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Chapter 13
Environmental Change, Ungulate Biogeography,
and Their Implications for Early Human Dispersals
in Equatorial East Africa
J. Tyler Faith, Christian A. Tryon, and Daniel J. Peppe
Abstract To better understand the potential role of envi-
ronmental change in mediating human dispersals across
equatorial East Africa, this study examines the biogeo-
graphic histories of ungulates, including a summary of
current knowledge and fossil evidence stemming from our
fieldwork in the Kenyan portion of the Lake Victoria basin.
Phylogeographic and paleontological evidence indicates that
vegetation changes across Quaternary climate cycles medi-
ated ungulate distributions and dispersals via the opening
and closing of biogeographic barriers in equatorial East
Africa. Dispersal capabilities would have been enhanced
during phases of grassland expansion and diminished during
phases of grassland contraction. We propose that the
distribution and dispersal of diagnostic technological mark-
ers in the archaeological record may be similarly influenced
by environmental changes. The Middle Stone Age record
from the Lake Victoria region provides intriguing examples
of possible environmentally mediated technological
dispersals.
Keywords Grasslands Lake Victoria Late Pleistocene
Middle Stone Age Paleoenvironments Phylogeography
Range shift
Introduction
The prehistory of early modern humans is characterized by
massive range expansions and population dispersals. Fossil
and genetic evidence indicate an African origin of our spe-
cies (Homo sapiens)*200 ka (McDougall et al. 2005;
Campbell and Tishkoff 2010; Brown et al. 2012; but see
Weaver 2012 for an alternate interpretation). These early
humans later diverged into multiple genetically (Campbell
and Tishkoff 2010) and morphologically diverse (Gunz et al.
2009) populations during the Late Pleistocene (126–12 ka),
likely reflecting intra-African population expansions and
dispersals (Soares et al. 2012). At least one of these popu-
lations migrated out of Africa *70 ka (Campbell and
Tishkoff 2010), setting the stage for the expansion of
humans across the globe and the replacement of all other
hominin species, including Neanderthals (Homo nean-
derthalensis) in Eurasia, and the enigmatic “hobbit”(Homo
floresiensis) in Indonesia.
East Africa features prominently in this history of early
modern humans, as it provides the earliest fossil remains of
Homo sapiens (Omo I and II from the Kibish Formation,
Ethiopia, at *195 ka; McDougall et al. 2005; Brown et al.
2012) and the probable source populations for their Late
Pleistocene dispersals out of Africa (Soares et al. 2012,2016).
Understanding the factors underlying these dispersals is the
subject of intense debate spanning the fields of archaeology,
paleontology, genetics, and climatology, among others
(Beyin 2011). One of the more common explanatory mech-
anisms includes climate-driven environmental change and its
effects on human demography and the opening and closing of
biogeographic barriers (e.g., Forster 2004; Finlayson 2005;
Vaks et al. 2007; Carto et al. 2009; Trauth et al. 2010;
Compton 2011; Blome et al. 2012; Eriksson et al. 2012;
Soares et al. 2012,2016). However, the precise role of envi-
ronmental change in East African human dispersals is poorly
understood, stemming from a lack of well-dated archaeolog-
ical sites associated with detailed paleoenvironmental
J.T. Faith (&)
School of Social Science, Archaeology Program, University of
Queensland, Brisbane, QLD 4072, Australia
e-mail: j.faith@uq.edu.au
C.A. Tryon
Department of Anthropology, Harvard University, Peabody
Museum of Archaeology and Ethnology, 11 Divinity Avenue,
Cambridge, MA 02138, USA
D.J. Peppe
Terrestrial Paleoclimatology Research Group, Department of
Geology, Baylor University, Waco, TX 76706, USA
©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_13
233
records. This is compounded by the high variability of the
regional Middle Stone Age (MSA), which limits our ability to
identify an archaeological signature of population movements
(Tryon and Faith 2013).
While many fundamental questions concerning the role of
environmental change in early human dispersals from East
Africa remain unclear, a growing body of evidence indicates
that Pleistocene environmental change played a central role
in the biogeographic histories of associated East African
faunas (e.g., Rodgers et al. 1982; Kingdon 1989; Marean
1992; Grubb et al. 1999; Wronski and Hausdorf 2008;
Lorenzen et al. 2012; Faith et al. 2013). Not only does this
imply that human population movements may be related to
broader biogeographic patterns, but also that these insights
into the relationships between changing climate, environ-
ments, and species ranges represent a useful starting point
for formulating new hypotheses about human dispersals. Our
aim here is to develop a framework for understanding the
potential role of environmental change in Late Pleistocene
human dispersals through an examination of ungulate bio-
geography. We focus on ungulates because they are well
represented in the fossil record and because many ungulate
species are clearly linked to specific habitats, making them
strong candidates for exploring how environmental change
mediates their biogeographic histories. This chapter provides
a brief review of current knowledge together with an
examination of paleontological and archaeological evidence
stemming from our ongoing fieldwork in the Kenyan portion
of the Lake Victoria basin.
