Genetic residues of ancient migrations: An end to biological essentialism and the reification of race

University of Pittsburgh Law Review

Volumn 68, Issue 2, 2006, Pages 387-447

  Richman, William M ^a  

^a UNIVERSITY OF TOLEDO   (United States)

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[No Author keywords available]

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EID: 36849003317     PISSN: 00419915     EISSN: None     Source Type: Journal    
DOI: 10.5195/LAWREVIEW.2006.87     Document Type: Review

References (295)

1. Quoted in LEE GREEN, SPORTSWIT 270 (1986).
2. Belloc's poem is an irresistible target for anthropologists and is quoted in WILLIAM W. HOWELLS, MANKIND IN THE MAKING 275 (1959). It may be unfair to mulct Belloc with this racist sentiment. The poem was intended as an ironic characterization of the British upper class. Belloc himself may or may not have been a bigot. He opposed philosophical anti-Semitism, but may have entertained anti-Semitic prejudices personally.
3. Some readers might suspect that the position that I have labeled "essentialism" is a caricature, a straw man. But see STEPHEN J. GOULD, THE MISMEASURE OF MAN 73-112 (1981) [hereinafter GOULD, MISMEASURE] (devoting 40 pages to exposing a cavalcade of scientific racists, among them the foremost scientists of their ages). Robert M. Yerkes, a personal favorite and the author (with others) of the Army LQ. test, evokes the verse by Belloc with his estimates of the national I.Q.s of different countries. See id. at 197-99. The most recent examples are still the subject of current controversy.
4. See RICHARD J. HERRNSTEIN & CHARLES MURRAY, THE BELL CURVE: INTELLIGENCE AND CLASS STRUCTURE IN AMERICAN LIFE (1994);
5. ARTHUR R. JENSEN, THE G FACTOR: THE SCIENCE OF MENTAL ABILITY (1998);
6. MICHAEL LEVIN, WHY RACE MATTERS (1997);
7. J. PHILIPPE RUSHTON, RACE, EVOLUTION, AND BEHAVIOR (1995). Rushton's particular brand depends on the r/K dichotomy and what he calls the Life-History theory. Organisms that are r-selected devote more energy to producing multiple offspring while those that are K-selected produce fewer and put more parental care and energy into each one. RUSHTON at 200-16. The result is a ranking of races with Asians at the top in intelligence and at the bottom in athletic ability and genital size, Africans in the opposite position and Europeans in the middle on all three scales. The supposed reason for these differences is that Asians and Europeans moved out of Africa and had to contend with more challenging environments and thus had to grow smarter. Id. at 217-33. You can't make this stuff up!
8. For a summary of some older ideas of scientific racism, see STEVE OLSON, MAPPING HUMAN HISTORY 178-83 (2002).
9. There are particular problems, legal and social, where the biology of race matters: the rights to ancestral remains, identification of the dead and living (forensic anthropology), and adoption. In adoption, for example, for good reasons or not, prospective parents may wish to choose to adopt a child based on particular phenotypes or placing agencies might use race to determine suitability of prospective parents. On the conflict over the reality of race among forensic anthropologists, see Diana Smay & George Armelagos, Galileo Wept: A Critical Assessment of the Use of Race in Forensic Anthropology, TRANSFORMING ANTHROPOLOGY, 2000, No. 2, at 19, 20.
10. See JEROME FRANK, LAW AND THE MODERN MIND 31 (3d prtg. 1935).
11. See, e.g., Old Chief v. United States, 519 U.S. 172 (1997).
12. See MARK RIDLEY, THE COOPERATIVE GENE 45 (2001) [hereinafter RIDLEY, COOPERATIVE GENE]
13. (discussing ARTHUR O. LOVEJOY, THE GREAT CHAIN OF BEING (1936)). The idea owed much to Platonic essentialism and the Aristotelian notion of "telos" or final cause" ("that for the sake of which"), which followed a model of life on earth as a progression toward a final and most perfect state.
14. See also Donald Braman, Of Race and Immutability, 46 UCLA L. REV. 1375 (1999);
15. Steven M. Wise, The Legal Thinghood of Nonhuman Animals, 23 B.C. ENVTL. AFF. L. REV. 471 (1996).
16. On the fallacy of essentialism in biological taxonomy, see STEPHEN JAY GOULD, FULL HOUSE 38-42 (1996).
17. Braman, supra note 7, at 1386 n.25.
18. Id.
19. Sarah A. Tishkoff & Kenneth Kidd, Implications of Biogeography of Human Populations for 'Race' and Medicine, NATURE GENETICS, Nov. 2004, at S21, S21.
20. CARLETON S. COON, THE ORIGINS OF RACES 3 (1962) [hereinafter COON, ORIGINS]. Earlier, he had proposed a system of six races, with the Pacific race added to the five he settled on in 1962.
21. CARLETON S. COON, THE HISTORY OF MAN 190-95 (1955) [hereinafter COON, HISTORY]. The 2000 Census used a similar classification. "In October 1997, the Office of Management and Budget (OMB) announced the revised standards for federal data on race and ethnicity. The minimum categories for race are now: American Indian or Alaska Native; Asian; Black or African American; Native Hawaiian or Other Pacific Islander; and White."
22. U.S. Census Bureau, Racial and Ethnic Classifications Used in Census 2000 and Beyond, http://www.census.gov/population/www/socdemo/race/racefactcb.html (last visited Oct. 28, 2006).
23. For a discussion of the history of racial categories in the United States, see Eric Lillquist & Charles A. Sullivan, The Law and Genetics of Racial Profiling in Medicine, 39 HARV. C.R.-C.L. L. REV. 391, 402-08 (2004).
24. COON, HISTORY, supra note 11, at 190.
25. On the identification of races as Platonic types, see Rick A. Kittles & Kenneth M. Weiss, Race, Ancestry and Genes: Implications for Defining Disease Risk, ANN. REV. GENOMICS HUM. GENETICS (2003), at 33, 35.
26. On essentialism generally in biology, see ERNST MAYR, WHAT EVOLUTION Is 74-75 (2001) [hereinafter MAYR, EVOLUTION IS];
27. ERNST MAYR, THIS IS BIOLOGY xii (1997) [hereinafter MAYR, BIOLOGY].
28. This brief exposition is, of course, a gross simplification of a great diversity of thought on the concept of race in America and Europe. For a rich treatment, see Braman, supra note 7, at 1384-92.
29. That folk wisdom is still alive and well regardless of what science has to say about its validity, see Pilar Ossorio & Troy Duster, Race and Genetics, AM. PSYCHOLOGIST, Jan. 2005, at 115, 116 (2005);
30. George J. Armelagos, Race, Reason and Rationale, 4 EVOLUTIONARY ANTHROPOLOGY 103 (1995) (reviewing four books on race, three of which advocate neo-racialist positions) [hereinafter Armelagos, Race].
31. Smay & Armalagos, supra note 4, at 20; Tishkoff & Kidd, supra note 10, at S21.
32. See HERRNSTEIN & MURRAY, supra note 3; JENSEN, supra note 3; LEVIN, supra, note 3; RUSHTON, supra note 3; David C. Rowe, Under the Skin: On the Impartial Treatment of Genetic and Environmental Hypotheses of Racial Differences, AM. PSYCHOLOGIST, Jan. 2005, at 60, 60-70 (2005).
