Faunal evidence for mid- and late Quaternary
environmental change in southern Africa
james s. brink
Southern Africa is differentiated from other centres of aridity inAfrica by the
presence of an extended island of elevated, essentially treeless habitat in the
central interior, known as the Highveld and the Karoo. This area coincides
botanically with the Nama-Karoo and the Grassland Biomes. The large
geographic extent of this habitat is unique to southern Africa, since it has
no exact equivalent in modern-day east or north Africa. This uniqueness is
reﬂected in the large herbivores of the central interior, the grazers and mixed
feeders adapted to permanently available open habitat, which deﬁnes the
endemic faunal character of the subregion. This contribution presents some
of the faunal evidence for the appearance of permanently open habitat in
central southern Africa, a process that formed part of a longer-term trend of
faunal adaptation to aridiﬁcation and global cooling thatwas initiated within
the last 1 Ma, in a time known as the Cornelian Land Mammal Age (LMA).
A secondary and overlapping theme deals with the appearance of lakes and
wetlands on a subregional scale during the Florisian LMA, which lasted
from c. 0.6 Ma to the end of the Pleistocene/early Holocene. The end of the
Florisian LMA coincided with the regional extinction of wetland faunas in
the interior and with the extinction of specialised grazing ungulates over the
entire subregion, leading into the semi-arid conditions seen in the larger part
of modern-day southern Africa.
Climate change is often cited as a causal factor in biotic turnover (Vrba, 1995;
Behrensmeyer, 2006). It has been suggested that in high latitude areas, temperature
is the dominant climatic variable governing environmental change, but that
precipitation may be more dominant in low latitudes (deMenocal and Bloemendal,
1995). The southern African faunal evidence appears to be in accord, pointing to a
long-term trend towards drier climates throughout the Cenozoic (Pickford and
Senut, 1999). Southern Africa shares with north and east Africa a semi-arid to
arid climate, which gave rise to savannah grasslands, open grasslands and semi-
deserts, providing the environmental basis for the evolution of characteristic large
mammal faunas (Bigalke, 1978; Geraads, 1981; Skinner and Smithers, 1990;
Dupont and Leroy, 1995; Kingdon, 1997). In the south, the appearance of aridity
can be linked to the initiation during the mid-Cenozoic of the cold Benguela
Current along the southwest coast of Africa, and to major tectonic uplift of the
central plateau, following the separation of South America and Africa and the
breakup of the Gondwana supercontinent (Axelrod and Raven, 1978, Pickford and
Senut, 1999; Partridge and Maud, 2000; Tinker et al., 2008; McCarthy, 2013). This
caused an island of essentially treeless habitat, an area commonly known as the
Highveld and the Karoo, which botanically coincides with the Nama-Karoo and
the Grassland Biomes (Mucina and Rutherford, 2006) (Fig. 18.1). The large
herbivores of the central interior are mainly grazers and mixed feeders adapted
to the permanently available open habitat of this region. They deﬁne the endemic
faunal character of the subregion, and the marker taxa are the black wildebeest
Connochaetes gnou, blesbok Damaliscus pygargus, and springbok Antidorcas
marsupialis (Skinner and Smithers, 1990). To the north of the central interior,
savannah grasslands are found, which contain some woody components
(Fig. 18.1), but they are in structure and composition not very different from those
of east Africa (Kingdon, 1997).
The aim of this chapter is to review the faunal evidence for the appearance of
permanently open habitat in central southern Africa. This process was initiated
during the Cornelian Land Mammal Age (LMA) at ~1 Ma, and formed part of a
long-term trend of faunal adaptation to aridiﬁcation and global cooling. Open
grasslands became fully established during the Florisian LMA, which is the
subsequent faunal stage (Tables 18.1–18.3). In addition, faunal evidence is pro-
vided for the appearance of wetlands on a subregional scale during the Florisian.
The Florisian LMA lasted from ~0.6 Ma to the beginning of the Holocene and is
deﬁned by a number of now-extinct grazing ungulates, occurring in both the
interior and coast (see below), and by a wetland faunal component in the interior.
Wetlands are not unique to any given time, but their geographic extent during the
mid- to late Quaternary was signiﬁcant and provided a unique faunal signal that
intersected the open habitat of the central interior and surrounding savannah, and
excluded only the Cape montane and coastal areas (Fig. 18.1).
The thin strip of montane and coastal vegetation, which fringes the central
plains to the south and east of southern Africa, has a distinctive mammalian fauna,
Faunal evidence for Quaternary environmental change 285
adapted to more closed, woody habitat, as seen in the Cape Fynbos, Albany
Thicket and Indian Ocean Coastal Belt (Fig. 18.1). This area is biogeographically
peripheral to the central plains, and particularly in the Fynbos Biome there is a
unique endemic mammal fauna (Klein, 1983, 1984; Klein et al., 2007; Faith,
2013). Throughout the glacials of the mid- and late Quaternary, the Cape coastal
zone periodically experienced incursions of plains game from the interior when the
exposed continental margin provided additional habitat for grazing ungulates
(Klein, 1983; Brink, 1993, 2005). Here, this area is referred to mainly for the
insight that it provides in the early to mid-Quaternary, speciﬁcally the older
Cornelian materials from Elandsfontein, known as ‘Elandsfontein Main’(Klein
et al., 2007). The Cape Florisian shares a number of open-habitat taxa with the
Fig. 18.1. Biomes of southern Africa (shaded), showing the fossil localities
(numbered) referred to in the text and ﬁgures: 1. Florisbad, 2. Erfkroon, 3. Buffalo
Cave, 4. Makapan Limeworks, 5. Gladysvale External Deposits, 6, Sterkfontein,
7. Swartkrans, 8. Cornelia-Uitzoek, 9. Cornelia-Mara, 10. Wonderwerk Cave, 11.
Kathu Pan, 12. Elandsfontein Main, 13. Boomplaas Cave, 14. Klasies River, 15.
