Quaternary Science Reviews 237 (2020) 106301

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Quaternary Science Reviews

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The palaeoecology of Klasies River, South Africa: An analysis of the

large mammal remains from the 1984e1995 excavations of Cave 1 and
1A
Jerome P. Reynard a, *, Sarah Wurz a, b
a
School of Geography, Archaeology and Environmental Studies, University of the Witwatersrand, Private Bag 3, WITS, 2050, South Africa
b
SFF Centre for Early Sapiens Behaviour (SapienCE), University of Bergen, Post Box 7805, 5020, Bergen, Norway

a r t i c l e i n f o a b s t r a c t

Article history: Given the large number of hominin and archaeological remains the site has yielded, Klasies River has
Received 12 December 2019 contributed significantly to our understanding of how humans developed and behaved during the
Received in revised form Middle Stone Age. Its extensive occupational sequence and the abundance of faunal remains recovered
24 March 2020
from the deposits also make it an important site in exploring palaeoenvironmental change during the
Accepted 30 March 2020
Available online 25 April 2020
Late Pleistocene. The mammalian fauna from the over 70 000 year long sequence at Klasies River
possibly extending from MIS 6 to 3 are useful in positioning the evolution of complex human behaviour
within an environmental context. Here, we use the large mammal fauna excavated in the 1980s and
Keywords:
Africa
1990s from Klasies River Cave 1 and 1A to test links between ungulate diversity and palaeoclimatic
Pleistocene change in the south-eastern Cape of South Africa. Fauna from extended Pleistocene sequences in the
Middle stone age south-eastern Cape are relatively rare and collections such as these are important proxies for assessing
Palaeogeography environmental change in this particular region. Our analysis indicates that the proportion of ungulate
Klasies river grazers, browsers and mixed-feeders shifts in accordance with glacial/interglacial fluctuations. We find
Data treatment significant correlations between grazer proportions and ungulate diversity through the sequence which
Data analysis may be linked to the effect of marine regressions on the landscape or shifting moisture availability. We
Ungulate diversity
compare the Klasies River data set with a selection of Middle Stone Age sites in the southern Cape. Our
analysis suggests that primary productivity is greater along the eastern southern Cape than the western
region. This study has broad implications for understanding the relationship between expanding
grasslands and ungulate richness during the Late Pleistocene.
© 2020 Elsevier Ltd. All rights reserved.

1. Introduction our understanding of the development of H. sapiens and con-

Klasies River main is one of the key sites in African Middle Stone
Age (MSA) research. Its significance is based in part on an almost
continuous Pleistocene sequence spanning ~70 000 years and on
the large number of pre-100 thousand year old (ka) early Homo
sapiens remains found at the main site (Rightmire, 1984; Rightmire
and Deacon, 1991; Grine et al., 1998; Grine et al., 2017). Variability
in the lithic technological sequence from Klasies River main site
(hereafter, KRM) has allowed researchers to explore changes in
artefact production during the Late Pleistocene (Wurz, 2002, 2013).
Information from KRM has an important part to play in informing
* Corresponding author.
E-mail address: Jerome.Reynard@wits.ac.za (J.P. Reynard).
textualising the role that the southern Cape of South Africa played
in behavioural expressions of early modern humans (Wurz, 2008).
KRM has also been a significant source of palaeoecological in-
formation (e.g., Klein, 1976; Butzer, 1978; Shackleton, 1982; Avery,
1987; Thackeray, 1988a, 1992, 2007; Nel et al., 2018; Van Pletzen-
Vos et al., 2019). Archaeo-fauna are an important means of infer-
ring palaeoecology and data from large mammal faunal remains are
necessary to understand palaeoenvironmental change and human
subsistence behaviour. The long sequence at KRM also makes this a
potentially useful proxy for environmental change. Previous ana-
lyses of the KRM fauna played a significant role in exploring the
palaeoecology of the southern Cape (e.g., Klein, 1976; Singer and
Wymer, 1982; Avery, 1987; Thackeray, 1988a) but also started an
important debate on early hominin subsistence behaviour (e.g.,
Binford, 1984; Thackeary and Binford, 1986; Thackeray, 1988b;

https://doi.org/10.1016/j.quascirev.2020.106301
0277-3791/© 2020 Elsevier Ltd. All rights reserved.
2 J.P. Reynard, S. Wurz / Quaternary Science Reviews 237 (2020) 106301

Klein, 1989; Milo, 1998; Bartrams and Marean, 1999; Klein et al.,
1999). The first major excavation at KRM was by Singer and
Wymer (1982, p. 7) from December 1966 to July 1968. Wymer
recovered a large collection of faunal remains but the sampling
method had been problematic because they used mesh sizes of
between 1 and ½ inch that did not retain the smaller sized remains.
Faunal remains were initially sorted in the field and bones “not
regarded as taxonomically identifiable were discarded” (Klein,
1976, p. 76). Much of the subsequent discussions about the use-
fulness of faunal data obtained from the Singer and Wymer exca-
vations, centred on data obtained from this sampling method. For
example, bovid skeletal-part profiles at KRM from the Singer and
Wymer material consist of a prevalence of skull remains and distal
elements for large bovids, with an under-representation of upper
long bones. Elements from smaller bovids were generally more
equitably distributed. This pattern e called the ‘Klasies pattern’ by
Bartrams and Marean (1999) e was the root of much of the debate
around the KRM fauna (e.g., Klein et al., 1999). While researchers
have generally attributed this pattern to a combination of excavator
and taphonomic bias (Turner, 1989; Marean et al., 1992, 2004;
Marean and Frey, 1997; Bartrams and Marean, 1999; Pickering et al.,
2003), others have argued that carnivore scavenging (Binford,
1984) or human transport decisions (Klein, 1989) may be the
cause. Van Pletzen-Vos et al. (2019) found bovid skeletal-profiles
significantly more equitable in the Deacon compared to the
Singer and Wymer assemblage suggesting that Singer and Wymer’s
sampling method played an important role in the Klasies pattern.
The southern Cape has yielded a number of archaeological sites
key to conversations of modern human evolution (Henshilwood,
2012; Wadley, 2015). The Greater Cape Floristic Region (GCFR) of
the southern and south-western Cape is dominated by fynbos
vegetation e an evergreen, sclerophyllous shrub that thrives in
winter rainfall. The temperate climate and the accessibility of
abundant food resources in this region such as geophytes (Deacon,
1993; Proches et al., 2005; Larbey et al., 2019), shellfish (Jerardino
and Marean, 2010; Langejans et al., 2012, 2017; Kyriacou et al.,
2014), and fauna (Klein, 1980; Skinner and Chimimba, 2005;
Thompson and Henshilwood, 2014) have led some researchers to
suggests that the environmental of the southern and southern-
western Cape has had a significant impact of human behavioural
development in southern Africa during the Late Pleistocene
(Marean, 2010; Parkington, 2010; Compton, 2011).
Unlike most other MSA sites in the southern Cape, e.g. Die
Kelders, Blombos Cave, Klipdrift Shelter, and Pinnacle Point, KRM is
situated to the east of the GCFR (Fig. 1). MSA sites on the whole, and
archaeo-faunal assemblages in particular, are rare from the eastern
part of this region e especially along the coast. The environment in
the eastern region of the GCFR is not dominated by fynbos

