juvenile individuals in a population. Astudy in the Karoo of South Africa found that juveniles (<2 years of age) make up 35% ofthe entire population, whereas in the Matobo National Park of Zimbabwe, following thedrought of 1991–1992, juveniles comprised 25 % of the population (Barry 1994). In theTsitsikamma National Park of South Africa, Fairall and Crawford (1983) found thatindividuals of less than 1 year of age made up 55–56 % of the total populations.

At Collingham Shelter, a Later Stone Age cave in KwaZulu-Natal, South Africa,nearly all the rock hyrax remains from Phase A, dating to about 1260 BP, are ‘fresh’ inappearance, with pieces of flesh still attached to some of the elements. The samplecontains complete skulls and partially articulated elements, and clearly did not relate tothe rest of the fauna. Based on tooth wear and eruption, this sample consists of 15(63 %) juveniles and sub-adults, and 9 (37 %) adults and mature individuals (Plug1992). The front and hind limbs (cf. Cruz-Uribe and Klein 1998) are nearly equallyrepresented, and crania are not particularly common (Table 1). These specimens mostlikely represent natural deaths at the site (Plug 1992).

Anthropogenic Accumulations of Rock Hyrax Remains

Various ethnographic and historical sources have documented hunting rock hyraxes insouthern Africa in the recent past (e.g., Raven-Hart 1971: 459, 482), and hyraxes havefeatured in hunter-gatherer folklore from parts of southern Africa (Bleek 1929). Their

Table 1 Rock hyraxes fromCollingham Shelter, phase A(NISP)

Elements NISP Percentage (%)

Crania 54 12

Teeth 297 67

Front limb 56 13

Hind limb 39 9

28 Afr Archaeol Rev (2014) 31:25–43

remains are often found at hunter-gatherer and farming sites (Plug and Badenhorst2001), and they are easily killed or captured by stoning (Bousman 2005: 218) or snares(Wadley 2010).

Taphonomic signatures on animal bones can indicate the presence of humans in cavesand shelters. Bones that show signs of butchering (cut and chop marks) are directindicators of human activities (e.g., Lyman 1994), but a lack of these marks does notcontradict a human presence. Ethnographic studies of hunter-gatherers show that smallanimals are usually brought intact to camps (e.g., Lee 1979; Harako 1981; Bartram et al.1991) and that little or no butchering is required to reduce the carcass tomoremanageablepieces, resulting in fewer cut and chop marks visible on bones (cf. Szuter 1991; Jones1993; Fernández-Jalvo et al. 1999). Bones from large animals that were butchered mayalso lack conspicuous evidence of cut and chop marks (Parsons and Badenhorst 2004).Butchery marks on small animals such as hyraxes are more easily seen under magnifi-cation. When not applied, the tell-tale signs of consumption by humans may be difficultto ascertain (Charles and Jacobi 1994, but see Lloveras et al. 2009; Reynard 2011).According to Cruz-Uribe and Klein (1998: 143), the presence of tortoises, seals and otherlarge animals, which were definitely accumulated by people along with hyraxes, is anindicator that the hyraxes are of anthropogenic origin in Stone Age deposits.

Hyraxes accumulated by humans have a large number of post-cranial remainsrelative to crania, and forelimb bones are more common than those from the hind limbin instances of poor preservation or leaching (Cruz-Uribe and Klein 1998: 140). In suchinstances, the more robust distal humerus accounts for the higher proportion offorelimbs. Other studies have found similar discrepancies in forelimb versus hind limbratios in human accumulated deposits. For example, in the Middle and Later Stone Agedeposits of Pomongwe Cave in Zimbabwe, hyrax remains are attributed to humancollectors, and post-crania NISP (n=719) outnumber crania (n=473), and forelimbs(n=332) outnumber the hind limbs (n=169) (Brain 1981).

Evidence for burning on bones in cave assemblages may indicate human activity (Brainand Sillen 1988). Dunemole rats that have been roasted on an open fire show characteristicburning patterns on the incisors and pre-maxillae (Henshilwood 1997); it is possible thatsimilar patterns would occur on hyrax pre-maxillae and incisors if cooked in the same way.

Blombos Cave

Blombos Cave (BBC) is a (∼55 m2) cave situated in a steep wave-cut calcrete cliff, 100 mfrom the Indian Ocean and 34.5 m above modern sea level (34°25′S, 21°13′E; Fig. 1). Thecave contains archaeological remains of both LSA and MSA occupations. The calcareousenvironment is at least partially responsible for the generally good preservation of theorganic remains that have been recovered (Henshilwood 2008a; Henshilwood et al. 2001b).

A hiatus indicated by a layer of undisturbed aeolian sand (DUN) dated to ∼70 kaseparates the LSA from the underlying MSA layers (Fig. 2; Jacobs et al. 2006). TheMSA is divided into four phases with a number of discrete layers within each phase.These phases are named, from youngest to oldest, M1, upper M2, lower M2 and M3.The MSA layers have been dated using a number of methods including thermolumi-nescence (TL), optically stimulated luminescence (OSL), electron spin resonance(ESR) and uranium-thorium (U/Th) (Jones 2001; Henshilwood et al. 2002, 2011;Jacobs et al. 2003a, b; Tribolo 2003; Tribolo et al. 2006).

Afr Archaeol Rev (2014) 31:25–43 29

The M1 phase has an age of ∼73 ka (Jacobs et al. 2006) and the upper M2 has anage of ∼77 ka (Jacobs et al. 2006; Tribolo et al. 2006; Henshilwood et al. 2011). Thesetwo phases contain artefact types associated with the Still Bay techno-tradition. Thelower M2 layers have an age of ∼85–82 ka (Jacobs et al. 2006). The age for the M3phase is between 101 and 94 ka (Henshilwood et al. 2011).