286 James S. Brink
Fig. 18.2. Revised Pleistocene biochronology for southern Africa, including the
name localities for the Land Mammal Ages (Makapan Limeworks, Cornelia-
Uitzoek and Florisbad). Data from Hendey (1974), Grün et al. (1996), Lacruz
et al. (2002), Herries et al. (2006, 2009); Chazan et al. (2008); Porat et al. (2010);
Brink et al. (2012) and Braun et al. (2013).
Faunal evidence for Quaternary environmental change 287
interior Florisian, of which it is an impoverished version. These taxa, including an
as yet unnamed caprine species, became extinct in the Cape ecozone at the end of
the Pleistocene, coinciding with a postglacial, eustatic rise in sea level and a
general reduction in grassland availability (Klein, 1983, 1984; Deacon et al.,
1984; Brink, 1999, 2005; Faith, 2013, 2014).
18.2 Mammalian turnover and biogeography in southern Africa
The mammalian fossil record of southern Africa is divided into a sequence of
Land Mammal Ages (LMA), which are periods of geologic time distinguished by
their distinctive faunal character over a large area, and are deﬁned by the presence
or absence of time-sensitive taxa (Savage and Russel, 1983). In descending order
of geological age and starting with the early Miocene, the southern African LMA
scheme includes the Namibian, Langebaanian, Makapanian, Cornelian, Florisian
and the Recent (Holocene) (Hendey, 1974; Klein, 1984) (Table 18.1). This
scheme was ﬁrst established during the mid-20th Century with attempts to create
chronological and spatial order out of the growing complexity of the late Ceno-
zoic fossil record (e.g. Cooke, 1952; Wells, 1962; Ewer and Cooke, 1964)
(Table 18.1). The use of local southern African names reﬂects the difﬁculty of
correlating with east Africa (Cooke, 1967), a trend that becomes more pro-
nounced after 1 Ma (Brink et al., 2012). More recent studies of Quaternary fossil
mammal studies in southern Africa have beneﬁtted from Electron Spin Reson-
ance (ESR), Optically Stimulated Luminescence (OSL) and palaeomagnetism
dating techniques that allowed a major revision of the chronology of mid- to
late Quaternary fossil mammal localities. It has now become possible to place the
name localities of the LMA scheme and other important assemblages in a more
accurate temporal order (Grün et al., 1996; Herries et al., 2009; Brink et al., 2012)
Table 18.1 Southern African Land Mammal Ages in relation to approximate geological
time periods, adapted from Hendey (1974).
Florisian Middle & late Pleistocene
Cornelian End-early Pleistocene
Langebaanian Early Pliocene
288 James S. Brink
The middle and late Pleistocene trend in southern African faunas towards
biogeographic uniqueness is tempered by evidence for some faunal interchange
with east Africa. This was ﬁrst noted by zoologists (e.g. Balinsky, 1962) who
proposed a southwest–northeast arid corridor linking the faunas of southwest
Africa and northeast Africa at various times in the past. Wells (1962) applied this
idea to the Pleistocene faunas of southern Africa. Previous biogeographic connec-
tions between southern and east Africa are evident also in historic and pre-
Holocene distribution patterns of plains zebra Equus quagga subspp., hartebeest
Alcelaphus buselaphus subspp, and blue wildebeest Connochaetes taurinus
subspp. (Geraads, 1981; Kingdon, 1997). Such taxa provide some basis for
correlation with east Africa during the mid- to late Quaternary. It is noteworthy
that genetic data support periodic biogeographic connections between southern and
east Africa. Changes between wetter and dryer climate periods may have allowed
dispersal corridors to develop, facilitating the migration of large mammals adapted
to varying palaeohabitats. In particular, the genetic data support the idea of
evolutionary stability during the late Quaternary in southern Africa, as opposed
to the biogeographic and evolutionary dynamism of east Africa at this time
(Lorenzen et al., 2012).
18.3 The southern African faunal record
18.3.1 The end of the early Pleistocene: the Cornelian LMA
Biogeographic exchange between southern and east Africa is more evident in the
earlier fossil record, such as the Makapanian assemblages (De Ruiter, 2003;
Gilbert, 2008; Bernor et al., 2010; Gentry, 2010; Werdelin and Peigné, 2010;
Reynolds and Kibii, 2011). The effects of these dispersals can be seen in the
herbivores of the Cornelian assemblages. Taxa shared by southern and east
Africa, but which became extinct in southern Africa before the Florisian, may
be considered archaic in the Cornelian context. Examples of archaic forms from
Cornelia-Uitzoek are the hipparion Eurygnathohippus cornelianus, the suid
genera Kolpochoerus and Metridiochoerus, and the hippo Hippopotamus gor-
gops (Van Hoepen, 1932, 1947; Cooke, 1974; Harris and White, 1979; Bernor
et al., 2010) (Table 18.2). At Elandsfontein, the archaic faunal component is
more pronounced and includes Theropithecus oswaldi,Megantereon whitei,the
genus Kolpochoerus,Hippotragus gigas, Numidocapra arambourgi and the
genera Sivatherium and Gazella (Gentry and Gentry, 1978; Klein et al., 2007).
Numidocapra arambourgi occurs at Buffalo Cave and Elandsfontein, but not at
Cornelia-Uitzoek or at Cornelia-Mara (Brink et al., 2012). The presence of a
gazelle and an extinct carnivore at Elandsfontein is particularly interesting, since
Faunal evidence for Quaternary environmental change 289
Table 18.2 Taxonomic list of Cornelian faunas: a comparison between Cornelia-Uitzoek,
Buffalo Cave and Elandsfontein Main. Extinct taxa also found in East Africa are
underlined. Faunal lists modiﬁed and adapted from Herries et al. (2006), Klein et al.
