Neanderthal

Figure 1

Figure 2

I ONCE had a science teacher who, for reasons I know not, refused to pronounce the ‘th’ in Neanderthal, pronouncing it instead as, Neander-tall. Even now, the thought of it makes my ears cringe. 

And his pronunciation of Australopithecus was even worse. Not a mis-pronunciation per se, but far too nasally, too much tongue against teeth, and .... uck, the crack of trapped saliva in the throat on his ‘icus’.

His speech was otherwise unaccented Canadian, with these peculiar deviations reserved solely for his pronunciation of species' names, at which point he suddenly became ... I don't know ... a Scandinavian avec un rhume, peut-être?

Strange is that, ears I mean, strange how extreme aural sensitivity can make me fall in love with a voice, or, alternatively, in loathe; how the wrong voice can completely ruin an otherwise interesting film, lecture, conversation, et cetera. Maybe that is part of the reason why I am so fond of books.

But I digress. Today’s entry is about the Neanderthals. I tried to round up as much research as I could from the past few years, but there was a tremendous amount of material to sift through, and alas, I do have a day job. 

Undoubtedly much remains unknown about Neanderthals et al, with huge theoretical swathes left open to speculation. But if the abundance of ancestry.com type websites, commercials and TV shows indicate anything, it is that we have an abundance of curiosity about our genetic make-up; about our origins and recent past; and about what goes into making us, us.

On the Subject of the Neanderthal 

• The Neanderthal is named after the site of its first discovery in Germany’s Neander valley. [56] 

• Neanderthals are the closest extinct relatives of modern humans. [2]

Figure 3

The Neanderthals are believed to have inhabited Europe and parts of western Asia between 230,000 and 28,000 years ago. [79]

Neanderthal fossils have been found across Europe and Asia, from Spain to Russia to Iraq. [3]

Figure 4

Puncture marks in the skull fragments of a Neanderthal child unearthed at the Cova Negra archaeological site in Spain are believed to have been caused by large carnivore attacks.  [48-50]

Figure 5

• Scientists dated this calcite-encrusted skeleton embedded in rock, deep inside a cave in Italy, to about 130,000 to 170,000 years old. 

• This makes him the oldest Neanderthal from which DNA has ever been extracted. [29]

• The DNA is too degraded to sequence the fossil’s genome using current technology. [29][30]

Jewelry

• Both ornaments and the pigment ocher have also been found at archaeological sites inhabited by Neanderthals. [7]

Figure 6

In March of this year, Davorka Radovčić et al, described how the remains of eight eagle 

talons Figure 6, dating to ~ 130,000 years ago, show that the Neanderthals were making jewelry well before the appearance of modern humans in Europe. [36] 

Diet

• Evidence collected from hardened dental plaque chipped off the teeth of a 50,000 year old Neanderthal from El Sidrón in Spain tells us Neanderthals consumed (or at least chewed) the strong flavored herbs, chamomile and yarrow (two anti-inflammatories). 

• Some researchers propose that these plants were eaten for medicinal purposes, while others suggest they were used to flavor their food. [35]

• Analysis of biomarkers from 50,000 year old samples of fecal matter has shown that Neanderthals predominantly consumed meat, but also had significant plant intake. [41]

• Archeological evidence found in Neanderthal teeth suggests that they had a diverse diet including: wild peas, acorns, pistachios, grass seeds, wild olives, pine nuts and date palms. [7]

• Increasing evidence suggests the Neanderthals were apex predators, [68][69] capable of bringing down a wide range of prey from deer, ibex and wild boar, to the occasional, mammoth, elephant and rhinoceros. [67][70]

• In 2010, an isotope analysis of Neanderthal teeth found traces of cooked vegetable matter. [71][72]

Gender Roles

• When Spanish researchers compared [52] deep dental grooves in fossilized Neanderthal teeth by sex, they found that women’s teeth had longer grooves in the lower portions of their incisors and canine teeth, while men had nicks in the upper portions.

• Although the fossil record demonstrates that Neanderthals used their teeth as a tool to aid in tasks, [53] the different dental grooves suggest different work was being done between the sexes. [51, 52]

Burial Practices 

• Researchers studying the burial practices of Neanderthals found that most of the dead were buried in places where the Neanderthals had lived, often in caves. [24][25] 

• More than a third of Neanderthal graves found contained children under the age of four, typically showing great care in their burials.

