The mammalian fauna associated with an archaic hominin skullcap and later Acheulean artifacts at ...

23 Pages·2012·1.79 MB·English

JournalofHumanEvolution52(2007)164e186
The mammalian fauna associated with an archaic hominin skullcap
and later Acheulean artifacts at Elandsfontein,
Western Cape Province, South Africa
Richard G. Klein a,*, Graham Averyb,c, Kathryn Cruz-Uribe d, Teresa E. Steele e,f
aPrograminHumanBiology,StanfordUniversity,Building80,InnerQuad,Stanford,CA94305,USA
bDepartmentofCenozoicStudies,NaturalHistoryDivision,IzikoSouthAfricanMuseum,P.O.Box61,CapeTown8000,SouthAfrica
cDepartmentofArchaeology,UniversityofCapeTown,PrivateBag,Rondebosch7700,SouthAfrica
dDepartmentofAnthropology,NorthernArizonaUniversity,Box15200,Flagstaff,AZ86001,USA
eDepartmentofHumanEvolution,MaxPlanckInstituteforEvolutionaryAnthropology,DeutscherPlatz6,Leipzig04103,Germany
fDepartmentofAnthropology,UniversityofCalifornia,OneShieldsAvenue,Davis,CA95616,USA
Received1February2006;accepted21August2006
Abstract
TheElandsfonteinsite,WesternCapeProvince,SouthAfrica,iswellknownforanarchaichomininskullcapassociatedwithlaterAcheulean
artifacts.Thesitehasalsoprovidednearly13,000mammalianbonesthatcanbeidentifiedtoskeletalpartandtaxon.Theassemblagederives
from49species,15ofwhichhavenohistoricdescendants.ComparisonstoradiometricallydatedfaunasineasternAfricaindicateanagebe-
tween1millionand600thousandyearsago.Uniquefeaturesofthefauna,includingthelateoccurrenceofadirk-toothedcatandasivathere,
mayreflectitsgeographicorigininaregionthatwasnotablehistoricallyforitsdistinctiveclimateandhighdegreeofbioticendemism.Together,
taxonomiccomposition,geomorphicsetting,andpollenextractedfromcoprolitesindicatetheproximityofalargemarshorpond,maintainedby
ahigherwatertable.Thesmallaveragesizeoftheblack-backedjackalsimpliesrelativelymildtemperatures.Thesumoftheevidenceplaces
bone accumulation duringone ofthemid-Pleistocene interglacials that were longer andcooler thanlater ones,includingthe Holocene.
Thegeomorphic context of the fauna presents no evidence for catastrophe, and most deaths probably resulted from attritional factors that
disproportionatelykilledtheyoungandold.However,onlythedental-ageprofileoflong-hornedbuffalosupportsthisdirectly.Fieldcollection
methodsbiasedskeletal-partrepresentation,butoriginally,itprobablyresembledthepatternintheyounger,marsh-edgeAcheuleanoccurrence
atDuinefontein2,45kmtothesouth.Excavationthereexposedmultiplevertebralspreads,whichprobablymarkcarcassesfromwhichhominins
orlargecarnivoresremovedthemeatierelements.Bonedamageatbothsitessuggeststhat,despiteabundantartifacts,homininsweremuchless
importantthancarnivoresintheboneaccumulation.Togetherwithlimitedobservationsfromothersites,ElandsfonteinandDuinefonteinpro-
visionally suggestthatAcheulean-age homininsobtained few largemammals, whether byhuntingorscavenging.
(cid:2)2006Elsevier Ltd.Allrightsreserved.
Keywords:Elandsfontein;Mid-PleistoceneAfricanmammals;Acheuleanecology
Introduction
The Elandsfontein farm and Acheulean site (33(cid:2)050S,
18(cid:2)150E)arelocatedapproximately18kmeastoftheAtlantic
* Correspondingauthor.
shoreand95kmnorth-northwestofCapeTownintheWestern
E-mailaddresses:[email protected](R.G.Klein),[email protected]
CapeProvinceofSouthAfrica(Fig.1).Thesiteissometimes
(G.Avery),[email protected](K.Cruz-Uribe),[email protected].
de(T.E.Steele). called Hopefield or Saldanha after towns located 11km
0047-2484/$-seefrontmatter(cid:2)2006ElsevierLtd.Allrightsreserved.
doi:10.1016/j.jhevol.2006.08.006
R.G.Kleinetal./JournalofHumanEvolution52(2007)164e186 165
Elands Bay Cave
Diepkloof Shelter
20º 25º 30º
0 300 km
25º 25º
Kathu Pan
Spreeuwal
Elandsfontein 30º SOUTH AFRICA 30º
Sea Harvest &
Hoedjies Punt
35º Klasies River Main 35º
Nelson Bay
20º 25º 30º
Duinefontein
A
T major outcrops
L
A CAPE TOWN of eolian sand
N
T
I
C 0 40 km
O
N C Swartklip
E
A
N FALSE
BAY
Die Kelders Cave
INDIAN OCEAN
Fig.1.ThelocationsoftheSouthAfricansitesmentionedinthetext[modifiedafterButzer(2004:Fig.1)].
northwest and 26km northeast, respectively. It is best known somewhat younger Acheulean site at Duinefontein (DFT) 2,
for a hominin skull cap that has been variously referred to about45kmsouthofEFTM.Asdiscussedbelow,theartifacts
Homorhodesiensis,‘‘archaic’’H.sapiens,orH.heidelbergen- andbonesatDFT2occurinasimilargeomorphiccontext,but
sis(Drennan,1953;Singer,1954;Rightmire,1998,2001).The they were still sealed in place, and they were recovered by
skull lay on a deflation surface or ‘‘bay’’ within an extensive excavation, mainly between 1996 and 2002 (Klein et al.,
dune field. Other similar bays have provided a fragment of 1999; Cruz-Uribe et al., 2003).
