Chapter 15. Diversity of Animals

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, 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.

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.

Figure 15.4. 

Animals exhibit different types of body symmetry. The (a) sponge is asymmetrical and has no planes of symmetry, the (b) sea anemone has radial symmetry with multiple planes of symmetry, and the (c) goat has bilateral symmetry with one plane of symmetry.

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).

Concept in Action

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

Most animal species undergo a layering of early tissues during embryonic development. These layers are called germ layers. Each layer develops into a specific set of tissues and organs. Animals develop either two or three embryonic germs layers (Figure 15.5). The animals that display radial symmetry develop two germ layers, an inner layer (endoderm) and an outer layer (ectoderm). These animals are called diploblasts. Animals with bilateral symmetry develop three germ layers: an inner layer (endoderm), an outer layer (ectoderm), and a middle layer (mesoderm). Animals with three germ layers are called triploblasts.

Figure 15.5. 

During embryogenesis, diploblasts develop two embryonic germ layers: an ectoderm and an endoderm. Triploblasts develop a third layer—the mesoderm—between the endoderm and ectoderm.

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.

Figure 15.6. 

Triploblasts may be acoelomates, eucoelomates, or pseudocoelomates. Eucoelomates have a body cavity within the mesoderm, called a coelom, which is lined with mesoderm tissue. Pseudocoelomates have a similar body cavity, but it is lined with mesoderm and endoderm tissue. (credit a: modification of work by Jan Derk; credit b: modification of work by NOAA; credit c: modification of work by USDA, ARS)

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.

Figure 15.7. 

Eucoelomates can be divided into two groups, protostomes and deuterostomes, based on their early embryonic development. Two of these differences include the origin of the mouth opening and the way in which the coelom is formed.

Despite their lack of complexity, sponges are clearly successful organisms, having persisted on Earth for more than half a billion years. Lacking a true digestive system, sponges depend on the intracellular digestive processes of their choanocytes for their energy intake. The limit of this type of digestion is that food particles must be smaller than individual cells. Gas exchange, circulation, and excretion occur by diffusion between cells and the water.

Sponges reproduce both sexually and asexually. Asexual reproduction is either by fragmentation (in which a piece of the sponge breaks off and develops into a new individual), or budding (an outgrowth from the parent that eventually detaches). A type of asexual reproduction found only in freshwater sponges occurs through the formation of gemmules, clusters of cells surrounded by a tough outer layer. Gemmules survive hostile environments and can attach to a substrate and grow into a new sponge.

Sponges are monoecious (or hermaphroditic), meaning one individual can produce both eggs and sperm. Sponges may be sequentially hermaphroditic, producing eggs first and sperm later. Eggs arise from amoebocytes and are retained within the spongocoel, whereas sperm arise from choanocytes and are ejected through the osculum. Sperm carried by water currents fertilize the eggs of other sponges. Early larval development occurs within the sponge, and free-swimming larvae are then released through the osculum. This is the only time that sponges exhibit mobility. Sponges are sessile as adults and spend their lives attached to a fixed substrate.

Concept in Action

Watch this video that demonstrates the feeding of sponges.

All cnidarians have two tissue layers. The outer layer is called the epidermis, whereas the inner layer is called the gastrodermis and lines the digestive cavity. Between these two layers is a non-living, jelly-like mesoglea. There are differentiated cell types in each tissue layer, such as nerve cells, enzyme-secreting cells, and nutrient-absorbing cells, as well as intercellular connections between the cells. However, organs and organ systems are not present in this phylum.

The nervous system is primitive, with nerve cells scattered across the body in a network. The function of the nerve cells is to carry signals from sensory cells and to contractile cells. Groups of cells in the nerve net form nerve cords that may be essential for more rapid transmission. Cnidarians perform extracellular digestion, with digestion completed by intracellular digestive processes. Food is taken into the gastrovascular cavity, enzymes are secreted into the cavity, and the cells lining the cavity absorb the nutrient products of the extracellular digestive process. The gastrovascular cavity has only one opening that serves as both a mouth and an anus (an incomplete digestive system). Like the sponges, Cnidarian cells exchange oxygen, carbon dioxide, and nitrogenous wastes by diffusion between cells in the epidermis and gastrodermis with water.

The phylum Cnidaria contains about 10,000 described species divided into four classes: Anthozoa, Scyphozoa, Cubozoa, and Hydrozoa.

The class Anthozoa includes all cnidarians that exhibit a sessile polyp body plan only; in other words, there is no medusa stage within their life cycle. Examples include sea anemones, sea pens, and corals, with an estimated number of 6,100 described species. Sea anemones are usually brightly colored and can attain a size of 1.8 to 10 cm in diameter. These animals are usually cylindrical in shape and are attached to a substrate. A mouth opening is surrounded by tentacles bearing cnidocytes (Figure 15.12).

Figure 15.12. 

Sea anemones are cnidarians of class Anthozoa. (credit: “Dancing With Ghosts”/Flickr)

Scyphozoans include all the jellies and are motile and exclusively marine with about 200 described species. The medusa is the dominant stage in the life cycle, although there is also a polyp stage. Species range from 2 cm in length to the largest scyphozoan species, Cyanea capillata, at 2 m across. Jellies display a characteristic bell-like body shape (Figure 15.13).

