Journal of Archaeological Science 38 (2011) 312e322

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Journal of Archaeological Science

journal homepage: http://www.elsevier.com/locate/jas

4500-Year old domesticated pearl millet (Pennisetum glaucum) from the Tilemsi
Valley, Mali: new insights into an alternative cereal domestication pathway
Katie Manning a, *,1, Ruth Pelling b, Tom Higham c, Jean-Luc Schwenniger c, Dorian Q. Fuller d
a
St Hughs College, University of Oxford, St Margarets Road, Oxford OX2 6LE, United Kingdom
b
Fort Cumberland, Fort Cumberland Road, Eastney, Portsmouth PO4 9LD, United Kingdom
c
Research Laboratory for Archaeology, Dyson Perrins Building, South Parks Road, Oxford OX1 3QY, United Kingdom
d
Institute of Archaeology, University College London, 31-34 Gordon Square, London WC1H 0PY, United Kingdom

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

Article history: We report here new evidence from the Lower Tilemsi Valley in northeastern Mali, which constitutes the
Received 7 April 2010 earliest archaeobotanical evidence for domesticated pearl millet (Pennisetum glaucum), predating other
Received in revised form nds from Africa or India by several centuries. These materials provide further morphological details on
1 September 2010
the earliest cultivated pearl millet. Our results demonstrate that pearl millet non-shattering evolved
Accepted 3 September 2010
earlier than the start of grain size increases and that once domesticated, pearl millet spread widely and
rapidly. Additional attention is given to the dating of these materials, highlighting potential aws in the
Keywords:
use of organic chaff tempered pottery to date occurrences of pearl millet. A revised chronology, based on
Pearl millet
Domestication
detailed Bayesian modelling, is presented for the Lower Tilemsi region.
Chaff-temper 2010 Elsevier Ltd. All rights reserved.
Agro-pastoralism
Tilemsi Valley

1. Introduction
Pennisetum glaucum, pearl millet, is a staple cereal of sub-Saharan
Africa and parts of India, where it is tolerant of the drier Sahelian/semi-
desert zones as well as thriving in savannas (Brunken, 1977; Brunken
et al.,1977; Harlan, 1992; Tostain, 1998). It is the only African cereal for
which existing archaeobotanical evidence is adequate for providing
some quantitative assessment of the domestication process: in
particular grain measurements from a series of sites in Africa and India
over time provides indications of grain size change under cultivation
(Fuller, 2007). On the basis of these data it has been suggested that the
domestication process of millet differed from that of other cereals,
such as wheat, barley and rice, as grain size increase appeared to be
delayed until after domestication rather than occur before or during
domestication (as dened by the evolution of non-shattering forms)
(Fuller, 2007; Kahlheber and Neumann, 2007). Archaeobotanical
* Corresponding author. 2 Ty Glan Conwy, Padog, Betws y Coed, Conwy LL24 0ST,
United Kingdom. Tel.: 44 (0) 7932 508280.
E-mail addresses: kat_mng@yahoo.co.uk (K. Manning), ruth.pelling@googlemail.
com (R. Pelling), thomas.higham@rlaha.ox.ac.uk (T. Higham), jean-luc.schwenninger@
rlaha.ox.ac.uk (J.-L. Schwenniger), d.fuller@ucl.ac.uk (D.Q. Fuller).
1 for early pearl millet in the Lower Tilemsi Valley, and if we are to
Present address: Institute of Archaeology, University College London, 31-34
Gordon Square, London WC1H 0PY, United Kingdom.
evidence from sub-Saharan West Africa suggests that pearl millet was
the predominant, or even the only, cultivated cereal across the region
including Mauretania, Mali, Ghana, Burkina Faso, and Cameroun (see
Neumann, 2005; DAndrea and Casey, 2002; Klee et al., 2004; Fuller
et al., 2007a). However, the date on these nds was never older than
the early Second Millennium BC, by which time domesticated pearl
millet had already spread to India, raising the likelihood that its origins
lay in earlier centuries (in the Third Millennium BC) somewhere in
West Africa (Fuller, 2003; Neumann, 2005: 258e259).
We report here new evidence from the Lower Tilemsi Valley in
Mali, which constitutes the earliest archaeobotanical evidence for
domesticated pearl millet predating other nds from Africa or India
by at least a few centuries. In the course of test excavating a sample of
Lower Tilemsi sites, samples were taken for the extraction of charred
plant remains, yielding only a few millet grains, whilst vegetable
tempered sherds were examined and found to contain a high
concentration of Pennisetum chaff. Antiquity of these remains was
conrmed by direct AMS dates on grains, on the organic fractions of
potsherds, and by comparative OSL dates on some sherds. It is often
the case in Saharan and sub-Saharan Africa that only one or two
independent AMS dates are presented in association with early pearl
millet remains. This study presents a detailed chronological sequence
accept the single-point dates at face value, domesticated pearl millet

0305-4403/$ e see front matter 2010 Elsevier Ltd. All rights reserved.
doi:10.1016/j.jas.2010.09.007
 K. Manning et al. / Journal of Archaeological Science 38 (2011) 312e322 313

Fig. 1. Map showing location of the Tilemsi valley in Mali and the sites sampled archaeobotanically in this study.

is conrmed in the mid-Third millennium BC. Bayesian modelling of
the chronological data, undertaken by TH and J-LS however, suggests
the AMS dates on the organic fractions of potsherds may be too old by
a few centuries, highlighting an important caveat when assessing the
temporal distribution of early pearl millet remains. These materials
also provide further morphological details on the earliest cultivated
pearl millet, offering insights into the evolution of the domestication
traits in this species, conrming that they differ in ordering from
better documented crop species such as wheat and barley.
The Tilemsi Valley is of key interest for the study of agricultural
development in sub-Saharan Africa. During the mid- to late Holo-
cene the Tilemsi provided a hydrological corridor between the
Saharan zones of West Africa, where the wild progenitors of pearl
millet are reported (Harlan, 1992; Tostain, 1998; Fuller, 2003), and
the West African Sahel. Enzyme similarity datasets have also fav-
oured southeast Mauretania (esp. Tostain, 1998) as a centre for
pearl millet domestication and/or a stretch from northeast Mali to
Lake Chad (see Fuller, 2003; Oumar et al., 2008), located close to the
northern terminus of the Tilemsi Valley. Early archaeological
eldwork in the Lower Tilemsi region reported the presence of
domesticated Pennisetum in pot sherd impressions from the site of
Karkarichinkat Sud (Smith, 1992: 74); then dated to c. 2000 BC on
the basis of stylistic association with stratigraphically unrelated
radiocarbon dates. The absence of larger scale, systematic archae-
obotanical sampling or direct-dating, however, has meant that,
until now, these nds were hard to substantiate, contributing little

