Hominin fossils from Dmanissi

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"Skull 3", D 2700 (original)
The lower jaw, probably belonging to "Skull 3", D 2700, D 2735 (original)

The hominid fossils from Dmanisi are the oldest outside of Africa discovered fossils from the form circle of Hominini , the closest ancestor of man . Her remains have been excavated since 1991 under the direction of Dawit Lortkipanidze near Dmanissi (international: Dmanisi ) in Georgia and interpreted as belonging to the genus Homo . Fund layers in the footwall were an age of nearly 1.85 million years dated .

The hominine fossils of Dmanissi are considered to be a possible link between the earliest representatives of the genus Homo from Africa and the later fossils of Homo erectus known from Asia . They show that representatives of the genus Homo reached Eurasia 300,000 years earlier than previously assumed.

At first, however, it remained unclear which species of the genus Homo the Dmanissi finds are to be assigned to. In the year 2000 the fossils were first placed near Homo ergaster by their discoverers . In 2002 they were referred to as Homo georgicus in French journals , and from 2006 onwards, due to further bone finds, they were placed in the vicinity of Homo habilis and Homo erectus . In 2013, the attribution of the finds to Homo georgicus was finally revoked and the fossils identified as a local variant of a subspecies of Homo erectus and - combined with far-reaching reinterpretations of the relationship between the early homo species - referred to as Homo erectus ergaster georgicus .

Find history

Dmanissi, Georgia; Homo georgicus 1999 discovery map.png
GeorgieKaart met Dmanisi.png

The archaeological excavations on the Dmanissi Plateau in southern Georgia (located 1171  meters above sea level) were originally - from 1983 - an abandoned medieval city. In the course of these excavations it was found that the remains of much earlier settlements lie beneath the city; In addition to fossil mammalian bones, stone tools of the very original Oldowan type (Mode 1) were also discovered. During excavations by the Georgian Academy of Sciences in cooperation with the Roman-Germanic Central Museum , Antje Justus discovered a lower jaw in 1991 (inventory number D211; geodata of the site: 41 ° 20 ′ 10 ″  N , 44 ° 20 ′ 38 ″  E ). This fossil was presented to the public for the first time in the same year during a symposium at the Senckenberg Research Institute . Its age (1.8 to 1.6 million years) and belonging to Homo erectus remained controversial , despite its great resemblance to the lower jaw of the Nariokotome boy . A hominine foot bone of the same age discovered in 1997 did not provide any clarification either.

The area around the site

In May 1999, after heavy rainfall, a brain skull was visible in the ground, which presumably belonged to a young adult (inventory number D2280). Accompanying finds of teeth of the fossil rodent genus Mimomys attest an age of 1.6 to 2.0 million years, and with the help of the argon-argon method , the fossil-bearing strata of the earth could be dated to 1.8 to 2.0 million years; other dating methods indicated an age of 1.77 million years.

In the years that followed, a total of five well-preserved skulls, four of which with the associated lower jaw, as well as other hominine bones were recovered and scientifically described, including the particularly well-preserved "Skull 3" (D 2700, see illustration). "Skull 5" (archive number D 4500 with the associated lower jaw D 2600, known since 2000), was discovered in 2005 and was also unusually well preserved; it was not described in detail until 2013; its peculiarities not seen in any previously known homo- fossil - in particular the small inner cranial volume of only around 450 cm³ in combination with a protruding, i.e. pronounced prognathic snout and large teeth - prompted the authors of the study to delimit the species Homo ergaster and Homo erectus as well as Homo rudolfensis and Homo habilis .

The special significance of Dmanisi discoveries consists in the fact that different ages were found from the same period on an area of just 20 x 20 meters at least seven individuals, the statements on inter-individual variability of these homo - Population allow. The reasons why these individuals died simultaneously or almost simultaneously have not yet been unequivocally reconstructed. In February 2008, however, after examining 30 soil samples, French researchers published the assumption that it might have been a family that was surprised by a volcanic eruption and buried under a layer of ash; the age of the skeletons was now given as 1.81 ± 0.05 million years.