• Some were buried with objects, including flint scrapers, a principal tool.

• Many had what appear to be ceremonial animal bones laid out carefully around the body.

• DNA analysis on the remains of one group burial plot found in Spain showed that the six adults, two juveniles and one infant were related. The three males were brothers, and two of the children were the offspring of two of the females. 

Figure 7

In another study, this one at the site of Marillac, in France, researchers found evidence of peri-mortem (shortly after death) manipulations on bones. Indicated on Figure 7 are cut marks on the femur of a Neanderthal child.

The reason behind the markings can only be speculated upon. [32-34]

Tools, Hunting & Travel

• The discovery of stone tools on the southern Ionian Islands has lead some researchers to speculate that Neanderthals may have been sailing the Mediterranean Sea as early as 110,000 years ago. [57][58]

• Quartz hand-axes, three-sided picks, and stone cleavers from Crete, dating to 170,000 have also been recovered. [59]

• Evidence from multiple archaeological sites in Europe suggest that Neanderthals hunted as a group, using the landscape to aid them. 

• They made spears by attaching stone points to wooden shafts. [7]

Figure 8

Figure 8 is an animation of the Levallois technique used by the Neanderthals.

Striking the platform separates a flake from the core with all of its edges sharpened by the trimming work.

Figure 9

The method provided greater control over the size and shape of the final flake which could then be used as is, or adapted into Lavallois points Figure 9. [70]

Speech & Expression

• The Neanderthal genome did include the FOXP2 gene which is required for proper speech and language development in humans. [60]

• The anatomy of their vocal tracts would have prevented them from sounding some vowels. [7]

Figure 10

Rock engravings at Gorham's Cave in Gibraltar could provide evidence of "symbolic thought" and "abstract expression." [78]

Eyes & Ears

• A 2013 study of Neanderthal skulls suggests that their eyesight may have been better than that of modern humans, owing to larger eye sockets and larger areas of the brain devoted to vision. [73]

Figure 11

Virtual 3D reconstruction of the auditory ossicle (ear bone) of a 2 year old Neanderthal child shows morphological differences between humans and the Neanderthals. 

It is not known how or if these morphological differences relate to differences in hearing capabilities.  [37][38]

• The gene ARHGAP11B, has been identified as playing a key role in the evolutionary expansion of the neocortex* in Homo sapiens. Neanderthals and Denisovans both possess this gene, Chimpanzees do not. [22][23]

*In humans, the neocortex is involved in higher functions such as sensory perception, generation of motor commands, spatial reasoning, conscious thought and language. [81]

Extinction 

• The Neanderthals gradually disappeared from Europe between 41,000 and 39,000 years ago. This time frame coincides with the start of a very cold period in Europe and is 5000 years after Homo sapiens are believed to have reached the continent. [61] 

• These millennia of co-existence created the opportunity for genetic and cultural exchange. [2][5]

Figure 12

Mitochondrial DNA-based simulation Figure 12 of modern human expansion in Europe starting (a) 1600 generations ago. 

Neanderthal range in light grey. [62] 

Extinction Theories

• Perhaps the longest running theory is that it was the superior intelligence of modern humans that drove their barrel chested cousins to extinction. [7][8][9] 

• Along the same lines as above is the theory that it was the Neanderthal's failure to master fire (to use it as "often or consistently" as modern humans) that led to their decline. [6] 

• Climate cooling caused by a volcanic super-eruption in Naples around 40,000 years ago has also been suggested. This is unlikely, however, as the European Neanderthal population was already in decline by that time, and already be extinct in Italy. [4]

• Scarcity of resources seems plausible as a contributing factor. [63]

• Studies on Neanderthal body structures have shown that they needed more energy/calories to survive than any other species of hominid. [64] 

• Dr John Stewart, is part of a team which found that while rabbits were a crucial part of the modern humans' diet, they were relatively under-utilised by Neanderthals. [12][13][14]

"Modern humans had more that they could do – they had more possibilities and were more able to cope with the deterioration of climate than Neanderthals were."

