Neanderthals in Southwest Asia

(Redirected from Levantine Neanderthal)

Southwest Asian Neanderthals were Neanderthals who lived in Turkey, Lebanon, Syria, Palestine, Iraq, and Iran - the southernmost expanse of the known Neanderthal range. Although their arrival in Asia is not well-dated, early Neanderthals occupied the region apparently until about 100,000 years ago. At this time, Homo sapiens immigrants seem to have replaced them in one of the first anatomically-modern expansions out of Africa. In their turn, starting around 80,000 years ago, Neanderthals seem to have returned and replaced Homo sapiens in Southwest Asia. They inhabited the region until about 55,000 years ago.[1]

The known range of Neanderthals in Southwest Asia (orange) and elsewhere. Sites that have yielded skeletal remains, tagged by country - Turkey (T1): Karain. Lebanon (L1-2): Ksâr 'Akil and El Masloukh. Palestine (Is1-5): Kebara, Tabun, Ein Qashish, Shovakh, and Amud, Palestine (WB1): Shuqba. Syria (S1): Dederiyeh. Iraq (Iq1): Shanidar. Iran (In1): Bisitun. Iran (In1):Wezmeh.

In Southwest Asia Neanderthals have left well-preserved skeletal remains in present-day Palestine, Syria, and Iraq. Remains in Turkey, Lebanon, and Iran are fragmentary. No Neanderthal skeletal remains have ever been found to the south of Jerusalem, and although there are Middle Palaeolithic Levallois points in Jordan and in the Arabian peninsula, it is unclear whether these were made by Neanderthals or by anatomically modern humans.[1] Neanderthals living further to the east, such as those found in present-day Uzbekistan[2][3] and Asian Russia[4][5] are known as Central and North Asian Neanderthals.

As of 2013, although many more Neanderthal remains have been discovered in Southwest Asia than in North Asia, where genetic studies have succeeded,[6] no attempt at extracting DNA from Southwest Asian Neanderthals has ever been successful.[7]

Early hominins (to 100 ka) edit

One of the southernmost Neanderthals: Homo neanderthalensis fossil from Tabun Cave, Israel. 120.000-50.000 BC. Israel Museum.

As the Levant is the landbridge to Eurasia, Dmanisi remains in Georgia from 1.81 Ma suggest that hominins passed through the Levant some time before this (unless they crossed the Bab el-Mandeb strait into Arabia). The oldest known hominin specimen from the Levant, and from all the Middle East, is the Zuttiyeh skull, found by Francis Turville-Petre near the Sea of Galilee in 1927. It has not been dated, but lithic industries suggest it dates from 350-250 ka (Bar-Yosef 1992).[8] Though it has some modern characteristics, it has been classified as a specimen of Homo heidelbergensis.

Other known hominin fossils in Southwest Asia older than 100 ka include Shanidar 2 and 4, as well as Tabun C1. All three have at one point been classified as Neanderthals, although the latter two are now usually called pre-Neanderthals. In part because none are well-dated these attributions have all been challenged.

Early AMHs in the Levant (100 to 80 ka) edit

It is believed that anatomically modern humans first entered the Levant by 130 – 100 ka (Grün 2005)[9] from Africa during the Last Interglacial, a hot, parched, period in Africa (Scholz et al. 2007).

Environment edit

Goat δ13C values from c. 92 ka collected at Qafzeh Cave show that goats were eating, depending on the season, between 20 and 50% of C4 plants. These plants are better adapted to dry and hot climates than the much more common C3 plants. Although it was already established that North Africa was arid in the early Last Glacial, Scholz et al. (2007)'s conclusions would extend this to the Levant, making the region in this period more arid than previously thought.[verification needed] Frumkin et al. (2011)[10] arrived at a similar conclusion of a hot and arid Levant[when?] by examining changes through time of lake levels and in the formation of cave formations.