33. Neil Risch, Esteban Burchard, Elad Ziv & Hua Tang, Categorization of Humans in Biomedical Research: Genes, Race, and Disease, GENOME BIOLOGY, July 1, 2002, at 2007.1, 2007.4, http://genomebiology.eom/ 2002/3/7/comment/2007.
34. See generally RIDLEY, COOPERATIVE GENE, supra note 7;
35. MARK RIDLEY, EVOLUTION (2d ed. 1996) [hereinafter RIDLEY, EVOLUTION].
36. RIDLEY, EVOLUTION, supra note 19, at 23 fig.2.2.
37. Id. at 80-81. They come in many forms with a bewildering array of names: single nucleotide polymorphisms, insertions, deletions, short tandem repeats, micro-satellite polymorphisms, etc. Each of these terms represents a particular way in which two genomes can differ; for our purpose it is sufficient to refer to them collectively as polymorphisms or markers (when used to trace migrations). See Sarah A. Tishkoff & Brian C. Verrelli, Patterns of Human Genetic Diversity, ANN. REV. GENOMICS HUM. GENETICS, 2003, at 293, 297-301.
38. MAYR, BIOLOGY, supra note 13, at 307.
39. Lillquist & Sullivan, supra note 11, at 410-11; RIDLEY, EVOLUTION, supra note 19, at 30-31.
40. MAYR, EVOLUTION Is, supra note 13, at 93, 290.
41. See RIDLEY, COOPERATIVE GENE, supra note 7, at 80.
42. See OLSON, supra note 3, at 35-37; RIDLEY, EVOLUTION, supra note 19, at 73-74.
43. LUIGI LUCA CAVALLI-SFORZA & FRANCESCO CAVALLI-SFORZA, GREAT HUMAN DIASPORAS 88-92 ( 1995) [hereinafter GREAT HUMAN DIASPORAS].
44. See RIDLEY, EVOLUTION, supra note 19, at 73-74.
45. For an explanation of natural selection, see RIDLEY, EVOLUTION, supra note 19, at 64-66; for sexual selection, see id. at 296-307.
46. See, e.g., id. at 69.
47. RIDLEY, EVOLUTION, supra note 19, at 127.
48. A classic example is the evolution of the peppered moth in England during the Industrial Revolution. Before industrial pollution, the moth had a peppered coloration (black and white) camouflaging it against the predominant background colors of the branches of the trees in its habitat. As industrial pollutants turned tree branches black, moths bearing melanistic mutations (which formerly had been disadvantageous) now enjoyed a selective advantage and began to increase in frequency. Id. at 64. Nearly as well known is the human sickle-cell mutation, which causes a change in the shape of red blood cells, a defect that is usually fatal. Thus, a child that is homozygous for the sickle-cell allele likely will not survive long enough to reproduce. All things being equal then the sickle cell allele should die out in a very few generations. It survives because an individual that is heterozygous for the gene (one normal-cell allele and one sickle-cell allele) does not develop the fetal flaw; moreover, it will enjoy a selective advantage (greater resistance to infection by certain strains of malaria) over those who are homozygous for the normal-cell allele. The frequency of the sickle cell allele in a population becomes a balance of the losses of homozygotes due to sickle-cell anemia versus the gains of heterozygotes from their increased malaria resistance. Thus the frequency of the allele in the population will become fixed at a percentage that is optimal based on that population's exposure to the risk of malaria. Id. at 110-12.
49. Besides mutation, drift and natural and sexual selection, there is one other obvious process that affects the genetics of a particular population - gene flow from another population. CHRISTOPHER STRINGER & ROBIN MCKIE, AFRICAN EXODUS 70 (1997) [hereinafter STRINGER & MCKIE, AFRICAN EXODUS].
50. See MERRITT RUHLEN, THE ORIGIN OF LANGUAGE (1994). Ruhlen's reconstruction of ancient migrations via linguistic evidence is largely similar to the reconstruction by the geneticists. See id. at 190-94.
51. See also LUIGI LUCA CAVALLI- SFORZA, GENES, PEOPLES, AND LANGUAGES 155-65 (2000) [hereinafter GENES, PEOPLES, AND LANGUAGES];
52. STRINGER & MCKIE, AFRICAN EXODUS, supra note 32, at 141.
53. On the history of using genetic markers to trace migrations, see L. Luca Cavalli-Sforza & Marcus W. Feldman, The Application of Molecular Genetic Approaches to the Study of Human Evolution, NATURE GENETICS, Mar. 2003, at 263.
54. See OLSON, supra note 3, at 37.
55. In addition to population, geneticists also have studied the genomes of some of our constant companions - rats, fleas, and Helicobacter pylori (bacterium responsible for stomach ulcers) - to fill in migrational details. See John Pickrell, Rat DNA Offers Clues to Pacific Colonization, Study Says, NATIONAL GEOGRAPHIC NEWS, June 9, 2004, http://news.nationalgeographic.com/news/2004/06/ 0609_040609_ratdna.html;
56. Noah Kerness Whiteman & Patricia G. Parker, Using Parasites to Infer Host Population History: A New Rationale for Parasite Conservation, 8 ANIMAL CONSERVATION 175 (2005). See also Risch et al., supra note 18, at 2007.5, for a listing of the various types of genetic markers studied.
57. See GENES, PEOPLES, AND LANGUAGES, supra note 33, at 17-19.
58. STRINGER & MCKIE, AFRICAN EXODUS, supra note 32, at 133. Coding DNA is DNA that contains instructions for the production of proteins that affect the organism's development and ultimate morphology and physiology. Non-coding DNA is DNA whose function and effect we do not (yet) know; it has no currently apparent effect on the organism's development. It had been termed "junk" DNA, but more recent research suggests that it does have significant functions. See Noncoding DNA, http://en.wikipedia.org/wiki/Noncoding_DNA (last visited Oct. 28, 2006).
59. SPENCER WELLS, THE JOURNEY OF MAN: A GENETIC ODYSSEY 21 (2002) [hereinafter WELLS, JOURNEY].
60. Id. at 70-71.
61. Id. at 21. The widow's peak example is actually an instance of coding DNA that codes for a selectively neutral feature; actual non-coding DNA has no known function or effect on the organism's development. The same principle of drift, rather than selection, controls the spread of non-coding and selectively neutral coding DNA.
62. See WELLS, JOURNEY, supra note 39, at 29;
63. L. LUCA CAVALLI-SFORZA, PAOLO MENOZZI & ALBERTO PIAZZA, THE HISTORY AND GEOGRAPHY OF HUMAN GENES 83 (1994).
64. GREAT HUMAN DIASPORAS, supra note 27, at 77;
65. WELLS, JOURNEY, supra note 39, at 29.
66. WELLS, JOURNEY, supra note 39, at 29. The reason it does not recombine is that it is a single circular structure, not a double helix. The reason it is so uniquely traceable is that it passes through the female line only. See the following note and its accompanying text.
67. Id. at 30. She was not the only woman alive at the time, just the only one lucky enough to have female offspring that continued to reproduce in an uninterrupted chain to the present.
68. On recombination, see MAYR, EVOLUTION IS, supra note 13, at 103.
69. GREAT HUMAN DIASPORAS, supra note 27, at 84.
70. WELLS, JOURNEY, supra note 39, at 42-43.
71. For an explanation of how the Y-chromosome behaves during reproduction see id.
72. The reason for the difference in coalescence times is the politics of human mating. Nearly every female can mate and has an upward limit on the number of mating opportunities that is a function of physiology. Not so for males; some do not get the opportunity to mate at all, but others mate with any number of females and produce huge numbers of offspring. An extreme example is Genghis Khan whose Y-chromosome appears in about .5 percent of the world total. Tatiana Zerjal et al., The Genetic Legacy of the Mongols, 72 AM. J. HUM. GENETICS 717, 717 (2003).
