Sunday, October 2, 2011

Early domesticated cowpea (Vigna unguiculata) from Central Ghana.

Early domesticated cowpea (Vigna unguiculata) from Central Ghana. From examining the remains of charred cowpeas from rock shelters inCentral Ghana, the authors throw light on the subsistence strategies ofthe Kintampo people of the second millennium BCE BCEabbr.1. Bachelor of Chemical Engineering2. Bachelor of Civil EngineeringBCEAbbreviation for before the Common Era. . Perhaps drivensouthwards from the Sahel by aridiflcation, the Kintampo operated asboth foragers and farmers, cultivating selected plants of the WestAfrican tropics tropics,also called tropical zone or torrid zone, all the land and water of the earth situated between the Tropic of Cancer at lat. 23 1-2°N and the Tropic of Capricorn at lat. 23 1-2°S. , notably cowpea cowpea,black-eyed pea,or black-eyed bean,annual legume (Vigna sinensis) of the pulse family. Introduced in the early 18th cent. , pearl millet and oil palm. Keywords: Ghana, Kintampo, domestication domesticationProcess of hereditary reorganization of wild animals and plants into forms more accommodating to the interests of people. In its strictest sense, it refers to the initial stage of human mastery of wild animals and plants. , cowpea, pearl millet, oilpalm, subsistence Introduction Recent research has raised new issues on the nature of subsistenceactivities practised by the Kintampo, a Later Stone Age (LSA) traditionof West Africa dating to 3600-3200 BP (Casey 2005; Gautier & VanNeer 2005; Watson 2005; D'Andrea et al. 2006). The relevant sitesfor the most part are found in present-day Ghana, distributed fromsouthern coastal regions to the far north (Figure 1). In addition toceramics, Kintampo peoples possessed a material culture that includeddaub architecture, rasps, microliths, chipped and ground projectile projectilesomething thrown forward.projectile syringesee blow dart.projectile vomitingforceful vomiting, usually without preceding retching, in which the vomitus is thrown well forward. points, ground stone axes, grinding stones and objets de parure (i.e.bracelets, pendants and quartz beads) (Watson 2005). Settlement patternsare quite varied, ranging from semi-sedentary villages to rock sheltersoccupied on a temporary basis (Davies 1962; Flight 1968; Dombrowski1980; Stahl 1985a & b; Casey 1993; 2000; Watson 2005). Kintamposubsistence has been of interest to many scholars since the early 1960s(e.g. Davies 1962; Flight 1976; Posnansky 1984; Andah 1993; Anquandah1993; Stahl 1993; Casey 2000; 2005; Watson 2005). Studies to date havedemonstrated that a wide array of wild animal and plant resources wereutilised by the Kintampo, many of which continued to be procured bylater populations until recent times (Carter & Flight 1972: 278;Rahtz & Flight 1974: 28; Stahl 1985b: 138-41; Casey 1993: 85-86;2000: 33-37; Gautier & Van Neer 2005: 203-6). Kintampo groups were also in possession of domesticated do��mes��ti��cate?tr.v. do��mes��ti��cat��ed, do��mes��ti��cat��ing, do��mes��ti��cates1. To cause to feel comfortable at home; make domestic.2. To adopt or make fit for domestic use or life.3. a. speciesincluding pearl millet (D'Andrea et al. 2001), a forerunner ofN'dama cattle and caprines (Carter & Flight 1972: 278; Stahl1985b: 138-40; Gautier & Van Neer 2005: 202; Watson 2005: 25).Furthermore, specialised techniques, involving arboricultural ar��bo��ri��cul��ture?n.The planting and care of woody plants, especially trees.arbo��ri��cul practiceshave been suggested for the exploitation of oil palm by Kintampo peoplesof south-central Ghana (D'Andrea et al. 2006). Here we report theidentification of an additional domesticated species, the cowpea,surviving in the form of charred remains recovered from rock shelters atthe 'B-sites' (Boase-sites) (Figure 1) in Central Ghana. Wethen discuss the implications of these new finds for interpretations ofprehistoric West African subsistence during the fourth millennium BP. [FIGURE 1 OMITTED] Archaeobotanical research at the B-sites The B-sites comprise six rock shelter loci loci[L.] plural of locus.lociPlural of locus, see there excavated under theauspices of the Kintampo Archaeological Research Project (KARP), whichbegan in 1998 (Watson & Woodhouse 2001; Watson 2003; 2005). They aresituated in the Buokem Hills, approximately 27km south-west of themodern village of Kintampo in Central Ghana (Figure 1) (Watson 2005:7-10). The local habitat consists of semi-deciduous forest/savannaecotone e��co��tone?n.A transitional zone between two communities containing the characteristic species of each.