How neandertals inform human variation

American Journal of Physical Anthropology

Volumn 139, Issue 1, 2009, Pages 91-102

  Wolpoff, Milford H ^a  

^a UNIVERSITY OF MICHIGAN   (United States)

Author keywords

Human variation; Neandertal; Race; Subspecies

Indexed keywords

ALLELE; ARTICLE; EVOLUTION; FOSSIL; GENE FLOW; GENETIC VARIABILITY; HOMINID; HUMAN; NEANDERTAL; PHYLOGENY; RACE;

ANIMALS; CONTINENTAL POPULATION GROUPS; EVOLUTION; GENE FLOW; GENETIC VARIATION; HOMINIDAE; HUMANS; PHENOTYPE; SKULL;

EID: 66949178029     PISSN: 00029483     EISSN: 10968644     Source Type: Journal    
DOI: 10.1002/ajpa.20930     Document Type: Article

References (111)

1. Ahern JCM. 2006. Non-metric variation in recent humans as a model for understanding Neanderthal-early modern differences: just how "unique" are Neanderthal unique traits? In: Havarti K, Harrison T, editors. Neanderthals revisited: new approaches and perspectives. Dordrecht: Springer. p 255-268.
2. Bazin E, Gleemin S, Galtier N. 2006. Population size does not influence mitochondrial genetic diversity in animals. Science 312:570-572.
3. Boule M. 1923. Les Hommes Fossiles. Elements de Paleeontolo- gie Humaine. Paris: Masson.
4. Brace CL. 1967 The stages of human evolution. Englewood Cliffs: Prentice-Hall.
5. Brauer G. 2008. The origin of modern anatomy: by speciation or intraspecific evolution? Evol Anthropol 17:22-37.
6. Briggs AW, Stenzel U, Johnson PLF, Green RE, Kelso J, Prufer K, Meyer M, Krause J, Ronan MT, Lachmann M, Paabo S. 2007. Patterns of damage in genomic DNA sequences from a Neandertal. Proc Nat Acad Sci USA 104:14616-14621.
7. Caspari R. 2003. From types to populations: a century of race, physical anthropology and the American Anthropological Association. Am Anthropol 105:63-74.
8. Churchill SE, Smith FH. 2000. Makers of the early Aurignacian of Europe. Yrbk Phys Anthropol 43:61-115.
9. Clark AG. 2008. Genome sequences from extinct relatives. Cell 134:388-389.
10. Coop G, Bullaughey K, Luca F, Przeworski M. 2008. The timing of selection at the human FOXP2 gene. Mol Biol Evol 25:1257-1259.
11. Crummett TL. 1994. The evolution of shovel shaping: regional and temporal variation in human incisor morphology, Ph.D. dissertation, University of Michigan. Ann Arbor: University Microfilms.
12. Curnoe DA, Thorne AG, Coate JA. 2006. Timing and tempo of primate speciation. J Evol Biol 19:59-65.
13. Deeb SS, J0rgensen AL, Battisi L, Iwasaki L, Motulsky AG. 1994. Sequence divergence of the red and green visual pigments in the great apes and man. Proc Natl Acad Sci USA 91:7262-7266.
14. D'Errico F. 2003. The invisible frontier: a multiple species model for the origin of behavioral modernity. Evol Anthropol 12:188-202.
15. D'Errico F, Julien M, Liolios D, Van Haren M, Baffier D. 2003. Many awls in our argument. Bone tool manufacture and use in the Chatelperronian and Aurignacian levels of the Grotte de Renne at Arcy-sur-Cure. In: Zilhao J, D'Errico F, editors. The chronology of the Aurignacian and of the transitional technocomplexes. Dating, stratigraphies, cultural implications. Trabalhos de Arqueologica 33:247-270.
16. Dobzhansky TH 1944. On species and races of living and fossil man. Am J Phys Anthropol 2:251-265.
