Animal exploitation strategies during the South African Middle Stone Age: Howiesons Poort and post-Howiesons Poort fauna from Sibudu Cave

Available online at www.sciencedirect.com

Journal of Human Evolution 54 (2008) 886e898

Animal exploitation strategies during the South AfricanMiddle Stone Age: Howiesons Poort and post-Howiesons

Poort fauna from Sibudu Cave

Jamie L. Clark a,*, Ina Plug b

a Museum of Anthropology, 4013 Museums Building, University of Michigan, Ann Arbor, MI 48109-1079, USAb Research Fellow, Department of Anthropology and Archaeology, University of South Africa, c/o P.O. Box 21022, Valhalla 0137, South Africa

Received 26 February 2007; accepted 11 December 2007

Abstract

As one of the few sites that preserve fauna from the Howiesons Poort (HP) and the immediately post-HP Middle Stone Age (MSA), SibuduCave provides a unique opportunity to explore the range of variability in subsistence behaviors during this important phase in human behavioralevolution. In addition to providing information on subsistence, the faunal assemblage serves as a means of reconstructing the environmentalconditions during these two periods. While the HP fauna is dominated by species that prefer closed (particularly forested) habitats, the faunafrom the upper-most layers of the post-HP MSA are largely representative of open conditions. These results largely coincide with macrobotan-ical analyses and may simply indicate that the extent of the riverine forest near the site was greater during the HP. Alternatively, the pattern couldbe indicative of a marked intensification in the exploitation of the environment in the immediate vicinity of the shelter during the HP, perhapsresulting from a decline in the productivity of adjacent regions. We also document variation in the frequency of the different bovid size classesover time. The evidence shows a declining focus on the smallest bovids after the HP, with a parallel increase in the frequency of large and verylarge bovids. Beyond a heavy focus on small bovids, small mammals and suids also occur at higher frequencies during the HP. Although the HPfaunal assemblage is largely unique as compared to the bulk of the MSA sequence at Sibudu, the evidence presented here suggests that thetransition between the HP and the post-HP MSA may have been more gradual than abrupt. Our results indicate that the HP and post-HPMSA inhabitants of Sibudu Cave were capable hunters; however, hunting strategies appear to show marked variation over time. We proposethat the variability in animal procurement strategies reflects a degree of behavioral plasticity beyond that generally attributed to MSApopulations.� 2007 Elsevier Ltd. All rights reserved.

Keywords: Zooarchaeology; Human behavioral evolution; Paleoenvironment; Late Pleistocene; Behavioral variability

Introduction

Although the growing database of comprehensively analyzedMiddle Stone Age (MSA) fauna has considerably expanded ourknowledge of the hunting abilities of the MSA inhabitants ofsub-Saharan Africa (e.g., Milo, 1998; Klein and Cruz-Uribe,2000; Marean et al., 2000; Assefa, 2006), much remains to be

* Corresponding author.

E-mail addresses: [email protected] (J.L. Clark), [email protected]

(I. Plug).

0047-2484/$ - see front matter � 2007 Elsevier Ltd. All rights reserved.

doi:10.1016/j.jhevol.2007.12.004

discovered about the range and nature of variability in subsis-tence behaviors within this important period of human evolution.Within the southern African MSA, two phases have figuredprominently in recent debates about the origins of modernhuman behavior: the Still Bay and the Howiesons Poort (HP).Characterized by the presence of bifacially-worked foliate orlanceolate points, the Still Bay is most recently known for theshell beads found in layers dating to w75 ka at Blombos Cave(d’Errico et al., 2005). Best known for its characteristic geomet-ric backed tools (cf. Singer and Wymer, 1982), the HP (typicallydated from w65 to w55 ka) has also received attention for theincised ostrich eggshell recovered at Diepkloof (Parkington

mailto:[email protected]

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Fig. 1. Map of southern Africa showing the location of Sibudu and other sites

mentioned in the text.

887J.L. Clark, I. Plug / Journal of Human Evolution 54 (2008) 886e898

et al., 2005; Wadley, 2006). The HP is of particular interest toour understanding of variability within the MSA because it isoverlain by horizons in which any classic signatures of ‘‘mod-ern’’ behavior have disappeared. Although a great deal hasbeen published on the significance of the Still Bay and HP forour understanding of human behavioral evolution (e.g.,McBrearty and Brooks, 2000; Henshilwood and Marean,2003; Mellars, 2006; McCall, 2007), remarkably little is knownabout subsistence behaviors during these phases (but see Klein,1976, for a discussion of the HP fauna from Klasies River).

With an extensive MSA sequence that includes Still Bay andHP horizons, and because of its excellent organic preservation,Sibudu Cave is an ideal setting for an exploration of subsistencebehaviors during the Middle Stone Age. We present here the firstreport on the macromammalian faunal assemblage from the HPlayers at the site. In addition to reporting on the HP fauna, thispaper includes a re-evaluation of the post-HP MSA assemblage,originally described in Plug (2004). Such a re-evaluation isrelevant both because refined chronological control resulted ina shift in some of the subdivisions employed in the previousanalysis and because results from the lowermost layers of thepost-HP MSA were unavailable at the time of that report. We fo-cus only on the macromammal (>300 g) remains as details onthe remainder of the faunal sample can be found elsewhere:Glenny (2006) reports on the microfauna; Plug (2006) considersthe aquatic animals; and Plug and Clark (in prep.) provide ananalysis of the avian remains. The rich faunal sequence at Si-budu has also been the subject of a number of other studies:Wells (2006) provides detailed analyses of fauna from layerRSp (located in the late MSA sequence), and Cain (2005,2006) reports on the nature of burning in the MSA horizonsand on the detailed taphonomic analysis of a selection of post-HP and late MSA layers.

The specific goals of this paper are to: (1) provide data onthe taxa represented in the HP and post-HP MSA assemblagesat Sibudu, (2) explore what the taxonomic data suggest aboutpast environmental conditions, (3) examine the bovid assem-blage in greater detail, focusing on what bovid exploitationpatterns can tell us about variation in subsistence strategiesover time, (4) evaluate the hunting abilities of Sibudu’s inhab-itants, and, finally, (5) provide hypotheses about the nature ofthe variability present in the Sibudu sample.

Background

Sibudu Cave is situated on a cliff above the Tongati River,approximately 40 km north of Durban in KwaZulu-Natal,South Africa (Fig. 1). Located at approximately 100 m abovemean sea level, the site is 15 km inland from the Indian Ocean.Sibudu was first excavated by Aron Mazel, who dug a trialtrench at the site in 1983. The present excavations, directedby Lyn Wadley, have been ongoing since 1998 (Wadley andJacobs, 2006). To date, a total of 21 m2 of MSA depositshave been excavated, although the lower-most layers (includ-ing all of the HP and much of the post-HP MSA) had onlybeen reached in a 2 m2 trial trench at the time of this analysis.Deposits are excavated following natural stratigraphy in 50 cm

quadrants. Initially, material was sieved through 2 mmscreens; however, since 2003, nested 2 mm and 1 mm sieveshave been employed in order to improve the collection ofseeds and micromammalian remains. All material from the2 mm sieves is sorted on site.