Ungulate Biogeography
In common use, the ungulates represent a paraphyletic group
of hoofed mammals, including in Africa the antelopes and
buffalo (Bovidae), suids (Suidae), zebras (Equidae), giraffes
(Giraffidae), rhinos (Rhinocerotidae), elephants (Elephanti-
dae), and hippos (Hippopotamidae), among others. Ungulate
diversity in Africa is exceptional (Turpie and Crowe 1994;du
Toit and Cumming 1999), with the bovids (the most speciose
family) alone represented by 82 extant species (IUCN 2012)
and more than 100 fossil species (Gentry 2010).
The greatest diversity of African ungulates is found in
equatorial East Africa (Turpie and Crowe 1994), which
encompasses the boundaries between the Somalian, Suda-
nian, and Zambesian biogeographic regions for mammals,
birds, reptiles, amphibians, and plants (Linder et al. 2012)
(Fig. 13.1). These biogeographic regions, which are broadly
similar to previously identified zones based on expert
opinion (e.g., White 1983; Burgess et al. 2004), were sta-
tistically defined by Linder et al. (2012) using cluster anal-
ysis of species occurrences across 1°×1°(latitude and
longitude) cells for plants and vertebrate groups. We rec-
ognize that these zones are only a coarse measure of broad
biogeographic patterns, and we use them here as a heuristic
device to illustrate how biogeographic patterns changed in
the past. Some of the ungulates characteristic of these
regions include Grevy’s zebra (Equus grevyi) in the xeric
shrublands of the Somalian region, kob (Kobus leche) in the
edaphic grasslands of the Sudanian region, and wildebeest
(Connochaetes taurinus) in the secondary grasslands in the
Zambesian region. All of these taxa are found in East Africa,
but their ranges do not overlap, contributing to the high
spatial turnover (beta diversity) observed more broadly for
all East African mammals (Linder et al. 2012).
The main factors underlying these broad biogeographic
regions include the complex interactions between climate
and geomorphology (e.g., tectonics, topography, soil type)
and their influence on the distribution of vegetation com-
munities, which in turn mediate the distribution of faunas
(e.g., Bell 1982;O’Brien and Peters 1999; Linder et al.
2012). For example, a drier climate characterized by bimodal
rainfall distinguishes the Somalian region from both the
Sudanian and Zambesian regions, which are wetter and
receive primarily uni-modal rainfall (O’Brien and Peters
1999). This aridity in the Somalian region translates to the
presence of dry scrub vegetation inhabited by arid-adapted
ungulates, including Grevy’s zebra and African wild ass
(Equus africanus). At the same time, species ranges are also
influenced by geomorphological factors. In addition to
mediating vegetation cover and habitat suitability through
altitudinal gradients, tectonically driven uplift and rifting can
create dispersal barriers to biotic communities, as evidenced
Fig. 13.1 The distribution of Sudanian, Somalian, and Zambesian
biogeographic regions for mammals (after Linder et al. 2012)
234 J.T. Faith et al.
by the close correspondence between the boundary of the
Sudanian and Somalian regions in Ethiopia (Fig. 13.1) and
the location of the Ethiopian Rift Valley, which bisects the
Ethiopian highlands. Lastly, soil type can also determine
broadscale biogeographic patterns. For example, the domi-
nance of low-nutrient soils weathered from basement rock
accounts for the Miombo (Brachystegia) woodlands and
low-biomass ungulate communities characteristic of the
Zambesian region, in stark contrast to the nutrient-rich vol-
canic soils in some portions of the Somalian region, which
are associated with grasslands and high-biomass ungulate
communities (Bell 1982; East 1984).
Not only does the combination of climatic and geomor-
phological variables in East Africa translate to biogeo-
graphic zonation of ungulate species, but the same is also
true for populations within species. Lorenzen et al. (2012)
show that many broadly distributed species, including war-
thog (Phacochoerus africanus), African buffalo (Syncerus
caffer), hartebeest (Alcelaphus buselaphus), and giraffe
(Giraffa camelopardalis), among others, display genetic
substructuring that distinguishes between populations from
the Sudanian savannas, and those in East Africa and
southern Africa. Among some species there is also evidence
for substructuring between East and southern African pop-
ulations. The overall picture indicates that East Africa
encompasses the boundaries of genetically distinct lineages
from multiple regions.
