Malaria and the peopling of early tropical Africa

Journal of World History

Volumn 16, Issue 3, 2005, Pages 269-291

  Webb Jr , James L A ^a  

^a Health Services and Department of Biology   (United States)

Author keywords

[No Author keywords available]

Indexed keywords

ARCHAEOLOGICAL EVIDENCE; COLONIZATION; DEMOGRAPHIC HISTORY; INFECTIOUS DISEASE; MALARIA;

AFRICA;

EID: 29644439511     PISSN: 10456007     EISSN: None     Source Type: Journal    
DOI: None     Document Type: Article

References (79)

1. For a recent effort to bring the early epochs into world history, see David Christian's fine introductory chapters "Beginnings: The Era of Foragers" and "Acceleration: The Agrarian Era" in the five-volume Berkshire Encyclopedia of World History, ed. William H. McNeill (Great Barrington, Mass.: Berkshire Publishing Group, 2005), 1:1-35.
2. William H. McNeill, Plagues and Peoples (New York: Anchor Books, 1977). McNeill's broad schema has been widely accepted. In a recent authoritative article, David E. Stannard cites McNeill repeatedly in portraying the broad outlines of human disease history.
3. See Stannard, "Disease, Human Migration, and History," in The Cambridge World History of Human Disease, ed. Kenneth F. Kiple (Cambridge: Cambridge University Press, 1993), pp. 35-44.
4. These themes have been reprised to great success by Jared Diamond in his book Guns, Germs, and Steel (New York: W. W. Norton & Company, 1997).
5. For the Cambridge historian John Iliffe, for example, the core theme in African history has been the struggle of Africans to overcome their environment to achieve positive rates of population growth. John Iliffe, The Africans: History of a Continent (Cambridge: Cambridge University Press, 1995). Some historians have found Iliffe's insistence on the primacy of this theme in the history of recent centuries to be excessive.
6. James L. A. Webb Jr., Desert Frontier: Ecological and Economic Change along the Western Sahel, 1600-1850 (Madison: University of Wisconsin Press, 1995).
7. Philip D. Curtin, "'The White Man's Grave': Image and Reality, 1780-1850," Journal of British Studies 1 (1961): 94-110,
8. Death By Migration (Cambridge: Cambridge University Press, 1989);
9. Dennis G. Carlson, African Fever: A Study of British Science, Technology, and Politics in West Africa, 1787-1864 (Canton, Mass.: Watson Publishing International, 1984).
10. There are two other human malarial parasites: Plasmodium, ovate and Plasmodium, malariae. Both are far less lethal than falciparum, and they have received less attention from researchers. Other malaria-protective genetic mutations have emerged within African populations, such as thalassemias, glucose-6-phosphate dehydrogenase deficiency (also known as G6PD deficiency) and hemoglobin C (a genetic variation that is allelic with that for sickle cell but that does not impose as severe costs for individuals who are homozygous for this variation). These polymorphisms are not as protective as Duffy negativity and the sickle cell mutation discussed in this paper. The historical issues concerning the emergence of these polymorphisms have not received as much scientific attention from the molecular biological community.
11. R. Carter and K. N. Mendis, "Evolutionary and Historical Aspects of the Burden of Malaria," Clinical Microbiology Reviews 115 (2002): 564-594.
12. The deep geographical origin of vivax is contested. One view holds that vivax malaria may have been present in the New World prior to European contact in the late fifteenth and early sixteenth centuries. Although most of the evidence suggests that this was not the case, recently scholars have called for biomolecular analysis of existing skeletal remains in the Amazon to reach a definitive conclusion. See Marcia Caldas de Castro and Button H. Singer, "Was Malaria Present in the Amazon Before European Conquest? Available Evidence and Future Research Agenda," Journal of Archaeological Science 32 (2005): 337-340.