As information on dating and stratigraphy are reported indetail elsewhere (Wadley and Jacobs, 2004, 2006; Jacobset al., in press), we only summarize the most relevant datahere. Across the site, MSA deposits occur directly belowIron Age layersdno Later Stone Age remains are present.Within the upper MSA sequence, OSL dating has indicatedthe presence of three distinct age clusters. Based on the lithicassemblage, these have been informally designated as the finalMSA (weighted mean age of 36.9� 1.2 ka, found in the east-ern excavations only); the late MSA (weighted mean age of49.7� 1.2 ka); and the post-HP MSA (weighted mean ageof 60.1� 1.5 ka; Wadley and Jacobs, 2006; Jacobs et al., inpress). Below these levels are the HP, the Still Bay, and thepre-Still Bay. Dates for the oldest layers are still being pro-cessed, although a preliminary OSL date of w61 ka for theupper-most HP layer was reported in Lombard (2005). Shouldthis preliminary date be confirmed, it would suggest that therewas little or no occupational hiatus between the HP and thepost-HP MSA at Sibudu. A detailed stratigraphic profile forthe HP and post-HP MSA, drawn from the northern wall ofthe trial trench, is found in Fig. 2.

Environmental context

Given the multidisciplinary nature of research at Sibudu,the faunal remains serve as only one of a number of meansof reconstructing past environmental conditions at the site.Although much of the natural vegetation around Sibudu hasbeen destroyed by sugar cane farming, remnant forest persistson the cliff faces and steep slopes in the vicinity of the site(Allott, 2006b). Wadley (2006) has pointed out the importance

Fig. 2. Stratigraphic profile of the north wall of the trial trench showing the post-HP MSA (layers BSp to B under YA2) and the HP (GR to PGS). Modified from

Wadley and Jacobs (2006).

888 J.L. Clark, I. Plug / Journal of Human Evolution 54 (2008) 886e898

of recognizing the mosaic of habitats that persist around thesite even today; while the cool, moist conditions created byshade from the cliff encourage forest growth below the shelter,the sunny northern slopes opposite the shelter are home toopen species such as Acacia spp. A significant proportion ofthe modern vegetation is comprised of riverine species thatare supported by the Tongati River, a perennial water source.

Macrobotanical analysis by Sievers (2006) indicates thepresence of sedges throughout the entire MSA sequence. Theirpersistence implies that a water source (likely the TongatiRiver) was present near the shelter during periods of occupa-tion. Given the continued presence of this water source, it isexpected that riverine taxa (both floral and faunal) will bepresent in some degree throughout the occupational sequence.Analyses of charcoal (Allott, 2006a,b) and other macrobotan-ical remains (Sievers, 2006) have shown this to be the case,although the relative proportion of riverine taxa varies overtime. We will return to these results when considering the im-plications of the faunal remains to reconstructions of the pale-oenvironment during the HP and post-HP MSA.

In addition to the botanical analyses, archaeomagnetic datahave also been used in reconstructing past climate at Sibudu.Herries (2006) reports on the analysis of magnetic susceptibil-ity (MS) and mineral magnetic measurements taken from thesection walls of the trial trench. Although YA2 was the lowestlayer sampled, and thus his analysis did not include the HP,Herries’ results are relevant to an understanding of conditionsduring the post-HP MSA. Within the post-HP MSA sequence,

the deepest layers (G1 to YA2) showed distinctly differentmagnetic characteristics and much lower MS values than theupper layers (BSp to P1dP1 is a small lens that is not visiblein the profile illustrated in Fig. 2; it is located directly abovelayer G1 where it occurs). Herries proposes that the archaeo-magentic signature in the lower layers reflects the cold, glacialclimate of OIS 4. Accordingly, it is argued that the majorchanges in magnetic mineralogy and the higher MS valuesbeginning with layer P1 represent the transition to OIS 3. AsHerries (2006) indicates, the dating of the post-HP MSA atSibudu is consistent with the age of the OIS 4 to 3 transitionprovided by the Vostok ice core. Although the consistent pres-ence of the Tongati River would have provided some degree ofa buffer against climate change, thus reducing the visibility ofsuch changes in the archaeological record, we expect that sucha major shift would be reflected in the flora and fauna from therelevant layers.

Materials and methods

In analyzing the fauna, we decided to split the post-HPMSA into two units: the uppermost layers (BSp to P1) arecalled the post-HP MSA 1, while the lower layers (G1 toYA2) are referred to as the post-HP MSA 2. This divisionallows for more resolution in exploring the nature of the tran-sition between the HP and the post-HP MSA and it is conve-niently located near the middle of the post-HP MSA sequence.While the decision to split the sample between layers P1 and

Table 1

Bovid size classes (adapted from Brain, 1974)

Size class Live weight

(kg)

Species (list not inclusive)

Bov I <23 Blue duiker, common duiker,

klipspringer, oribi

Bov II 23e84 Springbuck, mountain reedbuck,

blesbok, impala

Bov III 85e295 Red hartebeest, blue wildebeest,

waterbuck, roan antelope

Bov IV 295e950 African buffalo, eland

Bov V >950 Giant buffalo, giant

hartebeest (both extinct)


G1 was based in part on the archaeomagnetic data, a numberof other lines of evidence support this choice. For example,although the post-HP MSA layers are informally classifiedas belonging to a single lithic tradition, detailed analysis ofthe assemblage shows a distinctive turnover in raw materialtype during the course of the post-HP MSA (Cochrane,2006). Quartz and quartzite occur much more frequently inthe lower layers of the post-HP MSAdfrequencies peak at86% in the lens immediately below YA1. Although the propor-tion of quartz and quartzite begins a slow decline after thispoint, a marked shift occurs between layers Ch2 and Su2(G1 and P1 are small lenses located between these two layers).The contribution of quartz and quartzite falls from >10% inCh2 to <1% in Su2; in subsequent layers, hornfels and doler-ite completely dominate the assemblage, typically occurring atfrequencies in excess of 95%. Furthermore, certain stone tooltypes, including grindstones, an anvil, and a hammerstone, arerestricted to the lower portion of the post-HP MSA sequence;the uppermost of these tools was recovered from layer Ch2.

In addition to the evidence from the archaeomagnetic andlithic analyses, the macrobotanical evidence also tentativelysupports this subdivisiondan as yet unidentified seed type(currently called ‘‘Type 5’’) is found throughout the MSAsequence but is found in the greatest quantities in the lower-most layers of the post-HP MSA (Sievers, 2006). Finally,although OSL ages do not indicate a significant hiatus betweenthe deposition of layers G1 and P1, Herries (2006) proposesthat the presence of gypsum nodules in layers Ch2 to P1may reflect a hiatus in the sequence (see Schiegl and Conard,2006, for a more detailed discussion of implications of gyp-sum formation at Sibudu). While any one of these findingsmay not be significant on its own, the fact that several inde-pendent classes of data show a consistent pattern suggeststhat the division of the post-HP MSA into an upper and lowersection is warranted.

