Animal evolution


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Animal evolution

The theory that modern animals are the modified descendants of animals that formerly existed and that these earlier forms descended from still earlier and different organisms.

Animals are multicellular organisms that feed by ingestion of other organisms or their products, being unable to derive energy through photosynthesis or chemosynthesis. Animals are currently classed into about 30 to 35 phyla, each of which has evolved a distinctive body plan or architecture.

All phyla began as invertebrates, but lineages of the phylum Chordata developed the internal skeletal armature, with spinal column, which was exploited in numerous fish groups and which eventually gave rise to terrestrial vertebrates. The number of phyla is uncertain partly because most of the branching patterns and the ancestral body plans from which putative phyla have arisen are not yet known. For example, arthropods (including crustaceans and insects) may have all diversified from a common ancestor that was a primitive arthropod, in which case they may be grouped into a single phylum; or several arthropod groups may have evolved independently from nonarthropod ancestors, in which case each such group must be considered a separate phylum. So far as known, all animal phyla began in the sea. See Animal, Animal kingdom

Some features of the cells of primitive animals resemble those of the single-celled Protozoa, especially the flagellates, which have long been believed to be animal ancestors. Molecular phylogenies have supported this idea and also suggest that the phylum Coelenterata arose separately from all other phyla that have been studied by this technique. Thus animals may have evolved at least twice from organisms that are not themselves animals, and represent a grade of evolution and not a single branch (clade) of the tree of life. Sponges have also been suspected of an independent origin, and it is possible that some of the extinct fossil phyla arose independently or branched from sponges or cnidarians. See Coelenterata, Porifera, Protozoa

The earliest undoubted animal fossils (the Ediacaran fauna) are soft-bodied, and first appear in marine sediments nearly 650 million years (m.y.) old. This fauna lasted about 50 m.y. and consisted chiefly of cnidarians or cnidarian-grade forms, though it contains a few enigmatic fossils that may represent groups that gave rise to more advanced phyla. Then, nearly 570 m.y. ago, just before and during earliest Cambrian time, a diversification of body architecture began that produced most of the living phyla as well as many extinct groups. The body plans of some of these groups involved mineralized skeletons which, as these are more easily preserved than soft tissues, created for the first time an extensive fossil record. The soft-bodied groups were markedly diversified, though their record is so spotty that their history cannot be traced in detail. A single, exceptionally preserved soft-bodied fauna from the Burgess Shale of British Columbia that is about 530 m.y. old contains not only living soft-bodied worm phyla, but extinct groups that cannot be placed in living phyla and do not seem to be ancestral to them.

Following the early phase of rampant diversification and of some concurrent extinction of phyla and their major branches, the subsequent history of the durably skeletonized groups can be followed in a general way in the marine fossil record. The composition of the fauna changed continually, but three major associations can be seen: one dominated by the arthropodlike trilobites during the early Paleozoic, one dominated by articulate brachiopods and crinoids (Echinodermata) in the remaining Paleozoic, and one dominated by gastropod (snail) and bivalve (clam) mollusks during the Mesozoic and Cenozoic. The mass extinction at the close of the Paleozoic that caused the contractions in so many groups may have extirpated over 90% of marine species and led to a reorganization of marine community structure and composition into a modern mode. Resistance to this and other extinctions seems to have been a major factor in the rise of successive groups to dominance. Annelids, arthropods, and mollusks are the more important invertebrate groups that made the transition to land. The outstanding feature of terrestrial fauna is the importance of the insects, which appeared in the late Paleozoic and later radiated to produce the several million living species, surpassing all other life forms combined in this respect. See Annelida, Arthropoda, Insecta, Mollusca

The phylum Chordata consists largely of animals with a backbone, the Vertebrata, including humans. The group, however, includes some primitive nonvertebrates, the protochordates: lancelets, tunicates, acorn worms, pterobranchs, and possibly the extinct graptolites and conodonts. The interrelationships of these forms are not well understood. With the exception of the colonial graptolites, they are soft-bodied and have only a very limited fossil record. They suggest possible links to the Echinodermata in developmental, biochemical, and morphological features. In addition, some early Paleozoic fossils, the carpoids, have been classified alternatively as chordates and as echinoderms, again suggesting a link. In spite of these various leads, the origin of the chordates remains basically unclear. See Vertebrata

Chordates are characterized by a hollow, dorsal, axial nerve chord, a ventral heart, a system of slits in the larynx that serves variously the functions of feeding and respiration, a postanal swimming tail, and a notochord that is an elongate supporting structure lying immediately below the nerve chord. The protochordates were segmented, although sessile forms such as the tunicates show this only in the swimming, larval phase.