Table 1
A summary of the presence of diagnostic elements of Pennisetum sp. in casts examined by SEM. Shown are the counts of presence of the element on examined casts from each
site/context. **Rachis elements indicate the presence domesticated millet. Paired spikelet* indicated in notes also suggested domesticated millet. Possible wild involucre base
is indicated in notes. For full details see Supplementary Online Material.

Site name Context no. Involucre base Rachis** Bristle Uniseriate hairs Spikelet Lemma/palea frag. Grain (?) Notes
w/bristles
KN05 1 0 0 1 0 0 1 0
77 1 0 3 0 2 2 0
JS07-2 107 1 0 1 0 1 0 0
110 5 1** 3 1 2 1 0 paired spikelet(?)*
112 3 1** 1 0 3 1 0
EN07 80 5 1** 3 0 3 0 0 possible wild;
paired spikelet*
81 2 1** 4 2 1 3 1
82 1 1** 4 2 1 2 0 paired spikelet*
EB07 3 2 6** 8 5 10 2 1 paired spikelet*
5 0 0 2 1 1 0 0
Total % 24 11 27 10 23 11 2
39% 18% 44% 16% 38% 18% 3%
314 K. Manning et al. / Journal of Archaeological Science 38 (2011) 312e322

Table 2
Archaeobotanical macroremains recovered from otation in the Tilemsi Valley project.

Site Context Sample Pennisetum sp. Panicoid Indet. Poaceae Zizyphus Celtis sp. Indet. fruit Isolepis type
volume (litres) grass caryopses sp. (stone) stone
KN05 30 5 1 e e e e e e
18 5 1 e e e e e e
85 8 e e 1 e e e e
93 2 e e e 3 e e e
102 3 e e 1 e e 1 e
77 5 e e e e 5 1 e
47 2 e e 1 e e 1 e
166 6 e 1 e e e 2 e
EB07 2 5 e e e e 2 e e
3 5 e e e e 2 e e
3 20 e e e e 13 e e
TB07 9 3 e e e 3 e e e
10 20 e e e 5 e 1 e
11 3 e e e e 11 e e
15 10 e e e 7 e e e
16 3 e e e e 1 e e
TA07 53 0.5 e e e 1 e 1 e
53 5 e e e e e 1 1
EN07 81 5 e e e e 100 e e
82 5 e e e e 100 e e
85 5 e e e e 10 e e
86 5 e e e e 21 e e
38 5 e e e 2 35 e e
90 2 e e e e 3 e e
91 3 e e e e 1 e e
JS07-1 99 5 e e e 1 e e e
100 5 e e e 4 e 1 e
JS07-2 108 0.5 e e e e 1 e e
110 5 e e e e 14 e e
111 5 e e e e 12 e e
112 5 e e e e 5 e e

