第29章脊椎動物

Mammals are synapsids, meaning they have a single, ancestrally fused, postorbital opening in the skull. They are the only living synapsids, as earlier forms became extinct by the Jurassic period. The early non-mammalian synapsids can be divided into two groups, the pelycosaurs and the therapsids. Within the therapsids, a group called the cynodonts are thought to have been the ancestors of mammals (Figure 29.38).

As with birds, a key characteristic of synapsids is endothermy, rather than the ectothermy seen in many other vertebrates (such as fish, amphibians, and most reptiles). The increased metabolic rate required to internally modify body temperature likely went hand-in-hand with changes to certain skeletal structures that improved food processing and ambulation. The later synapsids, which had more evolved characteristics unique to mammals, possess cheeks for holding food and heterodont teeth, which are specialized for chewing, mechanically breaking down food to speed digestion, and releasing the energy needed to produce heat. Chewing also requires the ability to breathe at the same time, which is facilitated by the presence of a secondary palate (comprising the bony palate and the posterior continuation of the soft palate). The secondary palate separates the area of the mouth where chewing occurs from the area above where respiration occurs, allowing breathing to proceed uninterrupted while the animal is chewing. A secondary palate is not found in pelycosaurs but is present in cynodonts and mammals. The jawbone also shows changes from early synapsids to later ones. The zygomatic arch, or cheekbone, is present in mammals and advanced therapsids such as cynodonts, but is not present in pelycosaurs. The presence of the zygomatic arch suggests the presence of masseter muscles, which close the jaw and function in chewing.

In the appendicular skeleton, the shoulder girdle of therian mammals is modified from that of other vertebrates in that it does not possess a procoracoid bone or an interclavicle, and the scapula is the dominant bone.

Mammals evolved from therapsids in the late Triassic period, as the earliest known mammal fossils are from the early Jurassic period, some 205 million years ago. One group of transitional mammals was the morganucodonts, small nocturnal insectivores. The jaws of morganucodonts were “transitional,” with features of both reptilian and mammalian jaws (Figure 29.39). Like modern mammals, the morganucodonts had differentiated teeth and were diphyodonts. Mammals first began to diversify in the Mesozoic era, from the Jurassic to the Cretaceous periods. Even some small gliding mammals appear in the fossil record during this time period. However, most of the Jurassic mammals were extinct by the end of the Mesozoic. During the Cretaceous period, another radiation of mammals began and continued through the Cenozoic era, about 65 million years ago.

There are three major groups of living mammals: monotremes (prototheria), marsupials (metatheria), and placental (eutheria) mammals. The eutherians and the marsupials together comprise a clade of therian mammals, with the monotremes forming a sister clade to both metatherians and eutherians.

There are very few living species of monotremes: the platypus and four species of echidnas, or spiny anteaters. The leathery-beaked platypus belongs to the family Ornithorhynchidae (“bird beak”), whereas echidnas belong to the family Tachyglossidae (“sticky tongue”) (Figure 29.40). The platypus and one species of echidna are found in Australia, and the other species of echidna are found in New Guinea. Monotremes are unique among mammals because they lay eggs, rather than giving birth to live young. The shells of their eggs are not like the hard shells of birds, but have a leathery shell, similar to the shells of reptile eggs. Monotremes retain their eggs through about two-thirds of the developmental period, and then lay them in nests. A yolk-sac placenta helps support development. The babies hatch in a fetal state and complete their development in the nest, nourished by milk secreted by mammary glands opening directly to the skin. Monotremes, except for young platypuses, do not have teeth. Body temperature in the monotreme species is maintained at about 30°C, considerably lower than the average body temperature of marsupial and placental mammals, which are typically between 35 and 38°C.

Over 2/3 of the approximately 330 living species of marsupials are found in Australia, New Guinea, and surrounding islands, with the rest, nearly all various types of opossum, found in the Americas, especially South America. Australian marsupials include the kangaroo, koala, bandicoot, Tasmanian devil (Figure 29.41), and several other species. Like monotremes, the embryos of marsupials are nourished during a short gestational period (about a month in kangaroos) by a yolk-sac placenta, but with no intervening egg shell. Some marsupial embryos can enter an embryonic diapause, and delay implantation, suspending development until implantation is completed. Marsupial young are also effectively fetal at birth. Most, but not all, species of marsupials possess a pouch in which the very premature young reside, receiving milk and continuing their development. In kangaroos, the young joeys continue to nurse for about a year and a half.

Eutherians (placentals) are the most widespread and numerous of the mammals, occurring throughout the world. Eutherian mammals are sometimes called “placental mammals” because all species possess a complex chorioallantoic placenta that connects a fetus to the mother, allowing for gas, fluid, and nutrient exchange. There are about 6,000 species of placental mammals in 18 to 20 orders with various adaptations for burrowing, flying, swimming, hunting, running, and climbing. In the evolutionary sense, they have been incredibly successful in form, diversity, and abundance. The eutherian mammals are classified in two major clades, the Atlantogenata and the Boreoeutheria. The Atlantogeneta include the Afrotheria (e.g., elephants, hyraxes, and manatees) and the Xenarthra (anteaters, armadillos, and sloths). The Boreoeutheria contain two large groups, the Euarchontoglires and the Laurasiatheria. Familiar orders in the Euarchontoglires are the Scandentia (tree shrews), Rodentia (rats, mice, squirrels, porcupines), Lagomorpha (rabbits and hares), and the Primates (including humans). Major Laurasiatherian orders include the Perissodactyla (e.g., horses and rhinos), the Cetartiodactyla (e.g., cows, giraffes, pigs, hippos, and whales), the Carnivora (e.g., cats, dogs, and bears), and the Chiroptera (bats and flying foxes). The two largest orders are the rodents (about 2,000 species) and bats (about 1,400 species), which together constitute almost 60 percent of all eutherian species.