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Faunal evidence for mid- and late Quaternary environmental change in southern Africa

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Southern Africa is differentiated from other centres of aridity in Africa 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 that was 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.
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18
Faunal evidence for mid- and late Quaternary
environmental change in southern Africa
james s. brink
Abstract
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
reected in the large herbivores of the central interior, the grazers and mixed
feeders adapted to permanently available open habitat, which denes 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 aridication 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.
18.1 Introduction
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
284
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 dene 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 aridication and global cooling. Open
grasslands became fully established during the Florisian LMA, which is the
subsequent faunal stage (Tables 18.118.3). 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
dened 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 signicant 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, specically 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 gures: 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.
Cradock Spring.
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 dened 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 rst 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 reects the difculty 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 benetted 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)
(Fig. 18.2).
Table 18.1 Southern African Land Mammal Ages in relation to approximate geological
time periods, adapted from Hendey (1974).
Land Mammal
Age
Approximate geological
age
Recent Holocene
Florisian Middle & late Pleistocene
Cornelian End-early Pleistocene
Makapanian Plio-Pleistocene
Langebaanian Early Pliocene
Namibian Miocene
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 rst noted by zoologists (e.g. Balinsky, 1962) who
proposed a southwestnortheast 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 modied and adapted from Herries et al. (2006), Klein et al.
(2007) and Brink et al. (2012). = Extinct
Cornelia-
Uitzoek
Buffalo
Cave
Elandsfontein
Main
Rodentia
Bathyergus suillus -- X
Hystrix africaeasutralis -- X
Lagomorpha
Lepus capensis -- X
Primates
Homo sp. X - X
Cercopithecidae indet. - X -
Theropithecus oswaldi -- X
Pholidota
Manis sp. - - X
Carnivora
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
Proboscidea
Loxodonta atlantica zulu -X
Indet. X - -
Perissodactyla
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 - -
Artiodactyla
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 reect 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 reect early Pleistocene
endemism in southern Africa are the giant wildebeest Megalotragus eucornutus,
and the impala Aepyceros helmoedi (Van Hoepen, 1932, 1947; Brink, 2005;
Faith, 2014).
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.)
Cornelia-
Uitzoek
Buffalo
Cave
Elandsfontein
Main
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 reects 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 CorneliaUitzoek 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 rst
with morphological changes in the skull, dentitions and horns that reect 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
modied 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.
Florisbad
Spring
Erfkroon
L/I
Gladysvale
External Deposits
Primates
Homo helmei X- -
Homo sp. - - X
Papionini indet. X - -
Carnivora
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
Perissodactyla
Equus capensis †† XX X
Equus lylei †† XX -
Equus quagga subsp. X X X
Ceratotherium simum XX -
Artiodactyla
Hippopotamus amphibius XX -
Phacochoerus africanus X- -
Phacochoerus aethiopicus X- -
Phacochoerus sp. - X X
Girafdae 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 frontalssuture and 4) enlarged orbits. These features reect 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 reect 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 signicant and
suggests a connection with increased global cooling and the initiation of the
~100 kyr eccentricity-driven glacialinterglacial cycle soon after 1.0 Ma. It is
also likely that increased incidence of re, 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
grassland habitats.
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 CornelianFlorisian 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
CornelianFlorisian 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 Bonds
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-
Thorp, 1992).
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 Bonds 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, Bonds 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 aridication 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 Bonds springbok and lechwe Kobus leche
reects 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 Bonds 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 aridication 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 uctuations, which would have followed the glacialinterglacial
cycles, conditions remained sufciently 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
reected 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 aridication, 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 Bonds 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 Bonds 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 aridication
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 PleistoceneHolocene 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.
18.4 Summary
Aridication 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 reects 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 aridication
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.