to our understanding of the earliest history of this cereal. The
current study conrms the presence of domesticated pearl millet,
providing direct AMS and OSL dates on these nds, and expands the
known distribution of early pearl millet remains in this region.
2. Archaeobotany of the Tilemsi Valley project
In 2005 the Lower Tilemsi Valley Project was initiated by KM in
an attempt to rene the chronology, and investigate the emergence
of agro-pastoral communities in this region. Over the course of two
eldwork seasons in 2005/2006 and 2007 a total of 86 multi-period
sites were identied, including 65 occupation mounds and surface
scatter sites which, on the basis of surface nds, appear to date to
the Late Stone Age (LSA) between c. 3000 and 2000 BC, three
tumuli sites, also seeming to date to the LSA, and 17 iron working
sites dating to the mid- 1st millennium AD (Manning, 2008a). The
rst eld season involved a sustained excavation program at the
site of Karkarichinkat Nord (KN05 [see Manning, 2008b and
Finucane et al., 2008a for a more detailed description of the KN05
excavations and site plan]). Excavations at Karkarichinkat Sud
(KS05) were unfortunately abandoned due to the unconsolidated
nature of the deposits. The 2007 eld season was, therefore, dedi-
cated to test excavation at a sample of neighbouring occupation
mounds identied in the preceding survey (Fig. 1). All of the test-
excavated sites have been dated to between 2800 and 1800 BC with
the majority of dates clustering between 2400 and 2200 BC.
Bayesian modelling of all AMS and OSL dates, however, suggests
two possible phases of occupation: the rst represented at KN05
and possibly TB07, TA07 and JS07-1, dating to c. 2500 BC, and a later
phase of occupation represented at EB07, EN07 and JS07-2, dating
to c. 2000 BC. According to the chronological models for KN05,
JS07-2 and EN07 (see section 3.1 below and SOM), these sites
appear to have been occupied for less than 100 years each.
Overall, the faunal assemblage indicates a much wetter envi-
ronment than today, representing a Sahelian riparian environment,
possibly with gallery fringe forests and a likely rainfall of
>250e300 mm. Aquatic animals, including sh, crocodile and
freshwater turtle attest to an active hydrological system, although
osteometric reconstruction of certain sh species suggests the
principal channel of the Tilemsi River was already in decline by the
start of occupation (Manning, 2008a).
Bulk samples were taken from each site, trench and layer exca-
vated, with volumes ranging from 5 g to 20 L, but usually 5 L. Samples
were processed in the eld by manual water otation (wash-over
bucket otation) and ots collected on either a 250 micron or 500
micron mesh depending on the compaction of the soil matrix. Dried
ots were brought back to the UK and sorted by RP under a binocular
microscope at 10e20 magnication. Any seeds or chaff were
extracted and identied on the basis of morphological criteria and by
comparison with reference material held by the author or in the
Institute of Archaeology, University College London. Results are
discussed by site in the SOM and summarized in Table 1, which
includes only those samples which produced plant remains.
In addition to sampling for macro plant remains on site, analysis
of the pottery assemblage revealed that a large number of sherds
were tempered with chaff, constituting 55% of the EB07 assemblage
(N 494), 22% of the EN07 assemblage (N 165) and 37% of the JS07-
2 assemblage (N 181). At TB07 and TA07, meanwhile, chaff
tempering was absent and contributed only 0.7% of all sherds from
KN05 and 2.4% of all sherds from JS07-12. Forty potsherds were
sampled (Table 1) to assess the chaff component. A cast of the chaff
2
JS07 comprised 21 discrete mounds of surface scatter, each separated by
approximately 5m of sterile sand. Two of these mounds were test-excavated (JS07-
1 and JS07-2) in order to assess differential formation processes.
 K. Manning et al. / Journal of Archaeological Science 38 (2011) 312e322
Fig. 2. Image of Pennisetum seed recovered from context (018), KN05.
impression was made using a vinyl polysiloxane dental moulding
agent, which was applied to the surface of the pot sherd and left to
dry for approximately 10 min. Multiple casts were taken from each
sherd and from each impression site in order to assess the range of
threshing by-products in the clay matrix and to guarantee a clean
and anatomically detailed cast. These were then re-examined under
a stereomicroscope at 10e40 magnication and the most
promising in terms of anatomical details and morphological pres-
ervation were selected and mounted on metal stubs and sputtered
coated with gold for examination by Scanning Electron Microscopy,
with selected views saved as micrographs. Out of the original sample
19 of the sherds yielded promising inclusions. Some of the larger
sherds generated more than one impression site, making a total of 27
samples that were selected for SEM examination. The sampled
potsherds with unequivocal domesticated pearl millet chaff temper
were subsequently dated using a comparative AMS/OSL technique
(see below) in order to obtain a direct date on the sample.
3. Results
Seeds and fruit remains were present in small quantities at all
the sites sampled (Table 2 and SOM). Grass caryopses were found
Fig. 3. Diagram of domesticated millet ear and spikelet indicating parts.
315
only at Karkarichinkat Nord (KN05), which included Pennisetum sp.
grain morphology resembling the wild species, i.e. the grain lacked
the club-shape suggested as diagnostic based on modern compar-
isons (Fig. 2) (DAndrea et al., 2001; Zach and Klee, 2003), which has
signicant implications for the evolution of the domestication
syndrome in pearl millet (see below). Stones of Celtis sp., some-
times in large numbers, and occasionally of Zizyphus sp., suggest
some sort of harvesting and use of the fruits. These are common
species on archaeological sites across the Sahelian zone and
southern Sahara where few other remains survive. It is likely that
the general paucity of material at these sites is a reection of
preservation and particularly wind erosion, which has left only the
heavier fruit stones.
Sixty-one pearl millet fragments were identied in the SEM, and
can be divided into six plant part categories, (Table 2 and SOM).
These include the presence of involucre bases from which bristles
arise, the presence of the rachis which occurs below this bristle
base (and indicates a domesticate), bristle fragments (and a sub-
category of fragments with unicellular hairs on the bristle surface),
spikelets (i.e husk including lemma and/or palea) and smaller husk
fragments (Figs. 3 and 4). The most widespread, noted in 38% of the
casts were fragments of pearl millet bristles and of these more than
a third (10 of 27) preserved evidence of rows of unicellular
trichomes on their surface. This is likely to be an underestimate, as
many casts which only showed small bristle fragments were not
subjected to SEM investigation, and indeed based on this supercial
examination of the sherds it is probable that bristle fragments are
nearly ubiquitous. Spikelet impressions were noted on 38% but if
smaller fragments of husk are included this goes up to 54%. Invo-
lucre bases, with the base of the bristle clusters were noted in 38%
of the studied casts, but unfortunately only one third of these (9 of
24) showed evidence for the base, with either a rachis (8 cases) or
possible smooth scar of the wild type (1 case). A single impression
of a grain was noted (Fig. 4A). While the size of this grain
(3  1.7 mm) is slightly larger than expected for wild-type millet, it
should be kept in mind that if the grain was wet when added to the
clay its size may have expanded somewhat, and the metrics of this
one grain impression do not provide good evidence that grain size
increase had occurred by this date (see below).
The spikelet impressions were examined for evidence of paired
spikelets. Wild Pennisetum, normally has a single grain in each
bristly involucre, while domesticated forms often have multiples
grains. The study by Godbole (1925) of Indian pearl millet suggests
316 K. Manning et al. / Journal of Archaeological Science 38 (2011) 312e322
Fig. 4. SEMs of selected impression types.
w70% of involucres include two spikelets (each with a grain), while
w20% are single grained. The other w10% includes more than two
grains, with as many as 9 grains reported from a single involucre.
Archaeologically, early impressions of pearl millet indicate the
presence of paired spikelet, such as those previously reported from
Dhar Nema, Mauretania (see Fuller et al., 2007b). Four examples of
a pair of such spikelets were noted in the Tilemsi material (e.g.
Fig. 4B and C). The generally highly fragmentary nature of the chaff
impressions in this material does not permit quantication of the
proportion of single-spikelet versus paired spikelet forms.
However, the presence of this form does suggest that this feature
had evolved in the cultivar.
Eight involucre base impressions with preserved rachis frag-
ments were identied indicating the stalked, non-dehiscent mor-
photype of the domesticate (Fig. 4D and E), of which one was
ambiguous (Fig. 4B). These are distributed across three sites and six
contexts EB07 (3), JS07-2 (112) and (110), EN07 (80), (81) and (82).
Only one possible wild type involucre base was noted (Fig. 4F) from
JS07-2 (107), although this remains somewhat ambiguous. Despite
this sample size being small it suggests the predominance of the
domesticated form (i.e. 89% of preserved rachis remains). Based on
the evidence from other cereals that non-shattering evolved grad-
ually, on the order of 1000e2000 years (Fuller, 2007; Fuller et al.,
2009; Fuller and Allaby, 2010), we would infer cultivation to have
begun perhaps in the Fourth Millennium BC. This would imply that
when EB07, EN07 and JS07-2 were settled, the incoming population
brought with them an established economic suite, including
domesticated millet and domesticated cattle and ovicaprines.
3.1. Radiocarbon AMS and OSL-dating
Dating for the Tilemsi project was based largely on AMS radio-
carbon dates, as well as some optically stimulated luminescence
dates (OSL) on ceramics and sediments (Table S12, S13 and S14 in
SOM). For Karkarichinkat Nord,11 AMS dates are available, including
one direct date on a Pennisetum grain. Initial dates for other sites in
this study come from 14 radiocarbon dates on other materials, such
as charcoal, charred bone or wild seeds. Eight sherds with
 K. Manning et al. / Journal of Archaeological Science 38 (2011) 312e322
Table 3
Direct AMS and OSL dates on pearl millet remains from the Lower Tilemsi Valley (AMS dates calibrated using OxCal 4.1, using the IntCal 09 curve).
Site Lab no. Dating method
KN05 OxA16919 AMS
EN07 OxA-X-2287-26 AMS
EN07 OxA-X-2287-27 AMS
JS07-2 OxA-X-2287-29 AMS
EN07 X3318 OSL
EB07 X3315 OSL
identiable Pennisetum chaff impressions were chosen for dating.
Six of these contained the morphological traits of domestication.
