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)

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Journal of Anthropological Archaeology 27 (2008) 82–92

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)

Brian Finucane a,*, Kate Manning b, Mouktarde Toure c

a Yale University, 730 Whitney Avenue, Apt. 3a, New Haven, CT 06511, USAb Institute of Archaeology, Oxford University, United Kingdom

c Direction Regionale de la Jeunesse des Sports, Arts et Culture, Region de Gao, Republique du Mali

Received 11 August 2007; revision received 10 October 2007Available online 3 December 2007

Abstract

The pathways leading to the adoption of cereal cultivation and pastoralism in West Africa are poorly understood. In order to elu-cidate the transition to food production during the Late Stone Age in Mali’s Tilemsi Valley samples of ancient and modern humanand animal remains were selected for carbon and oxygen isotope analysis. Our results indicate the inhabitants of Karkarichinkat Nord(KN05) consumed considerable quantities (�85%) of carbon derived from C4 plants, either directly in the form of C4 grasses such as wildPanicum sp. and possibly domestic Pennisetum sp. or indirectly through the consumption of C4 grazers such as Bos sp. and Ovis sp.� 2007 Elsevier Inc. All rights reserved.

Keywords: Bioarchaeology; Stable isotopes; Agriculture; Pastoralism; Neolithic; Sahara; Millet; Cattle

Introduction

The transition to food production represents one of themost significant economic shifts in human prehistory.Although we are able to discern the broad outlines of thisprocess in some of the primary centers of domesticationsuch as the Fertile Crescent, China, Mexico, and the Cen-tral Andes, less is known about the adoption of agricultureand pastoralism in secondary centers such as sub-SaharanWest Africa. Both the absolute and relative timing of thetransition to sedentism, the adoption of animal husbandryand the transition to cereal cultivation in this region remainpoorly understood (but see Haour, 2003 for a review of theNeolithic in Niger). In order to elucidate the transition tofood production at the Late Stone Age (LSA) site of Kark-arichinkat Nord (KN05) in Mali’s Tilemsi Valley, stableisotope analysis of archaeological human and animal

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doi:10.1016/j.jaa.2007.10.001

* Corresponding author.E-mail address: [email protected] (B. Finucane).

remains from the site was undertaken. Whereas analysisof archaeological faunal and floral remains provides a mea-sure of the presence of resources, analysis of the stable iso-tope composition of an animal’s tissue can provide ameasure of the prevalence of these resources in an animal’sdiet. Although there have been a number of studies of aus-tralopith and early Homo dietary ecology utilizing biogeo-chemical techniques (see Lee-Thorp and Sponheimer, 2006for a review), our research represents the first applicationof stable isotope analysis to the investigation of the dietof prehistoric Homo sapiens in West Africa.

Environmental context

The Lower Tilemsi Valley is one of several palaeochan-nels extending north from the Niger River. Although cur-rent conditions are principally Saharan, the valley did atone time host an extensive river and tributary network,stretching from the Tanezrouft in the north, to the Nigerin the south.

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B. Finucane et al. / Journal of Anthropological Archaeology 27 (2008) 82–92 83

In the last 30 years, extensive research has been under-taken into the climatic and floral history of the Sahara,demonstrating a distinct humid phase in Africa’s later pre-history, beginning around 12,000 BP (Maley, 1977, 1980,1982; Lezine and Cassanova, 1989; Lezine, 1991; Marchantand Hooghiemstra, 2004; Hillaire-Marcel et al., 1983; Petit-Maire and Riser, 1981). Several optimal episodes occurredduring this period, notably between 9400 and 7700 BP, andagain from 7200 to 4500 BP. During the second of thesehumid phases, conditions were significantly more erratic,with periods of aridity at 6400 and 5500 BP. Between5000 and 4000 BP, conditions rapidly deteriorated, with adistinct dry episode around 4200/4000 BP, marking theend of the Holocene humid phase and the onset of currentarid conditions.

Today, the Tilemsi Valley is a semi-arid grassland whichsupports a few scattered acacia trees but where annual rain-fall is below the minimum required for dry agriculture.However, archaeological and palaeobotanical evidenceindicates that conditions during the later Holocene weremore comparable to those of the Middle Niger today,where dry agriculture reaches its northern extent, and irri-gated croplands are prevalent. The initial palaeobotanicalanalysis from KN05, undertaken by Dr. Ruth Pelling,has identified fragments of Vitex sp., Panicum sp., Zizyphys

sp., and several indeterminate fragments of Leguminosae.These species are more representative of a Sahelian or Sud-ano-Sahelian environment than of a Saharan one. The fau-nal remains from KN include aquatic mammals, deep andshallow water fish species, domestic livestock, and wildbovids. The remains of Lates niloticus measuring �1 m,recovered at KN05 attest to the fact that the Tilemsi con-tained a permanent deep water channel. Today, this sortof mixed faunal assemblages is only found south of theMiddle Niger.