The M1 and upper M2 phases contain more than 400 bifacial points, the fossiledirecteur of the Still Bay techno-tradition. Approximately half of the silcrete bifacialpoints were first heat-treated and then knapped using a pressure-flaking technique (Villaet al. 2009;Mourre et al. 2010). Beads made fromNassarius kraussianus shells, and bonetools, and engraved bone and ochre pieces were recovered from these phases(Henshilwood and Sealy 1997; d’Errico et al. 2001; Henshilwood et al. 2001a, b, 2002,2004, 2009; d’Errico and Henshilwood 2007). In the lower M2 phase, bone tools, bifacialpoints and shell beads are absent (Henshilwood et al. 2001b). A few ground and scrapedochre pieces are present, but none with deliberate engravings (Henshilwood et al. 2009).

A characteristic of the M3 phase is the dense shell deposits in the upper levels andmany basin-shaped hearths. Shell and bone are well preserved and the deposits arerelatively undisturbed. Lithics are common and the raw materials used include fine-grained silcrete and hornfels. Worked and unworked ochre are common and eight ochrepieces have deliberately engraved cross-hatched, Y-shaped or crenulated designs(Henshilwood et al. 2001b, 2009; Watts 2009). Two in situ ochre-processing toolkitswere found in the lowermost layer (CP). These toolkits include two abalone shells(Haliotis midae) in which an ochre-rich compound was mixed and stored(Henshilwood et al. 2011). The Later and Middle Stone Age fauna from BBC isdominated by small game animals such as rock hyrax, Cape dune mole rat, seals andsmall antelopes including steenbok and Cape grysbok (Henshilwood et al. 2001b).

Fig. 1 Location of BlombosCave

30 Afr Archaeol Rev (2014) 31:25–43

Rock Hyraxes

Rock hyraxes have a short and well-defined period of sexual activity triggered byphotoperiod, and not annual cycles in rainfall or temperature. The period of sexual activityvaries in southern Africa, and it is progressively later at lower latitudes. In the Western,Northern and Eastern Cape provinces, rock hyraxes currently give birth between Septem-ber and October. In the southern Cape, these birth peaks coincide with the local vegetation(fynbos) being in prime condition following the winter rains (Smithers 1983: 544, Skinnerand Chimimba 2005). A study at De Hoop Nature Reserve, which is close to BBC, foundthat captive females gave birth during the end of September and the first half of October(Millar 1971). As birth is dictated by photoperiod rather than temperature and rainfall, weassume in this study that the birth season of extant rock hyraxes was the same during theMSA, and that the modern rate of teeth eruption was similar during the MSA.

Eruption stages of teeth from neonate and juvenile individuals are useful fordetermining age classes at death, which in turn can indicate the season of occupation

Fig. 2 South section showing the stratigraphy and ages of the Middle Stone Age layers at Blombos Cave

Afr Archaeol Rev (2014) 31:25–43 31

of archaeological sites (e.g., Parkington and Poggenpoel 1971; Parkington 2001;Henshilwood 2008a). Rock hyraxes were common in the LSA and MSA levels exca-vated before 2001 at Blombos Cave (Henshilwood et al. 2001b), but the age classeswere not reported. The aging categories for teeth as established by Steyn and Hanks(1983) were applied. The age classes consider eruption stage, the presence or absence ofdeciduous and permanent teeth, and wear. In this study, we consider the eruptionsequences for animals less than 12 months old (classes I–VII), as only these can informon the month of death. Mandibles and maxillae, with several teeth present, wereanalysed to obtain precise ages at time of death. No isolated teeth were used. Althoughother aging classifications for hyraxes exist (see Steyn and Hanks 1983; Cruz-Uribe andKlein 1998), and while we recognize the slight discrepancies between these agingsystems, we opted for the most recent aging classifications (Steyn and Hanks 1983).

Materials and Methods

Hyrax remains from the CH to CL levels (Fig. 2) at BBC of the M3 phase wereanalysed using methods described by Driver (2005). During analysis, some bonespecimens lacked sufficient diagnostic morphological characteristics to allow speciesidentification. Many of these specimens were identified as ‘possible hyrax’ (cf.P. capensis), common in faunal analyses (e.g., Reitz and Wing 1999). We combinedboth firm and possible identifications of hyraxes, unless otherwise indicated, assumingthat the latter are in fact rock hyraxes.

Taphonomy was recorded using naked-eye observations due to time constrains. It isnot always feasible to subject every specimen to microscopic analysis (Reynard 2011).Breakage patterns were recorded as either spiral or transverse fractures. Breaks ofproximal and distal ends, if present, were counted separately. For example, a long bonewith an intact proximal end but a spiral break on the distal end was only counted as asingle spiral break. For the breakage patterns of long bones, only humeri, radii, femora,tibiae and metapodia were considered. For skeletal part representation, the scapulae,humeri, radii and ulnae were used for the forelimb, and the pelvis, femora and tibiae forthe hind limb (following Cruz-Uribe and Klein 1998), unless otherwise indicated.

The most basic form of quantification used in this study was the Number ofIdentified Specimens (NISP). For skeletal part representation, previous studies havefavoured the Minimum Number of Individuals (MNI) (Cruz-Uribe and Klein 1998) andthe Minimum Number of Elements (MNE) (De Ruiter et al. 2010). Due to theproblematic nature of the latter two methods (e.g., Reitz and Wing 1999; Lyman2008), only NISPs were used, except to quantify the minimum number of individualmandibles and maxillae for aging purposes.

It can be assumed that samples of naturally accumulated small mammals would bedominated by complete long bones (Badenhorst 2008). Long bone (humeri, tibiae andfemora only) fragmentation of rock hyraxes fromBlombos Cavewas compared to varioussamples of similarly sized cottontails (Sylvilagus sp.) and jackrabbits (Lepus sp.) from theopen-air human occupation sites Albert Porter Pueblo and Pueblo Bonito in the AmericanSouthwest (Badenhorst 2008). A previous study of hyrax remains fromMiddle and LaterStone Age sites in South Africa also used comparative jackrabbit samples from theAmerican Southwest (Cruz-Uribe and Klein 1998). In addition, rock hyrax, springhare

32 Afr Archaeol Rev (2014) 31:25–43

(Pedetes capensis) and hare (Lagomorph) samples from Iron Age farming and historicalsites in southern Africa were included. These samples are from Nanda (Plug 1993),Bosutswe (Plug 1996), Pont Drift, Schroda (Plug and Voigt 1985), Thulamela (Plug1997) and Doornbult (Badenhorst and Boshoff forthcoming). The small taxa from thesesites were most likely accumulated by humans and are useful proxies for this study.