Figure 15.13. 

Scyphozoans include the jellies. (credit: “Jimg944″/Flickr)

Concept in Action

Identify the life cycle stages of jellies using this video animation game from the New England Aquarium.

The class Cubozoa includes jellies that are square in cross-section and so are known as “box jellyfish.” These species may achieve sizes of 15–25 cm. Cubozoans are anatomically similar to the jellyfish. A prominent difference between the two classes is the arrangement of tentacles. Cubozoans have muscular pads called pedalia at the corners of the square bell canopy, with one or more tentacles attached to each pedalium. In some cases, the digestive system may extend into the pedalia. Cubozoans typically exist in a polyp form that develops from a larva. The polyps may bud to form more polyps and then transform into the medusoid forms.

Concept in Action

Watch this video to learn more about the deadly toxins of the box jellyfish.

Hydrozoa includes nearly 3,500 species,^[24] most of which are marine. Most species in this class have both polyp and medusa forms in their life cycle. Many hydrozoans form colonies composed of branches of specialized polyps that share a gastrovascular cavity. Colonies may also be free-floating and contain both medusa and polyp individuals in the colony, as in the Portuguese Man O’War (Physalia) or By-the-Wind Sailor (Velella). Other species are solitary polyps or solitary medusae. The characteristic shared by all of these species is that their gonads are derived from epidermal tissue, whereas in all other cnidarians, they are derived from gastrodermal tissue (Figure 15.14ab).

Figure 15.14. 

A (a) box jelly is an example from class Cubozoa. The (b) hydra is from class Hydrozoa. (credit b: scale-bar data from Matt Russell)

Free-living species of flatworms are predators or scavengers, whereas parasitic forms feed from the tissues of their hosts. Most flatworms have an incomplete digestive system with an opening, the “mouth,” that is also used to expel digestive system wastes. Some species also have an anal opening. The gut may be a simple sac or highly branched. Digestion is extracellular, with enzymes secreted into the space by cells lining the tract, and digested materials taken into the same cells by phagocytosis. One group, the cestodes, does not have a digestive system, because their parasitic lifestyle and the environment in which they live (suspended within the digestive cavity of their host) allows them to absorb nutrients directly across their body wall. Flatworms have an excretory system with a network of tubules throughout the body that open to the environment and nearby flame cells, whose cilia beat to direct waste fluids concentrated in the tubules out of the body. The system is responsible for regulation of dissolved salts and excretion of nitrogenous wastes. The nervous system consists of a pair of nerve cords running the length of the body with connections between them and a large ganglion or concentration of nerve cells at the anterior end of the worm; here, there may also be a concentration of photosensory and chemosensory cells (Figure 15.15).

Figure 15.15. 

This planarian is a free-living flatworm that has an incomplete digestive system, an excretory system with a network of tubules throughout the body, and a nervous system made up of nerve cords running the length of the body with a concentration of nerves and photosensory and chemosensory cells at the anterior end.

Since there is no circulatory or respiratory system, gas and nutrient exchange is dependent on diffusion and intercellular junctions. This necessarily limits the thickness of the body in these organisms, constraining them to be “flat” worms. Most flatworm species are monoecious (hermaphroditic, possessing both sets of sex organs), and fertilization is typically internal. Asexual reproduction is common in some groups in which an entire organism can be regenerated from just a part of itself.

Flatworms are traditionally divided into four classes: Turbellaria, Monogenea, Trematoda, and Cestoda (Figure 15.16). The turbellarians include mainly free-living marine species, although some species live in freshwater or moist terrestrial environments. The simple planarians found in freshwater ponds and aquaria are examples. The epidermal layer of the underside of turbellarians is ciliated, and this helps them move. Some turbellarians are capable of remarkable feats of regeneration in which they may regrow the body, even from a small fragment.

Figure 15.16. 

Phylum Platyhelminthes is divided into four classes: (a) Bedford’s Flatworm (Pseudobiceros bedfordi) and the (b) planarian belong to class Turbellaria; (c) the Trematoda class includes about 20,000 species, most of which are parasitic; (d) class Cestoda includes tapeworms such as this Taenia saginata; and the parasitic class Monogenea (not shown). (credit a: modification of work by Jan Derk; credit c: modification of work by “Sahaquiel9102”/Wikimedia Commons; credit d: modification of work by CDC)

The monogeneans are external parasites mostly of fish with life cycles consisting of a free-swimming larva that attaches to a fish to begin transformation to the parasitic adult form. They have only one host during their life, typically of just one species. The worms may produce enzymes that digest the host tissues or graze on surface mucus and skin particles. Most monogeneans are hermaphroditic, but the sperm develop first, and it is typical for them to mate between individuals and not to self-fertilize.