13. The two principal views are that vivax originated either in Africa or in Southeast Asia. Because of the large number of variables and the nature of the evidence, there is latitude for diverse interpretation, and it is unlikely that the issue will be definitively settled. For a recent argument that vivax originated in Southeast Asia, see Ananias A. Escalante, Omar E. Comejo, Denise E. Freeland, Amanda C. Poe, Ester Durrego, William E. Collins, and Altaf A. Lal, "A Monkey's Tale: The Origin of Plasmodium Vivax as a Human Malaria Parasite," Proceedings of the National Academy of Sciences 102, no. 6 (2005): 1980-1985.
14. In this view, vivax never was present in Africa; Duffy antigen negativity is either the result of a genetic accommodation to another, unknown agent or a random genetic variation that became fixed. On the African origins of vivax, see Richard Carter in "Speculations on the Origins of Plasmodium vivax Malaria," Trends, in Parasitology 19, no. 5 (2003): 214-219.
15. For an overview of some of the complications in unraveling the history of vivax, see Stephen M. Rich, "The Unpredictable Past of Plasmodium vivax Revealed in Its Genome," Proceedings of the National Academy of Sciences 101, no. 44 (2004): 15,547-15,548.
16. Duffy antigen negativity (FY*o) is one of three forms of mutation on the Duffy antigen receptor. Duffy A (FY*A) and Duffy B (FY*B) confer limited immunity to malaria. Neither FY*A or FY*B occur in tropical Africa. For more on Duffy mutations, see Martha T. Hamblin and Anna Di Rienzo, "Detection of the Signature of Natural Selection in Humans: Evidence from the Duffy Blood Group Locus," American Journal of Human Genetics 66 (2000): 1669-1679;
17. Martha T. Hamblin, Emma E. Thompson, and Anna Di Rienzo, "Complex Signatures of Natural Selection at the Duffy Blood Group Locus," American Journal of Human Genetics 70 (2002): 369-383. The biological process is termed "complement fixation." It involves the binding of a "complement" (a heat-sensitive, complex system in fresh human and other sera) to an antigen-antibody complex so that the complement is unavailable for subsequent reaction.
18. The issue of the evolution of virulence is complex and unresolved. For an introduction, see Alison P. Galvani, "Epidemiology Meets Evolutionary Ecology," Trends in Ecology and Evolution 18, no. 3 (2003): 132-139.
19. See also Margaret J. Mackinnon and Andrew F. Read, "Virulence in Malaria: An Evolutionary Viewpoint," Philosophical Transactions of the Royal Society of London, Series B, Biological Sciences 359, no. 1446 (2004): 965-986.
20. Human populations apparently were greatly reduced in number, perhaps to as few as five thousand females, around approximately 70,000 B.P. This "population bottleneck" coincides with a cataclysmic eruption of the Toba volcano in Sumatra, which produced a series of volcanic winters. See Stanley H. Ambrose, "Late Pleistocene Human Population Bottlenecks, Volcanic Winter, and Differentiation of Modern Humans," Journal of Human Evolution 34 (1998): 623-651;
21. Michael R. Rampino and Stanley H. Ambrose, "Volcanic Winter in the Garden of Eden: The Toba Supereruption and the Late Pleistocene Human Population Crash," in Volcanic Hazards and Disasters in Human Antiquity, ed. F. W. McCoy and G. Heiken, Geological Society of America special paper 345 (Boulder, Colo.: Geological Society of America, 2000).
22. For critical interpretations of the Toba eruption and a rejoinder, see F. J. Gathorne-Hardy and W. E. H. Harcourt-Smith, "The Super-Eruption of Toba, Did It Cause a Human Bottleneck?" Journal of Human Evolution 45 (2003): 227-230,
23. Stanley H. Ambrose, "Did the Super-Eruption of Toba Cause a Human Population Bottleneck? Reply to Gathorne-Hardy and Harcourt Smith," Journal of Human Evolution 45 (2003): 231-237.
24. M. T. Hamblin and A. Di Rienzo, the authors of a recent genetic analysis, have proposed that a "selective sweep" toward the fixation of Duffy negativity in sub-Saharan populations began between 97,200 and 6,500 years ago. See their "Detection of the Signature of Natural Selection in Humans," pp. 1669-1679.
25. For a survey of genetic responses to malarial pressure, including sickle cell (discussed later in the present paper), see Carter and Mendis, "Evolutionary and Historical Aspects", pp. 570-573.