To determine whether the rock hyrax sample is dominated by adults, juveniles orneonates, the proximal and distal fusion of post-crania was used, as few mandibulae,maxillae and isolated teeth could be aged. Each long bone received a two-letter code torecord the proximal and distal fusion. For example, a complete, fully fused femur wasrecorded as ‘FF’, indication that both articulation ends are fully fused. The codes wereadded for the long bones.

Results

The large mammal remains from levels CH to CL consists of 3,783 identified speci-mens, including a variety of large, medium and small animals. Of these, 1,059, or 28 %,are rock hyrax remains (including both P. capensis and cf. P. capensis). Rock hyraxesare the single most common taxon in the sample. No concentrations of rock hyraxes areevident in the levels, and they occur throughout the deposits in each of the layers(Table 2). Layer CH-CI is the thickest, and has produced the largest quantity of fauna,including rock hyrax remains. No articulating elements, which may suggest naturaldeposition (Plug 1992), were found.

Of the total rock hyrax sample, 8 % is burnt (Table 3). Amongst the burnt specimensis a mandible from Layer CH-CI that has light brown scorching on the corpusmandibulae. The anterior portion is a darker brown than the tooth row.

Few natural and anthropogenic modifications were noted (excluding burning) on therock hyrax sample. Both human (cut and chop marks) and natural (carnivore and rodentgnawing, digested and possible beak marks) damage are present, but in all instances, invery low numbers (Table 4) amounting to a single percentage or less. No ‘fresh’specimens were noted during analysis.

Table 5 lists the skeletal part representation of rock hyraxes for all specimens perNISP, and Table 6 lists only those elements used in a previous study (Cruz-Uribe andKlein 1998) using NISP, for comparative purposes. A number of aspects are worthhighlighting from these two tables. First, some groups of elements, most notably ribsand vertebrae, and, to a lesser extent, cranial remains, are represented by very lownumbers. This is due to the difficulty in differentiating between these elements ofsimilar sized taxa, and therefore most of these elements were identified during analysis

Table 2 Rock hyraxes from the different levels at BBC (NISP)

Taxa Common Name CH-CI CJ CK CL Total

Procavia capensis Rock Hyrax 548 324 51 52 975

cf. Procavia capensis Possibly Rock Hyrax 56 23 3 2 84

Total 604 347 54 54 1,059

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as ‘indeterminate small mammals’ and excluded from this study. Second, teeth, mostlyfound as isolated specimens, are most common. However, teeth are also numerous in asingle individual and, even after fragmentation, they are identifiable. As a result, thisaspect is not unusual. Third, crania (excluding teeth) are outnumbered by post-cranialremains. Fourth, forelimbs outnumber hind limbs. Fracture patterns indicate that spiralfractures outnumber transverse breakage (Table 7). Spiral fractures indicate freshbreaks, and transverse breaks would suggest dry fractures.

Complete long bones are rare, and most are fragmented (Table 8). Metapodia are themost common complete long bone element, but as these elements are relatively short,compact and dense, they are less prone to breakage. The long-bone fragmentation ratesfor humeri, femora and tibiae (following convention in the American Southwest, seeBadenhorst 2008) of rock hyraxes from BBC were compared to rock hyraxes,springhares and hares from Nanda, Bosutswe, Pont Drift, Schroda, Thulamela andDoornbult in southern Africa as well as cottontails and jackrabbits from Albert PorterPueblo and Pueblo Bonito from the American Southwest (Table 9). Long-bone frag-mentation is similar across all taxa, and indicates a clear dominance of fragmentedremains with few if any complete long bones. At Pont Drift, the number of completelong bones is relatively high compared to other sites. This is most likely due to eitherthe small sample size, or because some springhares are natural intrusions, as they makeburrows into soft sandy soils (Rautenbach 1982: 75). Nevertheless, the general patternfor the various small mammals shows that complete long bones are rare, or completelyabsent, in samples accumulated by humans.

Based on fusion stages of all long-bone epiphyses in the sample, young animals aremore common than adults (Table 10). Teeth also show a dominance of younger

Table 3 Burnt rock hyraxes per layer (NISP)

Taxa CH-CI CJ CK CL Total

Procavia capensis 45 27 2 3 77

cf. Procavia capensis 6 3 9

Total 51 30 2 3 86

% NISP of P. capensis 8 % 9 % 4 % 6 % 8 %

Table 4 Human and natural modifications (NISP)

Modification CH-CI CJ CK CL Total NISP of P. capensis (%)

Cut and chop 1 1 <1

Carnivore 4 4 8 1

Possible beak 1 1 <1

Rodent 1 1 <1

Digested 2 2 4 <1

34 Afr Archaeol Rev (2014) 31:25–43

individuals. Tooth eruption and replacement data indicate that the rock hyraxes diedduring different months of the year (Table 11).

Discussion

Hyrax Accumulation at BBC

Hyrax remains in archaeological deposits from cave and shelter sites could have beenaccumulated by humans, carnivores or raptors, died there naturally, or by a combinationof these agents. Due to the difficulties in determining the agent(s) of accumulation,often little discussion is offered to consider the presence of hyraxes in archaeologicaldeposits (but see Cruz-Uribe and Klein 1998; De Ruiter et al. 2010), except when aclear pattern is evident.

Blombos Cave yielded abundant evidence for human occupation and activities.Thick deposits of cultural artefacts, hearths and marine shells indicate that humanswere responsible for a considerable portion of the deposits. However, people did notnecessarily live at the site continuously over millennia (cf. Henshilwood 2008a), and itis conceivable that raptors and carnivores used the shelter from time to time. Some ofthe hyraxes could have died naturally in the shelter, and their remains could have beenincorporated into the anthropogenic deposits. Whether animals or humans were re-sponsible for the hyrax remains at BBC can be open to interpretation.

