Chapter 15: Diversity of Animals PDF

CHAPTER 15 Diversity of Animals FIGURE 15.1 The leaf chameleon (Brookesia micra) was discovered in northern Madagascar in 2012. At just over one inch long, it is the smallest known chameleon. (credit: modification of work by Frank Glaw, et al., PLOS) CHAPTER OUTLINE 15.1 Features of the Animal Kin...

CHAPTER 15 Diversity of Animals FIGURE 15.1 The leaf chameleon (Brookesia micra) was discovered in northern Madagascar in 2012. At just over one inch long, it is the smallest known chameleon. (credit: modification of work by Frank Glaw, et al., PLOS) CHAPTER OUTLINE 15.1 Features of the Animal Kingdom 15.2 Sponges and Cnidarians 15.3 Flatworms, Nematodes, and Arthropods 15.4 Mollusks and Annelids 15.5 Echinoderms and Chordates 15.6 Vertebrates INTRODUCTION While we can easily identify dogs, lizards, fish, spiders, and worms as animals, other animals, such as corals and sponges, might be easily mistaken as plants or some other form of life. Yet scientists have recognized a set of common characteristics shared by all animals, including sponges, jellyfish, sea urchins, and humans. The kingdom Animalia is a group of multicellular Eukarya. 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 currently living species of animals have been identified, scientists are continually discovering more species. The number of described living animal species is estimated to be about 1.4 1 million, and there may be as many as 6.8 million. Understanding and classifying the variety of living species helps us to better understand how to conserve and benefit from this diversity. The animal classification system characterizes animals 1 “Number of Living Species in Australia and the World,” A.D. Chapman, Australia Biodiversity Information Services, last modified August 26, 2010, http://www.environment.gov.au/biodiversity/abrs/publications/other/species- numbers/2009/03-exec-summary.html. 352 15 Diversity of Animals based on their anatomy, features of embryological development, and genetic makeup. Scientists are faced with the task of classifying animals within a system of taxonomy that reflects their evolutionary history. Additionally, 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. However, animals vary in the complexity of their organization and exhibit a huge diversity of body forms, so the classification scheme is constantly changing as new information about species is learned. 15.1 Features of the Animal Kingdom LEARNING OBJECTIVES By the end of this section, you will be able to: List the features that distinguish the animal kingdom from other kingdoms Explain the processes of animal reproduction and embryonic development Describe the hierarchy of basic animal classification Compare and contrast the embryonic development of protostomes and deuterostomes 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. FIGURE 15.2 All animals that derive energy from food are heterotrophs. The (a) black bear is an omnivore, eating both plants and animals. The (b) heartworm Dirofilaria immitis is a parasite that derives energy from its hosts. It spends its larval stage in mosquitos and its adult stage infesting the hearts of dogs and other mammals, as shown here. (credit a: modification of work by USDA Forest Service; credit b: modification of work by Clyde Robinson) 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 Access for free at openstax.org  15.1 Features of the Animal Kingdom 353 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. LINK TO LEARNING View this video (http://openstax.org/l/saving_life2) 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 Animals are classified according to morphological and developmental characteristics, such as a body plan. With the exception of sponges, the animal body plan is symmetrical. This means that their distribution of body parts is balanced along an axis. Additional characteristics that contribute to animal classification include the number of tissue layers formed during development, the presence or absence of an internal body cavity, and other features of embryological development. 354 15 Diversity of Animals VISUAL CONNECTION FIGURE 15.3 The phylogenetic tree of animals is based on morphological, fossil, and genetic evidence. Which of the following statements is false? a. Eumetazoa have specialized tissues and Parazoa do not. b. Both acoelomates and pseudocoelomates have a body cavity. c. Chordates are more closely related to echinoderms than to rotifers according to the figure. d. Some animals have radial symmetry, and some animals have bilateral symmetry. 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. Access for free at openstax.org  15.1 Features of the Animal Kingdom 355 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). LINK TO LEARNING Watch this video (http://openstax.org/l/symmetry2) 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 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. 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 356 15 Diversity of Animals 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) Protostomes and Deuterostomes 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. Access for free at openstax.org  15.2 Sponges and Cnidarians 357 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. 