Fig. 1. Klasies River main (KRM) and other Middle Stone Age sites in southern Africa. Shaded area corresponds to the Greater Cape Floristic Region. DRS ¼ Diepkloof Rockshelter,
DK ¼ Die Kelders, KDS ¼ Klipdrift Shelter, BBC ¼ Blombos Cave, PP ¼ Pinnacle Point, SI ¼ Sibudu.
 J.P. Reynard, S. Wurz / Quaternary Science Reviews 237 (2020) 106301
vegetation (Van Wijk et al., 2017), but instead occurs as part of a
mosaic of forest, thicket and coastal vegetation. Because of the long
sequence at KRM, the faunal remains are a valuable means of
assessing palaeoenvironmental change in the region. Given this, we
may be able to use these remains to examine regional differences
between the large mammal faunal signatures at KRM and other
southern MSA sites further to the west. By doing this we could
explore how far back into the past any environmental differences
between these areas extend.
While the Singer and Wymer (hereafter, SW) faunal data have
shown significant changes in ungulate habitat through the
sequence (Klein, 1976), smaller mammals such as size 1 bovids may
be underrepresented in this collection, affecting its worth as a
palaeoenvironmental proxy (Van Pletzen, 2000). The Deacon ex-
cavations from 1984 to 1995 recovered a large amount of faunal
material using smaller mesh sizes of 2 mm and, unlike the SW
excavations, all remains were collected and analysed. Although the
results of the faunal analyses (Van Pletzen, 2000) have recently
been summarized (Van Pletzen-Vos et al., 2019), it has not been
examined for its palaeoenvironmental implications. Given the
significance of the site, and the relevance fauna material from the
Deacon excavations may have to our understanding of palae-
oenvironmental conditions in the Late Pleistocene, it is crucial that
these data be interrogated further to explore the palaeoecology of
the MSA in the southern Cape.
Ungulate diversity can inform on habitat productivity and
environmental change (Thackeray, 1980; Klein, 1983). Ungulate
richness may be linked to primary environmental productivity and
precipitation (Coe et al., 1976; Thackeray, 1980; Radloff, 2008; Faith,
2013b). There is generally higher ungulate diversity with increased
precipitation in low to medium-rainfall biomes e with precipita-
tion up to ~750 mm/year e declining thereafter (Olff et al., 2002).
Faith (2011a) argues that decreasing ungulate richness in the
southern Cape is related to a decline in productivity and a loss of
available grasslands. This would likely be due to altered rainfall
regimes and shifting sea levels resulting from glacial/interglacial
climatic changes. Yet the effects of glacial/interglacial changes on
ungulate diversity in the south-eastern Cape is not fully under-
stood. In this paper, we assess the large mammal faunal remains
from the Deacon (hereafter, D) sample and discuss its role in the
palaeoecology of the southern and south-eastern Cape. Based on
hypotheses that link increasing ungulate richness to expanded
grasslands in the southern Cape (Klein, 1980; Faith, 2011a, 2013a;
Marean et al., 2014), we evaluate links between grazer abundance
and climatic change in the south-eastern Cape. Specifically, we
examine whether ungulate richness is associated with moisture
and grassland availability in this region during the late Pleistocene.
A comprehensive taphonomic analysis is in progress and without
this component few secure inferences can be made about subsis-
tence behaviour. Yet the stand-alone taxonomic data are a useful
resource for examining palaeoecological change through time in
this important region in modern human development.
2. Site background
KRM is situated 500 m from the Klasies River and consists of
Caves 1 and 1B (6 m above sea level [asl]), Cave 2 (at 18 m asl), and
Cave 1A, an overhang. The caves were cut into the Cape Supergroup
quartz arenite rocks more than 1 million years ago (Pickering et al.,
2013) into a steep cliff that marks the edge of the coastal platform
on the Tsitsikamma coastline. As Caves 1B and 1 became choked
with depositional debris, deposits accumulated against the Cave 1A
cliff face for the most extended period in the occupational history of
the site. Although mid-Holocene higher sea levels eroded much of
the deposits from KRM, more than 21 m of superimposed shell
3
middens remain. Wymer first excavated main site in 1967/8
through several cuttings, from the basal Layer 40 to the uppermost
Layer 1 in Cave 1A (Figure 2.1 in Singer and Wymer, 1982, Fig. 2, this
paper). Singer and Wymer recognised a number of lithic ‘industries’
or techno-complexes, including MSA l at the base, followed by MSA
ll, the Howiesons Poort (HP) and MSA lll. Deacon’s excavations, the
subject of this paper, followed in the 1980s and 1990s (Deacon and
Geleijnse, 1988; Deacon, 2004). This research sampled a column
through the sequence exposed by Wymer in a number of squares,
from PP 38 in Cave 1B to E50 in Cave 1A (Fig. 3). Fine meshed
screens were used to recover the smaller items and the focus was
on the contextual detail of the finds. Wurz is currently excavating
main site, following the protocol and broad stratigraphic divisions
of Deacon (Wurz et al., 2018). Deacon reconfigured the stratigraphy
in a number of members. From the base to the top, these are the
Light Brown Sand (LBS), Rubble Brown Sand (RBS), Shell and Sand
(SAS), Rock Fall (RF) and Upper members. The extent and charac-
teristics of the RBS member are currently under investigation.
Deacon recognised many more discrete units or layers than Singer
and Wymer and emphasised that the KRM deposits represent
multiple short-term human occupations separated by non-
occupation deposits, usually composed of sands.
This paper discusses the large mammal fauna excavated from
the Deacon excavations in Caves 1 and 1A in relation to the MSA l,
MSA ll Lower and Upper, HP and MSA lll techno-complexes
(Table 1). Below, the technological and stratigraphic context of
the Deacon faunal samples from Caves 1 and 1A is discussed. The
layers are grouped according to the scheme followed by Wurz
(2000, Figure 37, Table 11). Only a few ages for the KRM se-
quences are mentioned here, but recent summaries can be found in
Grine et al. (2017) and Wadley (2015).
The MSA 1 lithic techno-complex, also known as the Klasies
River techno-complex occurs in the LBS and the RBS members (SW
Layers 40-37) (Wurz, 2002). In the MSA l, a quartzite blade pro-
duction system was followed, in which elongated points and blades
from volumetric and two volume cores were produced (Wurz,
2002). There is evidence for more intensive reduction of cores in
the MSA l (Wurz, 2010). This techno-complex is, for example,
distinguished by intensive platform preparation in the form of step
flaking and rubbing, and a statistically significant difference in
platform size to length, compared to the other phases at KRM
(Wurz et al., 2003). The MSA l fauna analysed here are from Cave 1/
1A, Squares Z44-AA43, Layers SCB3S- SBS (Table 1; Fig. 3, see also
Wurz, 2000 Table 11). Deacon excavated these squares from Cave 1/
1A, the area where SW Main cutting square ‘a’ and East cutting
Square ‘P’ intersects (Fig. 3). Several ages are associated with the
half a metre thick LBS member, for example, 106.8 ± 12.6 ka
(thermoluminescence [TL] and optically stimulated luminescence
[OSL]),Feathers (2002) and 108.6 ± 3.4 ka (Uranium series [U-
series]), Vogel (2001). Deacon and Geleijnse (1988, p. 8) remarked
that the stalagmite formations in Cave 1 indicate that the LBS
member “spanned some thousands and even several tens of
thousands of years”. This hypothesis is currently being investigated
through U-series dating of the stalagmites in the cave.
The MSA ll techno-complex, also referred to as the Mossel Bay
phase at KRM, occurs in the SAS member. It contains elongated
quartzite products produced from Levallois-like cores (see also
Brenner and Wurz, 2019). The intensification evident during the
MSA l is not present here, and most of the products, particularly the
points, have large faceted platforms and prominent bulbs. There is
technological change through time in the MSA ll and therefore the
MSA ll Lower (SW Layer 16e17) and Upper (SW Layers 15-14) have
been distinguished. The MSA ll Upper products, especially the
points, are shorter and more standardized in terms length and
width than those from the MSA ll Lower (Thackeray and Kelly, 1988;
4 J.P. Reynard, S. Wurz / Quaternary Science Reviews 237 (2020) 106301
Fig. 2. Relative stratigraphy of Cave 1, 1A and 1B at Klasies River Main. See Table 1 for details of stratigraphic members (after Wurz et al., 2018).
Wurz, 2002).
The MSA ll Lower fauna described in this paper are from three
areas at KRM. In Cave 1 it is from the Witness Baulk, SAS Upper
(SASU) sub-member, layers HHH to D1. The Deacon excavation of
the Witness Baulk is adjacent to SW Main Cutting ‘south’ and West
cutting Square ‘H’. The faunal sample further originates from Cave
1/1A Squares AA43, Z44, Y45 and Y44, layers CL2-SCB2AS (Fig. 3;
Table 1). The fauna from the MSA ll Lower in Cave 1A are only from
a few layers from square T50, Layers SM5T to SM5LB. The MSA ll
Upper fauna are from the Witness Baulk e the SAS Wedge (SASW)
sub-member e from layers A1-H3 and in Cave 1A from squares L51
to T50, Layers YS- BS4L. Deacon excavated squares L51 to T50 off
SW’s Initial Cutting (see Fig. 3). Various ages, most from Cave 1, are
associated with the SAS member. The MSA ll Lower, for example is
linked to an age of 101 ± 12 ka (U-series and electron spin reso-
nance, Eggins et al., 2005), and the MSA ll Upper to U-Th ages of
between 100.8 ± 7.5 ka and 85.2 ± 2.1 ka (Vogel, 2001). In Cave 1A
the MSA ll Upper is associated with a U-Th date of 77.4 ± 7.0 ka
(Vogel, 2001).
In Cave 1A there is a substantial hiatus between the MSA ll
Upper layers and the HP, as evidenced in the Rock Fall (RF) member
(SW’s layer 22). The HP techno-complex occurs above this, in the
lower 1.8 m of Upper Member. It is a blade technology, containing a
higher proportion of smaller blades than in the rest of the MSA
sequence at KRM (Wurz, 2002; Villa et al., 2010), associated with
backed geometrics and notched artefacts (Wurz, 2002). The plat-
form preparation techniques evident in the MSA l appear again, but
this time in relation to even more intensive reduction of cores and
utilisation of a wider range of raw materials (Wurz, 2010). Non-
quartzite material is especially evident in the middle layers of the
HP at KRM where silcrete and quartz increase (Wurz, 2000). The HP
at KRM is associated with ages of for example 63.2 ± 2.7 ka (single
grain OSL, Jacobs and Roberts, 2017), 65.6 ± 5.3 ka (U-series Vogel,
2001) and a mean TL age of 53 ± 3 ka (Tribolo et al., 2013). The HP
fauna comes from three squares excavated off Wymer’s initial
cutting, squares J51, H51 and the lower section of E50, Layers YSx6-
CP5.
The MSA lll is quite an ephemeral techno-complex at KRM that
is perhaps best known for the absence of HP technological elements
and a return to the more typical MSA products of the MSA at KRM.
 J.P. Reynard, S. Wurz / Quaternary Science Reviews 237 (2020) 106301 5

Fig. 3. Site map of Klasies River main site.

Table 1
Stratigraphic relationship between units, layers and techno-complexes in Cave 1 & 1A at Klasies River.