The trematodes, or flukes, are internal parasites of mollusks and many other groups, including humans. Trematodes have complex life cycles that involve a primary host in which sexual reproduction occurs and one or more secondary hosts in which asexual reproduction occurs. The primary host is almost always a mollusk. Trematodes are responsible for serious human diseases including schistosomiasis, caused by a blood fluke (Schistosoma). The disease infects an estimated 200 million people in the tropics and leads to organ damage and chronic symptoms including fatigue. Infection occurs when a human enters the water, and a larva, released from the primary snail host, locates and penetrates the skin. The parasite infects various organs in the body and feeds on red blood cells before reproducing. Many of the eggs are released in feces and find their way into a waterway where they are able to reinfect the primary snail host.

The cestodes, or tapeworms, are also internal parasites, mainly of vertebrates. Tapeworms live in the intestinal tract of the primary host and remain fixed using a sucker on the anterior end, or scolex, of the tapeworm body. The remaining body of the tapeworm is made up of a long series of units called proglottids, each of which may contain an excretory system with flame cells, but will contain reproductive structures, both male and female. Tapeworms do not have a digestive system, they absorb nutrients from the food matter passing them in the host’s intestine. Proglottids are produced at the scolex and are pushed to the end of the tapeworm as new proglottids form, at which point, they are “mature” and all structures except fertilized eggs have degenerated. Most reproduction occurs by cross-fertilization. The proglottid detaches and is released in the feces of the host. The fertilized eggs are eaten by an intermediate host. The juvenile worms emerge and infect the intermediate host, taking up residence, usually in muscle tissue. When the muscle tissue is eaten by the primary host, the cycle is completed. There are several tapeworm parasites of humans that are acquired by eating uncooked or poorly cooked pork, beef, and fish.

In nematodes, the excretory system is not specialized. Nitrogenous wastes are removed by diffusion. In marine nematodes, regulation of water and salt is achieved by specialized glands that remove unwanted ions while maintaining internal body fluid concentrations.

Most nematodes have four nerve cords that run along the length of the body on the top, bottom, and sides. The nerve cords fuse in a ring around the pharynx, to form a head ganglion or “brain” of the worm, as well as at the posterior end to form the tail ganglion. Beneath the epidermis lies a layer of longitudinal muscles that permits only side-to-side, wave-like undulation of the body.

Concept in Action

View this video to see nematodes move about and feed on bacteria.

Nematodes employ a diversity of sexual reproductive strategies depending on the species; they may be monoecious, dioecious (separate sexes), or may reproduce asexually by parthenogenesis. Caenorhabditis elegans is nearly unique among animals in having both self-fertilizing hermaphrodites and a male sex that can mate with the hermaphrodite.

A unique feature of arthropods is the presence of a segmented body with fusion of certain sets of segments to give rise to functional segments. Fused segments may form a head, thorax, and abdomen, or a cephalothorax and abdomen, or a head and trunk. The coelom takes the form of a hemocoel (or blood cavity). The open circulatory system, in which blood bathes the internal organs rather than circulating in vessels, is regulated by a two-chambered heart. Respiratory systems vary, depending on the group of arthropod: Insects and myriapods use a series of tubes ( tracheae) that branch throughout the body, open to the outside through openings called spiracles, and perform gas exchange directly between the cells and air in the tracheae. Aquatic crustaceans use gills, arachnids employ “book lungs,” and aquatic chelicerates use “book gills.” The book lungs of arachnids are internal stacks of alternating air pockets and hemocoel tissue shaped like the pages of a book. The book gills of crustaceans are external structures similar to book lungs with stacks of leaf-like structures that exchange gases with the surrounding water (Figure 15.19).

Figure 15.19. 

The book lungs of (a) arachnids are made up of alternating air pockets and hemocoel tissue shaped like a stack of books. The book gills of (b) crustaceans are similar to book lungs but are external so that gas exchange can occur with the surrounding water. (credit a: modification of work by Ryan Wilson based on original work by John Henry Comstock; credit b: modification of work by Angel Schatz)

Phylum Arthropoda includes animals that have been successful in colonizing terrestrial, aquatic, and aerial habitats. The phylum is further classified into five subphyla: Trilobitomorpha (trilobites), Hexapoda (insects and relatives), Myriapoda (millipedes, centipedes, and relatives), Crustacea (crabs, lobsters, crayfish, isopods, barnacles, and some zooplankton), and Chelicerata (horseshoe crabs, arachnids, scorpions, and daddy longlegs). Trilobites are an extinct group of arthropods found from the Cambrian period (540–490 million years ago) until they became extinct in the Permian (300–251 million years ago) that are probably most closely related to the Chelicerata. The 17,000 described species have been identified from fossils (Figure 15.18).

The Hexapoda have six legs (three pairs) as their name suggests. Hexapod segments are fused into a head, thorax, and abdomen (Figure 15.20). The thorax bears the wings and three pairs of legs. The insects we encounter on a daily basis—such as ants, cockroaches, butterflies, and bees—are examples of Hexapoda.

Figure 15.20. 

In this basic anatomy of a hexapod, note that insects have a developed digestive system (yellow), a respiratory system (blue), a circulatory system (red), and a nervous system (purple).