73. On the use of the Y-chromosome to reconstruct human migrations, see Jaume Bertranpetit, Genome, Diversity, and Origins: The Y Chromosome as a Storyteller, 97 PROCEEDINGS NAT'L ACAD. SCI. U.S. 6927 (2000).
74. WELLS, JOURNEY, supra note 39, at 55.
75. On the genetic proximity of humans and African apes see also STRINGER & MCKEE, AFRICAN EXODUS, supra note 32, at 20.
76. National Center for Biotechnology Information, http://www.ncbi.nlm.nih. gov/ (last visited Oct. 29, 2006).
77. JARED DIAMOND, THE THIRD CHIMPANZEE 34-39 (1992) [hereinafter DIAMOND, THIRD CHIMPANZEE]; OLSON, supra note 3, at 19.
78. Id. at 33-34.
79. GREAT HUMAN DIASPORAS, supra note 27, at 42-45;
80. WILLIAM HOWELLS, GETTING HERE 102 (1997).
81. DIAMOND, THIRD CHIMPANZEE, supra note 54, at 37.
82. Id.
83. See STRINGER & MCKIE, AFRICAN EXODUS, supra note 32, at 47. For more on Neanderthals, see HOWELLS, supra note 56, at 141-55.
84. See Peter Forster, Ice Ages and the Mitochondrial DNA Chronology of Human Dispersals: A Review, 359 PHILOSOPHICAL TRANSACTIONS ROYAL SOC'Y LONDON 255, 257 (2004); Tishkoff & Verrelli, supra note 21, at 307. The term "Paleolithic" means literally "old stone age," and refers to a culture which was the only human lifestyle for about 150,000 years. There was no metallurgy, and most tools were fashioned from stone initially and later from animal bone or sinew; food was obtained by foraging. "Neolithic" refers to a more developed culture that also lacked metallurgy, but incorporated agriculture and herding. Tools were of natural materials but became more sophisticated and specialized.
85. Lev A. Zhivotovsky et al., Features ofEvolution and Expansion of Modern Human Populations from Genomewide Microsatellite Markers, 72 AM. J. HUM. GENETICS 1171, 1182 (2003). Before this population split, the species probably underwent a significant expansion event, starting with a population of about 6,600.
86. See David E. Reich & David B. Goldstein, Genetic Evidence for a Paleolithic Human Population Expansion in Africa, 95 PROC. NAT'L ACAD. SCI. U.S. 8119, 8123 (1998).
87. Pygmies no longer have their own language but speak the languages of the surrounding farmers. GREAT HUMAN DIASPORAS, supra note 27, at 9.
88. Tishkoff & Verrelli, supra note 21, at 309. For more on Africa, see generally Reich & Goldstein, supra note 61; Antonio Salas et al., The Making of the African mtDNA Landscape, 71 AM. SOC. HUM. GENETICS 1082 (2002). This view of human dispersal, now widely accepted is known as Recent African Origin (RAO); the opposite view, multi-regionalism posits an early migration of Erectus out of Africa and parallel evolution of modern humans on several different continents. The difference between the two theories is highly relevant to the subject of this paper as the RAO view postulates racial differences that are only 100,000 years old and thus relatively superficial. On the two theories,
89. see GENES, PEOPLES, AND LANGUAGES, supra note 33. The MR view depends upon a million-year-old separation, which would allow for fundamental racial differences.
90. See STRINGER & MCKIE, AFRICAN EXODUS, supra note 32, at 59-61. Some evidence for the MR view exists in the shared anatomic features of archaic and modern humans in particular geographic regions, e.g., shovel-shaped incisors among archaic and modern Asians. The principal difficulties with MR are (1) the genetic evidence suggesting RAO, (2) the improbability of nearly exact convergent evolution of archaics into moderns in several different, widely separated locales, (3) the improbability of significant genetic exchange among small, widely separated groups of archaics (the only alternative to coincidental convergent evolution), and (4) the high degree of genetic difference among African populations compared to non-African ones.
91. See Recent Single-Origin Hypothesis, http://en.wikipedia.org/wiki/ Recent_single-origin_hypothesis (last visited Dec. 1, 2006). The RAO model is clearly dominant. Milford Wolpoff is the most well-known proponent of multiregionalism.
92. See MILFORD WOLPOFF & RACHEL CASPARI, RACE AND HUMAN EVOLUTION: A FATAL ATTRACTION (1997).
93. Tishkoff & Verrelli, supra note 21, at 309.
94. There is, however, some evidence for migration along both routes, from the Horn and through the Levant. See J.R. Luis et al., The Levant Versus the Horn of Africa: Evidence for Bidirectional Corridors of Human Migrations, 74 AM. J. HUM. GENETICS 532 (2004).
95. Modern humans had moved out of Africa and into the Middle East earlier, but did not stay long. Cyclic warming and cooling of the area made it sometimes more hospitable to modern humans, and other times more habitable by cold-adapted Neanderthals. WELLS, JOURNEY, supra note 39, at 98-99.
96. See id. at 68.
97. Id.
98. About ten miles distance separates the Horn of Africa from Yemen and there are close genetic links between the populations of those areas, suggesting two-way migration over many years. The narrow inlet is called the Bab-el-Mandeb or "Gate of Tears." See Toomas Kivisild et al., Ethiopian Mitochondrial DNA Heritage: Tracking Gene Flow Across and Around the Gate of Tears, 75 AM. J. HUM. GENETICS 752, 752 (2004). This is known as the Out of Africa model, as contrasted with Multiregionalism.
99. See J. Lynn B. Jorde & Stephen P. Wooding, Genetic Variation, Classification and 'Race ', NATURE GENETICS, Nov. 2004, at 28.
100. WELLS, JOURNEY, supra note 39, at 72.
101. See OLSON, supra note 3, at 129.
102. Max Ingman & Ulf Gyllensten, Mitochondrial Genome Variation and Evolutionary History of Australian and New Guinean Aborigines, 13 GENOME RESEARCH 1600 (2003).
103. WELLS, JOURNEY, supra note 39, at 72-74.
104. Id. at 78.
105. Id.
106. Id. at 74.
107. Id. at 113; Peter A. Underhill, A Synopsis of Extant Y Chromosome Diversity in East Asia and Oceania, in THE PEOPLING OF EAST ASIA: PUTTING TOGETHER ARCHAEOLOGY, LINGUISTICS AND GENETICS 313 (2004) [hereinafter Underhill, Synopsis];
108. P.A. Underhill, Inferring Human History: Clues from Y-Chromosome Haplotypes, 68 COLD SPRING HARBOR SYMPOSIA ON QUANTITATIVE BIOLOGY 487, 489 fig.1 (2003) [hereinafter Underhill, Inferring].
109. See Kumarasamy Thangaraj et al., Genetic Affinities of the Andaman Islanders, a Vanishing Human Population, 13 CURRENT BIOLOGY 86, 86 (2003).
