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GUIDE FOR READING After you read the following sections, you will be able to 28-1 Introduction to Arthropods • Discuss the characteristics and classification of arthropods. • Explain how arthropods perform CHAPTER essential life functions. 28-2 Spiders and Their Relatives • Discuss the distinguishing characteristics of chelicerates. Arthropods 28-3 Crustaceans • Discuss the distinguishing characteristics of crustaceans. 28-4 Insects and Their Relatives • Describe and give examples of three classes in the subphylum Unifamia. • Discuss the anatomy of a typical insect. • Explain ways insects communicate. 28-5 How Arthropods Fit into the World Arthropods display a wide range of forms and habit* • Describe how arthropods interact with other living things. ^^*z:str'alherbiv°re-The v™" R theoceTflhe ^ ^ ^m ^ « ^long ho ocean floor on legs nearly a meter long, the lukewarm waters Journal Activity YOU AND YOUR WORLD beneath a^ ^ in^the COntr°'0,3 b^ow beneath a cypress tree.^And heart of a Florida swamp, 43 ^~ZZZ:.~Z;Z Bugs! Creepy crawly insects. The mere thought of them makes some people cringe. But not all insects are harmful. Pretend that you are a press agent for insects. Write a short press release informing the public about the benefits of sharing our world with insects. 28-1 Introduction to Arthropods Guide For Reading ¦ What are the four subphyla of arthropods? ¦ What are three important arthropod features? ¦ How are arthropods adapted for performing life functions? ¦ What is metamorphosis? To describe even a fraction of the living arthropods would take several books. More than a million arthropod species have been described, and scientists are certain there are many more that have not as yet been found! As you will soon learn, members of this phylum vary enormously in size, shape, and habits. Diversity and Evolution in Arthropods You already know about many common arthropods. In fact, you have probably even eaten a number of them! Today most biologists divide arthropods into four subphyla: • Trilohita This is thought to be the oldest subphylum of arthropods. Trilobites (TRlGH-loh-bights) were dwellers in ancient seas. They are now all extinct. • Chelicerata Chelicerates (keh-LlHS-er-ayts) include spiders, ticks, mites, scorpions, and horseshoe crabs. • Crustacea Crustaceans (kruhs-TAY-shuhnz) include such familiar (and edible) organisms as crabs and shrimp. 8 Uniramia IMramians (yoo-nih-RAY-mee-ahnz) include most arthropods: centipedes, millipedes, and all insects— including bees, moths, grasshoppers, flies, and beetles. Why are there so many different kinds of arthropods? One reason is that they have been evolving on Earth for a long time. The first arthropods appeared in the sea more than 600 million years ago. Since that time, these animals have experienced sev¬ eral adaptive radiations. Some arthropods have remained in polar regions to bene pools on the equator. Some arthropods are ong e most destructive animals on Earth. Others are beneficial even essentml. to the survival of other organisms. What are arthropods? What common characteristics do they the answers tQ the'^ ^ ^ ^ Y°u ^ ^over answers tg,these questions In the pages that follow. Figure 28-1 The scorpion, a chelicerate, is feeding on a hawk moth, a uniramian (left). The whitebooted shrimp is a crustacean (right). 607 Figure 28-2 The velvet worm is traditionally placed in its own phylum along with other organisms that have characteristics of both annelids and arthropods. Such animals support the hypothesis that modern annelids and uniramian arthropods descended from a common ancestor. However, some molecular biologists now feel that these animals should be placed in the phylum Arthropoda. the water, where they have colonized all parts of the sea and most freshwater habitats. Others were among the very first members of the animal kingdom to colonize the land. The de¬ scendants of those pioneers were on hand when the first flow¬ ering plants appeared millions of years later. The roots of the arthropod family tree are cloaked in mys¬ tery because the ancestors of the arthropods were soft-bodied animals that left few fossils. But by studying both living and fossil invertebrates, researchers have accumulated many clues to arthropod evolutionary history. Insects, centipedes, and mil¬ lipedes seem to have evolved from ancestors that were closely related to the ancestors of modern annelid worms. Living evi¬ dence for this line of descent can be found in wormlike animals that live in the tropics. See Figure 28-2. Other arthropods, in¬ cluding crustaceans, spiders, and the extinct trilobites, evolved from more ancient and more distantly related ancestors. The body form of the earliest arthropods is thought to be similar to that of the trilobites. A typical trilobite's body had a thick, tough outer covering and was composed of many seg¬ ments, each of which bore a pair of appendages. Each appen¬ dage was divided to form two branches, one a walking leg and one a featherlike gill. See Figure 28-3. Body segments Figure 28-3 The dorsal side of a fossil trilobite clearly shows the three lengthwise body lobes that give the animal its name (inset). An artist's rendering of the ventral side of a trilobite as it might have appeared when the animal was alive reveals numerous similarly shaped appendages. Most living arthropods exhibit two evolutionary