Subphylum Myriapoda includes arthropods with legs that may vary in number from 10 to 750. This subphylum includes 13,000 species; the most commonly found examples are millipedes and centipedes. All myriapods are terrestrial animals and prefer a humid environment (Figure 15.21).

Figure 15.21. 

(a) The centipede Scutigera coleoptrata has up to 15 pairs of legs. (b) This North American millipede (Narceus americanus) bears many legs, although not one thousand, as its name might suggest. (credit a: modification of work by Bruce Marlin; credit b: modification of work by Cory Zanker)

Crustaceans, such as shrimp, lobsters, crabs, and crayfish, are the dominant aquatic arthropods. A few crustaceans are terrestrial species like the pill bugs or sow bugs. The number of described crustacean species stands at about 47,000.^[25]

Although the basic body plan in crustaceans is similar to the Hexapoda—head, thorax, and abdomen—the head and thorax may be fused in some species to form a cephalothorax, which is covered by a plate called the carapace (Figure 15.22). The exoskeleton of many species is also infused with calcium carbonate, which makes it even stronger than in other arthropods. Crustaceans have an open circulatory system in which blood is pumped into the hemocoel by the dorsal heart. Most crustaceans typically have separate sexes, but some, like barnacles, may be hermaphroditic. Serial hermaphroditism, in which the gonad can switch from producing sperm to ova, is also found in some crustacean species. Larval stages are seen in the early development of many crustaceans. Most crustaceans are carnivorous, but detritivores and filter feeders are also common.

Figure 15.22. 

The crayfish is an example of a crustacean. It has a carapace around the cephalothorax and the heart in the dorsal thorax area. (credit: Jane Whitney)

Subphylum Chelicerata includes animals such as spiders, scorpions, horseshoe crabs, and sea spiders. This subphylum is predominantly terrestrial, although some marine species also exist. An estimated 103,000^[26] described species are included in subphylum Chelicerata.

The body of chelicerates may be divided into two parts and a distinct “head” is not always discernible. The phylum derives its name from the first pair of appendages: the chelicerae (Figure 15.23a), which are specialized mouthparts. The chelicerae are mostly used for feeding, but in spiders, they are typically modified to inject venom into their prey (Figure 15.23b). As in other members of Arthropoda, chelicerates also utilize an open circulatory system, with a tube-like heart that pumps blood into the large hemocoel that bathes the internal organs. Aquatic chelicerates utilize gill respiration, whereas terrestrial species use either tracheae or book lungs for gaseous exchange.

Figure 15.23. 

(a) The chelicerae (first set of appendages) are well developed in the Chelicerata, which includes scorpions (a) and spiders (b). (credit a: modification of work by Kevin Walsh; credit b: modification of work by Marshal Hedin)

Concept in Action

Click through this lesson on arthropods to explore interactive habitat maps and more.

This phylum is comprised of seven classes: Aplacophora, Monoplacophora, Polyplacophora, Bivalvia, Gastropoda, Cephalopoda, and Scaphopoda.

Class Aplacophora (“bearing no plates”) includes worm-like animals living mostly on deep ocean bottoms. These animals lack a shell but have aragonite spicules on their skin. Members of class Monoplacophora (“bearing one plate”) have a single, cap-like shell enclosing the body. The monoplacophorans were believed extinct and only known as fossils until the discovery of Neopilina galatheae in 1952. Today, scientists have identified nearly two dozen living species.

Animals in the class Polyplacophora (“bearing many plates”) are commonly known as “chitons” and bear an armor-like, eight-plated shell (Figure 15.25). These animals have a broad, ventral foot that is adapted for attachment to rocks and a mantle that extends beyond the shell in the form of a girdle. They breathe with ctenidia (gills) present ventrally. These animals have a radula modified for scraping. A single pair of nephridia for excretion is present.

Figure 15.25. 

This chiton from the class Polyplacophora has the eight-plated shell indicative of its class. (credit: Jerry Kirkhart)

Class Bivalvia (“two shells”) includes clams, oysters, mussels, scallops, and geoducks. They are found in marine and freshwater habitats. As the name suggests, bivalves are enclosed in a pair of shells (or valves) that are hinged at the dorsal side. The body is flattened on the sides. They feed by filtering particles from water and a radula is absent. They exchange gases using a pair of ctenidia, and excretion and osmoregulation are carried out by a pair of nephridia. In some species, the posterior edges of the mantle may fuse to form two siphons that inhale and exhale water. Some bivalves like oysters and mussels have the unique ability to secrete and deposit a calcareous nacre or “mother of pearl” around foreign particles that enter the mantle cavity. This property is commercially exploited to produce pearls.

Concept in Action

Watch animations of clams and mussels feeding to understand more about bivalves.

Gastropods (“stomach foot”) include well-known mollusks like snails, slugs, conchs, sea hares, and sea butterflies. Gastropods include shell-bearing species as well as species with a reduced shell. These animals are asymmetrical and usually present a coiled shell (Figure 15.26).

Figure 15.26. 