Return of Neanderthals to the Levant (80 to 55 ka) edit

Potential causes for the initial extinction of modern humans from the Levant and subsequent replacement by Neanderthals around 80 ka include a short-term extreme environmental change in the Levant and environmental changes in the regions neighbouring the Levant. The only long-term environmental change that has been found in the Levant at this period is an increase in rainfall (Hallin et al. 2012),[11] and some moderate cooling of the region (Bar-Yosef 1992,[8] Harrison and Bates 1991),[12] which would not predict a hominin extinction (Hallin et al. 2012).[11] Hence, a short-lived intense environmental change (such as a volcanic winter of 6 to 1000 years caused by the Toba supereruption, Rampino and Self 1992)[13] might have caused modern human extinction, especially if the modern human Levantine population was low (Shea 2008).[14] In this case, Neanderthals could have moved in during or after the modern human evacuation of the region.

Environment edit

Contrarily to modern humans who arrived in the Levant in a period particularly warm in both Eurasia and Africa, Neanderthals are thought to have entered the Levant in a period of increasing Eurasian coldness and aridity known as the Last Glacial Period (Tchernov 1989).[15] In the Levant, however, the conditions would have been wetter than those of today and of the preceding interglacial. This is what Hallin et al. (2012)[11] concluded in their study of the δ13C and δ18O[Note 1] values of goat and gazelle enamel from Amud cave in Neanderthals layers dated to 70-53 ka (Glacial Period), after comparing them to values from Qafzeh cave in a modern human layer dated to 92 ka. They found that at Amud, neither the goats nor the gazelle consumed arid-adapted C4 plants, indicating that the environment was not as dry as today. They found the magnitude and pattern of δ18O values in gazelle and goat teeth enamel to indicate that, unlike today, rain fell throughout the year at Amud in the Neanderthal period. Indeed, the variation in δ18O values in the oldest layers of the teeth, formed when the animal is only a few months old, suggested that goats and gazelles were born throughout the year, which today happens only in the Levant in wetter-than-normal conditions.

Belmaker et al. (2011)[16] compared the prevalence in caves of microfaunal bones brought back by Barn and Eagle owls, which are known to accumulate large quantities of rodent skeletons. Since owls are opportunistic hunters and will eat any small mammal, the bone assemblage they leave behind are thought to be accurate indicators of the environment, given that rodents, like beetles, are susceptible to small changes in temperature and humidity. The researchers did find changes in microfaunal bone assemblages, but all were expected by taphonomic bias. They therefore concluded that there were no significant changes in the environment of the Levant between 70 and 55 ka.

Return of AMHs to the Levant (after 55 ka) edit

Unless modern humans started expanding into Eurasia through Arabia (where stone tools but very few human bones have been found), the Levant was most likely the first place Neanderthals lost ground to expanding modern humans.

It is unclear why Neanderthals abandoned the Levant or lost it to modern humans. Shea (2008)[14] has argued that Neanderthals were climatically forced out of the region (as modern humans probably were about 75,000 years ago). This is contested. Belmaker et al. (2011)[16] have found that the proportions of microfaunal bones in the Levant between 70 and 55 kya seem stable, despite the fact that small fauna are sensitive to slight changes in temperature or humidity. There would be hence little reason to believe in a slow deterioration of the Neanderthal habitat driving them out by 45 kya. It is possible that climate change drove Neanderthals to extinction quickly some time after 55,000 BP, but Hallin et al. (2012)[11] found strong consistency through time in the δ13C and δ18O values in the teeth of both goats and gazelles, suggesting that the climate was stable in the Levant at this crucial period when Middle Palaeolithic technologies of Neanderthals and early modern humans were replaced by Upper Palaeolithic technology of later modern humans.

Although the environment at this period in the Levant seems to have been stable, Shea (2008)[14] suggested that a rapidly changing environment at the border between the Levant and the Sinai could have stimulated the development of the first Upper Palaeolithic technologies. In this scenario, modern humans would only have been able to displace Neanderthals once they had developed this technology. The advanced tools found in Europe in the millennia thereafter would have developed there and then.

Resource gathering strategy edit

It is believed that Neanderthals and modern humans in the Levant had different resource gathering behaviours. Lieberman and Shea (1994)[17] analysed their lithic hunting technologies and found their differences to suggest Neanderthals hunted more frequently than modern humans. They argue that Neanderthals would have used their intensive hunting strategy to cope with a biodepleted environment, much like the Inuit of the 20th century used to depend largely on hunting. Henry et al. (2017)[18] found that in the Levant the site exploitation territory of Neanderthals was narrow, as they limited themselves to rugged woodlands, while modern humans of the Upper Palaeolithic would have been generalists with large exploitation territories in both the rugged woodlands and levelled steppe landscapes.

