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Follow the Senqu: Maloti-Drakensberg Paleoenvironments and Implications for Early Human Dispersals into Mountain Systems


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The Maloti-Drakensberg Mountains are southern Africa’s highest and give rise to South Africa’s largest river, the Orange-Senqu. At Melikane Rockshelter in highland Lesotho (~1800 m a.s.l.), project AMEMSA (Adaptations to Marginal Environments in the Middle Stone Age) has documented a pulsed human presence since at least MIS 5. Melikane can be interrogated to understand when and why early modern humans chose to increase their altitudinal range. This paper presents the results of a multi-proxy paleoenvironmental analysis of this sequence. Vegetation shifts are registered against a background signal of C3-dominated grasslands, suggesting fluctuations in temperature, humidity and atmospheric CO2 within a generally cool highland environment with high moisture availability. Discussing Melikane in relation to other paleoenvironmental and archeological archives in the region, a model is developed linking highland population flux to prevailing climate. It is proposed that short-lived but acute episodes of rapid onset aridity saw interior groups disperse into the highlands to be nearer to the Orange-Senqu headwaters, perhaps via the river corridor itself.
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Chapter 14
Follow the Senqu: Maloti-Drakensberg Paleoenvironments
and Implications for Early Human Dispersals into Mountain
Brian A. Stewart, Adrian G. Parker, Genevieve Dewar, Mike W. Morley, and Lucy F. Allott
Abstract The Maloti-Drakensberg Mountains are southern
Africas highest and give rise to South Africas largest river,
the Orange-Senqu. At Melikane Rockshelter in highland
Lesotho (*1800 m a.s.l.), project AMEMSA (Adaptations
to Marginal Environments in the Middle Stone Age) has
documented a pulsed human presence since at least MIS 5.
Melikane can be interrogated to understand when and why
early modern humans chose to increase their altitudinal
range. This paper presents the results of a multi-proxy
paleoenvironmental analysis of this sequence. Vegetation
shifts are registered against a background signal of C
dominated grasslands, suggesting uctuations in tempera-
ture, humidity and atmospheric CO
within a generally cool
highland environment with high moisture availability.
Discussing Melikane in relation to other paleoenvironmental
and archeological archives in the region, a model is
developed linking highland population ux to prevailing
climate. It is proposed that short-lived but acute episodes of
rapid onset aridity saw interior groups disperse into the
highlands to be nearer to the Orange-Senqu headwaters,
perhaps via the river corridor itself.
Keywords Dispersals !Late Pleistocene !Later Stone
Age !Lesotho !Middle Stone Age !Mountain foraging !
Orange-Senqu River !Paleoenvironment
Diverse lines of evidence suggest that Africa between MIS
6-2 experienced a complex demographic history (Cornelissen
2002; Batini et al. 2007,2011; Tishkoff et al. 2007,2009;
Barham and Mitchell 2008; Behar et al. 2008; Atkinson et al.
2009; Castañeda et al. 2009; Pereira et al. 2010; Scheinfeldt
et al. 2010; Drake et al. 2011; Henn et al. 2011; Blome et al.
2012; Pickrell et al. 2012; Schlebusch et al. 2012,2013;
Soares et al. 2012; Veeramah et al. 2012; Barbieri et al. 2013;
Cancellieri and di Lernia 2013; Coulthard et al. 2013; Foley
et al. 2013; Lombard et al. 2013; Mercarder et al. 2013; Rito
et al. 2013). Frequent oscillations in climate, environment
and natural resources contributed to variability in human
adaptive strategies, including shifts in population sizes,
structures and movements. Demographic ux appears to have
been exacerbated in the continents higher latitudes where
environments are more water stressed than in equatorial East
and tropical Africa (Blome et al. 2012).
In southernmost Africa, resource-rich coastal environ-
ments closely neighbor those of the less productive, more
seasonal interior plateau. At the root of this ecological
imbalance are the subcontinents peripheral mountains that
divide the coast and interior. Rising steeply from the narrow
coastal forelands, the Great Escarpment and its associated
ranges form a 5000 km-long arc from northern Angola to
eastern Zimbabwe (Partridge and Maud 1987; Birkenhauer
B.A. Stewart (&)
Museum of Anthropological Archaeology, University of
Michigan, Ruthven Museums Building, 1109 Geddes Avenue,
Ann Arbor, MI 48109, USA
A.G. Parker
Human Origins and Palaeoenvironments Research Group,
Department Of Social Sciences, Faculty of Humanities and Social
Sciences, Oxford Brookes University, Headington, Oxford, OX3
G. Dewar
Department of Anthropology, University of Toronto, Scarborough
1265 Military Trail, Toronto, ON M1C 1A4, Canada
M.W. Morley
Centre for Archaeological Science, University of Wollongong,
Room 268, Building 41, Northelds Ave, Wollongong, NSW
2522, Australia
L.F. Allott
Institute of Archaeology, University College London, 31-34
Gordon Square, London WC1H 0PY UK
©Springer Science+Business Media Dordrecht 2016
Sacha C. Jones and Brian A. Stewart (eds.), Africa from MIS 6-2: Population Dynamics and Paleoenvironments,
Vertebrate Paleobiology and Paleoanthropology, DOI 10.1007/978-94-017-7520-5_14
1991; Moore and Blenkinsop 2006; Burke and Gunnell
2008; Clark et al. 2011). The mountains profoundly affect
climates and environments on either side by conning most
rainfall coastward and casting rain shadows far inland
(Wellington 1955). With relatively high precipitation and
low evaporation rates, their drainages are the subcontinents
major sources of fresh water, feeding all of southern Africas
perennial rivers (Clark et al. 2011). Within the mountain
zone itself, high geological, topographical and biological
diversity combine to produce microhabitats and high species
endemism (Kingdon 1990; Cowling and Hilton-Taylor
1994,1997; Davis et al. 1994; van Wyk and Smith 2001;
Carbutt and Edwards 2003,2006; Steenkamp et al. 2005;
Mucina and Rutherford 2006).
The peripheral mountainsstrong inuence on resource
distribution has major implications for prehistoric population
dynamics, something archeologists have long recognized
(e.g., Carter 1970,1976; Parkington 1972,1977,1980;
J. Deacon 1974; H. Deacon 1976,1979). Areas of broken
topography such as this are attractive to hunter-gatherers
because they provide enhanced resource diversity per unit
area of terrain and stable supplies of key resources
(Harpending and Davis 1977; J. Deacon 1974; Mitchell
1990). One well established pattern for Holocene
hunter-gatherer occupation of the mountains is to persist
and sometimes intensify during times of resource stress.
But their role in earlier prehistory including MIS 6-2 has
gone largely unexplored. These long isotope stages were
punctuated by numerous rapid climate change events that
would have placed premiums on areas where key resources
were comparatively stable. Surface water would have been
particularly vital during short-lived but acute arid pulses
with abrupt onsets (Partridge et al. 1993,2004; Scott 1999;
Holmgren et al. 2003). We expect that the well-watered,
ecologically diverse peripheral mountains often featured
prominently in human settlement decisions (cf. Mitchell
1990) through the instability of MIS 6-2.
To assess this, detailed paleoenvironmental frameworks
are needed from local terrestrial archives that are scaffolded
by robust chronologies. Previous efforts to reconstruct
pre-Holocene population dynamics anywhere on the sub-
continent have been impeded by a paucity of such archives
(Deacon and Thackeray 1984; Beaumont 1986; Mitchell
1990). Here we present a well-dated multi-proxy paleoen-
vironmental record from the sedimentary sequence at Meli-
kane, a rockshelter in the Maloti-Drakensberg Mountains of
highland Lesotho (Fig. 14.1). The Maloti-Drakensberg is the
highest and most widespread mountain system along the
Great Escarpment, and is the source of the subcontinents
largest river the Orange, known in Lesotho as the Senqu.
A new radiometric chronology for Melikane shows that the
sequence spans the last *83 ka (Stewart et al. 2012). The
site provides a record from MIS 5 onwards of foraging
behavior and regional environmental conditions in southern
Africas highest mountains. We analyzed plant opal phy-
toliths, δ
C of sediment organic matter (SOM) and arche-
ological charcoals. Taken in unison, these proxies provide a
comprehensive record of shifting environmental-vegetation
dynamics with important implications for understanding
early human dispersals into mountain systems.
Modeling Pleistocene Exploitation
of the Highlands
Determining when, why and how Pleistocene foragers
exploited highland Lesotho are key questions that have
remained unanswered since Patrick Carter pioneered the
areas archeology (Mitchell 2009; Stewart et al. 2012). For
the Holocene, Carter (1970,1976,1978) envisaged a system
of seasonal transhumance, with foragers spending summers
in the highland Maloti-Drakensberg and winters at lower
altitudes across the escarpment in the midlands of southern
KwaZulu-Natal and northeastern Eastern Cape (Fig. 14.1).
Some groups may have stayed entirely within the highlands
year-round, moving seasonally along the Senqu corridor and
its larger tributaries (Carter 1978). In contrast, Carter (1976)
suspected that highland environments in the Late Pleistocene
would have placed serious constraints on hunter-gatherers
that demanded more seasonally constrained exploitation
patterns, particularly during glacials and stadials when
temperatures, snowlines and sour alpine grasses were all
lower. During especially harsh climatic conditions, such as
the Last Glacial Maximum (LGM), people likely abandoned
the highlands altogether (Carter 1976).
Carters model of seasonal mobility, like others proposed
elsewhere along the Great Escarpment (Parkington 1972,
1977,1980; Deacon 1976,1979), met subsequent challenges
(Cable 1984; Mazel 1984; Opperman 1987) and no consensus
has yet emerged. In contrast, productive research has contin-
ued into the longer-term trends of Holocene population ux
(see review in Mitchell 2009). Whereas issues of taphonomy
and equinality still impede the precision necessary for
reconstructing seasonal movements, there is now a critical
mass of well-dated Holocene archeological and paleoenvi-
ronmental sequences to map broader paleodemographic pro-
cesses. Though much work remains before the same can be
achieved for MIS 6-2, the pace of research into the cultures and
environments of this period has recently accelerated. The
southeastern part of the subcontinent now boasts a compara-
tively dense concentration of sites that together cover much of
the Late Pleistocene (e.g., Deacon 1976; Carter 1978; Carter
et al. 1988; Mitchell 1988,1990,1992,1995,1996a,b,1998,
2008; Opperman 1988,1992,1996; Kaplan 1989,1990;
Opperman and Heydenrych 1990; Wadley 1997,2006;
248 B.A. Stewart et al.
Mitchell and Arthur 2010; Stewart et al. 2012; Roberts et al.
2013) (Fig. 14.1). We are now in a position to ask why during
specic periods of the Late Pleistocene lowland foragers chose
to increase their altitudinal range.
Ethnographic and ethnohistorical accounts of hunter-
gatherers engaging with mountain systems are rare (Alden-
derfer 1998). But a growing global corpus of archeological
data from such settings suggests that the motivations for, and
modes of, upland exploitation were myriad (e.g., Wright et al.
1980; Bender and Wright 1988; Bettinger 1991; Black 1991;
Benedict 1992; Madsen and Metcalf 2000; Walsh 2005;
Aldenderfer 2006; Brantingham 2006; Walsh et al. 2006;
Morgan 2009; Kornfeld et al. 2010; Morgan et al. 2012). The
conventional view that medium and high mountains were
strictly marginal habitats whose exploitation was a last resort
option by lowland groups facing climatic or environmental
deteriorationis ceding to models incorporating more real-
istic interplays of environmental and other (e.g., subsistence,
demographic, social) variables. Similarly, the reasons for the
Pleistocene exploitation of the Maloti-Drakensberg cannot be
reduced to a single overarching driver, but rather involved
diverse push and pull factors that varied through time and
space. However, the regions paucity of Pleistocene research
and deep antiquity of human settlement means that the
number and resolution of datasets remain too low to move
beyond climate-related inferences.
Elsewhere (Stewart et al. 2012) we have outlined the
general environmental conditions in which we expect a Late
Pleistocene human presence in the highlands to have inten-
sied. In the broadest sense, we predicted that this would
Fig. 14.1 Map of southeastern southern Africa with locales mentioned in the text. GRA: Grassridge; HM: Ha Makotoko; MEL: Melikane; NT:
Ntloana Tsoana; RCC: Rose Cottage Cave; SC: Sibudu Cave; SEH: Sehonghong; STB: Strathalan B; UMH: Umhlatuzana
14 Follow the Senqu: Mountain Dispersals 249
have coincided with phases when southeastern Africa was
relatively (1) warm and/or (2) arid. While higher tempera-
tures have obvious appeal to montane foragers, the latter
expectation is related to southern Africas uneven distribution
of freshwater. The subcontinent has extremely steep gradients
of rainfall and evaporation, with mean annual precipitation
decreasing from east to west and mean annual potential
evaporation increasing inversely (Lynch 2004; Schultz
2008). Roughly 60% of the subcontinents precipitation falls
on only 20% of its land surface and only *9% of annual
rainfall is converted into usable surface runoff or ground-
water recharge (Midgley et al. 1994). Precipitation and runoff
also vary markedly through time, both seasonally by rainfall
zone and inter-annually within rainfall zones. Wet and dry
years occur in cycles, with wet years heavily skewing the
mean (Eamus et al. 2006). The inter-annual variation in
runoff caused by this irregular rainfall is even more pro-
nounced (Schultz et al. 2001). Evaporation rates fall under
1600 mm only in the highland Maloti-Drakensberg and the
highest elevations of the southern Cape Fold Mountains
(Lynch 2004). With upwards of 2000 mm of mean annual
precipitation, the high uKhahlamba-Drakensberg escarpment
is thus one of the subcontinents only areas where evapora-
tion does not exceed precipitation. Moreover, areas with the
most precipitation and least evaporation are also those with
the lowest inter-annual variability of rainfall and runoff
(Schultz 2008). The freshwater resources generated by the
high escarpment are therefore some of southern Africas most
abundant and consistent.
This surplus runoff constitutes the bulk of the base ow
for a number of major rivers either side of the escarpment
watershed (Eamus et al. 2006) (Fig. 14.1). Draining the steep
eastern slopes are the Thukela, Mkhomazi, Mzimkhulu and
Mzimvubu. These rivers fall rapidly until they reach the
well-watered midlands of KwaZulu-Natal and Eastern Cape
where they slow and broaden (Rivers-Moore et al. 2007). All
runoff west of the escarpment contributes to the Orange
Senqu, which differs from these east-owing rivers in
important ways. The Senqu begins as high-altitude
(*3200 m a.s.l.) bogs and small fast owing streams in
the steep basalts of northeastern Lesotho, but soon slows and
widens as its bed cuts into the soft sandstones of south-
eastern Lesotho where Melikane is situated (Swanevelder
1981). The river then turns west to cross over 2000 km of
increasingly arid interior plateau into the Atlantic. Annual
rainfall falls rapidly from >1500 mm in the highland head-
waters to *300 mm in the central Karoo, then more grad-
ually to <50 mm at its hyper-arid mouth (Earle et al. 2005).
Before hydrological regulation began the late 19
the rivers middle and lower reaches experienced regular
droughts and some years ceased owing completely (Cam-
bray et al. 1986). Unlike the rivers owing east from the
escarpment, the Senqus upper catchment is thus vastly more
productive than any other part of the river basin. Despite
comprising only 5% of the basin, Lesotho receives over
twice its mean annual rainfall and supplies nearly half of the
rivers total streamow (Earle et al. 2005).