Find descriptions

1991: The first lower jaw

In the first description of the find from 1995, the lower jaw D211, discovered in 1991, was assigned to Homo erectus due to its age - 1.8 to 1.6 million years - as well as the fact that several features matched similarly old African and Asian fossil finds ; in 1998 it was assigned a proximity to Homo ergaster [ "Homo sp. indet. (aff. ergaster) " ] ascribed. Since no researcher had expected such an early appearance of representatives of the genus Homo in Eurasia , the dating was widely questioned; To some researchers, the shape of the jaw seemed too "modern" for such an old Homo erectus , so that they suspected that the lower jaw only got into the old layers of the soil much later - long after the death of the individual.

1999: Two well-preserved skulls

"Skull 2", D 2282 (reconstruction)

Only the two well-preserved skulls discovered in 1999 brought clarity. They were excavated two meters away from D211 and unequivocally dated to around 1.7 to 1.8 million years ago. One of the skulls D 2280, presumably belonging to an adult man, had an internal skull volume of only 775 cm³ ; parts of the skull and upper jaw including four teeth were preserved. The second skull D 2282, recovered from the same soil layer, presumably belonged to a young woman and had a volume of only 650 cm³ (for comparison: today's adult men have a volume of around 1500 cm³). Various other features of the skull were recognized as being consistent with the findings of Homo ergaster known from Africa . "Surprising" is the low level of correspondence between the finds, which are consistently referred to as "Dmanisi hominids", with later European and Asian homo finds.

2001: A homo ergaster with an extremely small brain?

Two other finds that were discovered in 2001 bore even less resemblance to the later representatives of the genus Homo : the "skull 3" D 2700 and the lower jaw D 2735 discovered a meter away from it; both fossils were excavated about 15 meters away from the known skull finds. The brain volume of only about 600 cm³ was significantly smaller than that of all previously known representatives of Homo erectus , it corresponded rather to the mean value of Homo habilis , so that Tim White , among others , pointed out that the Dmanissi finds might be a new support for the older one and the outdated theory that Homo habilis only developed into the Asian variant of Homo erectus outside of Africa . In addition to the small, evolutionarily " primitive " brain, the canines and the facial skull also turned out to be "original". In the scientific description of the Dmanissi finds, the authors explained in Science in July 2002 that the new finds could be interpreted - as before - as "representatives of Homo ergaster with an extremely small brain", but also as "the most primitive individuals to date, the erectus assigned "were. However, one could also argue that “this population is closely related to Homo habilis ”, as we know it from finds in Tanzania ( Olduvai Gorge ) and Kenya ( Koobi Fora ). According to the authors, the finds also show some anatomical features "which show a certain degree of isolation from related groups in Africa and the Far East". In particular, the lower jaw D2600, which had not yet been described and was excavated in September 2000, differs greatly from the known variants of Homo ergaster / Homo erectus . In 2014, an exact analysis of the teeth of this lower jaw was submitted, in which it was reported that all teeth are extremely worn, which suggests the consumption of a very hard and therefore largely abrasive vegetable diet, as is known from chimpanzees and gorillas , but not from comparably old finds of the genus Homo . In addition, the tooth roots show signs of inflammation.

2002: Homo ergaster becomes Homo georgicus

Find situation: fragments of the skull still embedded in the rock 5

Two months later, in September 2002, this suggestion of "isolation from related groups in Africa and the Far East" became the first description of the new species Homo by the doyen of Georgian paleoanthropology , Léo Gabounia and Abesalom Vekua, in a journal of the Académie des Sciences georgicus expanded; the lower jaw was named D2600 as the holotype . The fossils, now dated to an age of 1.81 ± 0.05 million years, were placed in the vicinity of Homo habilis , and it was postulated that they were the base group of all later representatives of Homo erectus in Europe and Asia.

In 2006, this assignment was confirmed, again in a French specialist journal, with reference to five skulls, four lower jaws and numerous other bone fragments. The age at the onset of death was also mentioned for the first time: an approx. 13 to 14 year old girl ("skull 3"), an 18 to 20 year old woman ("skull 2"), two male adults of approx. 25 up to 30 and 40 years of age ("skull 1" and "skull 5") and a toothless old man ("skull 4").