Although it does seem as if the environment favoured the initial success and eventual demise of Neanderthals, it is still unclear whether differences in their resource gathering strategies reflect differences in the environment, in anatomy, or in intelligence.

List of Southwest Asian Neanderthals edit

The remains of more than 70 Southwest Asian Neanderthals have been found. Sites are sorted west to east, first by country (westernmost site) then within countries. Important remains are in bold.

List of Southwest Asian Neanderthals
Present-day country (country of discovery) Site Principal Neanderthal finds MNI Geological age (ka) Descriptions Notes
Turkey Karain Four teeth 1 Senyürek (1949)[19][20]

Yalçınkaya (1988)[21][20]

Lebanon Ksâr 'Akil K2: Teeth and partial maxilla 1 Ewing (1963)[22] Ewing lost this specimen while transferring Ksar Akil material from Boston College to Fordham University.[23]
Lebanon El Masloukh Upper second molar[24] (1) ? Neanderthal attribution is stratigraphic, not morphological.[25]
Israel Kebara KMH1: 7-9 mo. old partial skel.

KMH2: Post-cranial adult ♂

Various fragments

KMH3: Milk tooth (m1-r)[26]
KMH4: 9 milk teeth, germ of 1 permanent tooth [27][26]
KMH5: Child mandibular symphysis fragment, no teeth[26]
KMH6: Right maxillary fragment with M1 and M2[26]
KMH7: Milk tooth (m?-r)[26]
KMH8: Milk tooth (m2-l)[26]
KMH9: Foot bone (4th right metatarsal)[26]
KMH10: Foot bone (1st toe distal phalanx)[26]
KMH11: Right clavicle fragment[26]
KMH12: Milk tooth (m?-r)[26]
KMH13: Milk tooth germ (m1-l)[26]
KMH14: Tooth (M2-l)[26]
KMH15: Milk tooth (m1-r)[26]
KMH16: Milk tooth (left i1)[26]
KMH17: Clavicle fragment[26]
(KMH18: Mandibular fragment with tooth [M2-r])[26]
(KMH19: Fragmentary tooth crown [M?-r])[26]
(KMH20: Parietal bone fragment)[26]
(KMH21: Germ of tooth [M1-l])[26]
(KMH22: Milk tooth [upper c-l])[26]
(KMH23: Milk tooth [i2-r])[26]
KMH24: Tooth (M3-l)[26]
(KMH25: 3 milk teeth germs [upper c-l, m1, m2])[26]
(KMH26: Tooth germ [i2-r])[26]
KMH27: Tooth (I2)[28][28]
KMH28: Tooth (I2)[28][28]
(KMH29: Milk tooth [i2-l])[26]
KMH30: Milk tooth (m1)[26][28]
(KMH31: Tooth [lower c])[28]

21 + (10) 64-59[29][30] KMH1: Smith et al. (1977)[31]

KMH2: Arensburg et al. (1985)[32]
KMH5-17, 24-31 : Tillier et al. (2003) [26]

Neanderthal attribution uncertain in KMH18-23, 25, 29, and 31[26][28]
West Bank (Mandatory Palestine) Shuqba S-D1: Tooth and cranial frags.[24] 1 Keith (1931)[33]
Israel (Mandatory Palestine) Tabun T C1: Nearly complete adult ♀

T C2: Toothed mandible missing I1 (♂)

Various fragments

T E1: Right femur shaft (♂?)
T E2: Tooth (M1 or M2, ♀?)
T C3: Right femur shaft (♀)
T C4: Distal right radius frag. (♀)
T C5: Right hamate bone
T C6: Right pisiform bone
T C7: Distal thumb phalanx
T B1: 10-11 year-old maxilla (♂?) with I2-r, M2-r
T BC2: Four teeth
(I2-l, M1-l, P3-r, M1-r)
T B3: One tooth (I2-r)
T B4: Four teeth
(I1-l, I2-l, M1-l, M3-r)
T B5: Two teeth (M2-l, M2-r)
T BC6: Two teeth (I1-l, M2-l)

15 ≈170-90


McCown (1936)
McCown and Keith (1939)
T C1: Neanderthal attribution is not universally accepted.[37]

