Abstract
Large-scale reconstructions of the spread of food production systems require fine-scale analyses of dietary evidence. One current impediment to understanding early African pastoralism is a lack of high-resolution portraits of herd management, specifically with respect to sheep (Ovis aries) and goat (Capra hircus), osteologically similar but behaviorally distinct caprines. In this study, we argue for the anthropological relevance of distinguishing sheep and goat remains in African pastoralist contexts, commenting upon implications for ecological settings and pastoralists’ strategies for production and risk management. We explain why sheep/goat distinction is rare in African zooarchaeological studies, particularly in comparison to Southwest Asia. We then apply three methods to distinguish caprines in an archaeofaunal sample from the Pastoral Neolithic site of Luxmanda, Tanzania, dated to c. 3000 BP: morphological identifications by two independent analysts, collagen-peptide mass fingerprinting (Zooarchaeology by Mass Spectrometry, ZooMS), and carbon stable isotope analyses. A comparative assessment of the results demonstrates the ability of biomolecular methods to improve resolution of faunal records, particularly where preservation is poor. We call for wider application of these methods to legacy collections, in order to refine existing regional models for the spread of herding in Africa, and to better understand ancient pastoralists’ herd-management decisions.
Similar content being viewed by others
Notes
“Indigenous” refers to unimproved breeds, rather than implying that the domestic goat and sheep are indigenous to Africa. Selective breeding of sheep and goat (i.e., breed improvement) has increased over the past few centuries and has involved introductions of breeds from other world regions.
References
Ambrose SH (1998) Chronology of the Later Stone Age and food production in East Africa. J Archaeol Sci 25:377–392
Ambrose SH, Sikes NE (1991) Soil carbon isotope evidence for Holocene habitat change in the Kenya Rift Valley. Science 253:1402–1405
Arbuckle BS, Atici L (2013) Initial diversity in sheep and goat management in Neolithic South-Western Asia. Levant 45:219–235. https://doi.org/10.1179/0075891413Z.00000000026
Atici L, Kansa SW, Lev-Tov J, Kansa EC (2013) Other people's data: a demonstration of the imperative of publishing primary data. J Archaeol Method Theory 20:663–681
Badenhorst S (2002) The ethnography, archaeology, rock art and history of goats (Capra hircus) in southern Africa: an overview. Anthropol South Afr 25:96–103
Badenhorst S (2006) Goats (Capra hircus), the Khoekhoen and pastoralism: current evidence from Southern Africa. Afr Archaeol Rev 23:45–53. https://doi.org/10.1007/s10437-006-9007-0
Badenhorst S (in press) Exploitation of sheep (Ovis aries) and goats (Capra hircus) by Iron Age farmers in southern Africa. Quat Int. https://doi.org/10.1016/j.quaint.2017.12.023
Badenhorst S, Plug I (2003) The archaeozoology of goats, Capra hircus (Linnaeus, 1758): their size variation during the last two millennia in southern Africa (Mammalia: Artiodactyla: Caprini). Ann Transv Mus 40:91–121
Balasse M, Ambrose SH (2005) Distinguishing sheep and goats using dental morphology and stable carbon isotopes in C4 grassland environments. J Archaeol Sci 32:691–702
Balasse M, Ambrose SH, Smith AB, Price TD (2002) The seasonal mobility model for prehistoric herders in the South-western Cape of South Africa assessed by isotopic analysis of sheep tooth enamel. J Archaeol Sci 29:917–932
Balasse M, Obein G, Ughetto-Monfrein J, Mainland I (2012) Investigating seasonality and season of birth in past herds: a reference set of sheep enamel stable oxygen isotope ratios. Archaeometry 54(2):349–368
Bar-Gal GK, Ducos P, Horwitz L (2003) The application of ancient DNA analysis to identify neolithic caprinae: a case study from the site of Hatoula, Israel. Int J Osteoarchaeol 13:120–131. https://doi.org/10.1002/oa.666
Barthelme JW (1985) Fisher-Hunters and Neolithic Pastoralists in East Turkana, Kenya. British Archaeological Reports International Series 254. British Archaeological Reports, Oxford, England
Bekure S, de Leauw PN, Grandin BE, Neate PJH (1991) Maasai herding: an analysis of the livestock production system of Maasai pastoralists in eastern Kajiado District, Kenya. ILCA Systems Study 4. International Livestock Centre for Africa, Addis Ababa
Blackburn H, Field C (1990) Performance of Somali blackhead sheep and Galla goats in northern Kenya. Small Rumin Res 3:539–549
Boessneck J (1969) Osteological difference between sheep and goats. In: Brothwell D, Higgs E (eds) Science in archaeology: a survey of progress and research. Thames and Hudson, Bristol, pp 331–358
Bollig M (2016) Adaptive cycles in the savannah: pastoral specialization and diversification in northern Kenya. J East Afr Stud 10(1):21–44
Bousman C, Mauldin R, Zoppi U, Higham T, Scott L, Brink J (2016) The quest for evidence of domestic stock at Blydefontein Rock Shelter. Southern African Humanities 28, 39-60.
