Tribal history of man

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Kladogramm the family of great apes (hominids) and its subfamilies Ponginae and Homininae and the still extant genera Pongo (orangutans) , Gorilla , Pan (chimpanzees) and Homo

As human evolution that is through evolution -related emergence of modern humans ( Homo sapiens ) and its closest relative of the common ancestor called. According to today's view, human tribal history began with the splitting up of the last common ancestral population of chimpanzees and humans. That of the two sub-populations from which humans emerged, as well as all of their extinct and still living descendants are referred to as hominini .

Research into tribal history "takes into account all fossil and current forms of life of the apes in the context of the doctrine of primates , primatology or primatology founded by TH Huxley as early as 1863 ".

A steadily growing number of well-preserved fossils from different regions of Africa , Asia and Europe has expanded knowledge about the ancestors of man, but has not led to any agreement about their lifespan and distribution areas . This is also due to the fact that the systematic allocation of numerous individual finds and the relationships of many prehistoric species are controversial, because research is currently "at the limits or even beyond the analytical capabilities of the data and the available methods".

The types of hominini are often referred to as follows: the australopithecines as pre-humans ; Homo habilis and Homo rudolfensis as prehistoric men ; all later species of the genus Homo (except Homo sapiens ) as early humans ; Homo sapiens as now man or anatomically modern man . Occasionally the hominini are preceded by the animal-human transition field named in 1958 by Gerhard Heberer .

Molecular biological and paleoanthropological findings on the origin of humans

On the basis of individual fossil finds, the existence of species that can be distinguished from one another can be proven for early humans . The exact point in time at which the body structure of individuals of a certain species has changed so much through evolution that their populations can be assigned to a later species can only be roughly estimated using individual fossils. This is due to the fact that a smooth transition from the original characteristics to the newly acquired characteristics can be expected. A sharp demarcation proves to be difficult and often completely impossible because of the gaps in the fossil evidence.

The so-called molecular clock , with the help of DNA analyzes in individuals of species living today , has proven to be an important tool for approximating the times at which species split up . “Today it is assumed that every person has around 50 base changes (mutations) that have arisen in the parents' germ line and were therefore not part of the parental genome. This results in a mutation rate of around 50 changes per generation per genome. Assuming that around 25 years pass per generation, the genetic differences between two populations can be used to calculate when they must have separated from each other. ”The results of such assessments are - not only for the early phase of primate development - in specialist circles However, it is controversial because the exact speed of the molecular clock, i.e. the frequency of mutations in past epochs, is unknown and its calibration depends on reliably dated fossil finds.

A time scale for the evolution of primates based on molecular biological models was first published in 1967; In a recalculation in 2012, this was clearly shifted to the older age, especially for the development of Homo sapiens .

Skulls of humans , chimpanzees , orangutans and macaques with information on the mean brain weight

The points in time calculated using the molecular clock and the points in time for the splitting of lines of development revealed by fossil finds also often differ by several million years. In 1985, based on DNA analyzes, a study was published according to which in the Cretaceous Period around 90 million years ago those lineages emerged from the Euarchontoglires that led to today's mice and primates . This dating was confirmed in 2009 and a time span of 80 to 116 million years was given for the split. On the other hand, based on fossil finds, a point in time around 56 million years ago - in the Paleocene - is most likely. A possible explanation for this large discrepancy was expelled in 2012 that the then living diminutive primates (the today's lemurs resembled) a shorter sequence of generations and therefore a higher mutation rate than did large-growing primates, since the majority of mutations in germ cells during the replication occurs ; the increase in the size of the primate species since the Paleocene was therefore presumably accompanied by a reduction in the "extraordinarily rapid" mutation rate in the earliest primates.

For the time of the splitting of the Old World monkeys into the humans and the vervet monkeys (this includes vervet monkeys , baboons and macaques ) around 23 million years ago, at the beginning of the Miocene , were calculated by DNA analysis . The human-like parted this date, according to some 15 million years ago in the great apes and gibbons on. The time of the split of the great apes into the Asian species (the ancestors of the orangutans ) and the African species was calculated as 11 million years, for the separation of the gorillas from the chimpanzees around 6.5 million years and for the separation of the chimpanzees from the hominini finally 5.2 ± 1.1 million years.

The following examples show how provisional these times are:

  • The "in earlier studies" based on mutation rates after decoding the gorilla genome sequence for the separation of the gorillas from the chimpanzees 5.95 million years; With reference to fossil finds, this dating was relativized and - based on the mutation rates in Homo sapiens populations living today - a lower mutation rate was assumed, which resulted in a separation of these lines of development 10 to 6 million years ago.
  • On the basis of fossil finds, Terry Harrison dated the separation of the chimpanzees from the hominini at the beginning of 2010 to 7.5 million years ago, while C. Owen Lovejoy dated this separation in 2009 to about 6 to 5 million years ago. After a revision of the assumptions about the frequency of mutations, a separation 8 to 7 million years ago was calculated again in 2012.
  • In 2013 and 2019, the authors of several studies came to the conclusion that the evolution rates were only half as high as numerous earlier publications assumed.

