Out of Africa I
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In paleoanthropology, Out of Africa I is the first hominin expansion into Eurasia, taking place between 1.8 and 0.8 million years ago. It is thought that Homo erectus developed a flexible adaptation to the open grounds, descending from the older Homo habilis lineage, which was strictly adapted to the dense woodlands. Such an adaptation would have allowed Homo erectus to leave Africa and expand its range into Eurasia. According to the recent African origin of modern humans hypothesis (Out of Africa II), the first hominin expansion out of Africa is followed by the dispersal into Eurasia and replacement of these previous hominins by anatomically modern humans, starting about 100,000 years ago. Without further specification, Out of Africa is usually held to mean Out of Africa II, the expansion of modern humans into Eurasia.
Movements out of Africa by early hominins seem to have occurred in at least three waves. Primitive chopper producers were first out by c. 1.8 Ma, followed by early Acheulean industries c. 1.4 Ma, and various cleaver-producing Acheulean groups around 0.8 Ma.[1]
Until the early 1980s, hominins were assumed to have been restricted to the African continent for the whole of the Early Pleistocene (until about 0.8 Ma), and so much archaeological effort has disproportionately focused on Africa. Compounded with hominins probably being rare out of East Africa in the Early Pleistocene,[2] we are left with a sequence of events broken in space and time.
Sites
Hominin sites are oldest in East Africa. The earliest evidence for retouched tools are from Kada Gona, Ethiopia, dating back to 2.6 – 2.5 Ma in the very early Pleistocene. They might be the product of Australopithecus garhi or Paranthropus aethiopicus, the two known hominins contemporary with the tools.[3]Homo habilis is the first member of the Homo line and could have descended from the Australopithecus as early as 2.3 Ma; it is first attested in Lake Turkana, Kenya. Homo erectus seems to appear a little later, its earliest remains dating to c. 1.9 – 1.6 Ma, from Koobi Fora, Kenya.[4] The two species would have lived face to face in East Africa for nearly half a million years.[5]
Well before Homo habilis disappeared (c. 1.4 Ma), Homo erectus had made it into Eurasia. The earliest well-dated Eurasian site is that of Dmanisi in Georgia, and is securely dated to 1.81 Ma.[6]
There, some evidence of caring for the old was found. The skull of an old Homo erectus had lost all but one teeth years before his death, and is perhaps unlikely to have survived on his own (but such caring is not yet certain – a partially paralysed chimpanzee at the Gombe reserve survived for years without help[7]).
Early Pleistocene sites in North Africa, the geographical intermediate of East Africa and Georgia, are in poor stratigraphic context. The earliest of the dated is Ain Hanech in northern Algeria (c. 1.8[8] – 1.2 Ma[9]), an Oldowan grade layer. It attests that early hominins have crossed the northern African tracts, which are usually hot and dry. Hominins were part of the East African biome, and so a flux in climate could have momentarily expanded their environment, giving them the chance to move north.
There is very little time between Homo erectus’ apparent arrival in South Caucasus, and its probable arrival in East and Southeast Asia. There is evidence of hominins in Yuanmou, China, dating to 1.7 Ma and in Sangiran, on Java, Indonesia, dating to 1.66 Ma.[10] It appears hominins took longer to move into Europe, with the earliest site in Barranco León in southeastern Spain dated to 1.4 Ma[11] and a more controversial Pirro Nord in southern Italy, allegedly from 1.3 – 1.7 Ma.[12]
In any case, by 1 Ma, hominins had settled in most of the Old World. In Western Europe, it is hard to say, however, if settlement was continuous or if successive waves repopulated the territory in glacial interludes. Early Acheulean tools were present at Ubeidiya by 1.4 Ma[13] and it seems likely that successive waves out of Africa after then would have brought Acheulean technology to Western Europe, but handaxes, which are typical of the Acheulean industry, are absent in early Western European sites.
Routes out of Africa
Sinai Peninsula
The Sinai Peninsula should be the African exit route par excellence, being since the Pliocene the only land bridge between the two continents of the Old World. As detailed below, unless we argue for boats on behalf of Homo erectus, it is surely the only way out. However, it was hard to access until the Middle Pleistocene. The Nile followed a different and pitiful course.There are two Eurasian entryways that take advantage of the Sinai. First, the Levantine corridor, which moves north along the Eastern Mediterranean. Second, down the eastern bank of the Red Sea. Archaeological efforts in Arabia is limited, and attention is usually given to the Levantine corridor.