The origins and development of agriculture and pastoralism

in sub-Saharan West Africa

Evidence at the site of Nabta Playa E-75-6 indicatesdomestic cattle were present in the eastern Sahara by�8000 BP. From here, domestic cattle appear to havespread westwards, across the Sahara, and southwards,along the Nile valley. Cattle are present at Gabrong andBaradigiue in the Tibesti where remains have been datedto 7455 ± 180 BP (Gautier, 1984; Barich, 1987), and atAdrar Bous in the Tenere desert of Niger, where they havebeen dated to �6000 BP. After 4000 BP, the deteriorationof conditions in the Sahara, prompted the movement ofcattle into sub-Saharan West Africa (Clark, 1976; Munson,1976, 1980; Casey, 1998). Shaw (1977) notes that anymovement of pastoral populations prior to this time wouldhave been restricted by the disease vectors of Sahelian WestAfrica and was only alleviated by a southward displace-ment of the tsetse barrier �3700 BP.

The earliest evidence for agriculture in West Africaappears much later than that for pastoralism. The earliest

direct evidence comes from the Sahara–Sahel margins. AtKarkarichinkat, de Wet (Smith, 1984, p. 89; Smith, 1992,p. 74) identified domesticated pearl millet (Pennisetum

glaucum) impressions on pottery, which Smith stylisticallydates to the end of the 3rd millennium BP. At Dhar Tichitt,in southwestern Mauritania, impressions of domesticatedpearl millet have been firmly dated to at least 3800 BP(Amblard, 1984, 1995; Amblard and Pernes, 1989). Morerecently domesticated millet grains have been dated to3490 ± 50 BP (1878–1744 cal BC) at Birimi in northernGhana (D’Andrea et al., 2005).

Based on the distribution of its wild progenitors, Harlan(1971, 1992) locates the center of millet domestication inthe Western Sahara. However, the earliest remains ofdomesticated millet have been found in western India. Atthe sites of Koethe, Gujarat, Babor Kot, and Surkotadadomesticated millet has been dated to the end of the 5thmillennium BP (Tostain, 1998; Fuller, 2003). The archaeo-logically evidence for the center of millet domesticationtherefore still awaits discovery in the vast unstudied areasof Western Sahara.

Archaeological context

The sites of Karkarichinkat Nord (KN05) and Sud(KS05) are located 80–85 km north of Gao, in EasternMali (Fig. 1). They were first recorded by Mauny (1952)who visited the lower Tilemsi Valley in 1952. Two decadeslater Andrew Smith excavated five test pits, which yieldedevidence of an agro-pastoral economy dating to the endof the 5th millennium BP (1974a,b, 2005). At KN05 Smithuncovered evidence for intensive occupation, includingpottery, domestic livestock, and fish. Smith also identifieddomesticated pearl millet (P. glaucum) impressions onceramics, which he associates with the earliest levels ofoccupation. However, these samples appear to have beencollected only from the surface. They do not provide con-clusive support for the notion that a fully developed agri-cultural economy existed during the site’s initialoccupation, nor do these impressions establish the preva-lence of millet in the diet of the site’s inhabitants. AlthoughGaussen and Gaussen (1988) undertook an extensive sur-face survey and described three broadly contemporaneouscultural facies, their results lack chronological resolution.

In order to expand upon Smith’s preliminary findings in2005 a team from the University of Oxford, working in col-laboration with the Direction National du PatrimoineCulturelle, began excavations at KN05. A total of eight2 · 2 m trenches were excavated, of which three reachedbasal deposits at 2.40 m. An additional eight trenches werelocated over eroding human burials (Fig. 2). A series of 10AMS dates were obtained from the main excavation area,dating the occupation of the site to between 2206 and2620 cal BC (Table 1). A further AMS date of 2479–2292 cal BC (OxA-16976) was obtained from trench 4, ona piece of charcoal directly below skeleton 1, indicating

Fig. 1. Map of Mali indicating location of Karkarichinkat Nord and Sud.

84 B. Finucane et al. / Journal of Anthropological Archaeology 27 (2008) 82–92

the main phase of occupation and the eroding surface skel-etons were contemporaneous (Fig. 3).

