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Chimpanzee–human last common ancestor

January 09, 2024

The chimpanzee–human last common ancestor (CHLCA) is the last common ancestor shared by the extant Homo (human) and Pan (chimpanzee and bonobo) genera of Hominini. Estimates of the divergence date vary widely from thirteen to five million years ago.

In human genetic studies, the CHLCA is useful as an anchor point for calculating single-nucleotide polymorphism (SNP) rates in human populations where chimpanzees are used as an outgroup, that is, as the extant species most genetically similar to Homo sapiens.

Taxonomy edit

Main article: Hominini
Hominoidea (hominoids, apes)
Hylobatidae (gibbons)
Hominidae (hominids, great apes)
Ponginae
(orangutans)
Homininae
Gorillini
(gorillas)
Hominini
Panina
(chimpanzees)
Hominina (humans)

The taxon tribe Hominini was proposed to separate humans (genus Homo) from chimpanzees (Pan) and gorillas (genus Gorilla) on the notion that the least similar species should be separated from the other two. However, later evidence revealed that Pan and Homo are closer genetically than are Pan and Gorilla; thus, Pan was referred to the tribe Hominini with Homo. Gorilla now became the separated genus and was referred to the new taxon 'tribe Gorillini'.

Mann and Weiss (1996), proposed that the tribe Hominini should encompass Pan and Homo, grouped in separate subtribes.[1] They classified Homo and all bipedal apes in the subtribe Hominina and Pan in the subtribe Panina. (Wood (2010) discussed the different views of this taxonomy.)[2] A "chimpanzee clade" was posited by Wood and Richmond, who referred it to a tribe Panini, which was envisioned from the family Hominidae being composed of a trifurcation of subfamilies.[3]

Richard Wrangham (2001) argued that the CHLCA species was very similar to the common chimpanzee (Pan troglodytes) – so much so that it should be classified as a member of the genus Pan and be given the taxonomic name Pan prior.[4]

All the human-related genera of tribe Hominini that arose after divergence from Pan are members of the subtribe Hominina, including the genera Homo and Australopithecus. This group represents "the human clade" and its members are called "hominins".[5]

Fossil evidence edit

No fossil has yet conclusively been identified as the CHLCA. A possible candidate is Graecopithecus, though this claim is disputed as there is insufficient evidence to support the determination of Graecopithecus as hominin.[6] This would put the CHLCA split in Southeast Europe instead of Africa.[7][8]

Sahelanthropus tchadensis is an extinct hominine with some morphology proposed (and disputed) to be as expected of the CHLCA, and it lived some 7 million years ago – close to the time of the chimpanzee–human divergence. But it is unclear whether it should be classified as a member of the tribe Hominini, that is, a hominin, as an ancestor of Homo and Pan and a potential candidate for the CHLCA species itself, or simply a Miocene ape with some convergent anatomical similarity to many later hominins.

Ardipithecus most likely appeared after the human-chimpanzee split, some 5.5 million years ago, at a time when gene flow may still have been ongoing. It has several shared characteristics with chimpanzees, but due to its fossil incompleteness and the proximity to the human-chimpanzee split, the exact position of Ardipithecus in the fossil record is unclear.[9] It is most likely derived from the chimpanzee lineage and thus not ancestral to humans.[10][11] However, Sarmiento (2010), noting that Ardipithecus does not share any characteristics exclusive to humans and some of its characteristics (those in the wrist and basicranium), suggested that it may have diverged from the common human/African ape stock prior to the human, chimpanzee and gorilla divergence.[12]

The earliest fossils clearly in the human but not the chimpanzee lineage appear between about 4.5 to 4 million years ago, with Australopithecus anamensis.

Few fossil specimens on the "chimpanzee-side" of the split have been found; the first fossil chimpanzee, dating between 545 and 284 kyr (thousand years, radiometric), was discovered in Kenya's East African Rift Valley (McBrearty, 2005).[13] All extinct genera listed in the taxobox[which?] are ancestral to Homo, or are offshoots of such. However, both Orrorin and Sahelanthropus existed around the time of the divergence, and so either one or both may be ancestral to both genera Homo and Pan.