A number of factors potentially contribute to this unique
taxonomic and genetic diversity. High spatial variability in
forage quantity and quality has been invoked as one possible
driver of ungulate species richness in East Africa (du Toit
and Cumming 1999), an argument supported by quantitative
analyses showing extreme spatial heterogeneity in East
African plant communities (Linder et al. 2012). However,
this does not explain the convergence of genetically distinct
lineages within species. In light of the phylogeographic
evidence, Lorenzen et al. (2012) propose that environmental
changes across late Quaternary glacial/interglacial cycles
played an important role in the taxonomic and genetic
diversity of East African ungulates. Equating glacial and
interglacial conditions with dry versus humid climates, they
suggest that the expansion of grassland vegetation during
glacial phases would have facilitated interchange of ungulate
populations throughout sub-Saharan Africa. During inter-
glacials, the expansion of tropical forests across equatorial
East Africa created a barrier that fragmented populations and
restricted gene flow (see also Cowling et al. 2008). If so,
then the distinct biogeographic regions and high spatial
heterogeneity of biotic communities found in East Africa
today (Linder et al. 2012) may be a Holocene phenomenon,
with a more homogenous supercommunity characteristic of
Pleistocene phases of grassland expansion.
The Fossil Record
Biogeographic evidence suggests that East Africa is a hub
around which ungulate ranges expand, contract, and frag-
ment across cycles of Quaternary environmental change.
One of the key predictions generated from this is that during
phases with expanded grassland cover, allopatric ungulate
species from both north and south of the equator will con-
verge in equatorial East Africa. The fossil record provides
the requisite empirical evidence for testing this hypothesis.
Compared to the record from southern Africa, which has
been a focus of modern human origins research for decades,
our understanding of the fossil history of East African
ungulates over the last *200 kyr is only beginning to come
to light (Marean and Gifford-Gonzalez 1991; Marean 1992;
Assefa 2006; Assefa et al. 2008; Faith et al. 2011,2012,
2013,2014). However, the emerging evidence provides
compelling examples of climate-driven range shifts that are
consistent with hypotheses derived from ungulate
biogeography.
The Records from Rusinga
and Mfangano Islands
Here we highlight some of the more prominent range shifts
documented in the fossil record, with an emphasis on the
Late Pleistocene ungulates from Rusinga and Mfangano
Islands in Kenya’s Lake Victoria basin (Fig. 13.2). We focus
on these sites because: (1) all of the ungulate taxa involved
in major Late Pleistocene range shifts are documented here
(and in some cases elsewhere); and (2) the Lake Victoria
region is strategically situated along the equator in East
Africa, with geological evidence documenting massive
expansions and contractions in lake area, including periodic
desiccation, in response to late Quaternary climate change
(Stager and Johnson 2008; Stager et al. 2002,2011; Tryon
et al. in press). Seismic profiles across Lake Victoria show at
least four desiccation surfaces (Johnson et al. 1996; Stager
and Johnson 2008), the most recent of which correspond to
extreme aridity during Heinrich Event 1 from 17 to 16 ka
and again during a subsequent dry phase from 15 to 14 ka
(Stager et al. 2002,2011); the other desiccation surfaces
correspond to undated phases of previous aridity and the
pre-lake surface. The vegetation of the Lake Victoria region
was historically characterized by bushland, thicket, and
forest (White 1983), although multiple lines of evidence
indicate that phases of lake contraction were associated with
expanded grasslands (Kendall 1969; Talbot and Laerdal
2000; Talbot et al. 2006; Tryon et al. in press).
13 Ungulate Biogeography and Human Dispersals 235
Details of our research on Rusinga Island and Mfangano
Island have been published elsewhere (Faith et al. 2011,2012,
2014; Tryon et al. 2010,2012,2014), and we provide only a
brief summary here. Rusinga and MfanganoIslands are located
within Lake Victoria (Fig. 13.2), with the former separated from
the mainland by a narrow channel and the latter situated
*10 km from the mainland. On both islands, the poorly con-
solidated Pleistocene deposits, known as the Wasiriya Beds
(Rusinga Island) and the Waware Beds (Mfangano Island), are
characterized by weakly developed paleosols and tuffaceous
fluvial sediments documenting a complex cut-and-fill system.
The age of the Wasiriya Beds is constrained tobetween 100 and
33 ka, with the minimum provided by radiocarbon dates for
intrusive gastropods and the maximum by geochemical analy-
sis of tephra deposits, which suggest derivation from East
African Rift System (EARS) volcanoes that began erupting
*100 ka (Blegen et al. 2015). The Waware Beds have a similar
minimum age, also provided by radiocarbon age estimates for
intrusive gastropods, whereas tephra correlations suggest a link
to the Wasiriya Beds and a comparable maximum age (Blegen
et al. 2015). The artifacts recovered from the Wasiriya Beds are
typologically MSA (Fig. 13.3), and the same is likely the case
for those from the Waware Beds, although a larger sample is
needed to be certain.