human mandibular ramus, more than 160 later Acheulean From our observations of the EFTM bones and basic simi-
bifaces,thousandsofassociatedflaketoolsandflakingdebris, larities between EFTM and DFT2, we conclude that, despite
and nearly 13,000 iron-mineralized mammalian fossils. The the abundance of Acheulean artifacts, hominins played little
objects are tightly associated with a calcareous duricrust that role in shaping the EFTM bone assemblage. Instead, like
we call Elandsfontein Main, or EFTM, to distinguish it from DFT2, EFTM appears to have been a place were artifacts
overlying horizons that have also provided bones and occa- and bones accumulated mostly independent of one another,
sional Middle Stone Age (MSA) artifacts. near a water hole that attracted both people and other large
We recently finished sorting the EFTM mammalian bones, mammals. Published reports on other Acheulean or Acheu-
and we aim here to summarize their implications for the geo- lean-agesitesinsimilargeomorphiccontextalsofailtoimpli-
logic antiquity, paleoenvironment, and paleoecology of the cate hominins importantly in the bone accumulation, and as
site.Morethan90%werecollectedunsystematicallyfromde- a hypothesis for further testing, we suggest that Acheuleans
flation surfaces in the 1950s and early 1960s, and to recon- andtheircontemporarieshadlittleimpact onthecontempora-
struct their original occurrence, we depend heavily on the neous large-mammal communities.
166 R.G.Kleinetal./JournalofHumanEvolution52(2007)164e186
Geomorphic setting
TheEFTMartifacts andbonesaccumulatedonquartz-and
shell-rich sands transported by wind from the Atlantic shore.
Similar sand bodiesoccur at various points along theWestern
Cape coast (Fig. 1), and carbonate in the comminuted shell
helped to buffer bones from dissolution. At Elandsfontein,
recentdeflationhasthoroughlymobilizedthesandmantle,ex-
posing, covering, and then re-exposing long-buried bones and
artifacts. In 1965e66, seventeen trial excavations to recover
Acheulean artifacts and bones before deflation were largely
fruitless. The single exception was ‘‘Cutting 10,’’ which ex-
posed456bonesand208artifacts,including50bifaces,below
a low mound of intact sands within a large deflation bay
(Klein, 1978b; Singer and Wymer, 1968). The biface concen-
Fig.2.TheElandsfonteinsitefromtheairin1976.
trationwasremarkableandanomalousforthesiteasawhole,
but the bone samplewas too small to illuminate the nature of
the overall bone-artifact association. Thus, for this purpose, axisofthesite.Itsoriginsandagearedebatable,butitprobably
we rely on the much larger sample from DFT2, about 45km representstheinduratedcoreofaPlioceneorearlyPleistocene
to the south. The DFT2 artifacts and bones occur within dune.Itunquestionably antedates the artifact-and-boneoccur-
a sand body that closely resembles the one at EFTM, but de- rences at the site, since these lie in sands that abut against it.
flation has been much less severe, and the bones and artifacts The sands are interrupted by two duricrustsdthe nodular,
remain in place over large areas, sealed in a 15e20-cm-thick calcareousonethatwecallEFTMandahigher-lyingcompact,
band that appears to mark a former land surface. ferruginous one. Two discontinuous ferricrete ridgesd1.5e
The circumstances that created the sand bodies at Elands- 5.0m high, 2.0m wide, and more than 1.0km longdsnake
fontein, Duinefontein, and other west-coast localities are per- acrossthesitewithinthesandsoverlyingtheupper(ferruginous)
haps most clearly revealed in the Geelbek Dunefield, about duricrust. They have been variously interpreted as the spines
5km west of Elandsfontein. Pedogenic horizons formed on of ancient dunes (Mabbutt, 1956), the podsolic B-horizons
andwithintheGeelbeksandsmarkstableperiodsbetweenre- of fossil soils on a dipping slope (Deacon, 1964; Partridge,
current episodes of sand transport inland, beginning in the 1982),orchannelorsubchannelexpressionsofformerstreams
mid-Pleistocene and extending into the Holocene (Felix- (Butzer,1973).
Henningsenetal.,2003;Kandeletal.,2003).Massmovement The lower, EFTM calcareous crust has provided all of the
inlandisgenerallythoughttohaveoccurredduringperiodsof Acheulean bifaces and the abundant iron-mineralized bones
lower sea level, when the exposed continental shelf provided onwhich we focus here.Thesurface ofthe upper ferruginous
a fresh sand supply. However, the most recent transport epi- crusthasdeliveredoccasionalbones,perhapsdeflatedfromthe
sode has been linked to a mid-Holocene high sea-level stand, overlyingsands,andextremelyrarebutwell-madeleaf-shaped
whensandserodedfromolder,previouslystabilizeddunesbe- (Still Bay) points and other likely MSA artifacts. The sands
came available for transport (Compton and Franceschini, above the ferruginous crust have produced artifact-poor, cop-
2005). Locally, any natural or cultural event that removed or rolite-rich bone accumulations that appear to represent fossil
reducedthevegetationcovercouldmobilizesand,particularly hyena dens (Avery, 1989). The most completely described is
duringsummer,whenstrongsoutherlywindsarecommon,the the ‘‘Bone Circle,’’ a roughly circular concentration about
sands are driest, and veldt fires reduce the vegetation cover. 4.5macross(InskeepandHendey,1966).Itcontainednumer-
The event that exposed bones and artifacts at Elandsfontein ous tooth-marked bones and carnivore (especially jackal)
occurredinorbefore1906,whenlocalresidentsfirstreported fossils that pinpoint hyenas as the bone accumulators (Klein,
fossils in deflation bays between dunes. The size of thewind- 1983).Thebonesinthevariousfossildensarewellpreserved,
eroded area has fluctuated through time, and it is rapidly but generally lack the characteristic iron mineralization of
shrinkingtoday,duemainlytotheencroachmentofintroduced bones from the EFTM crust.
