The place of the Neanderthals in hominin phylogeny

Journal of Anthropological Archaeology

Volumn 35, Issue 1, 2014, Pages 32-50

  White, Suzanna ^a   Gowlett, John A J ^a   Grove, Matt ^a  

^a UNIVERSITY OF LIVERPOOL   (United Kingdom)

Author keywords

Neanderthals; Species status; Taxonomy

Indexed keywords

EID: 84900329042     PISSN: 02784165     EISSN: 10902686     Source Type: Journal    
DOI: 10.1016/j.jaa.2014.04.004     Document Type: Article

References (192)

1. Abi-Rached L., Jobin M.J., Kulkami S., McWhinnie A., Dalva K., Gragert L., Babrzadeh F., Luo M., Plummer F.A., Kimani J., Carrington M., Middleton D., Rajalingam R., Beksac M., Marsh S.G.E., Maiers M., Guethlein L.A., Tavoularis S., Little A.-M., Green R.E., Norman P.J., Parham P. The shaping of modern human immune systems by multiregional admixture with archaic humans. Science 2011, 334:89-94.
2. Ackermann R.R. Phenotypic traits of primate hybrids: recognizing admixture in the fossil record. Evol. Anthropol. 2010, 19(6):258-270.
3. Ackermann R.R., Bishop J.M. Morphological and molecular evidence reveals recent hybridization between gorilla taxa. Evolution 2009, 64:271-290.
4. Ahern J.C.M. Non-metric variation in recent humans as a model for understanding Neanderthal-early modern human differences: just how "unique" are Neanderthal unique traits?. Neanderthals Revisited: New Approaches and Perspectives 2006, 255-268. Springer, Dordrecht. K. Harvati, T. Harrison (Eds.).
5. Ahern J.C.M., Lee S.-H., Hawks J.D. The late Neandertal supraorbital fossils from Vindija Cave, Croatia: a biased sample?. J. Hum. Evol. 2002, 43:419-432.
6. Alves I., Hanulová A.Š., Foll M., Excoffier L. Genomic data reveal a complex making of humans. PLoS Genet. 2012, 8:e1002837.
7. Arnold M.L., Meyer A. Natural hybridization in primates: One evolutionary mechanism. Zoo. 2006, 109:261-276.
8. Arnold M.L., Meyer A. Natural hybridization in primates: one evolutionary mechanism. Zoology 2006, 109:261-276.
9. Arthreya S. A comparative study of frontal bone morphology among Pleistocene hominin fossil groups. J. Hum. Evol. 2009, 57:786-804.
10. Avise J.C., Wollenberg K. Phylogenetics and the origin of species. Proc. Natl. Acad. Sci. 1997, 94:7748-7755.
11. Bailey S.E. A closer look at Neanderthal postcanine dental morphology: the mandibular dentition. Anat. Rec. 2002, 269:148-156.
12. Bailey S.E., Lynch J.M. Diagnostic differences in mandibular P4 shape between Neandertals and anatomically modern humans. Am. J. Phys. Anthropol. 2005, 126:268-277.
13. Balakrishnan R. Species concepts, species boundaries and species identification: a view from the tropics. Syst. Biol. 2005, 54:689-693.
14. Barton N.H. Evolutionary biology: how did the human species form?. Curr. Biol. 2006, 16:R647-R650.
15. Barton C.M., Riel-Salvatore J. Agents of change: modeling biocultural evolution in upper pleistocene western Eurasia. Adv. Compl. Syst. 2012, 15:1150003-1-1150003-24.
16. Baum D.A., Donoghue M.J. Choosing among alternative "phylogenetic" species concepts. Syst. Bot. 1995, 20:560-573.
17. Bayle P., Macchiarelli R., Trinkaus E., Duarte C., Mazurier A., Zilhão J. Dental maturational sequence and dental tissue proportions in the early Upper Paleolithic child from Abrigo do Lagar Velho. Portugal. Proc. Natl. Acad. Sci. 2010, 107:1338-1342.
18. Beauval C., Maureille B., Lacrampe-Cuyaubere F., Serre D., Peressinotto D., Bordes J.G., Cochard D., Couchoud I., Dubrasquet D., Laroulandie V., Lenoble A., Mallye J.-B., Pasty S., Primault J., Rohland N., Pääbo S., Trinkaus E. A late Neandertal femur from Les Rochers-de-Villeneuve. France. Proc. Natl. Acad. Sci. 2005, 102:7085-7090.
19. Beerli P., Edwards S.V. When did Neanderthals and modern humans diverge?. Evol. Anthropol. 2002, 11:60-63.
20. Belle E.M.S., Benazzo A., Ghirotto S., Colonna V., Barbujani G. Comparing models on the genealogical relationships among Neandertal, Cro-Magnoid and modern Europeans by serial coalescent simulations. Heredity 2009, 102:218-225.
21. Boggs C.L. Species and speciation. International Encyclopedia of the Social and Behavioral Sciences 2001, 14855-14861. Elsevier, Oxford. N. Smelser, P. Baltes (Eds.).
22. Bower B. Hybrid-driven evolution. Science 2006, 169:308-309.
23. Bräuer G., Broeg H., Stringer C.B. Earliest Upper Paleolithic crania from Mladeč, Czech Republic, and the question of Neanderthal-modern continuity: metrical evidence from the fronto-facial region. Neanderthals Revisited: New Approaches and Perspectives 2006, 269-290. Springer, Dordrecht. K. Harvati, T. Harrison (Eds.).