The post-HP MSA fauna reported by Plug (2004) was ini-tially sorted by students at the University of the Witwaters-rand, with those fragments being considered identifiablesubmitted to her for analysis. Given the difficulty in recogniz-ing identifiable material in such a highly fragmented assem-blage, all sorting of the HP material was completed by theauthors. In addition, J.L.C. re-sorted the post-HP MSA fauna,increasing the identified sample by several hundred specimens.Once the identifiable fragments were removed, the ‘‘unidenti-fiable’’ material was subjected to additional sorting. Whilemuch of the sample could not be identified to specific skeletalelement or taxon, it was possible to sort the material into sev-eral fragment categories: skull, enamel, ribs, vertebrae, longbone shaft, and miscellaneous; material less than and greaterthan 2 cm was sorted separately. Detailed analysis of theunidentified fraction will be reported in a later publication.

Specimens were identified with the aid of the large compar-ative collection housed at the Transvaal Museum. As bovidswere the most common taxon at the site and species level iden-tification was not often possible, bovid remains were identifiedto size class utilizing Brain’s (1974) classificatory scheme,with the addition of the class ‘‘Bov V’’ for the extinct

Pelorovis antiquus and Megalotragus priscus (Table 1). Allidentified material was weighed and complete or nearly com-plete specimens were measured with digital calipers accordingto the procedures outlined in von den Driesch (1976). Wherepossible, sex was determined from pelvic fragments andhorn cores. Age at death was estimated based on epiphysealfusion, the presence of degenerative joint disease, and tootheruption/wear. Since the majority of teeth in the assemblageare highly fragmented, the presence of unerupted enamel oftenserved to identify juveniles while aged animals were identifiedbased on the presence of very heavily worn enamel.

MNI (minimum number of individual) values reported inTable 2 were calculated based on the total assemblage withineach phase; although this likely results in underestimated MNIvalues, the decision to aggregate in this manner was based onthe complex stratigraphy of the post-HP MSA, in which therelationship between layers is often ambiguous (e.g., layersIv and BM were excavated as independent units but may beparts of a hearth within P, MY may actually be a hearth withinMa, etc.). Given that MNIs can be significantly underestimatedwhen fragmentation rates are high (Grayson, 1984; Lyman,1994), and because of the problems involved in using mini-mum number counts to calculate the relative abundance oftaxa (Plug and Plug, 1990), analyses reported herein are basedon the number of identifiable specimens (NISP). We provideMNI values primarily so that those who prefer to utilize thesedata can do so.

In considering element frequencies, limb shafts were includedin our analysis and were identified to element wherever possible.As demonstrated in Pickering et al. (2006), a consideration ofcross-sectional geometry can be very useful in identifying longbone shafts to element; however, in their blind tests, only47.9% of fragments that preserved less than 50% of the shaft cir-cumference and length could be identified to skeletal element.Among the limb shaft fragments at Sibudu, the vast majorityof the assemblage preserves less than this amount. Thus, theidentification of shaft fragments to element generally dependedupon the use of diagnostic features such as foramina and muscleattachments. As a result of the heavy fragmentation, the GISbased approach to determining shaft-based MNEs (minimumnumber of elements) as developed by Marean et al. (2001) isnot applicable to the Sibudu assemblage.

Although we recognize that NISP values can be affected byfragmentation (e.g., Klein and Cruz-Uribe, 1984), minimum

Table 2

Macromammalian taxa present in the HP and post-HP MSA at Sibudu Cave

Taxon post-HP MSA 1 post-HP MSA 2 HP

NISP* %NISP MNI NISP %NISP MNI NISP %NISP MNI

Orcyterops afer, aardvark e e e e e e 1 0.0 1

Procavia capensis, rock hyrax 2 0.4 2 e e e 22 0.9 2

Lepus cf saxatilis, Cape hare e e e e e e 3 0.1 1

Pronolagus crassicaudatus, Natal red rock rabbit e e e e e e 2 0.1 1

Pronolagus sp. e e e e e e 3 0.1 2

Lepus/Pronolagus e e e e e e 12 0.5 1

Hystrix africaeaustralis, Cape porcupine 2 0.4 1 e 2 0.1 1

Thryonomys swinderianus, greater canerat e e e 2 0.6 1 2 0.1 1

cf. Thryonomys swinderianus e e e e 1 <0.1 e

Cricetomys gambianus, Gambian giant rat e e e 1 0.3 1 43 1.8 3

Rodent large e e e e e e 8 0.3 1

Papio hamadryas, chacma baboon e e e e e e 8 0.3 1

Cercopithecus pygerythrus, vervet monkey 1 0.2 1 e e e 47 2.0 4

cf. Cercopithecus pygerythrus 1 0.2 e e e e e e eCercopithecus albogularis, Sykes’ monkey e e e e e e 21 0.9 3

Primate small-medium e e e e e e 3 0.1 1

Primate medium-large e e e e e e 5 0.2 1

cf. Genetta tigrina, South African large-spotted genet e e e e e e 1 <0.1 1

Felid (cheetah/leopard) e e e e e e 2 0.1 1

Felid small (serval/wild cat) e e e e e e 4 0.2 1

Viverrid large e e e e e e 2 0.1 1

Viverrid/mustelid e e e e e e 1 <0.1 1

Galerella sanguinea, slender mongoose e e e e e e 3 0.1 2

Galerella sp. e e e e e e 1 <0.1 1

Atliax palundinosus, marsh mongoose e e e e e e 3 0.1 1

Mongoose e e e e e e 4 0.2 1

Mongoose small e e e e e e 12 0.5 2

Mongoose medium e e e e e e 2 0.1 1

Mongoose large e e e e e e 7 0.3 2

Canid small e e e e e e 1 <0.1 1

Canid medium (jackal size) e e e e e e 3 0.1 1

Canid large e e e e e e 1 <0.1 1

cf. Ictonyx striatus, striped polecat e e e e e e 5 0.2 1

Mustelid e e e e e e 1 <0.1 1

Carnivore small e e e e e e 5 0.2 1

Carnivore medium 1 0.2 1 e e e 2 0.1 1

Carnivore medium-large (hyena size) 1 0.2 1 e e e 1 <0.1 1

Carnivore large 6 1.1 2 e e e e e e

Equus quagga, plains zebra 25 4.6 2 e e e 9 0.4 1

cf. Equus capensis, extinct Cape horse 3 0.6 1 1 0.3 1 e e eEquus sp. 16 3.0 2 2 0.6 1 e e e