26. "Carter and Mendis, "Evolutionary and Historical Aspects," pp. 577-578.
27. The last glacial maxima in tropical Africa have been estimated at between 22,000 and 12,000 B.P. See A. S. Brooks and P. T. Robertshaw, "The Glacial Maxima in Tropical Africa: 22,000-12,000 B.P.," in The World at 18,000 B.P., vol. 2, Low Latitudes, ed. O. Soffer and C. Gamble (London: Unwin Hyman, 1990), pp. 121-169.
28. The thesis that resistance to malaria is tied to the "agricultural revolution" in Africa is broadly accepted. See, for example, Nina L. Etkin, "The Co-Evolution of People, Plants, and Parasites: Biological and Cultural Adaptations to Malaria," Proceedings of the Nutrition Society 62 (2003): 311-317;
29. J. C. C. Hume, J. Lyons, and K. P. Day, "Human Migration, Mosquitoes and the Evolution of Plasmodium falciparum," Trends in Parasitology 19, no. 3 (2003): 144-149.
30. This argument was first advanced by F. B. Livingstone in his classic article "Anthropological Implications of Sickle Cell Gene Distribution in West Africa," American Anthropologist 60 (1958): 533-562.
31. Recently, M. Coluzzi has written in support of this thesis in "The Clay Feet of the Malaria Giant and Its African Roots: Hypotheses and Influences About the Origin, Spread, and Control of Plasmodium, falciparum," Parassitologia 41 (1999): 277-283.
32. John E. Yellen, Alison S. Brooks, Els Cornelissen, Michael J. Mehlman, and Kathlyn Stewart, "A Middle Stone Age Worked Bone Industry from Katanda, Upper Semliki Valley, Zaire," Science 268, no. 5210 (1995): 553-556.
33. For the interpretive comparison with Europe, see B. Bower, "African Finds Revise Cultural Roots," Science News 147, no. 17 (1995): 260.
34. Sally McBrearty and Alison S. Brooks, "The Revolution That Wasn't: A New Interpretation of the Origin of Modern Human Behavior," Journal of Human Evolution 39 (2000): 458.
35. Ibid., p. 532.
36. Populations that have no immediate experience with malaria can be subject to epidemic malaria. Epidemic malaria is also extremely dangerous. By definition, epidemic outbreaks are episodic; there are fewer prospects for genetic accommodation.
37. Carter and Mendis, "Evolutionary and Historical Aspects," p. 573.
38. For an ambitious, brilliant interpretation of the period 9000-1000 B.C.E., see Christopher Ehret, The Civilizations of Africa: A History to 1800 (Charlottesville: University of Virginia Press, 2002), pp. 59-158.
39. Some scholars estimate the period at five to seven million years ago for the divergence of modern falciparum malaria. A. A. Escalante and F. J. Ayala, "Phylogeny of the Malarial Genus Plasmodium, Derived from rRNA Gene Sequences," Proceedings of the National Academy of Sciences USA 91 (1994): 11,373-11,377.
40. Carter and Mendis, "Evolutionary and Historical Aspects," especially pp. 572 and 578.
41. See also Xin-zhuan Su, Jianbing Mu, and Dierdre Joy, "The 'Malaria's Eve' Hypothesis and the Debate Concerning the Origin of the Human Malaria Parasite Plasmodium falciparum," Microbes and Infection 5 (2003): 891-896;
42. Daniel L. Hard, "The Origin of Malaria: Mixed Messages from Genetic Diversity," Nature Reviews Microbiology 2 (2004): 15-22.
43. David J. Conway, "Tracing the Dawn of Plasmodium falciparum with Mitochondrial Genome Sequences," Trends in Genetics 19, no. 12 (2003): 671-674.
44. See also David J. Conway, Caterina Fanello, Jennifer M. Lloyd, Ban M. A.-S. Al-Joubori, Aftab H. Baloch, Sushela D. Somanath, Cally Roper, Ayoade M. J. Aduola, Bert Mulder, Marinete M. Povoa, Balbir Singh, and Alan W. Thomas, "Origin of Plasmodium falciparum Malaria is Traced by Mitochondrial DNA," Molecular and Biochemical Parasitology 111 (2000): 163-171.
45. "David J. Conway and Jake Baum issue a cautionary note on the problems of dating the recent emergence of modem falciparum. See their essay "In the Blood - The Remarkable Ancestry of Plasmodium, falciparum," Trends in Parasitology 18, no. 8 (2002): 351-355.
46. Hamblin and Di Rienzo, "Detection of the Signature of Natural Selection in Humans," p. 1677.
47. Carter and Mendis, "Evolutionary and Historical Aspects," p. 571.