17. Duarte C, Maurlcio J, Pettitt PB, Souto P, Trinkaus E, van der Plicht H, Zilhao J. 1999. The Early Upper Paleolithic human skeleton from the Abrigo do Lagar Velho (Portugal) and modern human emergence in Iberia. Proc Natl Acad Sci USA 96:7604-7609.
18. Enard W, Przeworski M, Fisher SE, Lai CSL, Wiebe V, Kitano T, Monaco AP, Paabo S. 2002. Molecular evolution of FOXP2, a gene involved in speech and language. Nature 418:869-872.
19. Eswaran V, Harpending H, Rogers AR. 2005. Genomics refutes an exclusively African origin of humans. J Hum Evol 49:1-18.
20. Evans PD, Gilbert SL, Mekel-Bobrov N, Vallender EJ, Anderson JR, Vaez-Azizi LM, Tishkoff SA, Hudson RR, Lahn BT. 2005. Microcephalin, a gene regulating brain size, continues to evolve adaptively in humans. Science 309:1717-1720.
21. Evans PD, Mekel-Bobrov N, Vallender EJ, Hudson RR, Lahn BT. 2006. Evidence that the adaptive allele of the brain size gene microcephalin introgressed into Homo sapiens from an archaic Homo lineage. Proc Natl Acad Sci USA 103:18178-18183.
22. Excoffier L. 2006. Neandertal genetic diversity: a fresh look from old samples. Cur Biol 16:R650-R652.
23. Eyre-Walker A. 2006. Size does not matter for mitochondrial DNA. Science 312:537-538.
24. Fraipoint J, Lohest M. 1887. La race humaine de Neanderthal ou de Canstadt en Belgique: recherches ethnographiques sur des ossements humaines, deecouverts dans depots, quatern- aires d'une grotte a Spy et determination de leur age geeologi- que. Arch Biol (Liege) 7:587-757.
25. Frayer DW. 1992. Evolution at the European edge: Neanderthal and the Upper Paleolithic relationships. Prehist Europeenne 2:9-69.
26. Frayer DW. 1997. Perspectives on Neanderthals as ancestors. In: Clark GA, Willermet CM, editors. Conceptual issues in modern human origins research. New York: Aldine de Gruyter. p 220-235.
27. Frayer DW, Jelinek J, Oliva M, Wolpoff MH. 2006. Aurignacian male crania, jaws, and teeth from the Mladec Caves, Moravia, Czech Republic. In: Teschler-Nicola M, editor. Early modern humans at the Moravian Gate: The Mladec caves and their remains. Wien: Springer. p 185-272.
28. Frayer DW, Wolpoff MH. 2008. The importance of the Prrcaron; edmosti human remains. In: Veleminska J, BriiZek J, editors. Early modern humans from Prmosti near Prerov, Czech Republic. Prague: Academia. Chapter 4, p 35-30.
29. Futuyma DJ. 1986. Evolutionary biology. Sunderland: Sinauer. Garrigan D, Hammer MF. 2006. Reconstructing human origins in the genomic era. Nat Rev Genet 7:669-680.
30. Garrigan D, Kingan SB. 2007. Archaic human admixture, a view from the genome. Cur Anthropol 48:895-902.
31. GarriganD,MobasherZ,KinganSB, WilderJA,HammerMF. 2005a. Deep haplotype divergence and long-range linkage disequilibrium at Xp21.1 provides evidence that humans descend from a structured ancestral population. Genetics 170:1849-1856.
32. Garrigan D, Mobasher Z, Severson T, Wilder JA, Hammer MF. 2005b. Evidence for archaic Asian ancestry on the human X chromosome. Mol Biol Evol 22:189-192.
33. Gorjanovic-Kramberger D. 1904. Potijc e li moderni covjek ravno od diluvijalonga Homo primigeniusa? Beograd: I Kon- gress srpskih lekara i Prirodnjaka. p 1-8.