According to this publication, the Dmanissi skulls are not only small, but also relatively short and narrow. The frontal bone is less developed than in Homo erectus and has a noticeable narrowing behind the eye openings. The skull tops are flatter than in Homo erectus and Homo ergaster , but higher than in Homo habilis ; they are most comparable to Homo rudolfensis . With regard to the height and the transverse development of the skull central portion (in the parieto temporal region) are the Dmanisi specimens between habilis and ergaster . The hollow volume of the skull is also between these two homo species. The temporal bone is long and flat, the mastoid part is short. The upper part of the occiput is low and narrow, the crests of the skull thin and less developed than in the homo-erectus group. The upper temporal ridges are at a high point, and a torus angularis is present in the specimen that has been interpreted as a male adult. According to the study, the gracefulness of the face, the narrowness of the occiput and the pattern of their skull base distinguish the Dmanissi skulls from Homo erectus .

The orthognathic orientation of the face distinguishes the specimens of Dmanissi from early Pleistocene hominins ( Homo habilis and Homo ergaster ) as well as from the first Eurasian homo erectus finds; however, the subnasal region of the face is still protruding. The morphology of the middle part of the face, with a pronounced frontal nasal pillars (between the eye sockets and the nasal opening), an inframalar curvature of the zygomatic bone and an anterior root of the zygomatico maxillary ridge indicate a strong chewing load.

The bones in the area of ​​the shoulder joint socket are large and sharp-edged.

After weighing the skull features and skull dimensions as well as the other bone finds , the Dmanissi skulls lie according to the publication in the transition between the older Homo-habilis / Homo-rudolfensis group and the younger Homo ergaster , whereby they are the older (especially the Homo rudolfensis- find ER 1470) are anatomically closer. However, since the Dmanissi skulls can also be distinguished from Homo rudolfensis on the basis of many characteristics , they were placed in the new species Homo georgicus despite their assigned taxonomic proximity to this species .

The discovery of another very well preserved skull and lower jaw, described in Nature in 2005 , whose owner had lost all but one of the teeth years before his death, allowed conclusions to be drawn about the social behavior of the Dmanissi people . Numerous accompanying finds (stone tools and bones with indentations from stone tools) as well as the climatic conditions 1.8 million years ago have been interpreted to mean that the Dmanissi people - at least in winter - mainly ate meat. The tooth sockets of this oldest, toothless hominine skull therefore lead to the conclusion that the individual - although he could no longer chew coarse food - must have been provided with severely chopped food and, despite his disability, must have been socially integrated.

In 2004, a statistical analysis of the internal volume of the skull showed that the previously known finds can be attributed to a single species despite very different volumes.

2006: Instead of Homo georgicus "Dmanisi hominins"

Dawit Lortkipanidze (2018)
View over the fossil site (September 2007)

In connection with the naming, it is noteworthy that Dawit Lortkipanidze , who was involved in the discovery of the first lower jaw as excavation manager as early as 1991, was co-author of the assignment of all Dmanissi finds to the species Homo georgicus . In the descriptions of the finds published by him as the main author in Nature , however, he did not use this species name chosen by his former superior Léo Gabounia, but instead described the finds in 2007 as, for example, “Dmanisi hominins”, which “is largely comparable with the earliest Homo (that is Homo habilis ) “Be. In addition, in a detailed study in 2006, the skulls were so named, placed between Homo habilis and Homo erectus based on their anatomical features and cautiously placed at the base of that homo population "from which Homo erectus evolved ."

In fact, the isolation of the Dmanissi finds from the species Homo georgicus - outside of the French-language literature - was not able to establish itself in specialist science. For example, Winfried Henke noted in an overview article on “Origin and Distribution of the Homo Genus ” that the species status propagated by Vekua and Gabunia is provisional. There were reservations about defining a new species mainly because it remained unclear whether it was defined in terms of a chronospecies , a morphospecies or even a biospecies . Lortkipanidze himself cautiously referred to the species name as "a proposal", the viability of which will only be proven on the basis of further finds.