Only the sequence of the "branches" in the human family tree is currently considered certain, but not the time interval between two branches.

Cladogram and time periods of the splitting of the Euarchontoglires into Euarchonta and Glires as well as into the Primatomorpha for the following taxa (according to Janečka et al., 2007):
Rodents : Tamias (chipmunk) - Castor (beaver) - Pedetes (jumping
hare ) Hares : Sylvilagus ( cottontail rabbit) - Ochotona (Whistling hares )
Giant gliders : Galeopterus - Cynocephalus
Primates : Lemurs - Microcebus (mouse lemurs ) - Otolemur (giant galagos) - Ateles (spider monkeys) - Macaca (macaques) - Homo
pointed squirrels : Tupaias - Urogale (Philippine pointed squirrel) - Ptilocercus ( ptilocercus

Advancement of the human

The differentiation of the human-like : To the border from the Eocene to Oligocene separates in the form of district Old World monkeys (catarrhines) leading to the Cercopithecoidea development line (this includes the Cercopithecidae ) from that of the human-like (Hominoidea). The human taxon includes the gibbons (Hylobatidae), the orangutans (Ponginae) and the African great apes (homininae: gorillas , chimpanzees , humans).
Reconstruction of Proconsul's physique

“The cradle of the hominoidea” - the superfamily of the human species - “lies in the early Miocene of East Africa ”, that is, from 23 to 16 million years ago. Their early Miocene forms are known as archaic or stem hominoids ; their relationship to one another and to later species is controversial because of the small number of known fossils. Furthermore, none of the previously known species of Miocene humans have the physical characteristics of the locomotion apparatus typical of great apes, "which apparently only developed with the increasing terrestrial way of life of the pliocene hominoid", that is around 5 million years ago.

Years ago, some 18 to 15 million, two lines of development separated: one led to the Gibbons (Hylobatidae), the other to some already in the Miocene back extinct species - as Afropithecus , kenyapithecus and Griphopithecus , Pierolapithecus , Dryopithecus and Oreopithecus - as well as the great apes , that is, to the orangutans (Ponginae) and to the African great apes (Homininae: gorillas , chimpanzees , humans).

It is assumed by some of the specialist authors that the great apes could have emerged from the superfamily of the Proconsul-like (Proconsuloidea) existing in this epoch , for which Proconsul was the namesake and which , according to the paleontological findings, “was an extraordinarily richly shaped taxon that exclusively occurred in tropical forests of Africa and the Arabian Peninsula ”. Other specialist authors consider a sister group of the proconsul-like to be more likely or completely dispense with speculations: "According to the current state of knowledge, however, none of the known Miocene hominoid taxa can be placed in the direct common ancestry of African great apes and the homininin line."

First colonization of Eurasia

Between 17 and 14 million years ago, the climate in Africa became drier, and at the same time there were greater temperature fluctuations over the course of the year. According to the current state of research, these changes in ecological conditions reduced the Proconsul-like diversity; at the same time, early relatives of the vervet monkey relatives and the humans (for example Kenyapithecus wickeri , Equatorius africanus and Nacholapithecus ) developed into the predominant genera. The fossils from this epoch show that adaptations to harder foods were developed: e.g. B. strong jaws as well as thick enamel layers on the molars and that 16 to 15 million years ago, areas outside Africa - large parts of Eurasia - were colonized by these species for the first time .

Around 16 million years ago, around the border from the early to the middle Miocene, the relatives of the orangutans in Asia separated from that of the other great ape species ; The family group of orangutans also includes the extinct genera Ramapithecus , Sivapithecus / Ankarapithecus , Lufengpithecus and Khoratpithecus as well as Gigantopithecus . Ramapithecus , whose first fossils were found in northern India in the early 1930s, was mistakenly believed to be the ancestor of the hominini and thus of humans in the 1960s and 1970s.

The lower jaw of Dryopithecus fontani

The number of human-like species in Europe increased 13 to 9 million years ago. Among other things, at least four Dryopithecus species are known from this period in addition to the two finds of Pierolapithecus and Anoiapithecus discovered in Spain . A comparable development took place in Asia in the type circle of which the only survivors are the orangutans.

9.6 million years ago, the so-called Vallesium Crisis led to considerable changes in ecosystems in Europe: starting in the Spanish Mediterranean region, the subtropical , evergreen forests in western and central Europe disappeared as a result of gradual cooling; in their place followed deciduous trees and, in some southern regions, steppes . This climatic change also brought about a profound change in the animal world - most of the European human species of this epoch became extinct; only Oreopithecus survived until about 7 or even 6 million years ago on an island in the region of Sardinia / Corsica . Furthermore, fossil footprints that are 5.7 million years old on Crete have been interpreted as evidence of a native great ape species, which can not be Oreopithecus because of the forward-facing position of the toes with the big toe and ball of the foot (coordinates of the site: 35 ° 30 ′ 55.1 "  N , 23 ° 37 '29.2"  E ). The approximately 50 footprints are assigned to a hominin . In an analysis of tooth roots published in 2017, the fossil genus Graecopithecus , the remains of which (a lower jaw and a tooth) were found near Athens and Bulgaria and dated 7.2 million years ago, was described as possibly the oldest known representative of the hominini .