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... In this context, one particularly understudied region of the South African interior is the Free State Province (henceforth Free State), between the Vaal and Orange Rivers (Fig. 1). Its Grassland Biome, today dissected by sporadic rivers and punctuated by springs and seasonally dry lakes (pans), was characterized by exceptionally productive grasslands and wetlands at various stages during the Middle and Late Pleistocene, supporting large game populations that provided humans with a resource-rich environment (Brink, 2016). The best example of this setting is Florisbad, a spring site at the margin of a pan in the western Free State that has produced the partial cranium of an archaic Homo sapiens dated to 259 ± 35 ka (Bruner and Lombard, 2020;Clarke, 1985;Grün et al., 1996), early MSA and pre-HP lithic assemblages including an intact occupation surface dated to 121 ± 6 ka (Brink and Henderson, 2001;Henderson, 2001;Kuman et al., 1999), and the type faunal assemblage of the Florisian Land Mammal Age (LMA; between~780 and 578 ka, and~10 ka) (Brink, 1987(Brink, , 1988(Brink, , 2016Brink and Lee-Thorp, 1992;Lacruz et al., 2002), currently termed Late Naivashan (Van Couvering and Delson, 2020). ...
... Its Grassland Biome, today dissected by sporadic rivers and punctuated by springs and seasonally dry lakes (pans), was characterized by exceptionally productive grasslands and wetlands at various stages during the Middle and Late Pleistocene, supporting large game populations that provided humans with a resource-rich environment (Brink, 2016). The best example of this setting is Florisbad, a spring site at the margin of a pan in the western Free State that has produced the partial cranium of an archaic Homo sapiens dated to 259 ± 35 ka (Bruner and Lombard, 2020;Clarke, 1985;Grün et al., 1996), early MSA and pre-HP lithic assemblages including an intact occupation surface dated to 121 ± 6 ka (Brink and Henderson, 2001;Henderson, 2001;Kuman et al., 1999), and the type faunal assemblage of the Florisian Land Mammal Age (LMA; between~780 and 578 ka, and~10 ka) (Brink, 1987(Brink, , 1988(Brink, , 2016Brink and Lee-Thorp, 1992;Lacruz et al., 2002), currently termed Late Naivashan (Van Couvering and Delson, 2020). The deep stratigraphic sequence at Florisbad, spanning the last~300,000 years, has allowed paleoenvironmental reconstructions based on sediments (Toffolo et al., 2017), pollen , phytoliths , and stable isotopes of herbivore tooth enamel (Codron et al., 2008). ...
... The small grazer Antidorcas bondi (Bond's springbok), another key extinct species of the Florisian LMA, has been found in Middle and Late Pleistocene deposits at Florisbad, Vlakkraal, Erfkroon, and Rose Cottage Cave (Brink and Lee-Thorp, 1992;. Based on faunal assemblages, these sites are thought to be part of a network of Florisian wetlands associated with paleolakes and rivers that encompassed the South African interior and extended into northern Botswana at various stages during the Middle and Late Pleistocene, in spite of the general trend towards greater aridity observed along the coast during MIS 4 (Brink, 2016). This resource-rich environment thus had the potential to support humans during periods of prolonged climatic stress. ...
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... In our sub-tropical research area, aridity and humidity are often interpreted from proxies rather than temperature, as moisture availability is crucial in semi-arid landscapes. Initially, faunal studies pointed to a wetter than present environment being a key environmental component of the Pleistocene in the elevated central interior of South Africa (see review in Brink 2016). This includes specialised grazing ungulates and wetland faunal species such as lechwe (Kobus leche) and waterbuck (Kobus ellipsiprymnus) which went extinct in the region at the end of the Pleistocene (Brink 2016). ...
... Initially, faunal studies pointed to a wetter than present environment being a key environmental component of the Pleistocene in the elevated central interior of South Africa (see review in Brink 2016). This includes specialised grazing ungulates and wetland faunal species such as lechwe (Kobus leche) and waterbuck (Kobus ellipsiprymnus) which went extinct in the region at the end of the Pleistocene (Brink 2016). Other proxies, such as stable isotopes and pan sediments, have since further confirmed the paradigm of a Pleistocene environment defined by open grassland vegetation within a system of active lakes, springs and rivers. ...