Buckley M, Collins M, Thomas-Oates J (2008) A method of isolating the collagen (I) α2 chain carboxytelopeptide for species identification in bone fragments. Anal Biochem 374:325–334
Buckley M, Collins M, Thomas-Oates J, Wilson JC (2009) Species identification by analysis of bone collagen using matrix-assisted laser desorption/ionisation time-of-flight mass spectrometry. Rapid Commun Mass Spectrom 23:3843–3854
Buckley M, Kansa SW (2011) Collagen fingerprinting of archaeological bone and teeth remains from Domuztepe, South Eastern Turkey. Archaeol Anthropol Sci 3:271–280. https://doi.org/10.1007/s12520-011-0066-z
Buckley M, Whitcher Kansa S, Howard S, Campbell S, Thomas-Oates J, Collins M (2010) Distinguishing between archaeological sheep and goat bones using a single collagen peptide. J Archaeol Sci 37:13–20
Cerling TE, Harris JM (1999) Carbon isotope fractionation between diet and bioapatite in ungulate mammals and implications for ecological and paleoecological studies. Oecologia 120:347–363
Connor RJ (1994) African animal trypanosomiases. In: Coetzer JAW, Thomson GR, Tustun RC, Kriek NPJ (eds) Infectious diseases of livestock with special reference to Southern Africa, vol 1. Oxford University Press, New York, pp 167–205
Dahl G, Hjort A (1976) Having herds. Pastoral herd growth and household economy. Stockholm University, Stockholm
De Boom HPA (1950) Die onderskeiding tussen skaap en bokbene/Differentiating between bones of the sheep and the goat The Nongqai July:801–813
Dyson-Hudson R, Dyson-Hudson N (1980) Nomadic pastoralism. Annu Rev Anthropol 9:15–61
Galvin KA (2009) Transitions: pastoralists living with change. Annu Rev Anthropol 38:185–198
Gifford D, Isaac G, Nelson CM (1980) Evidence for predation and pastoralism at Prolonged Drift: a Pastoral Neolithic site in Kenya. Azania 15:57–108
Gifford-Gonzalez D (2000) Animal disease challenges to the emergence of pastoralism in Sub-Saharan Africa. Afr Archaeol Rev 17:95–139
Gifford-Gonzalez D (2017) “Animal disease challenges” fifteen years later: the hypothesis in light of new data. Quat Int 436:283–293
Gifford-Gonzalez D, Hanotte O (2011) Domesticating animals in Africa: implications of genetic and archaeological findings. J World Prehist 24:1–23
Gifford-Gonzalez D, Hanotte O (2013) Domesticating Animals in Africa. In: Mitchell P, Lane PJ (eds) The Oxford Handbook of African Archaeology, pp. 491-505. New York: Oxford University Press.