Sub-stage Cave 1A Cave 1/1A Cave 1

Member Square and Layers Member Square and Layers Member Square and Layers

MSA III Upper E50S-E50YS3

Howiesons Poort Upper E50CP5–J51YSX6
MSA ll upper SAS L51 YS-T50 BS4L SASW A1-H3
MSA ll lower SAS T50 SM5T - SM5LB SAS Y44 SM51SHB - Y45 CL2 - AA43 SCB2AS SASU D1-HHH
MSA l LBS Z44 SCB3S - AA43 SBS

Elongated larger quartzite artefacts again dominate (Wurz, 2002; (Tribolo et al., 2013).
Villa et al., 2010). The MSA lll faunal sample discussed in this paper
comes from square E50, Layers E50S-AB, from only half a metre of
3. Regional environment
deposits. Ages for this section of the stratigraphy include for
example 43.4 ± 3.0 ka (OSL; Feathers, 2002) and 57.0 ± 4.0 ka
KRM occurs within the Eastern Fynbos Biome at the eastern
6 J.P. Reynard, S. Wurz / Quaternary Science Reviews 237 (2020) 106301
edge of the GCFR along the present-day south-eastern coast of the
Eastern Cape of South Africa. Vegetation in the present-day GCFR
consists mainly of fynbos (shrubs) while renosterveld and strandveld
(shrubs and grasses) are also present (Bergh et al., 2014). The site is
currently surrounded by Algoa Dune Strandveld and Southern Cape
Dune Fynbos interspersed with Cape Seashore vegetation. The
Klasies River itself is within a heavily thicketed valley dominated by
Southern Afrotemperate Forest (Mucina and Rutherford, 2011). The
result of these various interdigitated biomes is a mosaic of
ecological regions surrounding the site typical of the eastern GCFR
(Van Wijk et al., 2017). Climatic and topographical diversity across
the GCFR results in a gradual change in environmental conditions
from the more westerly, winter rainfall zone near Cape Town, to the
eastern year-round rainfall zone (YRZ) near Algoa Bay, and from the
drier, northern mountainous regions to the moister, southern
coastal strip (Campbell, 1983; Chase and Meadows, 2007; Ellis et al.,
2014). KRM lies within the YRZ and is influenced by relatively
moderate oceanic and atmospheric circulatory systems (Carr et al.,
2007; Chase and Meadows, 2007). The warm Agulhas Current from
the south-east results in relatively mild climate and temperate sea
surface temperatures (SST) throughout the year. The current mean
annual precipitation (MAP) for the Southern Cape Dune Fynbos
ecotone between the Tsistikamma region and Oyster Bay is
~757 mm with a slight peak in autumn and spring (Mucina and
Rutherford, 2011) and MAP for the KRM region is estimated to be
~1000 mm/year (Van Wijk et al., 2017). Sea level fluctuations be-
tween glacial and interglacial periods would have resulted in sig-
nificant landscape changes in some areas of the southern Cape, due
to the broad, shallow Agulhas Bank continental shelf off the
present-day coast of the southern Cape. This expanded Palaeo-
Agulhas Plain, however, would have been longitudinally much
broader further west off the southern Cape coast than off the coast
of KRM (Compton, 2011; Rautenbach et al., 2019).
The GCFR host a diverse range of both small and large endemic
fauna (Skinner and Chimimba, 2005). Historic accounts suggest
that large herbivores were common in the GCFR but consist mostly
of smaller bovids such as Cape grysbok, steenbok and grey rhebuck
(Skead, 1980; Faith, 2011b) (scientific names in Table 1). Larger
ungulates, such as zebra, black rhino, hippo, African elephant, and
wildebeest, although present in the Holocene, may have been more
common in the Pleistocene (Klein, 1980; Faith, 2011a; Reynard
et al., 2016a). In the low-lying areas east of the Overberg, Cape
buffalo, bontebok and bushbuck were present in the forest and
thicket patches (Skead, 1980; Mucina and Rutherford, 2011).
4. Materials and methods
This faunal sample is from the assemblage excavated by Hilary
Deacon and colleagues between 1984 and 1995. All material was
retained after screening through 2 and 3 mm mesh. Identification
of the bones was undertaken by James Brink and Van Pletzen-Vos
for Cave 1A and Van Pletzen-Vos for the Witness Baulk in Cave 1
(Van Pletzen-Vos et al., 2019). The assemblage was analysed
following Klein and Cruz-Uribe (1984). Mammals of a similar size or
larger than the Cape dune molerat were examined in this analysis.
Fauna was identified to taxa using the South African National
Museums comparative faunal collections in Bloemfontein. Our
assessment uses the number of identified specimens (NISP) as the
preferred analytical quantitative unit because NISP is a primary
quantitative unit (cf. Lyman, 2008). Moreover, the minimum
numbers of individuals (MNI) are generally small and are probably
underrepresented due to extensive fragmentation of the assem-
blage (cf. Plug and Plug, 1990; Marshall and Pilgram, 1993). Faunal
material was assigned to Class, Order or Family using teeth. The
cranial and post-cranial remains of bovids that could not be
consigned to Linnaean Family were assigned size classes based on
Klein (1976) where bovids under ~ 20 kg, such as Cape grysbok,
klipspringer are categorised as small bovids. Small-medium bovids
are those between 84 and 18 kg such as grey rhebuck and impala.
Large-medium bovids, are those between 300 and 77 kg (e.g., red
hartebeest, kudu) and bovids heavier than 360 kg (eland and buf-
falo) are classed as large bovids (Table 1). Very large bovids, those
heavier than 950 kg, such as the extinct long-horned buffalo and
giant hartebeest are combined with larger bovids in our study.
Animal habitats are based on modern data from Skinner and
Chimimba (2005). We investigate variations in the faunal assem-
blage by examining presence/absence data and taxonomic diversity
throughout the sequence which can be a useful indicator of
palaeoenvironmental conditions (cf. Rector and Verrelli, 2010;
Faith, 2013a). To infer the dominant vegetation of the KRM region,
ungulate taxa are categorised according to their dietary prefer-
ences, viz., grazers, browsers and mixed-feeders. To explore
palaeoenvironmental conditions, a range of faunal diversity indices
are calculated using the PAST software package (Hammer et al.,
2001). Taxonomic richness or the number of taxa (NTAXA;
measured at the genus level), indicates niche breadth and can be
used to infer the diet breadth of a faunal assemblage. Evenness
measures how equally taxa are distributed throughout the assem-
blage and heterogeneity is a simultaneous measurement of both
richness and evenness (Lyman, 2008). We use the Shannon-Wiener
heterogeneity index for heterogeneity, calculated as H ¼ -S Pi (ln
Pi), where P is the proportion of taxon i in the assemblage. High
values signify increased heterogeneity. Evenness is calculated using
the Shannon index as e ¼ H/ln S, where H is the Shannon-Weiner
heterogeneity index and S is NTAXA. For this index, values fall be-
tween 0 and 1, with 1 representing a perfectly even assemblage.
Simpson’s index of diversity (1-D) is used to measure the degree of
dominance (D) of a single taxon and is calculated as 1 e (S ni [ni/
n]2) where ni is the number of specimens from taxon i (Magurran,
1988). The values for Simpson’s index range from 0 to 1 where a
lower value indicates dominance by a single taxon, and a higher
value represents increased diversity. Fisher’s alpha (a) is a simple
measure of diversity relatively insensitive to sample size discrep-
ancies (Magurran, 2004; Faith, 2013a) and is calculated as
NTAXA ¼ a þ ln(1 þ N/a), where N equals assemblage sample size.
Higher a values signify increased diversity. Because larger faunal
assemblages are likely to produce higher NTAXA values, and the
relationship between sample size and NTAXA is logarithmic (Wolff,
1975), a richness index (NTAXA/log NISP) is calculated by dividing
NTAXA by the log of the number of specimens in the taxa under
consideration (Lyman, 2008). We also use an ungulate richness
index to measured ungulate diversity in this study. This richness
index also uses uNTAXA divided by the sum of log-transformed
ungulate sample sizes (uNTAXA/log uNISP) (Lyman, 2008). Ungu-
late residual analyses are also conducted but only when uNTAXA
correlates to log transform SuNISP. Residuals above or below
regression lines are measured as an additional compensation for
sample size differences.
5. Results
The total weight of the faunal remains is 182.7 kg with over
78 kg identified to at least the class level. Although no taphonomic
assessment was undertaken, a study of the taphonomy of the
assemblage is forthcoming. Preliminary taphonomic indicators
from the Howiesons Poort (HP), (Achieng et al., 2019), MSA lll
(Pearson et al., 2019) and MSA ll phases (Lap et al., 2019) suggest
that both humans and animals played a significant role in bone
collection. Small fauna from the MSA lll layers were likely prefer-
entially accumulated by non-humans (Van Pletzen-Vos et al., 2019)
 J.P. Reynard, S. Wurz / Quaternary Science Reviews 237 (2020) 106301 7

while some larger bovids in the MSA ll may have been accumulated
through scavenging (Lap et al. in prep.).
The faunal collection displays a diverse range of taxa with rock
hyrax (n ¼ 595) and seal (n ¼ 460) the most abundant identified
species (Table 2). Bovids are the most common class of animals
constituting 65% (n ¼ 3473) of identified taxa. Eland (n ¼ 80) and
Raphicerus sp. (Cape grysbok and/or steenbok; n ¼ 101) dominate
the identified bovid taxa with African buffalo (n ¼ 43) also com-
mon. Although terrestrial carnivores occur (n ¼ 54), identified
carnivore remains are generally rare. Most of the identified remains
were recovered from MSA ll Lower while MSA l yielded the least
number of specimens.
5.1. Environmental indicators
In general the abundance of eland and the presence of both
browsers and grazers indicate an environment of grasslands
interspersed with thicket and other types of closed environments.
Extinct taxa make up a significant component of the identified
specimens and modern genetic and morphological analogies are
used to infer their dietary prevalence (e.g., Klein, 1974; Gentry,
2010; Faith and Thompson, 2013). Extinct blue antelope were
most likely grazers based on their teeth structure and their close
relationship with roan and sable (Faith and Thompson, 2013).
Extinct long-horned buffalo were also probably gregarious grazers
and their size suggests that they occurred in large open landscapes.
On the other hand, Raphicerus are ubiquitous to KRM and most MSA
sites in the GCFR and their presence suggests fynbos and other
closed environments (Faith, 2011b). Blue and common duikers also
occur in closed, bushy environments. Hippopotamus and otter
indicate wetlands or riparian environments while the Cape dune
mole rat, common in sandy coastal sites, is relatively rare in this
sample.
Temporal trends in ungulate communities suggest changing
environmental conditions through the sequence. While the envi-
ronment is generally mosaic, there is a relative shift from more
grazers in MSA l to a prevalence of mixed-feeders and browsers in
MSA ll and back to some dominance of grass-loving species in the
HP and MSA lll layers (Fig. 4). However, some shifts occur within
the HP and between the HP and MSA lll. There is a significant dif-
ference between smaller (small and small-medium) and larger
(large-medium and large) bovids between the HP and MSA lll
(Fig. 5, c2 ¼ 5.603; df ¼ 1; p ¼ 0.02). In general, smaller bovids such
as Raphicerus are adapted to more closed bushy habitats while