The first vertebrates were fishlike animals in which the pharyngeal slits formed a series of pouches that functioned as respiratory gills. An anterior specialized mouth permitted ingestion of food items large in comparison with those of the filter-feeding protochordates. Vertebrates are first known from bone fragments found in rocks of Cambrian age, but more complete remains have come from the Middle Ordovician. Innovations, related to greater musculoskeletal activity, included the origin of a supporting skeleton of cartilage and bone, a larger brain, and three pairs of cranial sense organs (nose, eyes, and ears). At first the osseous skeleton served as protective scales in the skin, as a supplement to the notochord, and as a casing around the brain. In later vertebrates the adult notochord is largely or wholly replaced by bone, which encloses the nerve chord to form a true backbone. All vertebrates have a heart which pumps blood through capillaries, where exchanges of gases with the external media take place. The blood contains hemoglobin in special cells which carry oxygen and carbon dioxide. In most fishes the blood passes from the heart to the gills and thence to the brain and other parts of the body. In most tetrapods, and in some fishes, blood passes to the lungs, is returned to the heart after oxygenation, and is then pumped to the various parts of the body.

The jawless fish, known as Agnatha, had a sucking-rasping mouth apparatus rather than true jaws. They enjoyed great success from the Late Cambrian until the end of the Devonian. Most were heavily armored, although a few naked forms are known. They were weak swimmers and lived mostly on the bottom. The modern parasitic lampreys and deep-sea scavenging hagfish are the only surviving descendants of these early fish radiations. See Devonian

In the Middle to Late Silurian arose a new type of vertebrate, the Gnathostomata, characterized by true jaws and teeth. They constitute the great majority of fishes and all tetrapod vertebrates. The jaws are modified elements of the front parts of the gill apparatus, and the teeth are modified bony scales from the skin of the mouth. With the development of jaws, a whole new set of ecological opportunities was open to the vertebrates. Along with this, new swimming patterns appeared, made possible by the origin of paired fins, forerunners of which occur in some agnathans.

Four groups of fishes quickly diversified. Of these, the Placodermi and Acanthodii are extinct. The Placodermi were heavily armored fishes, the dominant marine carnivores of the Silurian and Devonian. The Acanthodii were filter-feeders mostly of small size. They are possibly related to the dominant groups of modern fishes, the largely cartilaginous Chondrichthyes (including sharks, rays, and chimaeras) and the Osteichthyes (the higher bony fishes). These also arose in the Late Silurian but diversified later. See Acanthodii, Chondrichthyes

The first land vertebrates, the Amphibia, appeared in the Late Devonian and were derived from an early group of osteichthyans called lobe-finned fishes, of which two kinds survive today, the Dipnoi or lungfishes, and the crossopterygian coelacanth Latimeria. They were lung-breathing fishes that lived in shallow marine waters and in swamps and marshes. The first amphibians fed and reproduced in or near the water. True land vertebrates, Reptilia, with a modified (amniote) egg that could survive on land, probably arose in the Mississippian. See Amniota, Amphibia, Crossopterygii, Dipnoi

By the Middle Pennsylvanian a massive radiation of reptiles was in process. The most prominent reptiles belong in the Diapsida: dinosaurs, lizards and snakes, and pterosaurs (flying reptiles). The birds, Aves, which diverged from the dinosaur radiation in the Late Triassic or Early Jurassic, are considered to be feathered dinosaurs, and thus members of the Diapsida, whereas older authorities prefer to treat them as a separate case. In addition, there were several Mesozoic radiations of marine reptiles such as ichthyosaurs and plesiosaurs. Turtles (Chelonia) first appeared in the Triassic and have been highly successful ever since. See Aves, Reptilia

The line leading to mammals can be traced to primitive Pennsylvanian reptiles, Synapsida, which diversified and spread worldwide during the Permian and Triassic. The first true mammals, based on characteristics of jaw, tooth, and ear structure, arose in the Late Triassic. Derived mammals, marsupials (Metatheria) and placentals (Eutheria), are known from the Late Cretaceous, but mammalian radiations began only in the early Cenozoic. By the end of the Eocene, all the major lines of modern mammals had become established. Molecular analyses (blood proteins, deoxyribonucleic acid, ribonucleic acid) of living mammals show that the most primitive group of placentals is the edentates (sloths, armadillos, and anteaters). An early large radiation included the rodents, primates (including monkeys, apes, and humans), and bats, possibly all closely related to the insectivores and carnivores. The newest radiations of mammals are of elephants and sea cows, while the whales are related to the artiodactyls (cattle, camels). See Mammalia

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