Gobalet KW (2001) A critique of faunal analysis; inconsistency among experts in blind tests. J Archaeol Sci 28:377–386
Greenfield HJ (2010) The secondary products revolution: the past, the present and the future. World Archaeol 42:29–54
Grillo K, Hildebrand E (2013) The context of early megalithic architecture in eastern Africa: the Turkana Basin c. 5000-4000 BP. Azania Archaeol Res Afr 48:193–217. https://doi.org/10.1080/0067270X.2013.789188
Grillo KM, Prendergast ME, Contreras DA, Fitton T, Gidna AO, Goldstein ST, Knisley MC, Langley MC, Mabulla AZP (2018) Pastoral Neolithic settlement at Luxmanda, Tanzania. J Field Archaeol 43:102–120
Halstead P, Collins P, Isaakidou V (2002) Sorting the sheep from the goats: morphological distinction between the mandibles and mandibular teeth of adult Ovis and Capra. J Archaeol Sci 29:545–553
Haruda AF (2017) Separating sheep (Ovis aries L.) and goats (Capra hircus L.) using geometric morphometric methods: an investigation of astragalus morphology from Late and Final Bronze Age Central Asian contexts. Int J Osteoarchaeol 27:551–562
Hauck S, Rubenstein DI (2017) Pastoralist societies in flux: a conceptual framework analysis of herding and land use among the Mukugodo Maasai of Kenya. Pastoralism 7:18
Helmer D, Gourichon L, Vila E (2007) The development of the exploitation of products from Capra and Ovis (meat, milk and fleeces) from the PPNB to the Early Bronze in the northern Near East (8700 to 2000 BC cal.). Anthropozoologica 42:41–69
Hildebrand EA, Grillo KM, Sawchuk EA, Pfeiffer SK, Conyers LB, Goldstein ST, Hill AC, Janzen A, Klehm CE, Helper M, Kiura P, Ndiema E, Ngugi C, Shea J, Wang H (2018) A monumental cemetery built by eastern Africa’s earliest herders near Lake Turkana, Kenya. Proc Natl Acad Sci 115(36):8942–8947
Hoeven E, Fidalis MN, SGA L, Geerts S, Hanotte O, Han J (2007) Introgression of the Sahelian breed into West African dwarf goats. In: Njogu AR (ed) Proceedings of the International Scientific Council for Trypanosomiasis Research and Control Conference. African Union/IBAR, Nairobi, pp 622–626
Holtzman J (2009) Uncertain tastes: memory, ambivalence, and the politics of eating in Samburu, Northern Kenya. University of California Press, Berkeley
Horsburgh KA (2018) A reply to Plug 2017: science requires self-correction. Azania Archaeol Res Afr 53(1):114–118
Horsburgh KA, Orton J, Klein RG (2016a) Beware the springbok in sheep’s clothing: how secure are the faunal identifications upon which we build our models? Afr Archaeol Rev 33:353–361
Horsburgh KA, Moreno-Mayar JV, Gosling AL (2016b) Revisiting the Kalahari debate in the highlands: ancient DNA provides new faunal identifications at Sehonghong, Lesotho. Azania Archaeol Res Afr 51:1–12
Horsburgh KA, Moreno-Mayar JV, Klein RG (2017) Counting and miscounting sheep: genetic evidence for pervasive misclassification of wild fauna as domestic stock. South Afr Humanit 30:53–69
Jerardino A, Fort J, Isern N, Rondelli B (2014) Cultural diffusion was the main driving mechanism of the Neolithic transition in Southern Africa. PLoS One 9:e113672
Kimura B, Marshall FB, Chen S, Rosenbom S, Moehlman PD, Tuross N, Sabin RC, Peters J, Barich B, Yohannes H, Kebede F, Teclai R, Beja-Pereira A, Mulligan CJ (2011) Ancient DNA from Nubian and Somali wild ass provides insights into donkey ancestry and domestication. Proc R Soc B Biol Sci 278:50–57
King JM, Sayers AR, Peacock CP, Kontrohr E (1984) Maasai herd and flock structures in relation to livestock wealth, climate and development. Agric Syst 13:21–56
Lamprey R, Waller R (1990) The Loita–Mara Region in historical times: patterns of subsistence, settlement and ecological change. In: Robertshaw P (ed) Early Pastoralists of Southwestern Kenya. British Institute in East Africa, Nairobi