Rusinga and Mfangano Islands have yielded some of the
larger and better preserved fossil assemblages from this time
period in East Africa (Table 13.1). Alcelaphine bovids and
equids dominate the assemblages, indicating the presence of
open grassland vegetation distinct from the historic vegeta-
tion (White 1983). Several extinct bovids are present,
including Rusingoryx atopocranion,Damaliscus hypsodon,
Megalotragus sp., Syncerus antiquus, and an unnamed
impala, all of which are characterized by dental or postcra-
nial adaptations to grazing in open habitats (Klein 1980,
1994; Faith et al. 2011,2012,2014). The presence of large
gregarious grazers on Mfangano Island, some of which are
migratory species, suggests a likely connection to the
mainland when the deposits accumulated, requiring a ≥25 m
reduction in lake level (Tryon et al. 2014). The precipitation
decline needed to drive this would have set in motion a
series of feedback mechanisms leading to a substantial
reduction, if not complete desiccation, of Lake Victoria
(Broecker et al. 1998; Milly 1999), leaving behind a topo-
graphically smooth grassy plain (Tryon et al. 2014). As
detailed below, there is strong evidence that the expansion of
grasslands and reduction in surface area of Lake Victoria
facilitated interchange of ungulates from north and south of
the equator.
Fig. 13.2 The location of Rusinga and Mfangano islands in Kenyan Lake Victoria and nearby archaeological sites mentioned in the text
236 J.T. Faith et al.
Grevy’s Zebra (Equus grevyi)
Grevy’s zebra historically ranged throughout arid to
semi-arid grasslands and shrublands in the Horn of Africa
(Fig. 13.4) (Williams 2002). Fossil remains are known from
Rusinga Island (Table 13.1) and many other Middle-to-Late
Pleistocene sites in southern Kenya and northern Tanzania,
including Karungu, Lukenya Hill, Lainyamok, and Kisese II,
documenting a south and westward range shift or expansion
of at least 500 km (Faith et al. 2013). Although the species is
primarily confined to the Somalian biogeographic region, its
fossil occurrences are found in areas that are today part of
the Somalian, Sudanian, and Zambesian biogeographic
regions. At the onset of the Holocene, Grevy’s zebra dis-
appeared from southern Kenya and northern Tanzania and
Fig. 13.3 Middle Stone Age artifacts from Rusinga and Mfangano Islands: a–cbifacial points, d–eLevallois blades variably retouched,
fLevallois core
Table 13.1 The presence/absence of ungulate species from Rusinga and Mfangano islands as of our 2012 field season
Family Taxon Common name Rusinga Mfangano
Elephantidae Elephantidae cf. Loxodonta africana Elephant X –
Rhinocerotidae Ceratotherium simum White rhinoceros X –
Equidae Equus quagga Plains zebra X X
Equus grevyi Grevy’s zebra X –
Suidae Kolpochoerus sp.†Extinct bushpig X –
Phacochoerus sp. Warthog X X
Potamochoerus sp. Bushpig X X
Hippopotamidae Hippopotamus sp. Hippopotamus X X
Bovidae Taurotragus oryx Eland X –
Tragelaphus scriptus Bushbuck X –
Tragelaphus strepsiceros Greater Kudu X –
Tragelaphus cf. imberbis ?Lesser Kudu X –
Oryx beisa Oryx X –
Redunca fulvorufula/redunca Reedbuck X X
Redunca arundinum/Kobus kob Southern reedbuck/Kob X X
Aepyceros sp. nov.†Large extinct impala X –
Connochaetes taurinus Wildebeest X X
Damaliscus hypsodon†Small extinct alcelaphine X X
Megalotragus sp.†Giant wildebeest X –
Rusingoryx atopocranion†Extinct alcelaphine X X
Alcelaphini cf. Alcelaphus buselaphus Hartebeest X X
Sylvicapra grimmia Common duiker X –
Gazella thomsoni Thomson’s Gazelle X X
Syncerus antiquus†Long-horn buffalo X X
Syncerus caffer African buffalo X X
Ourebia ourebi Oribi X X
Oreotragus/Raphicerus Klipspringer/Steenbok –X
Madoqua sp. Dik-dik –X
†= extinct
13 Ungulate Biogeography and Human Dispersals 237
became rare in the fossil record, likely reflecting increased
precipitation, the contraction of dry grassland habitats, and
competition with mesic-adapted grazers (Faith et al. 2013).