Australian wattles (Acacia spp.). Figure 2 shows the exposed FerruginousandcalcareoussedimentsintergradeatElands-
area as it appeared from the air in 1976, when it extended fontein, and iron often coats calcareous nodules in the EFTM
up to 3km from north to south and 1.6km from east towest. crust. The crust originated when dissolved carbonates precip-
No deep sections exist at Elandsfontein, and the geologic itated from rainwater in the subsoil, and the ferruginization
context of artifacts and bones must be inferred mainly from followed at a time when the water table was much higher
surfacegeomorphology.Figure3summarizesthelocalstratig- and shallow ponds or marshes filled surface depressions
raphy reconstructed by Butzer (1973) from his own observa- (Roberts, 1996). Anoxic, acidic water from the ponds dis-
tions and those of Mabbutt (1956, 1957). The oldest solved iron that had been brought up from bedrock below
conspicuous feature is a calcrete ridge, up to 10m high and (Roberts, 1996) or introduced as an aerosol (Butzer, 2004)
60m wide, that extends for about 1km along the north-south and then released it at points of contact with calcareous
R.G.Kleinetal./JournalofHumanEvolution52(2007)164e186 167
0
2 m
hypothetical land surface at time of ferricrete channels
modern surface
FERRICRETE multicylcic
RIDGE caliche
upper duricrust
oxidation horizons
fossils and Acheulean
artifacts in lower CALCRETE RIDGE
duricrust
Composite, generalized stratigraphy of the Elandsfontein fossil site
Fig.3.Composite,generalizedstratigraphyofElandsfontein[modifiedafterButzer(1973:235)].
(alkaline) near-surface sediments (or bones). Higher intergla- Insum,geomorphicobservationsprovidenoreasontosup-
cialsealevel,greaterrainfall,orbothcouldexplainthehigher pose that the EFTM fauna is a composite of two or more
water table. The mammalian fauna, described below, together faunas that are widely separated in time. The EFTM crust
with pollen identified in fourteen associated hyena coprolites could comprise different crusts exposed in different parts of
by E.M. van Zinderen Bakker and J.A. Coetzee (Singer and the site, particularly on opposite sides of the main calcrete
Wymer, 1968), confirms that water stood nearby when ridge,butthebasichomogeneityandlimitedmetricvariability
EFTM formed, and the attraction of water to people and ani- of species samples gathered from widely separated interdunal
mals could explain why artifacts and bones are so common. bays (Klein, 1982, 1986; Klein and Cruz-Uribe, 1991; see
Similar ponds or marshes that formed when the water table below) suggest that time differences are limited. The extraor-
intersected the surface probably explain the thorough ferrugi- dinary homogeneity of the biface sample (Singer and Craw-
nization of the sands at DFT2 and the EFTM-like occurrence ford, 1958; Malan, 1962; Singer and Wymer, 1968) also
of artifacts and bones. Screening and bulk sampling at DFT2 suggests accumulation over a relatively short interval.
yielded thousands of bones from eight amphibian species,
which corroborate the geochemical indications for fresh sur- Taxonomic composition
face water (Sampson, 2003).
Deacon (1975) and Partridge (1982) have argued that the Table1liststhelargermammaliantaxaintheEFTMfauna,
EFTM crust marks a former water table to which objects infiveprogressivelyyoungerregionalfaunas,andinthehistoric
were lowered by persistent deflation over a span of perhaps fauna. Following prior practice (e.g., Klein and Cruz-Uribe,
tensorevenhundredsofthousandsofyears.Ifthisisaccepted, 2000),wedefinealargermammalasoneinwhichaverageadult
individual bones and artifacts could vary widely in age. The weightequalsorexceeds0.75kg.Itisonlargermammalsthat
fauna, as described below, might suggest this too, partly be- hunter-gatherers are most likely to concentrate, and smaller
causeitisamongthemostdiverseassemblageseverrecovered mammalsdmainly small rodents and insectivoresdabound
at an African Pleistocene site, and partly because it includes only in sites or layers where a paucity of artifacts suggests
species that elsewhere appear to have succeeded one another thattheywerecollectedbyraptors(Avery,1982).Thedistinc-
over a period of a half-million to a million years. tion between larger and smaller mammals is essentially aca-
However, the surfaces of freshly exposed EFTM bones demic for EFTM, since the recovery procedure produced no
never exhibit traces of prior sand abrasion, and an erosional smaller-mammalbones.
lag is inconsistent with ‘‘Cutting 10,’’ where bones and arti- The EFTM list in Table 1 depends heavily on specialist
facts were sealed on the calcareous duricrust beneath intact studies of baboons (Singer, 1962; Dechow and Singer,
sands. Persistent deflation is also an improbable explanation 1984), carnivores (Ewer and Singer, 1956; Hendey, 1974),
for the broadly similar artifact-and-bone spreads at DFT2, elephants (Maglio, 1973: 25e29), equids (Churcher, 1986;
where thebonesoftenoccurinanatomicalornear-anatomical Eisenmann, 2000), rhinoceroses (Hooijer and Singer, 1960a),
order and the paleosurface on which they lie undulates and hippopotamuses (Hooijer and Singer, 1960b), suids (Singer
dips weakly from east to west. Butzer (1973) found no geo- andKeen,1955;KeenandSinger,1956;CookeandWilkinson,
morphological evidence for deflation or dune formation in 1978:473),giraffids(sivatheres)(SingerandBone´,1960),and
the interval between the development of the EFTM crust and bovids (Klein and Cruz-Uribe, 1991). We are responsible for
historic times, and he concluded that the crust is the subsoil thetaxonomicassignmentsintheotherfaunas.Thenomencla-
manifestation of a gently rolling, nondunal land surface. De- ture for extant forms follows Skinner and Smithers (1990).
pressions in this surface could have held the ponds whose Hereafter, we use mainly vernacular names and we provide
acidic waters dissolved iron and ferruginized the sediments Linnaean equivalents only for species that are not included
and the bones. in Table 1.