In 2007, Lordkipanidze published in Nature the description of more than 30 bones and bone fragments from the shoulder girdle , spine , upper arms , thighs and lower legs , which were recovered in Dmanissi between 2003 and 2005. Some of these postcranial finds could be assigned to the skulls already known and for the first time provided information about the presumed external appearance of the Dmanissi people. Based on these fossils, their body weight was estimated at 40 to 50 kg, their height at around 145 to 166 cm and their brain volume at almost half that of modern humans. Even in comparison to her small body size, her brain was as small as that of the very first representatives of the genus Homo from Africa and significantly smaller than the brain of the previously known finds of Homo erectus . On the basis of some African homo erectus finds - among other things after the discovery of the so-called Nariokotome boy - the paleoanthropologists suspected that the first non-African representatives of the genus Homo were much larger.

In contrast, the Dmanissi people already had body proportions similar to those of modern humans: their legs were much longer than their arms, and their thighs were longer than their upper arms. Found foot bones were interpreted to mean that they were able to walk on a springy, two-legged walk and were good runners thanks to an arch of the foot. The anatomy of the shoulders and arms, however, differed from modern humans: For example, when the forearms and hands were in a resting position - with the upper arms hanging down - they did not point towards the body, but forward. The construction of their arms probably made it easier for them to climb trees.

  • "Skull 1"
    (replica)
    adult man, found in 1999, brain volume 775 cm³

  • "Skull 2"
    (replica)
    young woman,
    jaw found 1991, skull found 1999, brain volume 650 cm³

  • "Skull 3"
    (replica)
    female adolescents, found in 2001, brain volume 600 cm³

  • "Skull 4"
    (replica)
    toothless elderly man, found in 2002/2003, brain volume 625–650 cm³

  • "Skull 5"
    (originals)
    adult man, skull found 2005, jaw found 2000, brain volume 546 cm³

2013: Homo erectus ergaster georgicus

In a study published in Science in October 2013 , the researchers of the hominine Dmanissi fossils finally revoked the attribution of the finds to Homo georgicus by formally withdrawing the designation of the lower jaw D2600 as a holotype of the species.

They saw themselves prompted to do so because of the characteristics of "Skull 5" (archive number D 4500 with associated lower jaw D 2600), which on the one hand shows typical characteristics of the genus Homo ; on the other hand, "Skull 5" has only a very small internal volume of the skull with 450 cm³, but a strongly pronounced snout and large teeth - a combination that was previously not known from any homo- fossil, nor from the other skull finds from Dmanissi. From the synopsis of all five skull finds, the researchers led by David Lordkipanidze therefore now derived that the early representatives of the genus Homo apparently had a very large, previously unknown, inter-individual phenotypic variability , comparable to the variability in people living today and with the chimpanzees living today . The variability evidenced by the Dmanissi fossils is so great, according to the study, that both the characteristics of the comparably old fossil finds from Africa and Asia do not exceed the limits of this variability; literally the researchers wrote:

"Viewed from the Dmanissi point of view, the morphological diversity of the approximately 1.8 million year old, African homo- fossils reflects the variation between local populations of a single evolving lineage , appropriately named H. erectus ."

The researchers suggested further specifying the nomenclature : The African fossils separated from Homo erectus by researchers other than Homo ergaster should be reunited with Homo erectus erectus against the background of the Dmanissi findings as Homo erectus ergaster - that is, as an early subspecies . In this context, the hominine fossils from Dmanissi were formally identified as Homo erectus ergaster georgicus , combined with the note that the addition of georgicus refers to the origin of the local Georgian ancestry. In addition, the new view of the variability of the shape of Homo erectus could also have an impact on the delimitation of Homo habilis and Homo rudolfensis from Homo erectus : It may be appropriate to also assign those fossils to the ancestral group erectus that were previously ascribed to these two species.