24. Briggs A.W., Good J.M., Green R.E., Krause J., Maricic T., Stenzel U., Lalueza-Fox C., Rudan P., Brajkovic D., Kucan Z., Gusic I., Schmitz R., Doronichev V.B., Golovanova L.V., de la Rasilla M., Fortea J., Rosas A., Pääbo S. Targeted retrieval and analysis of five Neandertal mtDNA genomes. Science 2009, 325:318-321.
25. Burrell A.S., Jolly C.J., Tosi A.J., Disotell T.R. Mitochondrial evidence for the hybrid origin of the kipunji, Rungwecebus kipunji (Primates: Papionini). Mol. Phylogenet. Evol. 2009, 51:340-348.
26. Caramelli D., Lalueza-Fox C., Vernesi C., Larl M., Casoli A., Mallegni F., Chiarelli B., Dupanloup I., Bertranpetit J., Barbujani G., Bertorelle G. Evidence for a genetic discontinuity between Neandertals and 24,000-year-old anatomically modern Europeans. Proc. Natl. Acad. Sci. 2003, 100:6593-6597.
27. Caramelli D., Lalueza-Fox C., Condemi S., Longo L., Milani L., Manfredini A., de Saint Pierre M., Adoni F., Lari M., Giunti P., Ricci S., Casoli A., Calafell F., Mallegni F., Bertranpetit J., Stanyon R., Bertorelle G., Barbujani G. A highly divergent mtDNA sequence in a Neandertal individual from Italy. Curr. Biol. 2006, 16:R630-R632.
28. Churchill S.E., Smith F.H. A modern human humerus from the early Aurignacian of Vogelherdhöhle (Stetten, Germany). Am. J. Phys. Anthropol. 2000, 112:251-273.
29. Cortés-Ortiz L., Duda T.F., Canales-Espinosa D., García-Orduña F., Rodríguez-Luna E., Bermingham E. Hybridization in large-bodied new world primates. Genetics 2007, 176:2421-2425.
30. Cracraft J. Species concepts and species analysis. Curr. Ornithol. 1983, 1:159-187.
31. Currat M., Excoffier L. Modern humans did not admix with Neanderthals during their range expansion into Europe. PLoS Biol. 2004, 2:2264-2274.
32. Currat M., Excoffier L. Strong reproductive isolation between humans and Neanderthals inferred from observed patters of introgression. Proc. Natl. Acad. Sci. 2011, 108:15129-15134.
33. Dabney J., Knapp M., Glocke I., Gansauge M.-T., Weihmann A., Nickel B., Valdiosera C., Garcia N., Pääbo S., Arsuaga J.-L., Meyer M. Proc. Natl. Acad. Sci. 2013, 110:15758-15763.
34. Dalén L., Orlando L., Shapiro B., Durling M.B., Quam R., Gilbert M.T.P., Fernández-Lomana J.C.D., Willerslev E., Arsuaga J.L., Götherstrom A. Partial genetic turnover in neandertals: continuity in the east and population replacement in the west. Mol. Biol. Evol. 2012, 29:1893-1897.
35. de Queiroz K. The general lineage concept of species, species criteria, and the process of speciation: a conceptual unification and terminological recommendations. Endless Forms: Species and Speciation 1998, 57-75. Oxford University Press, New York. D.J. Howard, S.H. Berloche (Eds.).
36. de Queiroz K. Different species problems and their resolution. BioEssays 2005, 27:1263-1269.
37. de Vos J. Receiving an ancestor into the phylogenetic tree: Neanderthal man, Homo erectus and Homo floresiensis: L'histoire se repete. J. Hist. Biol. 2009, 42:361-379.
38. Encyclopedia of Human Evolution and Prehistory 2000, Garland Publishing Inc., New York. second ed. E. Delson, I. Tattersall, J.A. Van Couvering (Eds.).
39. Di Vincenzo F., Churchill S.E., Manzi G. The Vindija Neanderthal scapular glenoid fossa: comparative shape analysis suggests evo-devo changes among Neanderthals. J. Hum. Evol. 2012, 62:274-285.
40. Dobzhansky T. A critique of the species concept in biology. Philos. Sci. 1935, 2:344-355.
41. Dobzhansky T. On species and races of living and fossil man. Am. J. Phys. Anthropol. 1944, 2:251-265.
42. Duarte C., Mauricio J., Pettitt P.B., Souto P., Trinkaus E. The early Upper Paleolithic human skeleton from the Abrigo do Lagar Velho (Portugal) and modern human emergence in Iberia. Proc. Natl. Acad. Sci. 1999, 96:7604-7609.
43. Durand E.Y., Patterson N., Reich D., Slatkin M. Testing for ancient admixture between closely related populations. Mol. Biol. Evol. 2011, 28:2239-2252.
44. Endicott P., Ho S.Y.W. A Bayesian evaluation of human mitochondrial substitution rates. Am. J. Hum. Genet. 2008, 82:895-902.
45. Endicott P., Ho S.Y.W., Stringer C. Using genetic evidence to evaluate four palaeoanthropological hypotheses for the timing of Neanderthal and modern human origins. J. Hum. Evol. 2010, 59:87-95.
