第29章脊椎動物

The family Hominidae of order Primates includes the hominoids: the great apes and humans (Figure 29.46). Evidence from the fossil record and from a comparison of human and chimpanzee DNA suggests that humans and chimpanzees diverged from a common hominoid ancestor approximately six million years ago. Several species evolved from the evolutionary branch that includes humans, although our species is the only surviving member. The term hominin is used to refer to those species that evolved after this split of the primate line, thereby designating species that are more closely related to humans than to chimpanzees. A number of marker features differentiate humans from the other hominoids, including bipedalism or upright posture, increase in the size of the brain, and a fully opposable thumb that can touch the little finger. Bipedal hominins include several groups that were probably part of the modern human lineage—Australopithecus, Homo habilis, and Homo erectus—and several non-ancestral groups that can be considered “cousins” of modern humans, such as Neanderthals and Denisovans.

Determining the true lines of descent in hominins is difficult. In years past, when relatively few hominin fossils had been recovered, some scientists believed that considering them in order, from oldest to youngest, would demonstrate the course of evolution from early hominins to modern humans. In the past several years, however, many new fossils have been found, and it is clear that there was often more than one species alive at any one time and that many of the fossils found (and species named) represent hominin species that died out and are not ancestral to modern humans.

Three species of very early hominins have made news in the late 20th and early 21st centuries: Ardipithecus, Sahelanthropus, and Orrorin. The youngest of the three species, Ardipithecus, was discovered in the 1990s, and dates to about 4.4 MYA. Although the bipedality of the early specimens was uncertain, several more specimens of Ardipithecus were discovered in the intervening years and demonstrated that the organism was bipedal. Two different species of Ardipithecus have been identified, A. ramidus and A. kadabba, whose specimens are older, dating to 5.6 MYA. However, the status of this genus as a human ancestor is uncertain.

The oldest of the three, Sahelanthropus tchadensis, was discovered in 2001-2002 and has been dated to nearly seven million years ago. There is a single specimen of this genus, a skull that was a surface find in Chad. The fossil, informally called “Toumai,” is a mosaic of primitive and evolved characteristics, and it is unclear how this fossil fits with the picture given by molecular data, namely that the line leading to modern humans and modern chimpanzees apparently bifurcated about six million years ago. It is not thought at this time that this species was an ancestor of modern humans.

A younger (c. 6 MYA) species, Orrorin tugenensis, is also a relatively recent discovery, found in 2000. There are several specimens of Orrorin. Some features of Orrorin are more similar to those of modern humans than are the australopithicenes, although Orrorin is much older. If Orrorin is a human ancestor, then the australopithicenes may not be in the direct human lineage. Additional specimens of these species may help to clarify their role.

Australopithecus (“southern ape”) is a genus of hominin that evolved in eastern Africa approximately four million years ago and went extinct about two million years ago. This genus is of particular interest to us as it is thought that our genus, genus Homo, evolved from a common ancestor shared with Australopithecus about two million years ago (after likely passing through some transitional states). Australopithecus had a number of characteristics that were more similar to the great apes than to modern humans. For example, sexual dimorphism was more exaggerated than in modern humans. Males were up to 50 percent larger than females, a ratio that is similar to that seen in modern gorillas and orangutans. In contrast, modern human males are approximately 15 to 20 percent larger than females. The brain size of Australopithecus relative to its body mass was also smaller than in modern humans and more similar to that seen in the great apes. A key feature that Australopithecus had in common with modern humans was bipedalism, although it is likely that Australopithecus also spent time in trees. Hominin footprints, similar to those of modern humans, were found in Laetoli, Tanzania and dated to 3.6 million years ago. They showed that hominins at the time of Australopithecus were walking upright.

There were a number of Australopithecus species, which are often referred to as australopiths. Australopithecus anamensis lived about 4.2 million years ago. More is known about another early species, Australopithecus afarensis, which lived between 3.9 and 2.9 million years ago. This species demonstrates a trend in human evolution: the reduction of the dentition and jaw in size. A. afarensis (Figure 29.47a) had smaller canines and molars compared to apes, but these were larger than those of modern humans. Its brain size was 380 to 450 cubic centimeters, approximately the size of a modern chimpanzee brain. It also had prognathic jaws, which is a relatively longer jaw than that of modern humans. In the mid-1970s, the fossil of an adult female A. afarensis was found in the Afar region of Ethiopia and dated to 3.24 million years ago (Figure 29.48). The fossil, which is informally called “Lucy,” is significant because it was the most complete australopith fossil found, with 40 percent of the skeleton recovered.

Australopithecus africanus lived between two and three million years ago. It had a slender build and was bipedal, but had robust arm bones and, like other early hominids, may have spent significant time in trees. Its brain was larger than that of A. afarensis at 500 cubic centimeters, which is slightly less than one-third the size of modern human brains. Two other species, Australopithecus bahrelghazali and Australopithecus garhi, have been added to the roster of australopiths in recent years. A. bahrelghazali is unusual in being the only australopith found in Central Africa.