Oryx (Oryx beisa)
The current distribution of oryx broadly corresponds to that
of Grevy’s zebra, but extends south into the Zambesian and
northwest into the Sudanian biogeographic regions
(Fig. 13.4). Oryx has a similar habitat preference as Grevy’s
zebra, and the two species are significantly associated in the
fossil record (Faith et al. 2013). The presence of oryx on
Rusinga Island (Table 13.1) indicates a *250 km westward
range shift or expansion. Like Grevy’s zebra, its range and
abundance declined throughout East African fossil sites at
the beginning of the Holocene (Faith et al. 2013).
White Rhinoceros (Ceratotherium simum)
The white rhinoceros includes two subspecies, the northern
white rhinoceros (C. simum cottoni) and southern white
rhinoceros (C. simum simum), although some authorities
Fig. 13.4 The geographic ranges of Grevy’s zebra (Equus grevyi),
oryx (Oryx beisa), white rhinoceros (Ceratotherium simum), kob
(Kobus kob), and southern reedbuck (Redunca arundinum) relative to
their fossil occurrences on Rusinga and Mfangano islands (star). Range
maps are from Williams (2002) for Grevy’s zebra and the IUCN Red
List (2012) for all other species. Boxes correspond to fossil occurrences
mentioned in the text: (1) Karungu, (2) Lainyamok, (3) Lukenya Hill,
(4) Kisese II, (5) Prolonged Drift, (6) Mumba shelter
238 J.T. Faith et al.
place the former in its own species (C. cottoni) (Groves et al.
2010). Their geographic ranges are discontinuous, with the
southern white rhinoceros historically known from southern
Africa and the northern white rhinoceros known from central
Africa northwest of Lake Albert (Fig. 13.4). Several white
rhinoceros fossils have been recovered from Rusinga Island,
*430 km southeast of its historic range (assuming they
belong to the northern taxon), with an even greater Late
Pleistocene range extension (*775 km) documented at
Mumba rockshelter in Tanzania (Mehlman 1989). As evi-
denced by an additional fossil occurrence from Prolonged
Drift in central Kenya, a small and probably isolated popu-
lation persisted in equatorial East Africa into the late
Holocene (Gifford et al. 1980). The cause of its prehistoric
disappearance from Kenya is not well understood, but could
include long-term vegetation change, competition for forage
and water with pastoralists and their livestock, or stochastic
processes owing to small population size and isolation.
Kob (Kobus kob) or Southern Reedbuck
(Redunca arundinum)
Both Rusinga and Mfangano Islands yield dental remains of
a medium-sized reduncine, larger than Bohor reedbuck
(Redunca redunca), but smaller than waterbuck (Kobus
ellipsiprymnus), that are morphologically similar to and
overlap in size with modern kob and southern reedbuck.
Both taxa have a similar habitat requirement for fresh
grasses and year-round access to water, which is consistent
with the fluvial nature of the fossil deposits. The ranges of
these two reduncines are allopatric (Fig. 13.4), with kob
occurring north and west of Lake Victoria in the Sudanian
savannas and southern reedbuck ranging from southern
Africa to just south of the Lake Victoria basin. Although it is
unclear whether the fossil specimens belong to kob, southern
reedbuck, or both, the occurrence of either implies a
*350 km range shift to the southeast (kob) or northeast
(southern reedbuck).
A Non-Analog Supercommunity
The convergence of various ungulate species with histori-
cally allopatric ranges on Rusinga and Mfangano Islands
supports the notion of a homogenous East African super-
community during Pleistocene phases of grassland expan-
sion. It follows that the distinct biogeographic zonation
found in East Africa today (Fig. 13.1; Linder et al. 2012)is
ephemeral—probably limited to the Holocene—and closely
linked to climate and its effects on vegetation structure.
Other variables that contribute to the establishment of bio-
geographic zonation in East Africa (e.g., tectonics, topog-
raphy, and soil type), unquestionably contributed to Late
Pleistocene vegetation structure, but these are expected to be
relatively stable over the timescale examined here, leaving
climate as the most likely driver of first-order vegetation
change.
While the non-analog supercommunity is consistent with
enhanced ungulate dispersal capabilities related to expanded
grasslands and a smaller Lake Victoria, it is also possible
that environments at the time were highly productive and
uniquely suited to supporting ungulate diversity. Across a
range of East and southern African ecosystems, ungulate
diversity and biomass peaks between 700 and 800 mm
annual precipitation (Coe et al. 1976; East 1984; Faith
2013). A decline in annual precipitation from the *1000 to
1200 mm/yr observed today, which is supported by the
inferred decline of Lake Victoria (Tryon et al. 2014), could
further underpin the diversity of the Late Pleistocene
ungulate community. Reduced atmospheric CO
2
concentra-
tions may have also contributed to the diversity, by
enhancing foraging nutrient content, in turn supporting
higher biomass large herbivore communities (Faith 2011).