48. Frank B. Livingstone, "Malaria and Human Polymorphisms," Annual Review of Genetics 5 (1971): 33-64.
49. For an overview of the ecological history of West Africa, see James L. A. Webb Jr., "Ecology and Culture in West Africa," in Themes in West Africa's History, ed. Emmanuel Akyeampong (Athens: Ohio University Press, 2005).
50. J. G. Vaughn and C. A. Geissler, The New Oxford Book of Food Plants (Oxford: Oxford University Press, 1999), pp. 24-25.
51. M. Adebisi Sowunmi, "The Significance of the Oil Palm (Elaeis guineensis Jacq.) in the Late Holocene Environments of West and West Central Africa: A Further Consideration," Vegetation History and Archaeobotany 8 (1999): 199-210. These conclusions are tentative, and Sowunmi calls for palynological studies of terrestrial cores and more extensive archaeological study to confirm or refute them.
52. For an interpretation that the expansion of the oil palm was due to climate change, see Jean Maley, with the collaboration of Alex Chepstow-Lusty, "Elaeis guineensis Jacq. (Oil Palm) Fluctuations in Central Africa during the Late Holocene: Climate or Human Driving Forces for This Pioneering Species," Vegetation History and Archaeobotany 10 (2001): 117-120.
53. D. G. Coursey, Yams: An Account of the Nature, Origins, Cultivation, and Utilisation of the Useful Members of the Dioscoreaceae (London: Longmans, 1967);
54. The Origins and Domestication of Yams in Africa," in Origins of African Plant Domestication, ed. J. R. Harlan, J. M. J. de Wet, and A. B. L. Stemler (Hague: Mouton, 1975), pp. 383-408.
55. Edmond Dounias, "The Management of Wild Yam Tubers by the Baka Pygmies in Southern Cameroon," African Study Monographs, suppl. 26 (March 2001): 135-156. Dounias defines the term "paracultivation" as "a combination of technical patterns and social rules which structure the expliotation of wild plants. This term characterizes a particular process of wildd plant harvesting which aims at encouraging plant reproduction, so that the plant can be repeatedly exploited. Furthermore, the plant is voluntarily kept within its original environment, in order to better respond to the seasonal mobility of forest dwellers. The maintenance of plants in the forest is the key difference between paracultivation and protocultivation" (p. 137).
56. Christopher Ehret, "Historical/Linguistic Evidence for Early African Food Production," in From Hunters to Farmer, ed. J.D. Clark and S. A. Brandt (Berkeley: University of California Press, 1984), pp. 26-39;
57. Civilizations of Africa, pp. 82-83.
58. The Middle Niger Valley is an important agricultural region in whch some dryland crops (notably pennisetum) and the wet crop "red rice" (Orzya glabberima) were domesticated. For an excellent overview, see Roderick McIntosh, The Peolples of the Middle Niger (Malden, Mass.: Blackwell Publishing, 1998).
59. Alhough it is not possible to specify exactly when the sickle cell mutation first developed or became common, specialists are generally agreed that the falciparum malarial parasites developed from within the tropical African woodland and rainforest environments and later spread into other biomes. The earliest human physical evidence of sickle cell anemia comes from the mummified remains of Egyptians from the fourth millennium B.C.E. (A. Marin, N. Cerutti, and E. Rabino Massa, "Use of the Amplification Refractory Mutation System [ARMS] in the Study of HBS in Predynastic Egyptian Remains," Bollettino della Società Italiana di Biologia Sperimentale 75, nos. 5-6 [1999]: 27-30.)
60. This evidence suggests that falciparum may have spread from the upper Nile region to the lower Nile region. The issue of whether or not the sickle cell mutation arose separately in the lower Nile valley remains an open question. Today, the incidence of sickle cell along the entire length of the Nile is very low (0.0-2.5 percent). Stuart Edelstein, The Sickled Cell: From Myths to Molecules (Cambridge, Mass.: Harvard University Press, 1986), fig. 7.3, p. 148.