Potamochoerus larvatus, bushpig 1 0.2 1 13 4.0 1 201 8.3 4

cf. Potamochoerus larvatus e e e e e e 4 0.2 e

Phacochoerus africanus, common warthog 4 0.7 1 1 0.3 1 e e eSuid 7 1.3 1 9 2.8 3 16 0.7 2

cf. Giraffa camelopardalis, giraffe 2 0.4 1 1 0.3 1 e e e

Pelorovis antiquus, giant bufallo 5 0.9 1 e e e e e e

cf. Pelorovis antiquus 12 2.2 1 e e e e e eSyncerus caffer, African buffalo 14 2.6 2 e e e 12 0.5 2

cf. Syncerus caffer 9 1.7 e 1 0.3 1 e e e

Tragelaphus strepsiceros, kudu 10 1.8 2 e e e e e ecf. Tragelaphus strepsiceros 3 0.6 e e e e e e e

Tragelaphus scriptus, bushbuck e e e 2 0.6 1 5 0.2 2

cf. Tragelaphus scriptus 1 0.2 e 1 0.3 0 e e e

Tragelaphus oryx, eland 2 0.4 1 1 0.3 1 3 0.1 1

cf. Tragelaphus oryx e e e 2 0.6 0 e e e

Tragelaphus sp. 1 0.2 1 e e e e e e

Megalotragus priscus, giant hartebeest e e e e e e 4 0.2 1

cf. Megalotragus priscus 2 0.4 1 e e e e e eConnochaetes taurinus, blue wildebeest 5 0.9 1 1 0.3 1 3 0.1 2

cf. Connochaetes taurinus 5 0.9 e 2 0.6 0 e e e

Alcelaphus buselaphus, red hartebeest 2 0.4 1 e e e e e e


Table 2 (continued )

Taxon post-HP MSA 1 post-HP MSA 2 HP

NISP* %NISP MNI NISP %NISP MNI NISP %NISP MNI

cf. Alcelaphus buselaphus 1 0.2 1 e e e e e eDamaliscus pygargus, blesbok 1 0.2 1 e e e e e e

Alcelaphine large 14 2.6 2 e e e 3 0.1 1

Hippotragus equinus, roan antelope e e e e e e 5 0.2 2

Hippotragus/Tragelaphus oryx 1 0.2 1 e e e 4 0.2 1

Philantomba monticola, blue duiker 2 0.4 1 11 3.4 2 810 33.6 17

Cephalophus natalensis, red duiker e e e e e e 3 0.1 1

cf. Cephalophus natalensis e e e e e e 1 0.0 e

Sylvicapra grimmia, common duiker e e e e e e 2 0.1 2

Cephalophus/Sylvicapra e e e e e e 5 0.2 1

Redunca fulvorufula, mountain reedbuck 1 0.2 1 e e e e e e

Redunca sp. 2 0.4 2 e e e 1 <0.1 1

Kobus ellipsiprymnus, waterbuck 1 0.2 1 e e e e e e

cf. Kobus ellipsiprymnus 1 0.2 1 e e e e e e

Pelea capreolus, grey rhebok e e e e e e 3 0.1 2

cf. Pelea capreolus e e e e e e 2 0.1 ePelea/Redunca 1 0.2 1 e e e e e e

Raphicerus campestris, steenbok e e e e e e 21 0.9 3

cf. Raphicerus campestris e e e e e e 2 0.1 e

Raphicerus/Oreotragus e e e e e e 2 0.1 1

Aepyceros melampus, impala e e e 1 0.3 1 4 0.2 1

cf. Aepyceros melampus 1 0.2 1 e e e e e e

Oreotragus oreotragus, klipspringer 2 0.4 1 2 0.6 1 e e e

cf. Oreotragus oreotagus e e e 2 0.6 e e e eBov I 19 3.5 2 35 10.9 1 300 12.5 9

Bov I/II 2 0.4 1 8 2.5 1 6 0.2 1

Bov II 96 17.7 3 118 36.6 3 461 19.1 5

Bov II/III 1 0.2 1 9 2.8 1 4 0.2 1

Bov III 216 39.9 4 61 18.9 2 221 9.2 4

Bov III/IV 7 1.3 1 8 2.5 1 4 0.2 1

Bov IV 24 4.4 3 19 5.9 1 19 0.8 2

Bov IV/V 1 0.2 1 1 0.3 1 1 <0.1 1

Bov V 3 0.6 1 e e e e e e

Mammal small 2 0.4 1 7 2.2 2 23 1.0 3

Grand total 542 100.0 60 322 100.0 31 2408 100.0 122

Unidentified bone<2 cm 57932 51886 99533

Unidentified bone>2 cm 7176 3610 5055

Total unidentified 65108 55496 104588

% Identifiable against all fragments 0.83** 0.58 2.25

% Identifiable against>2 cm unidentified only 7.03** 8.19 32.26

* NISP is the number of identified specimens and MNI is the minimum number of inviduals these specimens represent. MNI was estimated for identifications

designated as ‘cf.’ only if those specimens must have come from individuals in excess of that represented by the remains verified to come from the species (e.g, if

Syncerus caffer was represented only by adult remains but cf. Syncerus included bones from a juvenile, then cf. Syncerus would receive an MNI count).** Complete sorting of the unidentified material from the uppermost four (out of a total of 22) layers in the post-HP MSA 1 is not yet complete and, thus, the

percent of identified bone reported for this stratigraphic unit is slightly overestimated.


number counts are not immune to these effects, and, as dem-onstrated by Grayson and Fray (2004), NISP-based analysesof skeletal part frequencies ‘‘can and do replicate the resultsof such analyses based on MNE, MAU and single elementMNI values’’ (40). Furthermore, as the same authors pointout, in cases where fragmentation results in a mismatch be-tween NISP and MNE values, it will not necessarily be clearwhich statistic provides a more accurate measure of relativeskeletal abundance. Given that the calculation of NISP is rel-atively straightforward and because the measure is not subjectto the aggregation effects that plague minimum numbercounts, element frequency data are also reported using NISP.

Results

The total assemblage analyzed to date consists of 106,996fragments in the HP; 55,818 fragments in the post-HP MSA2; and 65,650 in the post-HP MSA 1 (Table 2). The percentof identifiable bones within the assemblages largely dependsupon whether or not the less-than-2 cm fraction of unidentifi-able bone is included. When considering all excavated fauna,regardless of fragment size, the total percentage of identifiablematerial is 2.25% for the HP; this number declines to less than1% in both units of the post-HP MSA. These values areremarkably low and emphasize the incredibly high degree of


fragmentation in the sample. If the unidentified bone that isless than 2 cm (a common cutoff in zooarchaeological analy-ses) is excluded, those values rise to 32.36% for the HP, and8.92% and 7.03% for the post-HP MSA units. However, theselatter numbers are quite misleading, particularly for the HP, inwhich a large percentage of the identified bone was less than2 cm. Given the remarkable difference between the two setsof results, it is clear that analysts should specify how the per-centage of identifiable bone was calculated, as the methodemployed will affect interpretations regarding the degree offragmentation in the assemblage.