(2007) and Brink et al. (2012). †= Extinct
Bathyergus suillus -- X
Hystrix africaeasutralis -- X
Lepus capensis -- X
Homo sp. X - X
Cercopithecidae indet. - X -
Theropithecus oswaldi †-- X
Manis sp. - - X
Ictonyx striatus -- X
Mellivora capensis -- X
Suricata suricatta -- X
Viverra civetta -- X
Herpestes ichneumon -- X
Atilax paludinosus -- X
Canis mesomelas -- X
Vulpes chama -- X
Lycaon pictus -- X
Crocuta crocuta -- X
Parahyaena brunnea -- X
Felis libyca -- X
Felis caracal/serval -- X
Panthera leo X cf. X
Panthera pardus -- X
Megantereon whitei †-- X
Loxodonta atlantica zulu †-X
Indet. X - -
Eurygnathohippus cornelianus †XX -
Hipparionini indet. †-X -
Equus capensis †X- X
Equus quagga X- X
Equus sp. X - -
Diceros bicornis -- X
Ceratotherium simum -- X
Rhinocerotidae indet. X - -
Hippopotamus gorgops †X- -
Hippopotamus amphibius -- X
290 James S. Brink
both are absent from Buffalo Cave and Cornelia-Uitzoek (Table 18.2). They
may represent evolutionary remnants atypical of the Cornelian in general, and
probably reﬂect the biogeographically distant position of the Cape coastal
zone, where archaic forms are known to occur, as seen in the early Pliocene
locality of Langebaanweg (Gentry, 1980). Taxa that may reﬂect early Pleistocene
endemism in southern Africa are the giant wildebeest Megalotragus eucornutus,
and the impala Aepyceros helmoedi (Van Hoepen, 1932, 1947; Brink, 2005;
Damaliscus niro is found at Olduvai (Gentry and Gentry, 1978), but at Cornelia-
Uitzoek it is in an intermediate, Cornelian, stage of evolution (Thackeray and
Table 18.2 (cont.)
Phacochoerus sp. X X X
Metridiochoerus modestus †X- -
Metridiochoerus compactus †X- X
Kolpochoerus paiceae †X- X
Sivatherium maurusium †X- X
Taurotragus oryx -- X
Tragelaphus strepsiceros -- X
Syncerus antiquus †X? X
Redunca arundinum -? X
Hippotragus gigas †-? X
Hippotragus leucophaeus -- X
Damaliscus aff. lunatus -- X
Damaliscus niro †X- X
Damaliscus sp. - X -
Numidocapra arambourgi †-X X
Parmularius sp. †- X cf.
Megalotragus eucornutus †X- -
Megalotragus priscus †-- X
Connochaetes gnou laticornutus X- X
Connochaetes sp. - X -
Budorcas makapaani †-X -
Caprini indet. †-- X
Gazella sp. †-- X
Antidorcas recki †X- X
Antidorcas australis †-- X
Antidorcas bondi †X- X
Aepyceros helmoedi †X- -
Sylvicapra grimmia X- -
Raphicerus melanotis -- X
Raphicerus sp. X - -
Faunal evidence for Quaternary environmental change 291
Brink, 2004). D. niro became extinct in east Africa during the middle Pleistocene,
but in the south it continued to evolve into a Florisian stage, before the lineage
became extinct at the end-Pleistocene. An underived form of the extinct grazing
springbok Antidorcas bondi is abundant at Cornelia-Uitzoek, but occurs in small
numbers in the Elandsfontein Main assemblage and is so far not recorded at
Buffalo Cave (Herries et al., 2006; Brink et al., 2012). Later, during the Florisian
LMA, A. bondi evolved into a specialised grazer, characterised by extreme hypso-
donty (Brink and Lee-Thorp, 1992; Brink et al., 2013). It is noteworthy that, with
the exception of the long-horned buffalo Syncerus antiquus, none of the Cornelian
herbivore taxa that have Florisian descendants and which became extinct at the end
of the Florisian LMA are recorded in east Africa during the middle and late
Pleistocene (Faith, 2014) (Table 18.3). This reﬂects the increasingly regionalised
character of the large mammal faunas of southern Africa in relation to those of east
Africa after 1 Ma.
The Cornelian LMA was a time of faunal transition and may have lasted from
~1 Ma to 0.6 Ma. The earliest-recorded full Florisian fauna is from the Gladysvale
External Deposits, with a minimum (maximum) age of around 0.58 (0.78) Ma
(Lacruz et al., 2002) (Fig. 18.2). Stable isotopes and the reduced taxonomic
diversity in the faunal assemblages from Cornelia suggest that the interior Corne-
lian tended towards semi-arid conditions, whereas the evidence from Elandsfontein
suggests that during the Cornelian the coastal areas may have been wetter than
during the Florisian, in contrast to the situation in the interior (Codron et al., 2008;
Brink et al., 2012; Braun et al., 2013).
18.3.2 Black wildebeest Connochaetes gnou as a proxy for open habitat
The black wildebeest is a southern African endemic with a remarkably complete
local fossil record, which starts in the Cornelian LMA (Brink, 1993, 2005). It does
not occur in east Africa during the time equivalent to the Cornelian LMA, or later
(Gentry, 2010; Faith, 2014), although there are small-bodied wildebeest fossils
from Olduvai, referred to C. africanus, which is considered to be similar to the
black wildebeest (Gentry and Gentry, 1978).
The earliest black wildebeest specimens from Cornelia–Uitzoek are very
close in morphology to contemporary blue wildebeest C. taurinus prognu,
from which it evolved, and which had a pan-African distribution during the
end-early Pleistocene and early-middle Pleistocene (Gentry and Gentry, 1978;
Geraads, 1981; Gilbert, 2008). This evolutionary process was associated ﬁrst
with morphological changes in the skull, dentitions and horns that reﬂect adapta-
tions to a specialised territorial reproductive behaviour; then by a reduction in
body size and changes in body proportions (Brink, 1993, 2005) (Fig. 18.3a, b).