These were submitted for both optically stimulated luminescence
dates, which should indicate the time the ceramic was red, and for
AMS radiocarbon dating of the organic fraction of the clay, which
may relate directly to the millet temper and/or organics in the source
clay. Of these only two produced completely reliable results for OSL-
dating (others lacked sediment for gamma measurement or had
anomalous U content). AMS dating failed on three sherds (I4, I6 and
I26) because after pre-treatment the remaining carbon samples
were too small for analysis. The ve successfully AMS-dated sherds
and the KN05 grain all produced dates in the Third Millennium BC,
mainly between 2500 and 2000 BC, although one sherd (I12) with
indeterminate Pennisetum chaff was earlier at 2850e2550 BC. These
results are therefore consistent with an age between the middle and
late part of the Third Millennium BC, (Table 3). The two sherds that
produced acceptable luminescence dates, have slightly later ages in
the early Second Millennium BC, but the wide early bars on such
dates, mean that they are also consistent with dates for pottery
making in the later Third Millennium BC. In order to analyse more
fully the chronometric data, we built a series of Bayesian models
using the OxCal 4.1 software (Bronk Ramsey, 2001, 2009). The t of
individual radiocarbon likelihoods within the Bayesian models was
evaluated through outlier detection analysis (Bronk Ramsey, 2009).
This was applied to enable an objective assessment of the probability
associated with individual measurements being demonstrable
outliers. Potentially erroneous determinations within the sequence
can be explicitly quantied, rather than remaining hidden. The
results suggested that the AMS dates on potsherds were consistently
older than their expected age based on other data, whereas the OSL
dates on the same sherds and associated sediments appeared in
good agreement with associated AMS dates of charcoal (Figure Sb).
Four of the six AMS dates run on Pennisetum chaff came back as
outliers, equaling 75% of the sample, in contrast to three out of the 20
AMS dates run on seeds/charcoal, equaling only 15% of the sample
(see SOM). This is consistent with a scenario in which there is a small
uptake of older carbon within the extracted temper used for the AMS
pre-treatment and dating (see SOM for more details). This probably
comes from some low carbon sediment fraction. Until now, inde-
pendent dates on pearl millet tempered pottery, or associated wood
charcoal, have provided a broad age bracket, often with notably wide
error margins (see Fig. 8) for the domestication of pearl millet in
West Africa. The results of this analysis demonstrate a secure
Fig. 5. Bayesian modelled range for the earliest domestic pearl millet remains in the Lower Tilemsi Valley, representing a terminus post quem for the domestication of pearl millet.
317
Description Date BP Calibrated range BC
Pennisetum sp. seed 4011  33 2620e2467
Pottery temper with Pennisetum glaucum 3782  28 2296e2063
Pottery temper with P. glaucum 3980  31 2579e2369
Pottery temper with P. glaucum 3604  30 2033e1888
Pottery sherd with P. glaucum temper 3680  250
Pottery sherd with P. glaucum temper 3830  280
terminus post quem of 2000 BC (Fig. 5), whilst the independent dates
and associated archaeological context suggest the domestication
process may have begun several centuries earlier.
4. Discussion: evolution of the domestication syndrome
4.1. Non-shattering and paired spikelets
The loss of natural (wild-type) seed dispersal is often taken as the
key mutation of domesticated cereals (Zohary and Hopf, 2000;
Fuller, 2007). Pennisetum is thought to have control of shattering
by the interaction of multiple linked genes that are tightly clustered
in two linkage groups (Poncet et al., 1998; 2000), and diversity of
mutations in these linkage groups amongst cultivars could support
more than one domestication episode (Poncet et al., 2002). Non-
shattering could have evolved by a single mutation to a gene for
pedicel length (PL) which causes the development of a rachis in place
of a dehiscence scar. Archaeobotanists have been able to document
that early occurrence and gradual increase in this trait in wheat,
barley (Tanno and Willcox, 2006; Fuller, 2007) and recently rice
(Fuller et al., 2009), and the evidence reported here makes a very
small contribution towards similar documentation of pearl millet
domestication. Eight domestic type non-shattering involucres,
indicated by rachis remains, were identied in the impressions by
comparison to just one possible wild type base. This high frequency
of domesticated types would indicate that the domestication
process was far advanced, and by comparison with other cereals we
would expect that pre-domestication cultivation had begun perhaps
a millennium earlier. While non-shattering domesticated types have
been recognized in impressions previously (e.g. Amblard and
Pernes, 1989; Klee et al., 2004; Fuller et al., 2007a), no quantitative
data on the proportions of wild to domesticated forms has been
available for tracking the domestication process.
Paired spikelets, or higher multiples, in pearl millet involucres
should be deleterious for wild type dispersal, equivalent to the 6-row
and 4-row forms of barley. This is evidence in the present material
from four examples of paired spikelets, and examples are also clearly
documented from later impression assemblages such as that from
Mauretania (Fuller et al., 2007a). We expect this trait to have been
selected for as part of general selection for increasing grain number,
on the one hand, while on the other hand it was permitted by the
relaxation of selection of natural dispersal aids, which also includes
the reduction of awns and bristles (cf. Fuller, 2007: 905).
318 K. Manning et al. / Journal of Archaeological Science 38 (2011) 312e322
Fig. 6. Metrical series of archaeological pearl millet grain size indicating grain breadth measurements in mm (for populations, mean and standard deviation, maxima and minima
are shown), plotted against approximate median age (primary data sources in SOM). Possible periods of rapid size increase (selection events) are indicated by dotted lines and
arrows. Measured sample size (n) is indicated.
4.2. Grain size and shape changes
A reasonable database of archaeological pearl millet grain
measurements is available from four West African sites (reviewed
in Fuller, 2007). To this we add a couple of measurements from the
Tilemsi, one from the charted grain in Fig. 2 and one from an
impression. Most measurements have been on charred grains,
which have probably undergone shrinkage during charring,
whereas one measurable grain impression in the Tilemsi sherd
material would not have undergone shrinkage and may have even
expanded in the wet claydthis measurement has therefore been
corrected by 10% and 20%. From these data we are able to assess
changes in grain size over time in West Africa, and we hypothesize
two distinct local episodes of selection for grain size increase in
Africa and one in South Asia (Fig. 6). Most early grain assemblages,
from 1700e1200 BC in Ghana or Nigeria, show the subtle change
towards domesticated grain shape, becoming apically thicker and
more club-shaped (DAndrea et al., 2001; Zach and Klee, 2003), but
these show no appreciable size increase, in grain length or breadth
that makes them different from the range of wild populations (Klee
et al., 2004). However, a single large grain reported from Boase in
Ghana (DAndrea et al., 2006) could indicate selection for enlarged
grains in some areas, although a larger sample size is needed to
conrm this.
One complicating factor in the interpretation of pearl millet grain
measurements is the range of variation within pearl millet. First
there are four recognized races of pearl millet, dened in part on the
basis of grain shape (Brunken et al., 1977). Secondly, there is the
potential presence of weedy varieties of millet, which possess a mix
of domesticated and wild traits, including grains smaller than most
domesticated millet (DAndrea et al., 2001; Zach and Klee, 2003). The
weed has evolved in part through cross-pollination between
domesticated and wild millet, and it is therefore unclear how early
during the domestication process such intermediate forms emerged.
Despite this the overall trend in millet grain size, even if archaeo-
logical samples are mixed, should be towards larger (thicker) grains
as selection under cultivation occurred. The case for this is
strengthened by evidence suggestive of multiple adaptive episodes
of grain enlargement in different regions across Africa and India.
Early pearl millet introduced to Gujarat, India by ca. 1700 BC,
also appears small, within the wild size range. By contrast, rather
later seeds of a North Indian (Gangetic) population from Narhan are
markedly larger, suggesting selection for larger-grained pearl millet
by 1400e1000 BC in India. Meanwhile in the Sahara (Libya and
Nubia) there can be suggested to be a trend towards larger grain
sizes by the early centuries AD. The more rapid increase in size on
arrival in the Ganges plain in India has been postulated to be due to
selection under intensive ard tillage (Fuller, 2007: 920), while the
more gradual or later selection in Africa at a later date is uncertain.
It was suggested that selection for larger grain sizes had to be
balanced against selection for increased grain number, which are
both components of overall yield and may have operated in
opposing ways (Fuller, 2007: 920); a comparable trade-off has been
described for Eragtostic tef domestication in northeast Africa
(DAndrea, 2008). These data do suggest that the dispersal of pearl
millet out of a West African Sahelian centre of origin occurred
before selection for increasing grain size, and it can therefore be
concluded that independent selection for increased grain size
occurred in parallel in several regions.
5. The dispersal of pearl millet across Africa and India
We have compiled a simple database of archaeobotanical pearl
millet, which includes 46 sites/phases from which it has been
reported in Africa (Fig. 7; see SOM). In India there are 15 sites/
phases with archaeobotanical evidence for Pennisetum (see SOM).
Their geographical and chronological distribution generally coin-
cides with that of other crops of African origin that arrive in India by
or shortly after 2000 BC (Fuller and Boivin, 2009). After the Third
millennium BC nds from the Tilemsi Valley, several nds from the
rst half of the Second Millennium BC are widely spread across
West Africa, including in Mauretania, Mali, Ghana, Burkina Faso and
Nigeria. Already in the early Second Millennium BC, pearl millet
had reached India, and this implies that it must have spread rapidly
eastwards across Africa in regions that as yet have been undocu-
mented by archaeobotany (Fuller, 2003), such as the northern
savannas through Niger, Chad and Sudan. Within India, the landfall
of this crop is likely to have been the Saurashtra peninsula with
 K. Manning et al. / Journal of Archaeological Science 38 (2011) 312e322 319