... The purported wet phase at Wonderwerk Cave Excavation 2 and at KP1, KP6 & KP9 broadly overlaps with a period of fluctuating water levels at Florisbad, between 300 and 20 ka (Toffolo et al., 2017;Scott et al., 2019) (Fig. 2). Florisbad is east of Wonderwerk and Kathu Pan in what is today the grassland biome, but was part of the same central interior ecosystem in the Mid-Pleistocene (Brink 2016). The record at Florisbad extends further into the past than that at Kathu Pan (Table 1), but it is worth comparing their overlapping portions since they are based on similar geoproxies (Toffolo et al., 2017). ...
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When considering the pan-African process of human evolution in the Pleistocene it is important to review our understanding of climate on a regional scale, including heterogeneity due to regional variability in the relative strengths of forcing mechanisms. Research in the Kuruman Hills at the southern edge of the Kalahari in South Africa has recently provided new insights into the complexity and distinctiveness of southern African palaeoclimate. Here we compare these records, obtained from Kathu Pan, Wonderwerk Cave, and Mamatwan Mine, focusing on the presence of water bodies from c. 2 million years before present to the end of the Pleistocene. Through the synthesis of multiple proxies we create a picture of localized water availability as it would have affected hominin adaptation. These records are then framed within a larger discussion of regional climate and environmental change. The emerging record from the southern Kalahari suggests that the distribution of zones of rainfall seasonality varied significantly throughout the Pleistocene and a hominin presence is particularly found in association with wetter phases which supported lakes, springs and pans.
... Finally, we do not consid er a differentiation between the quagga and plains zebra (Equus quagga) following Bronner et al. (2003). (Brink, 2016;Codron et al., 2008) possibly of low quality (Marean, 1992). E. capensis was closely related to E. quagga and the largest African equid to be found during the Quaternary weighing in excess of 450 kg. ...
... Megalotragus priscus (giant hartebeest) was a large ~647 kg (calculated from the regression equation in (Janis, 1990, r 2 = 0.905) using the length of the third molar ( M. priscus fed almost entirely on grasses (Brink, 2016;Codron et al., 2008) possibly of low quality (Marean, 1992). M. priscus is well documented from numerous late Pleistocene sites around Southern Africa (Brink, 2016;Brink et al., 2015Brink et al., , 1995Klein, 1983;Rector and Reed, 2010;Rector and Verrelli, 2010). ...
... Megalotragus priscus (giant hartebeest) was a large ~647 kg (calculated from the regression equation in (Janis, 1990, r 2 = 0.905) using the length of the third molar ( M. priscus fed almost entirely on grasses (Brink, 2016;Codron et al., 2008) possibly of low quality (Marean, 1992). M. priscus is well documented from numerous late Pleistocene sites around Southern Africa (Brink, 2016;Brink et al., 2015Brink et al., , 1995Klein, 1983;Rector and Reed, 2010;Rector and Verrelli, 2010). ...
Thesis
Understanding the variations in structure and abundance of animals and what leads to their distribution within the landscape has captured the attention of ecologists for centuries. Importantly, knowledge of current behaviour of large mammals can be used to inform historic population dynamics and is essential to understanding how early humans used large mammals as a foraging resource. Central to this thesis and improving our understanding of large herbivores is the Palaeo-Agulhas Plain (PAP) where large mammalian herbivores formed a key food resource for early humans. The PAP, now submerged off the southern Cape of South Africa, formed a novel ecosystem during lower sea levels. Characterised by large expanses of nutrient rich grasslands and large grazing herbivores, the PAP stands in stark contrast to the nutrient poor fynbos ecosystems that is in the southern Cape today. In this thesis I focus on the Last Glacial Maximum (LGM; ~20 ka) when the PAP was last fully exposed to answer questions relating to the habitat use and range distribution of large herbivores. Importantly, through the Paleoscape Project, modelled climate, soil and vegetation have made these recreations of large mammals possible. Using modelled climate and vegetation this thesis aims to model the large herbivore communities and understand the influence of early humans on the PAP during the LGM for successful integration into the PaleoscapeABM (the PAP agent-based model). To improve our understanding of large mammals on the PAP I identified five large herbivores that became extinct on the PAP since the LGM and modelled their behavioural and physical traits using k-Nearest Neighbour imputation. I predicted the biomass of large herbivores across the PAP using actual biomass of large herbivores from 39 protected areas across South Africa (spanning five functional groups to include the extinct species) across a rainfall gradient and different biomes. The distribution of large herbivores would likely have been driven by similar top-down and bottom-up drivers we see in large herbivore ecology today. Knowing this, I created a predictive model for large mammals by predicting the probability of occurrence of functional groups of large herbivores in relation to environmental drivers and humans. Results showed that all species (except Antidorcas australis) were adapted to the grassy environment of the PAP and these specialisations likely contributed to their extinction along with changing climates and intensified hunting from humans. When predicting herbivore viii biomass, biome was the most important factor influencing the relationship between herbivores and rainfall. In general, large herbivore biomass increased with rainfall across biomes, except for grassland. Finally, I showed the probability of occurrence of large herbivores was influenced by early humans, water availability and a landscape of fear on the PAP. Through this thesis I have successfully provided detailed accounts of the biomass and probability of occurrence of large herbivores on the PAP. Importantly, this information can be seamlessly integrated into the PaleoscapeABM. Finally, I highlight the importance of this knowledge in understanding early humans, the potential shortcomings of this study and resulting areas where research needs to be focused.