292 James S. Brink
Table 18.3 Taxonomic list of Florisian faunas: a comparison between Florisbad Spring,
Erfkroon Last Interglacial levels (L/I) and the Gladysvale External Deposits. Faunal lists
modiﬁed and adapted from Brink (1987, 1994), Churchill et al. (2000), Lacruz et al.
(2002), and Brink et al. (2015). The extinct taxa also found in east Africa are underlined.
†† = Extinct, †= Regionally extinct.
Homo helmei X- -
Homo sp. - - X
Papionini indet. X - -
Aonyx capensis X- -
Cynictis penicillata -X -
Galerella sanguinea XX -
Atilax paludinosus XX -
Canis mesomelas XX X
Vulpes chama -X -
Lycaon pictus X cf. -
Crocuta crocuta XX -
Panthera leo X cf. X
Equus capensis †† XX X
Equus lylei †† XX -
Equus quagga subsp. X X X
Ceratotherium simum XX -
Hippopotamus amphibius XX -
Phacochoerus africanus X- -
Phacochoerus aethiopicus X- -
Phacochoerus sp. - X X
Girafﬁdae indet. - - X
Taurotragus oryx X- X
Tragelaphus streptsiceros -- X
Syncerus antiquus †† XX X
Kobus leche †XX X
Kobus sp.†XX -
Redunca arundimum -- X
Redunca fulvorufula -- X
Hippotragus sp. †X- X
Damaliscus niro †† XX -
Damaliscus pygargus XX X
Alcelaphus buselaphus XX -
Connochaetes gnou XX -
Megalotragus priscus †† XX -
Caprini indet. †† -- X
Antidorcas bondi †† XX X
Antidorcas marsupialis XX -
Raphicerus campestris X- -
Faunal evidence for Quaternary environmental change 293
Fig. 18.3. (A) The temporal pattern of morphological change in the skulls of the
black wildebeest as it evolved from a blue wildebeest ancestor. (B) The charac-
teristic features of black wildebeest skulls are present in underived states in the
earliest fossils from Cornelia-Uitzoek: 1) forward curving horns, 2) enlarged basal
bosses 3) fused frontals’suture and 4) enlarged orbits. These features reﬂect the
adaptation of black wildebeest to extreme territorial behaviour, which can only
function in open, visually unobstructed habitat (adapted from Brink, 2005).
294 James S. Brink
Black wildebeest are permanently territorial, and males patrol their territories by
vision and defend them aggressively. This is seen in the increased size of the
orbits, evidently linked to the greater need for vision in patrolling territories, and
in the structural adaptations for increased stability of the skull, needed during
head-to-head contact in the defence of territories. In contrast, its living ancestor,
the blue wildebeest has a greater reliance on smell (Attwell, 1977; Brink, 2005).
The morphological adaptations in black wildebeest skulls and dentition, there-
fore, do not reﬂect a trophic shift (Codron and Brink, 2007), but can be directly
associated with adaptation towards greater territoriality in breeding behaviour,
which can only function in open, unobstructed habitat. These properties point to
the Highveld and Karoo regions of central southern Africa as the place of origin
of this species.
The speciation of black wildebeest can be directly dated based on fossil
evidence from Cornelia-Uitzoek, which has a palaeomagnetic age of around
1.0 Ma (Brink et al., 2012). This is supported independently by genetic studies
(Corbet and Robinson, 1991; Corbet et al., 1994). The timing is signiﬁcant and
suggests a connection with increased global cooling and the initiation of the
~100 kyr eccentricity-driven glacial–interglacial cycle soon after 1.0 Ma. It is
also likely that increased incidence of ﬁre, which is related to the development
of grasslands (Hoetzel et al., 2013), may have contributed towards greater
openness in the Cornelian grasslands, and would have counteracted the propa-
gation of woody plants (Bigalke and Willan, 1984). Regardless of causal
factors, the evolution of the black wildebeest is evidently closely linked to the
development of a Highveld-type open habitat. Thus, the presence of black
wildebeest in the fossil record can be used as a proxy for the presence of open
18.3.3 The middle and late Pleistocene: the Florisian LMA
Following the extinction of the Cornelian archaic component, an essentially
modern-looking mammal fauna emerged (Table 18.3), with most of the lineages
continuing into extant populations, excluding six specialised grazing ungulates that
became extinct in southern Africa at the end of the Pleistocene. In addition, an as
yet unnamed grazing caprine became extinct at the end of the late Pleistocene in
the Cango valley and by the mid-Holocene in the northeastern Cape (Brink, 1999).
The Cornelian–Florisian turnover brought about highly productive grasslands in
the central interior with wetlands covering most of the interior of southern Africa
(Brink, 1987; Brink and Lee-Thorp, 1992) (Fig. 18.4). It should be noted that a
sivathere (?Sivatherium maurusium) was reported to occur at Florisbad and an
extinct proboscidean (Elephas recki) probably survived for some time after the
Faunal evidence for Quaternary environmental change 295
Cornelian–Florisian transition (Klein, 1984; Porat et al., 2010). Also, an extinct
grazing pig Metridiochoerus compactus has been recorded from Florisian contexts
at Redcliff Cave. These occurrences are exceptional and not typical of the interior
Florisian, and they are not included in Table 18.3.