Domestic livestock, notably cattle, as well as sheep,goat, and fish were found in abundance throughout thesequence. However despite intensive palaeobotanical sam-pling, evidence for the exploitation of grains, either domes-tic or wild, is very limited. A single grain of millet(Pennisetum sp.), which has been identified as possibledomestic, was retrieved from the upper 0.50 m of deposit.Clearly, a sample size of one cannot be used conclusivelyto infer domestication. Nonetheless, grinding equipmentwas found in relative abundance (N = 18) and suggests thatcereals did play a role in the diet of the Karkarichinkatinhabitants. Whether these grasses were domesticated hasyet to be established.

The abundance of domestic livestock at the site is con-sistent with a pastoralist economy. Of particular note is aburial feature containing a fully articulated cow with aset of post holes surrounding the grave pit found cut intothe primary occupation levels in trench 1-B/C. The remainsof disarticulated cattle bone, often charred and with cutmarks, are indicative of dietary consumption. In contrastthe purposeful inhumation of a complete cow suggests thatcattle also played a role in the belief systems of KN05’s ear-liest inhabitants. Numerous cow figurines were also foundat KN05, attesting to a recognition of the lactating func-tion of cattle. Several rim sherds of pottery (N = 6) werealso found with spouts, which may support the idea thatmilk, or milk products were part of the KN05 diet.

Interestingly, sheep and goat disappear from thesequence at 2.0 m, despite the continuance of cattle, whilstthe number of fish remains significantly increases. The ini-tial observations from the faunal analysis therefore suggestthat in the earliest levels of occupation, cattle and fish wereimportant resources. Soon after occupation was estab-lished, sheep and goat are introduced into the economy,and may indicate a shift towards more sedentary occupa-tion. Similarly, grinding equipment is absent from thelower 0.70 m of deposit, and is generally more abundanton the surface, which may also be representative ofincreased sedentism and a shift towards a more diversediet.

Over two kilograms of fish bones were recovered fromKN05, almost half of which came from a single pit (078)cut into the primary occupation layer of Trench 1-A. Thetaxa represent in this osteological assemblage include Clar-

iidae, Synodontis, Tilapiini, L. niloticus, Bagriidae, Gymanr-

chus, Polypterus, Protopterus, and Mormyridaei. Thecatfish Clariidae and Synodontis alone account for over75% of the assemblage. These remains provide strong evi-dence that fish constituted an important component ofhuman diet at the site during prehistory.

Stable isotope background

The study of paleodiet through stable isotope analysisproceeds from the experimental observation that the isoto-pic composition of animal tissues generally reflects that of

Fig. 2. Site plan of Karkarichinkat Nord (KN05).

Table 1Radiocarbon determinations from KN05

Lab code Date BP Error 1r Material Context

OxA-16895 3894 32 Charcoal Context (032) at 0.60 mOxA-16896 3882 32 Euphorbiaceae sp. Context (050) at 1.30 mOxA-16897 3856 32 Charcoal Context (081) at 1.60 mOxA-16919 4011 33 Pennisetum sp. Context (031) at 0.40 mOxA-16920 3922 33 Charcoal Context (037) at 1.0 mOxA-16973 3988 32 Leguminosae sp./Vitis sp. Context (166) at 2.40 mOxA-16974 3853 32 Charcoal Context (077) at 0.40 mOxA-16975 3877 33 Charcoal Context (077) at 0.50 mOxA-16976 3913 33 Sterculia sp. Context (147) at 0.10 m (skeleton 1)OxA-16898 3889 32 Combretaceae sp. Context (085) at 4.0 mOxA-16977 3986 31 Charcoal Context (087) at 4.40 m


the diet they consume. Such analyses utilize the variation inthe ratios of the stable isotopes of carbon and nitrogenwithin ecosystems to measure the relative contribution ofdifferent resources to the diets of humans and other ani-mals (DeNiro and Epstein, 1978, 1981). As some foods,such as maize and millet, have distinctive isotopic signa-tures, it is often possible to identify the consumers of theseresources.

The stable isotope ratios of carbon (13C and 12C) andthose of nitrogen (15N and 14N) are expressed in per mil(&) as d values:

d ¼ ½ðRsample=RstandardÞ � 1� � 1000

where R = 13C/12C for the measurement of carbon and15N/14N for the measurement of nitrogen. The standardsto which samples are compared are the limestone Vienna

Fig. 3. Probability density distributions of calibrated radiocarbon mea-surements from KN05.