There was also opposition to the designation Homo erectus ergaster georgicus ; the structure of the teeth, the lower jaw and the skull suggest “taxonomic diversity”, and that speaks in particular for the validity of the previous name Homo georgicus . In addition, according to Ian Tattersall , the morphological features of skull 5 differ so greatly from the features of the other four skulls that the findings - had they been recovered from different locations - would be assigned to two species. In addition, there are no traceable derived characteristics between the Dmanissi finds and the type specimen of Homo erectus , the Trinil 2 skull roof from Java , so that the assignment to this species is taxonomically unfounded and only owed to the fact that the finds from Dmanissi are similar as old as the finds from Java.

The scientific significance of the finds

Oldowan tool from Dmanisi (right) next to it for comparison, a hand ax the Acheulean

The hominine fossils of Dmanissi refute the assumption, which was previously derived from African pre- human finds and generally accepted in paleoanthropology, that the first people who emigrated from Africa had a brain volume of at least 1000 cm³, were about 170 cm in size and had advanced cultural techniques . By Bernard Wood as a transitional form (transitional form) called combination of " anatomically original " and " anatomically modern " features (very small brain, on apes referring shoulders and arms, but of Homo sapiens barely distinguishable legs) demonstrate for the first time in a uniform Fund- The sequence of the evolutionary processes from the archaic representatives of the genus Homo to modern humans. During a visit to the Senckenberg Research Institute in November 2007, the discoverer of the fossils, Dawit Lortkipanidze, pointed out the amazing similarities between the Dmanissi finds and Homo floresiensis .

For the first time with such ancient human fossils, statements can also be made about the range of variation in anatomical features in a population, which could lead to a reassessment of the delimitation of species that - such as Homo erectus and Homo habilis - were often only defined on the basis of a single lower jaw . With reference to the Dmanissi fossils, the assumption that Homo erectus developed from Homo habilis in Africa was also questioned ; Homo erectus may not have developed until north of the Levant , and parts of the population later migrated back to Africa, others towards Asia and Europe.

literature

  • Gerhard Bosinski , David Lordkipanidze, Konrad Weidemann: The ancient Paleolithic site of Dmanisi (Georgia, Caucasus). Yearbook of the Roman-Germanic Central Museum. Bonn 42, 1995, ISSN  0076-2741 , pp. 21-203.
  • Léo K. Gabunia et al .: New hominid finds from the ancient Paleolithic site of Dmanisi (Georgia, Caucasus) in the context of current excavation results. In: Archaeological correspondence sheet. 29, 1999, pp. 451-488.
  • Léo K. Gabunia et al .: New prehistoric man finds from Dmanisi (East Georgia). Yearbook of the Roman-Germanic Central Museum Mainz. 46, 2000, pp. 23-38.
  • Léo K. Gabunia et al .: Découverte d'un nouvel hominid à Dmanissi (Transcaucasie, Géorgie). In: Comptesrendus de l'Académie des sciences Paris. Palevol 1, 2002, pp. 243-253.
  • T. Garcia: Cadres stratigraphique, magnétostratigraphique et géochronologique des hominidés fossiles du site de Dmanissi en Géorgie . Muséum national d'histoire naturelle, Paris 2004.
  • Olaf Jöris: The ancient Paleolithic site of Dmanisi (Georgia, Caucasus). Monographs of the Roman-Germanic Central Museum. Volume 74. Schnell and Steiner, Regensburg 2008, ISBN 978-3-7954-2140-3 .
  • G. Philip Rightmire and David Lordkipanidze: Fossil Skulls from Dmanisi: A Paleodeme Representing Earliest Homo in Eurasia. In: John G. Fleagle et al .: Out of Africa I. The First Hominin Colonization of Eurasia. Springer, 2010, pp. 225-243, ISBN 978-94-007-3308-4 .
  • Ann Gibbons: The Wanderers. In: Science. Volume 354, No. 6315, 2016, pp. 958–961, doi: 10.1126 / science.354.6315.958 .