As of 1975, the whereabouts of T BC2, B3, and BC6 are unknown.[24]: 146 

Israel Ein Qashish (EQH-2: Third molar)

EQH-3: Adult lower limbs

1 + (1)[38] 70-60[38] Been et al. (2017)[38] Discovered in 2013, these were the first diagnostically Neanderthal remains in Southwest Asia not found in a cave.[38]

EQH-2: 70% posterior probability that Neanderthal attribution is correct.[38]

Israel Shovakh (Tooth, M(3)-l[39][Note 2]) (1) S. Binford (1966)[40]

Trinkaus (1987)[39]

"[A]lthough within archaic and modern human ranges of variation, this complex occlusal morphology may suggest that it is more likely to have derived from a Neandertal than an early modern human". (Trinkaus 1987)[39]
Israel Amud A1: Adult full skeleton ♂

A2: Maxillary fragment

A7: 10-mo.-old partial skel.

3[Note 3][41] 61-53[41] A1: Suzuki et al. (1970)[42]

A7: Rak et al. (1994)[43]

Syria Dederiyeh D1: 19-30-month-old full skel.

D2: 21-30-month-old full skel.

17 D1: Akazawa et al. (1993)[44]

D2: Akazawa et al. (1999) [45]

Iraq Shanidar S1: Adult partial skel. ♂

S2: Adult crushed skel. ♂
S3: Post-cranial adult ♂
S4: Adult partial skel. (♂)
S5: Adult partial skel. (♂)
S6: Adult partial skel. (♀)
S7: 6-9-mo.-old crushed skel.
S8: Adult skeletal fragments (♀)
S9: 6-12-month-old vertebrae
S10: 17-25-month-old skel.

10 S2, S4: > 100

Others: 60

S1: Stewart (1959)[46]

S2: Stewart (1961)[47]
S3: Solecki (1960)[48]
S4: Stewart (1963)[49]
S5: Trinkaus (1977)[50]
Pomeroy et al. (2017)[51] S6: Same as S4
S7: Senyürek (1957)[52][53]
S8: Same as S4
S9: ?
S10: Cowgill et al. (2007)[54]

Shanidar 2 and 4 are sometimes not treated as Neanderthals.

All but Shanidar 3 and 10 (and fragments of 5 excavated in 2015-2016)[51] may have been destroyed in the 2003 invasion of Iraq.[55]

Iran Bawa Yawan Lower left deciduous canine 1 ~43,600-~41,500 years ago[56] Heydari-Guran et al (2021)[56]
Iran Wezmeh maxillary right premolar tooth 1 70-40[57] Zanolli et al. (2019)[57]
Iran Bisitun Adult radius shaft 1 Trinkaus and Biglari (2006)[58]
Total 71 + (13)
  1. ^ δ13C and δ18O are so-called isotopic signatures. They respectively measure the prevalence of the heavier carbon-13 (13C) and oxygen-18 (18O) isotopes over the more common carbon-12 and oxygen-16. C4-pathway plants have relatively more 13C (higher δ13C) than C3-pathway plants. Because C4 plants are better adapted to hot and dry climates than C3 plants, high δ13C values are associated with hot and dry environments. To the contrary, a greater prevalence of 18O (higher δ18O) in (land) animal bones reflects a colder environment: Because 16O is lighter than 18O, water molecules of the form H216O evaporate slightly more readily than those of the H218O form. This difference in evaporation is greater the colder the environment gets, so that rain and snow in colder regions have a lower δ18O value. Animals incorporate the δ13C and δ18O information of their environment and it is possible to extract it from their bones and teeth.
  2. ^ M?-r according to S. Binford (1966)[40].
  3. ^ Valladas, Mercier, and Froget (1999) write that "[o]f the remains of 18 individuals recovered at the Amud Cave, 15 were derived from unambiguous Middle Palaeolithic contexts, all of them located in the northern area of the excavation (Hovers et al., 1995). The stratigraphic distribution of these remains encompasses the layers B1 and B2, with only a single specimen derived from layer B4 (Sakura, 1970). Three individuals bear diagnostic characteristics which define them as Neanderthals. Amud I, the skeleton of an adult male was found at the top of layer B1, while the partial skeleton of the baby Amud 7 (Rak, Kimbet & Hovers, 1994) was recovered from the top of layer B2, just under the contact with the base of layer B1. Amud II, represented by a fragment of the right maxilla, was excavated from layer B2."