Table 2
Taxa based on the number of identified specimens (NISP) from Klasies River Cave 1 & 1A.

Techno-complex MSA III HP MSA II U MSA II L MSA I

Cave 1A 1A 1 & 1A 1/1A AA43/Z44 1/1A AA43/Z44 Total

Order Taxa Common name

Primate Papio ursinus Baboon 2 2
Primate: indet. Primate 1 1
Rodentia Bathyergus siullus Cape dune mole rat 2 1 1 5 9
Hystrix africaeaustralis Porcupine 2 2
Lagomorpha Lepus sp. (? Capensis) Hare 1 3 4
Leporidae Hare 2 2
Hyracoidea Procavia capensis Hyrax 121 18 87 340 29 595
Carnivora Aonyx capensis African clawless otter 1 2 3
Hyaena brunnea Brown hyena 4 4
Cetacea sp. Whale/dolphin sp. 1 1 2
Arctocephalus sp. Seal 21 62 110 254 13 460
Carnivore: indet. Carnivores 17 3 18 14 2 54
Tubulidentata Orycteropus afer Aardvark 4 6 22 32
Ruminantia Equus sp. Zebra 1 1 1 5 2 10
Suid sp. Bushpig/Warthog 2 2 1 1 6
Hippopotamus amphibus Hippopotamus 8 5 13
Taurotragux oryx Eland 2 2 14 61 1 80
Tragelaphine cf. 2 2
Tragelaphus strepsiceros Greater kudu 1 2 1 18 1 23
Tragelaphus scriptus Bushbuck 2 21 23
Syncerus antiquus Long-horned buffalo 9 3 6 2 20
Syncerus caffer African buffalo 6 2 8 25 2 43
Sylvicapra grimmia Common duiker 6 1 7
Kobus sp. Waterbuck/lechwe 1 1
Redunca cf. Reedbuck 6 6
Redunca arundinum Southern reedbuck 1 2 3
Hippotragus sp. Roan/sable/bluebuck 1 2 4 7
Hippotragus leucophaeus Bluebuck 2 1 19 1 23
Damaliscus pygragus Bontebok/blesbok 2 1 3
Alcelaphus/Connochaetes Hartebeest/wildebeest 1 5 6 11 3 26
Megalotragus cf. Priscus Giant hartebeest 1 1
Philantomba monticola Blue duiker 1 1
Raphicerus sp. Cape grysbok/steenbok 10 6 22 62 1 101
Pelea capreolus Grey rhebok 2 1 4 3 10
Antidorcas sp. Springbok 1 1 2
Bovidae: Indet. Bovids 52 156 9 217
Large 47 61 77 248 34 467
Large-medium 97 81 91 366 73 708
Small-medium 56 66 131 284 48 585
Small 187 100 350 422 26 1085
Mammal: indet. 189 469 24 682
Total 595 418 1178 2852 282 5325

HP ¼ Howiesons Poort; MSA ll U ¼ Upper and L ¼ Lower.

8 J.P. Reynard, S. Wurz / Quaternary Science Reviews 237 (2020) 106301

the earlier phase to a more mosaic habitat in later period (Fig. 5).
The MSA ll Upper period has significantly more smaller bovids than
the next highest period, MSA lll (c2 ¼ 14.770; df ¼ 1; p < 0.001).

5.2. Taxonomic diversity

Table 3 shows a range of diversity indices calculated for the KRM

Fig. 4. Ungulate dietary preference at Klasies River. Number of identified specimens
(NISP) in columns.
sample. There is a clear relationship between NTAXA and log-
transformed SNISP (rs ¼ 0.886, p ¼ 0.019). NTAXA is significantly
correlated to SNISP (rs ¼ 0.949; p ¼ 0.014) but not to the richness
index (rs ¼ 0.791; p ¼ 0.1107). This suggests that, although taxo-
nomic abundance is linked to sample size, other factors may also be
at play. Correlation tests for heterogeneity (H) versus NTAXA
(rs ¼ 0.143; p ¼ 0.787) and evenness (e) (rs ¼ 0.315; p ¼ 0.544)
indicate no clear relationship between those indices. Simpson’s
index is correlated with heterogeneity (rs ¼ 0.899; p ¼ 0.015) but
there is no significant association between Simpson’s index and
either evenness (rs ¼ 0.667, p ¼ 0.148) or the richness index
(rs ¼ 0.145; p ¼ 0.784). Richness is highest in MSA lll and lowest in
the HP. Simpson’s index is high in MSA l and ll (especially MSA ll
Lower) and low in MSA lll and the HP. Generally, MSA l has the
highest diversity indices with the highest heterogeneity, evenness,
residual and Fisher’s alpha values.

5.3. Ungulate diversity

Ungulate indices data for KRM are presented in Table 4. The

Fig. 5. Bovid size classes through the Klasies sequence. SB ¼ Small bovid; SMB ¼ small
medium bovid; LMB ¼ large medium bovid; LB ¼ large & very large bovids. Number of
identified specimens (NISP) in columns.
number of ungulate taxa (uNTAXA) is not significantly correlated to
the log-transformed number of identified ungulate specimens
(uNISP) (rs ¼ 0.618; p ¼ 0.119) which could be related to sample
sizes. It means, however, that residuals could not be calculated.
Fisher’s alpha is significantly correlated to Simpson’s index and the
ungulate richness index (rs ¼ 0.900; p ¼ 0.037) suggesting that
these indices provide reliable measures of ungulate diversity for
KRM. Simpson’s index, Fisher’s alpha and the ungulate richness
index indicate that MSA l is the richest and most diverse period
with the ensuing MSA ll Lower and Upper the least diverse and rich.
Ungulate heterogeneity is highest in MSA l and MSA lll. The HP and
MSA l have the most even distribution of ungulate taxa, while
Simpson’s index diversity values are high throughout the sequence
suggesting that no specific ungulate taxa dominates the KRM
assemblage.

larger ungulates are more prevalent in open grasslands (Kramer

and Prins, 2010). There is also a significant difference in the pro-
portion of larger (large-medium and large) and smaller (small and
small-medium) bovids between MSA l and MSA ll Lower
(c2 ¼ 10.125; df ¼ 1; p ¼ 0.002). The relatively high proportions of
larger bovids in MSA l (implying a more open environment) and the
more equitable distributions of bovid size classes in the HP and
MSA lll layers also suggests a shift from grassier environments in
6. The Klasies River palaeoenvironment
In this section, we link data from our analysis and that of other
studies to the KRM chronology. The deposits at KRM span over
70 000 years incorporating various techno-complexes. Given this
unusually long occupational sequence, we discuss changing envi-
ronmental conditions at KRM based on chronologies defined by the
lithic techno-complexes.

Table 3
General taxonomic diversity indices at Klasies River. Highest values emboldened.