Due to the scarcity of fossil evidence for CHLCA candidates, Mounier (2016) presented a project to create a "virtual fossil" by applying digital "morphometrics" and statistical algorithms to fossils from across the evolutionary history of both Homo and Pan, having previously used this technique to visualize a skull of the last common ancestor of Neanderthal and Homo sapiens.[14][15]

Age estimates edit

An estimate of TCHLCA of 10 to 13 million years was proposed in 1998,[note 1] and a range of 7 to 10 million years ago is assumed by White et al. (2009):

In effect, there is now no a priori reason to presume that human-chimpanzee split times are especially recent, and the fossil evidence is now fully compatible with older chimpanzee–human divergence dates [7 to 10 Ma...

— White et al. (2009), [17]

Some researchers tried to estimate the age of the CHLCA (TCHLCA) using biopolymer structures that differ slightly between closely related animals. Among these researchers, Allan C. Wilson and Vincent Sarich were pioneers in the development of the molecular clock for humans. Working on protein sequences, they eventually (1971) determined that apes were closer to humans than some paleontologists perceived based on the fossil record.[note 2]

This paradigmatic age has stuck with molecular anthropology until the late 1990s. Since the 1990s, the estimate has again been pushed towards more-remote times, because studies have found evidence for a slowing of the molecular clock as apes evolved from a common monkey-like ancestor with monkeys, with both humans and non-human apes evolving from a common ape-like ancestor.[19]

A 2016 study analyzed transitions at CpG sites in genome sequences, which exhibit a more clocklike behavior than other substitutions, arriving at an estimate for human and chimpanzee divergence time of 12.1 million years.[20]

Gene flow edit

A source of confusion in determining the exact age of the Pan–Homo split is evidence of a more complex speciation process than a clean split between the two lineages. Different chromosomes appear to have split at different times, possibly over as much as a 4-million-year period, indicating a long and drawn out speciation process with large-scale gene flow events between the two emerging lineages as recently as 6.3 to 5.4 million years ago, according to Patterson et al. (2006).[21]

Speciation between Pan and Homo occurred over the last 9 million years. Ardipithecus probably branched off of the Pan lineage in the middle Miocene Messinian.[10][11] After the original divergences, there were, according to Patterson (2006), periods of gene flow between population groups and a process of alternating divergence and gene flow that lasted several million years.[21] Some time during the late Miocene or early Pliocene, the earliest members of the human clade completed a final separation from the lineage of Pan – with date estimates ranging from 13 million[16] to as recent as 4 million years ago.[21] The latter date was in particular based on the similarity of the X chromosome in humans and chimpanzees, a conclusion rejected as unwarranted by Wakeley (2008), who suggested alternative explanations, including selection pressure on the X chromosome in the populations ancestral to the CHLCA.[note 3]

Complex speciation and incomplete lineage sorting of genetic sequences seem to also have happened in the split between the human lineage and that of the gorilla, indicating "messy" speciation is the rule rather than the exception in large primates.[23][24] Such a scenario would explain why the divergence age between the Homo and Pan has varied with the chosen method and why a single point has so far been hard to track down.

See also edit

Notes edit

  1. ^ Based on a revision of the divergence of Hominoidea from Cercopithecoidea at more than 50 Mya (previously set at 30 Mya). "Consistent with the marked shift in the dating of the Cercopithecoidea/Hominoidea split, all hominoid divergences receive a much earlier dating. Thus the estimated date of the divergence between Pan (chimpanzee) and Homo is 10–13 MYBP and that between Gorilla and the Pan/Homo linage ≈17 MYBP."[16]
  2. ^ "If man and old world monkeys last shared a common ancestor 30 million years ago, then man and African apes shared a common ancestor 5 million years ago..."[18]
  3. ^ "Patterson et al. suggest that the apparently short divergence time between humans and chimpanzees on the X chromosome is explained by a massive interspecific gene flow event in the ancestry of these two species. However, Patterson et al. do not statistically test their own null model of simple speciation before concluding that speciation was complex, and—even if the null model could be rejected—they do not consider other explanations of a short divergence time on the X chromosome. These include natural selection on the X chromosome in the common ancestor of humans and chimpanzees, changes in the ratio of male-to-female mutation rates over time, and less extreme divergence versions with gene flow. I, therefore, believe that their claim of gene flow is unwarranted."[22]