• Perhaps, although another research team working in Israel have found evidence that Neanderthals had a history of adapting to resource fluctuations.

• By reconstructing hunting ranges and comparing oxygen, carbon, and strontium isotopes from gazelle tooth enamel with modern isotope data from the Amud Cave region in Israel, researchers found that Neanderthals adjusted their hunting territories considerably in relation to varying environmental conditions. [39][40]

• In her book The Invaders How Humans and Their Dogs Drove Neanderthals to Extinction, Pat Shipman states that, [19][20][21]

“The synergy between climate change and the invasion of modern humans created an untenable situation for Neanderthals. ...
There was an ecosystem-wide crash involving many members of the predatory guild."

• As noted by Barbara J. King in her NPR piece on Shipman's book, the 'and Their Dogs ...', part of the title is misleading as the Neanderthals would have already been extinct by the time the wolf-dogs appeared. To this the author replied: [21]

"... The second theme is that once humans allied with wolf-dogs (dogs), we formed an unbeatable team that was far more formidable in hunting than either alone. If this proves to have occurred after Neanderthals went extinct, then so be it because the appearance of a new invasive predator (us) was enough to cause serious and possibly crippling pressure on Neanderthals. If, on the other hand, our pact with wolf-dogs appeared while Neanderthals were still around, albeit in smaller numbers and more restricted areas, the domestication of wolves into wolf-dogs would have been a fatal innovation."

Interbreeding

• In May 2010, a research team led by evolutionary biologist Svante Paabo announced that they had sequenced, “the whole Neanderthal genome from powdered bone fragments taken from three females who lived in Europe 40,000 years ago.” [66][67]

• The researchers compared the genome of the Neanderthals to 5 modern humans from China, France, sub-Saharan Africa, and Papua New Guinea.

• The sequencing results indicated that between 1% to 4% of the DNA in modern humans of non-African origin was inherited from Neanderthals. [55]

• By 2012, further research had narrowed the time frame during which interbreeding was likely to have occurred to between 47,000 and 65,000 years ago. [15][80]

• In 2014, the genome of a 45,000 year old modern human male from Siberia was sequenced, and was found to contain an estimated 2.3% Neanderthal DNA. [15][16]

• This enabled researchers to refine the time frame for interbreeding to approximately 50,000 to 60,000 years ago, which is close to the time of the major expansion of modern humans out of Africa and the Middle East. [15][16]

Figure 13

Writing in the journal Nature, in April of this year, Israel Hershkovitz et al [18] explain how dating a unique human skull Figure 13 found in Manot Cave, Israel to 55,000 years ago shows that modern humans arrived in the region when it was already well populated with Neanderthals. 

• That modern humans and Neanderthals shared the land around Manot cave 50,000 to 60,000 years ago provides the opportunity for interbreeding that left non-Africans with a genome containing ~ 2% Neanderthal DNA. [17]

Figure 14

In May of this year, researchers reported that DNA extracted from a 40,000 year old modern human jawbone fossil in Romania revealed the genome of a male containing 5 to 11 % Neanderthal DNA. [10][11]

• That this modern human had a Neanderthal in its family tree going back four to six generations (as in great-great-grandparent) raises the possibility that interbreeding was not isolated to an initial co-mingling in the Middle East more than 50,000 years ago; that it continued into Europe, and was occurring right up until the Neanderthals went extinct. [10][11]

• Researchers analyzing genetic variants in 846 people of non-African heritage, 176 people from sub-Saharan Africa, and a 50,000-year-old Neanderthal genome sequence published in 2013 found that remnants of Neanderthal DNA in modern humans were associated with genes affecting type 2 diabetes, Crohn’s disease, lupus, biliary cirrhosis, and smoking behavior. [42–47]

• Neanderthal ancestry was also found to be increased in genes affecting keratin filaments, a fibrous protein which improves the toughness of skin, hair, and nails. [42-46]

• Genes expressed in the testes and on the X chromosome had reduced Neanderthal ancestry, a pattern which has been linked to hybrid infertility in fruit flies. [42-46]

•  This suggests that after 500,000 years of evolutionary separation, some Neanderthal-human hybrids would have had reduced fertility, and others may have been completely sterile. [42-46]

East Asian Populations

• East Asians have, on average, higher levels of Neanderthal ancestry than do Europeans, suggesting that Asian population had an additional encounter with the Neanderthals after they had already gone extinct in Europe. 