The presence of extinct grazing bovids characterized by
massive body mass (Syncerus antiquus and Megalotragus
sp.) or extreme hypsodonty (Damaliscus hypsodon,Rusin-
goryx atopocranion, and the unnamed impala), of which D.
hypsodon and R. atopocranion are dominant, also implies
the presence of non-analog environments. We have previ-
ously interpreted their presence, together with that of Gre-
vy’s zebra and oryx, as evidence for aridity (Tryon et al.
2010,2012; Faith et al. 2011,2013), although this is at odds
with the high diversity (see Faith 2013). A more detailed
exploration of this conflict is beyond the scope of this study,
but possible explanations could include the presence of a
complex Late Pleistocene grazing succession (e.g., Brink
and Lee-Thorp 1992) or an extinct migratory system (e.g.,
Marean 2010; Faith and Thompson 2013).
Climatic and Tectonic Drivers
Range shifts documented on Rusinga and Mfangano Islands
illustrate the importance of vegetation change, particularly
the expansion of grassland vegetation, in the biogeographic
histories of ungulate populations. The mechanisms respon-
sible for the expansion and contraction of grassland vege-
tation in equatorial East Africa during the Pleistocene
include moisture availability, atmospheric CO
2
concentra-
tions, and tectonically driven topographic shifts. The com-
plex interaction of these variables would have played an
important role in the opening and closing of dispersal
13 Ungulate Biogeography and Human Dispersals 239
corridors for ungulate populations, and we provide a sum-
mary of these mechanisms below.
Changes in moisture availability are sometimes equated
with glacial versus interglacial climates, which are typically
generalized as reflecting more arid (glacial) versus more
humid (interglacial) conditions. However, although changes
in global moisture availability across glacial/interglacial
cycles may influence equatorial East African climate, we
caution that this oversimplification ignores the complexities
of regional climate dynamics. Moisture availability in
equatorial East Africa is driven by the complex interplay
between orbital forcing, the position of the Intertropical
Convergence Zone (ITCZ), high-latitude climate events,
tropical ocean temperature gradients, and regional tectonics
(e.g., DeMenocal 1995,2004,2011; Trauth et al. 2003,
2005,2007,2009; Feakins et al. 2005,2013; Sepulchre et al.
2006; Verschuren et al. 2009; Blome et al. 2012). In addition
to these global and regional drivers, local factors may also
play an important role in vegetation change. For example,
today as much as 90% of water loss in Lake Victoria is due
to evaporation and 80% of the input is from direct precipi-
tation (rather than inflow from streams and rivers) (Piper
et al. 1986; Crul 1995), such that precipitation change plays
a key role in lake size and moisture availability (Broecker
et al. 1998; Milly 1999). This sensitivity to precipitation
means that small changes in local rainfall patterns could
translate to more pronounced aridification and vegetation
change in the Lake Victoria region compared to other areas
of East Africa.
In addition to moisture availability, reduced atmospheric
CO
2
concentrations during Pleistocene glacial phases may
also facilitate the expansion of grasslands due to the com-
petitive advantage of C
4
plants under such conditions
(Prentice et al. 2011). For example, the paleoenvironmental
record from Lake Challa on the Kenya/Tanzania border
shows that both humid and arid intervals from 12 to 25 ka
were dominated by C
4
vegetation, most of which are prob-
ably grasses, whereas C
3
vegetation only expands at the
onset of the Holocene (Sinninghe Damstéet al. 2011). This
may indicate that the past distribution of grassland habitats
in equatorial East Africa is more sensitive to atmospheric
CO
2
than to precipitation.
Over longer timescales (>100 kyr) through the Pleis-
tocene, changes in ungulate ranges may also have been
influenced by regional tectonics. Extension and uplift asso-
ciated with the EARS has dramatically altered the East
African landscape since the Oligocene by creating large rift
lakes such as Lakes Malawi, Tanganyika, Kivu, Albert and
Edwards, causing the opening and closure of connections
between major lakes, changing flow directions of major
rivers, and creating significant differences in topography
(e.g., Rosendahl 1987; Chorowicz 2005). Lake Victoria,
though not a true rift lake, was created when extension and
uplift of the western and east arms of the EARS caused
backponding of rivers into a topographic low between the
two branches of the rift *1.6–0.4 Ma (e.g., Kent 1944;
Bishop and Trendall 1967; Ebinger 1989; Johnson et al.
1996; Talbot and Williams 2009). After its formation, con-
tinued rifting caused the lake basin to shift as much as 50 km
eastward (Doornkamp and Temple 1966; Stager and John-
son 2008). Through the creation of rift-influenced lakes and
significant topographic variability across the landscape, the
rifting of the EARS may have influenced ungulate biogeo-
graphic histories by opening and closing potential dispersal
corridors. Although precipitation and atmospheric CO
2
concentrations are the primary drivers of vegetation change,
tectonically driven topographic changes probably also con-
tributed to habitat fragmentation and variability across East
Africa, which could have affected the ranges of ungulate
taxa throughout the Pleistocene.