(a) Like many gastropods, this snail has a stomach foot and a coiled shell. (b) This slug, which is also a gastropod, lacks a shell. (credit a: modification of work by Murray Stevenson; credit b: modification of work by Rosendahl)

The visceral mass in the shelled species is characteristically twisted and the foot is modified for crawling. Most gastropods bear a head with tentacles that support eyes. A complex radula is used to scrape food particles from the substrate. The mantle cavity encloses the ctenidia as well as a pair of nephridia.

The class Cephalopoda (“head foot” animals) includes octopuses, squids, cuttlefish, and nautilus. Cephalopods include shelled and reduced-shell groups. They display vivid coloration, typically seen in squids and octopuses, which is used for camouflage. The ability of some octopuses to rapidly adjust their colors to mimic a background pattern or to startle a predator is one of the more awe-inspiring feats of these animals. All animals in this class are predators and have beak-like jaws. All cephalopods have a well-developed nervous system, complex eyes, and a closed circulatory system. The foot is lobed and developed into tentacles and a funnel, which is used for locomotion. Suckers are present on the tentacles in octopuses and squid. Ctenidia are enclosed in a large mantle cavity and are serviced by large blood vessels, each with its own heart.

Cephalopods (Figure 15.27) are able to move quickly via jet propulsion by contracting the mantle cavity to forcefully eject a stream of water. Cephalopods have separate sexes, and the females of some species care for the eggs for an extended period of time. Although the shell is much reduced and internal in squid and cuttlefish, and absent altogether in octopus, nautilus live inside a spiral, multi-chambered shell that is filled with gas or water to regulate buoyancy.

Figure 15.27. 

The (a) nautilus, (b) giant cuttlefish, (c) reef squid, and (d) blue-ring octopus are all members of the class Cephalopoda. (credit a: modification of work by J. Baecker; credit b: modification of work by Adrian Mohedano; credit c: modification of work by Silke Baron; credit d: modification of work by Angell Williams)

Members of the class Scaphopoda (“boat feet”) are known colloquially as “tusk shells” or “tooth shells.” Tooth shells are open at both ends and usually lie buried in sand with the front opening exposed to water and the reduced head end projecting from the back of the shell. Tooth shells have a radula and a foot modified into tentacles, each with a bulbous end that catches and manipulates prey (Figure 15.28).

Figure 15.28. 

Antalis vulgaris shows the classic Dentaliidae shape that gives these animals their common name of “tusk shell.” (credit: Georges Jansoone)

Jawless fishes are craniates (which includes all the chordate groups except the tunicates and lancelets) that represent an ancient vertebrate lineage that arose over one half-billion years ago. Some of the earliest jawless fishes were the ostracoderms (which translates as “shell-skin”). Ostracoderms, now extinct, were vertebrate fishes encased in bony armor, unlike present-day jawless fishes, which lack bone in their scales.

The clade Myxini includes 67 species of hagfishes. Hagfishes are eel-like scavengers that live on the ocean floor and feed on dead invertebrates, other fishes, and marine mammals (Figure 15.37a). Hagfishes are entirely marine and are found in oceans around the world except for the polar regions. A unique feature of these animals is the slime glands beneath the skin that are able to release an extraordinary amount of mucus through surface pores. This mucus may allow the hagfish to escape from the grip of predators. Hagfish are known to enter the bodies of dead or dying organisms to devour them from the inside.

Figure 15.37. 

(a) Pacific hagfishes are scavengers that live on the ocean floor. (b) These parasitic sea lampreys attach to their lake trout host by suction and use their rough tongues to rasp away flesh in order to feed on the trout’s blood. (credit a: modification of work by Linda Snook, NOAA/CBNMS; credit b: modification of work by USGS)

The skeleton of a hagfish is composed of cartilage, which includes a cartilaginous notochord, which runs the length of the body, and a skull. This notochord provides support to the fish’s body. Although they are craniates, hagfishes are not vertebrates, since they do not replace the notochord with a vertebral column during development, as do the vertebrates.

The clade Petromyzontidae includes approximately 40 species of lampreys. Lampreys are similar to hagfishes in size and shape; however, lampreys have a brain case and incomplete vertebrae. Lampreys lack paired appendages and bone, as do the hagfishes. As adults, lampreys are characterized by a toothed, funnel-like sucking mouth. Some species are parasitic as adults, attaching to and feeding on the body fluids of fish (Figure 15.37b). Most species are free-living.

Lampreys live primarily in coastal and fresh waters and have a worldwide temperate region distribution. All species spawn in fresh waters. Eggs are fertilized externally, and the larvae are distinctly different from the adult form, spending 3 to 15 years as suspension feeders. Once they attain sexual maturity, the adults reproduce and die within days. Lampreys have a notochord as adults.

Gnathostomes or “jaw-mouths” are vertebrates that have jaws and include both cartilaginous and bony fishes. One of the most significant developments in early vertebrate evolution was the origin of the jaw, which is a hinged structure attached to the cranium that allows an animal to grasp and tear its food. The evolution of jaws allowed early gnathostomes to exploit food resources that were unavailable to jawless fishes.