References edit

  1. ^ Mann, Alan; Mark Weiss (1996). "Hominoid Phylogeny and Taxonomy: a consideration of the molecular and Fossil Evidence in an Historical Perspective". Molecular Phylogenetics and Evolution. 5 (1): 169–181. doi:10.1006/mpev.1996.0011. PMID 8673284.
  2. ^ B. Wood (2010). "Reconstructing human evolution: Achievements, challenges, and opportunities". Proceedings of the National Academy of Sciences. 107 (Suppl 2): 8902–8909. Bibcode:2010PNAS..107.8902W. doi:10.1073/pnas.1001649107. PMC 3024019. PMID 20445105.
  3. ^ Wood and Richmond.; Richmond, BG (2000). "Human evolution: taxonomy and paleobiology". Journal of Anatomy. 197 (Pt 1): 19–60. doi:10.1046/j.1469-7580.2000.19710019.x. PMC 1468107. PMID 10999270.
  4. ^ "Out of the Pan, Into the Fire" in: Frans B. M. De Waal, ed. (2001). Tree of Origin: What Primate Behavior Can Tell Us About Human Social Evolution. Harvard University Press. pp. 124–126. ISBN 9780674010048.
  5. ^ Bradley, B. J. (2006). "Reconstructing Phylogenies and Phenotypes: A Molecular View of Human Evolution". Journal of Anatomy. 212 (4): 337–353. doi:10.1111/j.1469-7580.2007.00840.x. PMC 2409108. PMID 18380860.
  6. ^ Fuss, Jochen; Spassov, Nikolai; Begun, David R.; Böhme, Madelaine (2017). "Potential hominin affinities of Graecopithecus from the Late Miocene of Europe". PLOS ONE. 12 (5): e0177127. Bibcode:2017PLoSO..1277127F. doi:10.1371/journal.pone.0177127. PMC 5439669. PMID 28531170.
  7. ^ . Archived from the original on 5 November 2019. Retrieved 5 November 2019.
  8. ^ Dickinson, Kevin (20 April 2019). "New fossils suggest human ancestors evolved in Europe, not Africa". Big Think. Retrieved 23 October 2022.
  9. ^ Wood, Bernard; Harrison, Terry (2011). "The evolutionary context of the first hominins". Nature. 470 (7334): 347–35. Bibcode:2011Natur.470..347W. doi:10.1038/nature09709. PMID 21331035. S2CID 4428052.
  10. ^ a b Wood, Bernard; Harrison, Terry (2011). "The evolutionary context of the first hominins". Nature. 470 (7334): 347–52. Bibcode:2011Natur.470..347W. doi:10.1038/nature09709. PMID 21331035. S2CID 4428052.
  11. ^ a b Wolpoff, Milford H. (1996). Human Evolution. McGraw-Hill, Incorporated. ISBN 978-0070718333.
  12. ^ Sarmiento, E. E. (2010). "Comment on the Paleobiology and Classification of Ardipithecus ramidus". Science. 328 (5982): 1105, author reply 1105. Bibcode:2010Sci...328.1105S. doi:10.1126/science.1184148. PMID 20508113.
  13. ^ McBrearty, Sally; Nina G. Jablonski (2005). "First fossil chimpanzee". Nature. 437 (7055): 105–108. Bibcode:2005Natur.437..105M. doi:10.1038/nature04008. PMID 16136135. S2CID 4423286.