See also edit

Notes edit

References edit

  1. ^ a b E. Been; E. Hovers; R. Ekshtain; A. Malinski-Buller; N. Agha; A. Barash; D.E. Bar-Yosef Mayer; S. Benazzi; J.-J. Hublin; L. Levin; et al. (2017). "The first Neanderthal remains from an open-air Middle Palaeolithic site in the Levant". Scientific Reports. 7 (2958): 2958. Bibcode:2017NatSR...7.2958B. doi:10.1038/s41598-017-03025-z. PMC 5462778. PMID 28592838. 
  2. ^ A. Okladnikov (1949). "Исследование мустьерской стоянки и погребеня неандертальца в гроте Тешик-Таш, Южный Узбекистан (Средняя Азия)" [Investigation of the Mousterian site and burial of a Neanderthal in the cave of Teshik-Tash, Southern Uzbekistan (Central Asia)]. Тешик-Таш. Палеолитический человек (in Russian). Moscow. pp. 7–85.{{cite encyclopedia}}: CS1 maint: location missing publisher (link)
  3. ^ Michelle Glantz; Bence Viola; Patrick Wrinn; Tatiana Chikisheva; Anatoly Derevianko; Andrei Krivoshapkin; Uktur Islamov; Rustam Suleimanov; Terrence Ritzman (2008). "New hominin remains from Uzbekistan". Journal of Human Evolution. 55 (2): 223–237. doi:10.1016/j.jhevol.2007.12.007. PMID 18499228.
  4. ^ M.B. Mednikova (2011). "A proximal pedal phalanx of a Paleolithic hominin from Denisova Cave, Altai". Archaeology, Ethnology and Anthropology of Eurasia. 39 (1): 129–138. doi:10.1016/j.aeae.2011.06.017.
  5. ^ B. Viola; S.V. Markin; A.P. Buzhilova; M.B. Mednikova; A. Le Cabec; M.V. Shunkov; et al. (2012). "New Neanderthal remains from Chagyrskaya Cave (Altai Mountains, Russian Federation)". American Journal of Physical Anthropology. 147 (Suppl. 54): 293–294. doi:10.1002/ajpa.22033.
  6. ^ K. Prüfer; F. Racimo; N. Patterson; F. Jay; S. Sankararaman; S. Sawyer; A. Heinze; G. Renaud; P.H. Sudmant; C. de Filippo; et al. (2014). "The complete genome sequence of a Neanderthal from the Altai Mountains". Nature. 505 (7481): 43–49. Bibcode:2014Natur.505...43P. doi:10.1038/nature12886. PMC 4031459. PMID 24352235.(subscription required)
  7. ^ J. Hawks (2013). "Significance of Neandertal and Denisovan Genomes in Human Evolution". Annual Review of Anthropology. 42: 433–449. doi:10.1146/annurev-anthro-092412-155548.
  8. ^ a b Bar-Yosef O (1992). "The role of western Asia in modern human origins". Philosophical Transactions: Biological Sciences. 337 (1280): 193–200. Bibcode:1992RSPTB.337..193B. doi:10.1098/rstb.1992.0097. PMID 1357694.
  9. ^ Grün R., Stringer C., McDermott F., Nathan R., Porat N., Robertson S., McCulloch M. (2005). "U-series and ESR analyses of bones and teeth relating to the human burials from Skhul". Journal of Human Evolution. 49 (3): 316–334. doi:10.1016/j.jhevol.2005.04.006. PMID 15970310.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  10. ^ Frumkin A., Bar-Yosef O., Schwarcz H. P. (2011). "Possible paleohydrologic effect on hominin migration and occupation of the Levantine Middle Paleolithic". Journal of Human Evolution. 60 (4): 437–451. doi:10.1016/j.jhevol.2010.03.010. PMID 20541789.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  11. ^ a b c d Hallin K. A., Schoeninger M. J., Schwarcz H. P. (2012). "Paleoclimate during Neandertal and anatomically modern human occupation at Amud and Qafzeh, Israel: the stable isotope data". Journal of Human Evolution. 62 (1): 59–73. doi:10.1016/j.jhevol.2011.09.005. PMID 22115545.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  12. ^ Harrison, D. L., & Bates, P. J. J. (1991). Felidae. In The mammals of Arabia (2nd ed., pp. 156–172). Sevenoaks: Harrison Zoological Museum.