[eco- + Greek tonos, tension, tone; see tone. , similar to that in which the nearby rock shelters K1 and K6 arelocated (Flight 1968; 1976; Rahtz & Flight 1974; Stahl 1985b). TheB-sites probably represent a single multi-component site occupied atvarious rimes by Kintampo and another ceramic LSA population known asthe Punpun (Watson 2005: 10). Details of stratigraphy stratigraphy,branch of geology specifically concerned with the arrangement of layered rocks (see stratification). Stratigraphy is based on the law of superposition, which states that in a normal sequence of rock layers the youngest is on top and the oldest on the , excavationmethods and flotation sampling procedures are summarised elsewhere(Watson 2005: 7-10; D'Andrea et al. 2006). The B-sites have been intensively sampled for archaeobotanicalremains (Watson 2005: 10; D'Andrea et al. 2006; here Table 1). Atotal of 40 flotation samples was examined which produced fiveidentified taxa including oil palm endocarp and kernels (E. guineensis),incense tree endocarp (C. schweinfurthii), cowpea (Vigna unguiculata Vigna unguiculataplant in the legume family Fabaceae; high-protein legume; used as hay or the seed is used as a concentrate feed. Can cause nitrate-nitrite poisoning. Called also V. catjang, V. sesquipedalis, V. sinensis, cowpea. var. unguiculata), seeds belonging to the pea family (Fabaceae), andpearl millet (Pennisetum glaucum) (the term 'seeds' hereincludes a number of botanical elements such as cotyledons, fruits,pericarp, etc.). Other categories include unidentifiable seeds, probableoil palm endocarp (cf. E. guineensis), probable incense tree endocarp(cf. C. schweinfurthii), unidentifiable endocarp and unidentifiableplant remains. These archaeobotanical data indicate that the loci mayrepresent, in part, specialised sites for the production of palm oil(D'Andrea et al. 2006). One cowpea specimen from B6B, Horizon 4 wassubmitted for radiocarbon dating by AMS AMS - Andrew Message System (Accelerator Mass Spectrometry accelerator mass spectrometryn.Mass spectroscopy in which a particle accelerator is used to disassociate molecules, ionize atoms, and accelerate the ions. ),with a resulting date of 3410 [+ or -] 60 bp, 1830-1595 cal BC at 95.5%c.i. (TO 11883). This date confirms a Kintampo association and isconsistent with other AMS determinations obtained from B6B (Table 2). [FIGURE 2 OMMITTED] B-Sites cowpea remains A total of 16 fragmentary specimens identified as domesticatedcowpea (V. unguiculata var. unguiculata) is present in the B-sitesflotation samples. An additional 11 specimens have been identified asprobable (cf.) cowpea (Table 1). Illustrations of Bsites cowpeas arepresented in Figures 2A-C and 3A & B. Unfortunately only threefragile specimens were complete, making it possible to obtain no morethan minimal metrical met��ri��cal?adj.1. Of, relating to, or composed in poetic meter: metrical verse; five metrical units in a line.2. Of or relating to measurement. data. All three specimens fall into the sameranges as domesticated cowpeas identified from archaeological sites inBurkina Faso (Table 3) (Kahlheber 2004). The B-sites specimens displayseveral features indicative of domesticated cowpea (Figure 4A). Theylack lateral compression present in some accessions of the wildprogenitor pro��gen��i��torn.1. A direct ancestor.2. An originator of a line of descent.progenitorancestor, including parent.progenitor cellstem cells. , V. unguiculata var. spontanea (Figure 4B). Instead charredcotelydons are asymmetric in outline and orate o��rate?intr.v. o��rat��ed, o��rat��ing, o��ratesTo speak in a formal, often pompous manner.