The clade Chondrichthyes, the cartilaginous fishes, is diverse, consisting of sharks (Figure 15.38a), rays, and skates, together with sawfishes and a few dozen species of fishes called chimaeras, or ghost sharks. Chondrichthyes have paired fins and a skeleton made of cartilage. This clade arose approximately 370 million years ago in the middle Devonian. They are thought to have descended from an extinct group that had a skeleton made of bone; thus, the cartilaginous skeleton of Chondrichthyes is a later development. Parts of the shark skeleton are strengthened by granules of calcium carbonate, but this is not the same as bone.

Most cartilaginous fishes live in marine habitats, with a few species living in fresh water for some or all of their lives. Most sharks are carnivores that feed on live prey, either swallowing it whole or using their jaws and teeth to tear it into smaller pieces. Shark teeth likely evolved from the jagged scales that cover their skin. Some species of sharks and rays are suspension feeders that feed on plankton.

Figure 15.38. 

(a) This hammerhead shark is an example of a predatory cartilaginous fish. (b) This stingray blends into the sandy bottom of the ocean floor when it is feeding or awaiting prey. (credit a: modification of work by Masashi Sugawara; credit b: modification of work by “Sailn1″/Flickr)

Sharks have well-developed sense organs that aid them in locating prey, including a keen sense of smell and electroreception, the latter being perhaps the most sensitive of any animal. Organs called ampullae of Lorenzini allow sharks to detect the electromagnetic fields that are produced by all living things, including their prey. Electroreception has only been observed in aquatic or amphibious animals. Sharks, together with most fishes, also have a sense organ called the lateral line, which is used to detect movement and vibration in the surrounding water, and a sense that is often considered homologous to “hearing” in terrestrial vertebrates. The lateral line is visible as a darker stripe that runs along the length of the fish’s body.

Sharks reproduce sexually and eggs are fertilized internally. Most species are ovoviviparous, that is, the fertilized egg is retained in the oviduct of the mother’s body, and the embryo is nourished by the egg yolk. The eggs hatch in the uterus and young are born alive and fully functional. Some species of sharks are oviparous: They lay eggs that hatch outside of the mother’s body. Embryos are protected by a shark egg case or “mermaid’s purse” that has the consistency of leather. The shark egg case has tentacles that snag in seaweed and give the newborn shark cover. A few species of sharks are viviparous, that is, the young develop within the mother’s body, and she gives live birth.

Rays and skates include more than 500 species and are closely related to sharks. They can be distinguished from sharks by their flattened bodies, pectoral fins that are enlarged and fused to the head, and gill slits on their ventral surface (Figure 15.38b). Like sharks, rays and skates have a cartilaginous skeleton. Most species are marine and live on the sea floor, with nearly a worldwide distribution.

Members of the clade Osteichthyes, or bony fishes, are characterized by a bony skeleton. The vast majority of present-day fishes belong to this group, which consists of approximately 30,000 species, making it the largest class of vertebrates in existence today.

Nearly all bony fishes have an ossified skeleton with specialized bone cells (osteocytes) that produce and maintain a calcium phosphate matrix. This characteristic has only reverted in a few groups of Osteichthyes, such as sturgeons and paddlefish, which have primarily cartilaginous skeletons. The skin of bony fishes is often covered in overlapping scales, and glands in the skin secrete mucus that reduces drag when swimming and aids the fish in osmoregulation. Like sharks, bony fishes have a lateral line system that detects vibrations in water. Unlike sharks, some bony fish depend on their eyesight to locate prey. Bony fish are also unusual in possessing taste cells in the head and trunk region of the body that allow them to detect extremely small concentrations of molecules in the water.

All bony fishes, like the cartilaginous fishes, use gills to breathe. Water is drawn over gills that are located in chambers covered and ventilated by a protective, muscular flap called the operculum. Unlike sharks, bony fishes have a swim bladder, a gas-filled organ that helps to control the buoyancy of the fish. Bony fishes are further divided into two clades with living members: Actinopterygii (ray-finned fishes) and Sarcopterygii (lobe-finned fishes).

The ray-finned fishes include many familiar fishes—tuna, bass, trout, and salmon (Figure 15.39a), among others. Ray-finned fishes are named for the form of their fins—webs of skin supported by bony spines called rays. In contrast, the fins of lobe-finned fishes are fleshy and supported by bone (Figure 15.39b). Living members of lobe-finned fishes include the less familiar lungfishes and coelacanth.

Figure 15.39. 

The (a) sockeye salmon and (b) coelacanth are both bony fishes of the Osteichthyes clade. The coelacanth, sometimes called a lobe-finned fish, was thought to have gone extinct in the Late Cretaceous period 100 million years ago until one was discovered in 1938 between Africa and Madagascar. (credit a: modification of work by Timothy Knepp, USFWS; credit b: modification of work by Robbie Cada)

Amphibia comprise an estimated 6,500 extant species that inhabit tropical and temperate regions around the world. Amphibians can be divided into three clades: Urodela (“tailed-ones”), the salamanders and newts; Anura (“tail-less ones”), the frogs and toads; and Apoda (“legless ones”), the caecilians.