  13. ^ Rampino M. R., Self S. (1992). "Volcanic Winter and accelerated glaciation following the Toba super-eruption". Nature. 369 (6390): 50–52. Bibcode:1992Natur.359...50R. doi:10.1038/359050a0. S2CID 4322781.
  14. ^ a b c Shea, J. J. (2008). Transitions or turnovers ? Climatically-forced extinctions of Homo sapiens and Neanderthals in the East Mediterranean Levant. Quaternary Science Reviews, 1–18.
  15. ^ Tchernov, E. (1989). The Middle Paleolithic mammalian sequence and its bearing on the origin of Homo sapiens in the southern Levant. In O. Bar-Yosef & B. Vandermeersch (Eds.), Investigations in South Levantine Prehistory (pp. 25–42). Oxford: Archaeopress.
  16. ^ a b Belmaker M., Hovers E. (2011). "Ecological change and the extinction of the Levantine Neanderthals: implications from a diachronic study of micromammals from Amud Cave, Israel". Quaternary Science Reviews. 30 (21–22): 3196–3209. Bibcode:2011QSRv...30.3196B. doi:10.1016/j.quascirev.2011.08.001.
  17. ^ Lieberman D. E., Shea J. J. (1994). "Behavioral Differences between Archaic and Modern Humans in the Levantine Mousterian". American Anthropologist. 96 (2): 300–332. doi:10.1525/aa.1994.96.2.02a00030.
  18. ^ D.O. Henry; M. Belmaker; S.M. Bergin (2017). "The effect of terrain on Neanderthal ecology in the Levant". Quaternary International. 435 (Part A): 94–105. Bibcode:2017QuInt.435...94H. doi:10.1016/j.quaint.2015.10.023.
  19. ^ M. Şenyürek (1949). "Türk Tarih Kurumu Adına Yapılan Karain Kazısında Bulunan İki Fosil Dişe Dair Kısa Ön Rapor". Belleten (in Turkish). 52: 833–837. Cited in Aytek and Harvati (2016)
  20. ^ a b A.I. Aytek; K. Harvati (2016). "The Human Fossil Record from Turkey". In K. Harvati (ed.). Paleoanthropology of the Balkans and Anatolia: Human Evolution and its Context. Springer Netherlands.
  21. ^ I. Yalçınkaya (1988). "9. Kazı Sonuçları Toplantısı". 1986 Yılı Karain Kazıları (in Turkish). pp. 15–37. Cited in Aytek and Harvati (2016)
  22. ^ J.F. Ewing (1963). "A probable Neanderthaloid from Ksâr 'Akil, Lebanon". American Journal of Physical Anthropology. 21 (2): 101–104. doi:10.1002/ajpa.1330210202. PMID 14110686.
  23. ^ J.K. Williams; C.A. Bergman (2010). "Upper Paleolithic Levels XIII-VI (A and B) from the 1937-1938 and 1947-1948 Boston College Excavations and the Levantine Aurignacian at Ksar Akil, Lebanon". Paléorient. 36 (2): 117–161. doi:10.3406/paleo.2010.5391.
  24. ^ a b c K.P. Oakley; B.G. Campbell; T.I. Molleson (1975). Catalogue of fossil hominids: Part III. Americas, Asia, Australasia. London: British Museum (Natural History).
  25. ^ S. El Zaatari (2018). "The central Levantine corridor: The Paleolithic of Lebanon". Quaternary International. 466: 1–15. Bibcode:2018QuInt.466...33E. doi:10.1016/j.quaint.2017.06.047.
  26. ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab A.-M. Tillier; B. Vandermeersch; B. Arensburg; M. Chech (2003). "New human remains from Kebara Cave (Mount Carmel). The place of the Kebara hominids in the Levantine Mousterian fossil record". Paléorient. 29 (2): 35–62. doi:10.3406/paleo.2003.4764.