[Latin in cross-section. Thehilum hilum/hi��lum/ (hi��lum) pl. hi��la ? [L.] a depression or pit on an organ, giving entrance and exit to vessels and nerves.hi��lar hi��lumn. pl. is broad and relatively large (unlike those of Sesbania, Tephrosiaand Rhynchosia), located at or near the centre of the seed (unlike manywild Vigna species), and situated very dose to the chalaza (Figure 5).The ventral side tends to be flattened and the dorsal side extended,resulting in lower B/H (breadth/height) ratios in comparison to wild V.unguiculata. Other possible Fabaceae candidates native to tropicalforest margin areas have been excluded based on differences in seed sizeand shape, including Bambara ground nut (Vigna subterranea) andKersting's ground nut (Macrotyloma geocarpum) (Figure 4C & D).Another feature that appears to be characteristic of charred cowpea iswrinkles present on the inner surface of cotyledons, as illustrated inFigure 2C; however the diagnostic value of this criterion is unknown(Kahlheber 2004). [FIGURE 3 OMITTED] Cowpea ecology and domestication history Cowpea is a widely grown crop in the semi-arid and sub-humid zonesof Africa and Asia. It is the most significant legume legume(lĕ`gym, lĭgy crop grown inAfrica today, particularly in West and Central Africa which typicallydominate world production (Faris 1965: 433; Rawal 1975; Lambot 2002:368). Cowpea constitutes a major dietary source of protein for manysub-Saharan populations. The cotyledons contain approximately 22-27 percent protein and provide a well-balanced complement of essential aminoacids (Busson et al. 1959: 13-14; Lambot 2002: 368-9). When mixed withcereals, protein quality is significantly improved. Leaves are anexcellent source of beta-carotene, ascorbic acid and folacin folacin:see vitamin. (Johnson& Raymond 1964: 71-2; Bressani 1985: 356-7; Burkill 1995: 479;Nielsen et al. 1997: 328-9). Cowpeas are consumed by humans andlivestock in a variety of ways. In human diets, the seeds are eatenafter boiling or roasting and they can be ground into flour to makebreads. Green pods are consumed fresh and the leaves are grown as anutritious potherb or are dried and made into soups. Various parts ofthe plant are used medicinally, and in making dyes and textiles (Hebbs& Hill 1946; Burkill 1995: 479). As a green fodder, cowpea isparticularly advantageous because individual plants continue to grow andproduce leaves even after they have been cut back several times (Irvine1956: 35; Rachie & Roberts 1974: 61; Westphal 1974: 231; Burkill1995: 479). [FIGURE 4 OMITTED] Cowpea is primarily a savanna savannaor savannah(both: səvăn`ə), tropical or subtropical grassland lying on the margin of the trade wind belts. species, highly adapted to marginalenvironments and depauperate field conditions where other crops do notperform well. It is a dependable food security crop in semiarid andforest-margin regions of tropical West and Central Africa (Rachie &Roberts 1974: 45; Westphal 1974: 213; Skerman et al. 1988: 479; Lambot2002: 368). Early maturing varieties can produce a harvest in as littleas two or three months with green pods ready after 50 days (Purseglove1968: 326; Rachie & Roberts 1974: 45). Sizeable yields can beachieved without the use of fertilizers and with minimal tending(Skerman et al. 1988: 479-80). Rapid flowering may represent anadaptation to short and variable growing seasons of the northernsavannas. Cowpea is commonly intercropped with pearl millet and sorghum sorghum,tall, coarse annual (Sorghum vulgare) of the family Gramineae (grass family), somewhat similar in appearance to corn (but having the grain in a panicle rather than an ear) and used for much the same purposes. (Sorghum bicolor bicolora coat color of two colors. In dogs, usually black with tan markings but may be other combinations such as ticking on a white background. In cats, more than two spots of color on the body, either white and one basic color, or white with one tabby color. ) and also cultivated in pure stands (Rachie &Roberts 1974: 45; Purseglove 1968: 326; Westphal 1974: 230; Lush &Evans 1981: 583). [FIGURE 5 OMITTED] Ng (1995: 329) has proposed that humans initially collected wildcowpea as fodder for livestock. He posits that early cattle-herdersgathered