The abundance, stability and accessibility of the upper
Senqus surface water and associated resources would have
been attractive to foragers during pulses of heightened
regional aridity in MIS 6-2. We anticipate that at such times
the source populations for upland intensications derived
more often from the interior plateau than the better-watered
coastal forelands. The steep eastwest rainfall and evapora-
tion gradients that rendered interior groups susceptible to
aridity would have let them respond by dispersing east into the
mountains. The Orange-Senqu River Valley may have acted
as an important uvial corridor for populations moving
between the interior and the better-watered afromontane
landscape. In contrast, the nearness of the Eastern Cape and
Kwa-Zulu Natal to the subcontinents major source of pre-
cipitation the Indian Ocean means that even during dry
phases the forelands would have enjoyed greater humidity.
Groups living in these areas may have had less incentive than
inland populations to seek highland resources, and could
reorient their settlement strategies locally. During particularly
warm and/or humid phases, however, it is possible that some
foreland groups were pushed upland in response to population
growth. We thus posit a dual-source model of population
inux to the Maloti-Drakensberg contingent on prevailing
regional climate. We emphasize that this does not apply to
seasonal exploitation of the uplands (cf. Carter 1970,1978;
Cable 1984; Opperman 1987), but rather to the longer-term
patterns of intensication within which seasonal uses may
have occurred.
The Site and Research Area
Melikane is a large (44 ×21 m, 7.5 m high) rockshelter in
the Maloti-Drakensberg highlands of southeastern Lesotho
(Fig. 14.2). It is situated along the south side of the Melikane
River, a tributary of the Senqu, at an altitude of 1860 m a.s.l.
The Melikane River rises from the high Lesotho plateau
buttressing the uKhahlambaDrakensberg Escarpment. In
these upper reaches the river meanders through broad alpine
grassland valleys. But along its lower stretch, particularly
between the rockshelter and the Senqu conuence, the river
has cut a steep canyon-like ravine that offers a sheltered
environment for dense riverine scrub vegetation (Fig. 14.3).
The site was originally excavated in 1974 by Patrick
Carter (1978). In 2007, Carters trench was reopened to
obtain samples for a subcontinent-wide chronological study
250 B.A. Stewart et al.
using optically stimulated luminescence (OSL) (Jacobs and
Roberts 2008; Jacobs et al. 2008). The following year we
commenced new stratigraphic excavations at the site for an
ongoing project investigating the evolution of modern human
adaptive exibility (Dewar and Stewart 2012; Stewart et al.
2012). Thirty layers comprising four major depositional
modes were differentiated (Facies AD) (Stewart et al. 2012)
(Fig. 14.4). A vertical sediment column of bulk samples from
surface to bedrock was taken at 10 cm intervals; these form
the basis for the phytolith and SOM δ
C analyses
(Fig. 14.4). A new chronological framework based on
single-grain OSL and AMS
C shows that the sequence was
deposited in sporadic pulses *8380, *61, *50, *4638,
*24, *9 and *3 ka (Stewart et al. 2012) (Fig. 14.4). These
pulses appear to have been interspersed with long periods of
little or no sediment deposition or human occupation.
Landscape, Climate and Vegetation
The Maloti-Drakensberg Mountains extend for
*55,000 km
over most of Lesotho and adjacent parts of
South Africas KwaZulu-Natal and Eastern Cape Provinces
(Fig. 14.1). The tallest peaks exceed 3000 m a.s.l. Melikane is
one of numerous rockshelters in the Maloti-Drakensberg that
formed from the differential erosion of a suite of sandstones
(Clarens Formation) (Johnson et al. 1996; Schlüter 2006).
Deep uvial incision acting on these sandstones and over-
lying basalts created intricate mountain topography with
steep valleys and dramatic scarps. The most spectacular and
well-known escarpment is the uKhahlamba-Drakensberg,
which forms the eastern ank of the system and the inter-
national border between Lesotho and South Africa
(Fig. 14.1). However, the Maloti-Drakensberg is in fact
bounded on all sides by scarps, forming a well-dened
roughly rectangular mountain massif (Moore and Blenkinsop
2006). An important exception is the southwestern corner
through which the Senqu River exits the mountains via a
broad, deeply incised uvial corridor (Fig. 14.1).
Fig. 14.2 Views northeast from (a) and southwest into (b) Melikane Rockshelter
Fig. 14.3 A view west down the canyon-like, heavily wooded and
well sheltered lower Melikane River Valley
14 Follow the Senqu: Mountain Dispersals 251
The present-day climate and ecology of the
Maloti-Drakensberg varies widely by season, locale and
altitude. The overall climate is temperate, subhumid and
continental. Summers (October-March) are generally warm
and receive roughly three-quarters of the regions annual
rainfall (Tyson et al. 1976; Killick 1978). Winters (May
August) are cool to cold and extremely dry, though the
eastern highlands experience frequent snow from April to
September. Rainfall is highest along the uKhahlamba-
Drakensberg escarpments eastern slopes and summit,
where mean annual precipitation (MAP) can exceed
1600 mm (Killick 1963; Schulze 1979; Sene et al. 1998).
Temperatures vary drastically on both a seasonal and diurnal
basis, and by altitude. Mean annual temperatures range from
*15 °C in lowland Lesotho to 6°C in the high Drak-
ensberg (Grab 1994,1997). The lowlands produce mean
mid-summer maximum temperatures of 29 °C and
mid-winter minimum temperatures 4.3 °C, with respective
values for the highlands of 17 °C and 6.1 °C (Grab and
Nash 2010). Frost is widespread and ranges from *31 days
per year in the lowlands to *150 days per year in the
highlands (Schulze 2008). Ground freezing in the high
Maloti-Drakensberg is estimated to occur up to 200 days per
year (Grab 1997).
The Maloti-Drakensberg are lower in altitude but higher
in latitude than the mountain systems of tropical East and
northeast Africa, with comparatively cooler temperatures
reached at lower elevations. Consequently, vegetation dis-
tribution is mainly conditioned by altitude and aspect, vari-
ations in which produce an ecology with marked vertical
differentiation. Particularly signicant are altitudinal differ-
ences in the proportions of plant taxa following C
versus C
photosynthetic pathways. The dominance of C
or C
etation is primarily a function of temperature during the
growing season (Vogel et al. 1978; Ehleringer et al. 1997),
though precipitation and ambient CO
ratios are also
important (Ficken et al. 2002). In general, plants following
the C
pathway possess high tolerances to hot and dry
Fig. 14.4 The west prole of our excavation trench (left) and east prole of P. Carters excavation trench (right) at Melikane showing stratigraphy
and associated
C and OSL dates
252 B.A. Stewart et al.
climates with high irradiance and low atmospheric CO
) (Ehleringer et al. 1997; Retallack 2001). These
include tropical grasses, sedges and xeric herbs (OLeary
1981). C
plants are less CO
and water efcient, and thus
prefer cool, moist climates (Alexandre et al. 1997). They
include most woody taxa such as trees, herbs and shrubs, as
well as temperate grasses and sedges. As elsewhere in the
summer rainfall zone, the present-day Maloti-Drakensberg is
dominated by C
grasses with C
trees and shrubs at lower
altitudes and along the river corridors. However, the balance
shifts at high elevations (>2100 m a.s.l.) where C
have an adaptive advantage and outcompete C
The research areas major vegetation zones, from lowest to
highest altitude, are as follows. Along the boulder-strewn
Senqu River Valley and its many tributaries at *1600
1900 m a.s.l. is Senqu Montane Shrubland (Mucina and
Rutherford 2006). This is a Cymbopogon-Themeda-Era-
grostis grassland dominated by C
Panicoid grasses but with
some C
Chloridoid taxa also present. Important grass taxa
include Cymbopogon phrinodis,Heteropogon contortis,
Setaria abellate,Themeda trianda,Tristachya hispida (all C
Panicoids) and Eragrostis spp. (a C
Chloridoid). However,
numerous species of tree and evergreen shrub (all C
) also
thrive here (Bawden and Carroll 1968; Mucina and Rutherford
2006). These C
taxa are dominated by Rhus erosa,Olea
europaea and Diospyros austro-africana. Rarer thickets
dominated by Leucosidea sericea,Kiggelaria africana and
Rhamnus prinoides (all C
) occur in more sheltered valleys
(Mucina and Rutherford 2006). This zone is effectively an
eastern intrusion of lowland Lesotho taxa into the highlands
along the Senqu corridor (Jacot Guillarmod 1971). Cymbo-
pogon-Themeda-Eragrostis grassland also extends up onto the
gently rolling plateaux that overlook the deep river valleys,
though here trees and woody shrubs are comparatively rare.
Lesotho Highland Basalt Grassland occurs above the
river valleys and plateau shelves at altitudes of between
*19002900 m (Mucina and Rutherford 2006). This is a
dense, short C
Panicoid and Chloridoid grassland with
patchy shrublands dominated by Passerina montana
(Mucina and Rutherford 2006). Although some ericaceous
and composite taxa occur in the basalt grasslands, trees are
mostly absent. Due to its large altitudinal range (*1000 m),
this zone contains two altitude-specic vegetation belts. At
lower elevations, between *19002100 m on southern
(cooler) slopes but reaching up to *2700 m on northern
(warmer) slopes, grasses are dominated by C
notably Themeda triandra. This C
species is renowned for
its excellent pasturage quality. Other important C
taxa that
occur in this lower Themeda-dominated belt include
Andropogon spp. (a Panicoid), Eragrostis spp. and Micro-
chloa caffra (both Chloridoids).
In the upper belt of the basalt grasslands above
*2100 m on southern slopes and *2700 m on northern
slopes C
grasses become dominant. These Festuca-
Merxmuellera grasses are shorter and less palatable (sour
or letsiri in SeSotho) than Themeda.Festuca caprina is
dominant, but Festuca rubra and Festuca costata are also
important. Danthonia disticha,aC
Arundinoid, occurs on
thin, rocky soils. C
dominance continues at altitudes
exceeding 2900 m, although at these highest elevations the
basalt grasslands give way to Drakensberg Afroalpine
Heathland (Mucina and Rutherford 2006; termed Alpine
Heath by Killick 1990). This is a short heath and shrubland
dominated by Helichrysum trilineatum,Erica dominans and
Eumorphia sericea with dwarf bushes (all C
) (Killick 1978;
Mucina and Rutherford 2006). The heathlands are inter-
spersed with Merxmuellera-dominated C
grasses (Killick
1990). Embedded within this zone are patches of aquatic and
hygrophilous vegetation that include streambank communi-
ties and numerous alpine peat-forming bogs (Killick 1978,
1990). The latter occur either as basalt ssure seepage bogs
on mountain slopes or as thufur-covered sponge bogs at
riverheads that regulate headwater ow into Senqu River
system (Grab 1997; Jacot Guillarmod 1971; Killick 1978,
1990; van Zinderen Bakker and Werger 1974).
The Maloti-Drakensbergs alpine grass- and heathlands
are unique among southern Africas summer rainfall zones
as C
predominance is otherwise restricted to areas with
winter rainfall (the Western Cape) (Vogel et al. 1978). Such
close proximity of vegetation zones dominated by physio-
logically dissimilar taxa makes the region particularly well
suited to paleoenvironmental reconstruction. Because the
competitive balance between C
and C
is determined by
prevailing climate, shifts in the proportions of C
through time can be used to infer climatic changes acting to
alter local vegetation dynamics. In general, colder phases
encourage the down-slope expansion of C
taxa at the
expense of C
, whereas warmer phases provoke the reverse.
The result is a vertically migrating frontof C
nance, which registers in various proxies (e.g., phytoliths,
C signatures) at terrestrial archives including rockshelters.
In fact, the rst African rockshelter at which this principle
was applied was Melikane. In a seminal study, Vogel (1983)
investigated Late Pleistocene and Holocene paleoenviron-
ments using dietary δ
C composition of equid teeth from
Carters original excavation. His results were useful in
demonstrating the dominance during the Holocene and
Pleistocene of C
and C
taxa, respectively. However,
opportunities for obtaining better resolution were limited by
Carters coarse excavation methods and the limits of radio-
carbon dating. A series of more recent applications in the
Maloti-Drakensberg (Smith et al. 2002; Grab et al. 2005;
Parker et al. 2011; Roberts et al. 2013) and further aeld in
the highlands of East Africa (Ambrose and Sikes 1991;
Street-Perrott et al. 1997; Huang et al. 1999; Olago et al.
1999; Wooller et al. 2003) have focused almost exclusively
14 Follow the Senqu: Mountain Dispersals 253
on the Holocene/terminal Pleistocene. Since Vogels(1983)
pioneering study, therefore, very little research has been
conducted on deeper Pleistocene paleoenvironments in the
research area, and none using C
Materials and Methods
Recent work on phytolith morphotypes from modern vege-
tation in southern Africa has yielded new information for the
application of plant biogenic silica studies in paleoenviron-
mental reconstruction (Mercader et al. 2010; Cordova 2013).
These studies suggest that a number of morphotypes tradi-
tionally used to separate C
and C
vegetation, especially
grass silica short cell morphotypes, are found across a
number of C
and C
grass tribes. There is thus greater
redundancy in morphotypes than previously thought. Nev-
ertheless, some morphotypes and morphotype groups can be
attributed to particular grass subfamilies, sedges and woody
taxa. In particular, short cell short-saddle forms are attributed
to chloridoids, lobates to panicoids, papillae and achenes to
sedges, and circular rugose/globular granulates to dicot trees
and shrubs (Mercader et al. 2010,2013). It should be noted,
though, that Cordova (2013) designates some lobate forms
to C
grass subfamilies.
Subsamples were prepared for phytolith analysis using
the methods outlined in Parker et al. (2011). Ten grams of
sediment per sample were sieved through a 2 mm sieve in
order to remove the coarse fraction prior to phytolith
extraction. Carbonates were removed using 5% HCl fol-
lowed by 30% wv H
to remove organics. Samples were
deocculated using 50 ml 2% Calgon and 250 ml distilled
water, then shaken continuously for 30 min before being
passed through a 212 μm sieve. Heavy liquid separation
using sodium polytungstate (2.35 s.g.) was employed to
separate the phytoliths from the heavier inorganic residue.
The samples were diluted to a specic gravity of 1 and then
passed through a 5 μm vacuum ltration system (sensu
Theunissen 1994) to remove the clay and nest silt fractions.
Samples were mounted onto microscope slides using Canada
Balsam and identied at ×400 magnication using a Nikon
Eclipse E400 light microscope. Overall phytolith preserva-
tion was good, but in some samples phytoliths were either
entirely absent or had undergone dissolution rendering
viable counting impossible. The number of phytoliths
counted varied between 301 and 459 per sample.
Two phytolith indices were employed to help with data
interpretation: the D/P ratio and the climatic index (Ic%).
The D/P ratio is the ratio of ligneous dicotyledon morpho-
types (D) to Poaceae morphotypes (P), and is used as a
proxy of tree cover density (Alexandre et al. 1997). The
value 1 indicates maximum tree cover and zero none. The
climatic index (Ic%) indicates the inuence of climate on the
ratio of C
to C
morphotypes based on the proportion of
pooids to the sum of all short cell grass forms (Ic
% = pooid/pooid + chloridoid + panicoid) (Twiss 1987).