8 to 7 million years ago, the elevation of the Tibetan highlands also increased the duration and strength of the monsoons in Asia. The result - also in Europe - was a reduction in the frequency and duration of precipitation, demonstrable, among other things, by the spread of C4 grasses . These climatic changes gradually led to the complete extinction of human species in Europe and to a decline in biodiversity in Asia, from which only the orangutans and gibbons were ultimately spared.

Development in Africa

North and East Africa were also affected by the Vallesium Crisis 9.6 million years ago. However, so far only a few finds are known from these regions, which were dated to the time from 13 to 7 million years ago. This had at times led to the suspicion that the great apes had evolved in Asia and migrated back to Africa. In fact, however, the poorer conditions for the formation of fossils - the more humid climate - were one of the main reasons for the low number of African finds. With the almost 10 million year old Nakalipithecus nakayamai from Kenya , first described in 2007 , the somewhat older Samburupithecus , which has been known from Kenya for a long time, and the somewhat older Chororapithecus recovered in Ethiopia , it can now be proven that Africa in the middle and late Miocene of was populated by a variety of human species. In the late Miocene - 8 to 6 million years ago - the evolutionary lines of the gorillas and chimpanzees also separated from those of the hominini in Africa, whereby it is assumed that the last common ancestor of chimpanzees and hominini had approximately the body dimensions of today's chimpanzees.

A genetic analysis by the Broad Institute , a joint institute of the Massachusetts Institute of Technology and Harvard University , published in the summer of 2006 in the journal Nature , was interpreted to mean that the development of the earliest hominini was more unusual than is most assumed. In this study, 20 million base pairs of human DNA as well as chimpanzee and gorilla DNA were compared with one another using the molecular clock. According to the study, an early great ape separated itself from the ancestors of the hominini about 10 million years ago. However, these two populations reunited a few millennia later and formed a mixed population in which crosses occurred.

The fossil " Ardi ", of an Ardipithecus ramidus : Drawing based on the reconstruction drawing in Science .

According to the interpretations of the researchers, there was a succession of crossbreeds and divergent groups over four million years until the last permanent separation of the chimpanzee ancestors and the hominini took place about 6.3 to 5.4 million years ago. This last gene exchange is proven by the consistently very low age of the X chromosomes , which only developed in the form characteristic of humans at this later point in time and which were very similar to the X chromosomes of the chimpanzee. However, this scenario did not go unchallenged.

Fan out of the hominini

The published data on the duration of the existence of the hominini genera and species are estimated values ​​that were derived from the geological ( stratigraphic ) investigations of the fossil sites, i.e. from the determination of the age of individual sites. The so-called molecular clock cannot be used for the genera of the hominini, since modern humans are the only surviving species and genetic reference material is only available from them and the closely related Neanderthals .

Due to the few finds so far, it is completely unclear which African ancestors immediately preceded the 7 to 5 million year old genera Sahelanthropus , Orrorin and Ardipithecus and whether these three genera were rightly placed in the direct line of ancestors of the genus Homo by their discoverers .

The genera of the hominini :
The chronological sequence does not allow any conclusions to be drawn about their family relationships

Early days of the hominini

Sahelanthropus , holotype (copy)

After the discovery of the around 6 million year old Orrorin in 2000 and the 7 to 6 million year old Sahelanthropus in 2001 (in the location TM 266 ), both species were identified as the oldest known species of hominini that were already upright direct ancestors of man. However, this interpretation contradicts the findings calculated on the basis of the molecular clock, according to which the separation of the lines of development leading to homo and chimpanzees did not occur until 5 to 6 million years ago.

Fossils of Ardipithecus ramidus were discovered in Ethiopia as early as 1994 . They are 4.4 million years old and are also considered by many researchers to be the direct ancestors of humans. As the fossil Ardi in particular seems to show, individuals of this species could also walk on two legs on the ground. The family relationships between Sahelanthropus , Orrorin and Ardipithecus, as well as the relationships of these three species to the later hominini, are, however, controversial.

These finds come from the transition from Messinian to Zancleum , a geological epoch in which the world mean temperature was about 4 ° C higher than today. According to a study published in 2011, the habitats of the early hominini 6 million years ago were savannas with at most 40 percent canopy cover; 3.6 million years ago the tree cover was 40 to 60 percent, and in the transition from the Pliocene to the Pleistocene it decreased again, so that 1.9 million years ago areas with a tree cover of more than 50 percent had almost disappeared. In a study published in 2019 it was pointed out that the species communities in the hominini's habitat only began to resemble those in today's steppe areas around 700,000 years ago; in the six million years before there were many more species of extremely large herbivores ( megafauna ) than afterwards. These herbivore species were also mostly not ruminants , so that the grassland was eaten much more than today, for example in the Serengeti .

Australopithecines

Hypothesis on the evolution of the australopithecines , as represented by Friedemann Schrenk , for example, based on the current finds .