trends away from the trilobite form. First, many have far fewer body segments. The many segments found in their embryos fuse into larger segments during development. Second, arthropod ap¬ pendages have become increasingly specialized for feeding, locomotion, and other functions. Figure 28-4 Some arthropods, such as the spiny lobster (center) and the tick (top), have extremely hard, tough exoskeletons. The mouthparts of a tick (inset) are adapted for biting and hanging onto a host. Other arthropods, such as the emperor gum moth caterpillar (bottom), have flexible, leathery exoskeletons. Form and Function in Arthropods Although living arthropods are quite different from one an¬ other, all arthropods exhibit several key features. The three most important arthropod features are a tough exoskeleton, a series of jointed appendages, and a segmented body. Other characteristics of arthropods include a brain located in the dorsal part of the head, a ventral nerve cord, and an open cir¬ culatory system powered by a single heart. THE ARTHROPOD BODY PLAN The exoskeleton (exomeans outside) is a system of external supporting structures that are made primarily of the carbohydrate chitin (KlGH-tihn). Some exoskeletons, such as those of most insects, are leathery and flexible. Others, such as those of ticks and lobsters, are ex¬ tremely hard. These exoskeletons provide excellent protection from physical damage. The exosketetons of many terrestrial arthropods are waterproof. This adaptation restricts the loss of water from the body and makes it possible for arthropods to live in extremely dry environments such as deserts. The exo¬ skeleton also helps arthropods move efficiently and adapt to their environment in many other ways. Although the exoskeleton protects an arthropod's body like a suit of armor, it has the disadvantages you might expect from such a covering. Because an exoskeleton is a solid coat¬ ing, not a living tissue, it cannot grow as the animal grows. (You will learn how arthropods deal with this problem shortly.) And movement can occur only at the joints of the "armor." 609 iM, All arthropods have jointed appendages (arthro- means joint; -pod, which literally means foot, refers to the appen¬ dages) that enable them to move. In primitive arthropods, such as trilobites, every body segment carries a single pair of ap¬ pendages. But in species in which body segments are fused to¬ gether, some appendages have been lost. Over millions of years, the remaining appendages have evolved into marvelously versatile adaptations to different environments. Arthro¬ pod appendages include antennae, claws, walking legs, wings, flippers, and other specialized structures. All arthropods have segmented bodies. Some, such as mil¬ lipedes and centipedes, have long, wormlike bodies with many visible segments. Others, such as insects, spiders, and crabs, have lost some segments in the course of evolution or had the segments fuse together to form a few large body parts. FEEDING The appendages of arthropods have evolved in ways that enable these animals to eat almost any food you can imagine. Every mode of feeding is seen in arthropods—herbi¬ vores, carnivores, parasites, filter feeders, and detritus feeders. Figure 28-5 The waterflea is a tiny freshwater crustacean that lacks external segmentation and uses its antennae to propel it through the water. The round structures on the waterflea's back are its eggs. Although some herbivores, such as locusts, eat just about any¬ thing green, other herbivores are more selective. Some herbi¬ vores are specialized to eat specific parts of plants. Others feed exclusively on a particular kind of plant. Some carnivores, such as spiders, praying mantises, centipedes, and king crabs, catch and eat other animals. Other carnivores, such as many crabs and crayfish, feed primarily on animals that are already dead. External parasites—such as ticks, fleas, and lice—drink the blood and body fluids or nibble on the skin of other animals, including humans. Some internal parasites passively absorb nutrients through the body wall, whereas others eat away at the host from inside. Many marine arthropods are filter feeders that use comblike bristles on their mouthparts or legs to filter tiny plants and animals from the water. RESPIRATION Arthropods have evolved three basic types of respiratory structures—gills, book gills and book lungs, and tracheal tubes. Although most arthropods have only one of these types of respiratory structures, a few species have both book lungs and tracheal tubes. And some species com¬ pletely lack specialized respiratory organs. Many aquatic arthropods, such as crabs and shrimp, have gills that look like a row of feathers located just under cover of their exoskeleton. These gills are formed from part of the same appendages that form mouthparts and legs. Movement of the mouthparts and other appendages keeps a steady stream of water moving over the gills. Book gills (which are found in horseshoe crabs) and book lungs (which are found in spiders and their relatives) are unique to these arthropods. In both these structures, several sheets of tissue are layered like pages in a book. The many tis¬ sue layers increase the surface area for gas exchange. A horse¬ shoe crab's book gills are carried beneath its body, whereas a spider's book lungs are contained inside a sac within the body. An opening called the spiracle (SPlHR-ah-kuhl) connects the sac containing the book lungs with the fresh air outside. Most terrestrial arthropods—insects, some spiders, and millipedes, for example—have another respiratory device found in no other animals. From spiracles, long branching tra¬ cheal tubes reach deep into the animals' tissues. The network of tracheal tubes supplies oxygen by diffusion to all body tis¬ sues. As the arthropods walk, fly, or crawl, the movements of their body muscles cause the tracheae to shrink and expand, pumping fresh air in and out of the spiracles. Tracheal tubes work well only in small animals; large animals require a more efficient way to deliver oxygen and remove carbon dioxide. Figure 28-7 Although a fiddler crab spends much time on land, it uses gills for respiration. The male crab's large claw is used to attract females and to fight with other moles. Figure 28-8 The internal structures of a representative arthropod—a grasshopper—are shown here. . fli Figure 28-6 The praying mantis (left) is a carnivore. The lubber grasshopper (right) is a herbivore. Head Thorax P Abdomen i Digestive I tract Antenna Heart j Compound eye Nerve cord Walking legs 610 Tracheal tube Spiracle Malpighian tubules 611 Figure 28-9 The diamond beetle . J uses Malpighian tubules to get nd of nitrogen-containing wastes, whereas the hermit crab (right) uses its green glands and gills. h^H if m abdomen. In lobsters and crayfish the the hearTrf nt antd 1165 abOUt halfway down the body. When the heart contracts, it pumps blood through arteries iZ branch lnt0 Iler vessels and enter the ^ leaves the vessels and moves through spaces in the tis ue called smuses. Eventually, the blood collects ,^ a arge cav" Ity surrounding the heart, from which it re-enters he h i through small openings and is pumped around agafn spilsliiii RESPONSE Most arthropods have well-developed ner¬ vous systems. All have a brain that consists of a pair of ganglia in the head. These ganglia serve as central switchboards for in¬ coming information and outgoing instructions to muscles. From the brain, a pair of nerves runs around the esophagus and connects the brain to a nerve cord that runs along the ven¬ tral part of the body. Along this nerve cord are several more ganglia, usually one for each original body segment. These ganglia serve as local command centers to coordinate the movement of legs and wings. (That's why many insects can still walk or flap their wings after their heads are cut off!) Where many body segments have fused together, as in insects, there are several ganglia for each major body part. Arthropods have simple sense organs such as statocysts and chemical receptors. Most arthropods also have sophisti¬ cated sense organs such as compound eyes for gathering infor¬ mation from their environment. Compound eyes may have more than 2000 separate lenses and can detect color and mo¬ tion very well. In fact, insects can see certain things better than we can. (That's one reason it is so hard to swat flies and mos¬ quitoes!) For example, the blades of a quickly moving fan—just a blur to our eyes—are clearly visible to a fly. And many in¬ sects can see ultraviolet light, which is invisible to humans. Both crustaceans and insects have a well-developed sense of taste, although their taste receptors are located in strange places. The chemical receptors associated with the senses of taste and smell are located on the mouthparts, as might be ex¬ pected, but also on the antennae and legs! Flies, for example, know immediately whether a drop of water they step in con¬ tains salt or sugar. Crustaceans and insects have sensory hairs that detect movement in the air or water (another reason in¬ sects are hard to swat or catch). As an object moves toward them, they can feel the movement of the displaced air or water and respond appropriately. Many insects have well-developed ears that hear sounds above the human range. Insect ears are often in odd places. The eardrums in grasshoppers, for exam¬ ple, are behind their legs. Figure 28-10 The horsefly has huge compound eyes through which it sees poppy flowers much as they appear here (inset). Figure 28-11 A harlequin beetle (left) uses its long antennae as "feelers." The red hourglass on the abdomen of the black widow spider (right) warns of the spider's venomous bite. the body sinuses TheTh |0rgans' are bathed in blood inside concentrate them remove wastes from this blood, fore h leaves via'the ^ ^ t0 undigested food be- such as lobster. ^ r g S- Many ^ic arthropods. or to" ¦"p,"d 10"""" "»»"»»¦ per Growth and Development in Arthropods Figure 28-12 Can you find the grasshopper in the photograph? The stick grasshopper's shape is a form of camouflage (left). The markings and behavior of this caterpillar trick insect-eating birds into thinking it is a bird-eating viper (center). The harmless hover fly mimics a stinging honeybee (right). Figure 28-13 Muscles attached to the inside of an exoskeleton bend and straighten the joints. Exoskeleton An arthropod's well-developed sense organs help it detect and rrpeu predftors' The combination of these sense organs and a tough exoskeleton is enough to protect many arthropods But some arthropods have additional means of protection' Scorpions, bees, and some ants have venomous stings, and many spiders and centipedes have venomous bites. Lobsters and crabs can attack potential enemies with powerful claws And many insects and millipedes