Higher Ic% values indicate more C
pooid grasses and thus
cooler conditions.
Sediment Organic Matter δ
Sediment samples were prepared for isotopic analysis at
Oxford Brookes Universitys Human Origins and Paleoen-
vironments (HOPE) Research Group laboratory following
procedures adapted from Ambrose and Norr (1993) and
Smith (1997). After sieving each sample through 2 mm and
2µm mesh, the sediment (*1 g) was pretreated with 2 M
HCl to remove carbonates. The analyses were performed at
the Godwin Laboratory at the University of Cambridge. The
samples were rinsed with deionized water, freeze-dried,
weighed into aluminum cups and measured using an auto-
mated elemental analyzer coupled in continuous-ow mode
to an isotope-ratio-monitoring mass spectrometer (Costech
elemental analyser coupled to a Finnigan Delta V mass
spectrometer). Stable carbon isotope concentrations were
measured relative to the VPDB international scale and are
reported as permil δvalues. The IAEA standard of caffeine
and in-house laboratory standards of nylon, alanine and
bovine liver were employed for calibration. Replicate anal-
yses of international and laboratory standards suggest that
measurement errors for δ
C are less than ±0.2. However,
sediments are likely to produce greater intra-sample uncer-
tainty due to high variance in organic content (Parker et al.
2011; Roberts et al. 2013). Each sediment sample was run in
It is well established that decreased temperatures during
the growing season promote C
vegetation. However,
Bousman (1991) demonstrated that a positive relationship
also exists between C
grass percentage and rainfall.
Working in the westernmost extension of the
uKhahlamba-Drakensberg Escarpment (the Kikvorsberg;
Fig. 14.1), he showed that as available moisture increases C
grasses outcompetes C
grasses and other C
species. He developed a formula for predicting the propor-
tion of C
grasses in the landscape based on the bulk δ
values. We employ a version of Bousmans Index that is
modied to account for anthropogenic inuences on SOM
C in the Melikane sequence. Bousmans formula allowed
for alteration of δ
C between plant and SOM of 3%, but we
have used a more conservative 2% following Wedin et al.
(1995): % of C
plants = (δ
C2 + 12.5)/0.14. These
254 B.A. Stewart et al.
values are checked against the phytolith Ic%, which provides
an independent estimation of C
ratios as noted above.
The entire charcoal assemblage from Melikane was rst
examined and quantied to isolate deposits containing
fragments suitable for analysis. The initial goal was to
analyze 200 fragments from each stratigraphic unit to gain
an overview of woody vegetation represented (cf. Allott
2004,2005,2006). However, preliminary inspection
revealed high inter-layer variability in charcoal abundance
and poor preservation of internal charcoal structure. This is
likely a result of repeated throughput of water originating
from ssures in the rear shelter wall at times both during and
after the accumulation of occupation debris (Stewart et al.
2012). Although wood charcoal is comparatively inert to
weathering processes, the effects of saturation and drying
can be detrimental to preservation. Water ow or saturation
introduces small particles of sediment into wood charcoal
that can obscure and damage internal anatomical features
used for identication. Most layers produced fewer than 200
fragments suitable for identication. Therefore, rather than
examining changes in the relative abundance of specic
taxa, we rely on habitat preferences and requirements in
order to make inferences about past regional climate and
vegetation conditions.
Five layers were selected that span the bulk of the Late
Pleistocene sequence: Layers 24 (*61 ka), 13 (*42 ka), 11
(*42 ka), 8 (*41.3 ka) and 5 (24 ka). Where possible more
than 100 charcoal fragments were identied from each layer
for a total of 445 fragments. Nevertheless, the majority of
layers produced very few charcoal fragments with only three
layers (13, 11 and 8) of the ve containing in excess of 100
fragments. Fragments were fractured along three planes to
reveal transverse, tangential longitudinal and radial longi-
tudinal sections (Leney and Casteel 1975). Characteristics of
anatomical features visible in each were recorded with ref-
erence to the IAWA list of microscopic features for hard-
wood (Wheeler et al. 1989). Taxonomic identications have
been provided through comparison with modern reference
material developed at the University of the Witwatersrand
(Dowson 1988; Wadley et al. 1992; Esterhuysen 1996;
Allott 2004,2005) and reference atlases/resources (Kromh-
out 1975; Eichhorn 2002; Inside wood 2004 onwards).
Where possible, fragments were identied to family, genus
or species. Secure species identications from wood
anatomical features alone are rarely possible, though when a
genus is represented by a single species within the region it
has been named as the most likely origin of the charcoal.
Family identications are given when similarities between
genera within a family are too great for differentiation or
when anatomical features are not sufciently clear to rene
the identication.
The results of the phytolith results are presented in Figs. 14.5
and 14.6. The former shows all phytolith morphotypes
organized according to depth from surface, while the latter
presents phytolith family summaries chronologically with
ages of samples from undated layers interpolated using
depth. The SOM δ
C results are presented in Table 14.1
and Fig. 14.6, where they are presented graphically along-
side the summary phytolith results. Table 14.2 presents the
results of the charcoal analyses, while Table 14.3 gives
habitat preferences of the woody taxa represented.
Several general observations can be made. First, both the
phytolith and SOM δ
C results clearly show that the entire
Late Pleistocene portion of the Melikane sequence (Layers 3
and below) is heavily dominated by C
grassland taxa (Ic
6095%) (Fig. 14.6). Only with the onset of the Holocene
(Layer 2) do phytolith C
indicators outnumber C
(Ic 40%).
This is consistent with Vogels(1983) original ndings at
Melikane and recent SOM δ
C data from the Lesotho
lowlands (Roberts et al. 2013), and reafrms that Late
Pleistocene temperature depressions provoked the lowering
of C
alpine grasses. Considering that today C
grasses reach
up to 2700 m on north-facing slopes, and assuming a tem-
perature drop of 0.6 °C/100 m (Smith et al. 2002), this
suggests that temperatures were at least 5 °C cooler
throughout the Late Pleistocene at Melikane, a north-facing
shelter at an altitude of 1860 m a.s.l.. Second, with the
exception of the uppermost part of the sequence there is
good agreement between the phytolith and SOM δ
records, which show a close correspondence between the Ic
% values and the Bousman Index values (Fig. 14.6). This
gives us condence that these records reect changes in
landscape vegetation dynamics rather than purely stochastic
anthropogenic inputs (although the selective collection of
vegetation will be reected heavily in the charcoal record).
Third, there are differences between the records in the
magnitude of changes. The phytoliths and to a lesser extent
the SOM δ
C show variability through time, with changes
apparent within the C
dominated signals. In contrast, the
shrubs and trees identied in the wood charcoal assemblage
show a degree of continuity (Table 14.2). This may indicate
either overall stability of local vegetation or that humans
preferentially and consistently selected these woods for fuel
through time. Since shifts within the phytolith record make
vegetation stability unlikely, we favor selection continuity as
a more probable interpretation.
14 Follow the Senqu: Mountain Dispersals 255
MIS 5a
One sediment sample MLK 22 (*83 ka; Layer 29) was
taken from Melikanes deepest strata deposited during MIS
5a. The δ
C value for this sample is 23.42(Table 14.1).
This is a strongly C
signal, with the resulting Bousman
Index value suggesting >93% of the signal is derived from
vegetation with minor but extant C
input. This is sup-
ported by the phytoliths, which show a dominance of pooid
short cells and an Ic% value of >95% (Fig. 14.6). Only a
trace of panicoid types and no short-saddle chloridoids were
present. Also present in low abundances are papillae and
achene phytoliths from sedges. The tree cover density index
(D/P ratio), which indicates the proportion of woody taxa to
grasses, is low at *0.1 (Fig. 14.6). This suggests that this C
grass-dominated landscape hosted a low woodland element.
A suitable charcoal sample for MIS 5a could not be
MIS 4 is represented by three samples: MLK21 (*73 ka
extrapolated; Layer 26 lower), MLK20 (*67 ka extrapo-
lated; Layer 26 upper) and MLK19 (*61 ka). The oldest
MIS 4 sample (MLK21) registers a very slight positive shift
to 23.19(from 23.42in late MIS 5a) (Table 14.1),
with correspondingly small negative shifts in the Bousman
Index (*90%) and phytolith Ic% (*90 %) (Fig. 14.6).
There is then a 1.8positive shift in δ
C values to
21.41in MLK20, followed by a slight negative shift in
MLK19 back to 22.46. The corresponding Bousman
Index value for MLK20 suggests a shift in the vegetation
composition with an increase in the C
component to
*30%. The phytoliths corroborate this with Ic% values of
*70% (Fig. 14.6) The short cell grassland phytoliths in
MLK20 account for 20% of the assemblage. Panicoid
lobates increase slightly to *5% when compared with MIS
5a, along with a trace of short body saddle chloridoids.
Chloridoid phytoliths are absent from the youngest MIS 4
sample (MLK19). The presence of sedges was also noted but
in low numbers. There is an increase in ligneous (woody)
dicot phytoliths when compared with MIS 5a to *7% in
MLK20 and 5% in MLK19. The D/P ratio in MLK19 is also
higher than for MIS 5a, rising to 0.2 (*20% woody vege-
tation) (Fig. 14.6). This suggests more trees were present in
the landscape and/or that a greater proportion of woody taxa
were being deliberately brought into the site. However, the
overall proportion of woody phytoliths remains low.
A small but informative charcoal sample was obtained
from the youngest MIS 4 sample MLK19 (Layer 24). Four
taxa were recorded: Buddleja cf. salviifolia;Rhamnus sp.;
Leucosidea sericea; and Protea sp. (Table 14.2). Of these,
Table 14.1 SOM δ
C data for the Melikane samples showing depth from surface, mean, standard deviation and percentage of elemental carbon
for each set of triplicate samples
Sample Depth from surface (m) Mean δ
C() SD () C(%)
MLK1 0.1 ––
MLK2 0.2 24.01 0.05 0.71
MLK3 0.3 23.19 0.07 0.27
MLK4 0.4 22.48 0.04 2.84
MLK5 0.5 23.12 0.13 5.78
MLK6 0.6 22.91 0.09 2.73
MLK7 0.7 22.71 0.06 3.34
MLK8 0.8 22.58 0.10 5.69
MLK9 0.9 23.00 0.11 0.55
MLK10 1 22.77 0.08 0.47
MLK11 1.1 23.89 0.01 3.08
MLK12 1.2 23.41 0.06 1.63
MLK13 1.3 23.68 0.05 0.96
MLK14 1.4 23.43 0.04 1.97
MLK15 1.5 22.88 0.08 0.97
MLK16 1.6 23.42 0.12 0.48
MLK17 1.7 22.92 0.05 0.83
MLK18 1.8 23.41 0.10 1.46
MLK19 1.9 22.46 0.04 1.38
MLK20 2 21.41 0.04 4.90
MLK21 2.1 23.19 0.03 1.93
MLK22 2.2 23.42 0.01 0.71
256 B.A. Stewart et al.
Fig. 14.5 All phytolith morphotypes (arranged by depth from surface)
14 Follow the Senqu: Mountain Dispersals 257
Fig. 14.6 Summaries of phytolith families and morphotypes, and the phytolith D/P and Ic indices juxtaposed with the SOM δ
C results (C%, δ
C and Bousman Index). These results are arranged
chronologically with ages for undated layers interpolated by depth. Vostok data from Jouzel et al. (2007)
258 B.A. Stewart et al.
Table 14.2 Taxonomic identications of the charcoal samples selected for analysis by layer. Details on relevant
C and OSL dates for these layers are also shown
Layer C14 Date
(cal BP)
OSL Date (BP) Buddleja
Rosaceae Olea
5 24,20023,600 27,100 ±1800 1 14 10
5 24,20023,600 27,100 ±1800 1
5 24,20023,600 27,100 ±1800 11
8 41,76541,097
41,300 ±3000 32 49 19 3
11 Bracketed between
(Layer 14) and
(Layer 9)
Bracketed between 41,300
±3,000 (Layer 8) and
45,900 ±3,800 (Layer 14)
25 29 1 23 4 1 4
11 Bracketed between
(Layer 14) and
(Layer 9)
Bracketed between 41,300
±3,000 (Layer 8) and
45,900 ±3,800 (Layer 14)
14 15 47
13 Bracketed between
(Layer 14) and
(Layer 9)
Bracketed between 41,300
±3,000 (Layer 8) and
45,900 ±3,800 (Layer 14)
52 19 10 13 1 1 2 3
24 No 61,000 ±2500 23 1 3 14
14 Follow the Senqu: Mountain Dispersals 259
both Buddleja cf. salviifolia and Rhamnus sp. favor ever-
green forest margins or grow along watercourses
(Table 14.3). The Rhamnus specimens, although unidenti-
able to species, are almost certainly Rhamnus prinoides,
which is common in present-day afromontane forest settings
(including Lesotho) at medium and high altitudes. Buddleja
cf. salviifolia and Rhamnus sp. are found in similar habitats
to Leucosidea sericea, which is a pioneer species of higher
altitude (1000 m) mountainsides occurring along running
streams (Table 14.3). All three taxa prefer relatively humid
environments, consistent with the phytolith and δ
C results
for this later MIS 4 sample. The fourth taxon, Protea sp.,
tends to grow on slightly drier, rocky ground or mountain
grassland, though it too prefers sheltered locations where
moisture is available.
A*10 kyr hiatus in deposition occurred between MIS 4
(MLK19) and MIS 3 (MLK18). The MIS 3 strata at Meli-
kane (Layers 22-6) span from *50 to 38 ka and appear to
have been deposited two broad pulses: *50 ka and *46
38 ka. Thirteen samples were examined, four from the for-
mer pulse (MLK1815) and nine from the latter (MLK14-6).
Isotopic values *50 ka (MLK 18-15) range between
23.42 and 22.88(Table 14.1). The Bousman Index and
Ic% values show little variation in the proportion of C
(>90%) to C
(<10%). However, an increasing proportion of
the isotope C
signal likely derives from tree and shrub
elements of the vegetation. The phytolith record from the
lower three samples (MLK18-16) shows an increase D/P
from 0.2 to 0.3, indicating a greater proportion of woody
phytolith morphotypes (30%) to grassland (70%) (Fig. 14.6).
These values are the highest shown in the entire sequence
and suggest an increase in the regional woodland component
and/or an increase in the selection of woody material that
was brought into the site. The phytolith and SOM δ
values suggest a greater cover of woodland than in MIS 4,
with C
taxa dominating the grassland ora under cool,
wet/humid conditions. The uppermost sample (MLK15) in
the older MIS 3 sediments contained very poorly preserved
phytoliths with major dissolution noted (Fig. 14.5). Its iso-
topic value (22.88) gives some insight into a C
nated landscape with a C
component. Panicoid levels are
lower than those in MIS 4, while chloridoid phytoliths are
present but in trace amounts (Fig. 14.6).
Between *4638 ka (MLK14-6) the SOM δ
C values
range between 23.89 and 22.56(Table 14.1). The
values become more depleted from MLK14 to MLK11 to
reach the lowest values recorded in the Melikane sequence
Table 14.3 Habitat preferences of the woody taxa identied in the charcoal assemblage
Identied Taxa Habitat Preferences Known
Medium to tall trees
Occurs in evergreen forest, in riverine forest and sometimes in drier coastal and mountain forests
Protea sp. Drier, rocky ground or mountain grassland, sheltered locations with available moisture Y
Tall Shrubs or small trees
Buddleja cf.