  27. ^ i1-r, i1-l, m1-r, i2-r, lower c-r and c-l, m1-r, m1-l, m2-r, M1
  28. ^ a b c d e f g A. Le Cabec; P. Gunz; K. Kupczik; J. Braga; J.-J. Hublin (2013). "Anterior tooth root morphology and size in Neanderthals: Taxonomic and functional implications". Journal of Human Evolution. 64 (3): 169–193. doi:10.1016/j.jhevol.2012.08.011. PMID 23266488.
  29. ^ H. Valladas; J.L. Joron; G. Valladas; B. Arensburg; O. Bar-Yosef; et al. (1987). "Thermoluminescence dates for the Neanderthal burial site at Kebara in Israel". Nature. 330 (6144): 159–160. Bibcode:1987Natur.330..159V. doi:10.1038/330159a0. S2CID 4308952.
  30. ^ H.P. Schwarcz; W.M. Buhay; R. Grün; H. Valladas; E. Tchernov; O. Bar-Yosef; B. Vandermeersch (1989). "ESR dating of the Neanderthal site, Kebara Cave, Israel". Journal of Archaeological Science. 16 (6): 653–659. doi:10.1016/0305-4403(89)90029-0.
  31. ^ P. Smith; B. Arensburg (1977). "A Mousterian skeleton from Kebara Cave". In B. Arensburg; O. Bar-yosef (eds.). Moshe Stekelis Memorial Volume. Jerusalem: The Israel Exploration Society. pp. 164–176.
  32. ^ B. Arensburg; O. Bar-Yosef; M. Chech; P. Goldberg; H. Laville; L. Meignen; Y. Rak; E. Tchernov; A.-M. Tillier; B. Vandermeersch (1985). "Une sépulture néandertalienne dans la grotte de Kébara (Israël)". Comptes Rendus de l'Académie des Sciences (in French) (300): 227–230.
  33. ^ A. Keith (1931). New Discoveries Relating to the Antiquity of Man. London: William and Norgate. pp. 204–208. Retrieved 15 February 2017.
  34. ^ A. Jelinek. "The Middle Paleolithic in Southern Levant, with comments on the appearance of modern Homo sapiens". In A. Ronen (ed.). The transition from Lower to Middle Paleolithic and the origin of modern man.
  35. ^ R. Grün; C.B. Stringer (2000). "Tabun revisited: Revised ER chronology and new ESR and U-series analyses of dental material from Tabun C1". Journal of Human Evolution. 39 (6): 601–612. doi:10.1006/jhev.2000.0443. PMID 11102271.
  36. ^ B. Vandermeersch. "Neanderthal Geographical and Chronological Variation". In S. Condemi; G.-C. Weniger (eds.). Continuity and Discontinuity in the Peopling of Europe: One Hundred and Fifty Years of Neanderthal Study.
  37. ^ A.-M. Tillier (2005). "The Tabun C1 skeleton: A Levantine Neanderthal?". Mitekufat Haeven: Journal of the Israel Prehistoric Society. 35: 439–450.
  38. ^ a b c d e E. Been; E. Hovers; R. Ekshtain; A. Malinski-Buller; N. Agha; A. Barash; D.E. Bar-Yosef Mayer; S. Benazzi; J.-J. Hublin; L. Levin; et al. (2017). "The first Neanderthal remains from an open-air Middle Palaeolithic site in the Levant". Scientific Reports. 7 (2958): 2958. Bibcode:2017NatSR...7.2958B. doi:10.1038/s41598-017-03025-z. PMC 5462778. PMID 28592838. 
  39. ^ a b c E. Trinkaus (1987). "The Upper Pleistocene Human Molar from Me'arat Shovakh (Mugharet Esh-Shubbabiq), Israel". Paléorient. 13 (1): 95–100. doi:10.3406/paleo.1987.4419.
  40. ^ a b S. Binford (1966). "Me'arat Shovakh (Mugharet esh-Shabbabiq)". Israel Exploration Journal. 16 (2): 96–103. JSTOR 27925048.
  41. ^ a b H. Valladas; N. Mercier; L. Froget (1999). "TL dates for the Neanderthal site of the Amud Cave, Israel". Journal of Archaeological Science. 26 (3): 259–268. doi:10.1006/jasc.1998.0334.