fight back using nasty chemh P . arthroPods trick predators by creating a diversion or example, some crabs can drop a claw or leg. This body part keeps on moving to distract predators while the rest of the aniOtherCUrtheS ^ The Crab then grOWS back the lost PmbOther arthropods use visual trickery to fool predators. Some ide through camouflage—matching the color and texture of their surroundings so closely that they seem to disappear. Others imitate the warning coloration of poisonous or dangerous species—a phenomenon called mimicry. MOVEMENT Arthropods have well-developed muscle systems that are coordinated by the nervous system. Muscles genera e force by contracting, then transfer that force to the toflexthp'i" AI i0int' SOme mUSC,es are Positioned ° th „ , an 0 er ,Tlusdes t0 exten<J it. See Figure 2813. The pull of muscles against the exoskeleton allows arthro¬ pods to beat their wings against the air to fly, push their legs Exoskeletons, as useful as they are, present a problem in terms of growth. As a growing student, you can understand that problem. Imagine that you had a skin-tight suit of clothes tai¬ lored for you last year. Could you fit into it now? Probably not. You need larger clothes as you grow. Similarly, arthropods must replace their exoskeletons with larger ones in order to allow their bodies to increase in size as they mature. The prob¬ lem is not a simple one, however, because the exoskeleton not only covers all appendages and sense organs, but also lines the gut as far down as the stomach. In order to grow, all arthropods must molt, or shed, their exoskeletons. This complicated proc¬ ess is controlled by several important hormones, the most im¬ portant of which is called molting hormone. When molting time is near, the epidermis (the layer of cells that covers the outside of the body) digests the inner part of the exoskeleton and absorbs much of the chitin in order to re¬ cycle the chemicals in it. After it secretes a new exoskeleton in¬ side the old one, an arthropod pulls completely out of its old exoskeleton. Arthropods often eat what is left of the old exo¬ skeleton. The animal then expands to its new, larger size, and the new exoskeleton (which is still soft) stretches to cover it. The animal must then wait for the new exoskeleton to harden, a process that can take from a few hours to a few days. During this time, the new shell stays soft and the animal is quite vul¬ nerable. Thus arthropods hide from predators during molting. Most arthropods molt several times between hatching and adulthood. In most cases the process of growth and develop¬ ment involves metamorphosis, or a dramatic change in form. Some arthropods, such as grasshoppers, mites, and crusta¬ ceans, hatch from eggs into young animals that look much like the adults. However, these young animals lack functioning sex¬ ual organs and often lack other adult structures such as wings. As the young grow, they keep molting and getting larger until they reach adult size. Along the way, they gradually acquire the characteristics of adults. In insects, this kind of gradual change during development is called incomplete metamorphosis. Figure 28-14 The adult cicada is emerging from the molted exoskeleton of an immature stage. Arthropods molt in order to increase in size and also to change from one body form to another in the process of metamorphosis. Figure 28-15 The grasshopper undergoes incomplete metamorphosis, whereas the monarch butterfly undergoes complete metamorphosis. wrterto'swim.011"'1 t0 Walk' ^ beat tlleir nipperS against the Muscle that flexes the joint Exoskeleton sim^eP|VMDUCT!)0r Reproduction in most arthropods is simple. Males and females produce sperm and eggs, respecive y, and fertilization usually takes place inside the body of e female, n spiders and some crustaceans, the male deposits Muscle that extends the joint second t SPerm that the female picks up- ln most insects and crustaceans, however, the male uses a special reproductive organ to deposit sperm inside the female. Complete Metamorphosis Incomplete Metamorphosis Older larva Newly hatched larva Figure 28-16 Insect pupae are often surrounded by a protective covering. The bumblebee pupa in this photograph is surrounded by a wax case. Many caterpillars spin cocoons of silk. Many insects, such as bees, moths, and beetles, undergo a four-stage process of development called complete metamor¬ phosis. Refer to Figure 28-15 on page 615 as you read about the process of complete metamorphosis. The eggs of insects that undergo complete metamorphosis hatch into larvae that look nothing like their parents. As these larvae grow, they molt re¬ peatedly, growing larger each time but changing little in ap¬ pearance. When a larva reaches a certain age, it sheds its larval skin one last time and becomes a pupa (PYOO-pah; plural: pupae). During the pupal stage, the insect's body is totally rearranged—adult structures grow from tiny buds and larval structures are broken down to supply the raw materials for the adult structures. When metamorphosis is complete, the animal emerges as a fully grown adult with both internal and external body parts that are completely different from what it had be¬ fore. Not only does this adult look like a totally different ani¬ mal, it acts differently too. Metamorphosis is controlled by a complicated interaction of several hormones, including molting hormone. In insects that undergo