Large mammal turnover in Africa and the Levant between 1.0 and 0.5 Ma H.J. O’REGAN1, L.C. BISHOP1, A. LAMB1, S. ELTON2 & A. TURNER1 1School

of Biological & Earth Sciences, Liverpool John Moores University, Liverpool, L3 3AF, UK (e-mail: h.j.o’[email protected]) 2Department of Anatomy, Hull York Medical School, University of Hull, Hull, HU6 7RX, UK Abstract: Faunal change at the Early–Middle Pleistocene boundary in Europe has long been a topic for discussion. However, analyses of large mammal turnover at this time in Africa have been lacking, largely because of the low number of sites dated to this interval. Recent work, particularly in the last 10 years, has resulted in a much larger published sample of sites and we synthesize these data in this paper. In our multivariate (TWINSPAN) analyses of African and Levantine large mammal faunas we found that localities were subdivided by geographic regions, not by age. There were some small-scale changes with the appearance or extinction of particular taxa, but there was no large-scale turnover such as that seen in Europe. The Levant was included as a possible route for faunal interchange with east Africa, but no similarities were found between these areas. It therefore appears that the modern zoogeographic separation of the Levant and north Africa into the Palaearctic region and sub-Saharan Africa into the African region can be traced back to at least the Early–Middle Pleistocene boundary.

The interval between 1.0 and 0.5 Ma is of great interest to archaeologists and physical anthropologists. It encompasses the first well-dated hominin occupation of western Europe at Atapuerca in northern Spain at c. 0.8 Ma (Carbonell et al. 1999), with either Homo heidelbergensis, H. antecessor or H. erectus as the possible earliest immigrant species. The mid-Quaternary also sees a major turnover in the extensive mammalian fauna of western Europe, and while this turnover is probably more gradual than previous assumptions of a marked change at the Early–Middle Pleistocene boundary have implied (Turner 1995a), it is still clear that a significant shift in faunal composition took place. European hominins were part of that fauna, and whatever the taxonomic problems surrounding their study it is clear that their dispersion patterns, in particular, are likely to be best understood when viewed against those of the larger set of mammals. The continent of Africa has experienced some major turnovers in its mammalian fauna since the beginning of the Pliocene (Turner 1990, 1995b; Turner & Wood 1993; Vrba 1995, 1997, 1999; Turner & Antón 1999, 2004), but much of the evidence for this comes essentially from the Pliocene and is based on well-dated localities in eastern Africa. In contrast, the changes within the African Pleistocene are less well understood, largely because well-dated suitable deposits tend to be absent, and relatively little synthesis has been attempted. Indeed Potts & Deino (1995, p. 106) wrote ‘the pace of Mid Pleistocene faunal change [in Africa] has previously been undocumented’ largely in response to the lack

of sites dating to the Early–Middle Pleistocene in east and south Africa, but in the last eight years nine sites have been published, reanalysed or redated, which gives a total sample of 21 sites that have been included in the following analysis. On a broad scale Africa appears to have undergone its last major turnover in mammalian faunas towards the end of the Pliocene and into the earliest part of the Pleistocene. This occurred when the sabretoothed cats became locally extinct and the large predator guild took on essentially its present structure and composition, and when the archaic deinotheres, gomphotheres and chalicotheres disappeared along with robust australopithecines of the genus Paranthropus. Pigs and giraffes reduced in diversity and the numbers of three-toed hipparion horses dwindled (Turner & Antón 2004). At the same time species that had appeared on the continent during the later part of the Pliocene survived in greater numbers down to the present day. Africa therefore began to look modern at a substantially earlier date than Eurasia or the Americas, where the modern mammalian fauna is a Holocene phenomenon, while still retaining a larger proportion of its larger Pliocene mammals. In an effort to redress the imbalance of syntheses of the African Pleistocene fauna this study examines continental-scale patterns in turnover across the Early–Middle Pleistocene boundary, informally dated at 0.78 Ma (Bassinot et al. 1994; Richmond 1996), initially to see if any species could potentially be used as indicators of this event in Africa. Although lacking absolute dates, there are several

From: HEAD, M.J. & GIBBARD, P.L. (eds) 2005. Early–Middle Pleistocene Transitions: The Land–Ocean Evidence. Geological Society, London, Special Publications, 247, 231–249. 0305-8719/05/$15 © The Geological Society of London 2005.

232

H.J. O’REGAN ET AL.

Early–Middle Pleistocene assemblages recorded in north Africa and the Levant that may yield interesting results when compared with those from subSaharan Africa. These two areas are in separate modern zoogeographical regions, with north Africa and the Levant in the Palaearctic (which also includes Europe and much of Asia) and east and south Africa occupying the African region (Cox 2001). However, the faunal composition in the Levant is, at times, regarded as an extension of that of Africa (Pickford & Morales 1994), with the region providing a route for dispersal into and out of that continent (Tchernov 1992). We have restricted our discussion to the larger mammalian fauna as these are the animals most likely to have a range that extends across all four regions. We will also look for evidence of endemism or geographical isolation in any of the four regions of north, east and south Africa and the Levant.

Compilation of the data tables We have included in this analysis assemblages from ten east African, four south African, four north African and three Levantine localities, and these are shown on Figure 1. Sites were included in this analysis if they met two criteria: (1) the majority of their estimated dates fall within the 1.0–0.5 Ma boundaries that we are using to bracket the Early–Middle Pleistocene transition; and (2) a full faunal list is available. All mammals were included in our analysis except Rodentia, Lagomorpha, Hyracoidea, Insectivora and Chiroptera as many of these tend to be smaller bodied and often have more localized species. For east Africa the dates of some localities can be calculated using direct dating techniques (e.g. 40Ar/39Ar), but in most other areas the sites have been dated using biostratigraphy and palaeomagnetism (Table 1). The Plio–Pleistocene sites from south Africa were placed within a relative chronology using faunal resemblance indices by McKee et al. (1995) and we have followed their sequence here. Sites for which there is major disagreement over their ages have not been included, so that for example Kabwe in Zambia which has been placed in many different time intervals, from 0.125 Ma to 1.33 Ma (McBrearty & Brooks 2000), is not appropriate for inclusion as it might be placed anywhere along this continuum. We have used the program TWINSPAN (TwoWay INdicator SPecies ANalysis) to give an independent assessment of the similarity or differences between faunal lists from the 21 sites in our analysis. This is a multivariate hierarchical classificatory method originally developed to handle ecological data, in particular species distribution (Hill 1979), and as such it is very suitable for the analysis of pal-

Latamne Aïn Maarouf

Tighenif

Evron Quarry Gesher Benot Ya'aqov

Thomas Quarry 1

Palaearctic

Buia

African

Bodo Daka

Kasibos Kanjera

N

Kapthurin Olorgesailie

Olduvai

0

1500 km

Plovers Lake Gladysvale Cornelia Elandsfontein

Fig. 1. Outline map of Africa and the Levant showing the approximate locations of all localities included in our analysis. See Table 1 for further details of each locality.