46. Eriksson A., Manica A. Effect of ancient population structure on the degree of polymorphism shared between modern human populations and ancient hominins. Proc. Natl. Acad. Sci. 2012, 35:13956-13960.
47. Ferguson J.W.H. On the use of genetic divergence for identifying species. Biol. J. Linn. Soc. Lond. 2002, 75:509-516.
48. Finlayson C., Pacheco F.G., Rodríguez-Vidal J., Fa D.A., Gutierrez López J.M., Santiago Pérez A., Finlayson G., Allue E., Baena Preysler J., Cáceres I., Carrión J.S., Fernández Jalvo Y., Gleed-Owen C.P., Jimenez Espejo F.J., López P., López Sáez J.A., Riquelme Cantal J.A., Sánchez Marco A., Guzman F.G., Brown K., Fuentes N., Valarino C.A., Villalpando A., Stringer C.B., Martinez Ruiz F., Sakamoto T. Late survival of Neanderthals at the southernmost extreme of Europe. Nature 2006, 443:850-853.
49. Fitzpatrick B.M. Rates of evolution of hybrid inviability in birds and mammals. Evolution 2004, 58:1865-1870.
50. Foley R.A. How many species of hominid should there be?. J. Hum. Evol. 1991, 20:413-427.
51. Forhan G., Martiel J.-L., Blum M.G.B. A deterministic model of admixture and genetic introgression: the case of Neanderthal and Cro-Magnon. Math. Biosci. 2008, 216:71-76.
52. Frayer D.W., Jelínek J., Oliva M., Wolpoff M.H. Aurignacian male crania, jaws and teeth from the Mladeč Caves, Moravia, Czech Republic. Early Modern Humans at the Moravian Gate: The Mladeč Caves and Their Remains 2006, 185-272. Springer, Vienna. M. Tescheler-Nicola (Ed.).
53. Fu Q., Mittnik A., Johnson P.L.F., Bos K., Lari M., Bollongino R., Sun C., Giemsch L., Schmitz R., Burger J., Ronchitelli A.M., Martini F., Cremonesi R.G., Svoboda S., Reich D., Pääbo S., Krause J. A revised timescale for human evolution based on ancient mitochondrial genomes. Curr. Biol. 2013, 23:553-559.
54. Garrigan D., Kingan S.B. Archaic human admixture: a view from the genome. Curr. Anthropol. 2007, 48:895-902.
55. Garrigan D., Mobasher Z., Kingan S.B., Wilder J.A., Hammer M.F. Deep haplotype divergence and long-range linkage disequilibrium at Xp21.1 provide evidence that humans descend from a structured ancestral population. Genetics 2005, 170:1849-1856.
56. Ghirotto S., Tassi F., Benazzo A., Barbujani G. No evidence of Neandertal admixture in the mitochondrial genomes of early European modern humans and contemporary Europeans. Am. J. Phys. Anthropol. 2011, 146:242-252.
57. Ghiselin M.T. A radical solution to the species problem. Syst. Zool. 1974, 23:536-544.
58. Giles R.E., Blanc H., Cann H.M., Wallace D.C. Maternal inheritance of human mitochondrial DNA. Proc. Natl. Acad. Sci. 1980, 77:6715-6719.
59. Gould S.J. Why we should not name human races - a biological view. Ever Since Darwin: Reflections in Natural History 1991, 231-237. Penguin Books, St. Ives. S.J. Gould (Ed.).
60. Grant B.R., Grant P.R. Hybridization and speciation in Darwin's finches. Endless Forms: Species and Speciation 1998, 404-422. Oxford University Press, New York. D.J. Howard, S.H. Berloche (Eds.).
61. Green R.E., Krause J., Ptak S.E., Briggs A.W., Ronan M.T., Simons J.F., Du L., Egholm M., Rothberg J.M., Paunovic M., Pääbo S. Analysis of one million base pairs of Neanderthal DNA. Nature 2006, 444:330-336.
62. Green R.E., Malaspinas A.-S., Krause J., Briggs A.W., Johnson P.L.F., Uhler C., Meyer M., Good J.M., Maricic T., Stenzel U., Prufer K., Siebauer M., Burbano H.A., Ronan M., Rothberg J.M., Egholm M., Rudan P., Brajkovic D., Kucan Z., Gusic I., Wikstrom M., Laakkonen L., Kelso J., Slatkin M., Pääbo S. A complete Neandertal mitochondrial genome sequence determined by high-throughput sequencing. Cell 2008, 134:416-426.
63. Green R.E., Briggs A.W., Krause J., Prufer K., Lachmann M., Pääbo S. The Neandertal genome and ancient DNA authenticity. EMBO J. 2009, 28:2494-2502.
64. Green R.E., Krause J., Briggs A.W., Maricic T., Stenzel U., Kircher M., Patterson N., Li H., Zhai W., Fritz M.H.-Y., Hansen N.F., Durand E.Y., Malaspinas A.-S., Jensen J.D., Marques-Bonet T., Alkan C., Prüfer K., Meyer M., Burbano H.A., Good J.M., Schultz R., Aximu-Petri A., Butthof A., Höffner B., Siegemund M., Weihmann A., Nusbaum C., Lander E.S., Russ C., Novod N., Affourtit J., Egholm M., Verna C., Rudan P., Brajkovic D., Kucan Ž., Gušic I., Doronichev V.B., Golovanova L.V., Lalueza-Fox C., de la Rasilla M., Fortea J., Rosas A., Mullikin J.C., Slatkin M., Nielsen R., Kelso J., Lachmann M., Reich D., Pääbo S. A draft sequence of the Neandertal genome. Science 2010, 328:710-722.