• It is unknown if the reason for this is because extinction happened later in Asia, allowing for Neanderthals to breed with Asians a second time; or because Asians interbred with some other group of humans that had interbred with Neanderthals; or perhaps, because there was a dilution of Neanderthal lineages in Europeans by interbreeding with an unknown ancestral population. 

• As yet undiscovered fossil evidence may one day offer an explanation. [26][27][28]

Figure 15

Chris Stringer's hypothesis of the family tree of genus Homo, published 2012 in Nature: [65]

• Homo floresiensis (left of diagram) originated in an unknown location from unknown ancestors and reached remote parts of Indonesia. 

• Homo erectus spread from Africa to western Asia, then East Asia and Indonesia; its presence in Europe is uncertain, but it gave rise to Homo antecessor, found in Spain. 

• Homo heidelbergensis originated from Homo erectus in an unknown location and dispersed across Africa, southern Asia and southern Europe. 

• Homo sapiens spread from Africa to western Asia and then to Europe and southern Asia, eventually reaching Australia and the Americas. 

• In addition to Neanderthals and Denisovans, a third gene flow of archaic Africa origin is indicated at the far right of the diagram. 

• If you want to know how much Neanderthal DNA you carry, check out  National Geographic Society’s Genographic Project. [7]

AND LASTLY, (and in an attempt to maintain balance) a note on the reliability of ancient DNA analysis as provided in the following excerpts from A.J. Smuskiewicz’s, More mystery about Neanderthal and modern humans: How reliable is ancient DNA analysis : [55]

• As soon as any organism dies, its DNA begins to break down and decay as the nucleotide bonds fall apart. Chemical reactions with water in the ground accelerate the decay. After examining such factors, Danish and Australian researchers reported in 2012 that DNA has a half-life of only 521 years.

• Thus, the analysis of ancient, partially decayed, and fragmentary DNA inevitably includes making certain assumptions and decisions about missing and degraded pieces, in order to fill in the gaps as best as possible. Some decisions about assembling Neanderthal sequences are based on sophisticated algorithms involving the use of modern human and chimpanzee genomes as guides.

• These challenges and complexities raise the possibility of the inadvertent introduction of errors in the sequencing process—even when all precautions are taken by highly professional researchers. These issues also open up any obtained data to different interpretations and to possible limitations in the application of the data.

According to a model developed by Cambridge evolutionary ecologists Andrea Manica and Anders Eriksson, 

“There was an ancestor of both Neanderthal and modern humans—some archaeologists would call that Homo heidelbergensis—that would have covered Africa and Europe about half a million years ago.” [54]

In other words, it may just be a remnant of a common ancient African ancestor and not the result of interbreeding. [74][75]  

• *John D. Hawks, an associate professor of Anthropology at the University of Wisconsin–Madison, has argued that the genetic similarity to Neanderthals may indeed be the result of both structure and interbreeding, as opposed to just one or the other. [76] 

• *First Peoples, a series on the origins and spread of modern humans around the world premieres June 24 on PBS. 

•  Watch one of John D. Hawks clips from the upcoming series via this link: “The Secret to Our Success - Connectivity”.

UPDATES / RELATED

DNA from Neandertal relative may shake up human family tree / Scientific American / September 11, 2015

***

Fin

REFERENCES

[1] http://www.abc.net.au/science/articles/2015/04/27/4224475.htm     

[2] http://www.livescience.com/49290-volcano-did-not-kill-neanderthals.html

[3] http://science.howstuffworks.com/life/evolution/why-did-neanderthals-become-extinct.htm

[4] http://www.livescience.com/49290-volcano-did-not-kill-neanderthals.html

[5] Humans Did Not Wipe Out the Neanderthals, New Research Suggests

[6] Did Neanderthals Die Off Because They Couldn't Harness Fire?