aeontological data (e.g. Graham et al. 1996). The program searches for similarities and differences within the dataset and then subdivides the complete dataset into two groups. These groups are then reanalysed and divided into two again, and so on. The groups produced may be of unequal sizes, but once they include four sites or less they stop being subdivided (Kent & Croker 1992). Until then the program will continue to force a separation even when there is no obvious reason for doing so, and the majority of analyses are therefore stopped after three divisions. Information on the localities in this analysis, including the stratigraphic resolution of each faunal list, dates and method of dating are shown in Table 1. This information has been taken largely from the original site reports but more recent references on dates and faunal re-evaluations have also been included. These faunal lists reflect different levels of stratigraphic precision, with some site reports providing detailed breakdowns of the levels within a site, whilst others are more general. Thus at Thomas Quarry 1 resolution is sufficient for there to be faunal lists for each level, whilst at Olduvai Gorge many individual sites are represented in the faunal lists for Beds 3 and 4. The ideal would be for each faunal list to come from a well-dated level within each site, but in the majority of cases this is not possible. Many localities are only dated by member, or in some cases formation, and we have taken the best combination of dating and faunal resolution available for use in this analysis. In any case, for the purposes of this study the finer stratigraphic differences do not matter so long as the locality is not simply dated between 1.0

Morocco

Danakil (Afar), Eritrea Kanam East, Kenya Tanzania

Kenya

Middle Awash, Ethiopia Kenya

Tanzania

TQ1L

Tigh

Lain

TQ1G

Buia

Old3

Olo1p

Daka

Olo6/7

Old4

Level L, Thomas Quarry 1 Tighenif‡

L’Aïn Maarouf

Level G, Thomas Quarry 1§ Buia

Kanj

Bodo

KN-5, Kanjera North

Bodo

Member 6/7, Olorgesailie Bed IV, Olduvai

Kasi

Algeria

Lata

Latamne

Kasibos Formation Bed III, Olduvai Upper Member 1 palaeosol, Olorgesailie Daka

Morocco

Gesh

Gesher Benot Ya’aqov‡

Homa Peninsula, Kenya Middle Awash, Ethiopia

Morocco

Syria

Israel

Israel

Evro

Evron Quarry

Location

Code

Locality

Member – unit u & u-t

Member

Bed

Member

Member

Member

Bed

Formation

Layer

Level

Site

Site

Level

Site

Site

Site

Stratigraphic unit

Biostratigraphy, palaeomagnetism Biostratigraphy, palaeomagnetism Biostratigraphy


1.0


0.6

0.78

0.8–0.6


0.9–0.78

1.0–0.8

0.992–0.974

1.15–0.8

40Ar/39Ar, biostratigraphy

palaeomagnetism 40Ar/39Ar, palaeomagnetism Palaeomagnetism, sedimentation rate Palaeomagnetism

40Ar/39Ar,

Biostratigraphy, Palaeomagnetism Biostratigraphy, palaeomagnetism Palaeomagnetism, sedimentation rate 40Ar/39Ar


1.0 1.07–0.99

Biostratigraphy

0.7–0.6


0.7


0.7

0.7–0.5

Palaeomagnetism, biostratigraphy, archaeology Fauna

Palaeomagnetism

Dating method

0.78–0.74

1.0–0.78

Age (Ma)

Table 1. Information on all African and Levantine localities included in this analysis.

Seasonal river (Wadi)

Open, dry grassland, by a river channel. Swamp, returning to mudflats

Open grassland, water-margins floodplain

Open, dry grassland, by a river channel. Open grasslands, trees in patches

Savanna

Marine cave

Lake with woods and open savanna

Open, dry grassland with few trees. Marshy or lacustrine

Mixed, with grass, woodland and water. Wooded, around lake margin

Palaeoenvironment

Plummer (1991, pers. comm.); *Ditchfield et al. (1999). Kalb et al. (1982a;1982b); *Clark et al. (1994).

*Potts et al. (1999); Potts (pers. comm.) *Hay (1994); Leakey (1994).

Asfaw et al. (2002).

Potts (1994); *Potts et al. (1999); Potts (pers. comm.)

*Hay (1994); Leakey (1994).

Ditchfield et al. (1999).

Abbate et al. (1998).

Tchernov et al. (1994); *Ron et al. (2003). Geraads & Tchernov (1983); Bar-Yosef (1998); *Goren-Inbar et al. (2000). Guérin et al. (1993); *Bar-Yosef (1998). Raynal et al. (2001); Geraads (2002). *Dauphin et al. (1994); Geraads (2002). Geraads & Amani (1997); *Geraads (2002). Geraads (2002).

Source of faunal list

Elan

Elandsfontein ¶

Site

Formation

Site

Site

Bed

Stratigraphic unit

Fauna, archaeology


0.7–0.4


0.7

ESR, palaeomagnetism Biostratigraphy Moist, with grassland and bush

Mixed, open and closed environment faunas. Grassland and bush, with nearby water Open grassland

40Ar/39Ar

Fauna

Palaeoenvironment

Dating method

0.78–0.578


1.0

0.5420.004– 0.5090.009

Age (Ma)

Background information including stratigraphic information, dates and source of references for faunal lists for all localities included in this analysis. * Source of the date, if different from that of the faunal list. † Also known as Jisr Banat Yacub. ‡ Also known as Ternifine and Palikao. § Level G at Thomas Quarry 1 is also known as the Hominid Level.  Gladysvale external deposits (GVED), not those of the main cave. ¶ Elandsfontein Main site (EFTM) also known as Saldanha and Hopefield.
Approximate date. Blanks indicate that no information is available.

Corn

Glad

Gauteng, South Africa Gauteng, South Africa Free State, South Africa Cape Province, South Africa

Plov

Gladysvale (GVED)  Cornelia

Baringo, Kenya

Kapt

K3(ii), Kapthurin Formation Plovers Lake

Location

Code

Locality

Table 1 continued.

Butzer et al. (1974); *McKee et al. (1995). Klein & Cruz-Uribe (1991).

Lacruz et al. (2003).

Thackeray & Watson (1994).

McBrearty et al. (1996); *Deino & McBrearty (2002).

Source of faunal list

AFRICAN FAUNAL CHANGE

Ma and 0.5 Ma but is placed either before or after the 0.78 Ma boundary, as we are looking for broad patterns in faunal turnover at this time rather than changes within a single region or sequence. All specific and generic identifications have been included in Table 2. Subspecific designations have been ignored in the following analysis but are given in the Appendix. In Tables 1 and 2 the sites are shown in date order by geographic region. For brevity a four-letter code has been attached to each site in these tables and in the TWINSPAN analyses and these codes are given in Table 1. A full faunal list for each locality is presented in the Appendix, including those identifications that were only determinable to familial or tribal level. Where an attempt has been made in the faunal list to assign a specific designation then this has been counted as a species identification level as in Turner & Wood (1993) and McKee et al. (1995). Where a species has been found at only one or two sites then no judgement has been made on either its temporal or geographical range, with the exception of groups where there is an obvious limitation on their distribution such as the cervids, which have only been recorded from the three Levantine sites.