Due to the presence of Eurasian sites securely dated to the Early Pleistocene, to reject the Sinai is effectively to affirm that early hominins crossed straits, and this hypothesis has problems of its own.
Bab-el-Mandeb
The land connection with Arabia has disappeared in the Pliocene,[14] and though it may have reformed momentarily,[15] the evaporation of the Red Sea and associated increase in salinity would have left traces in the fossil record after just 200 years and evaporite deposits after 600 years. Neither have been detected.[16] A strong current flows from the Red Sea into the Indian Ocean and crossing would have been difficult without a land connection.
Oldowan grade tools are reported from Perim Island,[17] implying that the strait could have been crossed in the Early Pleistocene, though these finds have yet to be confirmed.[18]
Strait of Gibraltar
The Strait of Gibraltar is the Atlantic entryway to the Mediterranean, where Spanish and Moroccan banks are only 14 km apart. A decrease in sea levels in the Pleistocene due to glaciation would not have brought this down to less than 10 km. Water treadmills at Gibraltar. Deep currents push westwards, and surface water flows strongly back into the Mediterranean. The current would likely lose a swimmer or an unsteered raft.
Entrance into Eurasia across the strait of Gibraltar could explain the hominin remains at Barranco León in southeastern Spain dated to 1.4 Ma[11] and Sima del Elefante in northern Spain dated to 1.2 Ma.[19][20] But the site of Pirro Nord in southern Italy, allegedly from 1.3 – 1.7 Ma,[12] suggests a possible arrival from the East. Resolution is insufficient to settle the matter.[21]
Strait of Sicily
The modern Strait of Sicily separates Tunisia and Sicily by 145 km, but is shallow and would have been much narrower in glacial maxima. We have a poor understanding of plate tectonics of this area for the greater part of the Pleistocene. But while plate tectonics could have made the strait narrower than predicted by the lowering of sea levels alone, contrast of Pleistocene fauna strongly argues against an actual land bridge. Since the strait is only 400 km away from the North African hominin site of Ain Hanech in Algeria, dating to 1.8 Ma[8] or 1.2 Ma,[9] it remains a plausible route for Early Pleistocene expansion into Eurasia. But there is close to no evidence for an actual passing. Alimen based most of his argument[22] in favour of such migration on Bianchini’s discovery[23] of Sicilian Oldowan grade tools. But radiometric dates have not been produced, and the artefacts might as well be from the Middle Pleistocene.[24]
Crossing straits
Presence of hominin remains in Indonesian islands is good evidence for seafaring by Homo erectus late in the Early Pleistocene. Bednarik suggests that navigation had appeared by 1 Ma, possibly to exploit offshore fishing grounds.[25] He has reproduced a primitive dirigible raft to demonstrate the feasibility of faring across the Lombok Strait on such a device, which he believes to have been done before 850 ka. The strait has maintained a width of at least 20 km for the whole of the Pleistocene.Such an achievement by Homo erectus in the Early Pleistocene offers some strength to the suggested water routes out of Africa, as the Gibraltar, Sicilian, and Bab-el-Mandeb exit routes are harder to consider if boats are deemed beyond the capacities of Homo erectus.
It might be tempting to consider a one-off event getting a few hominins across a strait – perhaps an Homo erectus family drifting on flood debris to land on a Eurasian bank. But successful population of Eurasia by such a beginning is unlikely. There are biological constraints to the minimum size a population must maintain to avoid extinction. That is to say, if less than 50 hominins at once made it into Eurasia and loss contact with African hominins, the population would likely undergo an extinction vortex, in part due to inbreeding.[26]
Causes for hominin dispersals
Climate change and hominin flexibility
For a given species in a given environment, available resources will limit the amount of individuals that can survive indefinitely. This is the carrying capacity. Upon reaching this threshold, individuals may find it easier to gather resources in the poorer less exploited peripheral environment than in the preferred habitat. Homo habilis could have developed some baseline behavioural flexibility prior to its expansion into the peripheries (such as encroaching into the predatory guild[27][28]). This flexibility could then have been positively selected and amplified, leading to Homo erectus’ adaptation to the peripheral open habitats.[29] A new and more flexible hominin population could have come back to the old niche and replace the ancestral population.[30] Moreover, some step-wise shrinking of the woodland and the associated reduction of hominin carrying capacity in the woods around 1.8 Ma, 1.2 Ma, and 0.6 Ma would have stressed the carrying capacity’s pressure for adapting to the open grounds.[31][32]With Homo erectus’ new environmental flexibility, it is likely that it saw favourable climate fluxes open it the way to the Levantine corridor at least sporadically in the Early Pleistocene.[2]
Chasing fauna
Lithic analysis implies that Oldowan hominins weren't predators.[33] However, Homo erectus appears to have followed animal migrations to the north during wetter periods, likely as a source of scavenged food. The sabre-tooth cat Megantereon was an apex predator of the Early and Middle Pleistocene (before MIS 12). It went extinct in Africa c. 1.5 Ma,[34] but had already moved out through the Sinai, and is among the faunal remains of the Levantine hominin site of Ubeidiya, c. 1.4 Ma.[13] It couldn’t break bone marrow and its kills were likely an important food source for hominins,[35] especially in glacial periods.[36]In colder Eurasian times, the hominin diet would have to be principally meat-based and Acheulean hunters must have competed with cats.