entire plants by uprooting, an activity which would havecreated selection pressure for indehiscent pods. Cowpeas were eventuallyplanted for human consumption, at which time people selected fornon-shattering pods, reduced seed dormancy and larger seeds and pods. Ifcowpea resembles other pulses, however, increases in seed and pod sizesmay have been delayed (Ladizinsky 1987: 61-4; Fuller & Harvey 2006:256-7). Short-day cultivars (needing less than 12 hours of daylight inwhich to flower) probably evolved first in the savannas and were adaptedto cereal farming systems. These types are normally found in higherlatitudes where maturation occurs toward the end of the rainy season,allowing the plant to take full advantage of available moisture duringflowering (Rachie & Roberts 1974: 55; Ng 1995: 329). Day-neutralcultivars evolved later through introgression in��tro��gres��sion?n.Infiltration of the genes of one species into the gene pool of another through repeated backcrossing of an interspecific hybrid with one of its parents. with wild subspecies subspecies,also called race, a genetically distinct geographical subunit of a species. See also classification. present in the humid tropics, and were incorporated into yam-basedfarming systems (Steele & Mehra 1980: 398; Lush & Evans 1981:585; Ng 1995: 329). The scenario proposed by Ng (1995) may have somemerit, given that livestock were domesticated several thousand yearsbefore crops in Africa (Marshall & Hildebrand 2002). However, todate there is no archaeological evidence for domesticated cowpea amongearly agro-pastoralist populations in the savanna/Sahel regions,including at the site of Birimi which was extensively sampled formacrobotanical remains (D'Andrea & Casey 2002). Furthermore itmay be less likely that Kintampo groups were growing cowpea as a fodderbased on zooarchaeological data which indicate that livestock were notpresent in significant numbers (Casey 2005; Gautier & Van Neer2005). The domestication history of cowpea remains to be fully elucidated.Complicating factors include the enormous variability of morphologicalfeatures in both wild and domesticated forms and the wide geographicaldistribution of the wild progenitor. Hybridisation between cultivatedand wild types has also contributed to increasing diversification(Pasquet 1996; 1999; 2000; Coulibaly et al. 2002; Ba et al. 2004). Thisdegree of variation has resulted in the publication of several taxonomictreatments (e.g. Verdcourt 1970; Rawal 1975: 700; Mar,chal et al. 1978;Lush & Evans 1981; Ng & Mar,chal 1985; Pasquet 1993a & b;1997; 1999; Coulibaly et al. 2002). The absence of the wild progenitoroutside Africa indicates that cowpea was domesticated somewhere on theAfrican continent, and several domestication locales have been proposed(Murdock 1959; Faris 1965: 449; Sauer 1969; Vaillancourt & Weeden1992:1194). Phylogenetic phy��lo��ge��net��icadj.1. Of or relating to phylogeny or phylogenetics.2. Relating to or based on evolutionary development or history. studies and linguistic evidence indicate thatthe crop originated in tropical West Africa (Faris 1965; Rawal 1975:706; Mar,chal et al. 1978: 191-6; Ng & Mar,chal 1985; Vaillancourt& Weeden 1992:1198; Panella et al. 1993: 384; Blench blench?1?intr.v. blenched, blench��ing, blench��esTo draw back or shy away, as from fear; flinch.[Middle English blenchen, from Old English blencan, 1994-95: 92; Ng1995: 329), with a second possible centre of origin in Botswana (Panellaet al. 1993: 383-4). Recent studies using amplified fragment lengthpolymorphism Amplified fragment length polymorphism PCR, or "AFLP-PCR" (often AFLP), is a tool used in the study of genetics and in the practice of genetic engineering.Amplified Fragment Length Polymorphism (AFLP analysis (AFLP) have concluded that the two most likelyareas of domestication are tropical West or north-eastern Africa(Coulibaly et al. 2002: 365). This work and a subsequent study usingrandom amplified polymorphic DNA DNA:see nucleic acid. DNAor deoxyribonucleic acidOne of two types of nucleic acid (the other is RNA); a complex organic compound found in all living cells and many viruses. It is the