TWINSPAN results The results of a TWINSPAN analysis of specieslevel identifications only are shown in Figure 2. In this analysis Evron Quarry stood out at the first level as being very different from the rest of the fauna. This site has recently been redated to c. 1.0 Ma (Ron et al. 2003), but had previously been regarded as being of a similar age to ‘Ubeidiya (c. 1.4 Ma) in Israel (Tchernov et al. 1994). Therefore this division suggests that the fauna from Evron Quarry is very different from the later Levantine and African localities, but this is based on a sample of only four species-level identifications. The one animal that really sets Evron Quarry apart from the other Levantine sites is a unique suid, Kolpochoerus evronensis, which is the youngest species of this genus to be found outside Africa (Tchernov et al. 1994). In the second level of division the north African and Levantine sites (not including Evron Quarry) were split off from the sub-Saharan African localities, with the exception of Bed 3ii of the Kapthurin formation which clustered with the north African and Levantine sites. At the third level of division, the north African–Levantine group was separated into a north African (with Bed 3ii, Kapthurin) and a Levantine group, and the sub-Saharan African group was subdivided into a south Africa and an east African group. Bed 3ii of the Kapthurin Formation shares a large number of species with many of the north African sites, including Crocuta crocuta,

235

Panthera leo, Connochaetes gnou, Ceratotherium simum, Theropithecus oswaldi and Homo erectus (see Table 2). These are all large-bodied and fairly cosmopolitan species within Africa, but the north African sites and Bed 3ii of the Kapthurin Formation are the only places at which all these species cooccur. The initial similarity with the Levant is based on the possible presence of Hippopotamus amphibius at Gesher Benot Ya’aqov and the definite identification of this species from Bed 3ii at Kapthurin. The conclusion from this analysis is that on the basis of their specific-level identifications the localities are divided along geographic lines (with one exception), initially into their separate biogeographic regions – the Palaearctic versus the African – and then into regions within them. There is no evidence that the date of the localities affected their placement, in that they clustered by region rather than by age of site. A generic-level analysis was also performed (results not shown) which allowed many more identifications to be included. The discrimination of sites was very poor, since in this case five of the north African and Levantine sites – Evron Quarry, Gesher Benot Ya’aqov, Latamne, Thomas Quarry Level L and L’Aïn Maarouf – formed the first division from all of the other sites. The two remaining north African sites (Tighenif and Level G of Thomas Quarry) then formed a subgroup with Kapthurin Bed 3ii at the next level of division. Finally, the two members at Olorgesailie grouped with the south African localities, while all remaining east African localities grouped together. These divisions were not related to either site geography or date, while the faunal divisions did not appear to correspond with the site environment, or the environmental requirements of the fauna. Therefore the generic-level analysis did not produce meaningful results, probably because some genera such as Hippopotamus and Equus are ubiquitous and the lumping of their component species together lost information on their individual characteristics. Figure 3 shows the results for the analysis of all species-level identifications and genera that had no species assigned (e.g. Makapania sp. is the only identification of any members of that genus in Table 2) and four genera for which the specific and generic identifications were amalgamated (Ursus, Camelus, Hipparion and Stegodon). The pattern of divisions is very similar to that of Figure 2, except all of the Levantine sites group together first, followed by the distinction between north and sub-Saharan Africa at the second division. At the third division the sites again divided along geographic lines, with all south African localities in one group and all east African sites in the other. Again, these divisions emphasize geographic differences between the areas rather than any change through time.

Carnivora Canis adustus Canis mesomelas Canis sp. Lycaon magnus Lycaon pictus Ursus bibersoni Ursus sp. Lutra maculicollis Ictonyx striatus Aonyx capensis Aonyx sp. Mellivora capensis Mellivora sp. Genetta sp. Viverra civetta Herpestes ichneumon Suricatta suricatta Ichneumia sp. Crocuta crocuta Crocuta sp. Hyaena hyaena Parahyaena brunnea Pachycrocuta brevirostris Felis caracal Panthera pardus Panthera leo Homotherium sp. Megantereon cultridens Dinofelis piveteaui Monachus monachus Artiodactyla Hippopotamus sirensis Hippopotamus gorgops Hippopotamus amphibius Hippopotamus behemoth Hippopotamus sp.

Evro

X

Gesh

?

Lata

cf.

X

X

TQ1L cf.

Tigh X

? X

?

?

cf. ?

Lain cf.

X

TQ1G cf.

aff.

X

X

X

X

X

X

cf.

aff.

Buia X

Kasi X

Old3 X

Olo1p cf.

X

cf.

X

cf.

aff.

Daka

East Africa

cf.

X

Olo6/7

North Africa Old4 X

X

X

Kanj cf. cf.

Bodo cf.

Kapt X

X

X X

Plov X

X

Southern Africa

X

X

Glad

Levant

X

Corn

Table 2. List of all specific and generic identifications included in this analysis

X

X

X

X

X

X

X X X

X

X X

X

X

Elan

Hexaprotodon karumensis Kolpochoerus limnetes Kolpochoerus majus Kolpochoerus olduvaiensis Kolpochoerus evronensis Kolpochoerus sp. Metridiochoerus compactus Metridiochoerus modestus Metridiochoerus hopwoodi Metridiochoerus andrewsi Metridiochoerus sp. Phacochoerus africanus Phacochoerus aethiopicus Phacochoerus sp. Potamochoerus porcus Sus scrofa Camelus thomasi Camelus sp. Giraffa pomeli Giraffa camelopardalis Giraffa jumae Giraffa stillei Giraffa sp. Sivatherium maurusium Sivatherium olduvaiensis Cervus elephas Capreolus sp. Dama mesopotamica Praemegaceros verticornis Bovini Bos bubaloides Bos primigenius Bos sp. Pelorovis antiquus Pelorovis oldowayensis Pelorovis sp. Bison priscus Syncerus caffer Syncerus acoelotus Syncerus sp. Tragelaphini Tragelaphus algericus Tragelaphus strepsiceros Tragelaphus scriptus

cf.

cf. cf.

X

X

cf.

cf.

X

X

X

X

X

X

X

X

cf.

cf.

X

X

cf.

X

X

X

X

cf.

X

X

X

X

X

X X X

X X

X

X

cf. cf.

cf.

cf.

cf.

cf.

cf.

cf.

X X cf.

X X

X

X

cf.

X

X

X

X

X X

X

X X X

X X

X

?

X

X

X

X

X

X

X

cf.

X

cf.

X X

cf.

cf.

X

X

X

X

X

X

X

X

cf.

cf.

X

cf.

cf.

X X

X

X

X

X

X

X

Tragelaphus sp. Taurotragus oryx Taurotragus arkelli Taurotragus sp. Cephalophini Cephalophus sp. Sylvicapra sp. Neotragini Raphicerus campestris Raphicerus melanotis Ourebia sp. Oreotragus oreotragus Antilopini Gazella atlantica Gazella dracula Gazella gazella Gazella sp. Antidorcas recki Antidorcas australis Antidorcas bondi Antidorcas sp. Caprini Capra sp. Bouria anngettyae Ovibovini Makapania sp. Nitidarcus asfawi Alcelaphini Alcelaphus buselaphus Alcelaphus sp. Connochaetes taurinus Connochaetes gnou Connochaetes sp. Megalotragus priscus Megalotragus kattwinkeli Megalotragus sp.

Evro

cf.

cf.

Gesh

X

cf.

TQ1L cf.

Tigh X

X

cf. X

Lain X

TQ1G X

cf.

Old3 X

X

X

X

X

cf.

X

X

X

X

X

X

cf.

cf.