Occurs at the margins of or in evergreen forest, on rocky mountain slopes and along water courses at
higher altitudes
Rhamnus sp. Widespread and locally common at med-high alt, along watercourses, in riverine forest and at margins of
evergreen forest
Occurs over wide range of altitudes, often fringing evergreen forest, also found in wooded ravines, on
hillsides and rocky outcrops
Olea europea Variety of habitats usually near water, stream banks, riverine fringes also open woodland, among rocks and
in mountain ravines
Grewia sp. More typical of open grassland vegetation
Small Shrubs
Erica sp. Mountain ravines, rocky grassy slopes, moist places, sometimes near streams Y
In moist places, forest margins, rocky grassy slopes, up to 1800 m
Pioneer species of mountainsides, kloofs, valley bottoms and along streams at higher altitudes, often
occurs in dense stands of multistemmed, somewhat straggling shrubs
Gymnosporia sp. Occurs in rocky places in grassland associated with forest Y
Other taxa Identied to family
Rosaceae Includes a broad range of potential taxa
cf. Fabaceae Includes a broad range of potential taxa
260 B.A. Stewart et al.
(23.89). Values uctuate after this and become pro-
gressively more enriched, with a +1.31shift by MLK8
(Layer 9). Phytolith preservation was extremely variable in
these samples with a near complete absence of biogenic
silica in MLK14, 12 and 11 (Fig. 14.5). This may result from
the complete dissolution of phytoliths or the sediment being
constituted of decomposed and weathered roof collapse
material. Phytoliths show an increase in panicoid lobate
morphotypes (up to 20%) with a small trace of chloridoids.
The Bousman Index values show high C
values (9095%).
The Ic% shows close correlation up to MLK8 where it
diverges (55%) from Bousmans value (90%) (Fig. 14.6).
The most likely explanation for the difference in the two
values is an enhanced C
component in the isotope record
being derived from woody taxa collected for rewood. The
D/P values fall considerably over this time period to levels
<0.05 between *42 and 38 ka, suggesting major reductions
in regional woodland.
Suitable charcoal samples were obtained from the
younger MIS 3 pulses (*4638 ka). Charcoals from
MLK13 (Layer 14) and MLK11 (Layer 11) both *43
42 ka include a broader range of taxa (Table 14.2). In
addition to those mentioned from MIS 4, there is evidence
for further evergreen trees and shrubs that grow in riverine
forests. These include species such as Rapanea mela-
nophloeos, Olea europaea (Layer 13) and Heteromorpha
sp. (Layer 11), as well as several that prefer more open
grassland vegetation such as Grewia sp. and Fabaceae taxa
(Layer 13) (Table 14.3). Later, at *41 ka, there is continued
evidence for evergreen, riverine forest taxa in Layer 8
(MLK7), but also Erica sp., a genus that includes species
which are common components of high-altitude vegetation
in the Maloti-Drakensberg (Mucina and Rutherford 2006)
(Table 14.3). Shrubs within this large genus provide excel-
lent fuel and were likely targeted for rewood when they
became more accessible.
MIS 2 is represented by two samples MLK5 and 4 from
Layer 5 (*24 ka). δ
C values range from 23.12
(MLK5) to 22.48(MLK4) (Table 14.1). Both are still
strong C
pooid grassland signals, but with a slight increase
in the C
component. The Bousman Index values indicate
*8085% C
versus 2015% C
. The phytolith record
likewise shows an increase in chloridoid (5%) and panicoid
forms (*15%), along with sedges (*1%) (Fig. 14.6).
Woody taxa phytolith values drop to the lowest levels in the
sequence, accounting for <1% of the total sum. The D/P
values are also the lowest from the sequence at <0.1
(Fig. 14.6). Given the low D/P values it is suggested that the
component from woody taxa in the region is low.
However, as in late MIS 3, the presence of wood charcoal
may suggest that collected rewood has biased the C
slightly, enhancing it over a background grassland with an
increased C
The charcoal record from MIS 2 (Layer 5) differs slightly
from MIS 4 and 3. Buddleja cf. salviifolia and Rhamnus sp.,
which have been recorded throughout the earlier deposits,
are absent (Table 14.2). There is continued evidence for
Erica sp., Leucosidea sericea, Protea sp. and Olea europea.
With the exception of Protea sp., which is more indicative of
open scrub, each of these are common components of
riverine vegetation (Table 14.3). Although it is difcult to
interpret the absence of the taxa noted in the earlier layers, it
is interesting that some of those that do remain are frost
tolerant and pioneer species. All of the taxa noted in this
layer provide good sources of fuel.
Regional Paleoenvironments
and Population Flux
The Melikane sequence shows that humans were recurrently
visiting the broken topography of the Maloti-Drakensberg
highlands from at least MIS 5a. It is important to note,
however, that these were not the rst people to venture into
these uplands. A handful of handaxes and cleavers from
undated open-air contexts suggest a Middle Pleistocene
hominin presence, although this was extremely ephemeral
(Carter 1978). The phytolith and SOM δ
C data suggest
that highland environment *8380 ka was relatively stable,
humid and cool. The landscape around the site was a C
dominated grassland environment with some C
most likely derived from tall, hydric, heliophyte panicoid
grasses. Some woody elements from dicotyledonous trees
and shrubs were also present, but in low abundance and most
likely restricted to river corridors. The rockshelter itself also
appears to have been relatively stable at this time. Previously
published geoarchaeological data (Stewart et al. 2012)
indicate a low-energy rockshelter environment evidenced by
well-stratied sediments and sharp bounding surfaces
recorded in this part of the prole. The ssures present at the
rear of the rockshelter that today allow water ingress had not
developed at this time, and the interior of the site was a dry,
inactive environment well protected from mechanisms of
erosion related to water inux.
Paleoenvironmental conditions in wider southeastern
southern Africa for late MIS 5 remain poorly resolved. At
present the nearest continuous paleoenvironmental archive
to Melikane is the newly published Indian Ocean marine
14 Follow the Senqu: Mountain Dispersals 261
core CD154-17-17 K off the Eastern Capes east coast
(Ziegler et al. 2013) (Fig. 14.1). This record, which spans the
last *100 kyr, suggests that sea surface temperatures (SSTs)
in the southwestern Indian Ocean during late MIS 5 were
warmer than subsequent MIS 4-2, but cooler than today.
This is consistent with two other marine cores (RC17-69 and
MD 96-2077) extracted slightly further north off southern
KwaZulu-Natal (Prell and Hutson 1979; Bard and Rickaby
2009). Core CD154-17-17 K also shows that discharge from
rivers draining the southeastern escarpment and adjacent
forelands during late MIS 5 were highly variable. Sub-
stages MIS 5c and b appear to have been relatively moist
with several abrupt humid episodes registered by pulses of
uvial discharge. But at the outset of MIS 5a *83 ka there
is a shift to drier conditions that appear to persist for several
millennia (Ziegler et al. 2013).
This peak aridity event *8380 ka corresponds to the
earliest occupational pulse at Melikane. It is unclear what
caused this dryness as warm southwestern Indian Ocean
SSTs typically induce greater humidity across southeastern
Africa (Reason and Mulenga 1999; Chase 2010; Dupont
et al. 2011). However, this relationship is complex and at
times warm anomalies in the western Indian Ocean correlate
strongly with reduced (rather than increased) summer rain-
fall in southern Africa (Mason and Jury 1997; Landman and
Mason 1999). Similarly, Thackeray (1987,1988) has
demonstrated that no linear relationship exists between
temperature and rainfall in southern Africa. It is possible, for
example, that conditions *8380 ka were akin to the
mid-Holocene, when much of the subcontinent was simul-
taneously very warm and arid. A number of inland records
situated further north in the Kalahari Basin also register
lower precipitation *80 ka, including various cave spe-
leothems in the Kalahari that cease growing *8377 (Brook
et al. 1996,1997,1998). If areas both inland and coastward
of the Maloti-Drakensberg were arid, this contrasts with the
highland zone around Melikane, which was humid C
dominated grassland with a minor woody component. This
may suggest that some lowland populations were drawn into
the highlands to exploit this better-watered zone as a tem-
porary refugium. Clearly, though, more terrestrial archives
and archeological records are needed to develop a sharper
picture of regional ecology and population dynamics during
late MIS 5.
Melikane appears to have been largely or entirely aban-
doned over the next *20 kyr, an interval encompassing
terminal MIS 5 and most of MIS 4. Core CD154-17-17 K
suggests that after the dry phase *8380 ka the region
became increasingly humid in terminal MIS 5, with pulses of
heavy uvial discharge into the Indian Ocean towards the
transition to MIS 4 (Ziegler et al. 2013). Humidity remained
high throughout MIS 4, though with fewer peak periods of
increased discharge and sediment deposition than terminal
MIS 5. High moisture availability is consistent with the
relatively high SSTs estimated for MIS 4 in core MD79254
off Mozambique (van Campo et al. 1990; Chase 2010),
though more southerly cores closer to the research area give
colder SST estimates for this period (Prell and Hutson 1979;
Bard and Rickaby 2009). Still, the signature of high MIS 4
humidity in core CD154-17-17 K agrees well with the ter-
restrial archive from Sibudu (Fig. 14.1), where multiple
proxies suggest that during the Howiesons Poort occupations
in late MIS 4 (*6562 ka) the area hosted a woodland
savanna crosscut by riparian evergreen forests (Allott 2006;
Sievers 2006). Whereas winters appear to have been slightly
drier and colder than today, summer temperatures and pre-
cipitation levels were comparable (Bruch et al. 2012).
Melikanes abandonment *8061 ka need not imply
total depopulation of the highlands, but there is little to
suggest a human presence in the area at this time. This
interval saw two highly distinctive stone tool industries the
Still Bay and the Howiesons Poort appear throughout the
subcontinents coastal forelands (Henshilwood 2012; Sealy
2016). Yet no characteristic Still Bay bifacial points have
been found in the Maloti-Drakensberg despite several
intensive eld surveys (Carter 1978; Parkington and
Poggenpoel 1980; Bousman 1988; Mitchell 1996a,b;
Mitchell and Arthur 2010; Dewar and Stewart 2011), and the
only Howiesons Poort backed artifacts discovered to date are
those at Melikane and several lithic scatters noted by Carter
below the Eastern Cape Drakensberg near Kenegha Poort
(Mitchell 2009:130). According to Antarctic ice cores, the
bulk of MIS 4 saw depressed temperatures with the coldest
phase between *70 and *62 ka when mean temperature
was 8.7 °C below present (Jouzel et al. 2007). Such severe
cold earlier in MIS 4 probably discouraged highland
exploitation, and indeed this upland hiatus contrasts with
lower elevation sites either side of the Maloti-Drakensberg
such as Sibudu and Rose Cottage Cave (Fig. 14.1) where
humans were present *7062 ka (Jacobs et al. 2008;
Pienaar et al. 2008).
When Melikane was reoccupied at *61 ka, conditions
may have warmed sufciently for Howiesons Poort
tool-makers to make higher altitude incursions. The phy-
toliths show that the highland landscape at this time was still
dominated by grasses, but that temperatures and the pro-
portion of woodland had increased. The charcoal record
suggests that the latter included evergreen forest along the
riverine corridors, with pioneer, open ground woody vege-
tation also present. The SOM δ
C values and the Bousman
Index values likewise suggest the local vegetation signal was
still dominated by the C
component. This would have lar-
gely been sourced from pooid grass taxa, though an element
will have also been derived from C
tree and shrub taxa.
However, there was now more C
than was present in MIS
5a. The increase in the C
component (up to 30%) is
262 B.A. Stewart et al.
corroborated by the increase in morphotypes attributed to
panicoids along with the trace chloridoid signal (in MLK19).
More C
taxa suggests slightly warmer temperatures, a shift
in the vegetation belt with slightly drier conditions relative to
MIS 5a, or lowered pCO
levels (Street-Perrott et al. 1997).
We consider the latter more likely considering the higher
D/P ratio, the presence of charcoal from water-loving taxa
and evidence for widespread humidity across southern
Africa during MIS 4 (Chase 2010). The Melikane River
Valley directly below the site would have been a good
source of wood for fuel and other uses (Fig. 14.3).
Melikane witnessed a further 10 kyr period of abandon-
ment in terminal MIS 4 and early MIS 3 until *50 ka. There
is abundant evidence that the wider region experienced ini-
tial cold followed by increasingly warm and dry conditions
during this hiatus. Fluvial discharge in core CD154-17-17 K
diminishes markedly relative to MIS 4 and terminal MIS 5
(Ziegler et al. 2013). Though details vary, marine cores for
which SSTs have been estimated register a drop in temper-
atures at some point between *6050 ka (van Campo et al.
1990; Bard and Rickaby 2009). At Sibudu, makers of
post-Howiesons Poort (Sibudan) technology intensively
occupied the site across the MIS 4/3 transition around 58 ka.
Diverse proxy datasets suggest a riverine forest setting that
included frost tolerant afromontane vegetation signaling the
coldest conditions in the entire Sibudu sequence. Also
making a rst appearance are several bushveld taxa, which
today grow in more northerly, drier regions of the subcon-
tinent. Gradual warming and further drying through this
pulse at Sibudu is evidenced by the replacement of small,
solitary browsing ungulates by medium and large grazers
(Clark and Plug 2008), increasing frequencies of seeds from
deciduous trees (Sievers 2006) and higher magnetic sus-
ceptibility values for the upper sediments (Herries 2006).
Widespread and well-dated colluvial mantles (the Masotch-
eni Formation) deposited across inland KwaZulu-Natal
during phases of hillslope instability *5652 ka (Botha
and Partridge 2000) may indicate that this drying trend
continued after and might have played a role in Sibudus
Though these are the conditions (relatively warm and dry)
under which we expect highland settlement to intensify,
Melikane was not reoccupied until *50 ka. Humans were
present, however, at the nearby highland site of Sehonghong
at *58 ka, and at *56 ka at both Ntloana Tsoana and Rose
Cottage Cave farther west in lowland Lesotho and the
adjacent eastern Free State, respectively (Jacobs et al. 2008).
Moreover, it is important to note that virtually no rockshelter
sites anywhere in southern Africa have dated occupations
between *56 and *50 ka. One possibility is that settlement
preferences shifted away from rockshelters towards open-air
locales, perhaps in response to warmer temperatures and
reduced precipitation. Open-air sites with Late Pleistocene
artifacts abound in the highlands (Carter 1978; Bousman
1988; Mitchell 1996a,b; Dewar and Stewart 2011), but until
these can be dated the lack of highland occupation during
this interval this does not t model predictions.
Melikane was reoccupied in mid-MIS 3, rst at *50 ka
and later *4638 ka. These were the sites most enduring
human occupations, and the phytolith and SOM δ
records signal dramatic changes in the highland environ-
ment. The earlier pulse *50 ka appears to have been war-
mer and wetter than either MIS 5a or 4, supporting the
highest tree/shrub densities of any period registered at
Melikane. Higher temperatures, increased summer rainfall
and greater vegetation density are also inferred for the
broadly coeval (*48 ka) late MSA levels at Sibudu as
compared to earlier periods (Glenny 2006; Wadley 2006;
Bruch et al. 2012). The interior likewise appears to have
enjoyed higher precipitation at *50 ka, with rainfall levels
exceeding present day. Widespread warming and humidi-
cation may have encouraged exploitation of the highlands by
forelands groups, who introduced a Sibudu late MSA-like
lithic technology that included convergent Levallois points.