Indices MSA III HP MSA II U MSA II L MSA I Total KRM

NTAXA 19 14 17 22 15 28
NISP 191 107 270 893 66 1527
NTAXA/log NISP 8.33 6.90 6.99 7.46 8.24 8.79
Simpson’s Index (1-D) 0.58 0.63 0.72 0.76 0.75 0.75
Heterogeneity (h) 1.52 1.54 1.68 1.89 1.92 1.88
Evenness (e) 0.52 0.58 0.59 0.61 0.71 0.57
Fisher’s alpha 5.25 4.30 4.03 4.08 6.06 4.87
Residuals 1.38 1.32 2.00 1.78 1.61 2.11
 J.P. Reynard, S. Wurz / Quaternary Science Reviews 237 (2020) 106301
Table 4
Ungulate diversity indices at Klasies River. Highest diversity values in bold.
Indices MSA III HP
uNTAXA 13 11
uNISP 40 26
uNTAXA/log uNISP 8.12 7.77
Simpson’s Index (1-D) 0.85 0.87
Heterogeneity (H) 2.16 2.20
Evenness e 0.84 0.92
Fisher’s alpha 6.69 7.19
Grazer frequency 6.00 5.00
6.1. MSA l
In this phase, grazers are proportionally more common than in
other layers suggesting that the environment was likely more open
and grass-dominated. Higher ungulate diversity values at this time
also suggest a productive environment (Table 4). Micromammal
data show that the environment was likely drier (Nel et al., 2018).
Although their data still indicate an even distribution of shrub, bush
and grassy species, Nel et al. (2018) note that species which favour
more closed environments e such as golden moles (e.g., Amblyso-
mus hottentotus) e are also relatively prevalent in MSA l. This
probably reflects the mosaic environment surrounding KRM during
this time . Although Klein (1976, p. 78) argues that the proportions
of grazers, browsers and mixed-feeders during MSA l (SW layers
38-37) are likely similar to what was found in the Holocene, the
small bovid component was missing from this assemblage.
The MSA l phase is often linked to the Last Interglacial, MIS 5e
(Shackleton, 1982; Deacon and Geleijnse, 1988). Regionally, MIS 5e
was characterised by higher sea level stands and warmer temper-
atures (EPICA Community Members, 2004). Palaeo-coastal models
(Fisher et al., 2010) and data from the southern Cape (Carr et al.,
2010; Roberts et al., 2012; Cawthra et al., 2018) indicate that sea
levels may have been between 6 m and 8 m higher than present
between ~130 ka and ~125 ka. Benthic d18O records from the RC11-
83/ODP 1089 Southern Ocean marine core show large amplitude
oscillations during MIS 5e indicating warmer SST (Ninnemann
et al., 1999: 104). Data from the EPICA Dome C in Antarctica show
that this period was one of the warmest interglacial maxima
(Masson-Delmotte et al., 2010). A re-dating programme is in
progress that might change the temporal association with MIS 5e.
6.2. MSA ll
There is a significant difference between the taxa of MSA l and
MSA ll. Both general and ungulate taxonomic diversity is lowest in
MSA ll (Tables 3 and 4). Grazers become less abundant in this phase
with more mixed-feeders and a dominance of browsers. The SW
faunal data also show a decline in grazers and increase in taxa
associated with closed environments (Klein, 1976, p. 78). Interest-
ingly, micromammal data indicate that grass expanded from MSA l
through MSA ll while shrublands declined (Nel et al., 2018). Yet
both Avery (1987) and Nel et al. (2018) found that Otomys irroratus
(the Southern African vlei rat), a species associated with Afro-
montane forests, increases significantly from MSA l to MSA ll. The
presence of blue duiker (Philantomba monticola) (Table 2) e a forest
dwelling small bovid e in MSA ll supports a more forested envi-
ronment close to the site at this time. Wetland-linked species,
particularly hippopotamus are relatively prevalent in the MSA ll
layers in both the D and SW faunal samples (Klein, 1976). Estuarine
fish also increase in abundance in the SAS units which suggest the
presence of a lagoon or estuary nearby (Von den Driesch, 2004).
The Southern African vlei rat, blue duiker and the estuarine species
9
MSA II U MSA II L MSA I Total KRM
11 13 12 18
63 259 23 411
6.11 5.39 8.81 6.89
0.80 0.84 0.88 0.86
1.89 2.08 2.32 2.27
0.79 0.81 0.93 0.79
3.86 2.88 10.12 3.85
3.44 3.26 6.82 3.85
noted above imply more closed vegetation and a wetter environ-
ment perhaps related to taxonomic indices that show a decline in
diversity from MSA l to MSA ll (Nel et al., 2018).
The MSA ll at KRM is often associated with MIS 5d-a, a variable
period incorporating both warmer and cooler climatic phases
(EPICA Community Members, 2004; Wurz et al., 2018). There is
evidence of increased precipitation occurring in south-eastern
South Africa from ~120 ka to ~115 ka (Simon et al., 2015). Higher
precipitation at this time is also demonstrated in multiple core
datasets including marine core CD154-17-17K, situated near the
mouth of the Great Kei River (Simon et al., 2015), core CD154-
10e06P about 160 km off the KwaZulu-Natal coast (Ziegler et al.,
2013), core MD96-2094 on Walvis Ridge in the southeast Atlantic
Ocean (Stuut et al., 2002; Braun et al., 2019) and the Tswaing impact
crater sediment core near Pretoria (Partridge et al., 1997; Kirsten
et al., 2007). The Southern Ocean marine core RC11-83/1089
shows a long interval between d18O amplitude oscillations from
115 to 95 ka (Ninnemann et al., 1999) suggesting relatively stable
conditions then. This may correlate to a cold spell suggested by Van
Andel (1989, p. 141) between 110 ka and 105 ka based on micro-
mammal data analysed by Thackeray (1987) (Hillestad-Nel, 2013).
There is also some evidence of a short-lived sea-level stillstand at c.
107e108 ka that may have occurred after extensive dune deposi-
tion at c. 110 ka (Cawthra et al., 2018, p. 170). Pollen and charcoal
data from Vankervelsvlei suggest that the early MIS 5 phases are
associated with relatively warmer temperatures and mesic condi-
tions while data for MIS 5b-a reflect a shift to cooler temperature
(Quick et al., 2016). Speleothem data from Crevice Cave, Staircase
Cave and PP29 near Pinnacle Point show that winter rain and C3
grasses were dominant during MIS 5d-a, with the gradual expan-
sion of summer rain and C4 grass towards the beginning of MIS 4
(Bar-Matthews et al., 2010; Braun et al., 2019). Langejans et al.
(2017, p. 76) suggest that, during MSA ll Lower at KRM, shellfish
was probably accumulated during an interglacial phase when SST
were not significantly lower and sea levels were still relatively close
to present day levels. On the whole, although SST decreased slightly
during MIS 5 (Bard and Rickaby, 2009), they were generally stable
throughout this period (Loftus et al., 2017). Marine regressions
would have been affected by stadial and interstadial phases but was
likely never as extreme as during MIS 6 or 4 and the shoreline was
probably never more than 10 km away from the KRM coast (Dor,
2017).
6.3. Howiesons Poort
Ungulate and general taxonomic diversity is slightly higher in
the HP compared to MSA ll but not as high as MSA l (Tables 2 and 4).
Although samples sizes are small, grazers are, for all the HP layers
combined, more common during this period than in MSA ll (Fig. 4).
The SW faunal sample also shows an increase in grazing ungulates
from MSA ll to the HP suggesting a more open environment during
certain periods of the HP (Klein, 1976). This is supported by Avery’s
10 J.P. Reynard, S. Wurz / Quaternary Science Reviews 237 (2020) 106301
(1987) analysis which shows that micromammals indicative of
closed environments, especially vlei rats, become less common
through the HP. Large bovids are more prevalent in these layers
which appear to be corroborated by the abundance of eland, buffalo
and alcelaphines in the SW assemblage. However, both in the D and
SW samples browsing ungulates dominate the lower HP layers
suggesting that the earlier phases may have been more closed that
the later period (Klein, 1976; Van Pletzen, 2000; Van Pletzen-Vos
and Wurz, 2014). Although species favouring rocky shores still
dominate the shellfish assemblage, they are less common than in
the MSA ll (Thackeray, 1988a; Langejans et al., 2017). Langejans
et al. (2017, p. 77) argue that the increase in alikreukel and
turban shell suggests that tidal pools systems were more extensive
during this time e probably because of a more distant shoreline.
The HP at KRM is generally associated with MIS 4 (Deacon,
1989), a major glacial period encompassing significant de-
velopments in what has been termed ‘modern’ behaviour (Deacon,
1989; Henshilwood and Marean, 2003) or innovative complex
behaviour and cognition (Wadley, 2013; Wurz, 2013). Micro-
mammal data (Thackeray, 1987) and SST models (Ninnemann et al.,
1999; Bard and Rickaby, 2009; Braun et al., 2019) indicate that this
period was cooler than the preceding MSA ll phase. Chase and
colleagues (Chase and Meadows, 2007; Chase, 2010; Carr et al.,
2016) have argued that instead of being drier (Avery, 1983;
Deacon, 1989), conditions during MIS 4 were likely to have been
more humid due to the influence of warmer southern Indian Ocean
SST and the effects of westerly storm fronts. While a spike in Stoebe-
type pollen at c. 67 ka around Vankervelsvlei may be evident of
drought stress (Quick et al., 2016), data from marine core CD154-
17-17K show wetter conditions in the Eastern Cape between 65
and 60 ka; although this period was probably also interspersed
with arid events (Simon et al., 2015; Ziegler et al., 2013). Speleo-
them data from Crevice Cave and PP29 near Pinnacle Point indicate
unstable environmental conditions between ~73 and 64 ka with a
dominance of C4 plants during the latter part of MIS 4 suggesting
more regular summer rainfall then in that area (Bar-Matthews
et al., 2010; Braun et al., 2019). Phytolith evidence at Pinnacle
Point also shows a dominance of thicket and riparian vegetation e
in addition to fynbos and C4 grasses during MIS 4 (Esteban et al., in
press).
6.4. MSA lll
Long-horned buffalo e an extinct, probable grazer requiring
substantially open terrain e are proportionally more common in
these layers than any other phase of the D sample. Here, we also
find evidence of wetlands with southern reedbuck and a Kobus
specimen e very likely waterbuck e in the D sample, and reedbuck
and hippo recovered from the SW assemblage (Klein, 1976).
Although the sample is small, Cape dune mole rats are also pro-
portionally quite common in these layers (Table 1) suggesting
increased dune activity during this period. Micromammal data
show that this period was dominated by grass-indicator species
such as Hottentot golden moles (Amblysomus hottentotus), with a
corresponding decline in mole rats (Cryptomys hottentotus and
Georychus capensis) (Avery, 1987). In the SW assemblage, some of
the few examples of wildebeest from the southern Cape were
recovered from the MSA lll period suggesting an open environment
dominated by grasslands (Klein, 1976). The faunal data thus show
variable signals that may reflect fluctuating environmental
conditions.
Regionally, this period likely corresponds to the interstadial MIS
3, a variable phase marked by cooler periods with warm oscillations
(EPICA Community Members, 2004). SSTs would have probably
been slightly warmer compared to MIS 4 and the environment, in
general, may have been drier than in the Holocene (Ziegler et al.,
2013; Quick et al., 2016) although Agulhas Current SSTs Reynard
et al., 2016a show a steady decline through this period (Bard and
Rickaby, 2009). Benthic d18O records from marine core TN057-21
show general stability during MIS 3 with no discernible vari-
ability on millennial scales (Ninnemann et al., 1999, p. 104). The
Crevice Cave and PP29 speleothem records show a slight increase in
winter rain with a corresponding expansion of C3 vegetation (Bar-
Matthews et al., 2010; Braun et al., 2019). Sea levels would have
receded from ~60 ka to the Last Glacial Maximum (LGM) exposing a
wide Palaeo-Agulhas Plain for the southern Cape area, especially
west from KRM, during most of this period (Marean et al., 2014).
Evidence of grassland species from both the micromammal and
ungulate data seem to support this scenario.
7. Discussion
7.1. The Klasies sequence as environmental proxy
A key question that needs to be addressed is whether the faunal
data from the D sample are useful proxies for environmental
change in the southern Cape. It must be noted that archaeo-faunal
assemblages, such as those from KRM, would reflect human biases
in prey selection. Faunal remains in hyena-accumulated assem-
blages correlate well with animal abundance on the landscape
(Hayward, 2006). Human-selected prey may be influenced by
hunting preferences and strategies, nocturnal habits of prey and
technological capabilities (Discamps et al., 2011), and may not
directly reflect contemporaneous faunal communities. That said,
while it is difficult to discern the influence of human behaviour on
prey selection in the late Pleistocene, analyses of contemporary
Kwee forager assemblages in the Kalahari show that human
hunters exploit prey similar to their occurrence within local habi-
tats (Bunn, 1982).
Southern Cape archaeo-faunal assemblages reflect shifting un-
gulate dietary preferences that appear to indicate changing climatic
conditions. Fig. 6 shows a shift from a dominance of grazers in MIS
6, to more mixed-browsers at during MIS 5, and back to more
grazers in MIS 4 and 3. Generally in the southern Cape, sites dated
to MIS 6 (of which Pinnacle Point is the only one) show open
Fig. 6. Ungulate dietary preference in the southern Cape from MIS 4e6. Klipdrift
Shelter (KDS) data from Reynard et al. (2016a), b; Die Kelders (DK1) data from Klein
et al. (2000); Blombos Cave (BBC) data from Henshilwood et al. (2001); Pinnacle Point
(PP) 13B & 30 data from Rector and Reed (2010). Gr ¼ Grazers, Mb ¼ Mixed-browsers.
Number of identified specimens (NISP) in columns.
 J.P. Reynard, S. Wurz / Quaternary Science Reviews 237 (2020) 106301 11