The hyrax remains at BBC show limited evidence of raptor activity apart from asingle, possible beak mark on a specimen, as well as a few partially digested remains.

Table 5 Skeletal part representations of rock hyraxes (NISP) from the M3 phase at BBC

Element CH-CI CJ CK CL Total Percentage

Cranial 33 12 3 48 5

Mandible 31 20 4 5 60 6

Teeth 269 162 25 31 487 46

Remaining axial (ribs, vertebrae, sternum, sacrum) 9 7 1 1 18 2

Forelimb (scapula, humerus, radius, ulna, carpal, metacarpal) 116 66 7 7 196 19

Hind limb (pelvis, femur, patella, tibia, fibula, tarsal, metatarsal) 85 54 11 8 158 15

Remaining appendicular (metapodials, phalanges) 61 26 3 2 92 9

Total 604 347 54 54 1059 100

Table 6 BBC rock hyrax skeletal element frequencies, using only selected elements (NISP) (after Cruz-Uribeand Klein 1998)

Skeletal portion CH-CI CJ CK CL Total (%)

Front leg 92 45 5 5 147 (62)

Hind leg 48 31 6 5 90 (38)

Afr Archaeol Rev (2014) 31:25–43 35

The absence of fresh bone, the presence of rock hyrax remains in all layers togetherwith animals of small, medium and large size, the dominance of post-cranial hyraxremains (excluding teeth), and the higher frequency of forelimbs relative to hind limbssupport this argument.

The few chew marks and partially digested remains on hyraxes at BBC indicatesome carnivore activity, but the skeletal evidence suggests that these were unlikely tobe leopards. The presence and dominance of post-crania and the lack of large numbersof crania with distinct sheared-off damage suggest that leopards were not the mainagent of bone accumulation. Evidence for low carnivore activity at BBC is supportedby other studies (Henshilwood et al. 2001b; Thompson and Henshilwood 2011).

The lack of fresh bone, the overwhelming dominance of fragmented long bones andthe lack of articulated skeletons imply that hyraxes did not die naturally in BlombosCave. The dominance of young rock hyraxes at Blombos Cave, based on post-cranialfusion and teeth, is similar to other studies (Fourie 1983) of natural accumulations.However, the presence of young hyraxes is not necessarily an indication of naturalaccumulations, as humans could have specifically targeted young individuals (cf. Cruz-Uribe and Klein 1998). The skeletal part representation of rock hyraxes from BBC issimilar to the naturally accumulated sample from Collingham Shelter’s Phase A (Plug1992) (Tables 12 and 13).

Although the skeletal part representation of Blombos Cave and Collingham Shelterare similar, it may not imply that the hyraxes from the M3 phase at BBC died naturally.For example, the lack of articulating elements and absence of fresh specimens argueagainst natural hyrax deaths at BBC. The similarities in skeletal parts between BBC andCollingham Shelter may simply be a reflection of similar preservation at both sites.Both samples are dominated by teeth, a dense and numerous element which can easilybe identified despite heavy fragmentation, and both contain a high number of humeri.Distal humeri are dense and preserve exceptionally well in archaeological contexts.Despite these similarities at both sites, we suggest that predictions based only onskeletal part representation should be cautious. One example that illustrates our point

Table 7 Spiral and transverse breaks of rock hyrax long bones from BBC M3 phase (NISP)

Breakage CH-CI CJ CK CL Total Percentage

Spiral 30 11 2 – 43 63

Transverse 15 9 1 – 25 37

Table 8 Complete versusfragmented long bones of rock hy-raxes at BBC (NISP)

Element Complete (n) Incomplete (n) Complete (%)

Humeri 57 0

Radii 1 18 5

Femora 1 35 3

Tibiae 1 31 3

Metapodia 15 65 19

Total 18 206 8

36 Afr Archaeol Rev (2014) 31:25–43

is the finding of Brain (1981) that goat skeletal parts from Gobabeb villages in Namibiaresulted from human and dog activities, and similar skeletal parts from theMakapansgat site, a Plio-Pleistocene locality in South Africa, were the result of naturalagency.

Ages and Season of Death

At BBC young hyraxes dominate the assemblage (55 % juveniles and neonates basedon post-cranial fusion). At the two other sites where hyrax remains have been aged,21 % of the hyraxes (24 % of these are neonates) at Klasies River are younger than2 years, and 39 % of hyraxes are under 2 years (15 % neonate) at Die Kelders. Thesecalculations are based on tooth eruption sequences (Cruz-Uribe and Klein 1998). Poorpreservation at Die Kelders could have resulted in an underrepresentation of veryyoung hyraxes.

Few studies use animal remains to infer seasonal occupation of caves and shelters insouthern Africa during the MSA. Several large mammal species such as Cape buffalo,

Table 9 Long bone fragmentation (humeri, femora and tibiae only) for various small mammals (rock hyraxes,springhares, indeterminate hares, cottontails and jackrabbits) from Albert Porter Pueblo, Pueblo Bonito,Bosutswe, Schroda, Pont Drift, Doornbult, Nanda, Thulamela and BBC (NISP)

Taxa Site Complete Fragmented Complete (%)

Sylvilagus sp. Albert Porter Pueblo (AD 900–1275) 28 851 3

Sylvilagus sp. Pueblo Bonito (AD 1050–1105) 14 616 2

Lepus sp. Albert Porter Pueblo 2 219 1

Lepus sp. Pueblo Bonito 4 332 1

Lagomorph Bosutswea (AD 700–1450) 0 30 0

Lagomorph Schrodaa (AD 750–900) 0 85 0

Pedetes capensis Pont Drifta (AD 810–1110) 2 4 33

Pedetes capensis Schrodaa 0 6 0

Procavia capensis Doornbulta (AD twentieth century) 0 6 0

Procavia capensis Nandaa (AD sixth to seventh century) 0 5 0

Procavia capensis Schrodaa 0 6 0

Procavia capensis Thulamelaa (AD 1350–1750) 0 7 0

Procavia capensis Blombos Cave 2 123 2

a Unpublished data kept in the Archaeozoology and Large Mammal Section, Vertebrate Department, DitsongNational Museum of Natural History, Pretoria, South Africa. All data for all taxa listed in the table includeboth positive identifications and possible (e.g., cf. Procavia capensis) identifications