chemical substance of genes. (RAPD RAPD Randomly Amplified Polymorphic DNARAPD relative afferent pupillary defect (ophthalmology; aka Marcus-Gunn Pupil)) has ruled out the possibility ofdomestication in southern and eastern Africa (Ba et al. 2004: 547). Anorthern or north-eastern origin has been argued based on the absence oftrue ecologically wild cowpea in West Africa (Pasquet & Baudoin2001: 187-88; Coulibaly et al. 2002: 365). However, the paucity ofavailable accessions of the wild progenitor from Sudan, Eritrea andEthiopia (Pasquet 1999:1118), as well as questions about the efficacy ofAFLP and RAPD methods to infer crop origins (Allaby & Brown 2003;2004; Salamini et al. 2004) have precluded a final assessment. More dataare necessary, therefore, to demonstrate a specific area of origin forcowpea (Panella et al. 1993: 384; Pasquet 1999: 1118; Coulibaly et al.2002: 365; Ba et al. 2004: 547). In any case, it has been determinedthat a single domestication event occurred in a circumscribed region(Panella & Gepts 1992; Pasquet 1999; Coulibaly et al. 2002: 364; Baet al. 2004: 547). The B-sites AMS date of 1830-1595 cal BC represents the earliestarchaeological evidence to date of cowpea in Africa, and as such, thisdetermination lends some support to a West African origin for this crop.It is difficult, however, to build a strong case based on a singleradiocarbon age estimate. The B-sites specimens predate the earliestsecurely dated cowpea in India at 1700-1500 cal BC (Fuller 2003: 242-3).This points to an early and rapid spread of cowpea to India. However,additional genetic as well as baseline archaeological data are requiredto determine the location and timing of cowpea domestication in Africa. Cowpea and Kintampo subsistence Cowpeas may also be present at K6, where they are described asbeing very common in Kintampo deposits. Based on their abundance, Flight(1976: 217-18) argues that they were cultivated, but there wasinsufficient evidence insufficient evidencen. a finding (decision) by a trial judge or an appeals court that the prosecution in a criminal case or a plaintiff in a lawsuit has not proved the case because the attorney did not present enough convincing evidence. to determine whether seeds came from domesticatedplants. He further contends that the small size of the K6 cowpeas wasthe result of shrinkage due to charring (Flight 1976: 218). In laterexcavations at K6, Stahl (1985b: 141; 1994: 76) recovered four legumeseeds which bore a resemblance to cowpea, but at roughly 5mm in lengththey were judged to be smaller than available reference specimens. As aresult, they were conservatively identified to the family Leguminosae(=Fabaceae). Recent work has shown that a length of 5mm falls within therange of domesticated cowpea land races (Kahlheber 2004). In any case,low numbers of cowpeas in Kintampo archaeological contexts may notnecessarily reflect their economic insignificance in��sig��nif��i��cance?n.The quality or state of being insignificant.Noun 1. insignificance - the quality of having little or no significanceunimportance - the quality of not being important or worthy of note . Pulses are oftenunder-represented in archaeological sites because they can be highlysusceptible to deterioration by breaking down into two cotyledons(Gasser & Adams 1981: 183-4). It has recently been proposed that the Kintampo people were derivedfrom Sahelian foragers who were acculturated into an agro-pastoraleconomy during a long period of coexistence of forager-fishers andpastoralists (Watson 2005: 28-32). Modifications of subsistence regimesprobably took place as they occupied more humid zones. This would haveincluded the adoption of forest resources such as oil palm and incensetree (D'Andrea et al. 2006), and several species of wild fauna.North of the B-sites at Birimi, the cultivation of pearl millet amongKintampo occupants differed from those of agro-pastoral peoples presentfurther north in the Sahel (Klee et al. 2000: 232; Klee & Zach 1999:87; Neumann 1999: 78-9; Vogelsang et al. 1999: 60-1; Kahlheber &Neumann, in press). Late-maturing forms of pearl millet were selectedfrom a secondary centre of diversity that