There is little doubt that climate and tectonics strongly
influence East African paleoenvironments during the Pleis-
tocene. However, as emphasized by Blome et al. (2012),
determining the precise nature of environmental changes and
their underlying mechanisms in the Lake Victoria basin and
across East Africa will require improved chronological
control and more refined paleoenvironmental records.
Implications for Early Human Dispersals
Biogeographic evidence suggests that climate-driven vege-
tation changes across equatorial East Africa played an
important role in mediating the distribution of ungulate
species, with the expansion of grasslands facilitating inter-
change across East Africa (Lorenzen et al. 2012). This is
corroborated by the fossil records from Rusinga and Mfan-
gano Islands, which show that ungulate species with his-
torically allopatric ranges converged in equatorial East
Africa at a time when Lake Victoria was much reduced, if
not desiccated, and grasslands were widespread (Tryon et al.
2014). Presumably, the loss of grasslands and expansion of
the equatorial forest belt at the onset of the Holocene and
during previous humid intervals with high atmospheric CO
2
concentrations (Cowling et al. 2008) would have prompted
range shifts and fragmentations in species distributions
approaching those observed historically. What are the
implications of these patterns for early modern human
dispersals?
240 J.T. Faith et al.
In the Paleolithic archaeological record, diagnostic lithic
technology represents one of the few means of tracking
human dispersals. On its own, the extent to which the
movement of technological markers reflects the dispersal of
human populations (and their genes) is uncertain, but it at
least documents the dispersal of human behavioral reper-
toires. There is abundant ethnographic evidence showing
that the subsistence behaviors and associated technologies of
historic hunter-gatherers are very closely linked to the
environment (Oswalt 1973; Kelly 1995; Binford 2001;
Collard et al. 2005), in which case it is reasonable to suppose
that the movement of technological markers linked to
specific environments might parallel the broader patterns
observed in the ungulates. Below we explore some possible
links between the distributions of technologies in the Lake
Victoria region in relation to paleoenvironmental change.
The Lake Victoria MSA
Previous discussions of distinct technological markers from
the Lake Victoria region have focused on the Lupemban
industry, which represents one of the more distinct MSA
regional variants, characterized by the presence of large (>10
cm), thin, bifacially flaked lanceolate points. These lanceo-
lates are well known from central Africa (Barham 2000;
Mercader 2002), although their distribution extends to the
eastern margins of Lake Victoria, including near-shore
islands (Leakey and Owen 1945; Nenquin 1971; McBrearty
1988; Tryon et al. 2012; Taylor 2016). Where paleoenvi-
ronmental records are available, the Lupemban is typically
associated with forested vegetation (Barham 2000; Mercader
2002; Taylor 2016).
The vegetation of the Lake Victoria region today repre-
sents a mosaic of endemic taxa from neighboring regions
(White 1983), attesting to the past expansion and contraction
of vegetation communities, a model consistent with the
implications of the ungulates from Rusinga and Mfangano
Islands (Table 13.1), the distribution of modern faunas (e.g.,
Kingdon 1989; Wronski and Hausdorf 2008), and
paleo-vegetation models (e.g., Cowling et al. 2008). To the
extent that the distribution of Lupemban artifacts is tied to
forested paleoenvironments, we expect that its easternmost
occurrences in the Lake Victoria basin to be associated with
an expanded equatorial forest belt. As shown in Fig. 13.5, all
of the Lupemban sites in the eastern portion of the Lake
Victoria basin lie along now submerged portions of major
rivers that traversed the basin prior to peak lake size some-
time during the Middle or Late Pleistocene (Temple 1966;
Scholz et al. 1998). These sites represent the easternmost
limits of the Lupemban, and include Muguruk on the Winam
Gulf in Kenya (McBrearty 1988), sites reported by Leakey
and Owen (1945) in the Yala River valley of Kenya, and on
Bugaia Island in Uganda (Nenquin 1971). The nearby rivers
would have likely provided naturally wooded corridors
facilitating the expansion of forest-adapted taxa and tech-
nologies, perhaps including the Lupemban, around the
margins of Lake Victoria.