environments which may correspond to an expanded Palaeo-

Agulhas Plain during this glacial period. This appears similar to
grazer frequencies during MSA l at KRM. At Blombos Cave, dated to
between MIS 5d-4, grazers are not common and the environment is
generally closed and bushy, comparable to the MSA ll layers at KRM
(Badenhorst et al., 2016; Reynard and Henshilwood, 2019). The
Klipdrift HP layers (dated to MIS 4) show similar patterns to the HP
at KRM with a prevalence of grass-loving taxa (Reynard et al.,
2016a). The one exception is Die Kelders which is dated to be-
tween 70 and 60 ka (MIS 4; Feathers and Bush, 2000) and shows a
prevalence of browsers (Klein and Cruz-Uribe, 2000). However, Die
Kelders has an extraordinary abundance of Raphicerus (7609 out of
9107 ungulate taxa) and Klein and Cruz-Uribe (2000, p. 173) notes
that all extralimital species at Die Kelders such as wildebeest,
reedbuck, Cape zebra and long-horn buffalo were all grazers that
imply grassier vegetation. In their study on diet breadth in the MSA,
Clark and Kandel (2013) note that browsers are more prevalent in
MIS 4 and MIS 6. Unlike our analyses, their data includes faunal
assemblages from a range of southern African MSA sites not just the Fig. 7. The relationship between ungulate diversity and grazer frequency at Klasies
southern Cape, and Pinnacle Point 30 (a carnivore den dated to MIS River main. MSA ll U ¼ Upper, L ¼ Lower.
6) is also excluded from their analysis. They argue that the higher
proportion of browsers in MIS 4 in their dataset may not just reflect
bushy vegetation in MSA ll, shifting back to more grasslands in
environmental conditions but may be linked to subsistence
some parts of the HP and MSA lll (Klein, 1976). Larger ungulates are
intensification.
significantly more common in MSA l (Fig. 5). With the exception of
Ungulate dietary patterns from the various southern Cape sites
eland, larger herbivores generally consume lower quality forage
we examined suggest that Late Pleistocene faunal assemblages in
such as grass (Prins and Olff, 1998; Frits and Loison, 2006) which
this area are very likely time-averaged. Fig. 6 therefore represents a
suggests that grasslands were an important component of the KRM
palimpsest of data from MIS 6 to 3. Representations of data such as
ecology in MSA 1.
these tend to highlight more extreme shifts in environmental
We were unable to measure residual indices through the
conditions brought about by glacial/interglacial cycling. It is also
sequence because uNTAXA does not correlate to log uNISP in the D
likely that more severe environmental conditions could influence
sample, thus we cannot quantify the relationship between ungulate
taphonomic and post-depositional processes which would affect
diversity and precipitation. Yet, it is likely that shifting ungulate
faunal preservation. This is probably why at most sites the two
diversity and richness at KRM are associated with glacial and
more severe glacial periods e MIS 6 and 4 e show the most sig-
interglacial cycles which, in turn, is linked to moisture availability.
nificant changes in grazer representation (Fig. 6). This pattern is
MAP near KRM currently averages around 1000 mm/year (Van Wijk
also evident at KRM (Fig. 4) suggesting that the KRM dataset re-
et al., 2017, p. 15). If we assume that MAP did not drop below
flects a similar palimpsest. Notwithstanding these site formation
750 mm/year in the late Pleistocene, it is possible that increasing
biases, the extensive sequence at KRM shows how changing envi-
ungulate diversity for this sample could reflect less moisture
ronmental conditions affected ungulate abundance and reflects
availability (Thackeray, 1980; Faith, 2013b). In that case, increasing
trends across the southern Cape. Thus, despite the relatively small
ungulate richness may imply drier conditions. This is a reasonable
size of the D sample compared to the SW assemblage, proportions
suggestion since ungulate biomass is usually lower in forested re-
of grazers, mixed-feeders and browsers in our analysis generally
gions, increasing in more open, drier environs such as those in the
corresponds to patterns seen at other southern Cape sites (compare
Savannah and Grassland Biomes of the southern African interior (cf.
Figs. 6 and 4). This suggests that it is feasible to use the D sample to
Coe et al., 1976; Olff et al., 2002; Faith, 2013b). Thus, the current
infer palaeoenvironmental patterns and the KRM sequence may be
mosaic environment surrounding KRM would likely become more
a useful proxy for environmental change in this region.
forested with increasing precipitation, similar to the Afrotemperate
forests in the Klasies River valley and near present-day Knysna
7.2. Environmental change and ungulate diversity
further to the west (Mucina and Rutherford, 2011).
The high diversity values in MSA 1 may therefore suggest less
There is a clear link between environmental conditions and
available moisture during that period. This raises the question of
ungulate diversity at KRM (Fig. 7). Fisher’s alpha, the ungulate
whether we can link ungulate diversity in the MSA l layers to either
richness index and the frequency of grazers (tabulated as the per-
glacial or interglacial phases. While data from marine core CD154-
centage of grazers divided by ten) are significantly correlated
17-17K show no firm relationship between cooler/warmer phases
throughout the sequence (rs ¼ 0.900, p ¼ 0.037; Fig. 7). Simpson’s
in MIS 5 or 4, and humidity (Ziegler et al., 2013), there is substantial
index is also significantly correlated with Fisher’s alpha (rs ¼ 0.900,
evidence that glacial MIS 4 was humid in the south-western Cape
p ¼ 0.037). Although this index is not significantly associated with
(Stuut et al., 2004; Chase, 2010; Ziegler et al., 2013). However,
either the richness index or grazer frequency (rs ¼ 0.800, p ¼ 0.104),
conceptual models (Cockcroft et al., 1987; Partridge et al., 1990;
the correlation coefficient between these indices is very high. These
Engelbrecht et al., 2019) and proxy data (Avery, 1983; Deacon et al.,
indices peak in MSA l, dropping substantially in MSA ll before rising
1984; Deacon and Lancaster, 1988; Cowling et al., 1999; Meadows
slightly again in the HP and MSA lll. This relationship shows that
and Baxter, 1999; Chase and Meadows, 2007; Chase et al., 2017)
ungulate diversity is linked to changing vegetation and environ-
suggest the eastern part of the GCFR may have been dryer during
mental conditions. Specifically, it indicates that, as ungulate rich-
glacial periods. A recent climate model by Engelbrecht et al. (2019)
ness increases, grazers e and by implication, grasslands e become
projects a reduction in rainfall to the east of the Cape south coast
more prevalent. The SW sample also shows more grazers, and
during the LGM. They suggest that this may possibly be linked to
therefore a more open environment in the MSA l moving to closed,
12 J.P. Reynard, S. Wurz / Quaternary Science Reviews 237 (2020) 106301
the effects of a rain shadow along the south-north aligned Cape
Fold mountains during the northward displacement of westerly
frontal systems (Engelbrecht et al., 2019, p. 16) and more frequent
hot and dry berg wind conditions to the east (p. 24) in glacial pe-
riods. It is possible then that the implied dryer conditions in MSA l
is more likely to correspond to a glacial period. That said, Braun
et al. (2019) have suggested that moisture availability in the
southern Cape is highly dependent of river runoff from the interior
so diversity indices for the southern Cape may be linked to interior
rainfall regimes. Furthermore, limited land availability resulting
from glacial-linked marine regressions would also affect ungulate