Coevolved zoonotic diseases
Bar-Yosef and Cohen[1] suggest that the success of hominins within Eurasia once out of Africa is in part due to the absence of zoonotic diseases outside their original habitat. Zoonotic diseases are those that are transmitted from animals to humans. While a disease specific to hominins must keep its human host alive long enough to transmit itself, zoonotic diseases won't necessarily do so as they can complete their life cycle without humans. Still, these infections are well accustomed to human presence, having evolved alongside them. The higher an African ape's population density the better a disease fares. Fifty-five percent of chimps at the Gombe reserve die of disease, most of them zoonotic.[37] The majority of these diseases are still restricted to hot and damp African environments.Once hominins had moved out into dryer and colder habitats of higher latitudes, one major limiting factor in population growth was out of the equation.
Hominin biology
While Homo habilis was certainly bipedal, its long arms are indicative of an arboreal adaptation.[38] Homo erectus had longer legs and shorter arms, revealing a transition to obligate terrestriality, though it remains unclear how this change in relative leg length might have been an advantage.[39] Sheer body size, on the other hand, seems to have allowed for better walking energy efficiency and endurance.[40] A larger Homo erectus would also dehydrate more slowly and could thus cover greater distances before facing thermoregulatory limitations.[41] The ability for prolonged walking at a normal pace would have been a decisive factor for effective colonisation of Eurasia.[42]
Brain thermoregulation
Thermoregulation and dehydration are major problems that need to be dealt with to move into the open grasslands. In particular, vascularisation of the brain is crucial in maintaining it in the narrow frame of tolerable temperatures.
Bones of the higher cranium grow in response to expansion of cerebral mass, in such a way that brain tissue and blood vessels mold the inner brain case. Endocranial casts of fossil skulls allow
approximating brain vascularisation.[43] Dean Falk noticed that a single large vessel, the occipital marginal sinus, was responsible for irrigating most of the brain in early australopiths
(Australopithecus afarensis, Paranthropus robustus and boisei).[44] The vessel grew smaller with time so as to be progressively replaced by a network of small veins in later hominins, starting with Homo habilis and continuing well into Eurasia. She interprets the change as an adaptation to cool the brain,[45] which she uses to advance her “radiator theory” for accelerated encephalisation from Homo habilis onwards.[46] To Falk, bipedalism, which predates large brains, favoured a rewiring of cerebral blood vessels into a gravity-assisted irrigation network, itself allowing the cool down needed for encephalisation.
Endocranial casts of Homo habilis and Homo erectus differ in the organisation of the frontal lobe, in particular in the prefrontal cortex where higher mental functions of consciousness and abstraction occur.[47] By themselves, mental capacities have likely played a role in the success of Eurasian colonisation. They would have allowed for greater social complexity,[48] predation and sharing prey,[49] and an overall higher quality diet.[50] If we are to believe Bednarik and his seafaring Indonesian Homo erectus, then the brain must have played a role in crossing channels.
According to Wheeler,[51] loss of functional body hair would have helped prevent hyperthermia, since hair will obstruct air flow over the skin and restrict cooling by evaporation. He further suggests that body cooling due to hair loss has relieved a thermal constraint on brain size (but in a response to Falk’s radiator hypothesis, Ralph Holloway maintains that there is no evidence for a temperature constraint on brain size[52]). However, differences in body hair between Homo habilis and Homo erectus are impossible to test, and it will remain unclear whether hair loss was part of the hominin adaptation or preadaptation to Eurasia.