Web links

Individual evidence

  1. ↑ a b c Reid Ferring et al .: Earliest human occupations at Dmanisi (Georgian Caucasus) dated to 1.85–1.78 Ma. In: PNAS . Volume 108, No. 26, 2011, pp. 10432-10436, doi : 10.1073 / pnas.1106638108
  2. ^ A b G. Philip Rightmire, David Lordkipanidze and Abesalom Vekua: Anatomical descriptions, comparative studies and evolutionary significance of the hominin skulls from Dmanisi, Republic of Georgia. In: Journal of Human Evolution. Volume 50, No. 2, 2006, pp. 115-141, doi: 10.1016 / j.jhevol.2005.07.009
  3. ^ Ann Gibbons: First globetrotters had primitive toolkits. In: Science . Volume 323, No. 5917, 2009, p. 999, doi: 10.1126 / science.323.5917.999a , ISSN  0036-8075
  4. ↑ a b c d e David Lordkipanidze et al .: A Complete Skull from Dmanisi, Georgia, and the Evolutionary Biology of Early Homo. In: Science . Volume 342, No. 6156, 2013, pp. 326–331, doi: 10.1126 / science.1238484
  5. ^ Leo Gabunia, Abesalom Vekua: A Plio-Pleistocene hominid from Dmanisi, East Georgia, Caucasus. In: Nature . Volume 373, 1995, pp. 509-512, doi: 10.1038 / 373509a0
  6. ↑ 4th International Senckenberg Conference "100 Years of Pithecanthropus - The Homo-erectus Problem"
  7. ↑ Ann Gibbons: jawing with our Georgian ancestors. In: Science. , Volume 255, 1992, p. 401, doi: 10.1126 / science.255.5043.401
  8. ↑ The key fossils are dated by Ralf-Dietrich Kahlke from the Research Station for Quaternary Paleontology Weimar of the Senckenberg Society for Nature Research , see also: Jena paleontologists are preparing excavation project in Georgia. On: idw-online from August 6, 1998
  9. ↑ Michael Balter, Ann Gibbons: A Glimpse of Humans' First Journey Out of Africa. In: Science. Volume 288, 2000, pp. 948-950, doi: 10.1126 / science.288.5468.948 , ISSN  0036-8075
  10. ↑ "Skull 5" (D 4500 + D 2600) is the best preserved adult skull from the period between 2.6 - 0.9 million years ago. Source: Fred Spoor: Small-brained and big-mouthed . In: Nature. Volume 502, No. 7472, 2013, pp. 452-453, doi: 10.1038 / 502452a
  11. ^ Léo K. Gabunia, Abesalom Vekua, Marie-Antoinette de Lumley, David Lordkipanidze: A New Representative of Homo from the Lower Pleistocene of Dmanisi. In: Archeology, Ethnology and Anthropology of Eurasia. Volume 4, No. 12, 2002, pp. 145-153.
  12. ^ Marie-Antoinette de Lumley et al .: Impact probable du volcanisme sur le décès des Hominidés de Dmanissi. In: Comptes Rendus Palevol. Volume 7, No. 1, 2008, pp. 61-79, doi: 10.1016 / j.crpv.2007.09.002 ISSN  1631-0683
  13. ^ L. Gabunia, A. Vekua: A Plio-Pleistocene hominid from Dmanisi, East Georgia, Caucasus. In: Nature. Volume 373, 1995, pp. 509-512, doi: 10.1038 / 373509a0 , ISSN  0028-0836
  14. ^ Antonio Rosas and José María Bermúdez de Castro : On the taxonomic affinities of the Dmanisi mandible (Georgia). In: American Journal of Physical Anthropology. Volume 107, No. 2, 1998, pp. 145-162, doi : 10.1002 / (SICI) 1096-8644 (199810) 107: 2 <145 :: AID-AJPA2> 3.0.CO; 2-U
  15. ↑ Leo Gabunia, Abesalom Vekua, David Lordkipanidze et al .: Earliest Pleistocene Hominid Cranial Remains from Dmanisi, Republic of Georgia. Taxonomy, Geological Setting, and Age. In: Science. Volume 288, 2000, pp. 1019-1025, doi : 10.1126 / science.288.5468.1019 ISSN  0036-8075