The Australopithecines, especially the genus Australopithecus, are among the already upright ancestors of man, relatively well known from fossil finds . However , it is still unclear how their species are related to each other and to the species of the genus Homo . Australopithecus anamensis is now considered to be “the earliest undoubted hominini species”; one of the most important Australopithecus sites in South Africa is therefore considered to be the cradle of humanity . Particularly well-known individual finds are Lucy and the young, female skeleton DIK 1-1 (both belong to Australopithecus afarensis ) and the child of Taung (the skull of an Australopithecus africanus ). At Laetoli in the Olduvai Gorge in Tanzania , fossil footprints of several Australopithecus afarensis individuals were found, which clearly show that representatives of this species walked upright. However, it is disputed whether the Kenyanthropus, discovered by Meave Leakey in Kenya in 1999 , should be regarded as an independent genus or should be placed under the genus Australopithecus .

Paranthropus aethiopicus

The genus Paranthropus , whose representatives have extremely robust teeth, is also placed among the Australopithecines . The cause of this adaptation is a climate change (cooling) around 2.5 million years ago; It was triggered on the one hand by the glaciation of the Arctic , which began 2.7 million years ago, and on the other hand by the plate-tectonic uplift in East Africa. Both of these resulted in a decrease in the amount of precipitation and, as a result, an extensive desertification of the traditional habitat of the ancestors of Paranthropus . Such a savannah-like landscape primarily provided food for grass-eating cloven - hoofed ungulates and ruminants , which existed before, mostly in smaller forms, as foliage- grazing forest dwellers. These soon appeared in large herds, and because they became more numerous, predators and scavengers could also multiply. Two types of hominini thus differentiated themselves.

One type arose as a result of an adaptation to what is now - in comparison to the foliage of the forests - hard fibrous food in the savannah. In this ecological niche, Paranthropus boisei , Paranthropus robustus and Paranthropus aethiopicus developed huge masticatory muscles and correspondingly powerful molars . Her jaw muscles attached to the high crest of the skull.

The second type mitigated the effects of climate change by adopting a diet that increasingly included meat as food. Since these individuals neither had the ability to prey on larger prey as predators nor had claws or teeth that would have been suitable for killing or breaking open large prey, their diet is likely to have been limited to carrion and prey. It is very likely that stone tools were first used here , with stones being used to expose the marrow of long bones that were captured .

Controversial, any class and what types are to be assigned this oldest stone tools: "Some researchers is of the opinion that the production of Oldowan equipments the Australopithecus was due and that the use of tools not be regarded as exclusive indication of the human species may. It is possible that the Australopithecines were also capable of rough stone processing. ”However, a wide range of shapes of stone tools“ and their systematic production with the help of other tools, ie with artificially created devices ”can only be ascribed to the species of the homo genus .

A later representative of the genus Australopithecinen is Australopithecus sediba , which was first described by Lee Berger in 2010 and lived about 2 million years ago. It shows ape-like features as well as those of modern humans.

Genus Homo

From a species of the genus Australopithecus , the first representatives of the genus Homo developed three to two million years ago , the fossils of which were placed on Homo , especially on the basis of tool finds .

It is noticeable that this development also took place in an era in which the climate in East Africa - which had previously changed several times between relatively humid and relatively dry - again switched to drought. This is documented from 2.8 million years ago (with a maximum of 1.8 to 1.6 million years ago) by dust deposits, by the greater extent of savannas and by increased finds of horned beard such as antelopes ; the maximum of the dust deposits coincides with the oldest evidence of Homo erectus .

Evolutionary characteristics

feature Australopithecus homo
height small big
body shape long arms, short legs short arms, long legs
Locomotion biped and climbing two-legged
Jaw and teeth big small
(Embryonic) development fast slowly
Brain size small big

In paleoanthropology , a clear morphological or behavioral demarcation between the genera Australopithecus and Homo was first sought. The distinction could not be made on the basis of a single criterion, such as brain size or upright gait (bipedia). In particular, in addition to dinosaurs and birds, the upright gait emerged several times independently among the early great apes. Not every fossil that could walk upright therefore belongs in the ancestral line of humans. Therefore, a set of characteristics have been suggested as typical, including body shape and size, erect gait , large brain, small jaw and teeth , precision grip , hair reduction , sweat glands , flat fingernails and toenails , long embryo development , childhood elongation, and in behavior: sexual behavior , language , socialization and culture . The criteria are discussed controversially to this day. There is also disagreement about the individual criteria, such as the required brain size. Some researchers set this to be typical at 700 cubic centimeters, and for others only from 850 cubic centimeters. In the more recent point of view, there is now agreement that the incarnation was not a clear and rapid evolutionary step, but a parallel development of a multitude of characteristics, whereby in the process some were always ancestral, others were already more forward-looking ( mosaic evolution ).

