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    Evolution Before about it was widely thought that distinctively hominin fossils could be identified from 14 to 12 million years ago mya. However, during the s geneticists introduced the use of molecular clocks to calculate how long species had been separated from a common ancestor.


    The molecular clock concept is based on an assumed regularity in the accumulation of tiny changes in the genetic codes of humans and other organisms. Use of this concept, together with a reanalysis of the fossil record, moved the estimated time of the evolutionary split between apes and human ancestors forward to as recently as about 5 mya.

    Since then the molecular data and a steady trickle of new hominin fossil finds have pushed the earliest putative hominin ancestry back in time somewhat, to perhaps 8—6 mya. Possible pathways in the evolution of the human lineage. Announced inthis specimen is dated to the period between 7 and 6 mya. The distinctive mark of Hominini is generally taken to be upright land locomotion on two legs terrestrial bipedalism. The skull of S.

    The most remarkable aspect of this skull is the broadness and flatness of its face—something previously associated with much more recent hominins—in conjunction with a smaller, ape-sized braincase. This specimen also has small canine teeth compared with those of apes, thus aligning it with the hominins in an important functional regard.

    Sahelanthropus, then, emphasizes an evolutionary pattern that seems to have been a characteristic of the tribe Hominini from the very start—a pattern that aligns it with what is observed in most other evolutionarily successful groups of mammals.

    Human evolution, it appears, has consistently been a process of trial and error. As flattering to the modern human ego as this picture may be, it is evidently quite wrong.

    Instead, human evolution has been throughout its long history a matter of experimentation, with new species being constantly spawned and thrown into the ecological arena to compete and, more often than not, become extinct. Viewed this way, H. Fossils found since the early s have begun to hint at just how complex the hominin bush was in the three million years or so following the time of Sahelanthropus.

    Three other new genera of early hominins ArdipithecusOrrorin, and Kenyanthropus dating from 6 to 3 mya have been recovered from Kenya and Ethiopia.

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    Furthermore, during the latter half of the 20th century, new species were added to the long-established genera Australopithecus and Paranthropus, both known from South and East African sites. This early radiation diversification of hominins, of which the latest survivors lived as recently as about 1. In general, they were relatively small-bodied, with large chewing teeth but reduced sometimes highly reduced canine teeth. They had small ape-sized braincases and rather protruding faces.

    Replica of a 3. The design of her pelvis and feet are suggestive of bipedal locomotion. However, other skeletal elements indicate that she spent much of her time clambering through the branches of trees.

    These fossils, along with the slightly older trails of footprints found at LaetoliTanzania, prove that early hominins were upright bipeds when on the ground.

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    However, they also retained many reminders of their tree-dwelling ancestry, especially their rather long arms, short legs, narrow shoulders, and long grasping extremities. All these features would have made them agile upright foragers among tree branches, where they presumably sought food by day and sheltered at night even though they moved on two legs while on the ground.


    The environments in which these early hominins lived suggest that 1 they were still comfortable in the forest and 2 they were largely active at the forest edges and in the woodlands where the forest graded into more open savanna —a type of habitat that was expanding in their African homeland after about 7 mya as climates became drier and more seasonal.

    A trail of footprints probably left by Australopithecus afarensis individuals some 3. Moreover, although these ancient forms were clearly members of the same larger group, discerning exactly how any of them may have been connected to later species is problematic because of incomplete fossil evidence or different interpretations of the same evidence. Homo may have originated as early as about 2. A variety of incomplete or broken fossils from the period between about 2.

    Taken together, this hominin assemblage makes a rather odd assortment that is based more than anything else on a modest increase in the size of the brain compared with that of Australopithecus and its relatives. Even more important in the assignment of these fossils to Homo may be the occurrence in the same geologic deposits of very primitive stone tools.

    Decades later, the species responsible for producing the first stone tools remains unknown, but it likely was relatively small-brained, with a body proportioned quite differently from that of H.

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    Paleontologists speculate that, barring the finding of an as-yet-undiscovered species of Homo, the tools were likely constructed by members of Australopithecus or Kenyanthropus. Cranial remains dating to slightly less than 2 mya have been discovered at Koobi ForaKenya. These are thought to belong to the same species as the remarkably complete 1.

    The nature of the association between the two finds is not yet completely evident, as even partial hominin skeletons are almost vanishingly rare as researchers delve deeper into the past to a time before the introduction of burial practices. Discovered inthe slender-limbed, long-legged Nariokotome skeleton is the first solid evidence of an individual that resembled H.


    Here at last is a representative of a species that was definitely at home on the open savanna, emancipated from the forest and woodland environments to which its predecessors had been confined.