Very different conditions ensued at Melikane in the
longer pulse between *46 and 38 ka. The phytoliths indi-
cate a reduction in woodland and an expansion of grassland.
The increase in panicoids suggests more open conditions
with tall, heliophyte grasses. These may have been more
abundant due to slightly drier conditions and/or a lowering
of pCO
levels during mid-MIS 3. The charcoal record
shows evidence for evergreen trees and shrubs associated
with riverine forests, along with taxa that prefer more open
grassland conditions including Grewia sp. and Fabaceae
(Tables 14.2 and 14.3). From *43 ka woodland cover
diminishes further, with the absence of some evergreen
moist forest types and the appearance of taxa that may
suggest a depression in the alpine belt due to colder condi-
tions (Erica sp.). Reduced climatic stability is consistent
with evidence for major changes in site formation processes
at this time (Stewart et al. 2012). These include recurrent
inuxes of colluvial gravels alternating with episodes of roof
collapse, and substantial physical and chemical transforma-
tions of the sediments (including phytolith dissolution) from
increased water throughput via newly formed ssures.
This record of mid-MIS 3 instability accords well with
core CD154-17-17 K, which shows a transition at *50 ka to
a period of wetter pulses punctuated by drier episodes for the
remainder of MIS 3 (Ziegler et al. 2013). The timing of these
wet events in southern Africa has been linked to bipolar
responses to North Atlantic cold episodes (Heinrich Events),
with corresponding increases of uvial discharge into the
Indian Ocean (Ziegler et a. 2013). Partridge et al. (2004)
have shown that each Heinrich Event is preceded by *3
4 kyr of rapid onset arid phases linked to Antarctic cooling.
In KwaZulu-Natals Masotcheni Formation, these changes in
14 Follow the Senqu: Mountain Dispersals 263
aridity and wetness are reected in alternating phases of
colluviation and pedogenesis resulting from phases of hill-
slope activity and stability, respectively (Botha and Partridge
2000). Colluvial episodes signaling heightened aridity have
been OSL and
C dated to *4637 ka (Botha and Partridge
2000; Botha et al. 1992; Clarke et al. 2003; Temme et al.
2008). These are punctuated by numerous paleosols
reecting brief shifts to more humid conditions. The
Masotcheni sediments also appear to register the drop in
temperatures at *42 ka that is apparent at Melikane
(Temme et al. 2008). On the interior plateau there is likewise
evidence that this period was unstable if predominantly arid.
A recent multi-proxy analysis of a thoroughly dated (using
OSL) overbank alluvial-paleosol succession at Erkroon near
Bloemfontein indicate conditions between *46 and *32 ka
were more arid (estimated MAP: *200400 mm) than
present-day (MAP *400600 mm), with a brief phase of
intervening humidity at *42 ka (Lyons et al. 2014). This
agrees well with
C dated portion of the inland record at
Tswaing Crater further to the north, where a major reduction
in rainfall is apparent beginning *47 ka, followed by slight
humidication *42 ka and then another arid phase *40
35 ka (Kristen et al. 2007).
This period of instability and recurrent aridity corresponds
closely with Melikanes most intensive pulse of human
occupation *4638 ka. During this 8 kyr interval over a
third of the stratigraphic sequence was deposited. Recent
work at nearby Sehonghong (Fig. 14.1) suggests that a sub-
stantial portion of that sequence also dates to this time
(Jacobs et al. 2008). The concurrence between this intensi-
cation of highland exploitation and climatic instability in the
wider region is striking. We suggest that this could reect
dispersals of lowland populations towards the upper reaches
of the Senqu River in order to buffer against recurrent
shortfalls of accessible surface water and dependent plants
and animals. Preliminary support for this hypothesis comes
from the paleoenvironmental proxy results presented here.
While phytoliths and SOM δ
C often reect vegetation
changes in the broader landscape, charcoals are more rmly
tied to human selective processes and provide windows on
locally available fuel woods. Unlike the phytoliths and SOM
C which both show environmental changes, the charcoals
exhibit a high degree of continuity through the sequence. As
noted, most of the taxa represented are evergreen shrubs and
small trees that favor wooded ravines often in association
with watercourses. Ravines and crevices provide sheltered
locations with increased available atmospheric moisture, and
the presence of streams may have had an additional amelio-
rating effect enabling vegetation to remain at altitudes and in
climatic conditions that were otherwise unsuitable. The
Senqu and Melikane River Valleys could have provided such
a refuge, offering people reliable sources of surface water,
food, fuel and high-quality toolstone. Importantly, these
riparian charcoals continue even after 42 ka when the phy-
toliths and SOM δ
C suggest reduced temperatures, woody
vegetation and rainfall. The upper Senqu and its tributaries
like the Melikane thus appear to have sustained ample water
ow through this unpredictable time.
Humans were again absent from Melikane in late MIS 3
and earliest MIS 2 between *38 and 24 ka. After a pro-
nounced peak of river discharge *38 ka, core
CD154-17-17 K suggests slightly diminished variability in
this interval compared to *5038 ka (Ziegler et al. 2013).
The peak at 38 ka coincides with the termination of the
protracted mid-MIS 3 occupation at Melikane and a brief
occupational pulse at Sibudu by makers of nal MSA tech-
nology. A range of studies suggest that Sibudus setting at
this time was a complex mosaic of open savanna/woodland
with patches of riparian forest similar to today, albeit slightly
drier and cooler (Allott 2006; Glenny 2006; Bruch et al.
2012). At Erfkroon in the eastern interior, the aforementioned
arid phase *4632 was followed by humid conditions *32
28 ka, after which increasingly arid conditions again set in on
the lead up to the LGM. However, currently very little
additional paleoecological information is available for this
interval, though our ongoing work at Sehonghong is aug-
menting this picture (Loftus et al. 2015).
Occupation during MIS 2 at Melikane occurred just
before the LGM. The paleoenvironmental record from the
site suggests colder conditions than in MIS 3. Grassland
cover with a strong C
component most likely derived from
alpine sour grasses is present along with some panicoids,
which contribute to a low C
component. The presence of C
taxa may reect lower pCO
levels during the onset of the
LGM, as well as the presence of some tall, hydric, helio-
phyte grasses and sedges. The sediments from this level
(Layer 5) comprise anthropogenic materials mixed with
colluvial sediments and host bedrock attrition materials
derived from roof fall debris (Stewart et al. 2012). This
supports the notion of colder and drier conditions with
material derived from landscape erosion as well as
freeze-thaw and weathering processes. Tree cover is much
reduced, with trees and shrubs likely tightly restricted to the
river corridors or more sheltered areas of the landscape
where, as with the *4638 ka climatic downturn, there was
clearly sufcient surface water to support them. Cold and
frost tolerant taxa are present, including Leucosidea sericea
and Protea sp. At Rose Cottage Cave in the Caledon Valley,
late MIS 3 and early MIS 2 is also marked by Protea
sp. with Leucosidea sericea and other heathland species
(Wadley et al. 1992). The composition of the youngest layer
analyzed at Melikane is comparable with the addition of
Ericaceous taxa. It must be noted, however, that inferring
climate change from the presence of Erica sp. is limited by
our inability to identify this to the species level; it is possi-
ble, for example, that these specimens derive from Erica
264 B.A. Stewart et al.
cafforum, a streamside species that occurs in valleys together
with Buddleja and Leucosidea (Mucina and Rutherford
At Melikane there is an absence of human activity from
*24 ka until the start of the Holocene. Cold and semi-arid
to arid climatic conditions prevailed in the Drakensberg from
*24 ka, during what Lewis (1996) has named the Bottlenek
Stadial, which broadly corresponds with the LGM. During
this period there is evidence for niche glaciation and
extensive periglacial conditions across the region. Small
cirque glaciers were fed by snow-blow at altitudes as low as
2100 m at Mt. Enterprise in the Eastern Cape Drakensberg,
South Africa (Lewis and Illgner 2001), and 3100 m in the
Sekhokong range of eastern Lesotho. The onset of glaciation
in the Eastern Cape Drakensberg has been dated to *24 ka
(Lewis and Hanvey 1993), while in the Sekhokong range
glacial moraines have been dated between 19.4 and 14.7 ka
(Mills et al. 2009). Based on glacial landform elevations in
the Eastern Cape Drakensberg, Lewis and Illgner (2001)
estimate mean annual temperatures during the LGM were at
least *10 °C lower than present-day, with permanent
snowlines above 2100 m, and precipitation reduced by 70%
(Lewis 2008). Pollen data from Strathalan Cave B in the
Eastern Cape Drakensberg (Fig. 14.1) indicates alpine
environments after 24 ka at 1800 m. The site was abandoned
shortly thereafter and was not reoccupied until the early
Holocene (Opperman and Heydenrych 1990). A similar
occupational hiatus from early MIS 2 (28 ka) to the terminal
Pleistocene (12.5 ka) is apparent at the recently reexcavated
site of Ha Makotoko in lowland Lesotho to the west of our
research area (P. Mitchell, personal communication). The
cold, harsh conditions would have forced human populations
out of high elevation regions and displaced the vegetation
belts to much lower altitudes. It should be noted, however,
that a relatively ephemeral human presence during the LGM
is registered at nearby Sehonghong (Mitchell 1995).
Between Heinrich Stadial 2 and 1 core CD154-17-17 K
records very little discharge from rivers draining southeast-
ern southern Africa (Ziegler et al. 2013).
Finally, Melikanes uppermost layers reveal mixed and
reworked colluvial components that may have been formed
during the late glacial with reworking and bioturbation
during the Holocene. The higher chloridoid component is
more indicative of early Holocene environments, which is
supported by other δ
C records in the region (Grab et al.
2005; Roberts et al. 2013).
Following the Senqu
Most authors have assumed that prehistoric groups moved
between the highland Maloti-Drakensberg and the Eastern
Cape or KwaZulu-Natal midlands or coasts (Carter 1970,
1978; Cable 1984; Opperman 1987). However, we suspect
that the interior plateau also played an important role. Above
we put forward a model linking variability of population
movements into the highlands to prevailing climate. During
phases of heightened warmth or humidity we suggest that
incoming populations did usually derive from the forelands.
Such movements may have resulted in Melikanes initial
occupational pulse at *80 ka, and the subsequent pulses at
*61 and *50 ka. Greater ecological productivity at these
times might have led to higher population densities in the
southeastern forelands and simultaneously rendered the
highlands more hospitable, creating a push/pull effect. It
seems likely that such groups were more permanently based
along the coastal belt (e.g., Stapleton and Hewitt 1927,1928;
Davies 1975; Kaplan 1989,1990; Wadley 2006; Fisher et al.
2013) rather than in the midlands or Drakensberg foothills
where MSA sites are rare and often ephemeral (Deacon
1976; Derricourt 1977; Mazel 1982; Opperman 1987;
Opperman and Heydenrych 1990).
During drier periods, on the other hand, we anticipate that
dispersals more often originated on the interior plateau of
central South Africa. These may be responsible for Meli-
kanes most enduring pulse *4638 ka and the pre-LGM
pulse at *24 ka. Steep east-west rainfall and evaporation
gradients would have allowed interior populations evading
desiccation to move eastwards into the more stable, produc-
tive and heterogeneous mountain country closer to the
Orange-Senqu headwaters. The river itself may have served
as a conduit into the Maloti-Drakensberg via the natural gap
incised by its valley through the massifs southwest corner
(Fig. 14.1). Along this axis of movement, unlike that into the
forelands, there is abundant evidence for Late Pleistocene
activity. In the highlands themselves, dense clusters of
open-air and rockshelter sites including Melikane, Sehon-
ghong and others center on the uppermost Senqu and its
headwater tributaries (Carter 1978; Bousman 1988; Carter
et al. 1988; Mitchell 1996a,b; Dewar and Stewart 2011;
Stewart et al. 2012). Westward across the mountains similar
clusters occur along the Caledon and one of its major tribu-
taries, the Phuthiatsana, both of which ultimately feed the
Orange-Senqu (Mitchell and Steinberg 1992; Mitchell 1994;
Wadley 1997; Mitchell and Arthur 2010). Further down-
stream in the eastern and central Karoo, Sampson (1985) has
documented thousands of open-air MSA occurrences in his
comprehensive surveys of Orange River Scheme Area and
Seacow River Valley, another large tributary (Fig. 14.1).
These appear to reect successive humid-phase recoloniza-
tions of the central interior by MSA groups who through time
became increasingly tethered to the Orange-Senqu, its
tributaries and their conuences (Sampson 1985). We think it
plausible that in the drier abandonment phases between these
desert occupations some of these groups dispersed east
14 Follow the Senqu: Mountain Dispersals 265
into the Maloti-Drakensberg. Finally, we note that during the
Holocene when cultural connections and contacts are easier
to trace, links between the central Karoo and the Lesotho
highlands are numerous and well documented (Humphreys
1991; Mitchell 1996c,1999).
This paper has presented the results of several paleoenvi-
ronmental proxy analyses from the Late Pleistocene
sequence at Melikane Rockshelter in highland Lesotho. We
discussed these results in relation to other terrestrial and
marine archives to explore paleoenvironmental changes in
southeastern southern Africa and their possible implications
for regional population dynamics. Our expectations that a
human presence in the highland Maloti-Drakensberg inten-
sied when the wider region was relatively warm and/or arid
appear to be largely supported. Warm (and variably humid)
conditions conducive to upland exploitation seem charac-
terize the wider region *8380 ka, *50 ka and possibly
also *61 ka. In contrast, the occupational pulses *4638
and *24 ka seem to correspond to drier (and colder) phases.
However, deviations from our anticipated pattern also exist,
namely the occupational hiatuses during the periods of acute
aridity *5650 ka and across the LGM. While the former
hiatus may simply echo the wider absence of rockshelter
archeology across the subcontinent at this time, the latter is
probably rooted in the LGM temperature plunge. We also
posited a dual-source model of Pleistocene population
movements into the Maloti-Drakensberg linked to prevailing
climate. Testing this model is beyond the scope of this paper,
and will require further eldwork along with comparative
studies of lithic technology between highland sites and
others in possible lowland source areas. Integrating such
analyses with paleoenvironmental records like those pre-
sented here as well as new, more continuous regional
archives should yield further insights into the evolution of
human engagements with mountain systems.
Acknowledgments Our work at Melikane was made possible because
of an excavation permit generously granted (to BAS) by the Protection
and Preservation Committee (PPC) of the Lesotho Department of
Culture. We thank the PPC, and especially bo-Mme Moliehi Maneo
Ntene, Puseletso Moremi and the late Ntsema Khitsane for their sup-
port. The project Adaptations to Marginal Environments in the Middle
Stone Age(AMEMSA) is supported by grants from the McDonald
Institute for Archaeological Research, the University of Cambridge, the
British Academy, the Wenner-Gren Foundation, the Prehistoric Soci-
ety, and the Social Sciences and Humanities Research Council of
Canada. We also gratefully acknowledge Catherine Kneale for assis-
tance in processing the SOM δ
C samples. Finally, we thank Francis
Thackeray, Britt Bousman and a third anonymous reviewer, as well as
Peter Mitchell, for helping us substantially improve this paper.