  14. ^ "'Virtual fossil' reveals last common ancestor of humans and Neanderthals". 18 December 2015. Retrieved 30 June 2019.
  15. ^ Mounier, Aurélien; Mirazón Lahr, Marta (2016). "Virtual ancestor reconstruction: Revealing the ancestor of modern humans and Neandertals". Journal of Human Evolution. 91: 57–72. doi:10.1016/j.jhevol.2015.11.002. PMID 26852813.
  16. ^ a b Arnason U, Gullberg A, Janke A (December 1998). "Molecular timing of primate divergences as estimated by two nonprimate calibration points". Journal of Molecular Evolution. 47 (6): 718–27. Bibcode:1998JMolE..47..718A. doi:10.1007/PL00006431. PMID 9847414. S2CID 22217997.
  17. ^ White TD, Asfaw B, Beyene Y, et al. (October 2009). "Ardipithecus ramidus and the paleobiology of early hominids". Science. 326 (5949): 75–86. Bibcode:2009Sci...326...75W. doi:10.1126/science.1175802. PMID 19810190. S2CID 20189444.
  18. ^ Sarich, Vincent; Wilson, Allan (1967). "Immunological Time Scale for Hominid Evolution". Science. 158 (3805): 1200–1203. Bibcode:1967Sci...158.1200S. doi:10.1126/science.158.3805.1200. PMID 4964406. S2CID 7349579.
  19. ^ Venn, Oliver; Turner, Isaac; Mathieson, Iain; de Groot, Natasja; Bontrop, Ronald; McVean, Gil (June 2014). "Strong male bias drives germline mutation in chimpanzees". Science. 344 (6189): 1272–1275. Bibcode:2014Sci...344.1272V. doi:10.1126/science.344.6189.1272. PMC 4746749. PMID 24926018.
  20. ^ Moorjani, Priya; Amorim, Carlos Eduardo G.; Arndt, Peter F.; Przeworski, Molly (2016). "Variation in the molecular clock of primates". Proceedings of the National Academy of Sciences. 113 (38): 10607–10612. Bibcode:2016PNAS..11310607M. doi:10.1073/pnas.1600374113. ISSN 0027-8424. PMC 5035889. PMID 27601674.
  21. ^ a b c Patterson N, Richter DJ, Gnerre S, Lander ES, Reich D (June 2006). "Genetic evidence for complex speciation of humans and chimpanzees". Nature. 441 (7097): 1103–8. Bibcode:2006Natur.441.1103P. doi:10.1038/nature04789. PMID 16710306. S2CID 2325560.
  22. ^ Wakeley J (2008). "Complex speciation of humans and chimpanzees". Nature. 452 (7184): E3–4. Bibcode:2008Natur.452....3W. doi:10.1038/nature06805. PMID 18337768. S2CID 4367089.
  23. ^ Scally A, Dutheil JY, Hillier LW, et al. (March 2012). "Insights into hominid evolution from the gorilla genome sequence". Nature. 483 (7388): 169–75. Bibcode:2012Natur.483..169S. doi:10.1038/nature10842. PMC 3303130. PMID 22398555.
  24. ^ Van Arsdale, A.P. "Go, go, Gorilla genome". The Pleistocene Scene – A.P. Van Arsdale Blog. Retrieved 16 November 2012.