complete metamorphosis, the levels of juvenile hormone help regulate the stages of development. High levels of juvenile hormone keep an insect in its larval form each time it molts. As the insect matures, however, its production of juve¬ nile hormone decreases. At some point, the level of juvenile hormone drops below a certain critical point. The next time the insect molts, it becomes a pupa. And when no juvenile hor¬ mone is produced, the insect undergoes a pupa-to-adult molt. Because the balance of juvenile hormone, molting hor¬ mone, and other hormones is critical in arthropod develop¬ ment, it is possible to combat insects by tampering with their hormone levels. Certain plants defend themselves against her¬ bivorous insects by producing chemicals that prevent molting, cause insects to develop at the wrong rate, or keep insects from becoming functional adults. In recent years, researchers have developed chemicals that act in a similar manner. These chem¬ icals may eventually enable people to control crop-eating in¬ sects without using dangerous poisons. 28-2 Spiders and Their Guide For Reading ¦ What are several distinguishing Relatives characteristics of chelicerates? ¦ In what ways do spiders use silk? Spiders and their relatives—horseshoe crabs, ticks, and scorpions, for example-belong to the subphylum Chelicerata. Chelicerates are arthropods that are characterized by a twopart body and mouth parts called chelicerae. These arthro¬ pods also lack the sensory "feelers" that are found on the heads of most other arthropods. All chelicerates have a body that is divided into two parts: the cephalothorax (sehf-ah-loh-THOR-aks) and the abdomen. The anterior end of the cephalothorax contains the brain, eyes, mouth and mouthparts, and esophagus. The posterior end of the cephalothorax carries the front part of the digestive system and several pairs of walking legs. The abdomen contains-most of the internal organs. See Figure 28-17. All chelicerates have two pairs of appendages attached near the mouth that are adapted as mouthparts. The first pair of mouthparts are called chelicerae (keh-LlHS-er-ee; singular: chelicera) The second pair of mouthparts, which are longer than the chelicerae, are called pedipalps (PEHD-ih-palps). Both sets of mouthparts are adapted to serve different purposes in Figure 28-17 The internal structures of a typical spider are shown in this diagram. A jumping spider (inset) captures prey by pouncing on it, rather than by catching it in a web. feeding in different species. Horseshoe Crabs Among the oldest chelicerates are the horseshoe crabs. This name is somewhat misleading because these animals are not true crabs (which are crustaceans). Horseshoe crabs ap¬ peared in the Ordovician Period (more than 430 million years ago) and have not changed much since then—they are true "living fossils." Horseshoe crabs are heavily armor-plated, Pumping stomach Heart - |j 1 SECTION Malpighian tubules ll REVIEW 1. What are three characteristics of arthropods? Name the four subphyla of arthropods. 2. Compare complete and incomplete metamorphosis. 3. Describe the different types of organs that are used in arthropod respiration. 4. Critical Thinking—Making Inferences Terrestrial arthropods often have valves that can open and close the spiracles. How are these valves an adaptation to life on land? (Hint: What is the function of the stomata on leaves?) 616 Walking leg Poison gland Pedipalp Anus Nerve cord Spiracle Spinneret have five pa.rs of walking legs, and long spikelike tails The can grow up to 60 centimeters long-about the s ze I y Shape) of a frying pan. When they first hatch, however hors h reh are 0n'y about 1 cent™eter long. These newf hatched horseshoe crabs are called frilobite larvae bee™ y they look much like their extinct distant relatives. Arachnids rh,rilhe ^OSt faiTliJiar chelicerates are the arachnids, which in ude spiders, scorpions, ticks, and mites. AJI adult arachnid, have four pans of walking legs on their cephalothor^ ^ Figure 28-18 A horseshoe crab's tiny pincerlike chelicerae and five pairs of walking legs are visible when the animal is turned on its back. The platelike structures on the abdomen cover and protect the book gill's "pages." The long tail is not seen here because the horseshoe crab has pushed it into the sand to right itself. arachnids are carnivores that have pedipalps adapted for can unng and holding prey and chelicerae adapted for biting and sucking out their soft parts. 8 d sei.t!PIl?ERS Spiders are predators that usually feed on in sects. However, a few large tropical spiders are capable of Sn rie"18 anf eatl!!g Smal1 vertebrates. stJch as hummingbirds Spiders capture their prey in a variety of ways. Some spider, tehneSnnarre T". ^ WebS' 0thers stalk then pouTce on the prey. And some he in wait beneath the lid of a camouflaged Once a spider captures its prey, it uses its hollow fanglike c e icerae to inject paralyzing venom into it. When the prey is paralyzed, the spider's mouth introduces enzymes into toe wound5 made by the chelicerae. These enzymes break down tissue, with hUeS' ehabling the Spider 10 Suck up ,he ^^ficd tissues with Its esophagus and a specialized pumping stomach Figure 28-19 The wolf spider (left) ambushes prey from its silk-lined burrow. Large tarantulas (right) are capable of catching and devouring small vertebrates, such as lizards. The pumping stomach then forces the liquid