diversity (Marean, 2010; Faith, 2013b; Venter et al., in press) which
may also be one of the reasons why diversity indices are so high in
MSA 1.
7.3. Environmental trends at Klasies River
Diversity data may inform us on human occupational intensity
in the MSA lll. Singer and Wymer (1982, p. 21) note that the MSA lll
units (corresponding to Layers 1 to 9 in Cave 1A) generally consist
of ‘sandy scree’ with ‘laminated ash’ in the lower layers. The smaller
archaeological sample from the Deacon excavations (Wurz, 2000)
also implies that the MSA lll at KRM was marked by low occupa-
tional activity. The faunal sample bears some evidence of this and is
characterised by a significant increase in small mammals compared
to bovids (Van Pletzen-Vos et al., 2019). General taxonomic di-
versity (as defined by richness and Fisher’s alpha) is relatively high
in the MSA lll (Table 2). This is probably a result of the abundance of
small mammals in these layers which may have been accumulated
by carnivores rather than people (cf. Pearson et al., 2019).
Researchers often investigate the links between HP lithic as-
semblages and environmental conditions (e.g., Deacon, 1989;
Ambrose and Lorenz, 1990; Reynard et al., 2016a). Yet at KRM, these
links may be tenuous. As noted previously, the HP is associated with
MIS 4 when shorelines would have been relatively far from the
main site. Yet, marine core data show evidence of fluctuating
environmental conditions during MIS 4 (e.g., Ziegler et al., 2013)
and palaeogeographic and micro-climatic conditions would likely
have affected site-specific marine regressions (Carr et al., 2016).
Furthermore, the HP data presented here are a time-averaged,
lumped dataset that does not reflect the nuanced environmental
fluctuations during this period. This is probably why seal remains
dominate this phase even though the coastline must have been
relatively far away at certain periods of the HP.
Diversity patterns indicate a significant environmental shift
between MSA ll and l at KRM. Evidence suggests that SST were not
significantly variable from MIS 5d-a (Langejans et al., 2017; Loftus
et al., 2017). Marine core data also show relative climatic stability
from about 110 to 90 ka (Ninnemann et al., 1999; Simon et al., 2015).
Faunal data from the MSA ll layers at KRM and the M3 at Blombos
indicate a prevalence of mixed-browsers over grazers (Figs. 4 and
7). The shift from MIS 6 to 5e encompasses a much more signifi-
cant environmental change than variations within MIS 5 (e.g.,
Ninnemann et al., 1999; Masson-Delmont et al., 2010; Rector and
Reed, 2010; Marean et al., 2014). Research suggests that a signifi-
cant portion of the Palaeo-Agulhas Plain was reclaimed by the
ocean between ~137 ka and the beginning of MIS 5e (~128 ka)
(Marean et al., 2014) and that these marine reclamations were
likely rapid (Cawthra et al., 2018). Although KRM lies to the east of
Palaeo-Agulhas Plain and ungulate ranges would probably have
included areas north of KRM, this rapid sea-level rise would have
had a substantial impact on ungulate communities in this region,
decimating viable grazing pastures, disrupting riverine habitats and
possibly increasing bovid population pressure.
The presence of seal during the MSA l shows that KRM people
had access to the coast and also suggests that this period likely
occurred when the coastal plain was contracting. Furthermore, it
also reflects the importance of seal sculp (subcutaneous fat and
skin) for local foragers since this is a rich and highly valued source
of fat (Marean, 1986). Given the significance of seal as a fat resource,
it is likely that coastal foragers would have still transported seal
remains even when the shoreline was relatively far way. Research
on the intensity of seal field processing and transport decisions
could shed light on how valuable this resource was to KRM
populations.
Differences in ungulate diversity and species composition show
a significant shift in habitat between MSA l and MSA ll which may
correspond to a glacial/interglacial shift. While it is possible that
this could signify a change in environmental conditions from MIS
5e to 5d, it should be noted that the layer above deposits containing
MSA l artefacts e from the SASL sub-member, the base of the SAS
member e yielded dates of 110 ka from in situ speleothem material
(Brenner and Wurz, 2019). There is almost a meter of MSA l deposit
below this sub-member. Based on the trend of chronometric dates
in the layers above, it is likely that this meter-thick deposit would
correspond to at least several thousand years of occupation, as
suggested by Deacon and Geleijnse (1988, p. 8). It is therefore
possible that the age of the MSA l sub-member may date to beyond
120 ka, and be older than the accepted age range of MIS 5e (Deacon
and Geleijnse, 1988). As noted earlier, high ungulate diversity in
MSA l probably indicates dryer conditions; possibly corresponding
to glacial phases in the eastern GCFR (cf. Engelbrecht et al., 2019).
Furthermore, using Hutchinson’s weight ratio theory, Venter et al.
(in press) propose that species richness in the southern Cape
would have been high during low sea level conditions such as in the
glacial periods of MIS 6. We noted that the grazer-rich periods of
glacial MIS 4 and 6 bracket the browser-rich phase of MIS 5 in the
southern Cape. The differences in ungulate diversity and grazer
frequencies between the MSA l and MSA ll at KRM are remarkably
similar to changes between MIS 6 and 5 sites shown in Fig. 6.
Indeed, the similarities between proportions of grazers in the MSA
1 layers and Pinnacle Point assemblages dated to MIS 6 are also
striking. The Pinnacle Point 30 faunal assemblage is dominated by
grazers and, while the Lightly Cemented MSA Lower (LC-MSA)
horizon in Pinnacle Point 13B (with an age of 153e174 ka) has a
small sample of fauna, most of these are grazing alcelaphines
(n ¼ 4). The prevalence of grazers in these assemblages is probably
not a result of the differences in human and carnivore accumula-
tion. Given that the PP30 and LC-MSA assemblages were carnivore
and human-accumulated, respectively (Marean, 2010), it is likely
that these faunal remains are a reasonable reflection of MIS 6
environmental conditions, and are similar to what was found in the
MSA l layers at KRM.
7.4. Ungulate diversity at other southern Cape Middle Stone Age
sites
Comparing KRM to other MSA sites is complicated because of
KRM’s long archaeological sequence. Because of this (and for sta-
tistical reasons), we group the KRM sequence into two phases: the
post-80 ka period (MSA lll and the HP or upper KRM) and pre-80 ka
period (MSA ll and MSA l or lower KRM). Each of these periods is
then compared to contemporaneously dated MIS 4 to 5a sites. We
focus on ungulate diversity since this is used to explore precipita-
tion and habitat productivity (Thackeray, 1980; Olff et al., 2002;
Faith, 2013b). Table 5 shows the upper KRM (the MSA lll and HP)
compared to data from Klipdrift Shelter (Reynard et al., 2016a, b),
Blombos Cave’s M1 and M2 Phases (Henshilwood et al., 2001), the
Diepkloof Rock Shelter sequence from the post-HP to the Still Bay
layers (Steele and Klein, 2013) and the Die Kelders MSA layers
 J.P. Reynard, S. Wurz / Quaternary Science Reviews 237 (2020) 106301 13

Table 5
Ungulate diversity indices in the upper Klasies layers and at various MIS 3e4/5a sites. Highest diversity values in bold.

Indices KRM Upper BBC M1 & M2 KDS DRS (SB to-Post-HP) DK1

uNTAXA 15 10 14 20 15
uNISP 66 284 111 1757 9107
uNTAXA/log uNISP 8.24 4.08 6.85 4.92 3.79
Simpson’s index (1-D) 0.87 0.62 0.81 0.83 0.30
Heterogeneity (H) 2.33 1.38 2.07 1.11 0.80
Evenness (e) 0.86 0.60 0.78 0.37 0.29
Fisher’s alpha 6.06 2.02 4.24 3.16 1.75