X

X

cf.

X

Olo1p X

Daka

East Africa Olo6/7

North Africa Old4 X

X

X

X

X

X

Kanj X

Bodo X

Kapt cf. X

X

X

X

X X

X

Plov X X

X

X

Southern Africa

X

X

X

X

X

X

X

X

Glad

Levant

X

X

X

cf.

X

Corn

Table 2 continued.

X

X

X X X

X

X

Elan

Kasi

Buia

Lata

Rabaticeras arambourgi Rabaticeras sp. Damaliscus agelaius Damaliscus pygargus Damaliscus lunatus Damaliscus niro Damaliscus sp. nov. Damaliscus sp. Parmularius ambiguus Parmularius rugosus Parmularius angusticornis Parmularius sp. nov. Numidocapra crassicornis Aepycerotini Aepyceros melampus Aepyceros sp. Hippotragini Hippotragus gigas Hippotragus leucophaeus Hippotragus equinus Hippotragus niger Oryx gazella Oryx sp. Reduncini Kobus ellipsiprymnus Kobus kob Kobus leche Kobus sigmoidalis Kobus sp. Thaleroceros radiciformis Redunca arundinum Redunca fulvorufula Redunca sp. Peleini Pelea capreolus Perissodactyla Hipparion libycum Hipparion ethiopicum Hipparion sp. Equus mauritanicus Equus algericus Equus altidens Equus capensis

cf.

cf.

X

?

cf.

cf.

X

cf. cf.

X

?

cf.

X

X

X

cf.

X

X X

X

X

X

X

X

X

X X

cf.

X

X

X

X

X

cf.

?

X

X

X

X X

X X

X

X

X

cf.

cf.

cf.

X

X

X

cf.

X

X

X

X

X

X

X X

X

X X cf.

cf.

X

X

X

X

cf.

cf.

X

X

X

X X

?

aff. ? ?

X

Evro

X

X

Gesh

X

X

X

X

X

X

Lata

X

X cf.

X

TQ1L X

X

Tigh X

X

X

X

Lain X

X

X

TQ1G X

cf.

X

Buia X

X

X X

Kasi X

X

Old3 X

X

?

X

X

Olo1p X

X

X

X cf.

cf.

X

cf. cf.

X

X

X

X

X

X

X cf.

Old4 X

X

X

X

X

X

X

X X

X

X

X

X

X

Kanj

Differing levels of uncertainty are indicated by cf., ? and aff. Site code abbreviations are explained in Table 1. See Appendix for further details.

Equus burchelli Equus quagga Equus oldowayensis Equus grevyi Equus sp. Ceratotherium simum Ceratotherium sp. Dicerorhinus hemitoechus Diceros bicornis Proboscidea Loxodonta atlantica Loxodonta africana Elephas antiquus Elephas iolensis Elephas recki Elephas sp. Mammuthus trogontherii Stegodon trigonocephalus Stegodon sp. Pholidota Phataginus giganteus Phataginus sp. Tublidentata Orycteropus sp. Primates Cercopithecus sp. Cercopithecoidea kimeui Colobus sp. Theropithecus oswaldi Papio hamadryas Papio sp. Homo erectus Homo sp. Homo sp. (artefacts)

Daka

East Africa Olo6/7

North Africa Bodo X

X cf.

X

X cf.

Kapt X cf.

X cf.

X

X

X

X

Plov X

X

Southern Africa

X

X

Glad

Levant

X cf.

cf.

Corn

Table 2 continued.

X

X

X

X

X

X

?

Elan

AFRICAN FAUNAL CHANGE

241

Fig. 2. Results of a TWINSPAN analysis of all specieslevel identifications from the 21 localities included in our analysis. Site code abbreviations are explained in Table 1.

Fig. 4. Results of a TWINSPAN analysis of all specieslevel identifications from the 14 sub-Saharan sites included in our analysis. Site code abbreviations are explained in Table 1.

Fig. 3. Results of a TWINSPAN analysis of all specieslevel and isolated generic identifications (including four genera that were amalgamated at the species level – Ursus, Camelus, Hipparion and Stegodon) from the 21 localities included in our analysis. Site code abbreviations are explained in Table 1.

To see if we could detect any signs of a turnover within the fauna over time we restricted the analysis to the sub-Saharan localities, a total sample of 14 assemblages. The results of this can be seen in Figure 4. This shows that at the first level of division the south African sites were separated from the east African and there were no further divisions within the south African grouping, suggesting that these faunas are very similar. At the second level of division, Olduvai Beds 3 and 4, the Daka Member and Kasibos divided from the rest. The final division led to Buia, Bed 3ii of the Kapthurin Formation and the Bodo Formation being grouped together with a second group of Kanjera Member Kn-5, and Olorgesailie Upper Member 1 palaeosol and member 6/7. These groupings do not reflect the ages of the localities since Buia is dated to c. 1.0 Ma whereas Bed 3ii of the Kapthurin Formation is the youngest locality in the analysis having been dated to c. 0.5 Ma (see Table 1).

It appears from the above analyses that the pattern seen in the Early–Middle Pleistocene faunas of Africa is largely based on biogeography and that this signal overprints any more subtle changes in taxa that may have taken place. The idea that the Levant was a route of faunal exchange into and out of east Africa and should therefore have similar species occurrences to the east African localities is not supported by our results. The Levantine sites were the first to be separated from all others in each of the four analyses, suggesting a distinct biogeographic signal rather than a merging or exchange between two zones. There are some fine-level changes in faunal turnover that can be seen in Table 2 and these are discussed below.

Patterns of geographic and temporal distribution Carnivora The carnivores are largely uninformative because they are relatively rare and are often found in small numbers when they do occur at sites. Only Thomas Quarry 1, Level G and the Elandsfontein Main site have large numbers of carnivores. However, north Africa did provide the only records of bears (Ursus sp.) in our analysis. Although bears are frequently recorded from Quaternary sites in Europe and Asia they were not present at any of our sites in the Levant.

Geographic differences Several animals have interesting geographic distributions. The Levant and north Africa each have an