food through the rest of the spider's digestive system Whether or not they spin webs, all spiders produce a strong flexible protein called silk. Silk, which is produced in special glands located in the abdomen, is five times stronger than steel. It is strong enough, in fact, to withstand the equiva¬ lent of the impact of a jet fighter every time a spider's web traps a fly. Spiders spin silk into webs, cocoons for eggs, wrap¬ pings for prey, and other structures by forcing liquid silk through organs called spinnerets. As the liquid silk is pulled out of the spinnerets, it hardens into a single strand. Interest¬ ingly, the complicated behavior of web-spinning seems to be "preprogrammed" into a spider's brain. The spiders of webspinning species can build their intricate webs almost as soon as they hatch—without having to learn how. MITES AND TICKS Mites and ticks are small arachnids, many of which are parasites on humans, on farm animals, and on important agricultural plants. Most species are smaller than 1 millimeter, but some ticks can be as large as 3 centimeters. In many mites and ticks, the chelicerae are needlelike structures that are used to pierce the skin of their hosts. These chelicerae may also have large teeth to help the parasite keep a firm hold on the host. The pedipalps are often equipped with claws for digging in and holding on. Some species, such as spider mites, damage houseplants and are major agricultural pests on crops such as cotton. Others—including chiggers, mange, and sca¬ bies mites—cause painful itching rashes in humans, dogs, and other mammals. A whole host of ticks parasitize humans and the animals we raise. Tick bites are not just annoying—they can be dangerous. In the United States, ticks can spread Rocky Mountain spotted fever and Lyme disease. SCORPIONS Scorpions are widespread in warm areas around the world, including the southwestern United States. All scorpions are carnivores that prey on other invertebrates, usually insects. The pedipalps of scorpions are enormously en¬ larged into a pair of claws. The abdomen, which is long and segmented, terminates in a venomous barb used to sting prey. Usually, a scorpion grabs prey with its pedipalps, then whips Figure 28-20 Some spiders build webs to capture prey. Figure 28-21 Red velvet mites are similar in form to other members of their class (right). However, they ore unusual in that they are not parasites and are relatively large (about 1 centimeter long). The loser of a fight between two scorpions will be stung and eaten by the winner (left). Biologists can locate scorpions at night by shining ultraviolet (UV) light on the desert floor. Under UV light, scorpions glow brightly in the dark. Abdomen ——Cephalothorax i Second^antenna its abdomen over its head to sting the prey, thus killing or par¬ alyzing it. The scorpion then chews its meal with its chelicerae Because scorpions like to crawl into moist, dark places, people in areas with scorpions should check inside their shoes before putting them on in the morning. Most North American scor¬ pions have venom powerful enough to cause about as much pain as a wasp sting. However, the venom of one genus of scor¬ pions that lives in Mexico, New Mexico, and Arizona has killed small children who were stung accidentally. Carapace-^ j Heart Green gland | Brain f ^ Gonad (testis or ovary). Intestine. Figure 28-23 The internal and external structures of a crayfish are shown here. Can you now explain why crayfish, shrimp, lobsters, and crabs are sometimes known as Nerve cord decapods (deca- means ten)? Mandible Swimmerets Digestive SECTION gland Walking leg ¦ REVIEW 1. What are chelicerates? Name and give examples of the two main groups of chelicerates. 2. What is silk? How do spiders use silk? 3. Critical Thinking—Summarizing Information How are chelicerae modified for feeding in spiders? In ticks? Guide For Reading 28-3 Crusty ceans ¦ What are crustaceans? ¦ How are the body parts of crustaceans adaptations for survival? Figure 28-22 The pill bug is a terrestrial crustacean. When threatened, a pill bug curls into a ball to protect its soft underside. 620 The subphylum Crustacea contains over 35,000 species. Crustaceans are primarily aquatic, although there are some ter¬ restrial species. Crustaceans range in size from microscopic water fleas less than 0.25 millimeter long to Japanese spider crabs that are thought to grow up to 6 meters across and lob¬ sters that have a mass of more than 20 kilograms. And crusta¬ ceans vary in form as much as they vary in size! Although crustaceans adapted to different conditions are quite dissimilar in form, all crustaceans share a number of structural similarities. In general, crustaceans are character¬ ized by a hard exoskeleton, two pairs of antennae, and mouthparts called mandibles. As we examine a little of the enormous diversity of form and function in crustaceans, we will focus on a representative species, the crayfish. Refer to Figure 28-23 as you read about structure and function in crustaceans. The main crustacean body parts are the head, thorax, and abdomen. In crayfish, as in many other crustaceans, the head and thorax have fused into a cephalothorax that is covered by a tough shell called the carapace. Unlike most other arthropods, many large crustaceans have calcium carbonate (limestone) in the exoskeleton. This is what makes the shells of crustaceans such as crabs and lobsters