developed around Lake Chad(Tostain 1998:481). These forms, which are highly adapted to savannahabitats, were probably introduced to the region near Birimi during anarid phase when the crop was displaced south of the Sahel (D'Andreaet al. 2002: 160-2). Although the area of cowpea origin remains to bedetermined, if initially domesticated in a savanna habitat, andday-neutral cultivars evolved as the crop was introduced to the forestzone (Ng 1995: 329), the B-sites cowpeas may have arrived in centralGhana as a result of this process. Furthermore these introductions mayhave taken place over a similar time frame, with the earliest knownpearl millet at Birimi dating to 3460 [+ or -] 200 BP (D'Andrea etal. 2001: 343) and the B-sites cowpeas to 3410 [+ or -] 60 BE However,additional ANIS dates of Kintampo pearl millet and cowpea are necessaryto determine whether this was a widespread phenomenon. Subsistence links to savanna, Sahel and northern regions areevident in domesticated species identified in Kintampo contexts,including cattle, caprines, pearl millet and cowpea. Both pearl milletand cowpea are early-maturing, arid-adapted crops that tolerate poorenvironments, and are able to produce respectable harvests with minimaltending (Skerman et al. 1988: 479-80). Such crops would seem to beparticularly well-suited to populations with higher mobility, such asagro-pastoralists occupying savanna regions following the onset ofaridification c. 4500 BP, who may have been looking to minimise risk andestablish predictable food supplies (Gifford-Gonzalez 2005: 204-6;Marshall & Hildebrand 2002). Biological indicators at Kintampo sitesthat demonstrate the use of locally available forest resources are all,strictly speaking, wild species, including land snails, freshwater fish,reptiles, large rodents, antelopes, primates, oil palm, incense tree andCeltis (Stahl 1985b: 139-41; Gautier & Van Neer 2005:196-202;D'Andrea et al. 2006). The available archaeobotanical andzooarchaeological evidence attests to a heterogeneous origin of foodsprocured and produced by the Kintampo, representing a mosaic ofeffective solutions taken to obtain both wild and domesticatedresources. Farmers or foragers? How is one to characterise such a wide-ranging set of subsistencestrategies evidently practised by Kintampo peoples? Recently,investigators have drawn attention to the diversified nature of thesubsistence economy (Casey 2005: 236-9; Watson 2005: 30-1; D'Andreaet al. 2006), and some have argued that it is not possible to define theKintampo and other African prehistoric societies (Casey 2005: 240;Neumann 2005: 249) as strictly 'foragers' or'farmers'. This is a situation that is not unique to Africa,which, like many other regions of the world, provides unambiguousexamples of prehistoric foraging and farming societies, along withcultures that do not fit neatly into such categories (cf. Smith 2001;Zeder 2006). The existing evidence suggests that the Kintampo economy,at the very least, can be characterised by 'low level foodproduction with domesticates' (Smith 2001: 14-17); however it wouldseem premature at this stage to arrive ata conclusion on the economicimportance of wild vs. domesticated resources. Despite new dataavailable on species utilised by the Kintampo, overall only a handful ofsites has been systematically sampled for plant and animal remains.Archaeobotanical studies have been completed at only three sites (Stahl1985 a: 141; D'Andrea et al. 2001; 2006; D'Andrea & Casey2002), while zooarchaeological interpretations (Stahl 1985a: 210-4;Gautier & Van Neer 2005: 201-2) are based on very low MNIs (MinimumNumber of Individuals) of both domesticated and wild animals WILD ANIMALS. Animals in a state of nature; animals ferae naturae. Vide Animals; Ferae naturae. (Stahl1985a: Table 4.12). Although the agro-pastoral component of the economyis now known to have been significantly more complex than was previouslyunderstood, at this point in time there appears to be no firm basis uponwhich to conclude that either farming or foraging was the dominantsubsistence activity practised by the Kintampo. Acknowledgements Laboratory analysis and AMS determinations