When considering the interpretations that can be basedupon the data presented in Table 2, it is important to keep inmind two separate issues dealing with sample size; first, thelimited spatial extent from which the remains were excavated,and second, the extreme comminution of the sample. It is truethat the majority of the sample analyzed here derives from the2 m2 test trench. While we cannot completely discount theidea that the patterning in the fauna may reflect a samplingbiasdin other words, that we are simply capturing shifts inactivity areas over timedMSA sites from which larger hori-zontal exposures are available (including Rose CottageCave, Klasies River Shelter 1B, and Florisbad) do not showclear evidence for the existence of well-defined activity areas(Wadley, 2001). Rather, the evidence suggests the presence oflargely unstructured camp organization, although some refusedumping is indicated. This suggests that the sample derivingfrom the test trench is likely to be representative of what ispresent in the site as a whole; however, this cannot be certainuntil a larger spatial extent is excavated.

Given the extreme comminution of the sample, it is reason-able to ask whether differences in the degree of fragmentationacross taxa will influence the results of our analyses that relyon taxonomic abundances. For example, when looking at thetaxonomic information presented in Table 2, it becomes clearthat the HP assemblage contains a significantly higher propor-tion of the smallest bovids (including the species-level identifi-cation of the blue duiker, with an NISP of 810). Given that thebones of small animals are expected to be less affected by severefragmentation than those of larger species (in part because theyare more likely to retain their integrity during processingdseeKlein, 1989, for a discussion), it may be the case that small bo-vids are overrepresented in the HP assemblage. However, thesame principles should apply throughout the sequencedand de-spite the high resolution of this analysis, in which even the small-est fragments were analyzeddthe blue duiker and other smallbovids were identified in only very small quantities in thepost-HP MSA. This suggests that the patterning evidenced inthe sample is not simply a reflection of preservational bias.

Discussion

Contributors to the assemblage

As systematic taphonomic analysis of the full HP andpost-HP MSA sample is underway but not yet complete, ourconsideration of the taphonomic history of the assemblage is

by necessity limited; however, preliminary results indicate thathumans were the primary contributor to the assemblage. An ini-tial sample of 3062 identified and unidentified bones from theHP and 3054 from the post-HP MSA has been examined undera binocular microscope at 10� magnification in order to assesscortical preservation and to identify surface modifications. Al-though evidence for weathering is incredibly rare, cortical pres-ervation is negatively impacted by high levels of burningdforexample, more than 50% of the analyzed sample from thepost-HP MSA 2 showed at least some degree of calcination.Given the palimpsests of hearths found in the post-HP MSA, itis unlikely that this burning was a result of natural fires.

Poor cortical preservation is no doubt partly responsible forthe relatively low incidence of surface damage identified in theassemblage. Cut- or chopmarks were identified on 1.3% of theHP sample, and although percussion damage was slightlymore common than carnivore or rodent damage, none accountfor more than 1% of the sample. No surface modificationsattributable to birds of prey were found.

In the lower layers of the post-HP MSA, no forms of sur-face modification were found in frequencies higher than 1%;again, this is likely a factor of the intense burning in theselayers. Surface damage is slightly more common in the post-HP MSA 1dwhile less than 1% of the material analyzedshowed evidence of cut marks, 3% showed direct evidenceof percussion damage (in the form of pits with striae, impactflakes with bulbs of percussion, and negative flake scars). Car-nivore damage was incredibly rare, again occurring at a rate ofless than 1%, with no evidence of rodent damage or surfacemodifications attributable to birds of prey.

Although the number of specimens preserving surfacemodifications is low, given that human produced damage ismore common than carnivore damage, burning is present athigh frequencies, very few carnivore remains were recovered(accounting for less than 2% of the NISP), and because thereis a high NISP/MNI ratio [argued by Villa et al. (2004) to bea good signature for human produced assemblages], the evi-dence available thus far indicates that the fauna was predom-inantly accumulated by human activity.

Sedimentological analysis of a selection of later MSAlayers (Pickering, 2006) provides further evidence that thedeposits were produced by humans. These layers were foundto consist of angular sediment grains and abundant angularfragments of bone and charcoal. This, combined with a lackof classic sedimentary structures, led Pickering to argue for lit-tle or no water borne transport of the deposits and to suggestthat the deposits were almost entirely anthropogenic in origin.

Paleoenvironmental implications

Table 3 provides information on the habitat preferences ofthe identified taxa from the HP and post-HP MSA. The per-centages here are based upon the fraction of the sample thatcould be identified to species, not the overall NISP. Beyondthe reasons discussed above, NISP was chosen for this analysisbecause the sample sizes provided by the MNI counts are sosmall; however, it is worth noting that the same patterns are


manifested when the analysis is conducted using MNI datadthe only difference being that the trends are less marked.

The HP fauna is dominated by species that inhabit closed orsemi-closed habitats (91.4%). This supports the charcoal data,which showed a strong evergreen forest signature. The pre-dominant genus in the charcoal assemblage for the HP wasthe evergreen Podocarpus spp. (Allott, 2006b); mammalscharacteristic of modern Podocarpus spp. forests include thevervet monkey, blue duiker, red duiker, bushbuck, bushpig,and the banded mongoose (Mungos mungo; Cooper, 1985).With the exception of the banded mongoose, all of these spe-cies have been identified in the HP. While some of these alsooccur in the post-HP MSA, this particular combination of taxawas limited to the HP.

Although their numbers are small (8.6% of the assem-blage), the HP does include species more common in (orrestricted to) open environments, including African buffalo,blue wildebeest, and roan antelope. This suite of species, alongwith the presence of some equid remains, implies that huntershad at least some access to more open woodland and/orsavanna environments than are found in the region today.Again, this matches with the charcoal data, as the HP sampleincluded some species indicative of an environment more openthan at present, particularly Kirkia spp., which today occurs inwarm/dry habitats in northern South Africa (Allott, 2006a).

Unfortunately, the post-HP MSA 2 assemblage containedonly a very small number of remains that could be taken tothe species level (n¼ 37). Of that sample, a majority is fromspecies expected to occur in riverine forest. Species thatinhabit more open environments are rare but present, includingwarthog and eland. The available data are indicative of condi-tions intermediate to those present during the HP and the post-HP MSA 1 (described below); however, larger sample sizesare needed to evaluate the validity of this pattern. It is hopedthat future charcoal analysis will provide further indicationsof local conditions during the post-HP MSA 2.