34. Graves LL. 2001. The emperor's new clothes: biological theories of race at the millennium. New Brunswick: Rutgers University Press.
35. Green RE, Krause J, Ptak SE, Briggs AW, Ronan MT, Simons JF, Du L, Egholm M, Rothberg JM, Paunovic M, Paabo S. 2006. Analysis of one million base pairs of Neanderthal DNA. Nature 444:330-336.
36. Green RE, Malaspinas A-S, Krause J, Briggs AW, Johnson PLF, Uhler C, Meyer M, Good JM, Maricic T, Stenzel U, Prufer K Siebauer M, Burbana HA, Ronan M, Rothberg JM, Egholm M, Rudan P, Brajkovic D, Kucan Z, Gusic I, WilkstromM, Laakkonen L, Kelso J Slatkin M. 2008. A complete Neander- tal mitochondrial genome sequence determined by high- throughput sequencing. Cell 134:416-426.
37. Harding RM, Healy E, Ray AJ, Ellis NS, Flanagan N, Todd C, Dixon C, Sajantila A, Jackson IJ, Birch-Machin MA, Rees JL. 2000. Evidence for variable selective pressures at MC1R. Am J Hum Genet 66:1351-1361.
38. Hardy J, Pittman A, Myers A, Gwinn-Hardy K, Fung HC, de Silva R, Hutton M, Duckworth J. 2005. Evidence suggesting that Homo neanderthalensis contributed the H2 MAPT haplo- type to Homo sapiens. Biochem Soc Trans 33:582-585.
39. Hawks, J. 2006. Selection on mitochondrial DNA and the Neanderthal problem. In: Harvati K, Harrison T, editors. Neanderthals revisited: new approaches and perspectives. Dordrecht: Springer. p 221-238.
40. Hawks J, Cochran G. 2006. Dynamics of adaptive introgression from archaic to modern humans. PaleoAnthropol 2006:101-115.
41. Hawks J, Cochran GM, Harpending HC, Lahn BT. 2007a. A genetic legacy from archaic Homo. Trends Genet 24:19-23.
42. Hawks J, Wang ET, Cochran GM, Harpending HC, Moyzis RK. 2007b. Recent acceleration of human adaptive evolution. Proc Natl Acad Sci USA 104:20753-20758.
43. Hawks J, Wolpoff MH. 2003. Sixty years of modern human origins in the American Anthropological Association. Am Anthropol 105:87-98.
44. Henke W. 2008. Gorjanovic Kramberger's research on Krapina - its impact on paleoanthropology in Germany. In: Monge J, Mann A, Frayer DW, Radovcic J, editors. New insights on the Krapina Neandertals. Croatian Natural History Museum: Zagreb. p 5-18.
45. Holliday TW. 2006. Neanderthals and modern humans: an example of a mammalian syngameon? In: Harvati K, Harrison T, editors. Neanderthals revisited: new approaches and perspectives. Dordrecht: Springer. p 281-298.
46. Hrdlicka A. 1930. The skeletal remains of early man. Smithsonian Miscellaneous Collections, Vol. 83. Washington DC: Smithsonian Institution.
47. Huxley JS, Haddon AC. 1935. We Europeans: a survey of 'racial' problems. London: Jonathan Cape.
48. Jolly C. 2001. A proper study of mankind: analogies from the papionin monkeys and their implications for human evolution. Yrbk Phys Anthropol 44:177-204.
49. King W. 1864. On the reputed fossil of man of the Neanderthal. Quart J Sci (Lond) 1:88-97.
50. Klaatsch H, Hauser O. 1910. Homo aurignacensis Hauseri. Ein palaolithischer Skeletfund aus dem untern Aurignacien der Station Combe-Capelle bei Montferrand (Periogord). Praehist Zeitschr 1:273-338.
51. Krause J, Lalueza-Fox C, Orlando L, Enard W, Green RE, Bur- bano HA, Hublin J-J, Bertranpetit J, Hanni C, Fortea J, de la Rasilla M, Rosas A, Paabo S. 2007a. The derived FoxP2 variant of modern humans was shared with Neandertals. Cur Biol 17:1-5.