PeeDee Belemnite (VPDB) and atmospheric nitrogen(AIR) for carbon and nitrogen respectively. Most materi-als, including plant and animal tissues have less 13C thanthe VPDB, and their d13C values are typically negative.

Carbon enters terrestrial food webs through the uptakeof atmospheric CO2 by autotrophs. The fundamental vari-ation in the ratios of stable carbon isotopes in these foodwebs stems from differences in the photosynthetic path-ways of plants, categorized as C3, C4, and CAM. The over-whelming majority of plants utilize the Calvin Cycle (C3)pathway and have an average d13C value of �26.5&

(O’Leary, 1988). The d13C values of plants relying uponthe C4 pathway, mainly tropical grasses, are on average�12.5& (O’Leary, 1988; Van der Merwe and Tshauner,1999). Plants utilizing a third pathway, Crassulacean acidmetabolism (CAM), combine aspects of both C3 and C4

plants. The tissues of CAM plants have d13C values rang-ing from �11& when CO2 is absorbed at night, to�28& when CO2 is taken up during the day.

The carbon isotope signature of animal proteins, such ascollagen in bone and keratin in hair and nails, predomi-nantly reflect the protein component of diet, as amino acidsare preferentially routed from diet to be incorporated intobody tissue (Ambrose and Norr, 1993; Tieszen and Fagre,1993; Lee-Thorp et al., 1989). The proteins of animals areenriched in 13C relative to their diets by �4&. Consider-able variability is observed in d13C values of fauna in mar-ine and freshwater ecosystems as well as the consumers ofsuch protein. Much of this variation stems from differencesin the concentration of dissolved inorganic carbonates andthus the d13C values of aquatic plants within the ecosystem.

In general marine fish tend to be enriched in d13C, with iso-topic values resembling those of C4 plants whereas the iso-topic values of freshwater fish cover the spectrum from C3-like to C4-like (Norr, 1995; White et al., 1993).

In contrast the carbon isotope composition of the sec-ond major class of material analyzed in paleodietary stud-ies, carbonates, more accurately represents whole dietincluding carbohydrates, lipids, and protein (Ambroseand Norr, 1993; Tieszen and Fagre, 1993; Schwarcz,2000). Most of the mass of osseous remains is mineral, apa-tite and hydroxyapatite, 70% in bone and tooth dentin and98% in enamel. The approximate formula of this com-pound is Ca10(PO4)6(OH)2. Structural carbonate occurswhen –CO3 replaces –PO4 in the apatite crystals. This car-bonate is derived from blood bicarbonate generated by themitochondria during cellular metabolism and is on averageenriched in 13C by 9.5–12& relative to diet (Ambrose andNorr, 1993; DeNiro and Epstein, 1978; Lee-Thorp et al.,1989; Passey et al., 2005). Dental enamel provides a higherfidelity record of paleodiet than bone apatite as enamel isdenser, has larger crystals and is virtually devoid of organicmatter (LeGeros, 1991; Elliot, 1994). Enamel is depositedduring a discrete period early in an animal’s life and isnot subject to later reworking. Analysis of known fossilgrazers and browsers substantiate the reliability of enamelcarbonate as a record of paleodiet (Lee-Thorp and van derMerwe, 1987; Cerling et al., 1997).

The carbonate in the bioapatite of wild large herbivoresis enriched by 14.6& relative to diet, whereas the carbonateof small animals used in feeding experiments is enriched by9& relative to diet (Lee-Thorp et al., 1989; Tieszen andFagre, 1993; Passey et al., 2005). Swine, the best digestiveproxy available for humans, exhibit enrichment of carbon-ate by 10.2& relative to diet (Hare et al., 1991; Howlandet al., 2003). Variation in isotopic spacing between dietand the carbonates of bone and tooth enamel is attributedprimarily to differences in digestive physiology. The pro-duction of methane, CH4, by bacteria in the digestive tractresults in the loss of isotopically light carbon and a con-comitant enrichment of the remaining CO2 which entersthe bloodstream and is later incorporated in the structuralcarbonates of bone and enamel. The greater methane pro-duction in the digestive tracts of ruminants is invoked toexplain the enrichment of their bone and enamel carbon-ates vis-a-vis rodents (Hedges, 2003; Passey et al., 2005).

Using a linear mixing model, the percentage of C4

derived nutrients in an animal’s diet can be calculated usingthe following formula, adapted from Schwarcz et al.(1985):

%C4 ¼ ðdcarb � d3Þ=ðd4 � d3Þ � 100

where dcarb is the measured value of the carbonate sample,d3 = �16 (the value of carbonate of pure C3 consumer) andd4 = �2 (the value of carbonate of pure C4 consumer).