Of the three samples, the post-HP MSA 1 assemblage con-tains the lowest percentage of species that commonly occur inthe type of riverine forest present near the site (4.4%) and thelargest number of species that do not occur in the region today(n¼ 8). As argued by Plug (2004), the presence of these spe-cies, which include blue wildebeest, red hartebeest, and zebra,indicates drier conditions and a more open savanna environ-ment than present. The possible identification of giraffe (cf.Giraffa camelopardalis) further supports this argument. Thesedata provide a slightly different picture than that provided bythe charcoal sample, which, although variable, was dominatedby taxa from evergreen and riverine forest environments(Allott, 2006a). However, it seems reasonable to assume thatforagers would range farther afield to obtain meat than fire-wood, and it could simply be the case that plenty of fuelwoodwas available in the immediate vicinity of the site where foresttaxa predominated.

Taken as a whole, the data have at least two potential impli-cations. First, it is possible that the size of the forest remainedrelatively stable over time, with the faunal data indicatingmarked exploitation of the environment in the immediate

vicinity of the shelter during the HP. This type of intenseexploitation could potentially have resulted from a decline inthe productivity of adjacent regions due to increased aridityduring OIS 4. Alternatively, the extent of the riverine forestmay simply have been larger during the HP than in the post-HP MSA. It is expected that enlarged sample sizes, the resultsof taphonomic analysis, and an examination of lithic rawmaterial procurement strategies will all assist in choosing be-tween these alternatives.

Animal use: bovid exploitation strategies

Although bovids make up the majority of the assemblagethroughout the sequence, there are noticeable differences inthe representation of the different bovid size classes overtime (Table 4). The NISP values presented in Table 4 arehigher than those in Table 2 because the data include materialassigned to species or genus. As was the case in our consid-eration of paleoenvironment, the same trends are evidencedwhether NISP or MNI counts are utilized. In comparing therelative frequency of the different size classes, those frag-ments that were between classes were excluded, with theexception of the Bov IV/V fragments since the two largestbody-size categories were combined due to their small sam-ple sizes.

As a result of our increased sample sizes and the division ofthe post-HP MSA into two units, this analysis reveals patternsthat were not visible in Plug’s (2004) original study. With theexception of the Bov II size class, which occurs at the highestfrequency in the post-HP MSA 2 assemblage, the data showclear trends through time with the lower layers of the post-HP MSA being transitional between the HP and the post-HPMSA 1. The evidence shows a declining focus on the smallestbovids over time, with a parallel increase in the frequency oflarge and very large bovids. Given that the larger bovids pres-ent at the site tend to be those that prefer more open habitats,this pattern may in part be a reflection of the environmentaltrends discussed above.

In an attempt to further explore the significance of the pattern-ing in the size class data, we examined the element frequencydata by bovid size class and period. As density-mediated attritioncan have a significant effect on the interpretation of this class ofdata, we compared element representation for the medium tovery large bovids to the bone mineral density (BMD2) valuesprovided by Lam et al. (1999) for the blue wildebeest. Bov Iremains were excluded from this analysis because the most com-mon small bovid at the site, the blue duiker, is nearly two ordersof magnitude smaller than the wildebeest and thus we did notthink the wildebeest served as an appropriate analog. The bovidassemblage at Sibudu includes a large number of complete andnearly complete sesamoids (n¼ 232); however, Lam et al.(1999) did not provide density measurements for this element.The analysis was thus conducted twice; for the first run, densityvalues for the patella were used as a proxy measure for thesesamoids, while for the second run the sesamoids were ex-cluded. In both cases the results showed no significant corre-lation between element representation and mineral density

Table 3

Habitat preferences and frequency of occurrence (habitat data from Skinner and Chimimba, 2005)

Taxon Habitat preference Primary habitats Present today?* post-HP MSA 1 post-HP MSA 2 HP

Pronolagus crassicaudatus, Natal

red rock rabbit

Open Rocky areas Y e e 2 (0.2%)

Pronolagus sp. Grassy hillsides e e 3 (0.2%)

Hystrix africaeaustralis, Cape porcupine Both Most (except forest) Y 2 (2.2%) e 2 (0.2%)

Thyronomys swinderianus, greater canerat Open (w/ water) Tall grassy areas Y e 2 (5.4%) 2 (0.2%)

Reed/cane fields

Cricetomys gambianus, Gambian

giant rat

Closed Evergreen forest N e 1 (2.7%) 43 (3.5%)

Woodland savanna

Orcyteropus afer, aardvark Open Open woodland Y e e 1 (0.1%)

Grassland

Procavia capensis, rock hyrax Open Rocky outcrops Y 2 (2.2%) e 22 (1.8%)

Papio hamadryas, chacma baboon Open Woodland savanna Y e e 8 (0.6%)

Cercopithecus pygerythrus, vervet monkey Closed Riverine woodland Y 1 (1.1%) e 47 (3.8%)

Riparian savanna

Cercopithecus albogularis, Sykes’ monkey Closed Evergreen forest Y e e 21 (1.7%)

Riverine/coastal forest

Galerella sanguinea, slender mongoose Open Open areas Y e e 3 (0.2%)

Atilax paludinosus, marsh mongoose Both Close to water Y e e 3 (0.2%)

Equus quagga, plains zebra Open Savanna N 25 (27.5%) e 9 (0.7%)

Equus sp. Open woodland 16 (17.6%) 2 (5.4%) e

Potamochoerus larvatus, bushpig Closed Forest Y 1 (1.1%) 13 (35.1%) 201 (16.2%)

Riparian vegetation

Phacochoerus africanus, common warthog Open Open woodland Y 4 (4.4%) 1 (2.7%) e

Bushland

Syncerus caffer, African buffalo Open (w/ shade) Open woodland N 14 (15.4%) e 12 (1.0%)

Open vleis

Tragelaphus strepsiceros, kudu Open (w/ shade) Woodland savanna N 10 (11%) e e

Tragelaphus scriptus, bushbuck Closed Riverine underbrush Y e 2 (5.4%) 5 (0.4%)

Thickets

Tragelaphus oryx, eland Open Karoo Y 2 (2.2%) 1 (2.7%) 3 (0.2%)

Savanna

Connochaetes taurinus, blue wildebeest Open (w/ shade) Woodland savanna N 5 (5.5%) 2 (5.4%) 3 (0.2%)

Open woodland

Alcelaphus buselaphus, red hartebeest Open Grassland N 2 (2.2%) e e

Vlei

Damaliscus pygargus, blesbok Open Grassland N 1 (1.1%) e eHippotragus equinus, roan antelope Open Open savanna N e e 5 (0.4%)

Open grassland

Philantomba monticola, blue duiker Closed Forest Y 2 (2.2%) 11 (29.7%) 810 (65.4%)

Thicket

Cephalophus natalensis, red duiker Closed Forest Y e e 3 (0.2%)

Thicket

Sylvicapra grimmia, common duiker Semi-closed Bushland Y e e 2 (0.2%)

Tall grassland

Redunca fulvorufula, mountain reedbuck Semi-closed Rocky slopes Y 1 (1.1%) e e

Kobus ellipsiprymnus, waterbuck Both Vlei N 1 (1.1%) e e

Floodplain

Pelea capreolus, grey rhebok Open Rocky hills N e e 3 (0.2%)

Rocky slopes

Raphicerus campestris, steenbok Open Open grassland Y e e 21 (1.7%)

Open woodland

Aepyceros melampus, impala Open (w/ shade) Open woodland N e 1 (2.7%) 4 (0.3%)

Woodland savanna

Oreotragus oreotragus, klipspringer Both Rocky outcrops N 2 (2.2%) 2 (5.4%) e

Total NISP (%NISP)** 91 (100%) 37 (100%) 1238 (100%)

* This column indicates whether the species currently occurs in the region in which the site is located.** % NISP based upon this sample only, not the overall NISP.