External links edit

 
Wikispecies has information related to Hominini.
  • Human Timeline (Interactive) – Smithsonian, National Museum of Natural History (August 2016).
  • Locomotion and posture from the common hominoid ancestor to fully modern hominins, with special reference to the last common panin/hominin ancestor - R H Crompton E E Vereecke and S K S Thorpe, Journal of Anatomy, April 2008

January 09, 2024
chimpanzee, human, last, common, ancestor, chimpanzee, human, last, common, ancestor, chlca, last, common, ancestor, shared, extant, homo, human, chimpanzee, bonobo, genera, hominini, estimates, divergence, date, vary, widely, from, thirteen, five, million, ye. The chimpanzee human last common ancestor CHLCA is the last common ancestor shared by the extant Homo human and Pan chimpanzee and bonobo genera of Hominini Estimates of the divergence date vary widely from thirteen to five million years ago In human genetic studies the CHLCA is useful as an anchor point for calculating single nucleotide polymorphism SNP rates in human populations where chimpanzees are used as an outgroup that is as the extant species most genetically similar to Homo sapiens Contents 1 Taxonomy 2 Fossil evidence 3 Age estimates 4 Gene flow 5 See also 6 Notes 7 References 8 External linksTaxonomy editMain article Hominini Hominoidea hominoids apes Hylobatidae gibbons Hominidae hominids great apes Ponginae orangutans Homininae Gorillini gorillas Hominini Panina chimpanzees Hominina humans The taxon tribe Hominini was proposed to separate humans genus Homo from chimpanzees Pan and gorillas genus Gorilla on the notion that the least similar species should be separated from the other two However later evidence revealed that Pan and Homo are closer genetically than are Pan and Gorilla thus Pan was referred to the tribe Hominini with Homo Gorilla now became the separated genus and was referred to the new taxon tribe Gorillini Mann and Weiss 1996 proposed that the tribe Hominini should encompass Pan and Homo grouped in separate subtribes 1 They classified Homo and all bipedal apes in the subtribe Hominina and Pan in the subtribe Panina Wood 2010 discussed the different views of this taxonomy 2 A chimpanzee clade was posited by Wood and Richmond who referred it to a tribe Panini which was envisioned from the family Hominidae being composed of a trifurcation of subfamilies 3 Richard Wrangham 2001 argued that the CHLCA species was very similar to the common chimpanzee Pan troglodytes so much so that it should be classified as a member of the genus Pan and be given the taxonomic name Pan prior 4 All the human related genera of tribe Hominini that arose after divergence from Pan are members of the subtribe Hominina including the genera Homo and Australopithecus This group represents the human clade and its members are called hominins 5 Fossil evidence editNo fossil has yet conclusively been identified as the CHLCA A possible candidate is Graecopithecus though this claim is disputed as there is insufficient evidence to support the determination of Graecopithecus as hominin 6 This would put the CHLCA split in Southeast Europe instead of Africa 7 8 Sahelanthropus tchadensis is an extinct hominine with some morphology proposed and disputed to be as expected of the CHLCA and it lived some 7 million years ago close to the time of the chimpanzee human divergence But it is unclear whether it should be classified as a member of the tribe Hominini that is a hominin as an ancestor of Homo and Pan and a potential candidate for the CHLCA species itself or simply a Miocene ape with some convergent anatomical similarity to many later hominins Ardipithecus most likely appeared after the human chimpanzee split some 5 5 million years ago at a time when gene flow may still have been ongoing It has several shared characteristics with chimpanzees but due to its fossil incompleteness and the proximity to the human chimpanzee split the exact position of Ardipithecus in the fossil record is unclear 9 It is most likely derived from the chimpanzee lineage and thus not ancestral to humans 10 11 However Sarmiento 2010 noting that Ardipithecus does not share any characteristics exclusive to humans and some of its characteristics those in the wrist and basicranium suggested that it may have diverged from the common human African ape stock prior to the human chimpanzee and gorilla divergence 12 The earliest fossils clearly in the human but not the chimpanzee lineage appear between about 4 5 to 4 million years ago with Australopithecus anamensis Few fossil specimens on the chimpanzee side of the split have been found the first fossil chimpanzee dating between 545 and 284 kyr thousand years radiometric was discovered in Kenya s East African Rift Valley McBrearty 2005 13 All extinct genera listed in the taxobox which are ancestral to Homo or are offshoots of such However both Orrorin and Sahelanthropus existed around the time of the divergence and so either one or both may be ancestral to both genera Homo and Pan Due to the scarcity of fossil evidence for CHLCA candidates Mounier 2016 