The oxygen isotope composition of carbonate in toothenamel reflects the composition of the animal’s body waterand is affected by diet, climate, and physiology (Longinelli,


1984; Luz et al., 1984; Bocherens et al., 1996; Kohn, 1996).Body water is derived from atmospheric O2, the oxygenchemically bound in foods, and liquid water. As O2 inthe atmosphere is relatively constant (Dole et al., 1954;Kroopnick and Craig, 1972), the primary sources of varia-tion in body water composition are liquid water which isclimatically sensitive (Dangaard, 1964) and to a lesserextent oxygen in food.

Liquid water is ingested by herbivores and omnivoresnot only from surface water but also in roots, stems andleaves of plants. Although the isotopic composition ofwater contained within roots and stems is similar to meter-oric water, leaf water is generally enriched in 18O (Gonfian-tini et al., 1965; Epstein et al., 1977; Sternberg, 1989; Yakir,1992). In addition to free water, plant and animal tissuealso contains chemically bound oxygen with plants beingenriched in 18O relative to animal tissues.

The dietary patterning of animal oxygen isotope compo-sition is complex with browsers and mixed feeders showing18O enrichment relative to grazers in some environments(Kohn et al., 1996) and the opposite trend being observedin other environments (Bocherens et al., 1996). Sponheimerand Lee-Thorp (1999b) have suggested that this discrep-ancy may reflect the relative contribution of surface waterto an herbivore’s total water consumption. Althoughbrowsers and mixed feeders may obtain most of their waterfrom the 18O enriched water in plant leaves, if they drinksurface water on a daily basis their body water compositionmay not differ significantly from grazers which are obligatedrinkers. In general faunivores and many primates aredepleted in 18O relative to herbivores (Sponheimer andLee-Thorp, 1999b).

Methods and materials

The skeletal remains of at least 11 humans were recov-ered from nine contexts at the sites of KarkarichinkatSud (KS05) and Karkarichinkat Nord (KN05) (Figs. 1and 2). The human skeletal remains from KN05 wereexposed by deflation and were highly fragmented anddegraded due to bioturbation, four wheel drive vehiclesand exfoliation from wind and sun. Due to the disturbanceof the remains, the determination of the grave orientationwas not always possible, though in several cases skeletonsappeared to be interred in supine positions with both armsand legs extended in E–W orientation. Poor protein preser-vation precluded direct radiocarbon measurements ofhuman remains. In contrast the skeleton excavated atKS05 was both complete and better preserved than theremains from KN05. The skeletal remains from KS05 camefrom an intact burial cut into the Late Stone Age/Neolithicoccupation mound and ringed with stone. These remainswere estimated to date to the Islamic era (post AD1300)on the basis of the stratigraphy (the grave was cut throughthe LSA mound), their preservation (which included hairand cartilage), and the fact that the body was interred onits right side facing east towards Mecca.

Estimates of age and sex were made using standard oste-ological criteria (Buikstra and Ubelaker, 1994; Bass, 1987).Criteria considered included cranial and pelvic morphol-ogy, dental eruption and wear, cranial suture and epiphy-seal closure and overall robusticity. Due to the poorpreservation of the remains, broad age estimates were used.

In order to interpret human isotope values it was neces-sary to obtain an isotopic baseline using archaeological andmodern fauna as well as modern plant samples. Theremains of domestic Bos sp. and ovi-caprid (distinguishingbetween sheep and goats using skeletal remains is difficult)were recovered from both the surface and deep soundings.The skeletal remains of fish were abundant at the site andthe taxa sampled included Polypterus sp., Gymnarchus sp.,Claria sp., Tilapia sp., Synodontis sp., and L. niloticus. Inaddition modern fish from the Niger River and locallygrown crops were obtained for analysis from a market inthe town of Gao.

Collagen was extracted from bone and dentin followingprocedures described in Richards and Hedges (1999). Iso-topic analysis was conducted using a Carlo Erba 1108 car-bon and nitrogen elemental analyzer coupled to a EuropaGeo 20/20 mass spectrometer in continuous flow mode.The isotopic values of all samples were measured relativeto tertiary laboratory standards of nylon and alaninewhose isotopic values are calibrated with respect to IAEAand NBS standards which have internationally agreed val-ues relative to VPDB. All samples were analyzed in tripli-cates in separate batches. Analytical errors are of theorder of ±0.2& for d13C. The preparation and isotopicanalysis of modern fish bones and plant samples was con-ducted by Hannes Schroeder.