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endemic species of Hippopotamus (H. behemoth and H. sirensis). Hippopotamus gorgops is only found in sub-Saharan Africa, in particular east Africa, but there is a single record of this species from Cornelia in south Africa. The giraffid Sivatherium is only recorded from east and south Africa. Several animals are only found in the Levant, including the cervids of the genera Dama, Capreolus, Cervus and Praemegaceros, the bison (Bison priscus) and the extinct proboscidean Stegodon. This is an interesting distribution for the genus Stegodon, since although it persisted in Asia until relatively recently it became extinct in Africa in the Pliocene (Kalb 1995). In contrast, the archaic three-toed equid Hipparion was replaced in Europe by the monodactyl horses of the genus Equus at 2.4 Ma (Agustí & Antón 2002), and yet it was still present in the Bodo Formation in east Africa as late as 0.6 Ma. There are in fact three late records of Hipparion in Table 2: Bodo members u and u-t and Olduvai Bed 4 in east Africa, and Cornelia in south Africa. The Bodo members have been dated to 0.6 Ma using 40Ar/39Ar dating, Olduvai Bed 4 is thought to date to between 0.8 and 0.6 Ma whereas the Cornelia beds are not directly dated, but were placed by McKee et al. (1995) between Plovers Lake and the Elandsfontein Main site on the basis of faunal resemblance indices, suggesting an age of c. 0.7 Ma. Hipparion disappeared from Europe during the Late Pliocene, but is generally thought to have survived in Africa until c. 1.0 Ma. However, its presence at these localities, in particular Olduvai Bed 4, suggests that it did not disappear in Africa until after the Matuyama–Brunhes boundary, although the precise time of its final extinction is not known. The genus Equus was very successful, with different species being found on almost every site sampled. Two other predominantly European taxa are present in the Levantine sites: the rhinoceros Dicerorhinus (Stephanorhinus) hemitoechus and Mammuthus trogontherii. The finding of both these species together at Latamne in Syria may indicate time-averaging of the fauna, since D. hemitoechus is regarded as an interglacial species, whereas M. trogontherii is usually found in glacial faunas (Agustí & Antón 2002). The date of Latamne is also controversial; it is generally accepted as being 0.7–0.5 Ma (the date we have used here), but Tchernov et al. (1994) have suggested it may be closer in age to ‘Ubeidiyah (c. 1.4 Ma) and Evron Quarry (recently redated to c. 1.0 Ma; Ron et al. 2003). In our analyses it was grouped with Gesher Benot Ya’aqov (dated to c. 0.78 Ma), which lends support to the later date. Within the tribe Bovini, members of the genus Bos were only found in the Levant and north Africa, while species of the extinct genus Pelorovis and the extant genus Syncerus were only present in east and south Africa. However, a recent paper by

Alemseged & Geraads (2000) has reported a partial cranium from Eritrea which appears to be Bos; indeed they tentatively identified it as Bos cf. primigenius, commonly known as the aurochs. As this site has been dated to between 0.8 and 0.2 Ma, we have not included it in our analysis, but it is worth noting that this record marks the first finding of the genus Bos in sub-Saharan Africa prior to its arrival there as domestic cattle in recent times (Alemseged & Geraads 2000). Other species of note are those with pan-African distributions (found in north, east and south Africa), including the blue wildebeest (Connochaetes taurinus), the white rhinoceros (Ceratotherium simum), Rabaticeras arambourgi and Theropithecus oswaldi. Only two animals, the genus Oryx and the extinct elephant Loxodonta atlantica, have very disjunct distributions in the non-contiguous regions of north and south Africa. From this review and the results of our TWINSPAN analysis it seems that there are distinct differences between the regions in terms of their fauna, with the Levant in particular standing out as having a number of Eurasian taxa that do not appear to have migrated into Africa. Several animals are found throughout the African continent, and these tend to be the larger-bodied animals such as rhinoceroses, two large alcelaphines and the extinct theropithecine primates and Homo erectus. Sub-Saharan Africa contained several genera that had earlier become extinct elsewhere, such as the giraffid Sivatherium and the three-toed horse Hipparion.

Chronological patterns The chronology of species occurrences is of great interest as it is here that we may be able to pick up some of the fine-scale changes in taxon appearance and extinction that mark the Early–Middle Pleistocene boundary. However, these require more careful interpretation than geographic appearance of species (indicated by simple presence and absence), because the animals may be present both before and after this interval, but for some reason are absent from the faunal lists of the localities in this study. The case of the spotted hyaena (Crocuta crocuta) is an example; as can be seen in Table 2 it is only present at sites in our dataset after 0.7 Ma, yet we know that it was present in Africa long before this and that it is still extant (Turner 1990). The other cautionary note is sounded by the presence of Makapania sp. in the Gladysvale External Deposits, dated between 0.78 and 0.58 Ma (Lacruz et al. 2003). Prior to this discovery the youngest record of the genus was at Sterkfontein Member 4, dated to approximately 2.6–2.4 Ma (Vrba 1995). This indicates that any work of synthesis is likely to change relatively

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quickly, as the publication of one new locality can have a dramatic effect on our understanding of the geographic and temporal range of particular genera or species. Therefore some of the more detailed conclusions of this paper are likely to change as more sites are discovered. The late appearance of Dinofelis piveteaui in the Kasibos Formation is worth discussion. Prior to the publication of a partial skeleton from this formation (Ditchfield et al. 1999), the youngest accepted date for this species was c. 1.4 Ma (Turner & Antón 1999; Werdelin & Lewis 2001). The Kasibos Formation has normal polarity yet contains some older species such as Kolpochoerus limnetes and Metridiochoerus cf. modestus, while the presence of Kolpochoerus majus indicates that it postdates 1.8 Ma (Ditchfield et al. 1999). It has therefore been dated to the Jaramillo Subchron (1.07–0.99 Ma) as this is the only major interval of normal polarity in the Early Pleistocene before the Matuyama–Brunhes boundary. The published evidence for the date, based on both biostratigraphy and palaeomagnetic data, suggests no compelling reason why the Kasibos deposit should not be assigned to the normally magnetized Olduvai Subchron (c. 1.8–1.6 Ma). However, there are reasons to believe the date to be later as Acheulean bifaces have been found on the surface of the Kasibos Formation (Ditchfield et al. 1999). The earliest currently known Acheulean assemblage is dated to c. 1.4 Ma at Konso (Asfaw et al. 1992) and it is unlikely that the Kasibos Formation artefacts predate this. Various species have their last appearance in the 1.0–0.5 Ma interval. These need to be interpreted with caution, as Damaliscus pygargus ( D. dorcas (Wilson & Reeder 1993)) seems to have its last appearance at Olduvai Bed 4, yet it remains extant in Africa. However, there are some apparent definitive changes in the African fauna in this interval. Several new species appear that continue to the present day, including the modern hippopotamus (Hippopotamus amphibius), which first appears at approximately 0.78 Ma at Kanjera Member KN-5 and Gesher Benot Ya’aqov. The first appearance of the modern reedbuck (Redunca arundinum) is also at this time. According to Vrba (1995), Connochaetes gnou also appears around 0.7 Ma and this is confirmed by findings at Cornelia and Elandsfontein. A member of the genus Parmularius has also been reported from Elandsfontein, and the extinction of this group had hitherto been taken as c. 0.7 Ma (Vrba 1995). Elandsfontein is currently dated to between 0.7 Ma and 0.4 Ma (Klein & Cruz-Uribe 1991), but the presence of Parmularius would suggest that the site falls in the earlier part of this range. However, the assignment to Parmularius is extremely tentative and there is a suggestion that the specimen may even represent an entirely new member of the tribe

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Alcelaphini (Klein & Cruz-Uribe 1991). The date of Elandsfontein was assigned on the basis of the fauna present, but as it has the largest faunal list of any of the south African sites, includes many carnivores and is likely to be highly time-averaged, the date of 0.7–0.4 Ma must be regarded as uncertain. The same is true of Cornelia, where the fauna from several beds has been lumped into one list, and may include later contaminants, although Cooke (1974) thought this unlikely. The extinct alcelaphine Megalotragus kattwinkeli is last recorded in Olduvai Bed 4, whereas the first appearance of the modern hartebeest (Alcelaphus buselaphus) is at 0.6 Ma in the Bodo Formation. Several species occur from 1.0 Ma to 0.5 Ma, including Hippopotamus sirensis, Hippopotamus gorgops, Equus capensis, Ceratotherium simum, Loxodonta atlantica (although again, this may be an artefact of the young date given for Elandsfontein), Elephas recki, Homo erectus and Theropithecus oswaldi. Interestingly, there is a gap in the presence of Kobus ellipsiprymnus between 0.8 Ma and 0.6 Ma when it is absent from the nine sites dated to this interval. It therefore appears that some change in fauna did take place between 0.8 Ma and 0.7 Ma in Africa, but that this was not marked enough to override the strong geographic signal between and within the regions looked at in our TWINSPAN analysis.