61. See, for example, M. E. Kropp Dakubu, "The Peopling of Southern Ghana: A Linguistic Viewpoint," in The Archaeological and Linguistic Reconstruction of African History, ed. Christopher Ehret and Merrick Posnansky (Berkeley: University of California Press, 1982), pp. 245-255;
62. and M. E. Kropp Dakubu, "Linguistics and History in West Africa," in Akyeampong, Themes in West Africa's History.
63. On the expansion of yam farming in early West Africa, see Bassey W. Andah, "Identifying Early Farming Traditions of West Africa," in The Archaeology of Africa: Food, Metals and Towns, ed. Thurston Shaw, Paul Sinclair, Bassey Andah, and Alex Okpoko (New York: Routledge, 1995), pp. 240-254. The expansion of rice farming into the western regions of the West African rainforests is thought to have taken place in the first millennium C.E.
64. Kairn Klieman, "The Pygmies Were Our Compass" Bantu and Batwa in the History of West Central Africa, Early Times to c. 1900 C.E. (Portsmouth, N.H.: Heinemann, 2003).
65. E. De Langhe, R. Swennen, and D. Vuylsteke, "Plantain in the Early Bantu World," Azania 29-30 (1996): 318-323;
66. Christophe Mindzie Mbida, "Evidence for Banana Cultivation and Animal Husbandry during the First Millennium BC in the Forest of Southern Cameroon," Journal of Archaeological Science 27 (2000): 151-162;
67. Christophe Mindzie Mbida, "First Archaeological Evidence of Banana Cultivation in Central Africa During the Third Millennium Before Present," Vegetation History and Archaeobotany 10 (2001): 1-6.
68. De Langhe, Swennen, and Vuylsteke, "Plantain," p. 158,
69. citing Jan Vansina, Paths in the Rainforests (Madison: University of Wisconsin Press, 1999).
70. As Klieman notes, "Bantu populations grew in number, settled into larger more sedentary villages, and began to produce larger quantities and more diverse styles of ceramics. Iron tools and banana cultivation also allowed Bantu villagers to move into forested regions away from the original riverine routes of settlement. This phenomenon resulted in the formation of numerous new speech communities, especially during the Late Stone to Metal Age (1500-500 BCE). As was the case in other parts of Africa, the introduction of iron engendered a greater centralization of local economies and an increase in economic specialization" ("Pygmies Were Our Compass," pp. 123-1124).
71. Jan Vansina, "New Linguistic Evidence and the 'Bantu Expansion,'" Journal of African History 36, no. 2 (1995): 173-195;
72. Christopher Ehret, "Bantu Expansions: Re-Envisioning a Central Problem of Early African History," International Journal of African Historical Studies 34, no. 1 (2001): 5-41;
73. Roland Oliver, Thomas Spear, Kairn Klieman, Jan Vansina, Scott' MacEachern, David Schoenbrun, James Denbow, Yvonne Bastin, H. M. Batibo, and Bernd Heine in "Comments on Christopher Ehret, 'Bantu-History: Re-Envisioning the Evidence of Language,'" International Journal of African Historical Studies 34, no. 1 (2001): 43-87.
74. Jared Diamond has recently repopularized this notion of a Bantu military-industrial complex (Guns, Germs, and Steel, pp. 394-396).
75. McNeill, Pagues and peoples, pp. 69-131.
76. This third mutation was first identified in Senegal. Individuals with the Senegalese pattern of sickle cell anemia produce higher levels of hemoglobin F, and this anemia may therefore be less severe. See Edelstein, Sickled Cell, pp. 148-149.
77. Experts are not agreed on whether or not sickle cell hemoglobin mutations have a common ancestor. For the argument for a Middle Eastern origin of hemoglobin S and the diffusion of a single mutant, see F. B. Livingstone, "Who Gave Whom Hemoglobin S: The Use of Restriction Site Haplotype Variation for the Interpretation of the Evolution of the β-s-Globin Gene," American Journal of Human Biology 1 (1989): 289-302.
78. Edelstein, Sickled Cell, pp. 55-56, 147-148. In the 1980s, it was thought that the two Bantu expansions had taken place in the second and first millennia B.C.E., respectively.
79. For the argument that population pressure played a major role in prehistory, see Mark Nathan Cohen, The Food Crisis in Prehistory: Overpopulation and the Origins of Agriculture (New Haven, Conn.: Yale University Press, 1977).