[7] http://discovermagazine.com/2013/dec/22-20-things-you-didnt-know-aboutneanderthals By Jonathon Keats|Friday, September 05, 2014

[8] Neanderthals were not inferior to modern humans, study finds

[9] Paola Villa, Wil Roebroeks. Neandertal Demise: An Archaeological Analysis of the Modern Human Superiority Complex. PLOS ONE, 2014; 9 (4): e96424 DOI: 10.1371/journal.pone.0096424

[10] Analysis of bones found in Romania offer evidence of human and Neanderthal interbreeding in Europe / May 14, 2015 / phys.org

[11] Early European may have had Neanderthal great-great-grandparent / Nature / May 13, 2015

[12]What rabbits can tell us about Neanderthal extinction? / phys.org

Jun 09, 2015

[13] Why Didn’t Neanderthals Hunt Rabbits? / June 9, 2015

[14]"Rabbits and hominin survival in Iberia," Journal of Human Evolution, Volume 64, Issue 4, April 2013, Pages 233-241, ISSN 0047-2484, dx.doi.org/10.1016/j.jhevol.2013.01.002

[15] http://news.sciencemag.org/archaeology/2015/01/humans-and-neandertals-likely-interbred-middle-east

[16] Qiaomei Fu et al. Genome sequence of a 45,000-year-old modern human from western Siberia

Nature 514, 445–449 (23 October 2014) doi:10.1038/nature13810

[17] http://www.theguardian.com/science/2015/jan/28/ancient-skull-found-israel-sheds-light-human-migration-sex-neanderthals

[18] Hershkovitz, Israel et al. Levantine cranium from Manot Cave (Israel) foreshadows the first European modern humans Nature 2015/04/09/ http://dx.doi.org/10.1038/nature14134 10.1038/nature14134

[19] http://www.cbc.ca/radio/quirks/quirks-quarks-for-april-4-2015-1.3019735/how-dogs-helped-humans-drive-neanderthals-to-extinction-1.3019817

[20] http://news.nationalgeographic.com/news/2015/03/150304-neanderthal-shipman-predmosti-wolf-dog-lionfish-jagger-pogo-ngbooktalk/

[21] http://www.npr.org/sections/13.7/2015/03/26/395488202/what-drove-neanderthals-to-extinction-maybe-us

[22] Scientists Identify ‘Big Brain’ Gene in Humans, Neanderthals

[23] Marta Florio et al. Human-specific gene ARHGAP11B promotes basal progenitor amplification and neocortex expansion. Science, published online February 26, 2015; doi: 10.1126/science.aaa1975

[24] http://www.huffingtonpost.com/2014/04/29/neanderthals-parents-bone-study_n_5231842.html

[25] Penny Spikins et al. The Cradle of Thought: Growth, Learning, Play and Attachment in Neanderthal Children. Oxford Journal of Archaeology Volume 33, Issue 2, pages 111–134, May 2014

[26] http://blog.cosmosmagazine.com/blog/2015/2/20/why-asians-carry-more-neanderthal-dna

[27] A New Theory on How Neanderthal DNA Spread in Asia / February 19, 2015 / NYTimes

[28] http://www.cell.com/ajhg/abstract/S0002-9297%2815%2900014-2

[29] Oldest Neanderthal DNA Found in Italian Skeleton / Charles Q. Choi, Live Science Contributor   |   April 10, 2015

[30] The Neanderthal in the karst: First dating, morphometric, and paleogenetic data on the fossil skeleton from Altamura (Italy), Journal of Human Evolution, Available online 21 March 2015 DOI: 10.1016/j.jhevol.2015.02.007
[31] http://phys.org/news/2015-04-altamura-yields-oldest-neanderthal-dna.html

[32] Neandertal Post-Mortem Modification / Anthropology.net / April 14, 2015

[33] Dolores Garralda, M., Maureille, B. and Vandermeersch, B. (2014), Neanderthal infant and adult infracranial remains from Marillac (Charente, France). Am. J. Phys. Anthropol., 155: 99–113. doi: 10.1002/ajpa.22557

[34] http://www.natureworldnews.com/articles/14078/20150414/neanderthals-beat-up-their-deceased-brethren-study.htm

[35] http://www.newscientist.com/article/mg22630174.600-neanderthal-chefs-may-have-spiced-up-menus-with-wild-herbs.html#.VXo6A2jF_UU