52. Krause J, Orlando L, Serre D, Viola B, Prufer K, Ricards MP, Hublin J-J, Hanni C, Derevianko AP, Paabo S. 2007b. Neanderthals in central Asia and Siberia. Nature 449:902-904.
53. Krings M, Stone A, Schmitz RW, Krainitzid H, Stoneking M, Paabo S. 1997. Neandertal DNA sequences and the origin of modern humans. Cell 90:1-20.
54. Lieberman DE. 2008. Speculations about the selective basis for modern human cranial form. Evol Anthropol 17:55-68.
55. Marks J. 1995. Human biodiversity: genes, race, and history. New York: Aldine de Gruyter.
56. Martinez I, Arsuaga JL, Quam R, Carretero JM, Gracia A, Rodriguez L. 2008. Human hyoid bones from the Middle Pleistocene site of the Sima de los Huesos (Sierra de Atapuerca, Spain). J Hum Evol 54:118-124.
57. Martinez I, Rosa M, Arsuaga J-L, Jarabo P, Quam R, Lorenzo C, Gracia A, Carretero J-M, Bermudez de Castro J-M, Car- bonell E. 2004. Auditory capacities in Middle Pleistocene humans from the Sierra de Atapuerca in Spain. Proc Natl Acad Sci USA 101:9976-9981.
58. Mayr E. 1942. Systematics and the origin of species. New York: Columbia University Press. Mayr E. 1963. Animal species and evolution. Cambridge: Belknap Press of Harvard University Press. Mayr E. 1969. Principles of systematic zoology. New York: McGraw-Hill.
59. McCown TD, Keith A. 1939. The Stone Age of Mount Carmel: The fossil human remains from the Levalloiso-Mousterian, Vol. 2. Oxford: Clarendon Press. Montagu MFA, editor. 1964. The concept of race. New York: The Free Press.
60. Morral N, Nunes V, Casals T, Chillon M, Gimenez J, Bertranpetit J, Estivill X. 1993. Microsattelite haplotypes for cystic fi- brosis: mutation frameworks and evolutionary tracers. Hum Mol Genet 2:1015-1022.
61. Pearson K, Davin AG. 1924. On the biometric constants of the human skull. Biometrika 16(3/4):328-363.
62. Pearson OM. 2008. Statistical and biological definitions of ''anatomically modern'' humans: Suggestions for a unified approach to modern morphology. Evol Anthropol 17:38-48.
63. Plagnol V, Wall JD. 2006. Possible ancestral structure in human populations. PLoS Genetics 2:e105.
64. Quennell M, Quennell CHB. 1945. Everyday life in the Old Stone Age. London: Batsford.
65. Radovcic J. 1988. Dragutin Gorjanovic-Kramberger and Krapina Early Man: The foundation of modern paleoanthropology. Zagreb: Skolska knjiga and Hrvatski prirodoslovni muzej.
66. Ranke J. 1896. Der fossile Mensch und die Menschenrassen. Correspondenzblatt der Deutschen Gesellschaft fur Anthropologie, Ethnologie, und Urgeschichte. Archiv fur Anthropologie 27:151-156.
67. Relethford JH. 2008. The human species, 7th ed. Boston: McGraw-Hill.
68. van Riper AB. 1993. Men among the mammoths: Victorian science and the discovery of human prehistory. Chicago: University of Chicago Press.
69. Rougier H, Milota S, Rodrigo R, Gherase M, Sarcina L, Moldo- van O, Zilhao J, Constantin S, Franciscus RG, Zollikofer CPE, Ponce de Leon M, Trinkaus E. 2007. Pestera cu Oase 2 and the cranial morphology of early modern Europeans. Proc Natl Acad Sci USA 104:1165-1170.
70. Sackett JR. 2000. Human antiquity and the Old Stone Age: the nineteenth century background to paleoanthropology. Evol Anthropol 9:37-49.