  16. ↑ Abesalom Vekua, David Lordkipanidze et al .: A new skull of early Homo from Dmanisi, Georgia. In: Science. Volume 297, 2002, pp. 85-89, doi : 10.1126 / science.1072953 ISSN  0036-8075
  17. ↑ Michael Balter, Ann Gibbons: Were 'little people' the first to venture out of Africa? In: Science. Volume 297, 2002, pp. 26-27, doi : 10.1126 / science.297.5578.26 ISSN  0036-8075
  18. ↑ Laura Martín-Francés et al .: Palaeopathology of the Pleistocene specimen D2600 from Dmanisi (Republic of Georgia). In: Comptes Rendus Palevol. Advance online publication of January 18, 2014, doi: 10.1016 / j.crpv.2013.10.007
  19. ↑ Léo Gabounia, Marie-Antoinette de Lumley, Abesalom Vekua, David Lordkipanidze, Henry de Lumley: Découvert d'un nouvel hominidé à Dmanissi (Transcaucasie, Géorgie). In: Comptes Rendus Palevol. Volume 1, 2002, pp. 243-253, doi : 10.1016 / S1631-0683 (02) 00032-5 ISSN  1631-0683 and - with illus . - Sciencemag
  20. ↑ Marie-Antoinette de Lumleya, David Lordkipanidze: L'Homme de Dmanisi (Homo georgicus), il ya 1,810,000 ans. In: Comptes Rendus Palevol. Volume 5, 2006, pp. 273–281, doi : 10.1016 / j.crpv.2005.11.013 ISSN  1631-0683 - literally it says here: “Two new concepts can be retained: - the exodus from Africa took place earlier than previously thought, dating back to at least 1.8 Myr ago. It was carried out by Homo georgicus , a group close to Homo habilis ; - it is no longer valid to base explanations of Man's migratory capacity in terms of cranial development. "
  21. ↑ literally: "une adolescente d'environ 13-14 ans, une adulte jeune femelle de 18-20 ans, deux adultes mâles âgés de 25-30 ans et 40 ans et un vieillard édenté." In: Marie-Antoinette de Lumley, Léo Gabounia, Abesalom Vekua, David Lordkipanidze: Les restes humains du Pliocène final et du début du Pléistocène inférieur de Dmanissi, Géorgie (1991-2000). I - Les crânes, D 2280, D 2282, D ​​2700. In: L'Anthropologie. Volume 110, 2006, pp. 1–110, doi : 10.1016 / j.anthro.2006.02.001 ISSN  0003-5521
  22. ↑ David Lordkipanidze, Abesalom Vekua et al .: The earliest toothless hominin skull. In: Nature. Volume 434, 2005, pp. 717-718, doi : 10.1038 / 434717b ISSN  0028-0836
  23. ↑ David Lordkipanidze, Abesalom Vekua: A fourth hominin skull from Dmanisi, Georgia. In: The Anatomical Record Part A: Discoveries in Molecular, Cellular, and Evolutionary Biology . Volume 288A, 2006, No. 11, pp. 146-1157, doi : 10.1002 / ar.a.20379 ISSN  0003-276X
  24. ↑ Sang ‐ Hee Lee: Brief communication: Is variation in the cranial capacity of the Dmanisi sample too high to be from a single species? In: American Journal of Physical Anthropology. Volume 127, No. 3, 2005, pp. 263-266, doi: 10.1002 / ajpa.20105
  25. ↑ David Lordkipanidze, Abesalom Vekua et al .: Anthropology: The earliest toothless hominin skull. In: Nature. Volume 434, 2005, pp. 717-718, doi : 10.1038 / 434717b , ISSN  0028-0836
  26. ↑ David Lordkipanidze et al .: Postcranial evidence from early Homo from Dmanisi, Georgia. In: Nature. Volume 449, 2007, pp. 305-310, doi : 10.1038 / nature06134 , ISSN  0028-0836
  27. ↑ David Lordkipanidze, Abesalom Vekua, Philip G. Rightmire, and Ann Margvelashvili: Hominid Fossils from Dmanisi and Their Place Among the Early Hominids. In: Bulletin of the Georgian National Academy of Sciences. Volume 175, No. 1, 2007, pp. 104-111, full text
  28. ↑ In: Günther A. Wagner et al .: Homo heidelbergensis. Key find in human history. Konrad Theiss, Stuttgart 2007, p. 191, ISBN 978-3-8062-2113-8