Discussion: changing faunas at the Early–Middle Pleistocene boundary? This study has concentrated on faunal change at the Early–Middle Pleistocene boundary, in particular around the time of the switch from the Matuyama (reversed) to the Brunhes (normal) chrons in the palaeomagnetic timescale. The interval between 1.0 and 0.5 Ma also saw a major shift in the Earth’s climate from the relatively small-scale oscillations in the Pliocene and Early Pleistocene to the major oscillations that characterize the Middle and Late Pleistocene. This global transition (often referred to as the mid-Pleistocene transition, MPT) occurred between 0.92 Ma and 0.65 Ma. During the MPT, a large increase in global ice volume was followed by a change in the frequency of the Quaternary ice ages as the 41 ka glacial cyclicity was replaced by the 100 ka glacial cyclicity that continues to the present day (Shackleton & Opdyke 1976; Mudelsee & Schulz 1997). There is some disagreement over the speed of this transition from the shorter to a longer cyclicity. Some workers have suggested that the change was prolonged over several hundreds of thousands of years, with Ruddiman et al. (1989) arguing for a gradual transition between 0.9 Ma and 0.4 Ma although accepting major changes between 0.7 Ma and 0.6

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Ma. However, the current consensus seems to be for an abrupt change, at around 0.65 Ma, some time after the large increase in global ice volume at 0.92 Ma (Lau & Weng 1995; Mudelsee & Schulz 1997). During this major climatic perturbation, the global thermohaline circulation was weakened due to a reduction in North Atlantic Deep Water ventilation, which Schefuß et al. (2003) conclude led to warmer sea surface temperatures (SSTs) in the tropical Atlantic Ocean. The strength of the monsoon system that affects Africa depends on the degree of contrast between SSTs and continental landmass temperatures. Changes to tropical Atlantic SSTs will affect the nature and strength of the African monsoon and thus influence aridity on the African continent. Records of the nature of the African climate from this interval are found mostly from deep-sea cores and some isolated terrestrial records. Schefuß et al. (2003) used an ocean-core record of wind-transported plant waxes to infer changes in continental C3/C4 plant variations during the mid-Pleistocene transition. Large-scale vegetation changes on the African continent respond primarily to changes in aridity, temperature and CO2 concentration (see Schefuß et al. 2003, 2005 for review). Before 1.05 Ma, the African flora did not seem to respond to the glacial–interglacial cycles, but between 1.05 Ma and 0.9 Ma glacials (cold stages) resulted in decreases in river discharge in central Africa and the spread of Podocarpus into the central mountainous belt. During the MPT, between 0.9 Ma and 0.6 Ma, interglacials were generally warm and arid, whereas glacials were generally cool and humid in Africa (Dupont et al. 2001). After the transition to a 100 ka cycle, glacials on the continent eventually became characterized by colder and drier conditions and interglacials by warmer and humid conditions. Micromammalian evidence from the Early to Middle Pleistocene site of Kabwe, central Zambia, suggests that the Homo spp. present there occupied central Zambia only in conditions similar to today, i.e. interglacial periods (Avery 2003). This finding also seems to be true for other regions (e.g. northern south Africa). Alternatively, it may also be true that glacial–interglacial changes were not strong enough to cause much change in the low latitudes before 0.9 Ma (cf. Dupont et al. 2001) or that simply glacial-age occupation sites have yet to be found. This major change in climatic regime and terrestrial vegetation would presumably have had some effects on the animals that inhabited the African continent. There is certainly evidence from other parts of the world, notably Europe, that there was a major faunal shift beginning at 0.9 Ma, which has been termed the ‘end-Villafranchian event’ although it appears to have been of quite long duration (Azzaroli et al. 1988; Turner 1995a). A distinct turn-

over event has also been observed in Indonesia, where a major faunal change occurred between 0.8 Ma and 0.7 Ma, on both Java and Flores. The Javan faunal change was marked by the arrival of modern genera such as Elephas from the Asian mainland following the lowering of sea levels in the first major glacial episode (van den Bergh et al. 2001), while on Flores Stegodon sondaari was replaced by S. florensis (van den Bergh et al. 2001), a change that coincided with the arrival of Homo erectus on the island, as shown by the presence of stone artefacts (Morwood et al. 1998). In Africa, however, the change to 100 ka cyclicity does not appear to have greatly affected the faunas because, although there are some changes, the turnover is largely attritional rather than a marked event. At c. 0.7 Ma several new species do seem to appear, including the modern hippopotamus, reedbuck and black wildebeest, while members of the long-lived genus Parmularius become extinct. However, none of these appearances are definite enough to use as stratigraphic markers, because several of the localities have uncertain dates and, as discussed above, one new locality can completely change our perception of a species’ distribution in both time and space. The genus Homo is ubiquitous in these faunal lists because of the durability of the stone artefacts from which its presence can be inferred, and because archaeological sites tend to be preferentially published, which increases the bias associated with this faunal element. A further problem is that new hominin or archaeological sites tend to be published with very brief faunal lists, with no information on the specimens themselves, which makes evaluating the accuracy of the identifications impossible. However, from our observations it appears that the biogeography of the African fauna appears to be of much greater importance than the dates of the sites themselves (and therefore the climate at that time), hence the sites within different geographic regions cluster together and do not separate by age, even when the sample is restricted to sub-Saharan Africa.

Conclusions The Early–Middle Pleistocene boundary appears to have been a time of great change that had an effect on various mammal communities throughout the world. However, the African large mammal faunas, with the exception of the species discussed above, seem to have remained relatively stable throughout this interval in comparison with faunas from other continents. It does not therefore appear that the African large mammal record can be used to pinpoint the Early–Middle Pleistocene boundary. It would appear, however, that the Matuyama–Brunhes

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Chron boundary does provide a reliable signal for this boundary (despite the obvious problems of correlation) and it can be located world-wide, providing a baseline for researchers working on any of the seven continents. In summary, the African Early– Middle Pleistocene faunas seem to be heavily influenced by geographic locality and not by chronology. The observed patterns of distribution of the modern zoogeographic communities in the Levantine and north African areas of the Palaearctic and those of the sub-Saharan African region can be seen as far back as the Early Pleistocene. We would like to thank T. Plummer and R. Potts for allowing us to use unpublished data in this analysis, T. Clare and D. Wilkinson for help with TWINSPAN, D. Geraads for information on Thomas Quarry 1 and R. Lacruz for a preprint of the Gladysvale paper. Much of the research for this paper was conducted under a NERC grant within the EFCHED Programme.