Fig. 7. Map of the distribution of archaeological pearl millet reports in Africa (for database see Table S16). Sites numbered: 1. Tilemsi sites (this study), 2. Walad, 3. Cubalel, 4. Sincu
Bara, 5. Arondo, 6. Dhar Tichitt, 7. Oued Chebbi, Oued Bou Khzama, Dhar Oualata, 8. Dhar Nema sites: Djiganyai, 9. Dia-Shoma, 10. Jenne-Jeno, 11. Wind Koroji, 12. Burkina Faso sites:
Oursi, Ti-n-Akof, Saouga, 13. Birimi, 14. Boase B5C, 15. Gao, 16. Essouk, 17. Tinda B, 18. Zinchechra, 19. Jarma, 20. Nok sites: Janruwa, Janjala, Akura, 21. Bwambe-Sommet, 22. Abang
Minkoo, 23. Ganjiganna and Mege, 24. Kursakata, 25. Daima, 26. Qasr Ibrim, 27. Meroe, 28. Kabusanze, 29. Pemba sites: Tumbe, Kimimba, Chwaka, Kaliwa, 30. Nqoma, 31. Ziwa
(Inyanga), 32. Kgaswe, 33. Matlapaneng, 34. Silver Leaves, 35. Magogo, 36. Ndondonwane, 37. Shongweni, 38. Ounjougou. Desert margins represented by dotted line and savannah to
Sahel transition indicated by the solid line with a dotted fringe. Rainforest coverage represented by grey shaded area.