[36] Radovčić D, Sršen AO, Radovčić J, Frayer DW (2015) Evidence for Neandertal Jewelry: Modified White-Tailed Eagle Claws at Krapina. PLoS ONE 10(3): e0119802. doi:10.1371/journal.pone.0119802

[37] The stapes of a Neanderthal child points to the anatomical differences with our species      www.eurekalert.org / March 27, 2015

[38] Asier Gómez-Olivencia et al. La Ferrassie 8 Neandertal child reloaded: New remains and re-assessment of the original collection Journal of Human Evolution Volume 82, May 2015, Pages 107–126

[39] Neanderthals changed hunting strategy with climate change

[40] Gideon Hartman, et al., Isotopic evidence for Last Glacial climatic impacts on Neanderthal gazelle hunting territories at Amud Cave, Israel, Journal of Human Evolution, 7 May 2015 doi:10.1016/j.jhevol.2015.03.008

[41] Sistiaga A, Mallol C, Galván B, Summons RE (2014). The Neanderthal Meal: A New Perspective Using Faecal Biomarkers

PLoS ONE 9(6): e101045. doi:10.1371/journal.pone.0101045

(2014)

[42] Neanderthals gave us disease genes / By Paul RinconScience editor, BBC News website / 29 January 2014

[43] Modern human genomes reveal our inner Neanderthal / nature.com/ January 29, 2014.

[44] Ewen Callaway29 January 2014 Cross-breeding boosted Homo sapiens' ability to cope with cool climates, but the hybrids may have had trouble breeding. Nature doi:10.1038/nature.2014.14615

[45] Neanderthals’ DNA legacy linked to modern ailments January 29, 2014 / Stephanie Dutchen, Harvard Medical School Communications Science. 2014 Feb 28;343(6174):1017-21. doi: 10.1126/science.1245938. Epub 2014 Jan 29.

[46] Vernot B1, Akey JM. Resurrecting surviving Neandertal lineages from modern human genomes.Nature. 2014 Jan 2;505(7481):43-9. doi: 10.1038/nature12886.

[47] Prüfer K et al. The complete genome sequence of a Neanderthal from the Altai Mountains. Nature. 2014 Jan 2;505(7481):43-9. doi: 10.1038/nature12886. Epub 2013 Dec 18. PMID:24352235

[48] Ancient Carnivores Had a Taste for Neanderthal Meat / Helen Thompson /Smithsonian.com / MAY 26, 2015

[49] NEANDERTHALS WERE ATTACKED BY LARGE CARNIVORES / May 18, 2015

[50] Camarós, E., Cueto, M., Lorenzo, C., Villaverde, V., Rivals, F. “Large carnivore attacks on hominins during the Pleistocene: a forensic approach with a neanderthal example”, Archaeological and Anthropological Sciences. (2015).

[51] Neanderthals Divvied Up Chores by Sex / Erin Blakemore /smithsonian.com / February 20, 2015

[52] Almudena Estalrrich, , Antonio Rosas Division of labor by sex and age in Neandertals: an approach through the study of activity-related dental wear

Journal of Human Evolution Volume 80, March 2015, Pages 51–63

[53] Laura Pacey Mouth used as third hand by cavemen British Dental Journal 213, 336 (2012) 

Published online: 12 October 2012 | doi:10.1038/sj.bdj.2012.902

[54] Anders Eriksson1 and Andrea Manica1 Effect of ancient population structure on the degree of polymorphism shared between modern human populations and ancient hominins vol. 109 no. 35 13956–13960, doi: 10.1073/pnas.1200567109

[55] http://www.geneticliteracyproject.org/2015/05/12/more-mystery-about-neanderthal-and-humans-how-reliable-is-ancient-dna-analysis/

More mystery about Neanderthal and modern humans: How reliable is ancient DNA analysis? A. J. Smuskiewicz | May 12, 2015 | Genetic Literacy Project

[56] https://en.wikipedia.org/wiki/Neanderthal

[57] "Neanderthals beat modern humans to the seas by 50,000 years, say scientists".

[58] "Ancient Mariners: Did Neanderthals Sail to Mediterranean?".

[59] "Neanderthals May Have Sailed to Crete".

[60] Wade, Nicholas (19 October 2007)."Neanderthals Had Important Speech Gene, DNA Evidence Shows". The New York Times.