110. For more on Australia, see Underhill, Synopsis, supra note 77, and Ingman & Gyllensten, supra note 72.
111. WELLS, JOURNEY, supra note 39, at 112-13.
112. Id. at 108.
113. Id.
114. See id.
115. Id.
116. See id. at 182-83 fig.10.
117. Id. at 110-11.
118. See id. at 111.
119. Id. at 111.
120. Id. 112.
121. Id. at 112-13. For more on the genetic landscape of India, see T. Kivisild et al., The Genetic Heritage of the Earliest Settlers Persists Both in Indian Tribal and Caste Populations, 72 AM. J. HUM. GENETICS 313 (2003);
122. Malliya gounder Palanichamy et al., Phytogeny of Mitochondrial DNA Macrohaplogroup N in India, Based on Complete Sequencing: Implications for the Peopling of South Asia, 75 AM. J. HUM. GENETICS 966 (2004).
123. WELLS, JOURNEY, supra note 39, at 113.
124. Id. There are many examples of this asymmetrical mating pattern, including replacements of hunter/gatherer populations by Neolithic farmers, replacement of Native Americans by European colonizers and replacement of Australian Aborigines by Europeans. It may be function as a genocide marker. Not every asymmetry signals a genocide or even a conquest, however. When the asymmetry is weaker, a possibility is a dual-gender migration rather than a male-only conquest. See Bo Wen et al., Analyses of Genetic Structure of Tibeto-Burman Populations Reveals Sex-Biased Admixture in Southern Tibeto-Burmans, 74 AM. J. HUM. GENETICS 856 (2004).
125. WELLS, JOURNEY, supra note 39, at 182 fig.10.
126. Id. at 112.
127. Id. at 118.
128. See WELLS, JOURNEY, supra note 39, at 118-19. For more on the genetic landscape of East Asia, see Bo Wen et al., supra note 91;
129. Yong-Gang Yao et al., Phylogeographic Differentiation of Mitochondrial DNA in Han Chinese, 70 AM. J. HUM. GENETICS 635 (2002);
130. Yuan-Chun Ding et al., Population Structure and History in East Asia, 97 PROC. NAT'L ACAD. SCI. U.S. 14003 (2000).
131. See WELLS, JOURNEY, supra note 39, at 120-21.
132. Id. at 156-57; for more on the Neolithic expansions, see infra Section II.B.3.
133. See R. Spencer Wells et al., The Eurasian Heartland: A Continental Perspective on Y-Chromosome Diversity, 98 PROC. NAT'L ACAD. SCI. U.S. 10244, 10247-48 (2001) (fixing M45 as the source marker for the European and Native American populations) [hereinafter Heartland].
134. Mark Seielstad et al., A Novel Y-Chromosome Variant Puts an Upper Limit on the Timing of First Entry into the Americas, 73 AM. J. HUM. GENETICS 700 (2003).
135. See Peter Beerli & Scott V. Edwards, When Did Neanderthals and Modern Humans Diverge?, 2002 EVOLUTIONARY ANTHROPOLOGY SUPPLEMENT 60.
136. The best estimate is that the ancestors of modern humans and Neanderthals diverged genetically about 700,000 years ago. Id.
137. Forster, supra note 60, at 260.
138. See STRINGER & MCKIE, AFRICAN EXODUS, supra note 32, at 109-11.
139. See id. at 53.
140. Forster, supra note 60, at 260. Moreover, neither the Neanderthals nor any other population of archaic Homo sapiens has descendants among current human populations. Although they survived successfully for hundreds of thousands of years, they form an evolutionary dead end. For more on the making of the genetic landscape of Europe, see Lluis Quintana-Murci et al., Where West Meets East: The Complex mtDNA Landscape of the Southwest and Central Asian Corridor, 74 AM. J. HUM. GENETICS 827 (2004);
141. S. Plaza et al., Joining the Pillars of Hercules: mtDNA Sequences Show Multidirectional Gene Flow in the Western Mediterranean, 67 ANNALS HUM. GENETICS 312 (2003);
142. Martin Richards et al., In Search of Geographical Patterns in European Mitochondrial DNA, 71 AM. J. HUM. GENETICS 1168 (2002);
143. Siiri Rootsi et al., Phylogeography of Y-Chromosome Haplogroup I Reveals Distinct Domains of Prehistoric Gene Flow in Europe, 75 AM. J. HUM. GENETICS 128 (2004);
144. Kristiina Tambets et al., The Western and Eastern Roots of the Saami - the Story of Genetic "Outliers" Told by Mitochondrial DNA and Y Chromosomes, 74 AM. J. HUM. GENETICS. 661 (2004); Heartland, supra note 98.
145. The isolation of the marker is described in Seielstad, supra note 99.
146. There are some reports of M3 in extreme Northeast Siberia, suggesting a back migration from Beringia. See Maria-Catira Bortolini et al., Y-Chromosome Evidence for Different Demographic Histories in the Americas, 73 AM. J. HUM. GENETICS. 524 (2003).
147. WELLS, JOURNEY, supra note 39, at 136.
148. See generally J.M. ADOVASIO WITH JAKE PAGE, THE FIRST AMERICANS (2002);
149. THOMAS D. DILLEHAY, THE SETTLEMENT OF THE AMERICAS (2000).
150. Mark Sielstad et al., Letter to the Editor, A Novel Y-Chromosome Variant Puts an Upper Limit on the Timing of First Entry into the Americas, 73 AM. J. HUM. GENETICS 700 (2003) (fixing the date of entry by finding marker M242 between M45 and M3 on the Y-chromosome).
151. RUHLEN, supra note 33 at 169-70.
152. Jeffrey T. Lell et al., The Dual Origin and Siberian Affinities of Native American Y Chromosomes, 70 AM. J. HUM. GENETICS 192 (2002) (finding two waves of migration, one from South Central Asia and one from coastal East Asia);
153. Ripan S. Mahli et al., The Structure of Diversity within New World Mitochondrial DNA Haplogroups: Implications for the Prehistory of North America, 70 AM. J. HUM. GENETICS 905 (2002) (finding the population came in a single wave from a single source in the Altai Mountains of South Central Siberia);
154. Connie J. Mulligan et al., Population Genetics, History, and Health Patterns in Native Americans, 5 ANN. REV. GENOMIC & HUM. GENETICS 295 (2004) (finding American population results from a single migration);
155. Stephen L. Zegura et al., High-Resolution SNPs and Microsatellite Haplotypes Point to a Single, Recent Entry of Native American Y Chromosomes into the Americas, 21 MOLECULAR BIOLOGY & EVOLUTION 164 (2004) (finding one migration event about 17,000 years ago with a remote source in the Altai Mountains).
156. See Sandro Bonatto & Francisco M. Salzano, A Single and Early Migration for the Peopling of the Americas Supported by Mitochondrial DNA Sequence Data, 94 PROC. NAT'L ACAD. SCI. U.S. 1866 (1997) (originating from the Beringia model).
157. H.J. Bandelt et al., Identification of Native American Founder mtDNAs Through the Analysis of Complete mtDNA Sequences: Some Caveats, 67 ANNALS HUM. GENETICS 512 (2003) (finding support for a staggered series of migrations from Siberia to Beringia to the Americas).
158. See, e.g., Mulligan et al., supra note 111.
159. WELLS, JOURNEY, supra note 39, at 142.
160. For more on the population of the Americas, see Bonatto & Salzano, supra note 112; Keith Hunley & Jeffrey C. Long, Gene Flow across Linguistic Boundaries in Native North American Populations, 102 PROC.NAT'L. ACAD. SCI. U.S. 1312 (2005);
161. Elena B. Starikovskaya et al., Mitochondrial DNA Diversity in Indigenous Populations of the Southern Extent of Siberia, and the Origins of Native American Haplogroups, 2005 ANNALS HUM. GENETICS 67, 85-86 (comparing Amerind, Na Dene, Eskimo, and Chukchi populations, all of which were inhabitants of Beringia).