From Homo rudolfensis to Homo erectus

The lower jaw UR 501 of Homo rudolfensis is 2.4 million years old and is the second oldest known fossil of the genus Homo (original, Schrenk collection in the Senckenberg Natural History Museum )
Skull roof Sangiran II , original.
1.5 mya . Koenigswald collection in the Senckenberg Nature Museum

The two oldest homo species are Homo rudolfensis , which was named after Lake Rudolf - now Lake Turkana - in Kenya , and Homo habilis . The relationship between the two species and their relationship to previous and subsequent species of hominini have been controversial so far. In 1999, because of their very close anatomical proximity to Australopithecus , Bernard Wood even suggested that both species should be renamed Australopithecus rudolfensis and Australopithecus habilis . The anatomical features of the two species are, however, usually defined as differentiating them from older Australopithecus species.

The face of Homo habilis and Homo rudolfensis is lighter than that of Australopithecus , but the eyes of Homo habilis are even further apart than that of later homo species such as Homo erectus . The bulge above the eye , which is the continuous transverse bulge of the frontal bone above the bridge of the nose , is less pronounced than in Homo erectus . The skull of Homo habilis and Homo rudolfensis "does not narrow behind the eye sockets as much as in Australopithecus or Paranthropus , so that a larger brain volume results".

The upper and lower jaws of both species are also smaller than in Australopithecus , so the attachments of the masticatory muscles to the skull are less pronounced. According to Friedemann Schrenk , the distinguishing features for Homo habilis and Homo rudolfensis are : the greater brain volume in Homo rudolfensis ; the upper molars of Homo rudolfensis have 3 roots ( Homo habilis: 2), the lower 2 roots ( Homo habilis: 1); The wisdom teeth of Homo rudolfensis are smaller compared to Australopithecus ( not reduced in Homo habilis ), the thigh and foot of Homo rudolfensis are human-like, in Homo habilis they are similar to Australopithecus .

The skull D 2700 and the presumably associated lower jaw D 2735 from the Dmanisi site in Georgia (originals)

Homo ergaster developed about two million years ago . Its rank as a definable species is, however, controversial; various researchers assign its fossils as early specimens of the species Homo erectus . Homo erectus was the first species of the genus Homo to leave Africa and spread across the Middle East to Asia and Europe. The first evidence of the genus Homo outside of Africa comes from the hominine fossils of Dmanisi in Georgia , which were dated 1.8 million years ago and whose connection to the genus Homo is not known; possibly the Dmanisi fossils belong to Homo habilis .

It is also controversial whether the up to 1.2 million year old Spanish fossils called Homo antecessor by their discoverers were rightly designated as a separate species or should be classified as a local variant of Homo erectus .

Using genetic markers, it was calculated that 1.2 million years ago only around 18,500 individuals from the direct ancestral line of Homo sapiens lived.

Homo floresiensis , also jokingly called "Hobbit", the remains of which were discovered in 2003, is now mainly interpreted as a late dwarf form of Homo erectus . Members of this species lived on the Indonesian island of Flores until 60,000 years ago.

From Homo erectus to Homo neanderthalensis

About 800,000 years ago, Homo erectus developed into a form with a larger brain, which is usually referred to as Homo heidelbergensis , but was at times classified by some researchers as a subspecies of Homo erectus . From Homo heidelbergensis and Homo erectus heidelbergensis , the Neanderthals ( Homo neanderthalensis ) developed in Europe , while at the same time Homo sapiens emerged from the populations of Homo erectus that remained in Africa , today's humans .

In addition to the Neanderthals, around 40,000 years ago there was also a sister group of the Neanderthals in the Altai Mountains, the so-called Denisova people . Of them, only a molar, a finger and a toe bone from the Denissowa cave and the Xiahe lower jaw from Tibet have been scientifically described. Based on the analysis of mtDNA and cell nucleus DNA, the studies came to the conclusion that the Denisova people belong to a hominine group that is most closely related to the Neanderthals, but has a possibly 250,000 year long independent population history alongside the Neanderthals. Accordingly, it has in Central Asia in addition to homo sapiens and the Neandertaler a third, independent of these two types there immigrant population of the genus Homo added.

At least three species of the genus Homo - Neanderthals, Homo floresiensis and humans - as well as the Denisova humans therefore colonized Eurasia simultaneously over certain periods of time.

Gene flow between Homo sapiens and archaic human species

Since 2010, numerous archaeogenetic works have revealed a surprising inter-species gene flow between Neanderthals and Homo sapiens as well as between Denisovans and Homo sapiens . From 2013 to 2015, published archaeogenetic studies on the Homo sapiens finds from Peştera cu Oase in Romania and Ust-Ischim in Siberia revealed Neanderthal DNA in both fossils. Accordingly, a successful mating and gene flow between Neanderthals and Homo sapiens can be assumed not only in the Levant , but also in Eastern Europe and Siberia. Some nucleotide sequence variants ( haplotypes ) in the African ethnic groups San , Mandinka and Aka have been interpreted to mean that they could have "mixed" with a homo population around 35,000 years ago , which differed around 700,000 years ago which had separated the line of development leading to modern man. Since no fossils have yet been discovered from such an archaic African homopopulation , it has not yet been possible to prove to which population or species the nucleotide sequence variants can be assigned.