Fig. 18.4. A facilitating grazing succession suggested for the Florisian grass-
lands of the central interior of southern Africa, in which extinct and extant
larger-bodied grazers prepared and maintained grasslands in a state of regrowth,
providing niches for smaller-bodied grazers. The feeding niche of Bond’s
springbok Antidorcas bondi was that of the smallest of the specialised grazers
and provided the basis for suggesting the existence of unusually productive
grasslands during the Florisian that needed substantially increased precipitation
compared to modern conditions (see Fig. 18.5) (adapted from Brink and Lee-
296 James S. Brink
18.3.4 Florisian wetlands and the end-Pleistocene extinction
Florisian grasslands existed between 0.6 and 0.01 Ma (Fig. 18.2) and are primarily
indicated by the association of an evolved form of Bond’s springbok Antidorcas
bondi with a range of wetland and aquatic taxa, such as waterbuck (Kobus ellipsi-
prymnus), lechwe (K. leche) and hippo (Hippopotamus amphibius). During the
Florisian LMA, Bond’s springbok reached the climax of its adaptation as a special-
ised small-bodied grazer, and it has been suggested that this niche was facilitated by
larger-bodied grazers, both extinct and extant, and by markedly increased precipita-
tion (Brink and Lee-Thorp, 1992; Brink, 2005; Codron et al., 2008) (Fig. 18.4). The
Florisian wetland and aquatic taxa were closely associated with a palaeolake system
that extended across central southern Africa into eastern Zimbabwe, southern Zambia
and northern Botswana, where today a Florisian remnant wetland fauna survives.
The Florisian palaeolake system is today visible in the pan veld of the central and
western interior, such as around Florisbad (Loock and Grobler, 1988) (Fig. 18.5).
Fig. 18.5. A temporal trend in habitat types in the central interior of southern
Africa (A). Cornelian grasslands containing some woody and closed-habitat
component gave way to open grasslands and wetlands during the Florisian
LMA. The Florisian wetlands terminated at the end of the late Pleistocene due
to widespread aridiﬁcation of the central interior of southern Africa with the local
extinction of the wetland forms and the extinction of the six specialised grazers
(see Fig. 18.4). The Florisian grasslands were linked to an extensive palaeolake
system, which today survives as dry pans, of which an example is given from near
Florisbad (B). The fossil presence of Bond’s springbok and lechwe Kobus leche
reﬂects the existence of palaeolakes and wetlands across the landscape during the
Florisian LMA (adapted from Grobler and Loock, 1988; Brink, 2005).
Faunal evidence for Quaternary environmental change 297
It is also present in the Florisbad spring mound record, where high lake levels are
recorded until immediately before the start of the Holocene (Visser and Joubert,
1990). Based on the occurrence of lechwe and Bond’s springbok, it is clear that the
Florisian wetland system extended over much of the width of southern Africa and
from Cradock in the south to southern Zimbabwe in the north (Fig. 18.6). These
wetlands were anomalous and a temporary departure from the long-term trend
towards aridiﬁcation as seen in the sub-Saharan fossil record (Pickford and Senut,
1999; Bobe, 2006). The cause for the Florisian wetlands is still unclear, but in spite of
palaeoclimatic ﬂuctuations, which would have followed the glacial–interglacial
cycles, conditions remained sufﬁciently stable to allow taxonomic stasis and the
persistence of the wetland indicator taxa in this region over a period exceeding
0.5 Ma (Figs. 18.2, 18.5; Table 18.3). The stability in the Florisian record is also
reﬂected in ungulate phylogeographic data, suggesting a stable, long-standing south-
ern refuge (Lorenzen et al., 2012).
Fig. 18.5. (cont.)
298 James S. Brink
The Florisian ecosystem was disrupted towards the end of the Pleistocene by
intense aridiﬁcation, when primary productivity of the grasslands was dimin-
ished and the palaeo-lakes dried up to be transformed into the modern semi-arid
grassland and pan system (Loock and Grobler, 1988; Brink and Lee-Thorp,
1992). The interior fossil assemblages dating to around the Last Glacial
Maximum record this transformation by a decline in frequency and eventual
disappearance of wetland and aquatic fauna, such as Bond’s springbok, lechwe
and hippopotamus (Klein et al., 1991; Plug and Engela, 1992; Brink, 2005).
The extinction of the Florisian grazing ungulates coincided with the disappear-
ance of wetland elements at the end of the late Pleistocene/early Holocene
(Klein, 1984; Thackeray, 1984; Brink, 2005). Extinction affected only the more
specialised grazers, those with extremely large (Equus capensis,Syncerus
antiquus and Megalotragus priscus), or very small body size (Antidorcas
Fig. 18.6. The extent of late Quaternary fossil localities with Bond’s springbok
(Antidorcas bondi) and lechwe (Kobus leche). This distribution overlaps with
modern open habitat (Grassland and Nama-Karoo Biomes) and the savanna
grasslands (Savanna Biome) and gives a conservative impression of the geo-
graphic extent of the Florisian palaeolakes and wetlands.
Faunal evidence for Quaternary environmental change 299
bondi), or those with close competitors (D. niro and E. lylei) (Brink and Lee-
Thorp, 1992). The pattern of extinction is consistent with aridiﬁcation
and disruption of Florisian ecosystems through a marked reduction in primary
productivity of interior grasslands. Grassland reduction and impoverishment
also coincided with extinction in the Cape ecozone (Klein, 1983; Deacon et al.,
1984; Faith, 2013, 2014). The clear evidence for major regional ecological
shifts at the late Pleistocene–Holocene boundary argues against a human role in
the end-Pleistocene extinctions. Also, the stability in the southern African
archaeological record at this time (vide Deacon and Deacon, 1999) gives
further support to this being a natural process (see also Faith, 2014). By the
end of the Pleistocene and early Holocene, the process of extinction was
complete in both the interior and in the Cape coastal zone.