completed on the B-sitesmacro-remains were funded by a Simon Fraser University Simon Fraser University,main campus at Burnaby, British Columbia, Canada; provincially supported; coeducational; chartered 1963, opened 1965. The Harbour Centre campus in downtown Vancouver opened in 1989. InstitutionalGrant to ACD (Automatic Call Distributor) A computerized phone system that responds to the caller with a voice menu and connects the call to the appropriate agent. It can also distribute calls equally to agents. . Caroline Lefebvre and Meridith Sayre (Simon FraserUniversity) assisted in the sorting of B-sites samples, throughfinancial support provided by the Simon Fraser University Work StudyProgramme. Ali drawings were completed by Barbara Voss (University ofFrankfurt University of Frankfurt may refer to two (or three) German universities: the Johann Wolfgang Goethe University of Frankfurt am Main ("Johann Wolfgang Goethe-Universit?t Frankfurt am Main") in Frankfurt am Main ). We benefited immensely from several informative discussionsabout cowpeas with Dr Don Faris (formerly of the Research Branch,Agriculture Canada). In addition, we are grateful to Dr ThierryVanderborght (National Botanic Garden of Belgium, Meise) who providedVigna reference specimens for comparative purposes. We thank Dr LaurenceA. Pavlish (University of Toronto Research at the University of Toronto has been responsible for the world's first electronic heart pacemaker, artificial larynx, single-lung transplant, nerve transplant, artificial pancreas, chemical laser, G-suit, the first practical electron microscope, the first cloning of T-cells, ) who read and commented on variousdrafts of this paper and Shannon Wood (Simon Fraser University) forassistance with Figure 1. Helpful suggestions were also made by twoanonymous reviewers. Dr R,my Pasquet (Institut de Recherche pour leD,veloppement, Nairobi) kindly sent us difficult-to-acquire referencespecimens, and we appreciate his patience as we discussed at length theB-sites cowpeas. References ALLABY, R.G. & T.A. BROWN. 2003. AFLP Data and the Origins ofDomesticated Crops. Genome 46: 448-53. --2004. Reply to Comment by Salamini et al. on 'AFLP Data andthe Origins of Domesticated Crops'. Genome 47: 621-2. ANDAH, B.W. 1993. 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QualitasPlantarum et Materiae Vegetabiles 6:11-5. CARTER, P.L. & C. FLIGHT. 1972. A Report on the Fauna from theSites of Ntereso and Kintampo Rock Shelter Six in Ghana: With evidencefor the Practice of Animal Husbandry animal husbandry,aspect of agriculture concerned with the care and breeding of domestic animals such as cattle, goats, sheep, hogs, and horses. Domestication of wild animal species was a crucial achievement in the prehistoric transition of human civilization from During the Second Millennium BC.Man 7(2): 277-82. CASEY, J. 1993. The Kintampo Complex in Northern Ghana: LateHolocene Human Ecology on the Gambaga Escarpment escarpmentor scarp,long cliff, bluff, or steep slope, caused usually by geologic faulting (see fault) or by erosion of tilted rock layers. An example of a fault scarp is the north face of the San Jacinto Mts. in California. . Unpublished PhDThesis, University of Toronto. --2000. The Kintampo Complex: The Late Holocene on the GambagaEscarpment, Northern Ghana (BAR International Series 906, CambridgeMonographs in African Archaeology). Oxford: Archaeopress. --2005. Holocene occupations of the forest and savanna, in A.B.Stahl (ed.) African archaeology: 225-48. Oxford: Blackwell. COULIBALY, S., R.S. PASQUET, R. PAPA & P. GEPTS. 2002. AFLPAnalysis of the Phenetic phe��net��ic?adj.Of, relating to, or designating a system of classification of organisms based on overall or observable similarities rather than on phylogenetic or evolutionary relationships. Organization and Genetic Diversity of Vignaunguiculata L. Walp. Reveals Extensive Gene Flow Between Wild andDomesticated Types. Theoretical and Applied Genetics 104: 358-66. D'ANDREA, A.C. & J. CASEY. 2002. Pearl Millet and KintampoSubsistence. African Archaeological Review 19(3): 147-73. D'ANDREA, A.C., M. KLEE &J. CASEY. 