Appendix: Comprehensive lists of large mammal faunas for all localities included in this study Levels of uncertainty, such as ‘cf’ or ‘?’, reflect those of the original publications cited. Suid taxonomy follows Harris & White (1979). Levant Evron Quarry (Tchernov et al. 1994) Crocuta or Hyaena, Hippopotamus sp., Kolpochoerus evronensis, cf. Cervus (?) elaphus, cf. Capreolus sp., Bos cf. primigenius, Gazella cf. gazella, cf. Alcelaphus sp., Elephas sp., Stegodon sp. Gesher Benot Ya’aqov (Geraads & Tchernov 1983; Bar-Yosef 1998) Hippopotamus (?) amphibius, Sus scrofa, Cervus cf. elephas, Dama cf. mesopotamica, Bos sp., Gazella cf. gazella, Capra sp., Equus sp., Dicerorhinus hemitoechus, Elephas antiquus, Stegodon sp., Homo erectus. Latamne (Guérin et al. 1993) Canis sp. Crocuta crocuta, Hippopotamus cf. behemoth, Camelus sp., Giraffa camelopardalis, Praemegaceros verticornis, Bos primigenius, Bison priscus, Bovidae de type antilope, gen. et sp. indet. cf. Pontoceros (?), Equus cf. altidens, Dicerorhinus hemitoechus, Mammuthus trogontherii, Stegodon cf. trigonocephalus. North Africa Thomas Quarry 1, Level L (Geraads 2002) Hippopotamus cf. sirensis, Kolpochoerus sp., Gazella cf. atlantica, Alcelaphini indet., Equus cf. mauritanicus, Loxodonta atlantica, Homo erectus (inferred from archaeology).

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Tighenif (Geraads 2002) Canis cf. adustus?, Ursus?, Crocuta crocuta, Panthera leo?, Homotherium sp., Hippopotamus sirensis, Metridiochoerus compactus, Camelus thomasi, Giraffa cf. pomeli, Bos cf. bubaloides, Tragelaphus algericus, Gazella cf. atlantica, Gazella dracula, Gazella sp., Connochaetes taurinus prognu, Parmularius ambiguus, Hippotragus cf. gigas, Oryx cf. gazella, Kobus? sp., Equus mauritanicus, Ceratotherium simum, Loxodonta atlantica, Theropithecus oswaldi, Homo erectus. Aïn Maarouf (Geraads & Amani 1997) Crocuta sp. ( Crocuta crocuta), Hippopotamus cf. sirensis, Bovini indet., Tragelaphini indet.?, Connochaetes sp., Rabaticeras arambourgi?, Parmularius ambiguus, Equus cf. mauritanicus, Equus cf. algericus, Loxodonta atlantica, Elephas iolensis, Homo erectus Thomas Quarry 1, Level G (Homo erectus Cave) (Geraads 2002) Canis aff. adustus, Lycaon cf. magnus, Ursus bibersoni, Lutra cf. maculicollis, Mellivora sp., Crocuta crocuta, Hyaena hyaena, Panthera leo, Monachus aff. monachus, Hippopotamus cf. sirensis, Phacochoerus cf. africanus, Bovini indet., Gazella cf. atlantica, Connochaetes taurinus prognu, Alcelaphini indet., Oryx sp., Equus cf. mauritanicus, Ceratotherium simum, Elephantidae indet., Theropithecus cf. oswaldi, Homo erectus. East Africa Buia (Abbate et al. 1998) Hyaenidae indet., Hippopotamus gorgops, Hexaprotodon karumensis, Kolpochoerus cf. K. majus, cf. Kolpochoerus sp. (advanced form), Pelorovis sp., Kobus ellipsiprymnus, Bovidae indet., Hipparion sp., Equus sp., Ceratotherium simum, Rhinocerotidae indet., Elephas recki, Homo sp. Kasibos (Ditchfield et al. 1999) Canidae, Felidae, Dinofelis piveteaui, Hippopotamidae large, Kolpochoerus majus, Metridiochoerus cf. modestus, Bovini, Tragelaphini, Neotragini, Antilopini, Alcelaphini, Reduncini, Equus sp., Equidae, Elephantidae, Cercopithecidae. Olduvai Bed III (Leakey 1994) Canidae indet., Lutrinae indet., Hyaenidae indet., Hippopotamus gorgops, Kolpochoerus limnetes, Kolpochoerus major ( K. majus), Metridiochoerus compactus, Metridiochoerus modestus, Metridiochoerus hopwoodi, Sivatherium maurusium, Pelorovis oldowayensis, Syncerus aceolatus, Tragelaphus strepsiceros grandis, Antidorcas recki, Connochaetes taurinus olduvaiensis, Megalotragus kattwinkeli, Rabaticeras arambourgi, Damaliscus niro, Damaliscus agelaius, Parmularius rugosus, Hippotragus gigas, Kobus ellipsiprymnus, Kobus

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kob, Redunca sp., Hipparion cf. ethiopicum, Equus oldowayensis, Ceratotherium simum, Diceros bicornis?, Elephas recki, Colobinae indet., Theropithecus oswaldi. Olorgesailie Upper Member 1 palaeosol (R. Potts pers. comm.) Canis cf. mesomelas, Crocuta sp. nov., Hippopotamus cf. gorgops, Metridiochoerus cf. compactus, Metridiochoerus cf. hopwoodi, Phacochoerus cf. aethiopicus, Suidae cf. Phacochoerus, Suidae cf. M. compactus, Suidae cf. M. hopwoodi, Suidae indet., Giraffa sp., Pelorovis sp., Tragelaphus sp., Taurotragus sp., Antilopini cf. Gazella sp., Alcelaphini cf. Alcelaphus, Alcelaphini cf. Connochaetes, Connochaetes sp., Megalotragus sp., Redunca sp., Bovidae indet., Hipparion sp., Equus cf. burchelli, Equus oldowayensis, Equus cf. grevyi, Ceratotherium simum, Elephas recki, Theropithecus oswaldi. Daka (Asfaw et al. 2002) cf. Pachycrocuta aff. brevirostris, cf. Panthera leo, Hippopotamus gorgops, Kolpochoerus majus, Kolpochoerus olduvaiensis, Metridiochoerus compactus, Metridiochoerus modestus, Metridiochoerus cf. hopwoodi, Giraffa cf. camelopardalis, Sivatherium cf. maurusium, Pelorovis cf. antiquus, Tragelaphus cf. strepsiceros, Gazella sp., Bouria anngettyae, Nitidarcus asfawi, Connochaetes taurinus, Megalotragus kattwinkeli, Rabaticeras sp., Parmularius angusticornis, Numidocapra crassicornis, cf. Aepyceros sp., Kobus ellipsiprymnus, Kobus kob, Kobus sigmoidalis, Hipparion sp., Equus sp., Ceratotherium sp., Elephas recki recki, cf. Colobus sp., Cercopithecinae gen. indet., Theropithecus cf. oswaldi, Homo erectus. Olorgesailie Member 6/7 (R. Potts pers. comm.) Genetta sp., Hippopotamus cf. gorgops, Metridiochoerus cf. compactus, Suidae indet., Giraffa sp., Pelorovis sp., Tragelaphus sp., Taurotragus sp., Antilopini cf. Gazella sp., Alcelaphini cf. Connochaetes, Megalotragus sp., Aepyceros sp., Redunca sp., Bovidae indet., Hipparion sp., Equus cf. grevyi, Equus oldowayensis, Ceratotherium simum, Elephas recki, Theropithecus oswaldi. Olduvai Bed IV (Leakey 1994) Canidae indet., Aonyx sp., Viverridae indet., Hyaenidae indet., Panthera leo, Hippopotamus gorgops, Kolpochoerus limnetes, Kolpochoerus major (K. majus), Metridiochoerus compactus, Metridiochoerus modestus, Metridiochoerus hopwoodi, Phacochoerus antiquus (P. africanus), Giraffa jumae, Giraffa stillei, Sivatherium maurusium, Pelorovis antiquus, Syncerus aceolatus, Tragelaphus strepsiceros grandis, Tragelaphus aff. spekei or scriptus (Tragelaphus sp.), Taurotragus arkelli, Antidorcas recki, Antidorcas sp., Antilopini