Living salamanders (Figure 15.40a) include approximately 500 species, some of which are aquatic, others terrestrial, and some that live on land only as adults. Adult salamanders usually have a generalized tetrapod body plan with four limbs and a tail. Some salamanders are lungless, and respiration occurs through the skin or external gills. Some terrestrial salamanders have primitive lungs; a few species have both gills and lungs.

Figure 15.40. 

(a) Most salamanders have legs and a tail, but respiration varies among species. (b) The Australian green tree frog is a nocturnal predator that lives in the canopies of trees near a water source. (credit a: modification of work by Valentina Storti; credit b: modification of work by Evan Pickett)

Concept in Action

Watch this video about an unusually large salamander species.

Frogs (Figure 15.40b) are the most diverse group of amphibians, with approximately 5,000 species that live on all continents except Antarctica. Frogs have a body plan that is more specialized than the salamander body plan for movement on land. Adult frogs use their hind limbs to jump many times their body length on land. Frogs have a number of modifications that allow them to avoid predators, including skin that acts as camouflage and defensive chemicals that are poisonous to predators secreted from glands in the skin.

Frog eggs are fertilized externally, as they are laid in moist environments. Frogs demonstrate a range of parental behaviors, with some species exhibiting little care, to species that carry eggs and tadpoles on their hind legs or backs. The life cycle consists of two stages: the larval stage followed by metamorphosis to an adult stage. The larval stage of a frog, the tadpole, is often a filter-feeding herbivore. Tadpoles usually have gills, a lateral line system, long-finned tails, but no limbs. At the end of the tadpole stage, frogs undergo a gradual metamorphosis into the adult form. During this stage, the gills and lateral line system disappear, and four limbs develop. The jaws become larger and are suited for carnivorous feeding, and the digestive system transforms into the typical short gut of a predator. An eardrum and air-breathing lungs also develop. These changes during metamorphosis allow the larvae to move onto land in the adult stage (Figure 15.41).

Figure 15.41. 

A frog begins as a (a) tadpole and undergoes metamorphosis to become (b) a juvenile and finally (c) an adult. (credit: modification of work by Brian Gratwicke)

Caecilians comprise an estimated 185 species. They lack external limbs and resemble giant earthworms. They inhabit soil and are found primarily in the tropics of South America, Africa, and southern Asia where they are adapted for a soil-burrowing lifestyle and are nearly blind. Unlike most of the other amphibians that breed in or near water, reproduction in a drier soil habitat means that caecilians must utilize internal fertilization, and most species give birth to live young (Figure 15.42).

Figure 15.42. 

Caecilians lack external limbs and are well adapted for a soil-burrowing lifestyle. (credit: modification of work by “cliff1066″/Flickr)

Reptiles are tetrapods. Limbless reptiles—snakes—may have vestigial limbs and, like caecilians, are classified as tetrapods because they are descended from four-limbed ancestors. Reptiles lay shelled eggs on land. Even aquatic reptiles, like sea turtles, return to the land to lay eggs. They usually reproduce sexually with internal fertilization. Some species display ovoviviparity, with the eggs remaining in the mother’s body until they are ready to hatch. Other species are viviparous, with the offspring born alive.

One of the key adaptations that permitted reptiles to live on land was the development of their scaly skin, containing the protein keratin and waxy lipids, which prevented water loss from the skin. This occlusive skin means that reptiles cannot use their skin for respiration, like amphibians, and thus all must breathe with lungs. In addition, reptiles conserve valuable body water by excreting nitrogen in the form of uric acid paste. These characteristics, along with the shelled, amniotic egg, were the major reasons why reptiles became so successful in colonizing a variety of terrestrial habitats far from water.

Reptiles are ectotherms, that is, animals whose main source of body heat comes from the environment. Behavioral maneuvers, like basking to heat themselves, or seeking shade or burrows to cool off, help them regulate their body temperature,

Class Reptilia includes diverse species classified into four living clades. These are the Crocodilia, Sphenodontia, Squamata, and Testudines.

The Crocodilia (“small lizard”) arose approximately 84 million years ago, and living species include alligators, crocodiles, and caimans. Crocodilians (Figure 15.43a) live throughout the tropics of Africa, South America, the southeastern United States, Asia, and Australia. They are found in freshwater habitats, such as rivers and lakes, and spend most of their time in water. Some species are able to move on land due to their semi-erect posture.

Figure 15.43. 

(a) Crocodilians, such as this Siamese crocodile, provide parental care for their offspring. (b) This Jackson’s chameleon blends in with its surroundings. (c) The garter snake belongs to the genus Thamnophis, the most widely distributed reptile genus in North America. (d) The African spurred tortoise lives at the southern edge of the Sahara Desert. It is the third largest tortoise in the world. (credit a: modification of work by Keshav Mukund Kandhadai; credit c: modification of work by Steve Jurvetson; credit d: modification of work by Jim Bowen)

The Sphenodontia (“wedge tooth”) arose in the Mesozoic Era and includes only one living genus, Tuatara, with two species that are found in New Zealand. There are many fossil species extending back to the Triassic period (250–200 million years ago). Although the tuataras resemble lizards, they are anatomically distinct and share characteristics that are found in birds and turtles.