(Run 1: Spearman’s r¼�0.107, t¼�0.895, p¼ 0.374, d.f.¼ 69;Run 2: Spearman’s r¼�0.078, t¼�0.647, p¼ 0.519, d.f.¼ 68).These results suggest that patterns in the element frequencydata are not due to density-mediated attrition.

In exploring the element frequency data for the bovidassemblage, we divided the skeleton into six anatomical units:skull/head (cranial elements, horn, teeth, hyoid), axial (ribs,vertebrae, sternum, pelvis), upper limb (scapula, humerus,

Table 4

Variation in bovid size class frequency through time

post-HP MSA 1 post-HP MSA 2 HP

NISP %NISP NISP %NISP NISP %NISP

Bovid size class

I 23 5.0 50 19.2 1146 60.5

II 103 22.5 122 46.7 476 25.1

III 258 56.3 64 24.5 232 12.3

IV/V 74 16.2 25 9.6 39 2.1


femur, patella), lower limb (radius, ulna, tibia, metapodials,carpals, tarsals), and feet (phalanges, sesamoids). As discussedin the methodology, the frequencies shown in Fig. 3 are basedon %NISP. Sample sizes are presented below each chart.Given the difficulty of assigning highly fragmented axial ele-ments (particularly the ribs and vertebrae) to size class, weexpected that the frequency of these elements would be lowthroughout the assemblagedFig. 3 shows this to be the case.

post-HP MSA 1

Skull/horn

Axial

Upper limb

Lower limb

Feet

0%

10%

20%

30%

40%

50%

60%

70%

80%

90%

100%

0%

10%

20%

30%

40%

50%

60%

70%

80%

90%

100%

HP

Skull/horn

Axial

Upper limb

Lower limb

Feet

4 18 64 32

2

7 24 25 8

8 37 76 12

3 22 85 19

Bov I Bov II Bov III Bov IV/V

1 2 3

267 39 35 6

56 15 6 0

144 76 34 5

496 176 58 8

183 170 99 17


Fig. 3. Bovid elemen

The small sample sizes for certain anatomical units andbovid size categories inhibits our ability to conduct quantita-tive analyses; however, on qualitative grounds, the relativepaucity of Bov IV/V skull and horn elements for the HP andthe post-HP MSA 2 as compared to the post-HP MSA 1may imply that the inhabitants of the site had to range fartherto obtain the largest game during the two earlier periods, asheads are unlikely to be transported over large distances.That the HP preserves the lowest frequency of Bov I upperlimb elements is perhaps unexpected, given that the liveweight of the most common species in this category is onlyw4 kg (Skinner and Chimimba, 2005) and, thus, could haveeasily been transported to the site as a single package. How-ever, this could be an indicator of intensified processing duringthe HP; perhaps the long bones of even the smallest bovidswere processed for marrow during this phase, thus reducingthe identifiability of these elements. As the sample size im-proves, we will be better able to evaluate the significance ofthese and other trends in the element frequency data.

post-HP MSA 2

0%

10%

20%

30%

40%

50%

60%

70%

80%

90%

100%

Skull/horn

Axial

Upper limb

Lower limb

Feet

8 6 10 4

12 0 1

9 7 3 3

24 46 22 4

7 63 28 13


t frequency data.


Animal exploitation during the HP and post-HP MSA:general trends

The available data indicate that the HP and post-HP MSAinhabitants of Sibudu Cave were capable hunters of a widevariety of game. The age structure of the HP sample, in which9.3% of the bones were identified as representing young orvery old animals, implies a focus on prime-aged individuals.Atlhough small bovids dominate the HP assemblagedtheblue duiker alone comprises 33.6% of the total identifiedsampledsuids, particularly the bushpig, are also well-represented, accounting for 9.2% of the assemblage. Bushpigsare nocturnal and highly aggressive (Skinner and Chimimba,2005); their presence indicates a high level of proficiency atprocuring dangerous game. In contrast to the pattern Kleinhas identified in other MSA assemblages (Klein, 1995, 1998;Klein and Cruz-Uribe, 2000), Sibudu does not appear toshow a focus on less-dangerous large game such as the eland.In the HP, as in the rest of the assemblage considered here,eland are less well-represented than dangerous large game,such as the African buffalo, or by the smaller but fierce suids,such as the warthog and bushpig (this pattern also holds in thelate and final MSA at Sibudu; see Plug, 2004, and Wells, 2006,for more detail).

Moving up through the sequence, there is a trend towardsincreasing percentages of medium and large game in thepost-HP MSA, although 65.9% of the bovid assemblagefrom the post-HP MSA 2 still consists of animals smallerthan size class III. In contrast, the data from the post-HPMSA 1 show a strong bias in favor of large game, with72.5% of the bovid assemblage now deriving from animalsin size class III or larger. The available data indicate a contin-ued focus on prime adult individuals throughout the course ofthe post-HP MSA, with only 5.3% of the post-HP MSA 2sample and just under 10% of the post-HP MSA 1 derivingfrom juvenile or aged animals.

As a whole, the HP fauna appear to reflect very differentanimal procurement strategies than those evidenced in thepost-HP MSA, particularly when compared against the upperportion of that sequence. The focus on small game duringthe HP does not appear to have been limited to the bovid as-semblage, as other small mammalsdincluding hares, hyraxesand the Gambian ratdare also present. Although it is possiblethat these may have been brought into the rockshelter by rap-tors or mammalian carnivores, Skinner and Chimimba (2005)argue that the Gambian rat, at least, is only preyed upon by hu-mans. Furthermore, as previously discussed, mammalian car-nivores are poorly represented, and raptors are alsouncommon (Plug and Clark, in prep).

Given the high frequencies of blue duiker in the HP sam-ple, it is relevant to consider the animal’s ecology and behav-ior in more detail. The species prefers closed forests andis more common in evergreen than in deciduous forests(Seydack and Huisamen, 1999). Blue duiker are territorial;group sizes are small and generally consist of no morethan a pair and their offspring. Within their territories, theyoften have routine movement patterns such that marked paths

are created between bed sites and feeding grounds; peopleaware of these habits can catch them in snares set acrosssuch paths (Skinner and Chimimba, 2005). Research byLupo and Schmitt (2005) on small prey procurement amongthe Bofi and Aka, modern foragers living in the Central Af-rican Republic, shows that nets are by far the most commontechnology used for capturing the blue duiker. Of 353 docu-mented kills, 86.4% were taken with nets (and only 7% weretaken with spears).