Origin of Homo sapiens

The first migrations of modern humans, reconstructed by genetic markers in the Y chromosome of humans living today.
Four descendants of the first wave of emigration of Homo sapiens :
above Negrito woman ( Philippines ),
Aboriginal man ( Australia ),
below Papuan man ( New Guinea ),
Melanesian boy ( Vanuatu )

The archaic Homo sapiens originated between 300,000 and 200,000 years ago. The oldest finds attributed to him come from Ethiopia ( Bodo 1 ), Morocco ( Djebel Irhoud and Salé ), Zambia ( Kabwe 1 ), South Africa ( Florisbad 1 and Saldanha ) and Tanzania ( Ndutu 1 and Eyasi ), i.e. from northeast, Northwest, Southeast and South Africa. So far, however, no specific region has been identified that could be considered the region of origin. In a review published in 2018, it was argued that the anatomically modern human "does not come from a single founder population in a region of Africa", but from various hunter-gatherer groups scattered across the continent and largely isolated from one another: "Separated by deserts and dense forests they lived in different habitats. Thousands of years of separation resulted in an astonishing variety of human groups, the mixing of which ultimately shaped our species. "

70,000 years ago, Homo sapiens began to spread across Africa and the Middle East. 45,000 years ago it had settled all of Asia and Europe. This begs the question of what became of the prehistoric and early humans , particularly Homo erectus , the Denisovans, and the Neanderthals . There are two theories for this, the “mixing hypothesis” and the “displacement hypothesis”.

Proponents of the hypothesis of the multiregional origin of modern humans ("intermingling hypothesis ") take the view that populations of other prehistoric and early humans - such as Homo erectus and the Neanderthals, who have settled in Africa, Europe and Asia for a long time - are mixed with each other developed from the archaic Homo sapiens into an anatomically modern human being. However, genetic analyzes of the Y chromosome and the mitochondria in humans now support the out-of-Africa theory (see also: Adam of the Y chromosome and mitochondrial Eve ). A mix between Homo sapiens and the late representatives of Homo erectus in Asia is unproven, the mix between Homo sapiens and the Neanderthals was rather low with at most 1 to 4 percent and according to the previous analyzes was not reflected in externally visible features.

The fossil-based Out-of-Africa theory (“displacement hypothesis”) is widely accepted by paleoanthropologists today. According to this, Homo sapiens probably had offspring surviving more quickly and more frequently.

For decades, three finds were considered the oldest reliable evidence of Homo sapiens : the 195,000 year old fossils Omo 1 and Omo 2 as well as the 160,000 year old " Herto skull ", both discovered in northeastern Africa; Whether Homo sapiens also originated in this region or whether its origin lies elsewhere in Africa is currently the subject of scientific discussion - especially since the discovery of the 300,000 year old skull of Djebel Irhoud in Morocco .

Homo sapiens is the only human species to have colonized Australia (about 60,000 years ago) and America (about 15,000 to 11,500 years ago, in some opinion much earlier). Homo sapiens is also the last survivor of the genus Homo .

Important human finds in Europe

Important sites in Europe are the Sierra de Atapuerca in Spain , where colonization seems to have been secured for 1.2 million years, starting with Homo erectus / Homo antecessor through Homo heidelbergensis to Neanderthals and Homo sapiens ; the cave of Arago in southern France , in which 450,000 year old Homo heidelbergensis fossils were recovered, here called the human of Tautavel ; Peştera cu oasis in Romania (the place where the oldest Homo sapiens fossils were found in Europe) and Cro-Magnon in France, the namesake of the Cro-Magnon man .

Evolution of man in historical time

Human evolution does not stop. Several relatively younger, new features exist. Among the blood groups , blood group B emerged last, in Asia. Because of the rise in sea level, it was unable to spread to America, where the historically earlier population groups consisted exclusively of people with blood group 0.

A trend towards the reduction of the lower jaw and overbite has been observed since the Middle Ages . The cause is the nutritional lack of fiber-rich food. A reduction in wisdom teeth is observed in parallel. The change in skin color due to different melanin formation as a result of migrations in regions with different adaptation to solar radiation ( ultraviolet radiation ) is another new characteristic of Homo sapiens . An advantageous feature was the lactose tolerance , which has arisen several times independently in the past 10,000 years, i.e. the tolerance of milk in adulthood. Especially Europeans and Asians in the northern hemisphere (with the exception of the Chinese) have the underlying genetic mutation with a high proportion of the population. When settling down, it enabled people to tap into an additional, valuable source of food. Malaria resistance is present in heterozygous sickle cell anemia . In this form the disease is a selective advantage.

The adaptation to living conditions at high altitudes such as in the Himalayas, the Andes and East Africa took place several times independently through genetic changes in the oxygen processing in the blood or the formation of red blood cells . The regionally slightly different mutations are among the most recent physiological evolutionary adaptations of mankind.