Samples of carbonate were prepared from teeth accord-ing to procedures outlined in Finucane et al. (2006).Enamel was analyzed by Norman Charnley in the Depart-ment of Earth Sciences, Oxford University. Powdered sam-ples of enamel were reacted with 100% H3PO4 in a vacuumat 90 �C in a common acid bath for six minutes. Isotopicmeasurements were made using a VG Prism IRMS. Allmeasurements were made relative to the standardsNBS19 and IAEA CO1. Analytical precision is better than.1&.

Results

None of the archaeological samples of human and ani-mal bone or dentin yielded an appreciable quantity of col-lagen. However the modern samples from the Niger Riverprovide a reasonable, if rough proxy for the carbon isotopevalues of the ancient fish of the nearby Tilemsi drainage.The eight modern fish collagen samples analyzed have amean d13C of �18.4 ± 2.09& (Table 2). The carbon andoxygen isotope values of the human and animal toothenamel are presented in Tables 3 and 4, respectively, anddisplayed in Fig. 4. The eight samples of human toothenamel have a mean d13C value of �3.91 ± 1.31& and amean d18O of �1.5 ± 1.31&. The mean value of the 28

Table 3Stable isotope values of enamel carbonate of archaeological fauna from Karkarichinkat Nord

Sample Trench Context Taxon Tooth d13C d18O

KN59A B 5 Bos M3 �1.2 4.4KN22A A 15 Bos M3 �3.8 2.2KN53A A 15 Bos M3 2.0 1.1KN58A A 15 Bos M2 1.7 4.1KN21A B/C 31 Bos M1 1.9 0.6KN68A B/C 31 Bos M1 1.4 0.5KN16A B 34 Bos M1 1.6 2.3KN62A C 34 Bos M2 1.2 �0.5KN63A B 34 Bos M2 �0.8 1.9KN67A C 35 Bos M1 0.7 2.9KN70A B 36 Bos M1 �0.6 2.6KN69A B 40 Bos pm4 1.9 1.6KN57A C 48 Bos M1 2.2 3.0KN12A B 49 Bos M2 1.4 1.2KN23A B 50 Bos M3 0.2 2.3KN52A C 50 Bos M2 1.8 5.2KN56A B 50 Bos M2 2.3 4.7KN61A B 50 Bos M2 1.7 2.6KN54A B 71 Bos M3 2.6 2.4KN49A 3-A 72 Bos M1 0.8 2.1KN50A 3-A 73 Bos P4 1.8 4.5KN20A E 77 Bos M1 1.3 3.2KN64A B/C 81 Bos M2 1.5 1.3KN9A A 85 Bos M1 �0.3 6.0KN51A A 96 Bos M1 0.6 6.7KN15A B/C 166 Bos I1 2.1 6.1KN55A D 76/28 Bos M1 1.9 2.8KN60A G 77-SPIT4 Bos M2 0.4 3.7

Mean 1.0 2.91SD 1.4 1.8

KN19A A 16 Ovi-cap M1 �4.4 1.3KN10A B 31 Ovi-cap M �6.3 5.8KN66A B 31 Ovi-cap M2 �4.6 5.2KN65A D 124 Ovi-cap M2 2.0 4.4

Mean �3.3 4.21SD 3.7 2.0

Table 2Stable isotope values of human tooth enamel from Kakarichinkat Nord (KN) and Kakarichinkat Sud (KS)

Sample Trench Context Skeleton Sex Age Tooth d13C d18O

KN1H 160 8 M A LM3 �2.6 �1.9KN5H 7 151 10 F? A RP2 �4.3 �2.8KN11H 5 148 3 F YA LP1 �3.8 �2.3KN12H 159 7 F YA LM3 �3.1 �1.2KN13H 5 148 2 F? YA LM3 �3.1 �1.4KN17H 8 153 6 F? YA LM3 �3.7 �1.8KN20H 6 149 5 ? Child LM1 �4.3 �1.7KS2H 3 11 M 30–35 LM3 �6.4 1.6

Mean �3.9 �1.41SD 1.2 1.3


samples of Bos sp. tooth enamel have a mean d13C value of1.0 ± 1.37& and a mean d18O value of 4.2 ± 1.99&. Thefour ovi-caprid enamel samples have a mean d13C valueof �3.3 ± 3.66& and a mean d18O value of 4.2 ± 1.99&.Human and ovi-caprid enamel samples have d13C valueswhich are significantly lower than those of Bos sp. enamel

(t-test, p < .001). Human enamel samples are also signifi-cantly depleted in d18O relative to those of the fauna (t-test,p < .001), though the d18O values of enamel from Bos sp.and those of the ovi-caprids are not significantly different(t-test, p < .001). Carbon and oxygen isotope values arenot significantly correlated (d.f. = 37, r2 = .1502, p > .01).