Aridiﬁcation provided the environmental basis for the evolution of large
mammal faunas that are adapted to arid and semi-arid conditions in southern
Africa. By the end of the early Pleistocene, southern endemism became more
pronounced in Cornelian assemblages through extinction, dispersal and in loco
evolution. Black wildebeest evolved locally from a blue wildebeest ancestor,
and its presence since the Cornelian LMA reﬂects the appearance of open,
Highveld-type grasslands as a stable landscape component. During the Florisian
LMA, an essentially modern-looking fauna emerged, which included six extinct
specialised grazing ungulates and an as yet unnamed caprine species in montane
grassland areas. Florisian wetlands extended beyond modern biome boundaries
into the wooded grassland areas, covering most of southern Africa, but excluded
the coastal zones. Today some of the Florisian wetland forms survive as
biogeographic remnants in the Okavango Delta of northern Botswana and in
southern Zambia. The Florisian faunas were disrupted by extreme aridiﬁcation
towards the end of the late Pleistocene and early Holocene. This resulted in the
extinction of the specialised grazers, the local extinction of the wetland forms
and the emergence by the early Holocene of mainly arid and semi-arid adapted
faunas, as seen historically over most of southern Africa.
Atwell, C. A. M. (1977). Reproduction and population ecology of the blue wildebeest
(Connochaetes taurinus taurinus) in Zululand. Unpublished PhD dissertation, Univer-
sity of Natal, 303pp.
300 James S. Brink
Axelrod, D. I. and Raven, P. H. (1978). Late Cretaceous and Tertiary vegetation history of
Africa. In Biogeography and Ecology of Southern Africa, ed. M. J. A. Werger. The
Hague: W. Junk, pp. 77–130.
Balinsky, B. I. (1962). Patterns of animal distribution on the African continent. Annals
of the Cape Provincial Museums, 2, 299–310.
Behrensmeyer, A. K. (2006). Climate change and human evolution. Science, 311, 476–478.
Bernor, R., Armour-Chelu, M. J., Gilbert, H., Kaiser, T. M. and Schultz, E. (2010).
Equidae. In Cenozoic Mammals of Africa, eds. L. Werdelin and W. J. Saunders.
Berkeley: University of California Press, pp. 685–722.
Bigalke, R. C. (1978). Present-day mammals of Africa. In Evolution of African
Mammals, eds. V. Maglio and H. S. B. Cooke. Cambridge: Harvard University
Press, pp. 1–16.
Bigalke, R. C. and Willan, K. (1984). Effects of Fire Regime on Faunal Composition and
Dynamics. In Ecological Effects of Fire in Southern African Ecosystems, eds. P. de V.
Booysen and N. M. Tainton. Berlin: Springer-Verlag, pp. 255–271.
Bobe, R. (2006). The evolution of arid ecosystems in East Africa. Journal of Arid
Environments, 66, 564–584.
Braun, D. R., Levin, N. E., Stynder, D., Herries, A. I. R., Archer, W., Forrest, F., Roberts,
D. L., Bishop, L. C., Matthews, T., Lehmann, S. B., Pickering, R. and Fitzsimmons,
K. E. (2013). Mid-Pleistocene Hominin occupation at Elandsfontein, Western Cape,
South Africa. Quaternary Science Reviews, 82, 145–166.
Brink, J. S. (1987). The Archaeozoology of Florisbad, Orange Free State. Bloemfontein:
Memoirs van die Nasionale Museum, 24, 151pp.
Brink, J. S. (1993). Postcranial evidence for the evolution of the black wildebeest, Con-
nochaetes gnou: an exploratory study. Palaeontologia Africana, 30, 61–69.
Brink, J. S. (1994). An ass, Equus (Asinus) sp., from late Quaternary mammalian assem-
blages of Florisbad and Vlakkraal, central Southern Africa. South African Journal of
Science, 90, 497–500.
Brink, J. S. (1999). Preliminary report on a caprine from the Cape mountains, South Africa.
Archaeozologia, 10, 11–26.
Brink, J. S. (2005). The Evolution of the Black Wildebeest (Connochaetes gnou) and
Modern Large Mammal Faunas of Central Southern Africa. Unpublished DPhil
dissertation, University of Stellenbosch, 485pp.
Brink, J. S. and Lee-Thorp, J. A. (1992). The feeding niche of an extinct springbok,
Antidorcas bondi (Antilopini, Bovidae), and its palaeoenvironmental meaning. South
African Journal of Science, 88, 227–229.
Brink, J. S., Herries, A. I. R., Moggi-Cecchi, J., Gowlett, J. A. J., Bousman, C. B., Hancox,
J. P., Grün, R., Eisenmann, V., Adams, J. W. and Rossouw, L. (2012). First hominine
remains from a ~1.0 million year old bone bed at Cornelia-Uitzoek, Free State
Province, South Africa. Journal of Human Evolution, 63, 527–535.
Brink, J. S., de Beer, F., Hoffman, J. and Bam, L. (2013). The evolutionary meaning of
Raphicerus-like morphology in the dentitions and postcrania of Antidorcas bondi
(Antilopini). Zitteliana, 31 (Series B), 21.
Brink, J. S., Bousman, C. B. and Grün, R. (2015). A reconstruction of the skull of
Megalotragus priscus (Broom, 1909), based on a ﬁnd from Erfkroon, Modder River,
South Africa, with notes on the chronology and biogeography of the species.
Palaeoecology of Africa, 33, 71–94.
Chazan, M., Ron, H., Matmon, A., Porat, N., Goldberg, P., Yates, R., Avery, M., Sumner,
A., Horwitz, L. K. (2008). Radiometric dating of the Earlier Stone Age sequence in
Faunal evidence for Quaternary environmental change 301
Excavation I at Wonderwerk Cave, South Africa: preliminary results. Journal of
Human Evolution, 55, 1–11.
Codron, D. and Brink, J. S. (2007). Trophic ecology of two savanna grazers, blue
wildebeest Connochaetes taurinus and black wildebeest Connochaetes gnou.Euro-
pean Journal of Wildlife Research, 53, 90–99.