presented a project to create a virtual fossil by applying digital morphometrics and statistical algorithms to fossils from across the evolutionary history of both Homo and Pan having previously used this technique to visualize a skull of the last common ancestor of Neanderthal and Homo sapiens 14 15 Age estimates editAn estimate of TCHLCA of 10 to 13 million years was proposed in 1998 note 1 and a range of 7 to 10 million years ago is assumed by White et al 2009 In effect there is now no a priori reason to presume that human chimpanzee split times are especially recent and the fossil evidence is now fully compatible with older chimpanzee human divergence dates 7 to 10 Ma White et al 2009 17 Some researchers tried to estimate the age of the CHLCA TCHLCA using biopolymer structures that differ slightly between closely related animals Among these researchers Allan C Wilson and Vincent Sarich were pioneers in the development of the molecular clock for humans Working on protein sequences they eventually 1971 determined that apes were closer to humans than some paleontologists perceived based on the fossil record note 2 This paradigmatic age has stuck with molecular anthropology until the late 1990s Since the 1990s the estimate has again been pushed towards more remote times because studies have found evidence for a slowing of the molecular clock as apes evolved from a common monkey like ancestor with monkeys with both humans and non human apes evolving from a common ape like ancestor 19 A 2016 study analyzed transitions at CpG sites in genome sequences which exhibit a more clocklike behavior than other substitutions arriving at an estimate for human and chimpanzee divergence time of 12 1 million years 20 Gene flow editA source of confusion in determining the exact age of the Pan Homo split is evidence of a more complex speciation process than a clean split between the two lineages Different chromosomes appear to have split at different times possibly over as much as a 4 million year period indicating a long and drawn out speciation process with large scale gene flow events between the two emerging lineages as recently as 6 3 to 5 4 million years ago according to Patterson et al 2006 21 Speciation between Pan and Homo occurred over the last 9 million years Ardipithecus probably branched off of the Pan lineage in the middle Miocene Messinian 10 11 After the original divergences there were according to Patterson 2006 periods of gene flow between population groups and a process of alternating divergence and gene flow that lasted several million years 21 Some time during the late Miocene or early Pliocene the earliest members of the human clade completed a final separation from the lineage of Pan with date estimates ranging from 13 million 16 to as recent as 4 million years ago 21 The latter date was in particular based on the similarity of the X chromosome in humans and chimpanzees a conclusion rejected as unwarranted by Wakeley 2008 who suggested alternative explanations including selection pressure on the X chromosome in the populations ancestral to the CHLCA note 3 Complex speciation and incomplete lineage sorting of genetic sequences seem to also have happened in the split between the human lineage and that of the gorilla indicating messy speciation is the rule rather than the exception in large primates 23 24 Such a scenario would explain why the divergence age between the Homo and Pan has varied with the chosen method and why a single point has so far been hard to track down See also editHistory of hominoid taxonomy List of human evolution fossils with images Notes edit Based on a revision of the divergence of Hominoidea from Cercopithecoidea at more than 50 Mya previously set at 30 Mya Consistent with the marked shift in the dating of the Cercopithecoidea Hominoidea split all hominoid divergences receive a much earlier dating Thus the estimated date of the divergence between Pan chimpanzee and Homo is 10 13 MYBP and that between Gorilla and the Pan Homo linage 17 MYBP 16 If man and old world monkeys last shared a common ancestor 30 million years ago then man and African apes shared a common ancestor 5 million years ago 18 Patterson et al suggest that the apparently short divergence time between humans and chimpanzees on the X chromosome is explained by a massive interspecific gene flow event in the ancestry of these two species However Patterson et al do not statistically test their own null model of simple speciation before concluding that speciation was complex and even if the null model could be rejected they do not consider other explanations of a short divergence time on the X chromosome These include natural selection on the X chromosome in the common ancestor of humans and chimpanzees changes in the ratio of male to female mutation rates over time and less extreme divergence versions with gene flow I therefore believe that their claim of gene flow is unwarranted 22 References edit Mann Alan Mark Weiss 1996 Hominoid Phylogeny and Taxonomy a consideration of the molecular and Fossil Evidence in an Historical Perspective