... Common names are used when discussing fauna with scientific names in Table 2. Given the scope and number of palaeozoological sites in southern Africa, this is by no means an extensive documentation. The reader is referred to comprehensive regional reviews of southern African sites by others (e.g., Klein, 1980Klein, , 1983Plug and Badenhorst, 2001;Wadley, 2015;Brink, 2016). In the following section, each eco-region -including vegetation and moisture availability -is described and temporal trends based on large herbivore change are noted. ...
... Increased abundance of equids in Namibia also suggests more grasslands there, while the presence of klipspringer is indicative of more moisture in this region -even during the LGM (Thackeray, 1979;Vogelsang et al., 2010). Brink (2016) has argued that increased moisture (indicated by the presence of lechwe, hippo and Bond's springbok in the interior of southern Africa) can be linked to the remnants of Florisian-aged wetlands, dated to ca. 300 ka. These wetlands and associated grasslands correspond to a palaeo-lake system extending across central southern Africa into eastern Zimbabwe, southern Zambia and northern Botswana (Brink, 2016: 297). ...
Article
Analyses of faunal remains are a key means of inferring palaeoenvironmental change. In this paper, the use of faunal remains as a proxy for environmental conditions from Marine Isotope Stage 6 to the Holocene in southern Africa is reviewed. The focus of this review is on large herbivore abundance and how these fluctuate temporally and regionally in accordance with palaeo-climatic shifts. Here, southern Africa is divided into four eco-regions loosely based on climatic, biotic and zoogeographic traits: the Cape Floristic Region, the arid and semi-arid region, the savanna and grassland region, and the wetter eastern region. The relative abundance of large herbivores within these regions are noted, and temporal trends are inferred. On the whole, most eco-regions maintain similar herbivore compositions over time showing the regional ecological resilience of these taxa to local-scale environmental change. Yet some changes in faunal frequencies are apparent. The Cape Floristic Region shows evidence of significant faunal turnover from the Late Pleistocene to the Holocene. Here, grazers are significantly more abundant during glacial periods, probably linked to the terrestrial expansion of the palaeo-Agulhas coastal plain. Shifts in ungulate abundance in the currently xeric central interior, also indicate wetter periods in the Pleistocene. Holocene faunas are generally similar to historic distributions but shifts between xeric and mesic periods are also evident.
... It was the first site in Sub-Saharan Africa where the presence of Homo in the Pleistocene was established (Broom, 1913). The faunal remains were used to establish the Late Pleistocene Florisian Land Mammal Age (Hendey, 1974;Brink, 1987Brink, , 2005Brink, , 2016. Florisbad produced the first radiocarbon dates on a Stone Age site in Southern Africa in 1954 measured by Willard Libby (van Zinderen Bakker, 1957). ...