Larson G, Piperno DR, Allaby RG, Purugganan MD, Andersson L, Arroyo-Kalin M, Barton L, Climer Vigueira C, Denham T, Dobney K, Doust AN, Gepts P, Gilbert MTP, Gremillion KJ, Lucas L, Lukens L, Marshall FB, Olsen KM, Pires JC, Richerson PJ, Rubio de Casas R, Sanjur OI, Thomas MG, Fuller DQ (2014) Current perspectives and the future of domestication studies. Proc Natl Acad Sci 111:6139–6146. https://doi.org/10.1073/pnas.1323964111
Le Meillour L, Zazzo A, Cersoy S, Marie A, LeBon M, Lesur J, Pleurdeau D, Zirah S (2018) Tracing the introduction of domestic caprines to Southern Africa using palaeoproteomics. Paper presented at the 24th Biannual Meeting of the Society of Africanist Archaeologists, Toronto, June 19
Loreille O, Vigne J-D, Hardy C, Callou C, Treinen-Claustre F, Dennebouy N, Monnerot M (1997) First distinction of sheep and goat archaeological bones by the means of their fossil mtDNA. J Archaeol Sci 24:33–37
Makarewicz C, Tuross N (2006) Foddering by Mongolian pastoralists is recorded in the stable carbon (δ13C) and nitrogen (δ15N) isotopes of caprine dentinal collagen. J Archaeol Sci 33:862–870
Makarewicz CA (2014) Winter pasturing practices and variable fodder provisioning detected in nitrogen (δ15N) and carbon (δ13C) isotopes in sheep dentinal collagen. J Archaeol Sci 41:502–510
Marshall F (1986) Aspects of the advent of pastoral economies in East Africa. PhD thesis, University of California (Berkeley)
Marshall F (1990) Origins of specialized pastoral production in East Africa. Am Anthropol 92:873–890
Marshall F, Grillo KM, Arco L (2011) Prehistoric pastoralists and social responses to climatic risk in East Africa. In: Miller N, Moore K, Ryan K (eds) Sustainable lifeways: cultural persistence in an ever-changing environment. University of Pennsylvania Museum of Archaeology and Anthropology, Philadelphia, pp 38–73
Marshall F, Hildebrand E (2002) Cattle before crops: the beginnings of food production in Africa. J World Prehist 16:99–143
Marshall F, Stewart KM, Barthelme J (1984) Early domestic stock at Dongodien in Northern Kenya. Azania 19:120–127
Martin L, Edwards Y (2013) Diverse strategies: evaluating the appearance and spread of domestic caprines in the southern Levant. In: Colledge S, Conolly J, Dobney K, Manning K, Shennan S (eds) Origins and spread of domestic animals in Southwest Asia and Europe. Left Coast Press, Berkeley, pp 49–82
McCabe JT (1987) Drought and recovery: livestock dynamics among the Ngisonyoka Turkana of Kenya. Hum Ecol 15:371–389
Muigai AWT, Hanotte O (2013) The origin of African sheep: archaeological and genetic perspectives. Afr Archaeol Rev 30:39–50. https://doi.org/10.1007/s10437-013-9129-0
Mutundu K (2005) Domestic stock age profiles and herd management practices: ethnoarchaeological implications from Maasai settlements in southern Kenya. Archaeofauna 14:83–92
Newman ME, Parboosingh JS, Bridge PJ, Ceri H (2002) Identification of archaeological animal bone by PCR/DNA analysis. J Archaeol Sci 29:77–84
Österle M (2008) From cattle to goats: the transformation of East Pokot pastoralism in Kenya. Nomadic Peoples 12(1):81–91
Payne S (1969) A metrical distinction between sheep and goat metacarpals. In: Ucko PJ, Dimbleby GW (eds) The domestication and exploitation of plants and animals. Duckworth, London, pp 295–305
Payne S (1973) Kill-off patterns in sheep and goats: the mandibles from Asvan Kale. Anatol Stud 23:281–303
Payne S (1985) Morphological distinctions between the mandibular teeth of young sheep, Ovis, and goats, Capra. J Archaeol Sci 12:139–147
Pereira F, Queiros S, Gusmao L, Nijman IJ, Cuppen E, Lenstra JA, Consortium E, Davis SJM, Nejmeddine F, Amorim A (2009) Tracing the history of goat pastoralism: new clues from mitochondrial and Y chromosome DNA in North Africa. Mol Biol Evol 26:2765–2773. https://doi.org/10.1093/molbev/msp200