Table 10 Fusion of all long bones of rock hyrax at BBC (NISP)

Age CH-CI CJ CK CL Total Percentage

Neonate (amorphous) 22 5 1 28 10

Sub-Adult (unfused/just fused) 70 42 10 4 126 45

Adult (fused) 77 37 5 6 125 45

Afr Archaeol Rev (2014) 31:25–43 37

black and blue wildebeest and Cape fur seal have discrete birthing seasons that areuseful indicators of season of death, but large enough samples of these species are notfound in MSA assemblages (Klein and Cruz-Uribe 2000). Age profiles for the sealremains at BBC have not been established but the small sample size (Henshilwoodet al. 2001b; Thompson 2008) mitigates against a definitive seasonal argument.

Determining when prehistoric coastal sites were occupied is of significant interest inmapping the movement of ancient populations (Henshilwood 2008b). People maymake periodic use of shelters and caves for a variety of reasons, of which fluctuationsin the availability of animal and plant resources are probably the most important factors(Yellen 1976; Marean 1986; Reitz and Wing 1999; Parkington 2001; Jerardino 2003) inaddition to the presence of water. Little is known about the mobility of people duringthe MSA and the timing of their movements across the southern Cape landscape. Thesemovements could variously have been between coastal and inland locations, along thecoast only, or centered around the interior regions. The location of MSA archaeologicalsites containing Still Bay and Howiesons Poort type-artefacts suggests that these peoplewere able to move both along the coast and inland, but whether these movements wererelated to social activities or to take advantage of seasonally available plants andanimals is not known. Previous research on the age profiles of seal remains from thecombined MSA layers of Klasies River and Die Kelders indicate that MSA people in

Table 11 Age classes of young rock hyraxes based on tooth eruption and replacement

Age class Na/MNI Months of death Predominant season

I 0/0 September–January Summer

II 4/3 January–April Summer

III 5/4 April–July Winter

IV 3/2 July–January Summer

V 4/2 January–April Summer

VI 3/1 April–July Winter

VII 4/3 July–December Summer

VIII 3/2 n/a n/a

aN represents discrete mandibles and maxillae with intact teeth. For example, for Age Class II there are fourmandibles and maxillae with intact teeth, which represents at least three individuals

Table 12 Blombos Cave and Collingham Shelter skeletal part representation. Data from Table 5 and Plug(1992). All elements were included (NISP)

Element BBC BBC BBC BBC BBC Collingham Shelter

CH-CI CJ CK CL Total

% % % % % %

Crania 12 10 14 10 13 12

Teeth 50 52 50 61 59 67

Forelimb 22 21 14 14 18 13

Hind limb 16 17 22 16 11 9

38 Afr Archaeol Rev (2014) 31:25–43

this region did not specifically target seasonally abundant, (washed up) juvenile seals,as was the case in the Later Stone Age (LSA), and were thus not taking advantage ofseasonal opportunities (Klein and Cruz-Uribe 1996, 2000; Klein et al. 1999). However,it is possible that nearby colonies of seals were not available under different coastalconfigurations in the past.

At the Later Stone Age site of De Hangen in the southwestern Cape, the dominanceof juvenile hyrax individuals in a sample of 110 mandibulae suggest a late spring to latesummer occupation during the Late Holocene (Parkington and Poggenpoel 1971). Atthe contemporary Elands Bay Cave, the opposite pattern was observed with tootheruption stages indicating a winter occupation (Parkington 2001). An analysis of fourmandibulae and maxillae from hyraxes recovered in the ∼2 ka LSA levels at BlombosCave suggest the site was occupied during autumn in March and April (Henshilwood2008a). At a nearby site in the Blombosfontein Reserve, BBF 9 with an age of <2 ka,four hyrax specimens indicate the site was occupied during late winter in August/September (Henshilwood 2008a).

The teeth sample size of hyraxes from the 100 ka levels at BBC that could indicateseason of death is not large (n=23, MNI=15). In addition, aging criteria can beambiguous (Cruz-Uribe and Klein 1998, but see Steyn and Hanks 1983). Nevertheless,if it is accepted that the majority of hyraxes in the assemblage were acquired byhumans, then our data suggest both summer and winter visits to the cave during theM3 phase.

Evidence for human activities at BBC is overwhelming. Dense shell deposits, manybasin-shaped hearths, abundant lithics and other cultural artefacts, and the presence ofprey of various sizes clearly indicate human occupation of the shelter (e.g.,Henshilwood et al. 2001b). We conclude that despite some of the evidence for naturalaccumulation we have discussed above, that the hyraxes found at BBC were broughtthere by humans. The evidence is clear and includes the presence of cut marks, burntbone, the burnt mandible (e.g., Vigne and Marinval-Vigne 1983; Henshilwood 1997;Badenhorst 2008), the forelimbs outnumbering the hind limbs, the dominance of post-crania compared to crania (excluding teeth), and fragmentation of long bones. Theseasons of occupations during the M3 phase are more ambiguous but our evidencesuggests that year-round occupations are a probability.

Acknowledgments This research was funded through a grant to SB by the Palaeontological Scientific Trust(PAST) in South Africa. Financial support for the Blombos Cave project was provided to CSH and KvN by aEuropean Research Council Advanced Grant, TRACSYMBOLS No. 249587, awarded under the FP7programme at the University of Bergen, Norway and to CSH by a National Research Foundation/Department of Science and Technology funded Chair at the University of the Witwatersrand, South Africa.We are grateful to Denise Hamerton at the Iziko South African Museum in Cape Town who provided access tothe comparative collection, and Petro Keene, also at the same institution, who gave access to the Blombosfauna. Reviewers made useful suggestions.