15.2 Sponges and Cnidarians LEARNING OBJECTIVES By the end of this section, you will be able to: Describe the organizational features of the simplest animals Describe the organizational features of cnidarians The kingdom of animals is informally divided into invertebrate animals, those without a backbone, and vertebrate animals, those with a backbone. Although in general we are most familiar with vertebrate animals, the vast majority of animal species, about 95 percent, are invertebrates. Invertebrates include a huge diversity of animals, millions of species in about 32 phyla, which we can just begin to touch on here. The sponges and the cnidarians represent the simplest of animals. Sponges appear to represent an early stage of multicellularity in the animal clade. Although they have specialized cells for particular functions, they lack true tissues in which specialized cells are organized into functional groups. Sponges are similar to what might have been the ancestor of animals: colonial, flagellated protists. The cnidarians, or the jellyfish and their kin, are the simplest animal group that displays true tissues, although they possess only two tissue layers. Sponges Animals in subkingdom Parazoa represent the simplest animals and include the sponges, or phylum Porifera (Figure 15.8). All sponges are aquatic and the majority of species are marine. Sponges live in intimate contact with water, which plays a role in their feeding, gas exchange, and excretion. Much of the body structure of the sponge is dedicated to moving water through the body so it can filter out food, absorb dissolved oxygen, and eliminate wastes. 358 15 Diversity of Animals FIGURE 15.8 Sponges are members of the phylum Porifera, which contains the simplest animals. (credit: Andrew Turner) The body of the simplest sponges takes the shape of a cylinder with a large central cavity, the spongocoel. Water enters the spongocoel from numerous pores in the body wall. Water flows out through a large opening called the osculum (Figure 15.9). However, sponges exhibit a diversity of body forms, which vary in the size and branching of the spongocoel, the number of osculi, and where the cells that filter food from the water are located. Sponges consist of an outer layer of flattened cells and an inner layer of cells called choanocytes separated by a jelly-like substance called mesohyl. The mesohyl contains embedded amoeboid cells that secrete tiny needles called spicules or protein fibers that help give the sponge its structural strength. The cell body of the choanocyte is embedded in mesohyl but protruding into the spongocoel is a mesh-like collar surrounding a single flagellum. The beating of flagella from all choanocytes moves water through the sponge. Food particles are trapped in mucus produced by the sieve-like collar of the choanocytes and are ingested by phagocytosis. This process is called intracellular digestion. Amoebocytes take up nutrients repackaged in food vacuoles of the choanocytes and deliver them to other cells within the sponge. Access for free at openstax.org  15.2 Sponges and Cnidarians 359 FIGURE 15.9 The sponge’s basic body plan is shown. Physiological Processes in Sponges 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. LINK TO LEARNING Watch this video (http://openstax.org/l/sponge_feed) that demonstrates the feeding of sponges. Cnidarians The phylum Cnidaria includes animals that show radial or biradial symmetry and are diploblastic. Nearly all (about 99 percent) cnidarians are marine species. Cnidarians have specialized cells known as cnidocytes (“stinging cells”) containing organelles called nematocysts. These cells are concentrated around the mouth and tentacles of the animal and can immobilize prey with toxins. Nematocysts contain coiled threads that may bear barbs. The outer wall of the cell has a hairlike projection that is sensitive to touch. When touched, the cells fire the toxin-containing coiled threads that can penetrate and stun the predator or prey (see Figure 15.10). 360 15 Diversity of Animals FIGURE 15.10 Animals from the phylum Cnidaria have stinging cells called cnidocytes. Cnidocytes contain large organelles called (a) nematocysts that store a coiled thread and barb. When hairlike projections on the cell surface are touched, (b) the thread, barb, and a toxin are fired from the organelle. Cnidarians display two distinct body plans: polyp or “stalk” and medusa or “bell” (Figure 15.11). Examples of the polyp form are freshwater species of the genus Hydra; perhaps the best-known medusoid animals are the jellies (jellyfish). Polyps are sessile as adults, with a single opening to the digestive system (the mouth) facing up with tentacles surrounding it. Medusae are motile, with the mouth and tentacles hanging from the bell-shaped body. In other cnidarians, both a polyp and medusa form exist, and the life cycle alternates between these forms. FIGURE 15.11 Cnidarians have two distinct body plans, the (a) medusa and the (b) polyp. All cnidarians have two tissue layers, with a jelly- like mesoglea between them. Physiological Processes of Cnidarians 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 Access for free at openstax.org  15.2 Sponges and Cnidarians 361 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. Cnidarian Diversity 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). 362 15 Diversity of Animals FIGURE 15.13 Scyphozoans include the jellies. (credit: "Jimg944"/Flickr) LINK TO LEARNING Use this video (https://openstax.org/l/amazing_jelly2) to identify the life cycle stages of jellies. 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. LINK TO LEARNING Watch this video (http://openstax.org/l/box_jellyfish) to learn more about the deadly toxins of the box jellyfish. 2 Hydrozoa includes nearly 3,500 species, 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). 