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Introduction; The Process of Evolution; Characteristics, Classification, and Evolution of the Primates; The First Humans: Australopithecines; The Genus Homo; The Evolution of Cultural Behavior
By around 6 million years ago in Africa, an apelike species had evolved with two important traits that distinguished it from apes: (1) small canine, or eye, teeth (teeth next to the four incisors, or front teeth) and (2) bipedalism—that is, walking on two legs as the primary form of locomotion. Scientists refer to these earliest human species as australopithecines, or australopiths for short. The earliest australopith species known today belong to three genera: Sahelanthropus, Orrorin, and Ardipithecus. Other species belong to the genus Australopithecus and, by some classifications, Paranthropus. The name australopithecine translates literally as “southern ape,” in reference to South Africa, where the first known australopith fossils were found. The Great Rift Valley, a region in eastern Africa in which past movements in Earth’s crust have exposed ancient deposits of fossils, has become famous for its australopith finds. Countries in which scientists have found australopith fossils include Ethiopia, Tanzania, Kenya, South Africa, and Chad. Thus, australopiths ranged widely over the African continent.
Fossils from several different early australopith species that lived between 4 million and 2 million years ago clearly show a variety of adaptations that mark the transition from ape to human. The very early period of this transition, prior to 4 million years ago, remains poorly documented in the fossil record, but those fossils that do exist show the most primitive combinations of ape and human features. Fossils reveal much about the physical build and activities of early australopiths, but not everything about outward physical features such as the color and texture of skin and hair, or about certain behaviors, such as methods of obtaining food or patterns of social interaction. For these reasons, scientists study the living great apes—particularly the African apes—to better understand how early australopiths might have looked and behaved, and how the transition from ape to human might have occurred. For example, australopiths probably resembled the great apes in characteristics such as the shape of the face and the amount of hair on the body. Australopiths also had brains roughly equal in size to those of the great apes, so they probably had apelike mental abilities. Their social life probably resembled that of chimpanzees.
Most of the distinctly human physical qualities in australopiths related to their bipedal stance. Before australopiths, no mammal had ever evolved an anatomy for habitual upright walking. Australopiths also had small canine teeth, as compared with long canines found in almost all other catarrhine primates. Other characteristics of australopiths reflected their ape ancestry. They had a low cranium behind a projecting face, and a brain size of 390 to 550 cu cm (24 to 34 cu in)—in the range of an ape’s brain. The body weight of australopiths, as estimated from their bones, ranged from 27 to 49 kg (60 to 108 lb), and they stood 1.1 to 1.5 m (3.5 to 5 ft) tall. Their weight and height compare closely to those of chimpanzees (chimp height measured standing). Some australopith species had a large degree of sexual dimorphism—males were much larger than females—a trait also found in gorillas, orangutans, and some other primates. Australopiths also had curved fingers and long thumbs with a wide range of movement. In comparison, the fingers of apes are longer, more powerful, and more curved, making them extremely well adapted for hanging and swinging from branches. Apes also have very short thumbs, which limits their ability to manipulate small objects. Paleoanthropologists speculate as to whether the long and dexterous thumbs of australopiths allowed them to use tools more efficiently than do apes.
The anatomy of australopiths shows a number of adaptations for bipedalism, in both the upper and lower body. Adaptations in the lower body included the following: The australopith ilium, or pelvic bone, which rises above the hip joint, was much shorter and broader than it is in apes. This shape enabled the hip muscles to steady the body during each step. The australopith pelvis also had a bowl-like shape, which supported the internal organs in an upright stance. The upper legs angled inward from the hip joints, which positioned the knees to better support the body during upright walking. The legs of apes, on the other hand, are positioned almost straight down from the hip, so that when an ape walks upright for a short distance, its body sways from side to side. Australopiths also had shorter and less flexible toes than do apes. The toes worked as rigid levers for pushing off the ground during each bipedal step. Other adaptations occurred above the pelvis. The australopith spine had an S-shaped curve, which shortened the overall length of the torso and gave it rigidity and balance when standing. By contrast, apes have a relatively straight spine. The australopith skull also had an important adaptation related to bipedalism. The opening at the bottom of the skull through which the spinal cord attaches to the brain, called the foramen magnum, was positioned more forward than it is in apes. This position set the head in balance over the upright spine. Australopiths clearly walked upright on the ground, but paleoanthropologists debate whether the earliest humans also spent a significant amount of time in the trees. Certain physical features indicate that they spent at least some of their time climbing in trees. Such features include their curved and elongated fingers and elongated arms. However, their fingers, unlike those of apes, may not have been long enough to allow them to brachiate through the treetops. Study of fossil wrist bones suggests that early australopiths had the ability to lock their wrists, preventing backward bending at the wrist when the body weight was placed on the knuckles of the hand. This could mean that the earliest bipeds had an ancestor that walked on its knuckles, as African apes do.
Compared with apes, humans have very small canine teeth. Apes—particularly males—have thick, projecting, sharp canines that they use for displays of aggression and as weapons to defend themselves. The oldest known bipeds, who lived at least 6 million years ago, still had large canines by human standards, though not as large as in apes. By 4 million years ago australopiths had developed the human characteristic of having smaller, flatter canines. Canine reduction might have related to an increase in social cooperation among humans and an accompanying decrease in the need for males to make aggressive displays. The australopiths can be divided into an early group of species, known as gracile australopiths, which arose prior to 3 million years ago; and a later group, known as robust australopiths, which evolved after 3 million years ago. The gracile australopiths—of which several species evolved between 4.5 million and 3 million years ago—generally had smaller teeth and jaws. The later-evolving robusts had larger faces with large jaws and molars (cheek teeth). These traits indicate powerful and prolonged chewing of food, and analyses of wear on the chewing surface of robust australopith molar teeth support this idea. Some fossils of early australopiths have features resembling those of the later species, suggesting that the robusts evolved from one or more gracile ancestors.
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