168 R.G.Kleinetal./JournalofHumanEvolution52(2007)164e186
Table1
Thelarge-mammalspeciesrepresentedinthehistoricfaunanearElandsfonteinandinthefossilassemblagesfromElandsfonteinMain(EFTM),Duinefontein2
(DFT2),Spreeuwal(SPW),Swartklip1(ZW1),DiepkloofRockShelter(DRS),andElandsBayCave(EBC)
Taxa EFTM DFT2 SPW ZW1 DRS EBC Historic
Insectivora
Erinaceusfrontalis 1/1 152/20
Lagomorpha
Lepuscapensis,Capehare 74/5 146/8 X
Lepussaxatilis,scrubhare X
Lepusspp.,Capeand/orscrubhare 1/1 241/12 368/18 X
Rodentia
Bathyergussuillus,dunemolerat 173/14 2382/108 1/1 18/4 165/13 2973/157 X
Hystrixafricaeaustralis,porcupine 7/2 5/1 11/2 51/2 X
Pholidota
Phataginussp.,pangolin 2/1
Primates
yTheropithecusoswaldi,geladababoon 9/5
Papioursinus,chacmababoon 2/1 40/3 X
*Homoheidelbergensis,fossilhominin 2/1
H.sapiens,modernhumans 2/1 X
Pinnipedia
Arctocephaluspusillus,Capefurseal 6666/98 X
Carnivora
Canismesomelas,black-backedjackal 411/22 50/3 404/21 7/1 82/3 X
Vulpeschama,Capefox 22/4 35/2 1/1 X
Lycaonpictus,huntingdog 28/2 7/2
Ictonyxstriatus,stripedpolecat 3/1 18/4 1/1 43/6 X
Mellivoracapensis,honeybadger 62/6 8/2 7/1 X
Aonyxcapensis,clawlessotter 9/3 X
Viverracivetta,civetcat 2/1
Genettagenettaauttigrina,genetcat 4/1 1/1 X
Herpestesichneumon,Egyptianmongoose 25/2 7/1 6/1 4/1 X
Herpestespulverulentus,graymongoose 6/2 73/7 X
Atilaxpaludinosus,watermongoose 1/1 ?
Suricatasuricatta,slender-tailedmongoose 29/5
Crocutacrocuta,spottedhyena 4/1
Parahyaenabrunnea,brownhyena 43/9 2/1 21/3 X
Hyaenidae,spottedand/orbrownhyena 118/12 52/1 8/1 X
Protelescristatus,aardwolf X
Felislibyca,wildcat 14/2 2/1 31/4 20/1 22/2 X
Feliscaracalautserval,caracalorserval 244/12 4/1 74/4 22/3 43/2 X
Pantheraleo,lion 61/8 7/1 69/4 X
Pantherapardus,leopard (?)14/3 1/1 2/1 27/1 X
yMegantereonwhitei,dirk-toothedcat 15/2
Tubulidentata
Orycteropusafer,aardvark 35/2 X
Proboscidea
*Loxodontaatlantica,Atlanticelephant 383/14
Loxodontaafricana,Africanelephant 9/1 7/1 X
Hyracoidea
Procaviacapensis,rockhyrax 99/10 650/30 X
Perissodactyla
yEquuscapensis,Capezebra 2313/118 42/3 31/2 6/1 62/3
Equusquagga,quagga 36/7 63/3 12/2 7/1 15/1
Equuszebra,mountainzebra X
Equussp.indet.,indeterminatezebra 18/3 91/3
Dicerosbicornis,blackrhinoceros 79/13 1/1 14/3 7/1 X
Ceratotheriumsimum,whiterhinoceros 7/2 6/2 63/7
Rhinocerotidae,blackand/orwhiterhino 918/48 49/4 71/3 8/1 32/2 X
Artiodactyla
Hippopotamusamphibius,hippopotamus 171/6 5/1 6/2 5/1 27/2 X
Potamochoerusporcus,bushpig 1/1
yKolpochoeruspaiceae,extinctbushpig 69/13
yMetridiochoerusandrewsi,extinctwarthog 4/4
Phacochoerusaethiopicus,warthog ?
Suidaesp(p).,suids 83/16 6/1
R.G.Kleinetal./JournalofHumanEvolution52(2007)164e186 169
Table1(continued)
Taxa EFTM DFT2 SPW ZW1 DRS EBC Historic
ySivatheriummaurusium,sivathere 49/8
Taurotragusoryx,eland 184/54 8/2 8/2 18/6 7/2 121/11 X
Tragelaphusstrepsiceros,greaterkudu 64/8 65/10 14/3
Oryxgazella,gemsbok 16/7 1/1 6/1
yHippotragusgigas,‘‘gianthippotragine’’ 71/18
Hippotragusleucophaeus,blueantelope 43/6 14/3 5/2 58/11 1/1 6/1
Reduncaarundinum,southernreedbuck 164/52 17/4 2/1 132/38 3/1
Alcelaphusbuselaphus,Capehartebeest X
*Rabaticerasarambourgi,Arambourg’shartebeest 187/45
yDamaliscusaff.lunatus,tsessebe-likeantelope 113/19
Damaliscusdorcas,bontebok 7/2 X
y?Damaliscusniro,bontebok-likeantelope 112/14
y?Damaliscussp.nov.,bastardhartebeest 26/9
y?Parmulariussp.nov.,bastardhartebeest 39/5
Connochaetesgnou,blackwildebeest 130/46 146/20 19/4 100/15 3/2
Alcelaphinisp.indet.,wildebeestorhartebeest 5/2 29/4 X
yMegalotraguspriscus,‘‘gianthartebeest’’ 31/5 1/1
Sylvicapragrimmia,greyduiker 112/11 X
Peleacapreolus,Vaalribbok 1/1 X
*Gazellasp.,gazelle 26/11
*Antidorcasrecki,Reck’sspringbok 8/4
yAntidorcasaustralis,southernspringbok 12/7 77/13
Antidorcassp.(p).,springboks 32/12 1/1
Oreotragusoreotragus,klipspringer 8/1 6/1 X
Raphicerusmelanotis,Capegrysbok 310/51 62/7 119/27 67/20 X
Raphiceruscampestris,steenbok 1/1 70/18 X
Raphicerussp(p),grysbokand/orsteenbok 1/1 27/4 1071/83 X
yPelorovisantiquus,long-hornedbuffalo 470/52 150/20 15/3 2/1 6/1
Synceruscaffer,Capebuffalo 33/5 ?