semi-arid savannah conditions much like parts of Africa, and
dispersal then proceeded southwards down the savanna corridor of
the Indian peninsula. By the late Second Millennium BC, pearl
millet has also been reported from the Ganges basin, where it
shows its rst evidence for grain size increase in India (Fig. 6). In the
First Millennium BC, during the early West African Iron Age, we
have our rst hard evidence from Senegal and by the mid- to late
First Millennium BC, pearl millet had penetrated the Central African
forest area in southern Cameroun. In the late First Millennium BC it
had also been introduced to the previously winter-crop-focused
farming of the Fezzan (Southern Libya). Here it is likely to be
associated with the introduction of new irrigation systems and
a shift in settlement focus from elevated promontory sites to the
wadi oors. It occurs in Nubia slightly later at the end of the First
Millennium BC/early First Millennium AD. At present, nds from
eastern Africa start in the First Millennium AD associated with the
East African Iron Age. Historical linguistics suggests that pearl
millet was an established East African crop by this time both
amongst Bantu-speaking and Central Sudanic language groups
(Schoenbrun, 1993).
6. Conclusion: pearl millet as an alternative model for cereal
domestication
Africa has been suggested to provide an alternative to the
general trajectory towards agriculture that has been generalised
from Near Eastern data (Marshall and Hildebrand, 2002; Garcea,
2004). Rather than representing sedentary hunter-gatherers who
became farmers, the African trajectory was rst towards mobile
pastoralist-collectors, who cooked in ceramics, and then much later
the addition of cultivation with sedentism developing at or after
this time. One example of this was the domestication pathway of
pearl millet on the southern fringes of the Sahara somewhere in
West Africa, beginning prior to 2500 BC. Parallels can be found in
320 K. Manning et al. / Journal of Archaeological Science 38 (2011) 312e322