Squamata (“scaly”) arose in the late Permian; living species include lizards and snakes, which are the largest extant clade of reptiles (Figure 15.43b). Lizards differ from snakes by having four limbs, eyelids, and external ears, which are lacking in snakes. Lizard species range in size from chameleons and geckos that are a few centimeters in length to the Komodo dragon, which is about 3 meters in length.

Snakes are thought to have descended from either burrowing lizards or aquatic lizards over 100 million years ago (Figure 15.43c). Snakes comprise about 3,000 species and are found on every continent except Antarctica. They range in size from 10 centimeter-long thread snakes to 7.5 meter-long pythons and anacondas. All snakes are carnivorous and eat small animals, birds, eggs, fish, and insects.

Turtles are members of the clade Testudines (“having a shell”) (Figure 15.43d). Turtles are characterized by a bony or cartilaginous shell, made up of the carapace on the back and the plastron on the ventral surface, which develops from the ribs. Turtles arose approximately 200 million years ago, predating crocodiles, lizards, and snakes. Turtles lay eggs on land, although many species live in or near water. Turtles range in size from the speckled padloper tortoise at 8 centimeters (3.1 inches) to the leatherback sea turtle at 200 centimeters (over 6 feet). The term “turtle” is sometimes used to describe only those species of Testudines that live in the sea, with the terms “tortoise” and “terrapin” used to refer to species that live on land and in fresh water, respectively.

Data now suggest that birds belong within the reptile clade, but they display a number of unique adaptations that set them apart. Unlike the reptiles, birds are endothermic, meaning they generate their own body heat through metabolic processes. The most distinctive characteristic of birds is their feathers, which are modified reptilian scales. Birds have several different types of feathers that are specialized for specific functions, like contour feathers that streamline the bird’s exterior and loosely structured down feathers that insulate (Figure 15.44a).

Feathers not only permitted the earliest birds to glide, and ultimately engage in flapping flight, but they insulated the bird’s body, assisting the maintenance of endothermy, even in cooler temperatures. Powering a flying animal requires economizing on the amount of weight carried. As body weight increases, the muscle output and energetic cost required for flying increase. Birds have made several modifications to reduce body weight, including hollow or pneumatic bones (Figure 15.44b) with air spaces that may be connected to air sacs and cross-linked struts within their bones to provide structural reinforcement. Parts of the vertebral skeleton and braincase are fused to increase its strength while lightening its weight. Most species of bird only possess one ovary rather than two, and no living birds have teeth in their jaw, further reducing body mass.

Figure 15.44. 

(a) Primary feathers are located at the wing tip and provide thrust; secondary feathers are located close to the body and provide lift. (b) Many birds have hollow pneumatic bones, which make flight easier.

Birds possess a system of air sacs branching from their primary airway that divert the path of air so that it passes unidirectionally through the lung, during both inspiration and expiration. Unlike mammalian lungs in which air flows in two directions as it is breathed in and out, air flows continuously through the bird’s lung to provide a more efficient system of gas exchange.

Order Primates of class Mammalia includes lemurs, tarsiers, monkeys, and the apes, which include humans. Non-human primates live primarily in tropical or subtropical regions of South America, Africa, and Asia. They range in size from the mouse lemur at 30 grams (1 ounce) to the mountain gorilla at 200 kilograms (441 pounds). The characteristics and evolution of primates are of particular interest to us as they allow us to understand the evolution of our own species.

All primate species have adaptations for climbing trees, as they all descended from tree-dwellers, although not all species are arboreal. This arboreal heritage of primates resulted in hands and feet that are adapted for brachiation, or climbing and swinging through trees. These adaptations include, but are not limited to 1) a rotating shoulder joint, 2) a big toe that is widely separated from the other toes and thumbs that are widely separated from fingers (except humans), which allow for gripping branches, and 3) stereoscopic vision, two overlapping visual fields, which allows for the depth perception necessary to gauge distance. Other characteristics of primates are brains that are larger than those of many other mammals, claws that have been modified into flattened nails, typically only one offspring per pregnancy, and a trend toward holding the body upright.

Order Primates is divided into two groups: prosimians and anthropoids. Prosimians include the bush babies of Africa, the lemurs of Madagascar, and the lorises, pottos, and tarsiers of Southeast Asia. Anthropoids include monkeys, lesser apes, and great apes (Figure 15.47). In general, prosimians tend to be nocturnal, smaller in size than anthropoids, and have relatively smaller brains compared to anthropoids.

Figure 15.47. 

Primates can be divided into prosimians, such as the (a) lemur, and anthropoids. Anthropoids include monkeys, such as the (b) howler monkey; lesser apes, such as the (c) gibbon; and great apes, such as the (d) chimpanzee, (e) bonobo, (f) gorilla, and (g) orangutan. (credit a: modification of work by Frank Vassen; credit b: modification of work by Xavi Talleda; credit d: modification of work by Aaron Logan; credit e: modification of work by Trisha Shears; credit f: modification of work by Dave Proffer; credit g: modification of work by Julie Langford)