65. Hammer M.F., Woerner A., Mendez F.L., Watkins J.C., Wall J.D. Genetic evidence for archaic admixture in Africa. Proc. Natl. Acad. Sci. 2011, 108:15123-15128.
66. Harrison R.G. Linking evolutionary patter and process: the relevance of species concepts for the study of speciation. Endless Forms: Species and Speciation 1998, 19-31. Oxford University Press, New York. D.J. Howard, S.H. Berloche (Eds.).
67. Harvati K. The Neanderthal taxonomic position: models of intra- and inter-specific craniofacial variation. J. Hum. Evol. 2003, 44:107-132.
68. Harvati K., Frost S.R., McNulty K.P. Neanderthal taxonomy reconsidered: implications of 3D primate models of intra- and interspecific differences. Proc. Natl. Acad. Sci. 2004, 101:1147-1152.
69. Harvati K., Gunz P., Grigorescu D. Cioclovina (Romania): affinities of an early modern European. J. Hum. Evol. 2007, 53:732-746.
70. Hawks J. Selection on mitochondrial DNA and the Neanderthal problem. Neanderthals Revisited: New Approaches and Perspectives 2006, 221-238. Springer, Dordrecht. K. Harvati, T. Harrison (Eds.).
71. Hawks J.D., Wolpoff M.H. The accretion model of Neandertal evolution. The Human Evolution Source Book 2006, 458-470. Pearson Prentice Hall, Upper Saddle River, NJ. R.L. Ciochon, J.G. Fleagle (Eds.).
72. Hawks J., Cochran G., Harpending H.C., Lahn B.T. A genetic legacy from archaic Homo. Trends Genet. 2008, 24:19-23.
73. Hennig W. Phylogenetic Systematics 1966, University of Illinois Press, Urbana.
74. Hershkovitz I., Smith P., Sarig R., Quam R., Rodríguez L., García R., Arsuaga J.L., Barkai R., Gopher A. Middle Pleistocene dental remains from Qesem Cave (Israel). Am. J. Phys. Anthropol. 2011, 144:575-592.
75. Hey J. On the failure of modern species concepts. Trends Ecol. Evol. 2006, 21:447-450.
76. Hey J., Waples R.S., Arnold M.L., Butlin R.K., Harrison R.G. Understanding and confronting species uncertainty in biology and conservation. Trends Ecol. Evol. 2003, 18:597-603.
77. 1 Upper Paleolithic Neandertals. Proc. Natl. Acad. Sci. 2006, 103:553-557.
78. Holliday T.W. Species concepts, reticulation, and human evolution. Curr. Anthropol. 2003, 653-660.
79. Holliday T.W. Neanderthals and modern humans: an example of a mammalian syngameon?. Neanderthals Revisited: New Approaches and Perspectives 2006, 281-298. Springer, Dordrecht. K. Harvati, T. Harrison (Eds.).
80. Holmes E.C., Pesole G., Saccone C. Stochastic models of molecular evolution and the estimation of phylogeny and rates of nucleotide substitution in the hominoid primates. J. Hum. Evol. 1989, 18:775-794.
81. Huxley T.H. Evidence as to Man's Place in Nature 1863, Williams and Norgate, London.
82. Jensen-Seaman M.I., Hooper-Boyd K.A. Molecular clocks: Determining the age of the human-chimp divergence. Encyclopedia of Life Sciences (ELS) 2008, John Wiley & Sons, Ltd.,
83. Jolly C.J. Species, subspecies, and baboon systematic. Species, Species Concepts, and Primate Evolution 1993, 67-101. Plenum Press, New York. W.H. Kimbel, L.B. Martin (Eds.).
84. Jolly C.J. A proper study for mankind: analogies from the papionin monkeys and their implications for human evolution. Yearbook Phys. Anthropol. 2001, 44:177-204.
85. Kaufman D. Comparisons and the case for interaction among Neanderthals and early modern humans in the levant. Oxf. J. Archaeol. 2001, 20:219-240.
86. Keita S.O.Y., Kittles R.A., Royal C.D.M., Bonney G.E., Furbert-Harris P., Dunston G.M., Rotimi C.N. Conceptualising human variation. Nat. Genet. 2004, 36:S17-S20.
87. Kimbel W.H. Species, species concepts and hominid evolution. J. Hum. Evol. 1991, 20:355-371.
88. Kimbel W.H., Rak Y. The importance of species taxa in paleoanthropology and an argument for the phylogenetic species concept. Species, Species Concepts and Primate Evolution 1993, 461-484. Plenum Press, New York. W.H. Kimbel, L.B. Martin (Eds.).