yTribeindet.,gen.etsp.nov.,‘‘spiralhorn’’ 13/7
Smallbovids 479/51 420/8 6/1 556/27 438/9 5719/85 X
Small-mediumbovids 134/22 20/2 6/1 432/14 86/3 150/5 X
Large-mediumbovids 2564/207 1231/35 140/10 1943/60 114/2 38/4 X
Largebovids 1708/71 1647/21 130/7 220/11 60/2 483/15 X
Verylargebovids 2543/70 16/3 4/1 6/1
Notes:Table2providesbackgroundinformationonthesites.ThenumbersfortaxainthefossilfaunasaretheNISPs(numberofidentifiedspecimens)overthe
MNIs(minimumnumberofindividualstheNISPmustrepresent).Asterisksdesignateextinctspeciesthatprobablyorcertainlyevolvedintolivingforms.Daggers
markspeciesthatbecameextinctwithoutissue.Sources:Duinefontein2(Cruz-Uribeetal.,2003);Spreeuwal(KleinandAvery,unpublished);Swartklip1(Hendey
andHendey,1968;Klein,1983);DiepkloofRockShelter(ParkingtonandPoggenpoel,1987;Parkingtonetal.,2005;Triboloetal.,2005b;KleinandSteele,un-
published);andElandsBayCave(KleinandCruz-Uribe,1987;Parkington,1988).
TheWesternCapehasfurnishednumerousotherQuaternary layers, but the MNIs in the table were calculated as if each
faunaswecouldhavechosenforcomparison,butthoseinTable sitecontainedonlyonelayer,thatis,asifthebonesinthedif-
1representtherangeinbothtaxonomiccompositionandmode ferentlayersallcamefromthesameindividuals.TheMNIsare
of accumulation. Table 2 details their geologic context, age, thus conservative, since the alternative approachdto sum the
and artifactual associations, if any. Two of the faunasd MNIs for different layersdwould produce roughly the same
from DFT2 and Spreeuwaldaccumulated like EFTM in the relative taxonomic representation, but it would appear to be
open air near fresh water; onedfrom Swartklip 1doriginates based on much largernumbers.
from a fossil hyena den; and the remaining twodfrom Withineachfaunalassemblage,easeofidentificationvaried
DiepkloofRockShelterandElandsBayCavedcomefromar- fromspecies tospecies,depending onwhetherclosely related
chaeologicalcavedeposits[MSAandLaterStoneAge(LSA), speciesofsimilarsizewerepresentandonthedegreeofbone
respectively].Foreachtaxonateachsite,Table1presentsthe fragmentation. This is particularly great in the archaeological
numberofidentifiedspecimens(NISP)andtheminimumnum- cavesamples,duepartlytohumanbutcheryandpartlytopost-
berofindividuals(MNI)fromwhichthespecimensmusthave depositional leaching and profile compaction. In general, cra-
come.Figures4e6illustratetheproportionalrepresentationof nial parts allowed the most secure species identifications, and
each taxon at EFTM, based on its MNI. One or more of the postcranialboneswereoftenmoredifficulttoseparate,partic-
authors was involved in sorting each fauna, and the same ularly between species of hares, hyenas, rhinoceroses, zebras,
recording methodology was employed throughout. Klein and pigs, and bovids of approximately equal size. For each prob-
Cruz-Uribe (1984) described the assumptions behind the lematic species, Table 1 presents numbers based only on se-
MNI calculations. The archaeological sites contain multiple curely identified elements (mainly dentitions, or for the
170 R.G.Kleinetal./JournalofHumanEvolution52(2007)164e186
Table2
Thecontext,geologicage,andartifactassociationsatsouthernAfricansitesusedforcomparisontoEFTM
Site Context Geologicageandbasisforassessment Artifactassociations
DFT1 Assemblageexcavatedfromtunnelsthoughtto Undated,butfaunasimilartothatofZW1 Noartifacts;numeroushyenacoprolites,
representafossilhyenaden. impliesrelativelycool,grassyconditions tooth-damagedbones,andlarge-carnivore
duringalatePleistocenecoldphase. bonesimplyaccumulationbyhyenas.
DFT2 Excavatedspreadofartifactsandbonesona 270ka(opticallystimulatedluminescence). Acheulean.
paleosurfacewithinferruginizedeoliansands.
DRS Fillofalargequartziteoverhang,perchedabove >40ka(14Cassayofcharcoal);probably MultiplelayersandlenseswithMSA
thesouthbankoftheVerlorenvlei,aseasonal mainlylateIsotopeStage5andearlyStage artifactsdividedamongsixormore
riverthatenterstheAtlanticOceanatElandsBay, 3,betweenroughly115and55ka,basedon successiveculture-stratigraphicunits.