[61] "BBC News - New dates rewrite Neanderthal story". BBC News.  August 20, 2014

[62] Currat, Mathias; Excoffier, Laurent (2004). "Modern Humans Did Not Admix with Neanderthals during Their Range Expansion into Europe". PLoS Biology 2 (12): e421.doi:10.1371/journal.pbio.0020421. PMC 532389.PMID 15562317.

[63] "The Mysterious Downfall of the Neandertals", Scientific American, August 2009

[64]  Froehle, Andrew W.; Churchill, Steven E. (2009). "Energetic Competition Between Neandertals and Anatomically Modern Humans" (PDF). PaleoAnthropology: 96–116.

[65] Stringer, Chris (2012). "Evolution: What makes a modern human".Nature 485 (7396): 33–5. Bibcode:2012Natur.485...33S.doi:10.1038/485033a. PMID 22552077.

[66] Green, Richard E. et al (2010). "A Draft Sequence of the Neandertal Genome". Science 328 (5979): 710–22. Bibcode:2010Sci...328..710G. doi:10.1126/science.1188021.PMID 20448178.

[67] Rincon, Paul (2010). "Neanderthal genes 'survive in us'". BBC News (BBC).

[68] Papagianni, Dmitra; Morse, Michael (2013). The Neandethals Rediscovered. Thames & Hudson. ISBN 978-0-500-05177-1.

[69] Bocherens, Hervé; Drucker, Dorothée G.; Billiou, Daniel; Patou-Mathis, Marylène; Vandermeersch, Bernard (2005). "Isotopic evidence for diet and subsistence pattern of the Saint-Césaire I Neanderthal: Review and use of a multi-source mixing model".Journal of Human Evolution 49 (1): 71–87.doi:10.1016/j.jhevol.2005.03.003. PMID 15869783.

[70] Ghosh, Pallab. "Neanderthals cooked and ate vegetables." BBC News. December 27, 2010. Lichfield, John (September 30, 2006). "French dig up Neanderthal 'butcher's shop'". The New Zealand Herald.

[71] Richards, Michael P.; Pettitt, Paul B.; Trinkaus, Erik; Smith, Fred H.; Paunović, Maja; Karavanić, Ivor (2000). "Neanderthal diet at Vindija and Neanderthal predation: The evidence from stable isotopes". Proceedings of the National Academy of Sciences 97(13): 7663–6. Bibcode:2000pnas...97.7663r.doi:10.1073/pnas.120178997. JSTOR 122870. PMC 16602.PMID 10852955.

[72] Webb, Jonathan (25 June 2014). "Oldest human faeces show Neanderthals ate vegetables". BBC News.

[73]  "Neanderthal brains focused on vision and movement leaving less room for social networking". Science Daily. March 19, 2013.

[74]  Jha, Alok (14 August 2012). "Study casts doubt on human-Neanderthal interbreeding theory". The Guardian. Retrieved19 February 2015.

[75] Lowery, Robert K.; Uribe, Gabriel; Jimenez, Eric B.; Weiss, Mark A.; Herrera, Kristian J.; Regueiro, Maria; Herrera, Rene J. (2013). "Neanderthal and Denisova genetic affinities with contemporary humans: Introgression versus common ancestral polymorphisms".Gene 530 (1): 83–94. doi:10.1016/j.gene.2013.06.005.PMID 23872234.

[76] Hawks, John (2013). "Significance of Neandertal and Denisovan Genomes in Human Evolution". Annual Review of Anthropology 42: 433–49. doi:10.1146/annurev-anthro-092412-155548.

[77] https://en.wikipedia.org/wiki/Levallois_technique

[78] Neanderthals created cave art, researchers discover / September 2, 2014

[79] The stapes of a neanderthal child points to the anatomical differences with our species / phys.org / March 27, 2015.

[80] Sriram Sankararaman, Nick Patterson, Heng Li, Svante Pääbo, David Reich. The Date of Interbreeding between Neandertals and Modern Humans. PLOS Genestics Published: October 4, 2012. DOI: 10.1371/journal.pgen.1002947
[81] Lui, J. H.; Hansen, D. V.; Kriegstein, A. R. (2011). "Development and Evolution of the Human Neocortex". Cell 146 (1): 18–36.doi:10.1016/j.cell.2011.06.030. PMC 3610574. PMID 21729779.