162. See sources cited in notes 120-59, infra.
163. See generally George J. Armelagos & Kristen N. Harper, Genomics at the Origins of Agriculture, 14 EVOLUTIONARY ANTHROPOLOGY 68 (2005).
164. JARED DIAMOND, GUNS, GERMS AND STEEL 98-99 (1997) [hereinafter DIAMOND, GUNS].
165. Domestication means altering the genetic composition of a plant or animal in order to make it more useful to humans. Id. at 14.
166. Id. at 100. The reason why some areas developed food production early, some late and some never at all (Australia), has little if anything to do with the differences in various human populations and very much to do with the native suite of domesticable plants and animals available to them during the Upper Paleolithic period. Put simply, some plants and animals are simply much better candidates for domestication than others, and those upper Paleolithic hunter-gatherers fortunate enough to find themselves in the vicinity of the wild ancestors of future domesticates benefited from the luck of the draw. What made a wild plant a good candidate for domestication by upper Paleolithic hunter gatherers? First it would have to be edible in the wild state and give high yields compared to other gathered plants. Ease of cultivation, by mere sowing or planting, was required because humans had no more sophisticated horticultural techniques. A short growing time minimized the delay of the return on the energy investment, and ease of storage provided a steady year-round source of nutrition. In addition to these obvious physical traits, two more subtle biological properties are crucial. A good potential domesticate should be self pollinating and thus able to breed true and retain its advantage in future generations. Plants lacking this feature might lose their selected genetic advantage in the process of random pollination by less suitable wild specimens. Moreover in an ideal domestication candidate, these six desirable crop traits should be obtainable with relatively little genetic modification of the wild precursor. See id. at 124. Fortunately upper Paleolithic hunter-gatherers did not need to know enough about horticulture to choose consciously the wild plants that had these characteristics. Hunter-gatherers did not gather indiscriminately (if they had they would not have lived long enough to pass on their genes); rather they consciously and unconsciously sought out those plants that were easy to find, nutritious, and required little time and effort to transform into edibility. These they consumed selectively, i.e., largest, sweetest, most resistant to climate extremes, and excreted their seeds nearby, thus increasing the frequency of the choice specimens in the next generation in the wild. Eventually purposeful selective propagation followed, and by evolution and later selective breeding, these wild precursors became the staple crops of the earliest Neolithic farmers. See id. at 123-30.
167. GREAT HUMAN DIASPORAS, supra note 27, at 134. The population of the world is estimated at 6 million at the dawn of the age of agriculture, and by the year 1700 it had reached 500 million. OLSON, supra note 3, at 101.
168. DIAMOND, GUNS, supra note 119, at 88; DIAMOND, THIRD CHIMPANZEE, supra note 54, at 189.
169. Chance again played a huge role in determining which societies would domesticate animals and how early in their history that development would occur. Once again, some animals are much better candidates for domestication than others, and the societies located in the natural ranges of these candidates developed herding early. Again it is important to define domestication. It consists of much more than mere taming. In addition to making the animal tolerate the presence and activities of humans, domestication requires the ability of the animal to reproduce in captivity and the ability of its human captors to breed it selectively. Thus in many cases (the cheetah and the elephant) animals have played a role in human history without ever being truly domesticated; rather each new specimen is caught wild. See DIAMOND, GUNS, supra note 119, at 159, 169-70. In order to be fully useful to human agriculturalists a candidate must be large - large enough to provide enough food to make the project worthwhile and large enough to be useful for draft duties. It must be efficient to feed; otherwise the result of keeping the animal would be a net loss rather than a gain; as a result few carnivores have been bred for food. It must mature quickly because few incipient herders are willing to invest the massive amount of food required to raise an animal to an exploitable age of more than a few years. So much for elephants, which are caught wild and tamed by Indians and Africans, rather than bred in captivity. The animal must be willing to breed in captivity, and that excludes some animals that even modern zoo keepers must inseminate artificially. Another required characteristic is a relatively mild and tractable disposition; bears and hippopotamuses fail spectacularly, but so, less obviously, do zebras and elk. A hyperactive startle reflex also precludes useful domestication of species like gazelles and most types of deer. Finally herding animals with defined dominance hierarchies and overlapping ranges are much better candidates than other herding species. Id. at 173.
170. GREAT HUMAN DIASPORAS, supra note 27, at 133-34.
171. See OLSON, supra note 3, at 103.
172. GREAT HUMAN DIASPORAS, supra note 27, at 9.
173. See DIAMOND, GUNS, supra note 119, at 112.
174. See id. at 211.
175. Id.
176. Id.
177. Id. at 150-51.
178. See id. at 186.
179. Id.
180. See id. at 187.
181. Id.
182. See id. at 315.
183. See Heartland, supra note 98, at 5 (stating that marker M172 is the signal of the spread of Neolithic farmers through Europe).
184. See WELLS, JOURNEY, supra note 39, at 152.
185. Its most dramatic examples would occur thousands of years later when European agriculturalists displaced Native American, African and Australian hunter-gatherers. In some places where European crops flourished (North America, the Cape area of Africa, and parts of Australia and Tasmania) the replacement was almost total. In other areas where European crops and animals fared less well (Australia and parts of South America), the replacement was less thorough.
186. For genetic evidence of the expansion, see Stephanie Plaza et al., Insights into the Western Bantu Dispersal: mtDNA Lineage Analysis in Angola, 115 HUM. GENETICS 439 (2004); Salas et al., supra note 63.
187. See also Onella Semino et al., Ethiopians and Khoisan Share the Deepest Clades of the Human Y-Chromosome Phytogeny, 70 AM. J. HUM. GENETICS 265 (2002).
188. DIAMOND, GUNS, supra note 119 at 396-97.
189. Unlike the click language speakers, the two groups of pygmies lost their native languages and now speak the languages of their Bantu neighbors. See supra note 62.
190. DIAMOND, GUNS, supra note 119, at 394.
191. WELLS, JOURNEY, supra note 39, at 156.
192. They bear the Y-chromosome marker M130. See supra notes 71-78 and accompanying text. The remnants of that early migration exist now in Asia as population and linguistic isolates such as the Andaman Islanders and the Negritos of the Philippines. WELLS, JOURNEY, supra note 39, at 74-75. Many admixed populations display that marker as do native Australians and New Guineans.
193. WELLS, JOURNEY, supra note 39, at 179.
194. Bing Su et al., Polynesian Origins: Insights from the Y Chromosome, 97 PROC. NAT'L. ACAD. SCI. 8225 (2000).
195. For evidence of this pattern in the Bantu expansion into Southern and Eastern Africa, see Elizabeth T. Wood et al., Contrasting Patterns of Y Chromosome and mtDNA Variation in Africa: Evidence for Sex-Biased Demographic Processes, 13 EUR. J. HUM. GENETICS 867, 875
196. WELLS, JOURNEY, supra note 39, at 113; DIAMOND, GUNS, supra note 119, at 291.
197. GREAT HUMAN DIASPORAS, supra note 27, at 138-39; Joaquim Fort, Toni Pujol & Luigi Luca Cavalli-Sforza, Paleolithic Populations and Waves of Advance, 14 CAMBRIDGE ARCH. J. 53 (2004).