Table 13 Blombos Cave andCollingham Shelter front and hindlimbs (after Cruz-Uribe and Klein1998). Data from Table 6 and Plug(1992; NISP)

Element Blombos Cave (%) Collingham Shelter (%)

Front limb 62 59

Hind limb 38 41

Afr Archaeol Rev (2014) 31:25–43 39

References

Andrews, P. (1990). Owls, caves and fossils. Chicago: University of Chicago Press.Badenhorst, S. (2003). The archaeofauna from iNkolimahashi Shelter, a Later Stone Age shelter in the

Thukela Basin, KwaZulu-Natal, South Africa. Southern African Humanities, 15, 45–57.Badenhorst, S. (2008). The zooarchaeology of great house sites in the San Juan Basin of the American

Southwest. Unpublished PhD dissertation, Simon Fraser UniversityBarry, R. E. (1994). Synchronous parturition of Procavia capensis and Heterohyrax brucei during drought in

Zimbabwe. South African Journal of Wildlife Research, 24(1–2), 1–5.Bartram, L. E., Kroll, E. M., & Bunn, H. T. (1991). Variability in camp structure and bone food refuse

patterning at Kua San hunter-gatherer camps. In E. M. Kroll & T. D. Price (Eds.), The interpretation ofarchaeological spatial patterning (pp. 77–148). New York: Plenum.

Bleek, D. F. (1929). Bushman folklore. Africa: Journal of the International African Institute, 2(3), 302–313.Bothma, J., & Du, P. (1971). Food habits of some Carnivora (Mammalia) from southern Africa. Annals of the

Transvaal Museum, 27(2), 15–26.Bousman, C. B. (2005). Coping with risk: Later Stone Age technological strategies at Blydefontein Rock

Shelter, South Africa. Journal of Anthropological Archaeology, 24, 193–226.Braczkowski, A., Watson, L., Coulson, D., Lucas, J., Peiser, B., & Rossiet, M. (2012). The diet of caracal,

Caracal caracal, in two areas of the Southern Cape, South Africa as determined by scat analysis. SouthAfrican Journal of Wildlife Research, 42(2), 111–116.

Brain, C. K. (1981). The hunters or the hunted? An introduction to African cave taphonomy. Chicago:University of Chicago Press.

Brain, C. K., & Sillen, A. (1988). Evidence from the Swartkrans cave for the earliest use of fire. Nature, 336,464–466.

Burton, M., & Burton, R. (2002). The international wildlife encyclopedia (Vol. 9). New York: MarshallCavendish.

Charles, R., & Jacobi, R. M. (1994). The late glacial fauna from the Robin Hood Cave, Creswell Crags: A re-assessment. Oxford Journal of Archaeology, 13(1), 1–32.

Cruz-Uribe, K., & Klein, R. G. (1998). Hyrax and hare bones from modern South African eagle roosts and thedetection of eagle involvement in fossil bone assemblages. Journal of Archaeological Science, 25, 135–147.

De Ruiter, D. J., Copeland, S. R., Lee-Thorp, J., & Sponheimer, M. (2010). Investigating the role of eagles asaccumulating agents in the dolomitic cave infills of South Africa. Journal of Taphonomy, 8(1–2), 129–154.

D’Errico, F., & Henshilwood, C. S. (2007). Additional evidence for bone technology in the southern AfricanMiddle Stone Age. Journal of Human Evolution, 52(2), 142–163.

D'Errico, F., Henshilwood, C. S., & Nilssen, P. (2001). An engraved bone fragment from ca. 75,000-year-oldMiddle Stone Age levels at Blombos Cave, South Africa: Implications for the origin of symbolism andlanguage. Antiquity, 75, 309–318.

Ditlhogo, M. K., & Setshogo, M. P. (2001). A floristic and faunal survey of the Kgale Hills. Botswana Notesand Records, 33, 91–99.

Driver, J. C. (2005). Manual for description of vertebrate remains. Unpublished manual. Cortez: CrowCanyon Archaeological Center.

Fairall, N., & Crawford, C. J. M. (1983). Application of the Ronson-Whitlock technique to estimate hyraxProcavia capensis numbers. South African Journal of Wildlife Research, 13(1), 25–26.

Fernández-Jalvo, Y., Andrews, P., & Denys, C. (1999). Cut marks on small animals at Olduvai Gorge Bed-I.Journal of Human Evolution, 36, 587–589.

Fourie, L. J. (1983). The population dynamics of the rock hyrax Procavia capensis (Pallas, 1766) in theMountain Zebra National Park. Unpublished PhD dissertation, Rhodes University

Gargett, V. (1972). Observations at a black eagle nest in the Matopos, Rhodesia. Ostrich, 43, 77–108.Harako, R. (1981). The cultural ecology of hunting behavior among Mbuti Pygmies in the Ituri Forest, Zaire.

In R. S. O. Harding & G. Teleki (Eds.), Omnivorous primates (pp. 499–555). New York: ColumbiaUniversity Press.

Henshilwood, C. S. (1997). Identifying the collector: Evidence for human processing of the Cape dune mole-rat, Bathyergus suillus, from Blombos Cave, southen Cape, South Africa. Journal of ArchaeologicalScience, 24, 659–662.

Henshilwood, C. S. (2008a).Holocene prehistory of the southern Cape, South Africa: Excavations at BlombosCave and the Blombosfontein Nature Reserve (Cambridge Monographs in African Archaeology. BritishArchaeological Reports S1860, Vol. 75). Oxford: Archaeopress.

40 Afr Archaeol Rev (2014) 31:25–43

Henshilwood, C. S. (2008b). Winds of change: Palaeoenvironments, material culture and human behaviour inthe Late Pleistocene (c. 77–48 ka) in the Western Cape Province, South Africa. South AfricanArchaeological Society Goodwin Series, 10, 35–51.

Henshilwood, C. S., & Sealy, J. (1997). Bone artefacts from the Middle Stone Age at Blombos Cave, southernCape, South Africa. Current Anthropology, 38(5), 890–895.