Aldenderfer, M. S. (1998). Montane Foragers: Asana and the
South-Central Andean Archaic. Iowa City: University of Iowa
Aldenderfer, M. (2006). Modelling plateau peoples: The early human
use of the worlds high plateaux. World Archaeology, 38(3),
Alexandre, A., Meunier, J.-D., Lezine, A.-M., Vincens, A., &
Schwartz, D. (1997). Phytoliths: Indicators of grassland dynamics
during the late Holocene in intertropical Africa. Palaeogeography,
Palaeoclimatology, Palaeoecology, 136(1), 213229.
Allott, L. F. (2004). Changing environments in oxygen isotope stage 3:
Reconstructions using archaeological charcoal from Sibudu Cave.
South African Journal of Science, 100(34), 179184.
Allott, L. F. (2005). Palaeoenvironments of the Middle Stone Age at
Sibudu Cave, KwaZulu-Natal, South Africa: An analysis of
archaeological charcoal. Ph.D. Dissertation, University of the
Witwatersrand, 2005.
Allott, L. F. (2006). Archaeological charcoal as a window on
palaeovegetation and wood use during the Middle Stone Age at
Sibudu Cave. Southern African Humanities, 18(1), 173201.
Ambrose, S. H., & Norr, L. (1993). Experimental evidence for the
relationship of the carbon isotope ratios of whole diet and dietary
protein to those of bone collagen and carbonate. In J. B. Lambert &
G. Grupe (Eds.), Prehistoric human bone (pp. 137). Berlin:
Ambrose, S. H., & Sikes, N. E. (1991). Soil carbon isotope evidence for
Holocene habitat change in the Kenya Rift Valley. Science, 253
(5026), 14021405.
Atkinson, Q. D., Russell, D. G., & Drummond, A. J. (2009). Bayesian
coalescent inference of major human mitochondrial DNA hap-
logroup expansions in Africa. Proceedings of the Royal Society B:
Biological Sciences, 276(1655), 367373.
Barbieri, C., Vicente, M., Rocha, J., Mpoloka, S. W., Stoneking, M., &
Pakendorf, B. (2013). Ancient substructure in early mtDNA
lineages of southern Africa. The American Journal of Human
Genetics,92(2), 285292.
Bard, E., & Rickaby, R. E. (2009). Migration of the subtropical front as
a modulator of glacial climate. Nature, 460(7253), 380383.
Barham, L., & Mitchell, P. (2008). The rst Africans: African
archaeology from the earliest toolmakers to most recent foragers.
Cambridge: Cambridge University Press.
Batini, C., Coia, V., Battaggia, C., Rocha, J., Pilkington, M. M.,
Spedini, G., et al. (2007). Phylogeography of the human mitochon-
drial L1c haplogroup: Genetic signatures of the prehistory of Central
Africa. Molecular Phylogenetics and Evolution, 43(2), 635644.
Batini, C., Lopes, J., Behar, D. M., Calafell, F., Jorde, L. B., van der
Veen, L., et al. (2011). Insights into the demographic history of
African pygmies from complete mitochondrial genomes. Molecular
Biology and Evolution, 28(2), 10991110.
Bawden, M. G., & Carroll, D. M. (1968). The land resources of Lesotho.
London: United Kingdom Directorate of Overseas Surveys.
Beaumont, P. B. (1986). Where did all the young men go during 018
stage 2? Palaeoecology of Africa, 17, 7986.
Behar, D. M., Villems, R., Soodyall, H., Blue-Smith, J., Pereira, L.,
Metspalu, E., et al. (2008). The dawn of human matrilineal diversity.
The American Journal of Human Genetics, 82(5), 11301140.
Bender, S. J., & Wright, G. A. (1988). High-altitude occupations,
cultural process, and high plains prehistory: Retrospect and
prospect. American Anthropologist, 90(3), 619639.
Benedict, J. B. (1992). Footprints in the snow: High-altitude cultural
ecology of the Colorado front range. USA Arctic and Alpine
Research, 24(1), 116.
266 B.A. Stewart et al.
Bettinger, R. L. (1991). Aboriginal occupation at high altitude: Alpine
villages in the White Mountains of Eastern California. American
Anthropologist, 93(3), 656679.
Birkenhauer, J. (1991). The Great Escarpment of southern Africa and
its coastal forelands. A re-appraisal.München: Institut für
Geographie der Universität.
Black, K. D. (1991). Archaic continuity in the Colorado Rockies: The
Mountain Tradition. Plains Anthropologist, 36(133), 129.
Blome, M. W., Cohen, A. S., Tryon, C. A., Brooks, A. S., & Russell,
J. (2012). The environmental context for the origins of modern
human diversity: A synthesis of regional variability in African
climate 150,000-30,000 years ago. Journal of Human Evolution, 62
(5), 563592.
Botha, G. A., & Partridge, T. C. (2000). Colluvial deposits. Oxford
Monographs on Geology and Geophysics, 40, 8899.
Botha, G. A., Scott, L., Vogel, J. C., & Von Brunn, V. (1992).
Palaeosols and palaeoenvironments during the Late Pleistocene
hypothermal in Northern Natal. South African Journal of Science,
88(9), 508512.
Bousman, C. B. (1988). Prehistoric settlement patterns in the Senqun-
yane Valley, Lesotho. South African Archaeological Bulletin, 43
(147), 3337.
Bousman, C. B. (1991). Holocene paleoecology and Later Stone Age
hunter-gatherer adaptations in the South African interior plateau.
Ph.D. Dissertation, Southern Methodist University, 1991.
Brantingham, S., & Xing, G. (2006). Peopling of the Northern Tibetan
Plateau. World Archaeology, 38(3), 387414.
Brook, G. A., Cowart, J. B., & Marais, E. (1996). Wet and dry periods
in the Southern African summer rainfall zone during the last 300 kyr
from speleothem, tufa and sand dune age data. Palaeoecology of
Africa, 24, 147158.
Brook, G. A., Cowart, J. B., Brandt, S. A., & Scott, L. (1997).
Quaternary climatic change in southern and Eastern Africa during
the last 300 ka: The evidence from caves in Somalia and the
Transvaal region of South Africa. Zeitschrift für Geomorpholo-
gie NF Supplementband, 108, 1548.
Brook, G. A., Cowart, J. B., & Brandt, S. A. (1998). Comparison of
Quaternary environmental change in eastern and southern Africa
using cave speleothem, tufa and rock shelter sediment data. In A.
Alsharan, K. W. Glennie, G. L. Wintle, & C. G. St C. Kendall
(Eds.), Quaternary deserts and climate change (pp. 239250).
Rotterdam: Balkema.
Bruch, A. A., Sievers, C., & Wadley, L. (2012). Quantication of
climate and vegetation from Southern African Middle Stone Age
sites: An application using Late Pleistocene plant material from
Sibudu, South Africa. Quaternary Science Reviews, 45,717.
Burke, K., & Gunnell, Y. (2008). The African erosion surface: A
continental-scale synthesis of geomorphology, tectonics, and envi-
ronmental change over the past 180 million years. Geological
Society of America Memoirs, 201,166.
Cable, J. H. C. (1984). Economy and technology in Late Stone Age
southern Natal. BAR International Series, 201. Oxford: British
Archaeological Reports.
Cambray, J. A., Davies, B. R., Ashton, P. J., Agnew, J. D., De Moor, F.
C., & Skelton, P. H. (1986). The Orange-Vaal river system. In B.
R. Davies & K. F. Walker (Eds.), The ecology of river systems
(pp. 89161). Monographiae Biologicae 60. Dordrecht: Junk.
Cancellieri, E., & di Lernia, S. (2013). Middle Stone Age human
occupation and dispersals in the Messak Plateau (SW Libya, Central
Sahara). Quaternary International, 300, 142152.
Carbutt, C., & Edwards, T. J. (2003). The ora of the Drakensberg
Alpine Centre. Edinburgh Journal of Botany, 60(03), 581607.
Carbutt, C., & Edwards, T. J. (2006). The endemic and near-endemic
angiosperms of the Drakensberg Alpine Centre. South African
Journal of Botany, 72(1), 105132.
Carter, P. L. (1970). Late Stone Age exploitation patterns in southern
Natal. South African Archaeological Bulletin, 25, 5558.
Carter, P. L. (1976). The effects of climate change on settlement in
Eastern Lesotho during the Middle and Later Stone Age. World
Archaeology, 8, 197206.
Carter, P. L. (1978). The prehistory of eastern Lesotho. Ph.D.
Dissertation, University of Cambridge, 1987.
Carter, P. L., Mitchell, P. J., & Vinnicombe, P. (1988). Sehonghong:
The Middle and Later Stone Age industrial sequence at a Lesotho
rock-shelter. BAR International Series 409. Oxford: British
Archaeological Reports.
Castañeda, I. S., Mulitza, S., Schefuß, E., Lopes dos Santos, R. A.,
Sinninghe Damsté, J. S., & Schouten, S. (2009). Wet phases in the
Sahara/Sahel region and human migration patterns in North Africa.
Proceedings of the National Academy of Sciences, 106(48), 20159
Chase, B. M. (2010). South African palaeoenvironments during marine
oxygen isotope Stage 4: A context for the Howiesons Poort and
Still Bay industries. Journal of Archaeological Science, 37(6),
Clark, J. L., & Plug, I. (2008). Animal exploitation strategies during the
South African Middle Stone Age: Howiesons Poort and
post-Howiesons Poort fauna from Sibudu Cave. Journal of Human
Evolution, 54(6), 886898.
Clark, V. R., Barker, N. P., & Mucina, L. (2011). The Great
Escarpment of Southern Africa: A new frontier for biodiversity
exploration. Biodiversity and Conservation, 20(12), 25432561.
Clarke, M. L., Vogel, J. C., Botha, G. A., & Wintle, A. G. (2003). Late
Quaternary hillslope evolution recorded in Eastern South African
colluvial badlands. Palaeogeography, Palaeoclimatology, Palaeoe-
cology, 197(34), 199212.
Cordova, C. E. (2013). C
poaceae and restionaceae phytoliths as
potential proxies for reconstructing winter rainfall in South Africa.
Quaternary International, 287, 121140.
Cornelissen, E. (2002). Human responses to changing environments in
Central Africa between 40,000 and 12,000 BP. Journal of World
Prehistory, 16(3), 197235.
Coulthard, T. J., Ramirez, J. A., Barton, N., Rogerson, M., & Brücher,
T. (2013). Were rivers owing across the Sahara during the Last
Interglacial? Implications for human migration through Africa.
PLoS ONE, 8(9), e74834.
Cowling, R. M. & Hilton-Taylor, C. (1994). Patterns of plant diversity
and endemism in Southern Africa: An overview. In B. J. Huntley
(Ed.), Botanical diversity in southern Africa.Strelitzia,1, 3152.
Cowling, R. M. & Hilton-Taylor, C. (1997). Phytogeography, ora and
endemism. In R. M. Cowling, D. M. Richardson & S. M. Pierce
(Eds.), Vegetation of southern Africa (pp. 4361). Cambridge:
Cambridge University Press.
Davies, O. (1975). Excavations at Shongweni South Cave: The oldest
evidence to date for cultigens in southern africa. Southern African
Humanities, 22(2), 627662.
Davis, S. D., Heywood, V. H., & Hamilton, A. C. (Eds.) (1994).
Centres of plant diversity. A guide and strategy for their conser-
vation. Vol. 1. Europe, Africa, South West Asia and the Middle
East. Gland, Switzerland: IUCN and WWF.
Deacon, H. J. (1976). Where hunters gathered: A study of Holocene
Stone Age people in the Eastern Cape. Claremont: South African
Archaeological Society.
Deacon, H. J. (1979). Excavations at Boomplaas Cave: A sequence
through the Upper Pleistocene and Holocene in South Africa. World
Archaeology, 10(3), 241257.
Deacon, H. J., & Thackeray, J. F. (1984). Late Pleistocene environ-
mental changes and implications for the archaeological record in
Southern Africa. In J. C. Vogel (Ed.), Late Cainozoic palaeoclimates
of the southern hemisphere (pp. 375390). Rotterdam: Balkema.
14 Follow the Senqu: Mountain Dispersals 267
Deacon, J. (1974). Patterning in the radiocarbon dates for the
Wilton/Smitheld complex in Southern Africa. The South African
Archaeological Bulletin, 29,318.
Derricourt, R. M. (1977). Prehistoric man in the Ciskei and Transkei.
Cape Town: Struik.
Dewar, G., & Stewart, B. A. (2011). 2009 archaeological eld survey
of the Sehonghong and Senqu Rivers, Thaba-Tseka District.
Lesotho: Unpublished report.
Dewar, G., & Stewart, B. A. (2012). Preliminary results of excavations
at Spitzkloof Rockshelter, Richtersveld, South Africa. Quaternary
International, 270, 3039.
Dowson, T. A. (1988). Shifting vegetation zones in the Holocene and
later Pleistocene: Preliminary charcoal evidence from Jubilee
Shelter, Magaliesberg, Southern Transvaal. Palaeoecology of
Africa, 19, 233239.
Drake, N. A., Blench, R. M., Armitage, S. J., Bristow, C. S., & White,
K. H. (2011). Ancient watercourses and biogeography of the Sahara
explain the peopling of the desert. Proceedings of the National
Academy of Sciences, 108(2), 458462.
Dupont, L. M., Caley, T., Kim, J.-H., Castañeda, I., Malaizé, B., &
Giraudeau, J. (2011). Glacial-interglacial vegetation dynamics in
South Eastern Africa coupled to sea surface temperature variations
in the Western Indian Ocean. Climate of the Past, 7, 12091224.
Eamus, D., Hatton, T., Cook, P., & Colvin, C. (2006). Ecohydrology:
Vegetation function, water and resource management. Colling-
wood, Australia: CSIRO Publishing.
Earle, A., Malzbender, D., Turton, A., & Manzungu, E. (2005). A
preliminary basin prole of the Orange/Senqu River. Pretoria:
AWIRU, University of Pretoria.
Ehleringer, J. R., Cerling, T. E., & Helliker, B. R. (1997). C
photosynthesis, atmospheric CO
, and climate. Oecologia, 112(3),
Eichhorn, B. (2002). Anthrakologische untersuchungen zur vegeta-
tiongeshichte des Kaokolandes, nordwest Namibia. Ph.D. Disser-
tation, University of Köln, 2002.
Esterhuysen, A. B. (1996). Palaeoenvironmental reconstruction from
Pleistocene to present: An analysis of charcoal from sites in the
Eastern Free State and Lesotho. Ph.D. Dissertation, University of
the Witwatersrand, 1996.
Ficken, K. J., Wooller, M. J., Swain, D. L., Street-Perrott, F. A., &
Eglinton, G. (2002). Reconstruction of a subalpine grass-dominated
ecosystem, Lake Rutundu, Mount Kenya: A novel multi-proxy
approach. Palaeogeography, Palaeoclimatology, Palaeoecology,
177(1), 137149.