  29. ↑ Joint lecture with Friedemann Schrenk on November 21, 2007 at the Senckenberg Research Institute , Frankfurt am Main
  30. ↑ David Lordkipanidze et al .: Postcranial evidence from early Homo from Dmanisi, Georgia. In: Nature. Volume 449, 2007, pp. 305-310, doi : 10.1038 / nature06134 , ISSN  0028-0836
  31. ^ Ian J. Wallace, Brigitte Demes, William L. Jungers , Martin Alvero and Anne Su: The bipedalism of the Dmanisi hominins: Pigeon-toed early Homo? In: American Journal of Physical Anthropology. Volume 136, No. 4, 2008, pp. 375-378, doi: 10.1002 / ajpa.20827
  32. ↑ The first Europeans 1.7 million years ago could walk and walk like us. Explanations by Christoph Zollikofer on idw-online from September 19, 2007; Illustration of the skeletal parts
  33. ↑ In the original (p. 330): "When seen from the Dmanisi perspective, morphological diversity in the African fossil Homo record around 1.8 Ma probably reflects variation between demes of a single evolving lineage, which is appropriately named H. erectus."
  34. ↑ In the original (p. 330): “It remains to be tested whether all of the fossils currently allocated to the taxa H. habilis and H. rudolfensis belong to a single evolving Homo lineage. Although we regard this null hypothesis as parsimonious and fully compatible with new evidence from Dmanisi, alternative scenarios exist. ”
    See: eurekalert.org of October 17, 2013: Unique skull discovery refutes early human biodiversity.
  35. ↑ The family tree is shrinking. In. Süddeutsche Zeitung of October 18, 2013, p. 16, full text
  36. ↑ Jeffrey H. Schwartz , Ian Tattersall and Zhang Chi: Comment on "A Complete Skull from Dmanisi, Georgia, and the Evolutionary Biology of Early Homo". In: Science. Volume 344, No. 6182, 2014, p. 360, doi: 10.1126 / science.1250056
  37. ^ Ian Tattersall: The Strange Case of the Rickety Cossack - and Other Cautionary Tales from Human Evolution. Palgrave Macmillan, New York 2015, pp. 175-177, ISBN 978-1-137-27889-0
  38. ↑ For example, The Cambridge Encyclopedia of Human Evolution (1992) names individuals resembling the Nariokotome boy as the first colonizers in Europe and Asia
  39. ↑ Ann Gibbons: A New Body of Evidence Fleshes Out Homo erectus. In: Science. Volume 317, No. 5845, 2007, p. 1664, doi : 10.1126 / science.317.5845.1664 , ISSN  0036-8075
  40. ↑ Joint lecture with Friedemann Schrenk on November 21, 2007
  41. ↑ Ann Margvelashvili, Christoph PE Zollikofer, David Lordkipanidze et al .: Tooth wear and dentoalveolar remodeling are key factors of morphological variation in the Dmanisi mandibles. In: PNAS. Volume 110, No. 43, 2013, pp. 17278–17283, doi: 10.1073 / pnas.1316052110
  42. ^ G. Philip Rightmire, Ann Margvelashvili and David Lordkipanidze: Variation among the Dmanisi hominins: Multiple taxa or one species? In: American Journal of Physical Anthropology. Volume 168, No. 3, 2019, pp. 481-495, doi: 10.1002 / ajpa.23759
  43. ↑ Robin Dennell , Wil Roebroeks : An Asian perspective on early human dispersal from Africa. In: Nature. Volume 438, 2005, pp. 1099-1104, doi : 10.1038 / nature04259
  44. ↑ Meet the frail, small-brained people who first trekked out of Africa. On: sciencemag.org of November 22, 2016
This version was added to the list of articles worth reading on November 29, 2007 .