89. King W. The reputed fossil man of the Neanderthal. Q. Rev. Sci. 1984, 1:88-97.
90. Klein R.G. Whither the Neanderthals?. Science 2003, 299:1525-1527.
91. Klein R.G. Out of Africa and the evolution of human behavior. Evol. Anthropol. 2008, 17:267-281.
92. Knight A. The phylogenetic relationship of Neandertal and modern human mitochondrial DNAs based on informative nucleotide sites. J. Hum. Evol. 2003, 44:627-632.
93. Krause J., Fu Q., Good J.M., Viola B., Shunkov M.V., Derevianko A.P., Pääbo S. The complete mitochondrial DNA genome of an unknown hominin from southern Siberia. Nature 2010, 464:894-897.
94. Krings M., Geisert H., Schmitz R.W., Krainitzki H., Pääbo S. DNA sequence of the mitochondrial hypervariable region II from the Neandertal type specimen. Proc. Natl. Acad. Sci. 1999, 96:5581-5585.
95. Krings M., Capelli C., Tschentscher F., Geisert H., Meyer S., von Haeseler A., Grossschmidt K., Possnert G., Paunovic M., Pääbo S. A view of Neandertal genetic diversity. Nat. Genet. 2000, 26:144-146.
96. Krings M., Stone A., Schmitz R.W., Krainitzki H., Stoneking M., Pääbo S. Neandertal DNA sequences and the origin of modern humans. The Human Evolution Source Book 2006, 471-480. Pearson Prentice Hall, Upper Saddle River, NJ. R.L. Ciochon, J.G. Fleagle (Eds.).
97. Kupczik K., Hublin J.-J. Mandibular molar root morphology in Neanderthals and Late Pleistocene and recent Homo sapiens. J. Hum. Evol. 2010, 59:525-541.
98. Lalueza-Fox C., Sampietro M.L., Caramelli D., Puder Y., Lari M., Calafell F., Martinez-Maza C., Bastir M., Fortea J., de la Rasilla M., Nertranpetot J., Rosas A. Neandertal evolutionary genetics; mitochondrial DNA data from the Iberian Peninsula. Mol. Biol. Evol. 2005, 22:1077-1081.
99. Lalueza-Fox C., Krause J., Caramelli D., Catalano G., Milani L., Sampietro L., Calafell F., Martínez-Maza C., Bastir M., García-Tabernero A., De la Rasilla M., Fortea J., Pääbo S., Bertranpetit J., Rosas A. Mitochondrial DNA of an Iberian Neandertal suggests a population affinity with other European Neandertals. Curr. Biol. 2006, 16:R629-R630.
100. Lalueza-Fox C., Rosas A., de la Rasilla M. Issues from Neandertal genomics: diversity, adaptation and hybridisation revised from the El Sidrón case study. Quatern. Int. 2012, 247:10-14.
101. Langergraber K.E., Prüfer K., Rowney C., Boesch C., Crockford C., Fawcett K., Inoue E., Inoue-Muruyama M., Mitani J.C., Muller M.N., Robbins M.M., Schubery G., Stoinski T.S., Viola B., Watts D., Wittig R.M., Wrangham R.C., Zuberbühler K., Pääbo S., Vigilant L. Generation times in wild chimpanzees and gorillas suggest earlier divergence times in great ape and human evolution. Proc. Natl. Acad. Sci. 2013, 109(39):15716-15721.
102. Lindahl T. Facts and artifacts of ancient DNA. Cell 1997, 90:1-3.
103. Linnaeus C. A General System of Nature: Through the Three Kingdoms of Animals, Vegetables and Minerals 1802, Lackington, Allen and Co., London.
104. Livingstone F.B., Dobzhansky T. On the non-existence of human races. Curr. Anthropol. 1962, 3:279-281.
105. Lohse, K., Frantz, L.A.F., 2013. Maximum Likelihood Evidence for Neandertal Admixture in Eurasian Populations from Three Genomes. arXiv preprint arXiv: 1307.8263.
106. Long J.C., Kittles R.A. Human genetic diversity and the nonexistence of biological races. Hum. Biol. 2003, 75:449-471.
107. Loogväli E.-L., Kivisild T., Margus T., Villems R. Explaining the imperfection of the molecular clock of hominid mitochondria. PLoS ONE 2009, 4:e8260.
108. Lotsy J.P. Species or Linneon?. Genetica 1925, 7(5-6):475-486.
109. Lowery R.K., Uribe G., Jimenez E.B., Weiss M.A., Herrera K.J., Regueiro M., Herrera R.J. Neanderthal and Denisova genetic affinities with contemporary humans: introgression versus common ancestral polymorphisms. Gene 2013, 530:83-94.
110. Mallet J. A species definition for the modern synthesis. Tree 1995, 10:294-299.
111. Mallet J. Concepts of species. Encyclopedia of Biodiversity 2007, 427-440. Academic Press, New York. S.A. Levin (Ed.).
112. Masters J.C. Primates and paradigms: problems with the identification of genetic species. Species, Species Concepts, and Primate Evolution 1993, 43-64. Plenum Press, New York. W.H. Kimbel, L.B. Martin (Eds.).
113. Max Planck Institute for Evolutionary Anthropology, 2013. A High-quality Neandertal Genome Sequence. (accessed 03.10.13). http://www.eva.mpg.de/neandertal/index.html.