Fisher, E. C., Albert, R.-M., Botha, G., Cawthra, H. C., Esteban, I.,
Harris, J., et al. (2013). Archaeological reconnaissance for Middle
Stone Age sites along the Pondoland coast, South Africa.
PaleoAnthropology, 2013, 104137.
Foley, R. A., Maíllo-Fernández, J. M., & Lahr, M. M. (2013). The
Middle Stone Age of the central Sahara: Biogeographical oppor-
tunities and technological strategies in later human evolution.
Quaternary International, 300, 153170.
Glenny, W. (2006). Report on the micromammal assemblage analysis
from Sibudu Cave. Southern African Humanities, 18, 279288.
Grab, S. W. (1994). Thufur in the Mohlesi Valley, Lesotho, Southern
Africa. Permafrost and Periglacial Processes, 5(2), 111118.
Grab, S. W. (1997). Thermal regime for a thufa apex and its adjoining
depression, Mashai Valley, Lesotho. Permafrost and Periglacial
Processes, 8(4), 437445.
Grab, S. W., & Nash, D. J. (2010). Documentary evidence of climate
variability during cold seasons in Lesotho, Southern Africa,
1833-1900. Climate Dynamics, 34(4), 473499.
Grab, S., Scott, L., Rossouw, L., & Meyer, S. (2005). Holocene
palaeoenvironments inferred from a sedimentary sequence in the
Tsoaing River Basin, Western Lesotho. Catena, 61(1), 4962.
Harpending, H., & Davis, H. (1977). Some implications for
hunter-gatherer ecology derived from the spatial structure of
resources. World Archaeology, 8(3), 275286.
Henn, B. M., Gignoux, C. R., Jobin, M., Granka, J. M., Macpherson,
J. M., Kidd, J. M., et al. (2011). Hunter-gatherer genomic diversity
suggests a Southern African origin for modern humans. Proceed-
ings of the National Academy of Sciences, 108(13), 51545162.
Henshilwood, C. S. (2012). Late Pleistocene techno-traditions in
Southern Africa: A review of the Still Bay and Howiesons Poort, c.
7559 ka. Journal of World Prehistory, 25(34), 205237.
Herries, A. I. R. (2006). Archaeomagnetic evidence for climate change
at Sibudu Cave. Southern African Humanities, 18, 131147.
Holmgren, K., Lee-Thorp, J. A., Cooper, G. R. J., Lundblad, K.,
Partridge, T. C., Scott, L., et al. (2003). Persistent millennial-scale
climatic variability over the past 25,000 years in Southern Africa.
Quaternary Science Reviews, 22(2122), 23112326.
Huang, Y., Street-Perrott, F. A., Perrott, R. A., Metzger, P., & Eglinton,
G. (1999). Glacial-interglacial environmental changes inferred from
molecular and compound-specicδ
C analyses of sediments from
Sacred Lake, Mt. Kenya. Geochimica Et Cosmochimica Acta, 63
(9), 13831404.
Humphreys, A. J. B. (1991). On the distribution and dating of bifacial
tanged and barbed arrowheads in the interior of South Africa. The
South African Archaeological Bulletin, 46, 4143.
InsideWood (2004-onwards). Retrieved May 14, 2012 from http://
Jacobs, Z., & Richard, R. G. (2008). Testing times: Old and new
chronologies for the Howiesons Poort and Still Bay industries in
environmental context. South African Archaeological Bulletin
Goodwin Series, 10,934.
Jacobs, Z., Roberts, R. G., Galbraith, R. F., Deacon, H. J., Grün, R.,
Mackay, A., et al. (2008). Ages for the Middle Stone Age of
Southern Africa: Implications for human behavior and dispersal.
Science, 322(5902), 733735.
Jacot Guillarmod, A. (1971). Flora of Lesotho (Basutoland). Lehre:
Verlag Von Cramer.
Johnson, M. R., Van Vuuren, C. J., Hegenberger, W. F., Key, R., &
Show, U. (1996). Stratigraphy of the Karoo Supergroup in Southern
Africa: An overview. Journal of African Earth Sciences, 23(1),
Jouzel, J., Masson-Delmotte, V., Cattani, O., Dreyfus, G., Falourd, S.,
Hoffmann, G., et al. (2007). Orbital and millennial Antarctic climate
variability over the past 800,000 years. Science, 317(5839), 793
Kaplan, J. M. (1989). 45000 years of hunter-gatherer history in Natal as
seen from Umhlatuzana Rock Shelter. South African Archaeolog-
ical Society Goodwin Series, 6,716.
Kaplan, J. M. (1990). The Umhlatuzana Rock Shelter sequence: 100
000 years of Stone Age history. Natal Museum Journal of
Humanities, 2,194.
Killick, D. J. B. (1963). An account of the plant ecology of the
Cathedral Peak area of the Natal Drakensberg. Memoirs of the
Botanical Survey of South Africa, 34,1178.
Killick, D. J. B. (1978). The afro-alpine region. In M. J. A. Werger
(Ed.), Biogeography and ecology of Southern Africa (pp. 515560).
The Hague: Junk.
Killick, D. J. B. (1990). Aeld guide to the ora of the Natal
Drakensberg. Johannesburg: Jonathan Ball.
Kingdon, J. (1990). Island Africa: The evolution of Africas rare
animals and plants. London: Collins.
Kornfeld, M., Frison, G. C., Larson, M. L., & Bradley, B. A. (2010).
Prehistoric hunter-gatherers of the High Plains and Rockies.
Walnut Creek: Left Coast Press.
Kristen, I., Fuhrmann, A., Thorpe, J., Rohl, U., Wilkes, H., & Oberhansli,
H. (2007). Hydrological changes in Southern Africa over the last
268 B.A. Stewart et al.
200 ka as recorded in lake sediments from the Tswaing impact crater.
South African Journal of Geology, 110(23), 311326.
Kromhout, C. P. (1975). n Sleutel vir die mikroskopiese uitkenning
van die vernaamste inheemse houtesoorte van Suid-Afrika. Depart-
ment of Forestry Bulletin, 50,1140.
Landman, W. A., & Mason, S. J. (1999). Change in the association
between Indian Ocean sea-surface temperatures and summer rainfall
over South Africa and Namibia. International Journal of Climatol-
ogy, 19(13), 14771492.
Leney, L., & Casteel, R. W. (1975). Simplied procedure for
examining charcoal specimens for identication. Journal of
Archaeological Science, 2(2), 153159.
Lewis, C. A. (1996). Periglacial features. In C. A. Lewis (Ed.), The
geomorphology of the Eastern Cape, South Africa (pp. 103119).
Grahamstown: NISC.
Lewis, C. A. (2008). Late Quaternary climatic changes, and associated
human responses, during the Last *45 000 yr in the Eastern and
adjoining Western Cape, South Africa. Earth-Science Reviews, 88
(34), 167187.
Lewis, C. A., & Hanvey, P. M. (1993). The remains of rock glaciers in
Bottelnek, East Cape Drakensberg, South Africa. Transactions of
the Royal Society of South Africa, 48, 265289.
Lewis, C. A., & Illgner, P. M. (2001). Late Quaternary glaciation in
Southern Africa: Moraine ridges and glacial deposits at mount
enterprise in the Drakensberg of the Eastern Cape Province, South
Africa. Journal of Quaternary Science, 16(4), 365374.
Loftus, E., Stewart, B. A., Dewar, G., & Lee-Thorp, J. (2015). Stable
isotope evidence of MIS 3 to middle Holocene palaeoenvironments
from Sehonghong, eastern Lesotho. Journal of Quaternary Science,
30(8), 805816.
Lombard, M., Schlebusch, C., & Soodyall, H. (2013). Bridging
disciplines to better elucidate the evolution of early Homo sapiens
in Southern Africa. South African Journal of Science, 109(11/12),
Lynch, S. D. (2004). Development of a raster database of annual,
monthly and daily rainfall for Southern Africa: Report to the water
research commission. WRC Report No. 1156/1/03. Pretoria: Water
Research Commission.
Lyons, R., Tooth, S., & Duller, G. A. T. (2014). Late Quaternary
climatic changes revealed by luminescence dating, mineral mag-
netism and diffuse reectance spectroscopy of river terrace
palaeosols: A new form of geoproxy data for the Southern African
interior. Quaternary Science Reviews, 95, 4359.
Madsen, D. B., & Metcalf, M. D. (2000). Intermountain archaeology.
Salt Lake City: University of Utah Press.
Mason, S. J., & Jury, M. R. (1997). Climatic variability and change
over Southern Africa: A reection on underlying processes.
Progress in Physical Geography, 21(1), 2350.
Mazel, A. D. (1982). Evidence for pre-Later Stone Age occupation of
the Natal Drakensberg. Southern African Humanities, 25(1), 6165.
Mazel, A.D. (1984). Through the keyhole: A preliminary peep at the
lithic composition of Later Stone Age sites in the central and upper
Tugela River Basin, Natal, South Africa. In M. Hall, G. Avery, D.
M. Wilson, & A. J. B. Humphreys (Eds.), Frontiers: Southern
African archaeology today (pp. 182193). BAR International Series
207. Oxford: British Archaeological Reports.
Mercader, J., Astudillo, F., Barkworth, M., Bennett, T., Esselmont, C.,
Kinyanjui, R., et al. (2010). Poaceae phytoliths from the Niassa Rift,
Mozambique. Journal of Archaeological Science, 37(8), 1953
Mercader, J., Bennett, T., Esselmont, C., Simpson, S., & Walde, D.
(2013). Phytoliths from Middle Stone Age habitats in the Mozam-
bican Rift (10529 ka). Journal of Human Evolution, 64(5), 328336.
Midgley, D. C., Pitman, W. V., & Middleton, B. J. (1994). The surface
water resources of South Africa 1990, volumes 1 to 6. WRC Report
Nos. 298/1.1/94 to 298/6.1/94 (text) and 298/1.2/94 to 298/6.2/94
(maps). Pretoria: Water Research Commission.
Mills, S. C., Grab, S. W., & Carr, S. J. (2009). Recognition and
palaeoclimatic implications of late quaternary niche glaciation in
Eastern Lesotho. Journal of Quaternary Science, 24(7), 647663.
Mitchell, P. J. (1988). The Late Pleistocene early microlithic assem-
blages of Southern Africa. World Archaeology, 20(1), 2739.
Mitchell, P. J. (1990). A palaeoecological model for archaeological site
distribution in Southern Africa during the Upper Pleniglacial and
Late Glacial. In C. Gamble & O. Soffer (Eds.), The world at 18,000
BP: Low latitudes (pp. 189205). London: Unwin Hyman.
Mitchell, P. J. (1994). The archaeology of the Phuthiatsana-ea-Thaba
Bosiu Basin, Lesotho, Southern Africa: Changes in Later Stone Age
regional demography. Antiquity,68, 8383.
Mitchell, P. J. (1995). Revisiting the Robberg: New results and a
revision of old ideas at Sehonghong Rock Shelter, Lesotho. South
African Archaeological Bulletin, 50, 2838.
Mitchell, P. J. (1996a). The Late Quaternary landscape at Sehonghong
in the Lesotho Highlands, Southern Africa. Antiquity, 70, 623638.
Mitchell, P. J. (1996b). The Late Quaternary of the Lesotho Highlands,
Southern Africa: Preliminary results and future potential of ongoing
research at Sehonghong Shelter. Quaternary International, 33, 3543.
Mitchell, P. J. (1996c). Prehistoric exchange and interaction in
southeastern Southern Africa: Marine shells and ostrich eggshell.
African Archaeological Review, 13(1), 3576.
Mitchell, P. J. (1998). The archaeology of the Alfred County Cave,
KwaZulu-Natal. Natal Museum Journal of Humanities, 10,117.
Mitchell, P. J. (1999). Pressure-aked points in Lesotho: Dating,
distribution and diversity. South African Archaeological Bulletin,
54, 9096.
Mitchell, P. J. (2008). Developing the archaeology of marine isotope
Stage 3. South African Archaeological Society Goodwin Series, 10,
Mitchell, P. J. (2009). Gathering together a history of the People of the
Eland: Towards an archaeology of Maloti-Drakensberg
hunter-gatherers. In P. J. Mitchell & B. W. Smith (Eds.), The elands
people: New perspectives in the rock art of the Maloti-Drakensberg
Bushmen (pp. 99139). Johannesburg: Wits University Press.
Mitchell, P., & Arthur, C. (2010). Archaeological eldwork in the
Metolong Dam catchment, Lesotho, 200810. Nyame Akuma, 74,
Mitchell, P. J., & Steinberg, J. M. (1992). Ntloana Tsoana: A Middle
Stone Age sequence from Western Lesotho. South African
Archaeological Bulletin, 47, 2633.
Moore, A., & Blenkinsop, T. (2006). Scarp retreat versus pinned drainage
divide in the formation of the Drakensberg Escarpment, Southern
Africa. South African Journal of Geology, 109(4), 599610.
Morgan, C. (2009). Climate change, uncertainty and prehistoric
hunter-gatherer mobility. Journal of Anthropological Archaeology,
28(4), 382396.
Morgan, C., Losey, A., & Adams, R. (2012). High-altitude
hunter-gatherer residential occupations in Wyomings Wind River
Range. North American Archaeologist, 33(1), 3579.
Mucina, L., & Rutherford, M. C. (2006). The vegetation of South
Africa, Lesotho and Swaziland. Strelitzia 19. Pretoria: South
African National Biodiversity Institute.
Olago, D. O., Street-Perrott, F. A., Perrott, R. A., Ivanovich, M., &
Harkness, D. D. (1999). Late Quaternary glacial-interglacial cycle
of climatic and environmental change on Mount Kenya. Kenya.
Journal of African Earth Sciences, 29(3), 593618.
OLeary, M. H. (1981). Carbon isotope fractionation in plants.
Phytochemistry, 20(4), 553567.
Opperman, H. (1987). The Later Stone Age of the Drakensberg Range
and its foothills. BAR International Series 339. Oxford: British
Archaeological Reports.
14 Follow the Senqu: Mountain Dispersals 269
Opperman, H. (1988). An excavation of a Middle Stone Age deposit in
Grassridge Rockshelter, Sterkstroom District, Cape Province. Fort
Hare Papers, 9, 5161.
Opperman, H. (1992). A report on the results of a test pit in Strathalan
Cave B, Maclear District, North-Eastern Cape. Southern African
Field Archaeology, 1, 98102.
Opperman, H. (1996). Strathalan Cave B, north-eastern Cape Province,
South Africa: Evidence for human behaviour 2900026000 years
ago. Quaternary International, 33, 4553.
Opperman, H., & Heydenrych, B. (1990). A 22 000 year-old Middle
Stone Age camp site with plant food remains from the North-Eastern
Cape. South African Archaeological Bulletin, 45, 9399.
Parker, A. G., Lee-Thorp, J., & Mitchell, P. J. (2011). Late Holocene
Neoglacial conditions from the Lesotho Highlands, Southern
Africa: Phytolith and stable carbon isotope evidence from the
archaeological site of Likoaeng. Proceedings of the Geologists
Association, 122(1), 201211.
Parkington, J. E. (1972). Seasonal mobility in the Late Stone Age.
African Studies, 31(4), 223244.
Parkington, J. E. (1977). Follow the San: An analysis of seasonality in
prehistory of the South Western Cape. Ph.D. Dissertation, Univer-
sity of Cambridge, 1977.