18kmtothewest. fauna,artifacts,andthermoluminescence
readingsonheatedquartziteandsilcrete
artifacts.
EBC Excavatedfillofacoastalcave. Fossiliferousdepositsdatedbetween13.6 MultiplelayersandlenseswithLSA
and7.9kaimmediatelybelowonesdated artifactsdividedamongthreeormore
betweenabout4.3and0.3ka(14Cassay successiveculture-stratigraphicunits.
ofcharcoal.)
EFTBC Circularconcentrationofbonesinthesands Undated,butfaunaidenticaltothatofZW1 Noartifacts;numeroushyenacoprolites,
abovetheferruginousduricrustatElandsfontein. impliesaccumulationundersimilarlycool tooth-damagedbones,andlarge-carnivore
conditions. bonesimplyaccumulationbyhyenas.
HDP1 Boneassemblageexcavatedfromtwoor Between300and200ka(infraredstimulated Noartifacts;numeroushyenacoprolites,
moreadjacentfissureorcavityfillswithin luminescence,thermoluminescence),but tooth-damagedbones,andlarge-carnivore
calcareouseolianite. possiblymuchyounger(stratigraphic bonesimplyaccumulationbyhyenas.
context,associatedfauna).
NBC Excavatedfillofacoastalcave. 18e0.5ka(14Cassayofcharcoal,ostricheggshell, MultiplelayersandlenseswithLSA
andmarineshell). artifactsdividedamongfoursuccessive
culture-stratigraphicunits.
KRM Excavatedfillofacoastalcavecomplex(Caves1, Between127and57ka(electron-spin-resonance MultiplelayersandlenseswithMSA
1A,1B,and2)atthemouthoftheKlasiesRiver. onteeth;thermoluminescenceonheatedartifacts; artifactsdividedamongfourormore
geologiccontext;faunalassociations). successiveculture-stratigraphicunits.
RDC Fillofacaveexposedinthewallofa Undated,butprobably>40ka,basedonthe MSA
limestonequarry. artifactsandfauna.
SPW 2.5-m-thicksequenceoffossiliferousmarshorlake >56ka(U-seriesreadingonacappingcalcrete); ?MSA(3flakes,othersprobablymissed
depositsoutcroppingintheintertidalzoneadjacent intervaloflowersealevel(?GlobalIsotopeStage duringcollectionintheswashzone).
toLangebaanLagoon.Bonesoccurprimarilyin 6,later5,or4)basedonoccurrenceatsealevel
grayorblackhumifiedbands.Itemscollected andtheexclusivelyterrestrialcompositionofthe
mainlyfollowingwaveerosionofthedeposits. fauna.
ZW1 Fissurefill1.5mhighand6macross,exposed >40ka(14Cassayofassociatedostricheggshell); Noartifacts;numeroushyenacoprolites,
at16e18mabovesealevelinaneolianitecliff intervaloflowersealevelbasedontheexclusively tooth-damagedbones,andlarge-carnivore
frontingFalseBay. terrestrialcompositionofthefauna;possibly bonesimplyaccumulationbyhyenas.
IsotopeStage6[between186and130ka
(racemizationofostricheggshell)}orlaterIsotope
Stage5[(between115and71ka)(correlationof
sealevelchangesrecordedintheeolianite
sequencewiththeglobalmarinerecord)].
Sources:DiepkloofRockShelter(DRS)(ParkingtonandPoggenpoel,1987;Parkingtonetal.,2005;Triboloetal.,2005a,b);Duinefontein1(DFT1)(Klein,1983);
Duinefontein2(DFT2)(Kleinetal.,1999;Feathers,2002;Cruz-Uribeetal.,2003);ElandsBayCave(EBC)(KleinandCruz-Uribe,1987;Parkington,1988,in
press);ElandsfonteinBoneCircle(EFTBC)(InskeepandHendey,1966;Klein,1983);HoedjiesPunt1(HDP1)(Klein,1983;BergerandParkington,1995;Stynder
etal.,2001;Butzer,2004;Matthewsetal.,2005);KlasiesRiverMain(KRM)(Klein,1976;SingerandWymer,1982;DeaconandGeleijnse,1988;Deacon,1989,
1995;DeaconandShuurman,1992;Triboloetal.,2005b);NelsonBayCave(NBC)(Klein,1972a,b;Deacon,1984;Inskeep,1987);RedcliffCave(RDC)(Brain
andCooke,1967;Cooke,1978;Klein,1978a;Cruz-Uribe,1983);Spreeuwal(SPW)(AveryandKlein,unpublished);andSwartklip1(ZW1)(SingerandFuller,
1962;HendeyandHendey,1968;Klein,1975;Butzer,2004).
bovids,dentitionsandhorncores),anditpresentsparallelnum- springboks;large-mediumforkudu,gemsbok,‘‘gianthippotra-
bers for composite categories that also include elements gine,’’ blue antelope, southern reedbuck, hartebeests, bastard
(mainly postcranial bones) that could have come from two hartebeests,andwildebeest;largeforeland,‘‘gianthartebeest,’’
or more closely related species. The categories are the genus Capebuffalo,andlong-hornedbuffalo(atDFT2);andverylarge
Lepus, the families Hyaenidae, Suidae, and Rhinocerotidae, forlong-hornedbuffalo(atallsitesexceptDFT2).