Codron, D., Brink, J. S., Rossouw, L. and Clauss, M. (2008). The evolution of ecological
specialization in southern African ungulates: Competition- or physical environmental
turnover? Oikos, 117, 344–353.
Cooke, H. B. S. (1952). Mammals, ape-men and stone age men in Southern Africa. South
African Archaeological Bulletin,7(26),59–69.
Cooke, H. B. S. (1967). The Pleistocene sequence in South Africa and problems of
correlation. In Background to evolution in Africa, eds. W. W. Bishop and J. D. Clark.
Chicago: University of Chicago Press, pp. 175–184.
Cooke, H. B. S. (1974). The fossil mammals of Cornelia, O.F.S., South Africa. Memoirs
van die Nasionale Museum, Bloemfontein,9,63–84.
Corbet, S. W. and Robinson, T. J. (1991). Genetic divergence in South African wildebeest:
comparative cytogenetics and analysis of mitochondrial DNA. Journal of Heredity,
Corbet, S. W., Grant, W. S. and Robinson, T. J. (1994). Genetic divergence in
South African wildebeest: analysis of allozyme variability. Journal of Heredity,
De Ruiter, D. (2003). Revised faunal lists for Members 1–3 of Swartkrans, South Africa.
Annals of the Transvaal Museum, 40, 29–41.
Deacon, H. J. and Deacon, J. (1999). Human Beginnings in South Africa. Cape Town:
David Philip, 224pp.
Deacon, H. J., Deacon, J., Scholtz, A., Thackeray, J. F., Brink, J. S. and Vogel, J. C.
(1984). Correlation of palaeoenvironmental data from the Late Pleistocene and
Holocene deposits at Boomplaas Cave, southern Cape. In Late Cainozoic Palaeo-
climates of the Southern Hemisphere, ed. J. C. Vogel. Rotterdam: Balkema,
deMenocal, P. B. and Bloemendal, J. (1995). Plio-Pleistocene climatic variability in
subtropical Africa and the paleoenvironment of hominid evolution: A combined
data-model approach. In Paleoclimate and Evolution, with Emphasis on Human
Origins, eds. E. S. Vrba, G. H. Denton, T. C. Partridge and L. H. Burckle. New
Haven: Yale University Press, pp. 262–288.
Dupont, L. M. and Leroy, S. A. G. (1995). Steps towards drier climatic conditions in
northwestern Africa during the upper Pliocene. In Paleoclimate and Evolution, with
Emphasis on Human Origins, eds. E. S. Vrba, G. H. Denton, T. C. Partridge and L. H.
Burckle. New Haven: Yale University Press, pp. 289–298.
Ewer, R. F. and Cooke, H. S. B. (1964). The Pleistocene mammals of southern Africa.
Monographiae Biologicae, 14, 35–38.
Faith, J. T. (2013). Taphonomic and paleoecological change in the large mammal sequence
from Boomplaas Cave, western Cape, South Africa. Journal of Human Evolution, 65,
Faith, J. T. (2014). Late Pleistocene and Holocene mammal extinctions on continental
Africa. Earth-Science Reviews, 128, 105–121.
Gentry, A. W. (1980). Fossil bovidae (Mammalia) from Langebaanweg, South Africa.
Annals of the South African Museum, 79, 213–337.
Gentry, A. W. (2010). Bovidae. In Cenozoic Mammals of Africa, eds. L. Werdelin and W.
J. Saunders. Berkeley: University of California Press, pp. 741–796.
302 James S. Brink
Gentry, A. W. and Gentry, A. (1978). Fossil Bovidae (Mammalia) of Olduvai
Gorge, Tanzania. Part I. Bulletin of the British Museum (Natural History), 29,
Geraads, D. (1981). Bovidae et Girafﬁdae (Artiodactyla, Mammalia) du Pléistocène
de Terniﬁne (Algérie). Bulletin Muséum national d'Histoire naturelle, Paris,4(3),
Gilbert, W. H. (2008). Introduction. In Homo Erectus: Pleistocene Evidence from the
Middle Awash, Ethiopia, eds. W. H. Gilbert and B. Asfaw. Berkeley and Los
Angeles: University of California Press, pp. 1–12.
Grobler, N. J. and Loock, J. C. (1988). Morphological development of the Florisbad
deposit. Palaeoecology of Africa, 19, 163–168.
Grün, R., Brink, J. S., Spooner, N. A., Taylor, L., Stringer, C. B., Franciscus, R. B.
and Murray, A. (1996). Direct dating of the Florisbad hominid. Nature, 382,
Harris, J. M. and White, T. (1979). Evolution of the Plio-Pleistocene African Suidae.
Transactions of the American Philosophical Society, 69, 1–128.
Hendey, Q. B. (1974). Faunal dating of the late Cenozoic of southern Africa, with special
reference to the Carnivora. Quaternary Research, 4, 149–161.
Herries, A. I. R., Reed, K. E., Kuykendall, K. L. and Latham, A. G. (2006). Speleology and
magnetobiostratigraphic chronology of the Buffalo Cave fossil site, Makapansgat,
South Africa. Quaternary Research, 66, 233–245.
Herries, A. I. R., Hopley, P. J., Adams, J. W., Curnoe, D. and Maslin, M. A. (2009).
Geochronology and palaeoenvironments of the South African early hominin bearing
sites –a reply to Wrangham et al., 2009: “Shallow-Water Habitats as Sources of
Fallback Foods for Hominins”.American Journal of Physical Anthropology, 143,
Hoetzel, S., Dupont, L., Schefuß, E., Rommerskirchen, F. and Wefer, G. (2013). The
role of ﬁre in Miocene to Pliocene C
grassland and ecosystem evolution. Nature
Geoscience, 6, 1027–1030.