The MSA assemblages from the Wasiriya and Waware
Beds of Rusinga and Mfangano Islands are typologically
distinct from Lupemban assemblages in the Lake Victoria
region or elsewhere, which is not surprising given the asso-
ciated grassy paleoenvironment. In light of the evidence from
ungulate biogeography, it is possible that the expansion of
grasslands would facilitate the movement of people or tech-
nological traditions associated with these environments from
broad East African regions north or south of the equator. In
previous studies (Tryon et al. 2012; Tryon and Faith 2013),
we identified one possible example of this. The lithic
assemblages from Rusinga and Mfangano Islands (Fig. 13.3)
include very small (2–4 cm in length) bifacially flaked points
that overlap in size with those from MSA sites throughout the
East African Rift System. These include the Late Pleis-
tocene MSA or MSA/LSA assemblages from Nasera Rock-
shelter in Tanzania (*290 km southeast), site GvJm-16 at
Lukenya Hill in Kenya (*340 km east southeast), and
Aduma in Ethiopia (*1,390 km northeast). At all of these
sites, the associated faunas or other paleoenvironmental
records indicate grassland vegetation comparable to that
inferred for Rusinga and Mfangano Islands, raising the pos-
sibility that these small points represent either a human
population or (more likely) a technology associated with Late
Pleistocene tropical grasslands. This possible connection to
Rift Valley sites is further supported by the rare presence of
obsidian from the Wasiriya Beds, the nearest outcrops of
which are from Rift Valley sources *250 km to the east.
The small MSA points found on Rusinga and Mfangano
Islands and in the Rift Valley are rare to the west of Lake
Victoria, reported only from Kibwera in Tanzania (Reid and
Njau 1994), although unfortunately no illustrations or pho-
tographs are provided (Fig. 13.5). Assuming that points and
other artifacts produced by hunter-gatherers are to some
extent a reflection of the environment in which they were
used, the geographic limits of different types of MSA point
(small points versus Lupemban lanceolates) along the mar-
gins of present-day Lake Victoria (Fig. 13.5) is consistent
with the hypothesis that this area is a nexus of environmental
changes that facilitated the movement of people and their
technological markers.
13 Ungulate Biogeography and Human Dispersals 241
Conclusions
Biogeographic evidence provides a compelling argument
that environmental changes across late Quaternary climate
cycles mediated the distributions and dispersals of ungulate
species via the opening and closing of barriers and dispersal
corridors in equatorial East Africa. For a broad range of
savanna ungulates, dispersal capabilities would have been
enhanced during phases of grassland expansion and dimin-
ished during phases of grassland contraction, leading to
repeated range expansions and fragmentations.
These patterns are of importance to human origins
research, as genetic evidence documents multiple East
African dispersals of early modern humans during the Late
Pleistocene, both within and out of Africa (Campbell and
Tishkoff 2010; Soares et al. 2012,2016). We propose that,
as is clearly the case for ungulates, the distribution and
dispersals of diagnostic technological markers in the
archaeological record may also be mediated by environ-
mental changes. The archaeological record from the Lake
Victoria region provides some intriguing examples of pos-
sible environmentally mediated technological dispersals, but
it is clear that a better sampled archaeological record and
improved chronology is crucial to fleshing out these patterns.
Although such limitations are an ever-present problem that
can only be resolved through further fieldwork, the patterns
explored here provide an initial framework for exploring the
dispersal of early human populations migrations and
expansions during a critical stage in human evolution.
Acknowledgments We thank Sacha Jones and Brian Stewart for
inviting us to contribute to this volume and the many collaborators who
have worked with us in the Lake Victoria basin, including Emily
Beverly, Nick Blegen, Steve Driese, David Fox, Niki Garrett, Zenobia
Jacobs, Kirsten Jenkins, Renaud Joannes-Boyau, Cara Roure Johnson,
Kieran McNulty, Sheila Nightingale, David Patterson, and Alex Van
Fig. 13.5 Buried river channels across the Lake Victoria basin,
simplified after Temple (1966). Note the proximity of the easternmost
Lupemban lanceolate sites to major east-west flowing rivers that
traverse the basin. Smaller MSA points are not reported in the western
portion of the Lake Victoria basin except at Kibwera (illustrations
unavailable). Artifact sketches after Leakey and Owen (1945), Nenquin
(1971), and McBrearty (1988), and illustrated at same scale
242 J.T. Faith et al.
Plantinga. We also thank two anonymous reviewers and Sally Reynolds
(reviewer) for helpful comments on a previous version of this chapter.
Fieldwork was conducted under research permits
NCST/RCD/12B/012/31 issued to JTF and NCST/5/002/R/576 issued
to CAT and an exploration and excavation license issued by the
National Museums of Kenya (NMK). Our fieldwork is made possible
through the support of the NMK and the British Institute in East Africa
and with financial support from the National Science Foundation
(BCS-0841530, BCS-1013199, BCS-1013108), the Leakey Founda-
tion, the National Geographic Society, the University of Queensland,
New York University, and Baylor University. Lastly, none of this
would have been possible without the support of Cornel Faith, Rhonda
Kauffman, Violet Tryon, and Sholly Gunter.
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