Article
We have reassessed the palynological record of Equus Cave in the Savanna Biome of the southern Kalahari, one of the longest Late Quaternary pollen records for the semi-arid central interior of South Africa. We combined published pollen results from the cave, derived from hyena coprolites and the rubified deposits in which they occur, into a single sequence. By re-considering the chronology of this sequence, we critically evaluated the palaeoenvironmental record for the site. We compared the pollen evidence from Equus Cave to that from the longer Wonderwerk Cave records (stalagmite, sediments and dung), also located in the Savanna Biome. Then, we contrasted Equus and Wonderwerk records with other previously published pollen sequences derived from a range of sources from several sites in central South Africa. These sites follow a broad northwest to southeast transect of c. 500 km through the Grassland and Nama Karoo Biomes of the Free State and Eastern Cape. Applying Principal Components Analysis to the pollen data, we derived climatic signals at a regional scale to refine reconstructions of Late Quaternary changes for central South Africa.
... Rainfall reaches the south-western Cape from the Atlantic Ocean mainly in June to August, and from the Indian Ocean from the northern and eastern side of southern Africa in December to February. A small by generalist species of reduced diversity (Brink 2016;Faith 2013, Faith andBehrensmeyer 2013). Also, East Africa is characterised by its ancient grasslands and rich faunal grazer diversity of 50-12ka BP, dominated by especially ancestors and extant species of wildebeest, hartebeest, zebra, roan, gazelle, waterbuck, white rhino, buffalo and oribi (Kingdon 1982, Stuart 2015, Faith et al 2020. ...
Article
Full-text available
As described in Part 1 & 2 (Private Game Issue 1 & 2), Earth’s path follows a sequence of natural oscillations effecting global dynamics of constant vegetation and habitat change. Consequently, phyllo-biogeographic existence and evolution (distribution, performance, genetic integrity, survival, and speciation trades) of animal species are affected. Animals move, adapt, or die as a result of changing environmental conditions. The conservation applications of palaeozoological data are numerous (Lyman 2006). One such application concerns the determination whether or not a species is native to a region. Such insight is essential to informing species translocations and to restoring biodiversity in disturbed habitats.
... The Kathu fauna is mainly composed of grazers, including mega-grazers, while Wonderwerk includes a large component of Tragelaphini and Antidorcas sp., whose modern-day representatives are browsers and/or mixed feeders. The differences in species composition may relate to the agents of accumulation, with the Kathu Pan fauna perhaps representing natural mortalities (Klein, 1988), the Cornelia-Uitzhoek and Florisbad fauna representing primarily hyaena den assemblages (Brink 1988(Brink , 2004(Brink , 2016, while the Wonderwerk Cave assemblage attests to a mixture of biogenic agents, although primarily anthropogenic in origin . Consequently, herbivore composition associated with different agents of accumulation, could explain, at least in part, the lower overall δ 13 C values for Wonderwerk compared to the other three assemblages considered here. ...
Article
This study investigates the mid-Pleistocene paleoenvironment and dietary behaviour of ancient herbivores in the South African central interior, today part of the semi-arid Kalahari savanna. Analyses were undertaken of carbon (δ¹³C) and oxygen (δ¹⁸O) stable isotopes in tooth enamel carbonate of twelve fossil herbivore species from Layers 4b and 4a, associated with Earlier Stone Age (ESA) and transitional ESA-Middle Stone Age (Fauresmith) industries respectively, at the archaeological site of Kathu Pan 1. The data are compared with other early to mid-Pleistocene herbivore assemblages located in the central interior, namely Cornelia-Uitzoek, Wonderwerk Cave and the Florisbad Spring. Results indicate that the median δ¹³C values for all ungulate taxa at Kathu were >-4‰, indicating predominantly C4 based diets, although in certain taxa, some individuals included a significant C3 component in their diet. The δ¹⁸O values of most of species at Kathu were relatively low, suggesting a cooler and/or wetter climate. Carbon isotope evidence for C4 dominated habitats at Kathu, but with a larger C3 component amongst grazers than today, resembles the other early to mid-Pleistocene assemblages in the region. Similarly, δ¹⁸O values for Kathu supplement existing evidence that the region was substantially wetter than in modern times.