Kingdon, J. (1997). The Kingdon Field Guide to African Mammals. London: Academic
Klein, R. G. (1983). Paleoenvironmental implications of Quaternary large mammals in the
Fynbos region. In Fynbos Palaeoecology: A Preliminary Synthesis, eds. H. J. Deacon,
Q. B. Hendey and J. J. N. Lambrechts. Pretoria: South African National Scientiﬁc
Programmes Report 75, pp. 116–138.
Klein, R. G. (1984). The large mammals of southern Africa: Late Pliocene to recent. In
Southern African Prehistory and Palaeoenvironments, ed. R. G. Klein. Rotterdam:
A.A. Balkema, pp. 107–146.
Klein, R. G., Cruz-Uribe, K. and Beaumont, P. B. (1991). Environmental, ecological,
and paleoanthropological implications of the late Pleistocene mammalian fauna
from Equus Cave, northern Cape Province, South Africa. Quaternary Research, 36,
Klein, R. G., Avery, G., Cruz-Uribe, K. and Steele, T. E. (2007). The mammalian
fauna associated with an archaic hominin skullcap and later Acheulean artifacts at
Elandsfontein, Western Cape Province, South Africa. Journal of Human Evolution,
Lacruz, R. S., Brink, J. S., Hancox, P. J., Skinner, A. R., Herries, A., Schmid, P. and
Berger, L. R. (2002). Palaeontology and geological context of a Middle Pleistocene
faunal assemblage from the Gladysvale Cave, South Africa. Palaeontologia Africana,
Faunal evidence for Quaternary environmental change 303
Loock, J. C. and Grobler, N. J. (1988). The Regional Geology of Florisbad. Navorsinge
van die Nasionale Museum, Bloemfontein, 5, 489–497.
Lorenzen, E. D., Heller, R. and Siegismuld, H. R. (2012). Comparative phylogeography of
African savannah ungulates. Molecular Ecology, 21, 3656–3670.
McCarthy, T. S. (2013). The Okavango delta and its place in the geomorphological
evolution of southern Africa. South African Journal of Geology, 116, 1–54.
Mucina, L. and Rutherford, M. C. (2006). The vegetation of South Africa, Lesotho and
Swaziland. Strelitzia 19. Pretoria: South African National Biodiversity Institute,
Partridge, T. C. and Maud, R. R. (2000). Macro-scale geomorphic evolution of southern
Africa. In The Cenozoic of Southern Africa, eds. T. C. Partridge and R. R. Maud.
Oxford: Oxford University Press, pp. 3–18.
Pickford, M. and Senut, B. (1999). Geology and Palaeobiology of the Namib Desert.
Windhoek: Memoir of the Geological Survey of Namibia, 18, 155pp.
Plug, I. and Engela, R. (1992). The macrofaunal remains from recent excavations at
Rose Cottage Cave, Orange Free State. South African Archaeological Bulletin, 47,
Porat, N., Chazan, M., Grün, R., Aubert, M., Eisenmann, V. and Horwitz, L. K. (2010).
New radiometric ages for the Fauresmith industry from Kathu Pan, southern Africa:
Implications for the Earlier to Middle Stone Age transition. Journal of Archaeological
Science, 37, 269–283.
Reynolds, S. C. and Kibii, J. M. (2011). Sterkfontein at 75: Review of palaeoenvironments,
fauna, dating and archaeology from the hominin site of Sterkfontein (Gauteng
Province, South Africa). Palaeontologia Africana, 46, 59–88.
Savage, D. E. and Russel, D. E. (1983). Mammalian Paleofaunas of the World. Reading
(Massachusetts): Addison Wesley, 462pp.
Skinner, J. and Smithers, R. H. (1990). The Mammals of the Southern African Subregion,
2nd Ed. Pretoria: University of Pretoria, 768pp.
Thackeray, J. F. (1984). Man, Animals and Extinctions: The Analysis of Holocene Faunal
Remains from Wonderwerk Cave, South Africa. Unpublished PhD dissertation, Yale
Thackeray, J. F. and Brink, J. S. (2004). Damaliscus niro horns from Wonderwerk Cave
and other Pleistocene sites: Morphological and chronological considerations.
Palaeontologia Africana, 40, 89–93.
Tinker, T., de Wit, M. J. and Brown, R. (2008). Mesozoic exhumation of the southern
Cape, South Africa, quantiﬁed using apatite ﬁssion track themochronology. Tectono-
physics, 455, 77–93.
Van Hoepen, E. C. N. (1932). Voorlopige beskrywing van Vrystaatse soogdiere. Paleon-
tologiese navorsinge van die Nasionale Museum, Bloemfontein,2,63–65.
Van Hoepen, E. C. N. (1947). A preliminary description of new Pleistocene mammals of
South Africa. Paleontologiese navorsinge van die Nasionale Museum, Bloemfontein,
Visser, J. J. N. and Joubert, A. (1991). Cyclicity in the late Pleistocene to Holocene spring
and lacustrine deposits at Florisbad, Orange Free State. South African Journal of
Geology, 94, 123–131.
Vrba, E. S. (1995). On the connections between paleoclimate and evolution. In Paleocli-
mate and Evolution, with Emphasis on Human Origins, eds. E. S. Vrba, G. H.
Denton, T. C. Partridge and L. H. Burckle. New Haven: Yale University Press,
304 James S. Brink
Wells, L. H. (1962). Pleistocene faunas and the distribution of mammals in Southern
Africa. Annals of the Cape Provincial Museums,2,37–40.
Werdelin,L.andPeigné,S.(2010).Carnivora.InCenozoic Mammals of Africa,eds.L.
Werdelin and W. J. Saunders. Berkeley: University of California Press,
Faunal evidence for Quaternary environmental change 305