hard and stony. In crustaceans, the first two pairs of appendages are "feelers" called antennae, which bear many sensory hairs. An¬ tennae serve primarily as sense organs in crayfish, but in some other crustaceans they are used in filter feeding. Still other crustaceans, such as water fleas, use their antennae as oars to push them through the water. The third pair of appendages are mouthparts that are called mandibles. In many species of crustaceans, including crayfish, mandibles are short heavy structures designed for biting and grinding food. In other species, mand.bles are bristly structures used in filter feeding, probelike structures used for finding and picking up detritus, or needlelike structures used to suck blood from a host. The appendages on the thorax and abdomen vary greatly from one group of crustaceans to another. Some such as barnacles, have delicate, feathery appendages for filter feeding, others have legs for walking or paddles for swimming. Appen¬ dages may be modified for internal fertilization, carrying eggs, soearing prey, burrowing, or many other functions. " As you can see in Figure 28-23, the appendages on a cray¬ fish's thorax and abdomen are adapted for several different functions. A pair of large claws, which are used to catch prey and pick up, crush, and cut food, are located on the thorax. Four pairs of walking legs are also attached to the thorax. Flip¬ perlike appendages called swimmerets. which are used for swimming, are located on the abdomen. A large pair of paddlelike appendages are found on the second-to-last abdomina segment. The paddlelike appendages and the final abdomina segment together form a large, flat tail. When the muscles of the abdomen contract, the crayfish's tail snaps fo™rd- This provides a powerful swimming stroke that can rapidly pull the animal backward. Figure 28-24 The abdomen of a crab is tucked beneath its cephalothorax. A female crab uses its abdomen and the swimmerets attached to it to carry its reddishbrown eggs. The blue and white semicircle below the eggs is made up of the last few segments of the tail. 1. What is a cephalothorax? 2. Describe the types of appendages on crayfish and give their functions. 3. Critical Thinking—Applying Concepts Suppose you want to catch a crayfish with a net. Should you try to scoop it up head first or tail first? Explain. 621 Antenna Wings Guide For Reading What are the characteristics of the three classes in the subphylum Uniramia? What anatomical adaptations do insects have that enable them to survive? How do insects communicate with one another? 28-4 Insects and Their Relatives The subphylum Uniramia contains more species than all other groups of animals alive today. It includes centipedes, millipedes, and insects. Uniramians are characterized by one pair of antennae and appendages that do not branch (unimeans one; ramus means branch). (Recall that the appendages in crustaceans and trilobites have two branches—usually a gill and a leg.) These arthropods, which display a multitude of forms and habits, are thought to have evolved on land about 400 million years ago. They inhabit almost every terrestrial habitat on Earth. In addition, some species live in fresh water and a few other species live in marine environments. Centipedes and Millipedes Figure 28-25 A centipede (top) is a carnivore that has one pair of legs per body segment. A millipede (bottom) is a herbivore that has two pairs of legs per body segment. Centipedes and millipedes are many-legged animals. Com¬ pared to crustaceans and insects, these two classes of arthro¬ pods are quite small in number—there are approximately 3000 species of centipedes and 7500 species of millipedes. Centi¬ pedes and millipedes are characterized by a long, wormiike body composed of many leg-bearing segments. Because they lack closable spiracles and a waterproof coating on their exoskeleton, their bodies lose water easily. Thus they tend to live beneath rocks, in soil, or in other relatively moist areas. CENTIPEDES Centipedes are carnivores that have, in ad¬ dition to other mouthparts, a pair of poison claws in their head region. These poison claws are used to catch and stun or kill prey. Centipedes eat other arthropods, earthworms, toads, small snakes, and even mice. The North American centipedes that may be familiar to you are usually red-brown in color and about 3 to 6 centimeters long. Some tropical species are brightly colored and quite large—up to 26 centimeters long. Despite their name, which means 100 legs (centi- means hundred; -pede refers to legs), centipedes may have from 15 to 170 pairs of legs, depending on the species. Each segment that makes up the body of the centipede bears one pair of legs, ex¬ cept for the first segment (which bears the poison claws) and the last three segments (which are legless). Insects We know of more than 900,000 insects, and new ones are discovered in the tropics all the time. Insects are extremely varied in body shape and habits. However, all members of this class share basic structural similarities. Insects are character¬ ized by a body that Is divided into three parts—head, thorax, and abdomen—and that has three pairs of legs at¬ tached to the thorax. In addition, a typical insect has one pair of antennae and one pair of compound eyes on the head, two pairs of wings on the thorax, and uses a system of tracheal tubes for respiration. Insects get their name from the