Parkington, J. E. (1980). Time and place: Some observations on spatial
and temporal patterning in the Later Stone Age sequence in
Southern Africa. South African Archaeological Bulletin, 35, 7383.
Parkington, J. E., & Poggenpoel, C. (1980). Senqu: An archaeological
reconnaissance along the southern perimeter road. Unpublished
Partridge, T. C., & Maud, R. R. (1987). Geomorphic evolution of
Southern Africa since the Mesozoic. South African Journal of
Geology, 90(2), 179208.
Partridge, T. C., Kerr, S. J., Metcalfe, S. E., Scott, L., Talma, A. S., &
Vogel, J. C. (1993). The Pretoria Saltpan: A 200,000 year Southern
African lacustrine sequence. Palaeogeography, Palaeoclimatology,
Palaeoecology, 101(34), 317337.
Partridge, T. C., Scott, L., & Schneider, R. R. (2004). Between Agulhas
and Benguela: Responses of Southern African climates of the Late
Pleistocene to current uxes, orbital precession and the extent of the
circum-Antarctic vortex. In R. W. Battarbee, F. Gasse, & C.
E. Stickley (Eds.), Past climate variability through Europe and
Africa (pp. 4568). Berlin: Springer Verlag.
Pereira, L., Silva, N. M., Franco-Duarte, F., Fernandes, V., Pereira,
J. B., Costa, M. D., et al. (2010). Population expansion in the North
African Late Pleistocene signalled by mitochondrial DNA hap-
logroup U6. BMC Evolutionary Biology, 10(1), 390.
Pickrell, J. K., Patterson, N., Barbieri, C., Berthold, F., Gerlach, L.,
Güldemann, T., et al. (2012). The genetic prehistory of Southern
Africa. Nature Communications, 3, 1143.
Pienaar, M., Woodborne, S., & Wadley, L. (2008). Optically stimulated
luminescence dating at Rose Cottage Cave. South African Journal
of Science, 104(12), 6570.
Prell, W. L., & Hutson, W. H. (1979). Zonal temperature-anomaly
maps of Indian Ocean surface waters: Modern and ice-age patterns.
Science, 206(4417), 454456.
Reason, C. J. C., & Mulenga, H. (1999). Relationships between South
African rainfall and SST anomalies in the southwest Indian Ocean.
International Journal of Climatology, 19(15), 16511673.
Retallack, G. J. (2001). A 300-million-year record of atmospheric carbon
dioxide from fossil plant cuticles. Nature, 411(6835), 287290.
Rito, T., Richards, M. B., Fernandes, V., Alshamali, F., Cerny, V.,
Pereira, P., et al. (2013). The rst modern human dispersals across
Africa. PLoS ONE, 8(11), e80031.
Rivers-Moore, N. A., Goodman, P. S., & Nkosi, M. R. (2007). An
assessment of the freshwater natural capital in Kwazulu-Natal for
conservation planning. Water SA, 33(5), 665674.
Roberts, P., Lee-Thorpe, J. A., Mitchell, P. J., & Arthur, C. (2013).
Stable carbon isotopic evidence for climate change across the Late
Pleistocene to early Holocene from Lesotho, Southern Africa.
Journal of Quaternary Science, 28(4), 360369.
Sampson, C.G., (1985). Atlas of Stone Age settlement in the Central
and Upper Seacow Valley. Bloemfontein: Memoirs van die
Nasionale Museum Bloemfontein 20: 1112.
Scheinfeldt, L. B., Soi, S., & Tishkoff, S. A. (2010). Working toward a
synthesis of archaeological, linguistic, and genetic data for inferring
African population history. Proceedings of the National Academy of
Sciences, 107(Supplement 2), 89318938.
Schlebusch, C. M., Skoglund, P., Sjödin, P., Gattepaille, L. M.,
Hernandez, D., Jay, F., et al. (2012). Genomic variation in seven
Khoe-San groups reveals adaptation and complex African history.
Science, 338(6105), 374379.
Schlebusch, C. M., Lombard, M., & Soodyall, H. (2013). MtDNA
control region variation afrms diversity and deep sub-structure in
populations from Southern Africa. BMC Evolutionary Biology,
13(1), 56.
Schlüter, T. (2006). Geological atlas of Africa, with notes on
stratigraphy, tectonics, economic geology, geohazards and geosites
of each country. Berlin: Springer.
Schulze, R. E. (1979). Hydrology and water resources of the
Drakensberg. Pietermaritzburg: Natal Town and Regional Planning
Schulze, R. E. (2008). South African atlas of climatology and
agrohydrology. WRC Report No. 1489/1/06. Pretoria: Water
Research Commission.
Schulze, R., Meigh, J., & Horan, M. (2001). Present and potential
future vulnerability of eastern and Southern Africas hydrology and
water resources. South African Journal of Science, 97, 150160.
Scott, L. (1999). Vegetation history and climate in the Savanna Biome,
South Africa, since 190,000 ka: A comparison of pollen data from
the Tswaing Crater (the Pretoria Saltpan) and Wonderkrater.
Quaternary International, 57, 215223.
Sealy, J. (2016). Cultural change, demography, and the archaeology of
the last 100 kyr in Southern Africa. In S. C. Jones & B. A. Stewart
(Eds.), Africa from MIS 6-2: Population dynamics and paleoenvi-
ronments. (pp. 6575). Dordrecht: Springer.
Sene, K. J., Jones, D. A., Meigh, J. R., & Farquharson, F. A. K. (1998).
Rainfall and ow variations in the Lesotho Highlands. International
Journal of Climatology, 18(3), 329345.
Sievers, C. (2006). Seeds from the Middle Stone Age layers at Sibudu
Cave. Southern African Humanities, 18(1), 203222.
Smith, J. M. (1997). Stable isotope analysis of fauna and soils from
sites in the Eastern Free State and Western Lesotho, Southern
Africa: A palaeoenvironmental interpretation. MSc Thesis, Univer-
sity of Cape Town, 1997.
Smith, J. M., Lee-Thorp, J. A., & Sealy, J. C. (2002). Stable carbon and
oxygen isotopic evidence for late Pleistocene to middle Holocene
climatic uctuations in the interior of southern Africa. Journal of
Quaternary Science, 17(7), 683695.
Soares, P., Alshamali, F., Pereira, J. B., Fernandes, V., Silva, N. M.,
Afonso, C., et al. (2012). The expansion of MtDNA haplogroup L3
within and out of Africa. Molecular Biology and Evolution, 29(3),
Stapleton, P., & Hewitt, J. (1927). Stone implements from a rock
shelter at Howiesons Poort, near Grahamstown. South African
Journal of Science, 24, 574587.
Stapleton, P., & Hewitt, J. (1928). Stone implements from Howiesons
Poort near Grahamstown. South African Journal of Science, 25,
Steenkamp, Y., Van Wyk, A. E., Smith, G. F., & Steyn, H. (2005).
Floristic endemism in southern Africa: A numerical classication at
generic level. Biologiske Skrifter, 55, 253271.
270 B.A. Stewart et al.
Stewart, B. A., Dewar, G. I., Morley, M. W., Inglis, R. H., Wheeler,
M., Jacobs, Z., et al. (2012). Afromontane foragers of the Late
Pleistocene: Site formation, chronology and occupational pulsing at
Melikane Rockshelter, Lesotho. Quaternary International, 270, 40
Street-Perrott, F. A., Huang, Y., Perrott, R. A., Eglinton, G., Barker, P.,
Khelifa, L. B., et al. (1997). Impact of lower atmospheric carbon
dioxide on tropical mountain ecosystems. Science, 278(5342),
Swanevelder, C. J. (1981). Utilising South Africas largest river: The
physiographic background to the Orange River Scheme. GeoJour-
nal, 2(2), 2940.
Temme, A. J. A. M., Baartman, J. E. M., Botha, G. A., Veldkamp, A.,
Jongmans, A. G., & Wallinga, J. (2008). Climate controls on late
Pleistocene landscape evolution of the Okhombe Valley,
KwaZulu-Natal, South Africa. Geomorphology, 99(1), 280295.
Theunissen, J. D. (1994). A method for isolating and preparing silica
bodies in grasses for scanning electron microscopy. Biotechnic and
Histochemistry, 69(5), 291294.
Thackeray, J. F. (1987). Late Quaternary environmental changes
inferred from small mammalian fauna, Southern Africa. Climatic
Change, 10, 285305.
Thackeray, J. F. (1988). Quantication of climatic change during the
Late Quaternary in Southern Africa. Palaeoecology of Africa, 19,
Tishkoff, S. A., Gonder, M. K., Henn, B. M., Mortensen, H., Knight,
A., Gignoux, C., et al. (2007). History of click-speaking populations
of Africa inferred from mtDNA and Y chromosome genetic
variation. Molecular Biology and Evolution, 24(10), 21802195.
Tishkoff, S. A., Reed, F. A., Friedlaender, F. R., Ehret, C., Ranciaro,
A., Froment, A., et al. (2009). The genetic structure and history of
Africans and African Americans. Science, 324(5930), 10351044.
Twiss, P. C. (1987). Grass-opal phytoliths as climatic indicators of the
Great Plains Pleistocene. Kansas Geological Survey Guidebook
Series, 5, 179185.
Tyson, P. D., Preston-Whyte, R. A., & Schulze, R. E. (1976). The
Climate of the Drakensberg. Pietermaritzburg: Town and Regional
Planning Commission.
van Campo, E., Duplessy, J. C., Prell, W. L., Barratt, N., & Sabatier, R.
(1990). Comparison of terrestrial and marine temperature estimates
for the past 135 kyr off Southeast Africa: A test for GCM
simulations of palaecoclimate. Nature, 348(6298), 209212.
van Wyk, A. E., & Smith, G. F. (2001). Regions of oristic endemism
in Southern Africa: A review with emphasis on succulents. Pretoria:
Umdaus Press.
van Zinderen Bakker, E. M., & Werger, M. J. (1974). Environment,
vegetation and phytogeography of the high-altitude bogs of
Lesotho. Plant Ecology, 29(1), 3749.
Vogel, J. C. (1983). Isotopic evidence for the past climates and
vegetation of southern Africa. Bothalia, 14(34), 391394.
Vogel, J. C., Fuls, A., & Ellis, R. P. (1978). Geographical distribution
of kranz grasses in South Africa. South African Journal of Science,
74, 209215.
Veeramah, K. R., Wegmann, D., Woerner, A., Mendez, F. L., Watkins,
J. C., Destro-Bisol, G., et al. (2012). An early divergence of
KhoeSan ancestors from those of other modern humans is supported
by an ABC-based analysis of autosomal resequencing data.
Molecular Biology and Evolution,29(2), 617630.
Wadley, L. (1997). Rose Cottage Cave: Archaeological work 1987 to
1997. South African Journal of Science, 93(10), 439444.
Wadley, L. (2006). Partners in grime: Results of multi-disciplinary
archaeology at Sibudu Cave. Southern African Humanities, 18(1),
Wadley, L., Esterhuysen, A. B., & Jeannerat, C. (1992). Vegetation
changes in the eastern Orange Free State: The Holocene and later
Pleistocene evidence from charcoal studies at Rose Cottage Cave.
South African Journal of Science, 88(11), 558563.
Walsh, K. (2005). Risk and marginality at high altitudes: New
interpretations from eldwork on the Faravel Plateau,
Hautes-Alpes. Antiquity, 79(304), 289305.
Walsh, K., Richer, S., & De Beaulieu, J. L. (2006). Attitudes to altitude:
Changing meanings and perceptions within a marginalalpine
landscape - the integration of palaeoecological and archaeological
data in a high-altitude landscape in the French Alps. World
Archaeology, 38(3), 436454.
Wedin, D. A., Tieszen, L. L., Dewey, B., & Pastor, J. (1995). Carbon
isotope dynamics during grass decomposition and soil organic
matter formation. Ecology, 76(5), 13831392.
Wellington, J. H. (1955). Southern Africa: A geographical study.
Cambridge: Cambridge University Press.
Wheeler, E., Baas, P., & Gasson, P. (1989). IAWA list of microscopy
features for hardwood identication. IAWA Bulletin, 10(3), 219
Wooller, M. J., Swain, D. L., Ficken, K. J., Agnew, A. D. Q.,
Street-Perrott, F. A., & Eglinton, G. (2003). Late Quaternary
vegetation changes around Lake Rutundu, Mount Kenya, East
Africa: Evidence from grass cuticles, pollen and stable carbon
isotopes. Journal of Quaternary Science, 18(1), 315.
Wright, G. A., Bender, S., & Reeve, S. (1980). High country
adaptations. Plains Anthropologist, 25, 181197.
Ziegler, M., Simon, M. H., Hall, I. R., Barker, S., Stringer, C., & Zahn,
R. (2013). Development of Middle Stone Age innovation linked to
rapid climate change. Nature Communications, 4, 1905. doi:10.
14 Follow the Senqu: Mountain Dispersals 271
... AMEMSA's 14 C and OSL dates (pink), Bousman Index values (% C 3 , green), tree cover density ratio values (D/P, blue), and 13 C isotope values (red). Spits within contexts are designated by tick marks on the left (data from Stewart et al., 2016) Vegetation composition in the Lesotho highlands is largely determined by temperature, and thus correlates with altitude and aspect (Ehleringer et al., 1997;Patalano et al., 2023;Vogel, 1983). Higher altitudes host primarily C 3 plants, whereas lower altitudes favor C 4 -dominant communities (Vogel, 1983). ...
... Higher altitudes host primarily C 3 plants, whereas lower altitudes favor C 4 -dominant communities (Vogel, 1983). Melikane is surrounded primarily by C 4 vegetation today, but soil organic matter (SOM) δ 13 C values from the site suggest that C 3 grasses dominated the local landscape throughout the site's late Pleistocene sequence (Stewart et al., 2016). Four SOM and phytolith samples were taken from Melikane's ~27-23 ka levels at depth increments of 10 cm, corresponding to contexts 6-8:3, 6-8:1, 5:3, and 5:1 ( Fig. 4; Table 3). ...
... Four SOM and phytolith samples were taken from Melikane's ~27-23 ka levels at depth increments of 10 cm, corresponding to contexts 6-8:3, 6-8:1, 5:3, and 5:1 ( Fig. 4; Table 3). The ratio of woody to grass phytolith types (D/P ratio) falls to its lowest point in the entire sequence in samples MLK4 and MLK5 (contexts 6-8), reflecting a treeless, open grassland landscape (Stewart et al., 2016). Slightly higher D/P ratios in MLK 2 and MLK3 are still lower than values from most other occupation pulses (Stewart et al., 2016). ...
Full-text available
Melikane, a large sandstone rockshelter in the Maloti-Drakensberg Mountains of highland Lesotho, preserves an 80,000-year-old archaeological sequence including an occupation pulse dated to the onset of the Last Glacial Maximum (LGM), ~27–23 kcal BP. Paleoenvironmental proxies indicate that temperature depressions of ~6 °C below present values provoked changes in vegetation distribution around the site. The onset of the LGM also coincides with a global shift towards microlithization, expressed in southern Africa as the Later Stone Age Robberg bladelet industry. Bousman and Brink’s (Quaternary International 495:116–135, 2018) rapid replacement hypothesis asserts that this technocomplex was adopted nearly simultaneously across the subcontinent ~24 ka cal BP, replaci