Animal evolution began in the ocean over 600 million years ago with tiny creatures that probably do not resemble any living organism today. Since then, animals have evolved into a highly diverse kingdom. Although over one million extant (currently living) species of animals have been identified, scientists are continually discovering more species as they explore ecosystems around the world. The number of extant species is estimated to be between 3 and 30 million.

But what is an animal? While we can easily identify dogs, birds, fish, spiders, and worms as animals, other organisms, such as corals and sponges, are not as easy to classify. Animals vary in complexity—from sea sponges to crickets to chimpanzees—and scientists are faced with the difficult task of classifying them within a unified system. They must identify traits that are common to all animals as well as traits that can be used to distinguish among related groups of animals. The animal classification system characterizes animals based on their anatomy, morphology, evolutionary history, features of embryological development, and genetic makeup. This classification scheme is constantly developing as new information about species arises. Understanding and classifying the great variety of living species help us better understand how to conserve the diversity of life on earth.

Even though members of the animal kingdom are incredibly diverse, animals share common features that distinguish them from organisms in other kingdoms. All animals are eukaryotic, multicellular organisms, and almost all animals have specialized tissues. Most animals are motile, at least during certain life stages. Animals require a source of food to grow and develop. All animals are heterotrophic, ingesting living or dead organic matter. This form of obtaining energy distinguishes them from autotrophic organisms, such as most plants, which make their own nutrients through photosynthesis and from fungi that digest their food externally. Animals may be carnivores, herbivores, omnivores, or parasites (Figure 15.2). Most animals reproduce sexually: The offspring pass through a series of developmental stages that establish a determined body plan, unlike plants, for example, in which the exact shape of the body is indeterminate. The body plan refers to the shape of an animal.

Complex Tissue Structure

A hallmark trait of animals is specialized structures that are differentiated to perform unique functions. As multicellular organisms, most animals develop specialized cells that group together into tissues with specialized functions. A tissue is a collection of similar cells that had a common embryonic origin. There are four main types of animal tissues: nervous, muscle, connective, and epithelial. Nervous tissue contains neurons, or nerve cells, which transmit nerve impulses. Muscle tissue contracts to cause all types of body movement from locomotion of the organism to movements within the body itself. Animals also have specialized connective tissues that provide many functions, including transport and structural support. Examples of connective tissues include blood and bone. Connective tissue is comprised of cells separated by extracellular material made of organic and inorganic materials, such as the protein and mineral deposits of bone. Epithelial tissue covers the internal and external surfaces of organs inside the animal body and the external surface of the body of the organism.

View this video to watch a presentation by biologist E.O. Wilson on the importance of animal diversity.

Animal Reproduction and Development

Most animals have diploid body (somatic) cells and a small number of haploid reproductive (gamete) cells produced through meiosis. Some exceptions exist: For example, in bees, wasps, and ants, the male is haploid because it develops from an unfertilized egg. Most animals undergo sexual reproduction, while many also have mechanisms of asexual reproduction.

Sexual Reproduction and Embryonic Development

Almost all animal species are capable of reproducing sexually; for many, this is the only mode of reproduction possible. This distinguishes animals from fungi, protists, and bacteria, where asexual reproduction is common or exclusive. During sexual reproduction, the male and female gametes of a species combine in a process called fertilization. Typically, the small, motile male sperm travels to the much larger, sessile female egg. Sperm form is diverse and includes cells with flagella or amoeboid cells to facilitate motility. Fertilization and fusion of the gamete nuclei produce a zygote. Fertilization may be internal, especially in land animals, or external, as is common in many aquatic species.

After fertilization, a developmental sequence ensues as cells divide and differentiate. Many of the events in development are shared in groups of related animal species, and these events are one of the main ways scientists classify high-level groups of animals. During development, animal cells specialize and form tissues, determining their future morphology and physiology. In many animals, such as mammals, the young resemble the adult. Other animals, such as some insects and amphibians, undergo complete metamorphosis in which individuals enter one or more larval stages. For these animals, the young and the adult have different diets and sometimes habitats. In other species, a process of incomplete metamorphosis occurs in which the young somewhat resemble the adults and go through a series of stages separated by molts (shedding of the skin) until they reach the final adult form.

Asexual Reproduction

Asexual reproduction, unlike sexual reproduction, produces offspring genetically identical to each other and to the parent. A number of animal species—especially those without backbones, but even some fish, amphibians, and reptiles—are capable of asexual reproduction. Asexual reproduction, except for occasional identical twinning, is absent in birds and mammals. The most common forms of asexual reproduction for stationary aquatic animals include budding and fragmentation, in which part of a parent individual can separate and grow into a new individual. In contrast, a form of asexual reproduction found in certain invertebrates and rare vertebrates is called parthenogenesis (or “virgin beginning”), in which unfertilized eggs develop into new offspring.