indet., Connochaetes taurinus olduvaiensis, Megalotragus kattwinkeli, Rabaticeras arambourgi, Damaliscus niro, Damaliscus agelaius, Parmularius rugosus, Alcelaphini sp. 1, Thaleroceros radiciformis?, Hipparion cf. ethiopicum, Equus oldowayensis, Ceratotherium simum, Diceros bicornis, Elephas recki, Colobinae indet., Cercopithecoidea kimeui, Theropithecus oswaldi, Papionini indet., Homo erectus. KN-5, Kanjera (T. Plummer pers. comm.) Carnivora indet. (medium), Hippopotamidae indet. (pygmy), Hippopotamus cf. gorgops, Hippopotamus cf. amphibius, Kolpochoerus limnetes, Kolpochoerus sp., Metridiochoerus hopwoodi, Metridiochoerus sp., Phacochoerus sp., Giraffa sp., Pelorovis sp. ?, Syncerus acoelotus, Tragelaphus sp. (small), Tragelaphini (large), Antilopini indet., Alcelaphini indet. (medium), Kobus sp., Reduncini indet. (medium), Equus sp., Diceros bicornis, Elephas recki, Loxodonta africana, Phataginus giganteus, Colobus sp. Theropithecus oswaldi. Bodo (Kalb et al. 1982a; Clark et al. 1994) Carnivora indet., Hippopotamus cf. gorgops, Kolpochoerus limnetes, Kolpochoerus majus, Metridiochoerus cf. jacksoni (M. andrewsi), Giraffa sp., cf. Pelorovis sp., Syncerus acoelotus, Alcelaphus buselaphus, Damaliscus niro, Aepyceros cf. melampus, Kobus cf. ellipsiprymnus, cf. Kobus leche, Hipparion sp., Equus sp., Ceratotherium cf. simum, Rhinocerotidae gen. et sp. indet., Elephas recki, Theropithecus oswaldi, Papio cf. hamadryas, Homo sapiens cf. rhodesiensis. K3(ii), Kapthurin (McBrearty et al. 1996) Ichneumia sp., Crocuta crocuta, Panthera leo, Hippopotamus amphibius, Kolpochoerus majus, Phacochoerus aethiopicus (P. africanus), Potamochoerus porcus, cf. Syncerus caffer, Tragelaphus cf. scriptus, Tragelaphus sp., Cephalophus sp., Sylvicapra sp., Ourebia sp., Gazella sp., Antidorcas sp., Alcelaphus buselaphus, Alcelaphus sp., Damaliscus sp., Oryx sp., Kobus cf. ellipsiprymnus, Kobus sp., Redunca sp., Ceratotherium simum, Elephas sp., Orycteropus sp., Cercopithecus sp., Colobus sp., Theropithecus cf. oswaldi, Papio sp., Homo cf. erectus. South Africa Plovers Lake (Thackeray & Watson 1994) Canis mesomelas, Panthera pardus, Metridiochoerus andrewsi, Taurotragus oryx, Tragelaphus strepsiceros, Antidorcas bondi, Alcelaphus and/or Damaliscus, Connochaetes sp., Megalotragus priscus, Kobus ellipsiprymnus, Pelea capreolus, Equus capensis, Equus burchelli, Papio hamadryas robinsoni.

AFRICAN FAUNAL CHANGE

Gladysvale External Deposits (Lacruz et al. 2003) Canis mesomelas, Hyaenidae indet., Panthera leo, cf. Phacochoerus sp., Giraffidae indet., Pelorovis sp., Syncerus sp., Tragelaphus strepsiceros, Taurotragus oryx, Raphicerus campestris, Oreotragus oreotragus, Antidorcas bondi, Makapania sp., Connochaetes taurinus, Connochaetes sp., Megalotragus sp., Damaliscus dorcas (D. pygargus), Alcelaphini indet. (medium sized), cf. Aepyceros sp., Hippotragus equinus, Hippotragus niger, cf. Oryx gazella, Kobus leche, Redunca arundinum, Redunca fulvorufula, Equus capensis, Equus burchelli, Papionini indet., Homo sp. Cornelia (Cooke 1974; updated by McKee et al. 1995) Hippopotamus gorgops, Stylochoerus compactus (Metridiochoerus compactus), Tapinochoerus modestus (Metridiochoerus modestus), “Afrochoerus” nicoli (Metridiochoerus compactus), Kolpochoerus sinuosus ( K. limnetes), Giraffa cf. camelopardalis, cf. Sivatherium olduvaiensis, Pelorovis cf. bainii (P. antiquus), cf. Tragelaphus strepsiceros, Taurotragus cf. oryx, cf. Sylvicapra grimmia, Antidorcas wellsi (A. recki), Connochaetes laticornutus (C. gnou), Megalotragus eucornutus (M. priscus), cf. Damaliscus sp., Damaliscus niro, cf. Kobus venterae (Kobus cf. leche), Redunca cf. arundinum, “Gazella helmoedi” (Bovidae indet.), Hipparion (Stylohipparion) steytleri (H. libycum), Equus cf. burchelli, Equus capensis, Equus plicatus (Equus sp.), cf. Ceratotherium simum. Elandsfontein Main site (Klein & Cruz-Uribe 1991) Canis mesomelas, Lycaon pictus, Ictonyx striatus, Mellivora capensis, Aonyx capensis, Viverra civetta, Herpestes ichneumon, Suricatta suricatta, Hyaena brunnea (Parahyaena brunnea), Crocuta crocuta, Felis caracal, Panthera leo, Megantereon gracile (M. cultridens), Hippopotamus amphibius, Kolpochoerus paiceae (K. limnetes), Metridiochoerus andrewsi, Sivatherium maurusium, Pelorovis antiquus, Tragelaphus strepsiceros, Taurotragus oryx, Raphicerus melanotis, Gazella sp., Antidorcas recki, Antidorcas australis, Connochaetes gnou, Megalotragus priscus, Rabaticeras arambourgi, Damaliscus aff. lunatus,? Damaliscus niro,? Damaliscus sp. nov.,? Parmularius sp. nov., Hippotragus gigas, Hippotragus leucophaeus, Redunca arundinum, Tribe indet., gen. et sp. nov., Equus capensis, Equus quagga?, Ceratotherium simum, Diceros bicornis, Loxodonta atlantica, Phataginus sp., Theropithecus oswaldi, ‘archaic’ Homo sapiens (Homo sp.).

247

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