the savanna zones of India (Fuller, 2006: 58e60), and parallels
might also be drawn with the seasonal mobile societies who
apparently were involved in maize domestication in southern
Mexico (see Ranere et al., 2009). While African evidence provides
an alternative trajectory to food production from the Near East, the
specic case of pearl millet provides an alternative cereal domes-
tication pathway from the more oft-studied cases of wheat and
barley. In the case of pearl millet non-shattering evolved earlier
than the start of grain size increases, and the latter may have
occurred in parallel in several separate regions.
Once domesticated, pearl millet spread widely and rapidly. To
judge from nds in India, dating after 2000 BC but by 1700 BC, this
cereal must have spread eastwards right across the Sahelian/north
Savanna zone and reached India via sea contact within less than
1000 years of domestication. This rapid rate of spread may well have
been aided by the suitability of this grain for cultivation by mobile
pastoralist societies and the crops minimal water requirements.
Indeed the direct dating of the Pennisetum chaff tempered
pottery from the Lower Tilemsi Valley would go a long way to
supporting this model. A foundational cattle burial at KN05
(Manning, 2008b), dating to 3988  32 bp (OxA-16973) supports
the notion of a developed pastoral component in the very earliest
levels of occupation, and livestock clearly played an important
socio-economic role throughout the occupation of this region.
Although direct dating evidence on pearl millet remains them-
selves are fairly rare, there is a small corpus of dates, both on grains
and pottery clay organic fractions. As can be seen in Fig. 8, when
taken individually, existing dates whether on seeds or sherds, are