Pilaar Birch SE, Scheu A, Buckley M, Çakırlar C (2018) Combined osteomorphological, isotopic, aDNA, and ZooMS analyses of sheep and goat remains from Neolithic Ulucak, Turkey. Archaeol Anthropol Sci. https://doi.org/10.1007/s12520-018-0624-8
Pleurdeau D, Imalwa E, Détroit F, Lesur J, Veldman A, Bahain J-J, Marais E (2012) “Of sheep and men”: earliest direct evidence of caprine domestication in Southern Africa at Leopard Cave (Erongo, Namibia). PLoS One 7(7):e40340
Plug I (2018) Reply to Horsburgh et al. 2016: ‘revisiting the Kalahari debate in the highlands.’Azania Archaeol Res Afr 53(1):98–113
Pratt DJ, Gwynne MD (1977) Rangeland management and ecology in East Africa. Hodder & Stoughton, London
Prendergast ME (2010) Kansyore fisher-foragers and transitions to food production in East Africa: the view from Wadh Lang’o, Nyanza Province Azania Archaeol Res Afr 45:83–111
Prendergast ME (2011) Hunters and herders at the periphery: the spread of herding in eastern Africa. In: Jousse H, Lesur J (eds) People and animals in Holocene Africa: recent advances in archaeozoology. Africa Magna Verlag, Frankfurt, pp 43–58
Prendergast ME, Mabulla AZP, Grillo KM, Broderick LG, Seitsonen O, Gidna AO, Gifford-Gonzalez D (2013) Pastoral Neolithic sites on the southern Mbulu Plateau, Tanzania. Azania Archaeol Res Afr 48:498–520
Prendergast ME, Buckley M, Crowther A, Frantz L, Eager H, Lebrasseur O, Hutterer R, Hulme-Beaman A, van Neer W, Douka K, Veall MA, Quintana Morales EM, Schuenemann VJ, Reiter E, Allen R, Dimopoulos EA, Helm RM, Shipton C, Mwebi O, Denys C, Horton M, Wynne-Jones S, Fleisher J, Radimilahy C, Wright H, Searle JB, Krause J, Larson G, Boivin NL (2017) Reconstructing Asian faunal introductions to eastern Africa from multi-proxy biomolecular and archaeological datasets. PLoS One 12:e0182565. https://doi.org/10.1371/journal.pone.0182565
Prendergast ME, Grillo KM, Mabulla AZP, Wang H (2014) New dates for Kansyore and Pastoral Neolithic ceramics in the Eyasi Basin, Tanzania. J Afr Archaeol 12:89–98. https://doi.org/10.3213/2191-5784-10245
Prendergast ME, Sawchuk E (2018) Boots on the ground in Africa’s ancient DNA ‘revolution’: archaeological perspectives on ethics and best practices. Antiquity 92:1–13
Prummel W, Frisch H-J (1986) A guide for the distinction of species, sex and body side in bones of sheep and goat. J Archaeol Sci 13:567–577
Redding R (1981) Decision making in subsistence herding of sheep and goats in the Middle East. PhD thesis, University of Michigan
Rossel S (2009) The development of productive subsistence economies in the Nile Valley: Zooarchaeological analysis at El-Mahâsna and South Abydos, Upper Egypt. Harvard University
Salvagno L, Albarella U (2017) A morphometric system to distinguish sheep and goat postcranial bones. PLoS One 12:e0178543. https://doi.org/10.1371/journal.pone.0178543
Scott K, Plug I (2016) Osteomorphology and osteometry versus aDNA in taxonomic identification of fragmentary sheep and sheep/goat bones from archaeological deposits: Blydefontein Shelter, Karoo, South Africa. South Afr Humanit 28:61–79
Skoglund P, Thompson JC, Prendergast ME, Mittnik A, Sirak K, Hajdinjak M, Salie T, Rohland N, Mallick S, Peltzer A, Heinze A, Olalde I, Ferry M, Harney E, Michel M, Stewardson K, Cerezo-Roman J, Chiumia C, Crowther A, Gomani-Chindebvu E, Gidna AO, Grillo KM, Helenius IT, Hellenthal G, Helm R, Horton M, López S, Mabulla AZP, Parkington J, Shipton C, Thomas MG, Tibesasa R, Welling M, Hayes VM, Kennett DJ, Ramesar R, Meyer M, Pääbo S, Patterson N, Morris AG, Boivin N, Pinhasi R, Krause J, and Reich D (2017) Reconstructing prehistoric African population structure. Cell 171 (1):59–71.e21
Tieszen LL, Senyimba MM, Imbamba SK, Troughton JH (1979) The distribution of C3 and C4 grasses and carbon isotope discrimination along an altitudinal and moisture gradient in Kenya. Oecologia 37:337–350