Research history

Early views up to the beginning of the 19th century

With reference to Pliny , Andreas Vesalius (1514–1564), the founder of modern anatomy and morphological thought, had already developed a theory of the descent of humans from apes through pygmies in the 16th century , and the English anatomist Edward Tyson had the similarities in 1699 described by chimpanzee and human brains. Apparently, as early as the 17th century, scholarly circles were aware of the great similarity between humans and apes. However, discoveries of human fossils in the 18th and early 19th centuries either found mistakes, like a as Homo diluvii testis designated tertiary giant salamander , or their " diluvial " finding connection was not recognized. The Thuringian geologist Ernst Friedrich von Schlotheim , who in his "Petrefactenkunde" published in 1820–1822, mentioned fossil human remains ("anthropolites") from Bad Cannstatt , Bilzingsleben , Meißen and Köstritz , was a proponent of the diluvial human being, which was controversial at the time . The first well-preserved and confirmed find of a Pleistocene human being was a skeleton discovered in 1823 on the Gower Peninsula ( Wales ), dated to 31,000 BP by means of C14 dating (corresponds to about 35,000 calibrated calendar years before today), which is known as " Red Lady of Paviland ”. This is a male burial sprinkled with red ocher (the sex was initially wrongly determined) of an anatomically modern human, the so-called Cro-Magnon human .

Dissent about the origin in the 19th century

Only with the discovery of the Neanderthal man from the Neanderthal in 1856 - three years before the publication of Darwin's main work on The Origin of Species - did indications emerge that there could have been another type of human besides Homo sapiens . Quarry workers had uncovered 16 bone fragments in a section of the Neandertal that had fallen victim to limestone mining today. In 1864 the Irish geologist William King attributed it to a species that can be distinguished from modern humans, the "Homo Neanderthalensis King". Nevertheless, it remained controversial until the end of the 19th century whether the find from the Neandertal was a “ prehistoric man ” or a deformed modern person.

Independently of this research dispute, Charles Darwin argued in his 1871 work The Descent of Man and Sexual Selection that man probably evolved in Africa because his closest relatives - chimpanzees and gorillas - are native there. The exact anatomical studies that Thomas Henry Huxley had presented in 1863 in his paper Evidence as to Man's Place in Nature suggested the origin of man in Africa.

The human family tree according to Ernst Haeckel

Ernst Haeckel , however, took a completely different approach . As early as 1868 he had written in his Natural Creation Story that "most of the signs pointed to southern Asia". Haeckel based his conjecture primarily on the comparison of hair, skin color and skull shape of the then primitive peoples of Africa and Asia , now called indigenous , with the great apes . At the same time, however, Haeckel conceded: “Perhaps Eastern Africa was also the place where primitive man first emerged from human-like apes; perhaps also a continent now sunk beneath the level of the Indian Ocean, which in the south of what is now Asia stretched east to the Sunda Islands on the one hand , and to Madagascar and Africa on the other . "

Haeckel's hypothesis that the Sunda Islands are the remainder of the sunken continent of Lemuria , on which ancient great apes developed into the ancestors of humans and the other modern great apes , fascinated the young Dutch military doctor Eugène Dubois . He was therefore transferred to Sumatra in 1887 to look for fossils in the area of ​​the Malay Archipelago . In his book The Early Period of man describes Friedemann Schrenk Dubois' approach as follows: "Obsessed with his idea, he started at a point in Java to dig, that would apply to today's ideas as completely hopeless. He dug in an area where within a radius of thousands of kilometers the slightest hint of remnants of a prehistoric man had never been found - and he dug in the right place to the centimeter. ”Dubois, however, knew tips from farmers who worked there Found animal fossils.

Haeckel's anatomical comparisons, Dubois' fossil of the Java man from 1891 and the discovery of the Peking man in the 1920s led to Darwin's reference to Africa as the cradle of mankind being ignored - instead, Asia was considered by the leading researchers than the region in which modern man developed. The discovery of a well-preserved fossil skull around two million years old, which was discovered in 1924 in Taung , which is now part of South Africa , could not change this . Raymond Dart , an anatomist from the University of the Witwatersrand in Johannesburg , recognized the importance of the find and published it in early 1925 in the journal Nature under the new species and genus name Australopithecus africanus .

The child from Taung

First paleoarchaeological evidence of the origin of man from Africa

Thanks to his medical, neuroanatomical training, Raymond Dart had recognized that the so-called child of Taung had an ape-like face; however, his brain and teeth were human-like. Therefore, Dart argued that important features of small skull as if more human-like ape-like, "The cheekbones , the zygomatic arches , maxillary leave and lower jaw delicate detect human-like characteristics." The same is true of the brain, the human-like also more than apelike features comprising: the The Taung child can therefore be classified as a mosaic , that is, as a member of "an extinct sex of monkeys that represents an intermediate link between the great apes of the present day and humans". However, Dart's interpretation of the fossil was not taken seriously until the 1940s.