Fig. 8. Chronological distribution of AMS and OSL dates for the earliest palaeobotanical remains of domesticated pearl millet or potsherds/deposits directly associated with
domesticated pearl millet.
 K. Manning et al. / Journal of Archaeological Science 38 (2011) 312e322
centuries later than individual dating results from the Tilemsi. For
example the earliest direct grain dates from Birimi and Ounjougou
are probably 1700e1800 BC, while the grain from KN05 is earlier
than 2400 BC. Equally the Tilemsi sherd radiocarbon dates gave
results centuries older than those from elsewhere. However, the
Bayesian modelling undertaken here, demands a more critical
review of the dating evidence. The use of both AMS and OSL dating
and comparative modelling suggests that the AMS dates on
potsherds were consistently older than the OSL dates on the same
sherds and their associated sediments. Instead, it is probable that
the mid-Third millennium date range relates to a small uptake of
older carbon within the extracted temper. The terminus post quem
for domestic pearl millet as ceramic temper, based on the modelled
date range for sites in the Tilmesi Valley (Fig. 5), is therefore only
slightly earlier, by c. 200 years, than other known nds from the
western Sahara-Sahel borderlands. Nevertheless similar date
modelling exercises are necessary before the reported Pennisetum
dates can be taken at face value. Previously the earliest suggested
date came from wood charcoal from Wind Koroji, considered to be
associated with domestic pearl millet (MacDonald, 1996). Economic
and technological correlates between the Tilemsi Valley and Wind
Koroji may indicate cultural connections between these two areas
revealing a process of agro-pastoral exchange that likely charac-
terised the Sahara-Sahel borderlands towards the end of the Third
Millennium BC. Nonetheless, comparison of the independent dates
and their calibrated error margins highlights the separation of the
Tilemsi dates from other regional nds. It is likely that further
sampling in this region will reveal a greater number of domesti-
cated pearl millet remains and a more precise chronological
sequence. Indeed, a direct date on the Pennisetum grain from KN05,
although morphologically indeterminate, of ca. 2400 cal. BC (OxA-
168919) indicates consumption centuries earlier. Comparative
examples of the slow evolution of cereal domestication traits (e.g.
wheat, barley, rice) suggest that this could be an early domesticate
or in the pre-domestication cultivation stage (see Fuller, 2007;
Fuller and Allaby, 2010).
There are other clues to suggest that domestication occurred
earlier than the current evidence would suggest. The material
culture from the Lower Tilemsi Valley signies little in the way of
technological or stylistic divergence, indicating a degree of
cultural continuity between the earlier and later sites. The small
number of chaff tempered sherds found at KN05 and JS07-1
appear to comprise solely of Pennisetum, with no other plant
species. This is to be expected if pearl millet was the only seed
crop being repeatedly harvested as weeds are rarely incorporated
in the crop-processing by-products of pearl millet on account of its
dense spikes which are readily harvested alone (Reddy, 1997). Due
to the rare occurrence of chaff temper in this early phase of
occupation, it is likely that the pottery was being produced and
imported from outside the Tilemsi Valley, perhaps reecting
seasonal movement between the Tilemsi and millet-growing
regions further aeld. Carbon 13 stable isotopes of the human and
animal bone from KN05 support a major C-4 input from plants
such as millet or sorghum, into a mixed diet (Finucane et al.,
2008b). The presence of pearl millet, therefore, albeit of an inde-
terminate status, is likely to have been the source of C-4 in the
KN05 diet. Finally, the high frequency of domesticated types at
EB07, EN07 and JS07-2 suggest that the domestication process was
far advanced, and that pre-domestication cultivation had begun at
an earlier point in time.
Due to the generally poor state of preservation in the Sahelian
regions of Africa, plant impressions are often the only source of
evidence for past cultivation practices. The detailed chronological
modelling presented here issues an important warning for dating
the organic fraction in chaff tempered pottery. As shown in Table S2
321
(SOM), the AMS sampled pottery produced very low carbon yields,
i.e. <2% compared to c. 50% yield on stratigraphically related
charcoal and carbonized seed remains, increasing the likelihood of
sediment-derived carbon becoming a factor in the age determina-
tion. It is imperative that we proceed with caution when assessing
the reliability of tempered-derived dating (see also Higham et al.,
2009), and construct, where possible, detailed chronological
models that contextualise single-point dates, in order to avoid
misrepresenting the direction and timing of agricultural
development.
Acknowledgements
The eldwork undertaken by KM was supported by the Arts
and Humanities Research Council, the British Institute in Eastern
Africa, the University of Oxford and the Royal Anthropological
Institute. All dating was funded by the ORADS scheme
Appendix. Supplementary material
Supplementary data associated with this article can be found in
the online version, at doi:10.1016/j.jas.2010.09.007.
References
Amblard, S., Pernes, J., 1989. The identication of cultivated pearl millet (Pennise-
tum) amongst plant impressions on pottery from Oued Chebbi (Dhar Oualata,
Mauritania). African Archaeological Review 7, 117e126.
Bronk Ramsey, C., 2001. Development of the radiocarbon calibration program.
Radiocarbon 43 (2A), 355e363.
Bronk Ramsey, C., 2009. Bayesian analysis of radiocarbon dates. Radiocarbon 51 (1),
337e360.
Brunken, J.N., 1977. A systematic study of Pennisetum sect. Pennisetum (Graminae).
American Journal of Botany 64, 161e175.
Brunken, J., De Wet, J.M.J., Harlan, J.R., 1977. The morphology and domestication of
pearl millet. Economic Botany 31, 163e174.
DAndrea, A.C., 2008. Tef (Eragrostis tef) in ancient agricultural systems of highland
Ethiopia. Economic Botany 62, 547e566.
DAndrea, A.C., Casey, J., 2002. Pearl millet and Kintampo subsistence. African
Archaeological Review 19, 147e173.
DAndrea, A.C., Klee, M., Casey, J., 2001. Archaeobotanical evidence for pearl millet
(Pennisetum glaucum) in sub-Saharan West Africa. Antiquity 75, 341e348.
DAndrea, A.C., Logan, A.L., Watson, D.J., 2006. Oil Palm and prehistoric subsistence
in Tropical Africa. Journal of African Archaeology 4 (2), 195e222.
Finucane, B., Manning, K., Tour, M., 2008a. Prehistoric dental modication in West
Africa e early evidence from Karkarichinkat Nord, Mali. International Journal of
Osteoarchaeology 18 (6), 632e640.
Finucane, B., Manning, K., Tour, M., 2008b. Late Stone Age subsistence in the
Tilemsi Valley, Mali: stable isotope analysis of human and animal remains from
the site of Karkarichinkat Nord (KN05) and Karkarichinkat Sud (KS05). Journal
of Anthropological Archaeology 27, 82e92.
Fuller, D.Q., 2003. African crops in prehistoric South Asia: a critical review. In:
Neumann, K., Kahlheber, S., Butler, E.A. (Eds.), Food, Fuel and Fields:
Progress in African Archaeobotany. Heinrich-Barth Institut, Cologne, pp.
239e271.
Fuller, D.Q., 2006. Agricultural origins and frontiers in South Asia: A working
synthesis. Journal of World Prehistory 20, 1e86.
Fuller, D.Q., 2007. Contrasting patterns in crop domestication and domestication
rates: recent archaeobotanical insights from the old world. Annals of Botany
100, 903e924.
Fuller, D.Q., Allaby, R., 2010. Seed dispersal and crop domestication: shattering,
germination and seasonality in evolution under cultivation. Annual Plants
Reviews 38, 238e295. doi:10.1002/978144314557.ch7.
Fuller, D.Q., Boivin, N.L., 2009. Crops, cattle and commensals across the Indian:
current and potential archaeobiological evidence. Etudes Ocean Indien 42e43,
13e46.
Fuller, D.Q., Macdonald, K., Vernet, R., 2007a. Early domesticated pearl millet in
Dhar Nema (Mauritania): evidence of crop processing waste as ceramic temper.
In: Cappers, R. (Ed.), Field of Change. Proceedings of the 4th International
Workshop for African Archaeobotany. Barkhuis & Groningen University Library,
Groningen, pp. 71e76.
Fuller, D.Q., Boivin, N., Korisettar, R., 2007b. Dating the Neolithic of South India: new
radiometric evidence for key economic, social and ritual transformations.
Antiquity 81, 755e778.
Fuller, Dorian Q, Qin, Ling, Zheng, Yunfei, Zhao, Zhijun, Chen, Xugao, Aoi Hosoya, Leo,
Sun, Guo-ping, 2009. The domestication process and domestication rate in rice:
spikelet bases from the Lower Yangtze. Science 323, 1607e1610.
322 K. Manning et al. / Journal of Archaeological Science 38 (2011) 312e322
Garcea, E.A.A., 2004. An alternative way towards food production: the perspective
from the Libyan Sahara. Journal of World Prehistory 18 (2), 107e154.
Godbole, S.V., 1925. Pennisetum typhoideum. In: Studies on the Bajra Crop; I: The
Morphology of Pennisetum typhoideum. Memoirs of the Department of Agricul-
ture in India, vol. 14. Agricultural Research Institute, Pusa. 247e268.
Harlan, J.R., 1992. Indigenous African agriculture. In: Cowan, C.W., Watson, P.J.
(Eds.), The Origins of Agriculture: An International Perspective. Smithsonian
Press, Washington, D.C., pp. 59e70.
Higham, C.F.W., Kuzmin, Y.V., Burr, G.S., 2009. The AMS 14C dating of Iron Age rice
chaff ceramic temper from Ban Non Wat, Thailand: rst results and its inter-
pretation. Nuclear Instruments and Methods in Physics Research Section B:
Beam Interactions with Materials and Atoms 268, 1022e1025.
Kahlheber, S., Neumann, K., 2007. The development of plant cultivation in semi-arid
West Africa. In: Denham, T.P., Iriarte, J., Vrydaghs, L. (Eds.), Rethinking Agri-
culture: Archaeological and Ethnoarchaeological Perspectives. Left Coast Press,
Walnut Creek, pp. 320e346.
Klee, M., Zach, B., Stika, H.-P., 2004. Four thousand years of plant exploitation in the
Lake Chad basin (Nigeria), part III: plant impressions in potsherds from the Final
Stone Age Gajiganna culture. Vegetation History and Archaeobotany 13,
131e142.
MacDonald, K.C., 1996. The Wind Koroji Complex: evidence for the peopling of the
eastern Niger Delta (2100-500 BC). Prhistoire Anthropologie Mditerranenes
5, 147e165.
Manning, K., 2008a. Mobility, climate change and cultural development. Arevised
view from the Lower Tilemsi Valley, Northeastern Mali. Unpublished PhD
thesis, University of Oxford.
Manning, K., 2008b. Mobility strategies and their social and economic implications
for Late Stone Age Sahelian pastoral groups: a view from the Lower Tilemsi
Valley, eastern Mali. Archaeological Review from Cambridge 23 (2), 125e145.
Marshall, F., Hildebrand, E., 2002. Cattle before crops: the beginnings of food
production in Africa. Journal of World Prehistory 16, 99e143.
Neumann, K., 2005. The romance of farming: plant cultivation and domestication in
Africa. In: Stahl, A.B. (Ed.), African Archaeology. A Critical Introduction. Black-
well Publishing Ltd, Oxford, pp. 249e275.
Oumar, I., Marciac, C., Pham, J.-L., Vigouroux, Y., 2008. Phylogeny and origin of pearl
millet (Pennisetum glaucum [L.] R. Br) as revealed by microsatellite loci. Theo-
retical and Applied Genetics 117, 489e497.
Poncet, V., Lamy, F., Enjalbert, J., Joly, H., Sarr, A., Robert, T., 1998. Genetic analysis of
the domestication syndrome in pearl millet (Pennisetum glaucum L, Poaceae):
inheritance of the major characters. Heredity 81, 648e658.
Poncet, V., Lamy, F., Devos, K., Gale, M., Sarr, A., Robert, T., 2000. Genetic control of
domestication traits in pearl millet (Pennisetum glaucum L., Poaceae). Theoret-
ical and Applied Genetics 100, 147e159.
Poncet, V.F., Martel, E., Allouis, S., Devos, K.M., Lamy, F., Sarr, A., Robert, T., 2002.
Comparative analysis of QTLs affecting domestication traits between two
domesticated x wild pearl millet (Pennisetum glaucum L., Poaceae) crosses.
Theoretical and Applied Genetics 104, 965e975.
Ranere, A.J., Piperno, D.R., Holst, I., Dickau, R., Irarte, J., 2009. The cultural and
chronological context of early Holocene maize and squash domestication in the
Central Balsas River Valley, Mexico. Proceedings of the National Academy of
Sciences (USA) 106 (13), 5014e5018.
Reddy, S.N., 1997. If the threshing oor could talk: integration of agriculture and
pastoralism during the late Harappan in Gujarat, India. Journal of Anthropo-
logical Archaeology 16, 162e187.
Schoenbrun, D.L., 1993. We are what we eat: ancient agriculture between the great
lakes. Journal of African History 34, 1e31.
Smith, A.B., 1992. Pastoralism in Africa. Origins and Development Ecology. Hurst &
Company, London.
Tanno, K.-I., Willcox, G., 2006. How fast was wild wheat domesticated? Science 311,1886.
Tostain, S., 1998. Le mil, une longue histoire: hypotheses sur domestication et ses
migrations. In: Chastenet, M. (Ed.), Plantes et Paysages dAfrique. Une Histoire
a explorer. Editions Karthala, Paris, pp. 461e490.
Zach, B., Klee, M., 2003. Four thousand years of plant exploitation in the Chad Basin
of NE Nigeria II: discussion on the morphology of caryopses of domesticated
Pennisetum and complete catalogue of the fruits and seeds of Kursakata.
Vegetation History and Archaeobotany 12, 187e204.
Zohary, D., Hopf, M., 2000. Domestication of Plants in the Old World, third ed.
Oxford University Press, Oxford.