van der Sluis LG, Hollund HI, Buckley M, De Louw PGB, Rjsdjik KF, Kars H (2014) Combining histology, stable isotope analysis and ZooMS collagen fingerprinting to investigate the taphonomic history and dietary behaviour of extinct giant tortoises from the Mare aux Songes deposit on Mauritius. Palaeogeogr Palaeoclimatol Palaeoecol 416:80–91
Vila E, Helmer D (2014) The expansion of sheep herding and the development of wool production in the ancient Near East: an archaeozoological and iconographical approach. In: Breniquet C, Michel C (eds) Wool economy in the ancient Near East and the Aegean. Oxbow, Oxford, pp 22–40
Wolfhagen J, Price MD (2017) A probabilistic model for distinguishing between sheep and goat postcranial remains. J Archaeol Sci Rep 12:625–631
Wright DK (2017) Accuracy vs. precision: understanding potential errors from radiocarbon dating on African landscapes. Afr Archaeol Rev 34:303–319. https://doi.org/10.1007/s10437-017-9257-z
Young HJ, Young TP (1983) Local distribution of C3 and C4 grasses in sites of overlap on Mount Kenya. Oecologia 58:373–377
Zeder MA (2006) Reconciling rates of long bone fusion and tooth eruption and wear in sheep (Ovis) and goat (Capra). In: Ruscillo D (ed) Recent advances in ageing and sexing animal bones. Oxbow, Oxford, pp 87–118
Zeder MA, Lapham HA (2010) Assessing the reliability of criteria used to identify postcranial bones in sheep, Ovis, and goats, Capra. J Archaeol Sci 37:2887–2905
Zeder MA, Pilaar SE (2010) Assessing the reliability of criteria used to identify mandibles and mandibular teeth in sheep, Ovis, and goats, Capra. J Archaeol Sci 37:225–242
Acknowledgments
Fieldwork at Luxmanda was conducted in collaboration with Drs. Audax Mabulla and Agness Gidna of the National Museum of Tanzania. Permission to excavate in 2015 was granted by the Tanzania Commission for Science and Technology (2015-119-ER-2012-50) and by the Division of Antiquities of the Ministry of Natural Resources and Tourism; the latter also gave permission to export faunal samples (License 03/2015/2016). We are grateful to our team members and University of Wisconsin–La Crosse field school students, and in particular to Gemma Zahradka for her assistance with faunal analysis. We thank Aradhna Tripati, Ben Elliot, Dyke Andreasen, and Colin Carney for the laboratory support. We gratefully acknowledge Shaw Badenhorst and two anonymous reviewers, whose comments on an earlier version of this manuscript improved the final product.
Funding
Funding for the 2015 field season and for some lab analyses was provided by a Faculty Research Grant (FRG) to KMG and by the College of Liberal Studies at the University of Wisconsin–La Crosse. Additional lab analyses were funded by a grant to MEP from the Radcliffe Institute for Advanced Study at Harvard University, and by a grant to AJ from the Cotsen Institute of Archaeology at the University of California Los Angeles. MB was supported by a Royal Society fellowship.
Author information
Authors and Affiliations
Corresponding author
Electronic supplementary material
Online Resource 1
Pastoral Neolithic sites with published faunal data including remains of caprines, as illustrated in Fig. 1. (XLSX 14 kb)
Online Resource 2
(XLSX 10 kb)
Online Resource 3
(XLSX 14 kb)
Online Resource 4
(XLSX 16 kb)
Online Resource 5
Complete dataset with morphological, ZooMS, and isotopic identifications, and comparisons among these lines of evidence. (XLSX 17 kb)
Rights and permissions
About this article
Cite this article
Prendergast, M.E., Janzen, A., Buckley, M. et al. Sorting the sheep from the goats in the Pastoral Neolithic: morphological and biomolecular approaches at Luxmanda, Tanzania. Archaeol Anthropol Sci 11, 3047–3062 (2019). https://doi.org/10.1007/s12520-018-0737-0
Received:
Accepted:
Published:
Issue Date:
DOI: https://doi.org/10.1007/s12520-018-0737-0