Henshilwood, C. S., D'Errico, F., Marean, C. W., Milo, R. G., & Yates, R. (2001a). An early bone tool industryfrom the Middle Stone Age at Blombos Cave, South Africa: Implications for the origins of modern humanbehaviour, symbolism and language. Journal of Human Evolution, 41, 631–678.

Henshilwood, C. S., Sealy, J. C., Yates, R., Cruz-Uribe, K., Goldberg, P., Grine, F. E., Klein, R. G.,Poggenpoel, C. A., van Niekerk, K., & Watts, I. (2001b). Blombos Cave, Southern Cape, SouthAfrica: Preliminary report on the 1992–1999 excavations of the Middle Stone Age levels. Journal ofArchaeological Science, 28, 421–448.

Henshilwood, C. S., D'Errico, F., Yates, R., Jacobs, Z., Tribolo, C., Duller, G. A. T., Mercier, N., Sealy, J. C.,Valladas, H., & Watts, I. (2002). Emergence of modern human behavior: Middle Stone Age engravingsfrom South Africa. Science, 295(5558), 1278.

Henshilwood, C. S., D'Errico, F., Vanhaeren, M., Van Niekerk, K. L., & Jacobs, Z. (2004). Middle Stone Ageshell beads from South Africa. Science, 304(5669), 404–404.

Henshilwood, C. S., D'Errico, F., & Watts, I. (2009). Engraved ochres from the Middle Stone Age levels atBlombos Cave, South Africa. Journal of Human Evolution, 57, 27–47.

Henshilwood, C. S., D'Errico, F., Van Niekerk, K. L., Coquinot, Y., Jacobs, Z., Lauritzen, S. E., Menu, M., &GarciaMoreno, R. (2011). A 100,000-year-old ochre-processing workshop at Blombos Cave, SouthAfrica. Science, 334(6053), 219–222.

Jacobs, Z., Wintle, A. G., & Duller, G. A. T. (2003a). Optical dating of dune sand from Blombos Cave, SouthAfrica: I—Multiple grain data. Journal of Human Evolution, 44(5), 599–612.

Jacobs, Z., Duller, G. A. T., & Wintle, A. G. (2003b). Optical dating of dune sand from Blombos Cave, SouthAfrica: II—Single grain data. Journal of Human Evolution, 44(5), 613–625.

Jacobs, Z., Duller, G. A. T., Wintle, A. G., & Henshilwood, C. S. (2006). Extending the chronology ofdeposits at Blombos Cave, South Africa, back to 140 ka using optical dating of single and multiple grainsof quartz. Journal of Human Evolution, 51, 255–273.

Jerardino, A. (2003). Pre-colonial settlement and subsistence along sandy shores south of Elands Bay, WestCoast, South Africa. The South African Archaeological Bulletin, 58(178), 53–62.

Jones, K. T. (1993). The archaeological structure of a short-term camp. In J. Hudson (Ed.), From bones tobehavior: Ethnoarchaeological and experimental contributions to the interpretation of faunal remains(Center for Archaeological Investigations, Occasional Paper, Vol. 21, pp. 101–114). Carbondale: SouthernIllinois University.

Jones, H. L. (2001). Electron spin resonance dating of tooth enamel at three Palaeolithic sites. UnpublishedM.Sc. dissertation, McMaster University

Klein, R. G., & Cruz-Uribe, K. (1996). Exploitation of large bovids and seals at Middle and Later Stone Agesites in South Africa. Journal of Human Evolution, 31, 315–334.

Klein, R. G., & Cruz-Uribe, K. (2000). Middle and Later Stone Age large mammal and tortoise remains fromDie Kelders Cave 1, Western Cape Province, South Africa. Journal of Human Evolution, 38(1), 169–195.

Klein, R. G., Cruz-Uribe, K., & Skinner, J. D. (1999). Fur seal bones reveal their variability in prehistorichuman seasonal movements on the southwest African coast. Archaeozoologia, 10, 181–188.

Lee, R. B. (1979). The !Kung San: Men, women and work in a foraging society. Cambridge: CambridgeUniversity Press.

Levinson, M. (1982). Taphonomy of microvertebrates—From owl pellets to cave breccia. Annals of theTransvaal Museum, 33(6), 115–121.

Lloveras, L., Moreno-García, M., & Nadal, J. (2009). Butchering, cooking and human consumption marks onrabbit (Oryctolagus cuniculus) bones: An experimental study. Journal of Taphonomy, 7(2–3), 179–201.

Lyman, R. L. (1994). Vertebrate taphonomy. Cambridge: Cambridge University Press.Lyman, R. L. (2008). Quantitative paleozoology. Cambridge: Cambridge University Press.Maclean, G. L. (1985). Robert’s birds of southern Africa. Cape Town: John Voelcker Bird Book Fund.Marean, C. W. (1986). Seasonality and seal exploitation in the southwestern Cape, South Africa. African

Archaeological Review, 4(1), 135–149.McMahon, C. R., & Thackeray, J. F. (1994). Plio-Pleistocene Hyracoidea from Swartkrans Cave, South

Africa. South African Journal of Zoology, 29, 40–45.Millar, R. P. (1971). Reproduction in the rock hyrax (Procavia capensis). Zoologica Africana, 6(2), 243–261.Mourre, V., Villa, P., & Henshilwood, C. S. (2010). Early use of pressure flaking on lithic artifacts at Blombos

Cave, South Africa. Science, 330(6004), 659.

Afr Archaeol Rev (2014) 31:25–43 41

Norton, P. M., Lawson, A. B., Henley, S. R., & Avery, G. (1986). Prey of leopards in four mountainous areasof the south-western Cape Province. South African Journal of Wildlife Research, 16, 47–52.