    Turkana Boy was 1. His skeleton bears the basic hallmarks of H. His face, however, was quite projecting, and his brain was little more than half the size of that of H. Cranial traits notwithstanding, this individual clearly deserves to be classified with H. However, he is often assigned by authorities to either H. Once modern human body proportions had been achieved, such species could indulge their newfound wanderlust.

    In their new territories they diversified, as might be expected, with new species emerging in different regions. Africa appears to have been the source of not just one but successive waves of hominin emigrants, including H. In Europe an early representative of H. Throughout there was a tendency for new hominin species to acquire ever larger brains.

    This increase must have come at a cost, because brain tissue expends significant amounts of energy. There must have been benefits of a larger brain, but what those benefits were can only be guessed; quantifying human intelligence is problematic even among living humans, let alone extinct ones.

    The increase in hominin cranial capacity over time. The oldest known remains, found at Jebel Irhoud, Morocco, were unearthed in the first decade of the s. This collection of specimens was made up of skull fragments, a complete jawbone, and stone tools, all of which date to about kya, which may suggest that the species was widely dispersed throughout Africa by that time.

    The second oldest remains date to approximately kya in East Africa.


    This estimate is shaped by the discovery in of the oldest remains attributed to H. The remains, made up of two skulls Omo 1 and Omo 2were initially dated to kya, but through the application of more-sophisticated dating techniques in the remains have been more accurately dated to kya. Most evidence points to H. Behavioral influences The story of hominin evolution is one of increasing behavioral complexity, but, because behaviour does not leave direct fossil evidence, clues must be sought in other sources.

    The most obvious candidates are in the archaeological recordwhich has traditionally begun with the appearance of Paleolithic Old Stone Age tools about 2. See also Stone Age ; however, that date is complicated by the discovery of tools in that date back to 3.

    Early tools were simple indeed: But, for all their simplicity, they marked a major advance in lifestyle: These tools also signify a cognitive advance in hominins; even with intensive training, no ape has yet mastered the notion of hitting one rock with another at precisely the angle needed to detach a sharp flake. Furthermore, the early toolmakers had the ability to anticipate their needs, since they often carried suitable rocks long distances before making them into tools.

    Replica stone tools of the Acheulean industry, used by Homo erectus and early modern humans, and of the Mousterian industry, used by Neanderthals.

    Top, left to right Mid-Acheulean bifacial hand ax and Acheulean banded-flint hand ax. Centre Acheulean hand tool.

    Bottom, left to right Mousterian bifacial hand ax, scraper, and bifacial point. Moreover, behavioral novelties have tended not to coincide with the appearance of new species. For almost a million years following the introduction of stone tools, the methods used for making them remained largely unchanged. It is only after about 1. Shaped carefully on both sides to a standard and symmetrical form, it was usually teardrop- or egg-shaped.

    It is the characteristic tool of the Acheulean industry. Although the notion has been contested, it does seem fairly clear that these implements bear witness to another cognitive advance: Hand axes were manufactured in Africa by the thousands—sometimes at apparent workshops—until quite recent times.

    Stone tools of this kind have always been rare in eastern Asia. It is only at about — kya that another major technological and possibly cognitive advance is found.

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    The great masters of this technique see also Mousterian industry were the Neanderthals, whose possession of language has long been debated. Regardless, it has been demonstrated in studies with people that language is not required for the transmission of the skills needed to make tools of this kind. The stone tool record is well-preserved, but it is only an indirect reflection of overall lifestyle and cognitive capacities.

    It is still unknown, for example, whether the earliest tool users hunted extensively or merely scavenged animal remains. It is likely that, if they hunted, it was for small prey. Nonetheless, metabolic studies of bone suggest that some Australopithecus may have eaten substantially more meat than chimpanzees do today.

    Most authorities had guessed that efficient ambush hunting was an invention of H. Unlike thrusting spears, which must be used at close range and with considerable risk, these 2-metre 6. The age of the location at which these spears were found puts them within the period of H.

    Also at kya there is the first convincing evidence of two other innovations: At Terra Amata in southern France, traces of large huts have been found. The huts were formed by embedding saplings into the ground in an oval and then bringing their tops together at the centre. Stones placed in a ring around the hut braced the saplings. Some of these huts were found to contain hearths scooped in the ground and lined with burned stones and blackened bones.

    These sites represent some of the earliest definitive proof of fires deliberately maintained and used for cookingalthough nearly ,year-old hearths are reported from a site in Israel.

    Prior to the advent of H. In the dwelling places of behaviorally modern early H. The earliest intimations of such partitioning are found at the South African site of Klasies River Mouthdating to more than kya.

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