198. WELLS, JOURNEY, supra note 39, at 154.
199. WELLS, JOURNEY, supra note 39, at 169.
200. See infra Part IV.A. An interesting satellite inquiry is whether the lifestyle of the remaining hunter-gatherers changed after the development of agriculture by their neighbors. Are today's remaining foragers remnants of an earlier way of life that was once ubiquitous, or are they occupying niches that did not even exist before their neighbors adopted food production? See Frank W. Marlowe, Hunter-Gatherers and Human Evolution, 14 EVOLUTIONARY ANTHROPOLOGY 54 (2005).
201. Tatiana Zerjal et al., The Genetic Legacy of the Mongols, 72 AM. J. HUM. GENETICS 717, 720 (2003) (about 8% of the region's male population and .5% of the world's male population carry the Y-chromosome of Genghis Khan and his male-line descendants).
202. See Antonio Salas et al., The African Diaspora: Mitochondrial DNA and the Atlantic Slave Trade, 74 AM. J. HUM. GENETICS 454, 463-64 (2004). Most of the transported populations (about 60%) came from western Africa with smaller portions coming from west-central Africa (about 30%) and southeastern Africa (less than 10%). The pervasive effect of the Bantu dispersals makes the original genetic homelands of those populations more difficult to pinpoint.
203. WELLS, JOURNEY, supra note 39 at 184-87.
204. Braman, supra note 7, at 1386 n.25.
205. Tishkoff & Kidd, supra note 10.
206. COON, HISTORY, supra note 11, at 190.
207. On Platonic types, see Kittles & Weiss, supra note 13.
208. This brief exposition is, of course, a gross simplification of a great diversity of thought on the history of the concept of race in America and Europe. For more complete treatment, see George J. Armelagos & Alan H. Goodman, Race, Racism and Anthropology, in BUILDING A NEW BIOCULTURAL SYNTHESIS: POLITICAL- ECONOMIC PERSPECTIVES ON HUMAN BIOLOGY 359 (Alan H. Goodman & Thomas L. Leatherman eds., 1998) [hereinafter Armelagos & Goodman]; Braman, supra note 7, at 1384-93; Lillquist & Sullivan, supra note 11, at 408-26.
209. Ossorio & Duster, supra note 15, at 116. The folk wisdom is alive and well and sometimes rationalized by pseudo science. See sources cited supra note 3. See also Armelagos, Race, supra note 15.
210. Tishkoff & Kidd, supra note 10, at S21; Smay & Annelagos, supra note 4, at 26.
211. See Armand Marie Leroi, Op-Ed., A Family Tree in Every Gene, N. Y. TIMES, Mar. 14, 2005, at A21;
212. Hua Tang et al., Genetic Structure, Self-Identified Race/Ethnicity, and Confounding in Case-Control Association Studies, 76 AM. J. HUM. GENETICS 268, 273-74 (2005) [hereinafter Hua Tang]; Rowe, supra note 17, at 60; Risch et al., supra note 18, at 2007.1.
213. See sources cited supra note 3.
214. DIAMOND, GUNS, supra note 119, at 36.
215. DIAMOND, THIRD CHIMPANZEE, supra note 54, at 23.
216. STRINGER & MCKIE, AFRICAN EXODUS, supra note 32, at 20.
217. They are more closely related to each other than to us, but we are only twice as different genetically from them as they are from each other. DIAMOND, THIRD CHIMPANZEE, supra note 54, at 24.
218. Id.
219. This view is known as the Recent African Origin Model. An alternative is the increasingly disfavored Multiregional Theory, which holds that homo-erectus and archaic humans migrated out of Africa and colonized other continents where each separate group independently evolved into different races of fully modern homo sapiens. See Cavalli-Sforza & Feldman, supra note 34.
220. STRINGER & MCKIE, AFRICAN EXODUS, supra note 32, at 117.
221. Id. at 20; Henry C. Harpending & Elise Eller, Human Diversity and Its History, in BIODIVERSITY (M. Kato & N. Takahata eds.) (forthcoming 2006) (manuscript at 2, available at http://www.continuitas.com/harpending_humandiversity.pdf);
222. Henry C. Harpending et al., Genetic Traces of Ancient Demography, 95 PROC. NAT'L. ACAD. SCI. 1961, 1961 (1998).
223. See supra Part III.B.2.
224. See supra Part III.B.3.
225. Ossorio & Duster, supra note 15, at 117; See also OLSON, supra note 3, at 29 (chimpanzees have more than twice as much variety in their mitochondrial DNA as do all six billion people); see generally Anne Fischer et al., Evidence for a Complex Demographic History of Chimpanzees, 21 MOLECULAR BIOLOGY & EVOLUTION 799 (2004) (characterizing patterns of genomic variation in central chimpanzees).
226. See Jorde & Wooding, supra note 69, at S28. For more on the homogeneity of human genomes, see E. Ya. Tetushkin, Genetics and the Origin of Human 'Races', 37 RUSSIAN J. GENETICS 853 (2001). Shortly before this article went to press, a research team led by J. Craig Ventner published the entire six-billion-letter sequence of one human, Ventner.
227. See Samuel Levy, Granger Sutton, Pauline C. Ng, Lars Feuk, Aaron L. Halpern et al., The Diploid Genome Sequence of an Individual Human, Vol. 5, No. 10 PLoS BIOLOGY 1371 (2007). One of the key findings that emerged from that project is that there is much more genetic variation among humans than had been thought previously. This finding runs counter to the results of nearly all previous studies on human genetic variation, from the early work of Lewontin on proteins, see infra note 180, to contemporary genomic studies, see, e.g., Jorde & Wooding, supra note 69, but the finding has yet to be replicated. Similarly the implications of this finding for intra-ethnic versus inter-ethnic genetic variation have yet to be determined.
228. Physical anthropologists have obsessed over measuring and cataloguing every sort of difference among humans. See GOULD, MISMEASURE, supra note 3; Olson, supra note 3, at 179.
229. See Richard C. Lewontin, Confusions About Human Races, Apr. 20, 2005, http:// raceandgenomics.ssrc.org/Lewontin/.
230. Ossorio & Duster, supra note 15, at 117; Cavalli-Sforza & Feldman, supra note 34, at 268-69. See also Noah A. Rosenberg et al., Genetic Structure of Human Populations, 298 SCI. 23 81 (2002) (placing the amount of intra-group variation as high as 95%).
231. While there is some agreement on the finding of 15% variation among human groups there is less agreement on the proportion of that variation that can be attributed to differences between continents versus differences between populations within each continent. Compare Ossorio & Duster, supra note 15, at 117 (attributing only 5% to inter-continental difference and 10% to intra-continental variation) with Robert J. Sternberg, Elena L. Grigorienko & Kenneth K. Kidd, Intelligence, Race, and Genetics, 60 AM. PSYCHOLOGIST 46, 54 (2005) (finding 9% of the variation within continental races and 6-7% between them), and Lewontin, supra note 180 (finding the 15% variation about evenly split between inter-continental and intra-continental differences). In the remainder of this article, I use this last estimate; although it is the most conservative, it still provides strong support for the thesis.
232. See Ossorio & Duster, supra note 15, at 117; Lewontin, supra note 180, at 1; Cavalli-Sforza & Feldman, supra note 34, at 269.