Olds, N., & Shoshani, J. (1982). Procavia capensis. Mammalian Species, 171, 1–7.Palmer, R., & Fairall, N. (1988). Caracal and African wild cat diet in the Karoo National Park and the

implications thereof for hyrax. South African Journal of Wildlife Research, 18(1), 30–34.Parkington, J. (2001). Presidential address: Mobility, seasonality and southern African hunter-gatherers. The

South African Archaeological Bulletin, 56(173/174), 1–7.Parkington, J., & Poggenpoel, C. (1971). Excavations at De Hangen, 1968. The South African Archaeological

Bulletin, 26(101/102), 3–36.Parsons, I., & Badenhorst, S. (2004). Analysis of lesions generated by replicated Middle Stone Age lithic

points on selected skeletal elements. South African Journal of Science, 100, 384–387.Plug, I. (1992). The macrofaunal remains of Collingham Shelter, a Late Stone Age site in Natal. Natal

Museum Journal of Humanities, 4, 53–59.Plug, I. (1993). The faunal remains from Nanda, an early Iron Age site in Natal. Natal Museum Journal of

Humanities, 5, 99–107.Plug, I. (1996). Seven centuries of Iron Age traditions at Bosutswe, Botswana: A faunal perspective. South

African Journal of Science, 92, 91–97.Plug, I. (1997). Faunal samples from Thulamela 2231 AC2, Kruger National Park, South Africa. Research by

the National Cultural History Museum, 6, 78–93.Plug, I., & Badenhorst, S. (2001). The distribution of macromammals in southern Africa over the past 30 000

years as reflected in animal remains from archaeological sites. Transvaal Museum Monograph No. 12.Pretoria: Transvaal Museum.

Plug, I., & Keyser, A. W. (1994). Haasgat Cave, a Pleistocene site in the central Transvaal: Geomorphological,faunal and taphonomic considerations. Annals of the Transvaal Museum, 36(9), 139–145.

Plug, I., & Voigt, E. A. (1985). Archaeozoological studies of Iron Age communities in southern Africa. In F.Wendorf & A. Close (Eds.), Advances in World Archaeology 4 (pp. 189–238). London: Academic.

Plug, I., Mitchell, P., & Bailey, G. (2003). Animal remains from Likoaeng, an open-air river site, and its placein the post-Wilton of Lesotho and eastern Free State, South Africa. South African Journal of Science, 99,143–152.

Rautenbach, I. L. (1982). Mammals of the Transvaal. Pretoria: Ecoplan Monograph No. 1. Ocoplan.Rautenbach, T. (2010). Assessing the diet of the Cape leopard (Panthera pardus) in the Cederberg and Gamka

Mountains, South Africa. Unpublished MSc thesis, Nelson Mandela Metropolitan UniversityRaven-Hart, R. (1971). Cape of Good Hope 1652–1702: The first fifty years of Dutch colonisation as seen by

callers. Cape Town: Balkema.Reitz, E. J., & Wing, E. S. (1999). Zooarchaeology. Cambridge: Cambridge University Press.Reynard, J. (2011). The unidentified long bone fragments from the Middle Stone Age Still Bay layers at

Blombos Cave, Southern Cape, South Africa. Unpublished MSc thesis, University of the WitwatersrandShaffer, B. S., & Neely, J. A. (1992). Intrusive Anuran remains in pit house features: A test of methods. Kiva,

57(4), 343–351.Skinner, J. D., & Chimimba, C. T. (2005). The mammals of the southern African subregion. Cambridge:

Cambridge University Press.Smithers, R. H. N. (1983). The mammals of the southern African sub-region. Pretoria: University of Pretoria.Steyn, D., & Hanks, J. (1983). Age determination and growth in the hyrax Procavia capensis (Mammalia:

Procaviidae). Journal of Zoology, 201(2), 247–257.Szuter, C. R. (1991). Hunting by prehistoric horticulturalists in the American Southwest. New York: Garland.Thompson, J. (2008). Zooarchaeological tests for modern human behavior at Blombos Cave and Pinnacle Point

Cave 13B, southwestern Cape, South Africa. Unpublished PhD dissertation, Arizona State UniversityThompson, J. C., & Henshilwood, C. S. (2011). Taphonomic analysis of the Middle Stone Age larger mammal

faunal assemblage from Blombos Cave, southern Cape, South Africa. Journal of Human Evolution,60(6), 746–767.

Tribolo, C. (2003). Apports des méthodes de la luminescence à la chronologie de techno-faciès du MiddleStone Age associés aux premiers Hommes modernes d’Afrique du Sud. Unpublished PhD dissertation,Université Bordeaux-1.

Tribolo, C., Mercier, N., Selo, M., Valladas, H., Joron, J. L., Reyss, J., Henshilwood, C., Sealy, J., & Yates, R.(2006). TL dating of burnt lithics from Blombos Cave (South Africa) and the antiquity of modernbehaviour. Archaeometry, 48, 341–357.

Vigne, J. D., & Marinval-Vigne, M. C. (1983). Methode pour la mise en evidence de la consummation du petitgibier. In J. Clutton Brock & C. Grigson (Eds.), Animals and archaeology: 1. Hunters and their prey (pp.239–242). Oxford: British Archaeological Reports International Series.

42 Afr Archaeol Rev (2014) 31:25–43

Villa, P., Soressi, M., Henshilwood, C. S., & Mourre, V. (2009). The Still Bay points of Blombos Cave (SouthAfrica). Journal of Archaeological Science, 36(2), 441–460.

Wadley, L. (2010). Were snares and traps used in the Middle Stone Age and does it matter? A review and acase study from Sibudu, South Africa. Journal of Human Evolution, 58, 179–192.

Watson, V. (2004). Composition of the Swartkrans bone accumulations, in terms of skeletal parts and animalsrepresented. In C. K. Brain (Ed.), Swartkrans: A cave’s chronicle of early man (Transvaal MuseumMonograph, Vol. 8, pp. 35–73). Pretoria: Transvaal Museum.

Watts, I. (2009). Red ochre, body painting and language: Interpreting the Blombos ochre. In R. Botha & C.Knight (Eds.), The cradle of language (pp. 62–92). Oxford: Oxford University Press.

Yellen, J. E. (1976). Settlement patterns of the !Kung: An archaeological perspective. In R. B. Lee & I. Devore(Eds.), Kalahari hunter-gatherers: Studies of the !Kung San and their neighbors (pp. 47–72). Cambridge:Harvard University Press.

Afr Archaeol Rev (2014) 31:25–43 43