71. Sampietro ML, Gilbert MTP, Lao O, Caramelli D, Lari M, Bertranpetit J, Lalueza-Fox C. 2006. Tracking down human contamination in ancient human teeth. Mol Biol Evol 23:1801-1807.
72. Schmitz RW. 2006. A romantic valley and a discovery that changed the world. In: Schmitz RW, editor. Neanderthal 1856-2006. Mainz am Rhein: Verlag Philipp von Zabern. p 35-44.
73. Schoetensack O. 1908 Der Unterkiefer des Homo heidelbergen- sis aus den Sanden von Mauer bei Heidelberg. Leipzig: W. Engelmann.
74. Schwalbe G. 1901. Der Neanderthalschadel. Bonner Jahrbucher 106:1-72.
75. Schwalbe G. 1906 Studien zur Vorgeschichte des Menschen. I. Zur Frage der Abstammung des Menschen. Stuttgart: Schwei- zerbartsche Verlagsbuchhandlung. p 10-80.
76. Sergi S. 1953. Morphological position of the ''Prophaneran- thropi'' (Swanscombe and Fontechevade). In: Howells WW, editor. Ideas on human evolution. Cambridge: Harvard University. p 507-520. (Reprinted and translated, 1962.)
77. Soficaru A, Catalin P, Dobos A, Trinkaus E. 2007. The human cranium from the Pestera Cioclovina Uscata;. Romania. Current Anthropology 48:611-619.
78. Soficaru A, Dobos A, Trinkaus E. 2006. Early modern humans from the Pestera Muierii. Baia de Fier, Romania. Proc Natl Acad Sci USA 103:17196-17201.
79. Stefansson H, Helgason A, Steinthorsdottir GTV, Masson G, Bernard J, Baker A, Jonasdottir A, Ingason A, Gudnadottir VG, Desnica N, Hicks A, Gylfason A, Gudbjartsson DF, Jons- dittir GM, Sainz J, Agnarsson K, Birgisdottir B, Ghosh S, Olafsdottir A, Cazier JB, Kristjansson K, Frigge ML, Thor- geirsson TE, Gulcher JR, Kong A, Stefansson K. 2005. A common inversion under selection in Europeans. Nat Genet 37:129-137.
80. Stolyhwo K. 1908. Homo primigenius appartient-il a une espeece distincte de Homo sapiens? L'Anthropol 19:191-216.
81. Stolyhwo K. 1937. Les Preeneeanderthaloides et leurs rapports avec la race de Neanderthal. Bulletin Ethnologique de Museee d'Ethnographie de Ljubljana 10:147-168.
82. Swed FS, Eisenhart C. 1943. Tables for testing randomness of grouping in a sequence of alternatives. Ann Math Stat 14:66-87.
83. Szombathy J. 1925. Die diluvialen Menschenreste aus der Fuirst-Johanns-Hoihle bei Lautsch in Maihren. Die Eiszeit 2:1-34, 73-95.
84. Tattersall I. 2000. Paleoanthropology: the last half-century. Evol Anthropol 9:2-16.
85. Tattersall I, Schwartz JH. 2008. The morphological distinctive- ness of Homo sapiens and its recognition in the fossil record: clarifying the problem. Evol Anthropol 17:49-54.
86. Templeton AR. 1998. Human races: a genetic and evolutionary perspective. Am Anthropol 100:632-650.
87. Templeton AR. 2005. Haplotype trees and modern human origins. Yearbk Phys Anthropol 48:33-59.
88. Trinkaus E. 2006. Modern human versus Neandertal distinctiveness. Curr Anthropol 47:597-620.
89. Trinkaus E. 2007. European early modern humans and the fate of the Neandertals. Proc Natl Acad Sci USA 104:7367-7372.
90. Trinkaus E, Shipman P. 1993. The Neanderthals: changing the image of mankind. New York: Knopf.
91. Trinkaus E, Milota S, Rodrigo R, Mircea G, Moldovan O. 2003. Early modern human cranial remains from the Pestera cu Oase. Romania. J Hum Evol 45:245-253.