233. All humans are 99.9% similar. If the Blair/Putin pair displays 90% of the total human diversity (.1%), they will be 99.910% [99.9% + (10% of .1%)] similar. If the Blair/Anan pair displays 20% of the available human diversity, they will be 99.980% [99.9% + (80% of .1%)] similar. Thus the Blair/Putin difference (100% - .01% = .09%) will be 4.5 times as great as the Blair/Anan difference (100% - .02% = .02); i.e., Blair will be 4.5 times more similar to Anan than to Putin. Genetic diversity occurs not only in the most visible "racial" traits such as skin color and hair form, but also with a host of "invisible" differences having to do with the production of hormones such as those controlling growth, adrenaline production, height, resistance to tooth decay, dyslexia, resistance to various kinds of cancer, etc. The point is that the generic variation that gives rise to the visible "racial" markers is only a small part of the total variation. Thus, while Blair is closer to Putin on the visible "racial" traits, he might be closer to Anan on a host of less visible but more significant traits, the distribution of which is not a function of geography. This useful clarification was suggested by David F. Armstrong.
234. On lumpers and splitters, see generally GOULD, MISMEASURE, supra note 3. The extreme among splitters was a group of taxonomists known as polygenists (literally, "many beginnings") that attributed separate creation of the several races. The opposite group monogenists believed in a single creation, but arranged races along a hierarchy. Both groups embraced biological essentialism and thus racism. Id. at 39.
235. OLSON, supra note 3, at 63. The measure is of most interest to conservation biologists who must determine what, if any, unique genetic stock exists in a particular threatened habitat. S.O.Y. Keita et al., Conceptualizing Human Variation, 36 NATURE GENETICS SUPPLEMENT S17 (2005).
236. OLSON, supra note 3, at 63.
237. Lillquist & Sullivan, supra note 11, at 409-10.
238. Id. at 410.
239. See WELLS, JOURNEY, supra note 39 at 191-92.
240. OLSON, supra note 3, at 49. Some writers have overstated this point and concluded that all human groupings are social constructs. That exaggeration, so obviously belied by observation (pygmies really are short), has produced skepticism about the truth and political motivation for the very different, and factually correct, assertion that continental race is not a necessary or even a useful way to sort human genetic variation.
241. See Smay & Armelagos, supra note 4, at 23.
242. Tishkoff & Kidd, supra note 10, at S24, S25 tbl.1. One commentator suggests that the greater climatic hardships endured by Asian migrants forced them to develop better intellects in order to survive. Africans, who remained in the relatively benign tropics, did not need the extra brain power. RUSHTON, supra note 3, at 263. Of course there is no empirical data to support this hypothesis, nor any way of quantifying the difficulties that different environments pose for human settlement. Nor is there any mention of the difficulty that greater population density and resource competition posed for Africans. Finally, if Asians endured climatic hardships, Native Americans presumably endured worse during hundreds of generations in Northern Siberia and Berrengia, but Rushton does not place them at the top of the intellectual pyramid. This is not science. Armelagos, Race, supra note 15, at 106-07.
243. Other factors also help explain the different appearances of different peoples. For example, sexual selection may explain differences that center on features that have cultural norms of beauty (breast, lips and buttocks shape). DIAMOND, THIRD CHIMPANZEE, supra note 54, at 120. Chance also plays a role. If a disproportionate percentage of a small founding population possesses a trait, a similarly disproportionate number of the much larger descendant population will possess it also. This may explain the concentration of blonds in far northern Europe and its absence in similar regions in Asia.
244. Pygmies, Negritos and Amazonian Indians all have developed small size but are phylogenetically distant.
245. See MAYR, EVOLUTION IS, supra note 13, at 156.
246. SCOTT WIEDENSAUL, RAPTORS: THE BIRDS OF PREY 6-7 (1995).
247. Several species of large mammals have developed sub-speciation because their former ranges have been split by human development. Examples are North American cougars, http://www.cotf.edu/ete/modules/everglades/FEpanther.html; gorillas, http://nationalzoo.si.edu/Animals/Primates/Facts/FactSheets/Gorillas/ default.cfm; and worldwide populations of grey wolves, http://www. kerwoodwildlife.com/GRAYWOLFSUBSPECIES.htm. For a general discussion of isolation mechanisms that can produce speciation, see OLSON, supra note 3, at 21-23.
248. For example, there is considerable supra-Saharan (especially Moroccan) admixture in some West African populations, including some that eventually ended up in the Americas via the Middle Passage. See generally Alexandra Rosa et al., MtDNA Profile of West Africa Guineans: Towards a Better Understanding of the Senegambia Region, 68 ANNALS HUM. GENETICS 340 (2004).
249. WELLS, JOURNEY, supra note 39 at 113, 166.
250. Lillquist & Sullivan, supra note 11, at 420.
251. Tishkoff & Kidd, supra note 10, at S22; Jeffrey Long & Rick A. Kittles, Human Genetic Diversity and the Nonexistence of Biological Races, 75 HUM. BIOLOGY 449, 467 (2003); Tishkoff & Verrelli, supra note 21, at 305.
252. They do, however, display a component of human diversity not contained in Africa. It is composed of the mutations that occurred during the migrations that peopled the rest of the world. The migrational history of those non-African populations is short (about 60,000 years) compared to the age of African populations, which continued to accumulate mutations during the same period. Thus, while there is a uniquely non-African component of total human diversity, it is much smaller than the diversity component that is uniquely African.
253. See supra Part III.B.
254. The question marks in the diagram are designed to show that human genetic differentiation did not stop at the identified groups; it continued on the left-hand branches of the diagram much as it did on the right. The vagueness of the descriptions on the left side results from much less study of the substructure and phylogeny of those populations. While this diagram is more accurate than the six-race model, it is still flawed. First it is very poor in resolution. Compare the much greater resolution in Underhill, Inferring, supra note 77, at 487, 489 fig.1. Moreover, the diagram purports to represent human phylogeny only, not the entire range of human genetic or morphological variation. A diagram more representative of total human genetic variation would have much fuzzier, more overlapping groups and many more crisscrossing lines, revealing the constant history of population intermixture. A Venn diagram could capture the relationship of human genetic variation to continent of ancestry. Imagine six nearly congruent circles (representing the six inhabited continents), each about six inches in diameter. Now suppose that the center points of each of the six fit within a one millimeter square. The huge area common to all six circles would represent the portion of the human genome that does not vary by continent of ancestry. The slivers near the outside boundary of the figure would be shaped by their inclusion within one circle but not the others. These slivers would then portray proportionally the amount of human genetic variation that is accounted for by continent of origin. Such a diagram would show accurately the contribution of continent of origin to total human genetic diversity, but it would not represent phylogeny, which the dendrogram does.
255. Traditionally, taxonomists used phenotypy to classify organisms. Today, however, scientists have the additional tools of genomics and generally accept the primacy of phylogeny (evolutionary history) over phenotypy. See generally Keita, supra note 186.
256. Id. at S18.
257. Multi-ethnic classifications do add an additional layer of complication. It is, of course, only the groupings that are not problematic; no one would claim that there are no social problems generated by and incident to race in the United States.
258. By "incoherent" I do not mean "not understandable" or "confused," but rather the older literal meaning of the term in which a group is incoherent when its members do not cohere; in this case they do not cohere genetically.
259. Hua Tang, supra note 165, at 273; Keita, supra note 186, at S19.
260. See supra Part III.B.2.
261. Ossorio & Duster, supra note 15, at 18; Keita, supra note 186, at S19; cf. Hua Tang, supra note 165, at 268 (continuous range of European admixture averaging 10-20%).
262. See generally Rosa, supra note 199 (detailing the many lineages that make up the population of one of many West African regions).
263. That is a nationwide average; but admixture differs substantially by region, averaging 10% in the South and 50% in the North. There is also some Native American and Hispanic admixture. GENES, PEOPLES, AND LANGUAGES, supra note 33, at 74-75.