Molecular Phylogenetics and Evolution 5 1 169 181 doi 10 1006 mpev 1996 0011 PMID 8673284 B Wood 2010 Reconstructing human evolution Achievements challenges and opportunities Proceedings of the National Academy of Sciences 107 Suppl 2 8902 8909 Bibcode 2010PNAS 107 8902W doi 10 1073 pnas 1001649107 PMC 3024019 PMID 20445105 Wood and Richmond Richmond BG 2000 Human evolution taxonomy and paleobiology Journal of Anatomy 197 Pt 1 19 60 doi 10 1046 j 1469 7580 2000 19710019 x PMC 1468107 PMID 10999270 Out of the Pan Into the Fire in Frans B M De Waal ed 2001 Tree of Origin What Primate Behavior Can Tell Us About Human Social Evolution Harvard University Press pp 124 126 ISBN 9780674010048 Bradley B J 2006 Reconstructing Phylogenies and Phenotypes A Molecular View of Human Evolution Journal of Anatomy 212 4 337 353 doi 10 1111 j 1469 7580 2007 00840 x PMC 2409108 PMID 18380860 Fuss Jochen Spassov Nikolai Begun David R Bohme Madelaine 2017 Potential hominin affinities of Graecopithecus from the Late Miocene of Europe PLOS ONE 12 5 e0177127 Bibcode 2017PLoSO 1277127F doi 10 1371 journal pone 0177127 PMC 5439669 PMID 28531170 Graecopithecus freybergi Oldest Hominin Lived in Europe not Africa Archived from the original on 5 November 2019 Retrieved 5 November 2019 Dickinson Kevin 20 April 2019 New fossils suggest human ancestors evolved in Europe not Africa Big Think Retrieved 23 October 2022 Wood Bernard Harrison Terry 2011 The evolutionary context of the first hominins Nature 470 7334 347 35 Bibcode 2011Natur 470 347W doi 10 1038 nature09709 PMID 21331035 S2CID 4428052 a b Wood Bernard Harrison Terry 2011 The evolutionary context of the first hominins Nature 470 7334 347 52 Bibcode 2011Natur 470 347W doi 10 1038 nature09709 PMID 21331035 S2CID 4428052 a b Wolpoff Milford H 1996 Human Evolution McGraw Hill Incorporated ISBN 978 0070718333 Sarmiento E E 2010 Comment on the Paleobiology and Classification of Ardipithecus ramidus Science 328 5982 1105 author reply 1105 Bibcode 2010Sci 328 1105S doi 10 1126 science 1184148 PMID 20508113 McBrearty Sally Nina G Jablonski 2005 First fossil chimpanzee Nature 437 7055 105 108 Bibcode 2005Natur 437 105M doi 10 1038 nature04008 PMID 16136135 S2CID 4423286 Virtual fossil reveals last common ancestor of humans and Neanderthals 18 December 2015 Retrieved 30 June 2019 Mounier Aurelien Mirazon Lahr Marta 2016 Virtual ancestor reconstruction Revealing the ancestor of modern humans and Neandertals Journal of Human Evolution 91 57 72 doi 10 1016 j jhevol 2015 11 002 PMID 26852813 a b Arnason U Gullberg A Janke A December 1998 Molecular timing of primate divergences as estimated by two nonprimate calibration points Journal of Molecular Evolution 47 6 718 27 Bibcode 1998JMolE 47 718A doi 10 1007 PL00006431 PMID 9847414 S2CID 22217997 White TD Asfaw B Beyene Y et al October 2009 Ardipithecus ramidus and the paleobiology of early hominids Science 326 5949 75 86 Bibcode 2009Sci 326 75W doi 10 1126 science 1175802 PMID 19810190 S2CID 20189444 Sarich Vincent Wilson Allan 1967 Immunological Time Scale for Hominid Evolution Science 158 3805 1200 1203 Bibcode 1967Sci 158 1200S doi 10 1126 science 158 3805 1200 PMID 4964406 S2CID 7349579 Venn Oliver Turner Isaac Mathieson Iain de Groot Natasja Bontrop Ronald McVean Gil June 2014 Strong male bias drives germline mutation in chimpanzees Science 344 6189 1272 1275 Bibcode 2014Sci 344 1272V doi 10 1126 science 344 6189 1272 PMC 4746749 PMID 24926018 Moorjani Priya Amorim Carlos Eduardo G Arndt Peter F Przeworski Molly 2016 Variation in the molecular clock of primates Proceedings of the National Academy of Sciences 113 38 10607 10612 Bibcode 2016PNAS 11310607M doi 10 1073 pnas 1600374113 ISSN 0027 8424 PMC 5035889 PMID 27601674 a b c Patterson N Richter DJ Gnerre S Lander ES Reich D June 2006 Genetic evidence for complex speciation of humans and chimpanzees Nature 441 7097 1103 8 Bibcode 2006Natur 441 1103P doi 10 1038 nature04789 PMID 16710306 S2CID 2325560 Wakeley J 2008 Complex speciation of humans and chimpanzees Nature 452 7184 E3 4 Bibcode 2008Natur 452 3W doi 10 1038 nature06805 PMID 18337768 S2CID 4367089 Scally A Dutheil JY Hillier LW et al March 2012 Insights into hominid evolution from the gorilla genome sequence Nature 483 7388 169 75 Bibcode 2012Natur 483 169S doi 10 1038 nature10842 PMC 3303130 PMID 22398555 Van Arsdale A P Go go Gorilla genome The Pleistocene Scene A P Van Arsdale Blog Retrieved 16 November 2012 External links edit nbsp Wikispecies has information related to Hominini Human Timeline Interactive Smithsonian National Museum of Natural History August 2016 Locomotion and posture from the common hominoid ancestor to fully modern hominins with special reference to the last common panin hominin ancestor R H Crompton E E Vereecke and S K S Thorpe Journal of Anatomy April 2008 Portals nbsp Evolutionary biology nbsp Primates nbsp Science Retrieved from https en wikipedia org w index php title Chimpanzee human last common ancestor amp oldid 1191278122, wikipedia, wiki, book, books, library,

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