Article
Full-text available
Sea levels rise and retreat sporadically in response to changes in global climate, as described in Part 1 & 2 of this series (Private Game Issue 1 & 2). Consequently, the size of low-lying coastal land decline or increase accordingly, and with huge effect on the movement, distribution and speciation of large and medium-sized animals of greater than 20kg body mass. In South Africa such areas include the sandy plains of northern KwaZulu-Natal and the presently submerged Agulhas Bank, that stretches from East London in the Eastern Cape to Saldanha/Langebaan on the West Coast. The most southern continental shelf of the African continent. It is better described as the Palaeo-Agulhas Plain. Animal Migration Portals of the Agulhas Plains and movement of associated game species, and of most recent significance is the native zoo-geographics of the endangered Bontebok.
Article
Teeth are usually targeted for dating archaeological sites because they are the least prone to dissolution, in comparison with bones. However, despite this apparent resistance, teeth do undergo diagenesis, which needs to be accounted for in order to obtain accurate ages. In particular, the uptake of trace elements such as uranium in dental tissues needs to be considered for dose rate determination when dated using electron spin resonance (ESR). Characterising the mineralogy and structural integrity of samples prior to dating may thus provide important information related to their state of preservation, especially in the case of teeth whose U content can significantly affect the dose rate. In this study, we dated five teeth of small-sized bovids using combined ESR/U-series dating. They were collected at the Middle Stone Age site of Lovedale, located in the central interior of South Africa. Micromorphology provided sedimentary context to the samples, which were recovered from a layer of gravel rich in faunal remains. Using cathodoluminescence, laser-induced fluorescence, Fourier transform infrared spectroscopy and Raman micro-spectroscopy we assessed the degree of preservation of the enamel. Results reveal that carbonate hydroxyapatite underwent post-depositional alteration, based on its molecular structure and elemental composition. Although the teeth all originate from the same layer and were sampled in the same 1-m square and at a similar elevation, U-content in the enamel differs highly from one tooth to the other, with values ranging from 1.7 to 29.6 ppm. These values are correlated with equivalent doses (De) from 228 to 923 Gy and are consistent with variations in crystallinity determined with vibrational spectroscopy. We also investigated the possible saturation of the ESR signal, by repeating measurements with microwave power values from 1 to 20 mW. Despite such diversity in U-content, the ages calculated assuming an early uptake of U all fall within the same range, from 63 ± 8 ka to 68 ± 15 ka and may only represent a minimum estimate.
Article
The coexistence of multiple species competing for a finite set of resources is a widely debated topic in community ecology. Species with strongly overlapping niches are expected to drive each other towards exclusion, but such species may also coexist if they have similar competitive abilities. This compromise can lead to a peculiar pattern of clumped coexistence, where multiple species share similar niches, leaving gaps open in the theoretically available niche space. Large mammal herbivores may be a good example of this, where species' dietary niches clump as either grazers, browsers, or intermediate-feeders, rather than being continuously distributed over the resource gradient. Here, we develop a model of such emergent neutrality amongst species competing for a set of predefined resources each distributed along a finite niche axis. The model is commensurate with stable isotope niches, thus allowing us to compare its predictions with empirical evidence for changes in community niche structure over evolutionary time. We present stable carbon and oxygen isotope evidence for six discontinuous Late Quaternary assemblages from the central interior of South Africa, demonstrating the emergence of a clumped niche structure from an initial pattern of strongly overlapping diets. We show that species tend to cluster on parts of the niche axis where resource availability is highest, mirroring the proliferation of grazer species as landscapes became more grass-dominated. However, the presence of competition means that species’ niches continue to differentiate, explaining the persistence of browser and intermediate-feeder species even in these open, predominantly treeless landscapes. These results highlight that species interactions are a necessary factor for robust inferences about the evolutionary dynamics of palaeocommunities.