114. Mayr E. Systematics and the Origin of Species: From the Viewpoint of a Zoologist 1964, Dover Publications Inc., UK.
115. Mayr E. What is a species, and what is not?. Philos. Sci. 1996, 63:262-277.
116. Mayr E., Ashlock P.D. Principles of Systematic Zoology 1991, McGraw-Hill, New York. second ed.
117. Mayr E., Gorton Linsley E., Usinger R.L. Methods and Principles of Systematic Zoology 1953, McGraw-Hill Book Company Inc., New York.
118. Mendez F.L., Watkins J.C., Hammer M.E. A haplotype at STAT2 introgressed from Neanderthals and serves as a candidate of positive selection in Papua New Guinea. Am. J. Hum. Genet. 2012, 91:265-274.
119. Meyer M., Kircher M., Gansuage M.-T., Li H., Racimo F., Mallick S., Schraiber J.G., Jay F., Prüfer K., de Filippo C., Sudmant P.H., Alkan C., Fu Q., Do R., Rohland N., Tandon A., Siebauer M., Green R.E., Bryc K., Briggs A.W., Stenzel U., Dabney J., Shendure J., Kitzman J., Hammer M.F., Shunkov M.V., Derevianko A., Patterson N., Andrés A.M., Eichler E.E., Slatkin M., Reich D., Kelso J., Pääbo S. A high-coverage sequence from an archaic denisovan individual. Science 2012, 338:222-226.
120. Miller W. The structure of species, outcomes of speciation and the 'species problem': ideas for paleobiology. Palaeogeogr. Palaeoclimatol. Palaeoecol. 2001, 176:1-10.
121. Minugh-Purvis N., Viola T.B., Teschler-Nicola M. The Mladeč 3 infant. Early Modern Humans at the Moravian Gate: The Mladeč Caves and Their Remains 2006, 357-383. Springer, Vienna. M. Tescheler-Nicola (Ed.).
122. Neves, A.G.M., Serva, M., 2011. Extremely Rare Interbreeding Events can Explain Neanderthal DNA in Modern Humans. arXiv:1103.4621v2.
123. Nixon K.C., Wheeler Q.D. An amplification of the phylogenetic species concept. Cladistics 1990, 6:211-223.
124. Noonan J.P., Coop G., Kudaravalli S., Smith D., Krause J., Alessi J., Chen F., Platt D., Pääbo S., Pritchard J.K., Rubin E.M. Sequencing and analysis of Neanderthal genomic DNA. Science 2006, 314:1113-1118.
125. Nordberg M. On the probability of Neanderthal ancestry. Am. J. Hum. Genet. 1998, 63:1237-1240.
126. Orlando L., Darlu P., Toussaint M., Bonjean D., Otte M., Hanni C. Revisiting Neandertal diversity with a 100,000 year old mtDNA sequence. Curr. Biol. 2006, 16:R400-R402.
127. O'Rourke D.H., Hayes M.G., Carlyle S.W. Ancient DNA studies in physical anthropology. Annu. Rev. Anthropol. 2000, 29:217-242.
128. Ovchinnikov I.V., Gotherstroms A., Romanova G.P., Kharitonov V.M., Liden K., Goodwin W. Molecular analysis of Neanderthal DNA from the northern Caucasus. Nature 2000, 404:490-493.
129. Pääbo S. The mosaic that is our genome. Nature 2003, 421:409-412.
130. Paterson H.E.H. The continuing search for the unknown and unknowable: a critique of contemporary ideas on speciation. S. Afr. J. Sci. 1981, 77:113-119.
131. Patterson N., Richter D.J., Gnerre S., Lander E.S., Reich D. Genetic evidence for complex speciation of humans and chimpanzees. Nature 2006, 44:1103-1108.
132. Pettitt P. The Palaeolithic Origins of Human Burial 2011, Routledge, London.
133. Prufer K., Racimo F., Patterson N., Jay F., Sankararaman S., Sawyer S., Heinze A., Renaud G., Sudmant P.H., de Filippo C., Li H., Mallick S., Dannemann M., Fu Q., Kircher M., Kuhlwilm M., Lachmann M., Meyer M., Ongyerth M., Siebauer M., Theunert C., Tandon A., Moorjani P., Pickrell J., Mullikin J.C., Vohr S.H., Green R.E., Hellmann I., Johnson P.L.F., Blanche H., Cann H., Kitzman J.O., Shendure J., Eichler E.E., Lein E.S., Bakken T.E., Golovanova L.V., Doronichev V.B., Shunkov M.V., Derevianko A.P., Viola B., Slatkin M., Reich D., Kelso J., Paabo S. The complete genome sequence of a Neanderthal from the Altai Mountains. Nature 2014, 505:43-49.
134. Pulquério M.J.F., Nichols R.A. Dates from the molecular clock: how wrong can we be?. Trends Ecol. Evol. 2007, 22:180-184.
135. Quicke D.L.J. Principles and Techniques of Contemporary Taxonomy 1993, Blackie Academic & Professional, Tertiary Level Biology, Glasgow.
136. Quintyn C. The naming of new species in hominin evolution: a radical proposal - a temporary cessation against new names. Homo 2009, 60:307-341.
137. Rak Y. On the differences between two pelvises of mousterian context from the Qafzeh and Kebara Caves, Israel. Am. J. Phys. Anthropol. 1990, 81:323-332.