and five successive size classes among the Bovidae. With WecalculatedtheMNIsforeachcompositecategoryasifit
respect to the bovid species in Table 1 and Fig. 5, the size comprisedonlyasinglespecies.WithregardtotheHyaenidae,
classes are: small for klipspringer, grysbok, and steenbok; for example, the MNI estimates assume that dentitions and
small-medium for grey duiker, Vaalribbok, gazelle, and the limb bones came from one species, even at sites where
R.G.Kleinetal./JournalofHumanEvolution52(2007)164e186 171
“warthog”
“bushpig” (0.6 ) sivathere
(1.8 ) (1.1 )
hippo
(0.8 )
small bovids
(7.2 )
white
rhino
(2 ) small-medium
bovids (3.1 )
black
rhino
(1.8 )
quagga
(1.0 )
Cape zebra large-medium
(16.6 ) bovids (29.1 )
elephant
(2.0 )
carnivores
(13.1 )
gelada
baboon large bovids
(0.7 ) (10.0 )
very large
bovids (9.8 )
pangolin hare
(0.1 ) porcupine (0.7 )
(0.3 )
Elandsfontein Main large mammals
Fig.4.Therelativerepresentationoflarge-mammaltaxaintheEFTMfauna,basedontheminimumnumbersofindividuals(MNI)represented.Thepercentages
werecalculatedonatotalMNIof712.Figures5and6illustratetheproportionalrepresentationofbovidsandcarnivores,respectively.
dentitions show that the spotted and brown hyenas were both likelihoodthatitalsooccurredatEFTM,butthereisthenagging
present. At EFTM, DFT2, Spreeuwal, and Swartklip 1, we problemthatitistheonlyEFTMbovidforwhichhorncoresare
could identify some rhinocerotid and bovid postcranial bones lacking.
to species, at least tentatively, and we have boxed them
accordinglyintheSouthAfricanMuseum.However,wemain- Geologic age and assemblage homogeneity
tain the composite categories to accommodate specimens we
couldnotidentifybelowthefamily(orfamily-and-size)level. Table1andFigs.4e6revealtheremarkablediversityofthe
Thecompositesalsohelptocompensatefordifferencesinthe EFTM fauna, which comprises 48 species of larger mammals
number of minimally identifiable specimens among samples. (excluding the hominin specimen). This number exceeds not
Thenumberisparticularlyhighinthefragmentedarchaeolog- only the number in theotherfossilfaunas,butalso inthe his-
ical samples. toricfauna,whichincludedonly34.Theexceptionaldiversity
ThespecieslistforEFTMdiffersslightlyfromapreviouslist of EFTM is due partly to the presence of three extralimitary
(Klein and Cruz-Uribe, 1991) because of taxonomic reassess- speciesdpangolin, slender-tailed mongoose (suricate), and
ments. Most important, we have transferred small dentitions civet catdthat are known in the Western Cape only at
previouslyassignedto‘‘gianthippotragine’’togemsbok.Gems- EFTM,butitismainlytheresultoftheoccurrence of15spe-
bokand‘‘gianthippotragine’’dentitionsdifferonlyinsize,and cies that have no historic descendants. Four of thesedthe
sizevariationintheEFTMsamplesuggeststhatbothspeciesare ‘‘Cape zebra,’’ ‘‘giant hartebeest,’’ southern springbok, and
present.Historically,gemsbokwerecommononlyinmorearid long-hornedbuffalodoccur insomeoftheotherfossilfaunas
settings, 700km or more to the north and east (Skinner and in Table 1 (and in additional, taxonomically similar assem-
Smithers, 1990: 678), although individuals may occasionally blages like the one for the Elandsfontein Bone Circle that
havevisitedtheElandsfonteinregion(Skead,1980).Therecent we have not listed). All four appear to have survived locally
recovery of gemsbok at DFT2 and Spreeuwal increases the until 12e9ka (Klein, 1984). In the Western Cape, the
172 R.G.Kleinetal./JournalofHumanEvolution52(2007)164e186
eland
“spiral horn” (12.8 ) greater
long-horned (1.7 ) kudu
buffalo (1.9 )
(12.4 )
gemsbok
(1.7 )
Cape grysbok
(12.1 ) “giant
southern hippotragine”
springbok (4.3 )
(1.7 )
blue
Reck’s antelope
springbok (1.4 )
(1.0 ) reedbuck
gazelle (12.4 )
(2.6 )
Arambourg’s
“giant hartebeest” hartebeest
(1.2 ) (10.9 )
black wildebeest tsessebe-like
(10.9 ) antelope
(4.5 )
?Damaliscus
niro
?Parmularius sp. (3.3 )
(1.2 ) ?Damaliscus
sp. nov.
(2.1 )
Elandsfontein Main bovids
Fig.5.TherelativerepresentationofbovidspeciesintheEFTMfauna,basedontheminimumnumberofindividuals(MNI)represented.Thepercentageswere
calculatedonatotalbovidMNIof421.