  42. ^ Suzuki, Hisashi; Takai, F. (1970). The Amud man and his cave site. Tokyo: Keigaku Publishing Co.
  43. ^ Y. Rak; W.H. Kimbel; E. Hovers (1994). "A Neandertal infant from Amud Cave, Israel". Journal of Human Evolution. 26 (4): 313–324. doi:10.1006/jhev.1994.1019.
  44. ^ T. Akazawa; Y. Dodo; S. Muhesen; A. Abdul-Salam; Y. Abe (1993). "The Neanderthal Remains from Dederiyeh Cave, Syria: Interim Report". Anthropological Science. 101 (4): 361–387. doi:10.1537/ase.101.361.
  45. ^ T. Akazawa; S. Muhesen; H. Ishida; O. Kondo; C. Griggo (1999). "New Discovery of a Neanderthal Child Burial from the Dederiyeh Cave in Syria". Paléorient. 25 (2): 129–142. doi:10.3406/paleo.1999.4691.
  46. ^ T.D. Stewart (1959). "Restoration and study of the Shanidar I Neanderthal skeleton in Baghdad, Iraq". Year Book of the American Philosophical Society for 1958: 274–278.
  47. ^ T.D. Stewart (1961). "The Skull of Shanidar II". Sumer. 17: 97–106.
  48. ^ R.S. Solecki (1960). "Three adult Neanderthal skeletons from Shanidar cave, northern Iraq". Annual Report of the Smithsonian Institution for 1959: 603–635.
  49. ^ T.D. Stewart (1963). "Shanidar skeletons IV and VI". Sumer. 19: 8–26.
  50. ^ E. Trinkaus (1977). "The Shanidar 5 Neanderthal skeleton". Sumer. 33: 35–41.
  51. ^ a b E. Pomeroy; M.M. Lahr; F. Crivellaro; L. Farr; T. Reynolds; C.O. Hunt; G. Barker (2017). "Newly discovered Neanderthal remains from Shanidar Cave, Iraqi Kurdistan, and their attribution to Shanidar 5" (PDF). Journal of Human Evolution. 111: 102–118. doi:10.1016/j.jhevol.2017.07.001. PMID 28874265.
  52. ^ M.S. Şenyürek (1957). "The skeleton of the fossil infant found in Shanidar cave, northern Iraq". Anatolia. 2: 49–55.
  53. ^ M.S. Şenyürek (1957). "A further note on the paleolithic Shanidar infant". Anatolia. 2: 111–121.
  54. ^ L.W. Cowgill; E. Trinkaus; M.A. Zeder (2007). "Shanidar 10: A Middle Paleolithic immature distal lower limb from Shanidar Cave, Iraqi Kurdistan". Journal of Human Evolution. 53 (2): 213–223. doi:10.1016/j.jhevol.2007.04.003. PMID 17574652.
  55. ^ B. Wood, ed. (2013). Wiley-Blackwell Encyclopedia of Human Evolution. Wiley-Blackwell. ISBN 978-1-118-65099-8.[page needed]
  56. ^ a b Heydari-Guran Saman; Benazzi Stefano; Talamo Sahra; Ghasidian Elham; Hariri Nemat; Oxilia Gregorio; Asiabani Samran; Azizi Faramarz; Naderi Rahmat; Safaierad Reza; Hublin Jean-Jacques; Foley Robert A.; Lahr Marta M. (2021). "The discovery of an in situ Neanderthal remain in the Bawa Yawan Rockshelter, West- Central Zagros Mountains, Kermanshah". PLOS ONE. 16 (8): 1-24. doi:10.1371/journal.pone.0253708. PMID 34437543.
  57. ^ a b Zanolli Clément; Biglari Fereidoun; Mashkour Marjan; Abdi Kamyar; Monchot Herve; Debue Karyne; Mazurier Arnaud; Bayle Priscilla; Le Luyer Mona; Rougier Hélène; Trinkaus Erik; Macchiarelli Roberto (2019). "Neanderthal from the Central Western Zagros, Iran. Structural reassessment of the Wezmeh 1 maxillary premolar". Journal of Human Evolution. 135: 102643. doi:10.1016/j.jhevol.2019.102643. PMID 31421316. S2CID 201057857.
  58. ^ E. Trinkaus; F. Biglari (2006). "Middle Paleolithic Human Remains from Bisitun Cave, Iran". Paléorient. 32 (2): 105–111. doi:10.3406/paleo.2006.5192.