138. Rak Y. Morphological variation in Homo neanderthalensis and Homo sapiens in the Levant: a biogeographic model. Species, Species Concepts, and Primate Evolution 1993, 523-536. Plenum Press, New York. W.H. Kimbel, L.B. Martin (Eds.).
139. Reich D., Green R.E., Kircher M., Krause J., Patterson N., Durand E.Y., Viola B., Briggs A.W., Stenzel U., Johnsonm P.L.F., Maricic T., Good J.M., Marques-Bonet T., Alkan C., Fu Q., Mallick S., Li H., Meyer M., Eichler E.E., Stoneking M., Richards M., Talamo S., Shunkov M.V., Derevianko A.P., Hublin J.-J., Kelos J., Slatkin M., Pääbo S. Genetic history of an archaic hominin group from Denisova Cave in Siberia. Nature 2010, 468:1053-1060.
140. Reydon T.A.C. Why does the species problem still persist?. BioEssays 2004, 26:300-305.
141. Rougier H., Milota Ş., Rodrigo R., Gherase M., Sarcinǎ, Moldovan O., Zilhão J., Constantin S., Franciscus R.G., Zollikofer C.P.E., Ponce de León M., Trinkaus E. Peştera cu Oase 2 and the cranial morphology of early modern Europeans. Proc. Natl. Acad. Sci. 2007, 104:1165-1170.
142. Sánchez-Quinto F., Botigué L.R., Civit S., Arenas C., Ávila-Arcos M., Bustamente C.D., Comas D., Lalueza-Fox C. North African populations carry the signature of admixture with Neandertals. PLoS ONE 2012, 7:e47765.
143. Sankararaman S., Patterson N., Li H., Pääbo S., Reich D. The date of interbreeding between Neandertals and modern humans. PLoS Genet. 2012, 8:e1002947.
144. Santa Luca A.P. A re-examination of presumed Neandertal-like fossils. J. Hum. Evol. 1978, 7:619-636.
145. Schmitz R.W., Serre D., Bonani G., Fein S., Hillgruber F., Krainitzki H., Pääbo S., Smith F.H. The Neandertal type site revisited: interdisciplinary investigations of skeletal remains from the Neander Valley, Germany. Proc. Natl. Acad. Sci. 2002, 99:13342-13347.
146. Scholz M., Bachmann L., Nicholson G.J., Bachmann J., Giddings I., Ruschoff-Thale B., Czarnetzki A., Pusch C.M. Genomic differentiation of neanderthals and anatomically modern man allows a fossil-DNA-based classification of morphologically indistinguishable hominid bones. Am. J. Hum. Gen. 2000, 66:1927-1932.
147. Schwartz J.H., Tattersall I. Significance of some previously unrecognized apomorphies in the nasal region of Homo neanderthalensis. Proc. Natl. Acad. Sci. 1996, 93:10854-10952.
148. Serre D., Pääbo S. The fate of European Neanderthals: results and perspectives from ancient DNA analyses. Neanderthals Revisited: New Approaches and Perspectives 2006, 211-220. Springer, Dordrecht. K. Harvati, T. Harrison (Eds.).
149. Serre D., Langaney A., Chech M., Teschler-Nicola M., Paunovic M., Mennecier P., Hofrieter M., Possnert G., Pääbo S. No evidence of Neandertal mtDNA contribution to early modern humans. PLoS Biol. 2004, 2:313-317.
150. Shaw K.L. Species and the diversity of natural groups. Endless Forms: Species and Speciation 1998, 44-56. Oxford University Press, New York. D.J. Howard, S.H. Berloche (Eds.).
151. Simpson G.G. Criteria for genera, species, and subspecies in zoology and paleozoology. Ann. N. Y. Acad. Sci. 1943, 44:145-178.
152. Simpson G.G. The species concept. Evolution 1951, 5:285-298.
153. Simpson G.G. Principles of Animal Taxonomy 1961, Columbia University Press, New York.
154. Sites J.W., Marshall J.C. Operational criteria for delimiting species. Annu. Rev. Ecol. Evol. Syst. 2004, 35:199-227.
155. Smith A.B. Systematics and the Fossil Record: Documenting Evolutionary Patterns 1994, Blackwell Scientific, Oxford.
156. Soficaru A., Dobos A., Trinkaus E. Early modern humans from the Peştera Muierii, Baia de Fier, Romania. Proc. Natl. Acad. Sci. 2006, 103:17196-17201.
157. Soficaru A., Petrea C., Doboş A., Trinkaus E. The human cranium from the Peştera Cioclovina Uscatǎ, Romania: context, age, taphonomy, morphology and paleopathology. Curr. Anthropol. 2007, 48:611-619.
158. Spoor F., Hublin J.-J., Braun M., Zonneveld F. The bony labyrinth of Neanderthals. J. Hum. Evol. 2003, 44:141-165.
159. Stringer C. The Origins of Our Species 2012, Penguin Group, England.
160. Stringer C.B., Andrews P. Genetic and fossil evidence for the origin of modern humans. Science 1988, 239:1263-1268.
161. Tattersall I. Species recognition in human paleontology. J. Hum. Evol. 1986, 15:165-175.
162. Tattersall I. Species concepts and species identification in human evolution. J. Hum. Evol. 1992, 22:341-349.
163. Tattersall I. Speciation and morphological differentiation in the genus Lemur. Species, Species Concepts, and Primate Evolution 1993, 163-176. Wiley, New York. W. Kimbel, L. Martin (Eds.).