(Klein et al., 2000). Most diversity values are higher at KRM while relationships between landscape, precipitation, evapotranspiration
Die Kelders is the least rich. These periods generally correspond to and vegetative regimes (Cowling, 1992; Cowling and Lombard,
MIS 5a (a warm phase), MIS 4 (a cooler phase) and e at KRM e MIS 2002). Moisture availability is probably similar between the
3 (an intermediate phase). southern and western regions of the GCFR (Bradshaw and Cowling,
For comparison with the lower KRM layers (MSA ll and MSA l), 2014) but primary productivity may be higher in the southern/
we limit our analysis to the ‘Mike’ and ‘Lower’ layers at Diepkloof eastern part (Cowling et al., 1999). In the Holocene and Pleistocene,
Rock Shelter (DRS; Steele and Klein, 2013) dated to ~ 100 ka, the M3 ungulate taxonomic richness was probably higher in the southern
phase at Blombos Cave (BBC; Henshilwood et al., 2001; Badenhorst part of the GCFR than the western region (Skead, 1987; Klein, 1983;
et al., 2016; Roberts et al., 2016), (dated to ~ 110-90 ka), Pinnacle Rector and Verrilli, 2010; Reynard and Henshilwood, 2017; Luyt
Point (PP; Rector and Reed, 2010) 13B (~135-90 ka) and PP 30 (~151 et al., 2019) and our data appear to support this (Tables 5 and 6).
ka) (Table 6). Various indices indicate that the lower KRM sequence Fossil trackways, for example, show that very large herbivores such
and PP30 have the most diverse assemblages, with DRS the least as giraffe and elephant were present in the southern Cape in the
rich. This is probably because fauna from DRS is extensively frag- Pleistocene indicating a mesic environment capable of sustaining
mented and identification was difficult (Steele and Klein, 2013). It large animal biomass (Roberts et al., 2008; Helm et al., 2019).
must be noted that PP30 is a hyena-accumulated den and these are
generally richer than human-selected assemblages (Rector and
Reed, 2010). To further examine diversity between these sites, a 8. Conclusion
regression analysis was conducted where residuals from the
reduced major axis regression between log-transformed ungulate In this study, we examine the palaeoecologiocal implications of
sample sizes and uNTAXA. This could be undertaken because log- the large mammal fauna from the Deacon excavations of KRM. We
transformed uNISP are significantly correlated to uNTAXA be- test hypotheses that propose that higher ungulate diversity corre-
tween the various MSA sites (rs ¼ 0.900; p ¼ 0.037). This metric sponds to increased grazer abundance in the southern Cape during
shows that the lower KRM layers have the richest ungulate the late Pleistocene (Faith, 2013a, 2013b). The length of the
assemblage of all the sites measured while ungulate richness in the sequence e spanning over 70 000 years e make this an important
M3 phase at BBC is the lowest (Table 6). Given that MSA ll has low proxy in understanding environmental change in this part of the
diversity values, the increased diversity of the lower KRM layers is southern Cape. Most southern Cape MSA environmental proxies are
likely a result of high ungulate richness/diversity in MSA l. It is from the south/south-western region of the Fynbos Biome. The
interesting to note that BBC M3 and MSA ll Lower e which are position of KRM near the eastern border of the GCFR means that we
broadly contemporaneous e both reflect low ungulate diversity, are able to explore environmental conditions in the later Pleisto-
and browsers dominate both phases (Fig. 4; Henshilwood et al., cene in the more environmentally-mosaic eastern region.
2001). On the whole, ungulate diversity indices for both the up- Although the D sample as a whole (and the MSA l assemblage in
per and lower KRM assemblages show that the environment at particular) is relatively small, the fact that proportions of ungulate
KRM was likely more productive than at the other southern Cape species are generally similar to the SW sample suggests that we can
sites in our sample. Faunal proxies indicate that environments were use these data to infer environmental change. The problematic
generally richer in the southern Cape further east: similar to sampling methods of the SW assemblage means that the D sample
present-day conditions (Rector and Verrelli, 2010; Faith and may be even more representative of faunal communities at KRM.
Thompson, 2013; Reynard and Henshilwood, 2017; Nel and Indeed, shifts between grazers and mixed-browsers through the
Henshilwood, 2016). KRM sequence corresponds to similar changes in other southern
Climatic variability within the GCFR is a result of complex Cape faunal assemblages (Fig. 6). Beside the ubiquitous hyrax, seal
are the most common taxa. With regard to ungulates, eland and
Raphicerus dominate the assemblage. The prevalence of those taxa,
and the presence of both browsers and grazers throughout the
Table 6
sequence indicate that, generally, KRM was a mosaic environment
Ungulate diversity indices in the lower Klasies layers and at various MIS 5 sites.
Highest diversity values in bold. of grasslands interspersed with thicket. Our data show a general
shift from more grazers in the MSA l to a prevalence of mixed-
Indices KRM lower BBC M3 PP13B PP30 DRS (M&L)
feeders in the MSA ll, to a dominance again of grazers in the HP
uNTAXA 15 9 10 13 5 and MSA lll layers (Fig. 4). This suggests that the environment
uNISP 345 96 105 343 18
changed from grass-dominated, to mixed-browse and back to
uNTAXA/log uNISP 5.91 4.54 4.95 5.13 3.98
Simpson’s index (1-D) 0.85 0.54 0.84 0.88 0.77 grassier.
Heterogeneity (H) 2.17 1.20 1.98 2.23 1.52 While the diversity of species during the HP and MSA lll show a
Evenness e 0.80 0.54 0.86 0.87 0.94 mosaic environment, the relative abundance of grazers also sug-
Fisher’s alpha 3.20 2.43 2.72 2.67 2.29 gests an increase in grasslands during these periods. The HP is
Residuals 1.24 0.80 0.08 0.74 0.38
normally linked to the MIS 4 and its associated marine regressions.
14 J.P. Reynard, S. Wurz / Quaternary Science Reviews 237 (2020) 106301
Yet our data show an abundance of seal remains during this period
which suggests that, at least for some of the time, the site was close
to the coast. This may be a product of time-averaged faunal as-
semblages over a relatively long HP occupational sequence (Wurz,
2000). In all likelihood, the HP included periods of both close and
distant shorelines. The MSA lll layers show high diversity values but
the prevalence of small mammals probably contributes to the
higher richness index in those layers. This and other archaeological
evidence (cf. Singer and Wymer, 1982) suggest that these layers
were likely infrequently occupied.
Ungulate diversity data from various MSA sites show that the
southern Cape was not a homogenous environmental region during
the late Pleistocene. With the exception of the hyena-den PP30,
ungulate diversity for both the upper and lower KRM sequence is
significantly higher than at any other sites in our sample. Our data
indicates that the region surrounding KRM may have had even
higher habitat productivity than other areas of the southern GCFR,
suggesting that primary productivity may increase along a gradient
from west to east.
Environmental change at KRM is likely associated with glacial/
interglacial shifts. The data show significant relationships between
diversity indices and grazer proportions through the KRM
sequence, suggesting that ungulate diversity is affected by chang-
ing vegetation regimes. It is possible that variation in ungulate di-
versity and richness may reflect changes in moisture availability
through the KRM sequence. However, because uNTAXA does not
correlate to log uNISP through the sequence, we were unable to test
this using residual indices. Because forested environments are
generally not associated with high herbivore biomass, it could
mean that increasing ungulate diversity may relate to a decrease in
moisture availability. However, the abundance of ungulate species
may have also been affected by a fluctuating Palaeo-Agulhas Plain.
On the whole, most general taxonomic indices are highest in the
MSA 1, while all ungulate indices are higher in this phase.
We suggest that the MSA 1 sub-member may be older than MIS
5e based on the following lines of evidence. Firstly, the basal SASL
sub-member e the layer above deposits containing MSA l artefacts
e yields dates of 110 ka from in situ speleothem material. Given the
almost meter thick layer of MSA l deposit below this material that
may be of a duration of several thousands of years, it suggests that
the MSA 1 layers are older than 120 ka. Secondly, the substantial
loss of habitat that would have occurred along the southern coast
between MIS 6 and 5e, would have resulted in major environmental
changes and disruptions in ungulate populations in the region, and
this appears to be reflected in the significant difference in ungulate
diversity values between MSA ll and MSA l. Finally, the similarities
between the KRM MSA 1 faunal sample and the oldest Pinnacle
Point faunal assemblages may also imply that this unit dates to MIS
6.
This study has significant implications for understanding MSA
hunting strategies by showing how ungulate diet breadth may be
closely linked to environmental conditions in this region of the
southern Cape. Data such as these from MSA sites in other regions
may also help us understand the relationship between hunting
behaviour and environmental conditions. That said, it is critical to
explore the taphonomy of these faunal assemblages. The soon-to-
be-published taphonomic analyses at KRM will also be important
in elucidating the subsistence behaviour of early modern humans
during the MSA. Currently, MSA l deposits at KRM are being re-
dated and the excavations conducted in the lower Witness Baulk
layers are yielding an abundance of fauna. Data from these as-
semblages will contribute much to our understanding of faunal
exploitation patterns and the palaeoecology of people during the
earlier sequence at KRM and may unpack a more detailed account
of environmental change during the early Late Pleistocene.
Author contributions
Both authors have made substantial contributions to the paper.
Dr. Reynard initiated and conceptualized the research question,
analysed the faunal data and co-wrote the paper. Prof. Wurz co-
wrote the paper, obtained the necessary permits to excavate the
site, assisted in the review of literature and refined the conclusions.
Both authors contributed to funding the research project and have
approved the re-submitted manuscript.
Declaration of competing interest
The authors declare that they have no known competing
financial interests or personal relationships that could have
appeared to influence the work reported in this paper.
Acknowledgements
This paper is dedicated to the memory of James Brink
(1957e2019). We thank him and Liezl van Pletzen-Vos for allowing
us access to their data. JR is funded by a South African Department
of Science and Technology (DST)-National Research Foundation
(NRF) Thuthuka grant (grant no. 107082) and an Enabling Grant
through the Diversifying of the Academy from the University of the
Witwatersrand. Any opinion, finding, conclusion or recommenda-
tion expressed in this article are those of the authors and the NRF
does not accept any liability in this regard. SW would like to
acknowledge the support of the DST-NRF Centre of Excellence in
Palaeosciences (CoE-Pal) towards this research. Opinions expressed
and conclusions arrived at, are those of the author and are not
necessarily to be attributed to the CoE-Pal.
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