Table 4Carbon isotope values of modern fish from the Niger River at Gao, Mali

Genus d13C (&)

Bagrus �19.4Clarias �17.7Hydrocynus �18.7Lates �15.9Mormyridae �16.5Synodontis �12.6Teradon �17.4Tilapia �17.4

Mean �16.9SD 2.1

Values have been corrected for the industrial effect by adding 1.5&.


The d13C values of modern samples of millet and sorghum(Table 5) are consistent with the known values of C4 plants.In order to compare the modern resources with the prehis-toric samples the carbon isotope values of fish and plantshave been adjusted by +1.5& to compensate for the indus-trial effect (Indermuhle et al., 1999).

Discussion

The observed d13C and d18O enamel values are consis-tent with biogenic rather than diagenetic signals. If theseisotopic signals were diagenetic, then diagenesis has differ-entially altered the carbon and oxygen isotope signatures ofhumans, Bos sp. and ovi-caprids. Given that these remainswere recovered in close proximity in similar burial contexts,there is no obvious reason for differential diagenesis oftooth enamel. These findings are consistent with patternsobserved in other studies of fossil tooth enamel (Lee-Thorpand van der Merwe, 1987; Bocherens et al., 1996; Cerlinget al., 1997; Sponheimer and Lee-Thorp, 1999a,b; Vander Merwe et al., 2003; Lee-Thorp and Sponheimer,2003; Kingston and Harrison, 2007) strengthening theproposition that enamel preserves dietary isotope signa-tures with high fidelity.

Fig. 4. Bivariate plot of carbon and nitrogen isotope values

The carbon isotope signatures of the human remains areconsistent with the consumption of a mixed diet incorpo-rating terrestrial animal protein (i.e. Bos sp. and ovi-capr-ids), riverine fish, and C4 plants such as millet andsorghum. Using the formula presented earlier, humans atKN05 consumed diets that were on average �85% derivedfrom C4 resources. Due to the fact that the domestic ani-mals at the site consumed diets comprised primarily ofC4 plants (see below) the d13C value of their body proteinis similar to that of millet and sorghum. As a result of thisisotopic equifinality it is difficult to assess the precise extentto which human tissues were enriched in 13C directlythrough the consumption of C4 plants such as millet andsorghum or indirectly through the consumption of C4 grasseating animals (see Table 6). By establishing the relativetrophic level of humans at KN05, analysis of the nitrogenisotope values of bone collagen could distinguish betweenthe contributions of these resources. Unfortunately proteinpreservation at the site is poor.

However, it should be noted that human consumptionof lean protein is limited to �29–41% of energy consump-tion. At higher levels of protein consumption, hyperammo-nemia and hypereaminoacidemia are likely to resultproducing the clinical symptoms described as ‘‘rabbit star-vation’’ (Cordain et al., 2000). Although animal proteinwas likely supplemented by fats, it is extremely improbablethat the consumption of animal products alone could haveproduced the observed isotopic signatures. For humans toexhibit the observed d13C values solely as the result of car-nivory, they would have to have consumed quantities of C4

animal protein comparable that of the big cats recovered inMember 2 at Swartkrans (Lee-Thorp et al., 1989).

Rather it is more probable that C4 plants directly con-sumed by humans were the source of the enrichment in13C of enamel carbonate. It appears that C4 grasses suchas wild Panicum sp. as well as possibly domestic Pennise-

tum sp. were a significant component of human diet.Humans are significantly depleted in 18O relative to the

fauna at KN05. This finding is consistent with the acquisi-

of human and animal remains from KN05 and KS05.