164. Tattersall I. Neanderthals, Homo sapiens, and the question of species in paleoanthropology. J. Anthropol. Sci. 2007, 85:139-146.
165. Tattersall I., Schwartz J.H. Morphology, paleoanthropology, and Neanderthals. Anatom. Rec. 1998, 253:113-117.
166. Tattersall I., Schwartz J.H. Hominids and hybrids: the place of Neanderthals in human evolution. Proc. Natl. Acad. Sci. 1999, 96:7117-7119.
167. Tattersall I., Schwartz J.H. The distinctiveness and systematic context of Homo neanderthalensis. Neanderthals Revisited: New Approaches and Perspectives 2006, 9-22. Springer, Dordrecht. K. Harvati, T. Harrison (Eds.).
168. Templeton A.R. The meaning of species and speciation: A genetic perspective. Speciation and its Consequences 1989, 3-27. Sinauer, Sunderland, Massachusetts. D. Otte, J.A. Endler (Eds.).
169. Templeton A.R. Species and speciation: geography, population structure, ecology, and gene trees. Endless Forms: Species and Speciation 1998, 32-43. Oxford University Press, New York. D.J. Howard, S.H. Berloche (Eds.).
170. Templeton A.R. Human races: A genetic and evolutionary perspective. Am. Anthropol. 1999, 100:632-650.
171. Templeton A.R. Haplotype trees and modern human origins. Yearbook Phys. Anthropol. 2005, 48:33-59.
172. Tetushkin E.Y. Genetics and the origin of human "races". Russ. J. Genet. 2001, 37:853-867.
173. Trinkaus E. Early modern humans. Annu. Rev. Anthropol. 2005, 34:207-230.
174. Trinkaus E. European early modern humans and the fate of the Neanderthals. Proc. Natl. Acad. Sci. 2007, 104:7367-7372.
175. Trinkaus E., Duarte C. The hybrid child from Portugal. Sci. Am. 2000, 282:102-103.
176. Trinkaus E., Moldovan O., Milota Ş., Bîlgǎr A., Sarcina L., Athreya S., Bailey S.E., Rodrigo R., Mircea G., Higham T., Ramsey C.B., van der Plicht J. An early modern human from the Peştera cu Oase, Romania. Proc. Natl. Acad. Sci. 2003, 100:11231-11236.
177. Valdiosera C., García N., Dalén L., Kahlke R.-D., Lidén K., Angerbjörn A., Arsuaga J.L., Götherström A. Typing single polymorphic nucleotides in mitochondrial DNA as a way to access Middle Pleistocene DNA. Biol. Lett. 2006, 2:601-603.
178. Van Valen L. Ecological species, multispecies, and oaks. Taxon 1976, 25:233-239.
179. Vernot B., Akey J.M. Resurrecting surviving neandertal lineages from modern human genomes. Science 2014, 343:1017-1021.
180. Wakeley J. Complex speciation of humans and chimpanzees. Nature 2008, 452:E3-E4.
181. Wall J.D. Detecting ancient admixture in humans using sequence polymorphism data. Genetics 2000, 154:1271-1279.
182. Wall J.D., Lohmueller K.E., Plagnol V. Detecting ancient admixture and estimating demographic parameters in multiple human populations. Mol. Biol. Evol. 2009, 26:1823-1827.
183. Wall J.D., Yang M.A., Jay F., Kim S.K., Durand E.Y., Stevison L.S., Gignoux C., Woerner A., Hammer M.F., Slatkin M. Higher levels of Neanderthal ancestry in East Asians than in Europeans. Genetics 2013, 194:199-209.
184. Webster M.T. Patterns of autosomal divergence between the human and chimpanzee genomes support an allopatric model of speciation. Gene 2009, 443:70-75.
185. Wiens J.J., Servedio M.R. Species delimitation in systematics: inferring diagnostic differences between species. Proc. R. Soc. Lond. B 2000, 267:631-636.
186. Wild E.M., Teschler-Nicola M., Kutschera W., Steier P., Trinkaus E., Wanek W. Direct dating of Early Upper Palaeolithic human remains from Mladeč. Nature 2005, 435:332-335.
187. Wiley E.O. The evolutionary species concept reconsidered. Syst. Zool. 1978, 27:17-26.
188. Wolpoff M.H., Frayer D.W., Jelínek J. Aurignacian female crania and teeth from the Mladeč caves, Moravia, Czech Republic. Early Modern Humans at the Moravian Gate: The Mladeč Caves and Their Remains 2006, 273-340. Springer, Vienna. M. Tescheler-Nicola (Ed.).
189. Wood B., Lonergan N. The hominin fossil record: taxa, grades and clades. J. Anat. 2008, 212:354-376.
190. Wood B.A., Richmond B.G. Human evolution taxonomy and paleobiology. J. Anat. 2000, 197:19-60.
191. Yang M.A., Malaspinas A.-S., Durand E.Y., Slatkin M. Ancient structure in Africa unlikely to explain Neanderthal and Non-African genetic similarity. Mol. Biol. Evol. 2012, 29:2987-2995.
192. Yokley T.R., Churchill S.E. Archaic and modern human distal humeral morphology. J. Hum. Evol. 2006, 51:603-616.