A major reason for this was the rejection by the then leading American paleontologist Henry Fairfield Osborn , who, as a critic of Darwin's theory of evolution , had popularized the theoretical concept of a so-called Dawn Man ( early man or man of the dawn ). Among other things, Osborn claimed that the modern human brain is so complex that two or three million years could not have been enough to produce it from an ape-like brain. The small brain of Australopithecus africanus with human-like teeth was thus a find that was not in line with the hypotheses of the scientific establishment of the time: Almost all researchers therefore followed Osborn's thesis in the 1920s. They assumed a period of at least 20 to 25 million years since the separation of the hominini from the other great apes : a hypothesis that was by no means new, because even Rudolf Virchow had not recognized the Neanderthals as a fossil species based on similar considerations. This hypothesis, which now seems completely arbitrary, found plausible support in the 1920s in the Piltdown man , whose bones had been collected in Sussex from 1912 and who was only exposed as a fake in 1953. Its head consisted of the skull of a modern human, to which the lower jaw of a monkey had been adapted. The forgery met the prevailing view of the time, according to which the ancestors of man had had a particularly large brain for a long time, and thus blocked the then leading British and American paleontologists from seeing the actual facts.

The turning point in favor of recognizing the Australopithecins as pre-humans only occurred when the informative value of other fossils was reassessed. More and more Neanderthals had been discovered whose physique was uniform and thus made the thesis that the first specimen found was a deformed modern human being untenable. From 1936 onwards, further Australopithecus fossils had been discovered in Africa, which a younger generation of researchers no longer primarily interpreted against the background of older doctrines on the duration of the evolution of the brain.

Establishing the out-of-Africa theory

Particularly significant fossils were discovered from the late 1950s / early 1960s by the couple Louis Leakey and Mary Leakey and later by their son Richard Leakey and his wife Meave Leakey in the Olduvai Gorge in what is now Tanzania and on Lake Turkana in Kenya discovered. Since the 1970s, researchers working with Yves Coppens , Donald Johanson and Tim White have unearthed numerous fossils in the Afar Triangle in Ethiopia . In 1991, the German researcher Friedemann Schrenk discovered in Malawi - after the fossil LD 350-1 - the second oldest fossil attributed to the genus Homo , the complete lower jaw UR 501 of Homo rudolfensis . The most complete skeleton of an early representative of the hominini discovered to date is the Little Foot fossil from Sterkfontein (South Africa), edited by Ronald J. Clarke , which has been uncovered since 1997.

Excavation work in the cave of Arago

In particular, the decades of research of Leakey, the elaboration of a reliable dating method for East African discoveries by Frank Brown and the comparative anatomical studies of Günter Bräuer contributed significantly to the out-of-Africa theory of the origin of modern humans compared to the multi-region theory to establish and consolidate. The Out of Africa theory has been the recognized theory of human regional origins since the 1980s; in fact, to date only hominine fossils are known outside Africa that are less than two million years old. Recently there has also been talk of a modified Out-of-Africa theory, which takes into account that the evolution of modern humans on the path of inter-species gene flow also took place outside of Africa to a small extent.

See also

literature

Web links

Commons : Human evolution  - collection of images, videos and audio files

Individual evidence

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  91. ↑ William King: The reputed fossil man of the Neanderthal. In: Quarterly Journal of Science. Volume 1, 1864, pp. 88–97, Online
    King had previously suggested the name elsewhere - quasi unofficially - see William King: On the Neanderthal Skull, or Reasons for believing it to belong to the Clydian Period and to a species different from that represented by Man. In: British Association for the Advancement of Science, Notices and Abstracts for 1863, Part II. London 1864, p. 81 f.
  92. ^ Charles Darwin: The Descent of Man, and Selection in Relation to Sex. John Murray, London 1871, Volume 1, p. 199: “In each great region of the world the living mammals are closely related to the extinct species of the same region. It is therefore probable that Africa was formerly inhabited by extinct apes closely allied to the gorilla and chimpanzee; and as these two species are now man's nearest allies, it is somewhat more probable that our early progenitors lived on the African continent than elsewhere. "
  93. ↑ Ernst Haeckel: Natural history of creation. Common scientific lectures on the theory of evolution in general and that of Darwin, Goethe and Lamarck in particular, on the application of the same to the origin of man and other related fundamental questions of natural science. Georg Reimer, Berlin 1868, chapter 19 ( full text )
  94. ↑ Friedemann Schrenk : The early days of man. The way to Homo sapiens. CH Beck, 1997, p. 81.
  95. ^ The Discovery of Java Man in 1891. In: Athena Review. Volume 4, No. 1: Homo erectus.
  96. ^ Raymond A. Dart: Australopithecus africanus: The Man-Ape of South-Africa. In: Nature. Volume 115, 1925, pp. 195–199, doi: 10.1038 / 115195a0 , (full text; PDF; 456 kB) .
  97. ^ R. Dart: "an extinct race of apes intermediate between living anthropoids and man." In: Nature. 1925, p. 195.
  98. ^ Charles Dawson, Arthur Smith Woodward: On the Discovery of a Palæolithic Human Skull and Mandible in Flint-Bearing Gravel Overlying the Wealden (Hastings Beds) at Piltdown, Fletching (Sussex). In: Quarterly Journal of the Geological Society. Volume 69, March 1913.
  99. ↑ KE Langergraber: Kinship and cooperation in primates. In: JC Mitani, J. Call, PM Kappeler, R. Palombit, J. Silk (Eds.): The Evolution of Primate Societies. University of Chicago Press, 2012, pp. 491-513.
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