Article
Full-text available
Palaeontological and geological research at the Gladysvale Cave during the last decade has concentrated on de-roofed deposits located outside the Main Chamber. This area has been termed the Gladysvale External Deposit (GVED) and consists of fossil-rich calcified and decalcified sediments. Here we report on the recent analysis of both the faunal material and the geological context of this deposit. The faunal assemblage, excavated from the decalcified sediments contains 29 mammal species including taxa rare or absent in the Witwatersrand Plio-Pleistocene fossil record (e.g. Pelorovis and Kobus leche). Carnivores and porcupines are identified as accumulating agents of the bones. No new hominin findings can be reported from this deposit, and no cultural remains have been recovered. Geologically the calcified and decalcified breccias represent part of a large talus cone that is relatively unexposed. Uniquely for a cave fill in the Witwatersrand hominin-bearing sites, the sediments are horizontally stratified and form a number of flowstone bound sequences. The dating of the in situ cemented sediments is based on electron spin resonance (ESR) and palaeomagnetism. Recent results indicate that the deposits are of Middle-Pleistocene age.
Article
Seventy-five years after Robert Broom's discovery of the first adult Australopithecus in 1936, the Sterkfontein Caves (Gauteng Province, South Africa) remains one of the richest and most informative fossil hominin sites in the world. The deposits record hominin and African mammal evolution from roughly 2.6 million years (Ma) until the Upper Pleistocene. Earlier excavation efforts focused on the Member 4 australopithecine-bearing breccia and the Member 5 stone tool-bearing breccias of Oldowan and Early Acheulean age. Ronald J. Clarke's 1997 programme of understanding the cave deposits as a whole led to the discovery of the near-complete StW 573 Australopithecus skeleton in the Member 2 deposit of the Silberberg Grotto, and the exploration of lesser known deposits such as the Jacovec Cavern, Name Chamber and the Lincoln Cave. Our aim is to produce a cogent synthesis of the environments, palaeodietary information, fauna and stone artefacts as recorded in the Sterkfontein sequence. We begin with an overview of the site and early accounts of the interpretations of the site-formation processes, after which we discuss each Member in turn and summarize the various types of evidence published so far. Finally, we review the most pertinent debates about the site, including the ages of Sterkfontein Member 2 and 4, and the types of habitats represented at the site through time.
Chapter
In view of the apparent largescale occurrence of fire over evolutionary time in much of South Africa, the fauna is, not unexpectedly, often well adapted to it. Such adaptation has apparently taken a number of forms, and ranges from escape mechanisms which are typical of some species, to a range of survival mechanisms practised by others. Such differences in the strategies which permit the different species to survive in a fire-prone environment inevitably give rise to a wide range of short and long-term responses to fire. The populations of some species, and in particular those which escape fire, may be largely unaffected directly by a fire event, but their survival in the long term may be dependent on the habitat which is induced by fire. In contrast, the populations of some species may be reduced considerably by a fire event, but they may possess the capacity to recover rapidly in the post-fire environment. Yet other species occupy only those sites from which fire has been excluded for some time, and such species are therefore confined largely to fire refuges. However, even within the three examples of broad behavioural classes mentioned, large differences in the behaviour of individual species may exist.
Article
This volume, the first in a series devoted to the paleoanthropological resources of the Middle Awash Valley of Ethiopia, studies Homo erectus, a close relative of Homo sapiens. Written by a team of highly regarded scholars, this book provides the first detailed descriptions, photographs, and analysis of the fossil vertebrates-from elephants and hyenas to humans-from the Daka Member of the Bouri Formation of the Afar, a place renowned for an abundant and lengthy record of human ancestors. These fossils contribute to our understanding human evolution, and the associated fauna provide new information about the distribution and variability of Pleistocene mammals in eastern Africa. The contributors are all active researchers who worked on the paleontology and geology of these unique deposits. Here they have combined their disparate efforts into a single volume, making the original research results accessible to both the specialist and the general reader. The volume synthesizes environmental backdrop and anatomical detail to open an unparalleled window on the African Pleistocene and its inhabitants.
Article
Black wildebeest fossils from the interior of South Africa and the Cape coastal zone are compared to modern specimens in order to trace the pattern of morphological change and the distribution of the species through time. Measurements taken on selected postcranial skeletal elements suggest that the evolution of the black wildebeest was marked by a general reduction in body size. It appears that the evolution of Connochaetes gnou from a blue wildebeest-like (C. taurinus) ancestor is best documented in areas to the south of the Vaal River. -from Author