Classification Features of Animals

Scientists have developed a classification scheme that categorizes all members of the animal kingdom, although there are exceptions to most “rules” governing animal classification (Figure 15.3). Animals have been traditionally classified according to two characteristics: body plan and developmental pathway. The major feature of the body plan is its symmetry: how the body parts are distributed along the major body axis. Symmetrical animals can be divided into roughly equivalent halves along at least one axis. Developmental characteristics include the number of germ tissue layers formed during development, the origin of the mouth and anus, the presence or absence of an internal body cavity, and other features of embryological development, such as larval types or whether or not periods of growth are interspersed with molting.

Body Symmetry

Animals may be asymmetrical, radial, or bilateral in form (Figure 15.4). Asymmetrical animals are animals with no pattern or symmetry; an example of an asymmetrical animal is a sponge (Figure 15.4a). An organism with radial symmetry (Figure 15.4b) has a longitudinal (up-and-down) orientation: Any plane cut along this up–down axis produces roughly mirror-image halves. An example of an organism with radial symmetry is a sea anemone.

Bilateral symmetry is illustrated in Figure 15.4c using a goat. The goat also has upper and lower sides to it, but they are not symmetrical. A vertical plane cut from front to back separates the animal into roughly mirror-image right and left sides. Animals with bilateral symmetry also have a “head” and “tail” (anterior versus posterior) and a back and underside (dorsal versus ventral).

Watch this video to see a quick sketch of the different types of body symmetry.

Layers of Tissues

Most animal species undergo a layering of early tissues during embryonic development. These germ layers are formed during gastrulation, and each germ layer typically gives rise to specific types of embryonic tissues and organs. Animals develop either two or three embryonic germs layers (Figure 15.5). The animals that display radial, biradial, or rotational symmetry develop two germ layers, an inner layer (endoderm or mesendoderm) and an outer layer (ectoderm). These animals are called diploblasts, and have a nonliving middle layer between the endoderm and ectoderm (although individual cells may be distributed through this middle layer, there is no coherent third layer of tissue). The four clades considered to be diploblastic have different levels of complexity and different developmental pathways, although there is little information about development in Placozoa. More complex animals (usually those with bilateral symmetry) develop three tissue layers: an inner layer (endoderm), an outer layer (ectoderm), and a middle layer (mesoderm). Animals with three tissue layers are called triploblasts.

Each of the three germ layers is programmed to give rise to specific body tissues and organs, although there are variations on these themes. Generally speaking, the endoderm gives rise to the lining of the digestive tract (including the stomach, intestines, liver, and pancreas), as well as to the lining of the trachea, bronchi, and lungs of the respiratory tract, along with a few other structures. The ectoderm develops into the outer epithelial covering of the body surface, the central nervous system, and a few other structures. The mesoderm is the third germ layer; it forms between the endoderm and ectoderm in triploblasts. This germ layer gives rise to all specialized muscle tissues (including the cardiac tissues and muscles of the intestines), connective tissues such as the skeleton and blood cells, and most other visceral organs such as the kidneys and the spleen. Diploblastic animals may have cell types that serve multiple functions, such as epitheliomuscular cells, which serve as a covering as well as contractile cells.

Presence or Absence of a Coelom

Triploblasts may develop an internal body cavity derived from mesoderm, called a coelom (pr. see-LŌM). This epithelial- lined cavity is a space, usually filled with fluid, which lies between the digestive system and the body wall. It houses organs such as the kidneys and spleen, and contains the circulatory system. Triploblasts that do not develop a coelom are called acoelomates, and their mesoderm region is completely filled with tissue, although they have a gut cavity. Examples of acoelomates include the flatworms. Animals with a true coelom are called eucoelomates (or coelomates) (Figure 15.6). A true coelom arises entirely within the mesoderm germ layer. Animals such as earthworms, snails, insects, starfish, and vertebrates are all eucoelomates. A third group of triploblasts has a body cavity that is derived partly from mesoderm and partly from endoderm tissue. These animals are called pseudocoelomates. Roundworms are examples of pseudocoelomates. New data on the relationships of pseudocoelomates suggest that these phyla are not closely related and so the evolution of the pseudocoelom must have occurred more than once (Figure 15.3). True coelomates can be further characterized based on features of their early embryological development.

Embryonic Development of the Mouth

Bilaterally symmetrical, triploblastic eucoelomates can be divided into two groups based on differences in their early embryonic development. Protostomes include phyla such as arthropods, mollusks, and annelids. Deuterostomes include the chordates and echinoderms. These two groups are named from which opening of the digestive cavity develops first: mouth or anus. The word protostome comes from Greek words meaning “mouth first,” and deuterostome originates from words meaning “mouth second” (in this case, the anus develops first). This difference reflects the fate of a structure called the blastopore (Figure 15.7), which becomes the mouth in protostomes and the anus in deuterostomes. Other developmental characteristics differ between protostomes and deuterostomes, including the mode of formation of the coelom and the early cell division of the embryo.