remaining 11 extinct species occur only at EFTM, and since northern Tanzania. Bed IV was once believed to incorporate
the other faunas in Table 1 contain no extinct species that the boundary between the Matuyama Reversed Chron and
are absent at EFTM, the implication is that the other faunas Brunhes Normal Chron at about 780ka, butit is now thought
are much younger. EFTM must thus be significantly older to be reversed throughout, and it may precede the Jaramillo
than270kyr,theageoftheoldestradiometricallydatedfauna Normal Subchron between 1.07 and 0.99Ma (Tamrat et al.,
(from DFT2) among the others. It is probably older than 1995). The implication is that P. antiquus probably emerged
330kyr, the minimum age of a fauna with 42 taxa from Lai- close to or somewhat later than 1Ma. The earliest-known re-
nyamok,Kenya,that is the oldest diverse,directly datedAfri- cordofAlcelaphusisatBodo,MiddleAwash,Ethiopia,where
can assemblage to consist entirely of extant forms (Potts and it is tentatively dated to about 600ka (Clark et al., 1994;
Deino, 1995). Greater precision is impossible, because Renne, 2000). The co-occurrence of Pelorovis antiquus and
EFTMitselfhas providednomaterialsuitable forradiometric Rabaticeras thus fixes EFTM between about 1Ma and
dating,andthedepositscannotbelinkedtoanydatedexternal 600ka. This interval is consistent with the presence of eland,
stratigraphy.Atpresent,comparisonoftheEFTMfaunatora- Taurotragus oryx, which may have evolved from a kudu-like
diometricallyconstrainedfaunasineasternAfricaprovidesthe species (Tragelaphus sp.) after the emergence of P. antiquus
most secure foundation for dating. (Gentry, 1978, 1990; Gentry et al., 1995), and it is in keeping
Based mainly on the bovid species, Klein and Cruz-Uribe with the presence of the gelada baboon (Theropithecus os-
(1991)previouslyconcludedthatifEFTMoccurredindatable waldi), the extinct suids (Metridiochoerus andrewsi and Kol-
geological context in eastern Africa, its age would fall be- pochoerus paiceae), the reedbuck (Redunca arundinum), and
tween 700ka and 400kyr. Recent east African findings now Reck’s springbok (Antidorcas recki), all of which are repre-
suggest a somewhat older interval, between roughly 1Ma sented by the same or closely related forms at east African
and 600ka. The most useful species for such calculations sites dated between roughly 1Ma and 600ka (Geraads
are those whose presumed ancestors or descendants provide et al., 2004). The age implications of other EFTM taxa are
maximum and minimum age estimates, respectively. At lesscertain, butthere are nospecies thatunquestionablyorig-
EFTM,thetwomostrelevantspeciesarethelong-hornedbuf- inated after 600ka.
falo, Pelorovis antiquus, which probably descended from P. A major purpose in dating EFTM is to estimate the age of
oldowayensis,andtheextinctArambourg’shartebeest,Rabati- thefossilhomininskullandtheassociatedAcheuleanartifacts,
ceras,whichprobablygaverisetotheextanthartebeestgenus, but if, for the sake of argument, these are used independently
Alcelaphus (Gentry, 1978, 1990). to date the fauna, both suggest an age nearer 600kyr than
TheevolutionofP.antiquusfromP.oldowayensisoccurred 1Myr. The skull is clearly derived with respect to the 1-
subsequent to the formation of Bed IV at Olduvai Gorge, myr-old skulls from Buia, Eritrea (Abbate et al., 1998) and
R.G.Kleinetal./JournalofHumanEvolution52(2007)164e186 173
leopard (3.7 )
dirk-toothed cat
(2.5 )
lion (9.9 )
black-backed jackal
(22 )
caracal or
serval (14.8 )
Cape
fox (4.9 )
wildcat
(2.5 ) hunting
dog
(2.5 )
polecat
(1.2 )
honey
badger (7.4 )
brown hyena
(11.1 )
civet cat (1.2 )
suricate
spotted hyena (6.2 )
(1.2 )
water mongoose Egyptian mongoose
(1.2 ) (2.5 )
Elandsfontein Main carnivores
Fig.6.TherelativerepresentationofcarnivorespeciesintheEFTMfauna,basedontheminimumnumberofindividuals(MNI)represented.Thepercentageswere
calculatedonatotalcarnivoreMNIof81.
Daka-Bouri, Ethiopia (Asfaw et al., 2002), or the 970e900- speciesarescattered widelyacrossthesite,andtheindividual
kyr-old fragmentary cranium from Olorgesailie (Potts et al., species samples are remarkably homogeneous (Klein, 1982,
2004), all of which more closely fit the concept of Homo 1986;KleinandCruz-Uribe,1991).Noneexhibitgreatermor-
ergaster or African H. erectus. Similarly, the EFTM bifaces phological or metrical variability than species samples from
tend to be more finely shaped than those from Olduvai Beds other sites where context or direct dating excludes substantial
IIeIV (Leakey and Roe, 1994) or Daka-Bouri (Schick and temporal mixture.
Clark, 2000), and they are perhaps on a par with those from Figure 7 shows that this conclusion applies to three com-
Bodo (Clark and Schick, 2000). mon speciesdthe black-backed jackal, the long-horned buf-
The circumstances of faunal exposure and recovery at falo, and the Cape grysbokdthat are known to have varied
EFTM allow for the possibility, alluded to previously, that significantly in size through Quaternary time. Relative (or in-
the fauna comprises a mix of two or more assemblages, per- trinsic) variability around the mean is similar between the
haps one that is closer to 1Ma and one that is closer to EFTM samples for each species and the samples from other
600ka. This might not only help explain the unusually high sites, even though the means for the samples often differ
species diversity, but also the occurrence of two archaic significantly. (In this instance and others below, a statistically
elementsdthe sivathere and the dirk-toothed catdwhich at significant difference is assumed when the 95% confidence
600ka would be the youngest known African representatives limits for the means of two samples fail to overlap.) A proce-
of their lineages. In eastern Africa, sivathere fossils have not dure described by Lewontin (1966) revealed only one signifi-
been found at any site that postdates 1Ma (Churcher, 1978; cant difference (at the 0.05 level or below) in relative
Geraads et al., 2004), while remains of the dirk-toothed cat variabilitybetweenanEFTMsampleanditscounterpartsdthe
are unknown after about 1.5Ma (Arribas and Palmqvist, EFTM and NBC grysbok.
1999;TurnerandAnto´n,2004).TheEFTMfaunaistightlyas- In advance, it seems unlikely that the fossils of sivathere
sociatedwiththelower(calcareous)duricrust,butitcouldstill and dirk-toothed cat happened to come from a portion of the
mix two or more assemblages if the crust actually comprises crust that was radically different in age from the crust(s)
crusts of widely different ages exposed in different parts of that provided the bones of jackal, buffalo, grysbok, and other
the site. This possibility may never be conclusively rejected, especiallycommonspecies,andEFTMthussuggeststhatsiva-
but fossils of the most abundant ungulate and carnivore there and dirk-toothed cat persisted in southern Africa long