IMAGE CREDITS

Figure 1: "Neanderthalensis" by Claire Houck from New York City, USA - Neanderthal Skeleton. Licensed under CC BY-SA 2.0 via Wikimedia Commons - https://commons.wikimedia.org/wiki/File:Neanderthalensis.jpg#/media/File:Neanderthalensis.jpg

Figure 2: "Homo sapiens neanderthalensis" by Luna04 - Own work. Licensed under CC BY 2.5 via Wikimedia Commons - https://commons.wikimedia.org/wiki/File:Homo_sapiens_neanderthalensis.jpg#/media/File:Homo_sapiens_neanderthalensis.jpg

Figure 3: "Carte Neandertaliens" by 120 - my own work 120. Licensed under CC BY-SA 3.0 via Wikimedia Commons - https://commons.wikimedia.org/wiki/File:Carte_Neandertaliens.jpg#/media/File:Carte_Neandertaliens.jpg

Figure 4: Cova Negra right parietal with punctures /  NEANDERTHALS WERE ATTACKED BY LARGE CARNIVORES / iphesnes.wordpress.com / May 18, 2015

Figure 5: "Altamura man" by Source. Licensed under Fair use via Wikipedia - https://en.wikipedia.org/wiki/File:Altamura_man.jpg#/media/File:Altamura_man.jpg

Figure 6: Radovčić D, Sršen AO, Radovčić J, Frayer DW (2015). Evidence for Neandertal Jewelry: Modified White-Tailed Eagle Claws at Krapina. PLoS ONE 10(3): e0119802. doi:10.1371/journal.pone.0119802 (Fig 3. Krapina 385.5, a right talon 1)

Figure 7: (Photo: M.D. Garralda et al) via anthropology.net / April 14, 2015

Figure 8: "Levallois Preferencial-Animation" by José-Manuel Benito Álvarez. Licensed under CC BY-SA 2.5 via Wikimedia Commons - https://commons.wikimedia.org/wiki/File:Levallois_Preferencial-Animation.gif#/media/File:Levallois_Preferencial-Animation.gif

Figure 9: "Pointe levallois Beuzeville MHNT PRE.2009.0.203.2.fond" by Didier Descouens - Own work. Licensed under CC BY-SA 3.0 via Wikimedia Commons - https://commons.wikimedia.org/wiki/File:Pointe_levallois_Beuzeville_MHNT_PRE.2009.0.203.2.fond.jpg#/media/File:Pointe_levallois_Beuzeville_MHNT_PRE.2009.0.203.2.fond.jpg

Figure 10: Neanderthal rock engraving at Gorham's Cave in Gibraltar. (Courtesy of Stuart Finlayson/Journal/Associated Press) via cbc.ca

Figure 11: 3-D Reconstruction of the Auditory Ossicle of a Two-Year-Old Neanderthal UPV/EHU via eurekalert.org

Figure 12: "CurratExcoffierNeandethalmtDNA" by Mathias Currat, Laurent Excoffier - http://dx.doi.org/10.1371/journal.pbio.0020421.g002. Licensed under CC BY 2.5 via Wikimedia Commons - https://commons.wikimedia.org/wiki/File:CurratExcoffierNeandethalmtDNA.png#/media/File:CurratExcoffierNeandethalmtDNA.png

Figure 13: Photograph: Tel Aviv University and University of Vienna, via Skull discovery suggests location where humans first had sex with Neanderthals / theguardian.com / January 28, 2015

Figure 14: (Pinterest) Pinterest Source: Ewen Callaway (2015) Early European may have had Neanderthal great-great-grandparent / nature.com /   (REF 1 – PNAS)

Figure 15: "Homo-Stammbaum, Version Stringer-en" by Homo-Stammbaum, Version Stringer.jpg: Chris Stringerderivative work: Conquistador - This file was derived from: Homo-Stammbaum, Version Stringer.jpg:. Licensed under CC BY-SA 3.0 de via Wikimedia Commons - https://commons.wikimedia.org/wiki/File:Homo-Stammbaum,_Version_Stringer-en.svg#/media/File:Homo-Stammbaum,_Version_Stringer-en.svg