Research among modern hunter-gatherers in southeasternCameroon (Yasuoka, 2006) has shown that bushpig, whichare prone to attack when flushed out of hiding, are also com-monly taken by setting snares or traps along their walkways.Snares and nets are unlikely to preserve in the archaeologicalrecord and have not yet been documented for the MSA. How-ever, there is a lack of traditional MSA hunting technology inthe HP at Sibududstone points are absent. Although this maybe an artifact of the limited spatial extent of the excavations,the absence of points may reflect a focus on a different typeof hunting technology. Micro residue analysis by Lombard(2007) demonstrates that the backed segments in the HPwere hafted and used to process animal material; she combinesthis knowledge with the results of use-wear and macrofractureanalysis to argue that the segments served as inserts for com-posite hunting weapons. The faunal data raise the possibilitythat this technology may have been augmented by one basedupon perishable organic materials, such as wooden spears orsnares, traps, and nets.

When considering the evidence from Sibudu within thebroader picture of hunting strategies during the southern Afri-can MSA, it becomes clear that the assemblage is not entirelyunique. Preliminary reports on the fauna from Diepkloofindicate similar patterns with small game, such as molerats,hyraxes, and hares occurring more frequently in the HP (TLdated to 65e55ka), and a focus on large grazing animals inthe post-HP MSA (Parkington et al., 2005; Rigaud et al.,2006). Although somewhat earlier in time, the results of pre-liminary faunal analysis from the Still Bay levels at BlombosCave (77e75 ka) also suggest a focus on small bovids, withpercussion damage commonly occurring even on Bov Iremains (Thompson, 2006). The significance of these trendsis as yet unclear.

It is proposed here that the focus on smaller game duringthe HP and the greater frequencies of large game in thepost-HP MSA at Sibudu are linked to changes in environmen-tal conditions. In this case, the HP record may be indicative ofa marked intensification in the exploitation of the environmentin the immediate vicinity of the shelter during the HP, perhapsresulting from a decline in the productivity of adjacent regionsduring OIS 4. Future work on the Sibudu fauna will allow fora more thorough evaluation of these hypotheses. However,even if climatic change (such as the transition from OIS 4 to3) resulted in a shift in the availability of local resources,the variability in animal procurement strategies evidenced inthe record at Sibudu appears to reflect a higher degree ofbehavioral plasticity than that generally attributed to MSApopulations.


Conclusions

With its extensive MSA sequence and its excellent organicpreservation, Sibudu provides an ideal setting for explorationsinto the range and variability of subsistence behaviors duringthe MSA; we focused here on a comparison of the faunalassemblages from the HP and immediately post-HP MSA.As archaeomagnetic data indicated that a dramatic climaticshift (perhaps reflecting the transition from OIS 4 to 3) oc-curred within the post-HP MSA, and because the lithic datashowed a marked shift in raw material preference occurringat the same time, the decision was made to split the post-HPMSA assemblage into an upper and lower section. Analysesof the HP and post-HP MSA assemblages showed the faunato be intensively fragmented; when all unidentified materialwas included in the calculations, the percentage of identifiedbone was as low as 0.58% in the post-HP MSA 2. We are stillinvestigating the significance of the high degree of fragmenta-tion, which may in part be a reflection of the heavy burningdocumented in the assemblage. Preliminary taphonomic anal-ysis indicates that the fauna was predominantly accumulatedby human agency.

The environmental signature provided by the faunal remainslargely coincides with that from charcoal and other macrobotan-ical analyses. The HP assemblage contains the largest proportionof species that prefer closed or semi-closed environments; thesemake up 91.4% of the total assemblage. However, the presenceof species that prefer more open habitats, such as Equus sp. andthe roan antelope, suggests at least some access to savanna envi-ronments. The post-HP MSA 2 fauna also consists predomi-nantly of species that could be expected to occur in theriverine forest around the perennial Tongati River; however,sample sizes for this period are particularly small. In contrast,the presence of large grazers such as the blue wildebeest, harte-beest, and zebra in the upper layers of the post-HP MSA assem-blage indicates a strong savanna/open woodland componentduring the post-HP MSA 1.

Data on variability in the frequency of the different bovidsize classes over time seem to largely reflect these environ-mental trends, as larger bovids of the type that dominate thepost-HP MSA 1 assemblage also tend to be those occurringin more open habitats. As with the environmental data, thepost-HP MSA 2 appears to be transitional between the HPand the post-HP MSA 1dthe only deviation being that me-dium sized bovids are most common in the post-HP MSA. 2Although the limited sample from this phase restricts our abil-ity to make any absolute statements, the data presented heresuggests that, at least as far as the faunal evidence is con-cerned, the transition between the HP and post-HP MSA wasmore gradual than abrupt. The implications of this to ourunderstanding of the changes in material culture at the endof the HP are not yet clear.

The available data indicate that the HP and post-HP MSAinhabitants of Sibudu Cave were capable hunters who focusedon a wide variety of game: both dangerous and docile, largeand small. We propose that the variability in subsistencebehavior evidenced at Sibudu demonstrates an ability to

successfully adapt to long-term changes in the environment.Ongoing research at Sibudu, such as charcoal and lithic anal-yses from the lower MSA horizons, and continued faunal stud-ies, including analysis of the Still Bay and pre-Still Bay faunalremains, will be critical to confirming our findings and will nodoubt shed further light on the significance of this variabilityto our understanding of human behavioral evolution duringthe MSA.

Acknowledgements

First and foremost, we would like to thank Lyn Wadley formaking the material available for study and for her continuedsupport. We also thank John Speth, Richard Redding, RobertWhallon, Adam Van Arsdale, Francis Thackeray, MilfordWolpoff, the editors of JHE, and two anonymous reviewersfor their helpful comments on earlier draftsdtheir input hasmade this a stronger paper. Any mistakes that remain are, ofcourse, our own. Thanks also to Grant Cochrane for providingunpublished data on the post-HP MSA lithic assemblage.Wendy Voorvelt drew the original base map and profile onwhich our Figs. 1 and 2 are based. J.L.C. would like to thankthe staff at the Transvaal Museum, particularly Teresa Kear-ney, for her assistance and generosity. Research by J.L.C.was made possible by NSF Dissertation Improvement Grant# 0612606 and by support from the James B. Griffin scholar-ship fund from the University of Michigan Museum ofAnthropology.

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Documents

Animal exploitation strategies during the South African Middle Stone Age: Howiesons Poort and post-Howiesons Poort fauna from Sibudu Cave