Table 5Carbon isotope values of modern plant samples obtained in Gao, Mali

Species d13C (&)

Sorghum bicolor �9.1Sorghum arundinaceum �9.0Pennisetum glaucum �9.5Oryza glabberima �25.3Oryza glabberima �25.3

Table 6Estimated average carbon isotope values of the diets consumed atKarkarichinkat Nord

Taxa Mean carbonate Mean collagen Plant Mean diet

Homo �4.0 �14.2Bos 0.8 �8.2 �13.8Sheep? 2.0 �7.0 �12.6Goat? �5.1 �14.1 �19.7Fish �16.8Millet �9.1

Modern fish and millet values have been adjusted by +1.5& to correct forthe industrial effect.


tion of water from distinct sources. Whereas, both Bos sp.and ovi-caprids may have obtained much of their waterfrom plant foods enriched in 18O, humans likely obtainedthe overwhelming majority of their water from the Tilem-si’s river. The lone sample from KS05, which dates to theIslamic era long after the desertification of the region, isan outlier with respect to both its oxygen and carbon iso-tope values. The higher d18O value of this individual mayreflect greater reliance upon plant and animal products assources of moisture relative to surface and ground water.The individual’s lower d13C likely reflects the growingimportance of C3 cereals such as rice (Oryza glabberima).The contemporary Tuareg and Arab inhabitants of theTilemsi Valley obtain rice through exchange with popula-tions living along the Niger River and it is likely their pre-decessors of the Islamic era did the same. The increasedconsumption of animal products derived from C3 consum-ing browsers (i.e. goats) as a result of the region’s increas-ing aridity may also have contributed to the observed shift.Needless to say, generalization from a sample size of one isunwarranted.

There is also an outlier within the ovi-caprid taxon.Whereas three of the ovi-caprids have d13C significantlylower than those of the grazing Bos sp., reflecting a dietof predominantly C3 browse, one individual has a d13Cvalue indistinguishable from the cattle. Modern sheepand goats can be distinguished by the bimodal distributionof carbon isotope values as grazing sheep enriched in 13Crelative to browsing goats (Balasse and Ambrose, 2005).The discrepancy in carbon isotope values observed in thefauna of KN05 may also reflect the ecological differencesbetween goats and sheep. The depleted animals are likelygoats, whereas the single enriched animal is probably asheep.

Conclusions

The evidence from KN05 suggests the site’s inhabitantsexploited a mixed economy of wild and domestic resources.Although we do not yet have conclusive evidence that theC4 cereals consumed at KN05 were domestic, it is clearfrom both the carbon isotope signatures of tooth enameland the grinding slabs that grain was a significant compo-nent of human diet. These findings are consonant with evi-dence for the exploitation of cereals (wild or domestic) bythe Tenerean Culture of neighboring Niger from �5500to 4000 BP (Haour, 2003). However, these d13C values alsospeak to the fact that C3 based resources, including fishfrom the Tilemsi palaeochannel (see the discussion of fishbones in Archaeological Context), contributed to the sub-sistence of KN05’s occupants.

The exploitation of domestic livestock was clearly amajor component of the subsistence at KN05, but the evi-dence from the Tilemsi postdates by several millennia theadoption of animal husbandry in Niger where the remainsof morphologically domestic cattle have been found atTakene Bawat and in the Azawagh dating to �6000 BP(Haour, 2003). The d13C signatures of three of the ovi-capr-ids (probably goats) indicate that the environment sur-rounding KN05 was not exclusively dominated by C4

grasses but also contained an appreciable quantity of C3

plants. Such a finding is consistent with the existence of aSahelian environment in the Tilemsi during the 3rd millen-nium BC and is consonant with paleoenvironmentalrecords from other regions of eastern Mali which indicatethat the Sahel once stretched to the north of the country.

Further research will be necessary to refine our under-standing of the transition to food production in the TilemsiValley. Additional investigation of the materials fromKN05, including the analysis of residues on ceramics, willbe required in order determine whether the grasses con-sumed at the site were wild or domestic and whether milkfrom domestic animals was being consumed by the site’sinhabitants. The excavation of additional stratified con-texts will also be needed in order to obtain a sequence ofhuman remains and thus a biogeochemical record of dietdocumenting diachronic shifts in diet.

Acknowledgments

B.F.’s postgraduate research was funded by the RhodesTrust and K.M.’s postgraduate research was funded by anaward from the Arts and Humanities Research Council(AHRC). The fieldwork was made possible by further con-tributions from the AHRC, the British Institute in EasternAfrica, and the University of Oxford. We also thank theDirection Regionale de la Jeunesse des Sports et Arts, inGao, our local excavation team, and Ayouba Ag Moussi-lim and Souleymane Ag Kiyou for all their invaluable help.Hannes Schroeder prepared the modern fish bones ana-lyzed in this study and helped supply references cited in this


study. Thanks also to Peter Ditchfield for his assistance inthe analysis of enamel carbonate.

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Documents

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)