92. Weidenreich F. 1943a. The ''Neanderthal Man'' and the ancestors of ''Homo sapiens''. Am Anthropol 45:39-48.
93. Weidenreich F. 1943b. The skull of Sinanthropus pekinensis:a comparative study of a primitive hominid skull. Palaeontolo- gia Sinica, New Series D, number 10 (whole series number 127).
94. Weidenreich F. 1946. Generic, specific, and subspecific characters in human evolution. Am J Phys Anthropol 4:413-431.
95. Weidenreich F. 1947. Are human races in the taxonomic sense ''races'' or ''species''? Am J Phys Anthropol 5:369-371.
96. Weinert H. 1925. Der Schaidel des eiszeitlichen Menschen von Le Moustier. Berlin: Springer.
97. Wilson EO, Brown WL. 1953. The subspecies concept and its taxonomic application. Syst Zool 2:97-111.
98. Wolpoff MH, Caspari R. 1996. An unparalleled parallelism. Anthropologie (Brno) 34:215-223.
99. Wolpoff MH, Caspari R. 1997. Race and human evolution. New York: Simon and Schuster.
100. Wolpoff MH, Hawks JD, Caspari R. 2000. Multiregional, not multiple origins. Am J Phys Anthropol 112:129-136.
101. Wolpoff MH, Hawks JD, Frayer DW, Hunley K. 2001. Modern human ancestry at the peripheries: a test of the replacement theory. Science 291:293-297.
102. Wolpoff MH and Lee Sang-Hee. 2007. Herto and the Neandertals: what can a 160,000-year-old African tell us about European Neandertal evolution? In: Sankhyan AR, Rao VR, editors. Human origins, genome and people of India: genomic, palaeontological and archaeological perspectives. New Delhi: Allied Publishers. p 329-336.
103. Wolpoff MH, Mannheim B, Mann A, Hawks J, Caspari R, Rosenberg KR, Frayer DW, Gill GW, Clark GA. 2004. Why not the Neandertals? World Archaeol 36:527-546.
104. Wolpoff MH, Wu Xinzhi, Thorne AG. 1984. Modern Homo sapiens origins: a general theory of hominid evolution involving the fossil evidence from east Asia. In: Smith FH, Spencer F, editors. The origins of modern humans: a world survey of the fossil evidence. New York: Alan R Liss. p 411-483.
105. Zaingl U. 2006. Hermann Schaaffhausen (1816-1893) and the Neanderthal finds of the 19th Century. In: Schmitz RW, editor. Neanderthal 1856-2006.
106. Mainz am Rhein: Verlag Philipp von Zabern. p 45-53.
107. Zietkiewicz E, Yotova V, Gehl D, Wambach T, Arrieta I, Batzer M, Cole DE, Hechtman P, Kaplan F, Modiano D, Moisan JP, Michalski R, Labuda D. 2003. Haplotypes in the dystrophin DNA segment point to a mosaic origin of modern human diversity. Am J Hum Genet 73:994-1015.
108. Zilhao J. 2001. Neandertal/modern human interaction in Europe. In: Thacker P, Hays M, editors. Questioning the answers: resolving fundamental problems of the Early Upper Paleolithic. British Archaeological Reports International Series 1005:13-19.
109. Zilhao J. 2006. Neandertals and moderns mixed, and it matters. Evol Anthropol 15:183-195.
110. Zilhao J, d'Errico F, Bordes J-G, Lenoble A, Texier J-G, Rigaud J-P. 2006. Analysis of Aurignacian interstratification at the Chatelperronian-type site and implications for the behavioral modernity of Neandertals. Proc Natl Acad Sci USA 103: 12643-12648.
111. Zilhao J, Trinkaus E, editors. 2002. Portrait of the artist as a child. The Gravettian human skeleton from the Abrigo do Lagar Velho and its archeological context, Vol. 22. Lisbon: Trabalhos de Arqueologia.