2 “The Hydrozoa Directory,” Peter Schuchert, Muséum Genève, last updated November 2012, http://www.ville-ge.ch/mhng/hydrozoa/ hydrozoa-directory.htm. Access for free at openstax.org  15.3 Flatworms, Nematodes, and Arthropods 363 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) 15.3 Flatworms, Nematodes, and Arthropods LEARNING OBJECTIVES By the end of this section, you will be able to: Describe the structure and systems of flatworms Describe the structural organization of nematodes Compare the internal systems and the appendage specialization of arthropods The animal phyla of this and subsequent modules are triploblastic and have an embryonic mesoderm sandwiched between the ectoderm and endoderm. These phyla are also bilaterally symmetrical, meaning that a longitudinal section will divide them into right and left sides that are mirror images of each other. Associated with bilateralism is the beginning of cephalization, the evolution of a concentration of nervous tissues and sensory organs in the head of the organism, which is where the organism first encounters its environment. The flatworms are acoelomate organisms that include free-living and parasitic forms. The nematodes, or roundworms, possess a pseudocoelom and consist of both free-living and parasitic forms. Finally, the arthropods, one of the most successful taxonomic groups on the planet, are coelomate organisms with a hard exoskeleton and jointed appendages. The nematodes and the arthropods belong to a clade with a common ancestor, called Ecdysozoa. The name comes from the word ecdysis, which refers to the periodic shedding, or molting, of the exoskeleton. The ecdysozoan phyla have a hard cuticle covering their bodies that must be periodically shed and replaced for them to increase in size. Flatworms The relationships among flatworms, or phylum Platyhelminthes, is being revised and the description here will follow the traditional groupings. Most flatworms are parasitic, including important parasites of humans. Flatworms have three embryonic germ layers that give rise to surfaces covering tissues, internal tissues, and the lining of the digestive system. The epidermal tissue is a single layer of cells or a layer of fused cells covering a layer of circular muscle above a layer of longitudinal muscle. The mesodermal tissues include support cells and secretory cells that secrete mucus and other materials to the surface. The flatworms are acoelomate, so their bodies contain no cavities or spaces between the outer surface and the inner digestive tract. Physiological Processes of Flatworms 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. 364 15 Diversity of Animals 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. Diversity of Flatworms 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. Access for free at openstax.org  15.3 Flatworms, Nematodes, and Arthropods 365 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. 366 15 Diversity of Animals Nematodes The phylum Nematoda, or roundworms, includes more than 28,000 species with an estimated 16,000 parasitic species. The name Nematoda is derived from the Greek word “nemos,” which means “thread.” Nematodes are present in all habitats and are extremely common, although they are usually not visible (Figure 15.17). FIGURE 15.17 (a) An scanning electron micrograph of the nematode Heterodera glycines and (b) a schematic representation of the anatomy of a nematode are shown. (credit a: modification of work by USDA, ARS; scale-bar data from Matt Russell) Most nematodes look similar to each other: slender tubes, tapered at each end (Figure 15.17). Nematodes are pseudocoelomates and have a complete digestive system with a distinct mouth and anus. The nematode body is encased in a cuticle, a flexible but tough exoskeleton, or external skeleton, which offers protection and support. The cuticle contains a carbohydrate-protein polymer called chitin. The cuticle also lines the pharynx and rectum. Although the exoskeleton provides protection, it restricts growth, and therefore must be continually shed and replaced as the animal increases in size. A nematode’s mouth opens at the anterior end with three or six lips and, in some species, teeth in the form of cuticular extensions. There may also be a sharp stylet that can protrude from the mouth to stab prey or pierce plant or animal cells. The mouth leads to a muscular pharynx and intestine, leading to the rectum and anal opening at the posterior end. Physiological Processes of Nematodes 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 Access for free at openstax.org  15.3 Flatworms, Nematodes, and Arthropods 367 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. LINK TO LEARNING View this video (http://openstax.org/l/nematode) 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. Arthropoda The name “arthropoda” means “jointed legs,” which aptly describes each of the enormous number of species belonging to this phylum. Arthropoda dominate the animal kingdom with an estimated 85 percent of known species, with many still undiscovered or undescribed. The principal characteristics of all the animals in this phylum are functional segmentation of the body and the presence of jointed appendages (Figure 15.18). As members of Ecdysozoa, arthropods also have an exoskeleton made principally of chitin. Arthropoda is the largest phylum in the animal world in terms of numbers of species, and insects form the single largest group within this phylum. Arthropods are true coelomate animals and exhibit prostostomic development. FIGURE 15.18 Trilobites, like the one in this fossil, are an extinct group of arthropods. (credit: Kevin Walsh) Physiological Processes of Arthropods 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). 368 15 Diversity of Animals 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) Arthropod Diversity 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). Access for free at openstax.org  15.3 Flatworms, Nematodes, and Arthropods 369 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 3 47,000. 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 4 subphylum is predominantly terrestrial, although some marine species also exist. An estimated 103,000 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. 3 “Number of Living Species in Australia and the World,” A.D. Chapman, Australia Biodiversity Information Services, last modified August 26, 2010, http://www.environment.gov.au/biodiversity/abrs/publications/other/species-numbers/2009/03-exec-summary.html. 4 “Number of Living Species in Australia and the World,” A.D. Chapman, Australia Biodiversity Information Services, last modified August 26, 2010, http://www.environment.gov.au/biodiversity/abrs/publications/other/species-numbers/2009/03-exec-summary.html.