264. Historically it was advantageous for some such people to violate the "one-drop" rule and "pass" as white.
265. The criteria for membership in the socio-cultural category is fuzzy and is determined by each tribe. Some tribes require only one third ancestry. See Race (United States Census), http://en.wikipedia.org/wiki/ Native_American_%28U.S._Census%29.
266. GENES, PEOPLES AND LANGUAGES, supra note 33, at 23.
267. Id. at 76.
268. The villain is Hugo Drax; the movie is Moonraker (1979); Drax is addressing James Bond, and referring to Bond's repeated efforts to thwart Drax's plans.
269. See supra note 7 and accompanying text.
270. Lewontin, supra note 180, ¶ 6; Tishkoff & Kidd, supra note 10, at S26 (37 populations studied using over 80 independent genome loci); Hua Tang, supra note 165(10,527 participants grouped using 326 genetic markers).
271. The program, known as "Structure," is succinctly described by Tishkoff & Kidd, supra note 10, at S25.
272. See Hua Tang, supra note 165; Joanna L. Mountain & Neil Risen, Assessing Genetic Contributions to Phenotypic Differences Among 'Racial' and 'Ethnic' Groups, 36 NATURE GENETICS SUPPLEMENT S48, S48 (2004); Risch et al., supra note 18, at 2007.
273. Leroi, supra note 165 ("[O]ne thing was clear; the consensus about social constructs was unraveling."); Risch et al., supra note 18; Rowe, supra note 17.
274. There is yet another variable that may affect the extent of continental clustering observed in these studies: the genetic markers selected for study. When studies are limited to coding regions of the DNA (those that have a recognizable effect on the organism's structure and function), continental clustering is not very pronounced; when instead the selected markers are non-coding - especially those that show the highest inter-population variability - clustering is more evident. See Tishkoff & Kidd, supra note 10, at S24; JOSEPH L. GRAVES, JR., THE RACE MYTH: WHY WE PRETEND RACE EXISTS IN AMERICA 112-17 (2004) [hereinafter GRAVES, MYTH]; Lewontin, supra note 180, ¶ 7; Kittles & Weiss, supra note 13, at 48-53.
275. See GRAVES, MYTH, supra note 225, at 112-17.
276. Joseph L. Graves, Jr., What We Know and What We Don't Know: Human Genetic Variation and the Social Construction of Race, ¶ 12, Apr. 25, 2005, http://raceandgenomics.ssrc.org/Graves/[hereinafter Graves, Variation].
277. See Jorde & Wooding, supra note 69, at S29; Tishkoff & Kidd, supra note 10, at S25; Kittles & Weiss, supra note 13, at 38.
278. Troy Duster, Race and Reification in Science, 307 Sci. 1050, 1050-51 (2005).
279. Note, for instance, the close genetic relationship of the populations of Northeast Ethiopia and the portion of Yemen just across the Red Sea at Bab-el-Mandeb. Kivisild, supra note 69.
280. See Kittles & Weiss, supra note 13, at 38; Tishkoff & Kidd, supra note 10, at S25; Jorde & Wooding, supra note 69, at S29. At least some evidence for this prediction comes from studies that show that Middle Easterners and Central Asians had partial membership in multiple clusters. Tishkoff & Kidd, supra note 10, at S25. Strangely, few studies sample Indians, prompting one pair of commentators to ask, "Who are those pesky billion? One race? A mix of the already-sampled races? A multiplicity of races as has often been suggested?," Kittles & Weiss, supra note 13, at 38. Also artificially exaggerating the clustering effect is the inclusion of only one or two African populations, in spite of the well-recognized finding that intra-African genetic diversity exceeds that of all the rest of the world's populations combined. Charles N. Rotimi, Are Medical and Nonmedical Uses of Large-Scale Genomic Markers Conflating Genetics and 'Race'?, 36 NATURE GENETICS SUPPLEMENT S43, S44 (2004).
281. Rosenberg et al., supra note 181, at 2382.
282. Jonathon Marks summarizes the methodological difficulties associated with using cluster studies to establish the reality of races: [A] cluster analysis . . . is sensitive to the population samples chosen, the individual people representing them, the demographic history of the populations, the assumptions of the particular algorithm, and the patterns of contact among the populations. In other words, the species still doesn't come prepackaged . . . ; you still have to decide, given the fact of difference, how much and what kind is meaningful and how much and what kind is not. Jonathan Marks, The Realities of Races, ¶ 14, Apr. 20, 2005, http://raceandgenomics.ssrc.org/Marks/.
283. See Keita, supra note 186, at S19.
284. The tendency to assume that the markers cluster concordantly relies on the fallacy that because differences are genetic that they also are pan-racial. Armelagos & Goodman, supra note 162, at 369.
285. Christen Brownlee, Code of Many Colors: Can Researchers See Race in the Genome?, 167 SCI. NEWS 232 (2005), available at http://www.sciencenews.org/articles/20050416/bob.asp ¶ 30 (2005) (paraphrasing Rosenberg et al., supra note 181).
286. Indeed, studies often do not assign an individual to a cluster absolutely; rather, they affix a probability of, say 90%, to membership in the assigned group and some smaller percentage probability of membership in one or more of the other groups. Jorde & Wooding, supra note 69, at S32.
287. See text following note 202, supra.
288. See Jorde & Wooding, supra note 69, at S30 (no clear boundaries); Long & Kittles, supra note 202, at 468; Rotimi, supra note 231, at S44.
289. See supra Part III.A.
290. For an explanation of the phenomenon, see Tishkoff & Kidd, supra note 10, at S23; Graves, Variation, supra note 227, ¶ 17.
291. See supra Part III.A
292. It has spawned a debate in the medical and pharmacological communities, over whether the geographically-mediated differences in disease frequency and drug susceptibility within our species are sufficient to justify medical treatment and pharmacological research tailored to particular groups. It also affects the way forensic anthropologists go about the task of identifying unknown bodies and the way genetic identity evidence can be used in criminal cases.
293. Braman, supra note 7.
294. Lillquist & Sullivan, supra note 11.
295. What then has driven a socio-cultural construction of race that is so far removed from any biologically justified model? One of many reasons (the one important to the theme of this article) is historical; modernity in science coincided with the era of European exploration and colonialization. Journeys of thousands of miles by ship gave the explorers and their accompanying naturalists a very different view of human variation than previous forays conducted over land. Land travelers see the country side between their homes and their destinations in detail and at close resolution; human variation appears continuous and clinal. Populations vary, of course, but the pattern of that variation appears as a gradient, not as a set of discontinuities separated by clear boundaries. By contrast long ocean voyages do not reveal clines of human variation; they drop the traveler into a completely different geographical environment populated by humans who look very different indeed. That artifact of modern travel and exploration goes a long way toward explaining the tendency of Europeans to see humanity as set of discrete and fundamentally distinct clades. See STRINGER & MCKIE, AFRICAN EXODUS, supra note 32, at 191-92. At the same time political events encouraged racial essentialism. The enlightenment, the rise of capitalism, egalitarianism and meritocracy led Jefferson, among others, to proclaim all men equal and equally possessed of natural rights. That noble sentiment did not fit in the same moral universe created by European colonization in Asia, Africa and the Americas and the development of a new and especially brutal form of chattel slavery in America. Racial essentialism was an ideal doctrine to cure the psychic dissonance. If there were fundamental differences in talent and character among groups, then even titular egalitarians could justify different treatment. Thus race, while biologically tenuous, became culturally constructed and entrenched.