2001. ArchaeobotanicalEvidence for Pearl Millet (Pennisetum glaucum) in Sub-Saharan WestAfrica. Antiquity 75: 341-8. A.C. D Andrea (1), S. Kahlheber (2), A.L. Logan (3) & D.J.Watson (4) (1.) Department of Archaeology, Simon Fraser University, Burnaby,British Columbia, Canada V5A 1S6 (2.) J.W. Goethe-Universitat, Institut fur ArcheologischeWissenschafien, Gruneburgplatz 1, D-60323 Frankfurt, Germany (3.) Museum of Anthropology, University of Michigan (body, education) University of Michigan - A large cosmopolitan university in the Midwest USA. Over 50000 students are enrolled at the University of Michigan's three campuses. The students come from 50 states and over 100 foreign countries. , 1109 GeddesAvenue, Ann Arbor, Michigan “Ann Arbor” redirects here. For other uses, see Ann Arbor (disambiguation).Ann Arbor is a city in the U.S. state of Michigan and the county seat of Washtenaw County. , USA 48109 (4.) 183 Castlemilk Drive, Glasgow G45 9JT, UKTable 1. Macrobotanical remains from 134C, B5C and 136B. Seed Count Pennisetum Vigna cf. V. glaucum unguicilata unguicculata CONTEXTLOCUS B4C Layer 1 -- -- -- Layer 2 -- -- -- Layer 3 -- -- -- B4C Total 0 0 0LOCUS B5C Horizon 1 -- -- -- Horizon 2 -- -- -- Horizon 3 1 -- 2 B5C Total 1 0 2LOCUS 13613 Horizon 1 -- -- -- Horizon 2 -- 1 Horizon 3 -- -- 0 Horizon 4 -- 15 6 B6B Total 0 16 16TOTAL 1 16 11 ALL SITES Unidentifiable Total Seed CONTEXT Fabaceae Seed Fragments CountLOCUS B4C Layer 1 -- -- 0 Layer 2 -- -- 0 Layer 3 -- 1 1 B4C Total 0 1 1LOCUS B5C Horizon 1 -- 14 14 Horizon 2 -- -- 4 Horizon 3 -- 25 28 B5C Total 0 43 46LOCUS 13613 Horizon 1 -- -- 0 Horizon 2 5 Horizon 3 11 3 Horizon 4 6 48 B6B Total 922 56TOTAL 9 66 103 ALL SITES Elaeis cf. E. guineensis E. guineensis guineensis CONTEXT endocarp kernels endocarpLOCUS B4C Layer 1 3.30 -- 0.85 Layer 2 2.69 -- 1.12 Layer 3 0.13 -- -- B4C Total 6.12 0.00 1.97LOCUS B5C Horizon 1 4.87 -- 2.10 Horizon 2 29.10 -- 2.37 Horizon 3 15.22 -- 0.00 B5C Total 49.19 0.00 4.47LOCUS 13613 Horizon 1 26.05 -- 1.31 Horizon 2 46.15 -- 2.55 Horizon 3 47.20 -- -- Horizon 4 500.70 0.18 2.75 B6B Total 620.10 0.18 6.61TOTAL 675.41 0.18 13.05 ALL SITES Weight (g) Canarium cf. E. schweinfurthii schweinfurthii Unidentiafiable CONTEXT endocarp endocarp EndocarpLOCUS B4C Layer 1 -- -- -- Layer 2 -- -- -- Layer 3 -- -- -- B4C Total 0.00 0.00 0.00LOCUS B5C Horizon 1 1.62 0.09 3.29 Horizon 2 0.42 0.62 7.74 Horizon 3 1.03 0.20 21.28 B5C Total 3.07 0.91 32.31LOCUS 13613 Horizon 1 0.28 0.01 9.24 Horizon 2 0.56 -- 12.72 Horizon 3 -- -- 0.38 Horizon 4 2.21 0.38 30.12 B6B Total 3.05 0.39 52.46TOTAL 6.12 1.30 84.77 ALL SITES Total Unidentiafiable CONTEXT Endocarp Charcoal Plant RemainsLOCUS B4C Layer 1 4.15 1.99 0.07 Layer 2 3.81 10.93 -- Layer 3 0.13 11.67 0.77 B4C Total 8.09 24.59 0.84LOCUS B5C Horizon 1 11.97 3.83 0.02 Horizon 2 40.25 2.91 0.02 Horizon 3 37.73 8.54 nil B5C Total 89.95 15.28 0.04LOCUS 13613 Horizon 1 36.89 3.30 0.18 Horizon 2 61.98 1.68 0.28 Horizon 3 47.58 0.30 -- Horizon 4 536.34 1.26 0.35 B6B Total 682.79 6.54 0.81TOTAL 780.83 46.41 1.69 ALL SITES Average E. guinnensis CONTEXT Density SedimentLOCUS B4C Layer 1 0.22 20.15 Layer 2 0.37 10.25 Layer 3 0.02 8.00 B4C Total -- 38.40LOCUS B5C Horizon 1 0.09 66.19 Horizon 2 1.51 32.26 Horizon 3 0.38 67.64 B5C Total -- 166.09LOCUS 13613 Horizon 1 2.38 14.00 Horizon 2 3.60 11.52 Horizon 3 4.44 10.65 Horizon 4 12.53 27.54 B6B Total -- 63.71TOTAL -- 268.19 ALL SITESTable 2. Locus B6B AMS determinations.Unit Horizon Lab. No. Material2 5 GX 28710 b.v. collagen2 4 TO 11883 cowpea cotyledon2 4 TO 11238 oil palm endocarp1 2 GX 29106 charcoalUnit Depth (cm) Date (uncal. bp)2 50 3230 [+ or -] 402 70 3410 [+ or -] 602 70 3460 [+ or -] 501 101 3470 [+ or -] 40Unit cal. BC * cal. BP *2 1610-1410 3560-33602 1830-1595 3780-35452 1885-1680 3835-36301 1890-1680 3840-3630* 95.5% confidence interval.Table 3. Metrical data for West African V. unguiculatavar. unguiculata.Dimensions (mm) B-sites Burkina Faso (Kahlheber 2004)Length 5.0-6.3 3.7-9.7Breadth 3.6-4.6 2.4-6.6Height 3.4-4.2 3.0-6.1Length:Height 1.5-1.6 0.9-2.1Breadth:Height 0.9-1.1 0.7-1.4Length:Breadth 1.4-1.8 1.0-2.3

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