Latin word insectum, mean¬ ing notched, which refers to the division of their body into three main parts; head, thorax, and abdomen. In many insects, such as ants, the three body parts are clearly separated from each other by narrow connections. In other insects, such as grasshoppers, the divisions between the three body parts are not as sharply defined. The essential life functions in insects are carried out in ba¬ sically the same ways as they are in other arthropods. How¬ ever, insects show a variety of interesting adaptations in feeding, movement, and behavior that deserve a closer look. FEEDING Insects have three pairs of appendages that are used as mouthparts, including a pair of mandibles. Mouthparts can take on an enormous variety of shapes in species adapted to feed on different foods. For example, a grasshopper's mouth¬ parts are designed to cut and chew plant tissues into a fine pulp. A female mosquito's mouthparts form a sharp tube that is used to pierce skin and suck blood. A butterfly's mouthparts Grasshopper Moth Head Abdomen Thorax Figure 28-26 An insect is characterized by a three-part body, six legs, two pairs of wings, one pair of antennae, and one pair of compound eyes. Figure 28-27 Although insect mouthparts are adapted for many different eating habits, they all evolved from the same basic Housefly MILLIPEDES Although millipedes do not have a thousand legs (milli- means thousand), they do seem to have twice as many as centipedes. Each millipede body segment is formed from the fusion of two segments in the embryo and thus bears two pairs of legs. Millipedes are timid creatures that live in damp places under rocks and in decaying logs. They feed on dead and decaying plant material. When disturbed, many milli¬ pedes roll up into a ball to protect their softer undersides. Some can also defend themselves by secreting unpleasant or toxic chemicals. 623 Si~S=SH5SS Figure 28-29 Leaf-cutter ant workers carry pieces of leaves and flower petals to their underground nest. Certain chambers in the nest are "farms" in which the ants grow rnmmm edible fungi on the bits of vegetation. ting. (Unfortunately for h^manVand ^ Clot- saliva also contains chemicals thAt animals, mosquito reaction. SSSSi have a number of adAntAti^r... I ^ Honeybees storing food. The legs and bodieT ® ™g' Processing, and with hairs that coUect nollen Ph ^ ^ ^ COVered change nectar inte^a more digest?ble™rmS b ^ fT helP on the abdomen secrete wal which ^ y' d glanc[s chambers for food and other struck: X': belhivT^ Figure 28-28 Insects, along with birds and bats, are the only living organisms capable of unassisted tght. 1 he hard brown wing covers on the may bug ore modified forewings; in may bugs and other for fiTghL^ ^ hindwin^ are used during flight is k - oy these muscles r l-jt=====-" efficiendye^wrenhler^r^ and ~ nies'which Sctln^ fy ^ ^ ^ — that live together Several tvr>ps TldUa'S 0f the Same SPecies invertebrates in that thev fn 0 lnsects are un'que among as a society In fsoc etv Z * ZZ tyPe 0f COlony known on one another for^^fva ^3 are.dependent 624 7 million individuals. Within such societies there is division of labor: Different individuals perform the tasks necessary for the survival of the entire group. There are several castes, or types of individuals, within insect societies. Each caste has a body that is specialized for its functions and is therefore distinctly different from that of another caste. The basic castes are repro¬ ductive females, reproductive males, and workers. The reproductive females, which are called queens, lay eggs that hatch into new individuals for the society. Most in¬ sect societies have only one queen, who is typically the largest individual in the colony. Termite queens, for example, may be 14 centimeters long (more than 10 times longer than a worker) and 3.5 centimeters wide. Most of a termite queen's body con¬ sists of a grotesquely swollen abdomen that contains enlarged reproductive organs. A termite queen can produce more than 30,000 eggs a day! The reproductive males function only to fertilize the queen's eggs. In some insect societies, such as those of ter¬ mites, a single reproductive male stays with the queen as a per¬ manent member of the colony. In other societies, such as those of bees, the queen receives all the sperm she needs for her eggs after a single mating with one or more reproductive males. The successful males die after mating, and the unsuccessful males are ejected from the colony and soon perish. The workers perform all the colony's tasks except for re¬ production: They care for the queen, eggs, and young; they gather, store, and even grow food; they build, maintain, and de¬ fend the colony's home; and they perform all other necessary jobs. In societies of ants, bees, and wasps, the workers are all females; in those of termites, there are both male and female workers. Bee and wasp workers are capable of performing all of their societies' tasks for workers. Ant and termite workers are specialized and are able to carry out only their specific tasks, such as defending the colony or storing food. Figure 28-30 Mature termite queens are approximately the size and shape of a hot dog. The large termite next to the queen is a reproductive male. The smaller brown termites with the large heads are called soldiers. The tiny white termites are workers. INSECT COMMUNICATION All insects use sound, visual ofThT and 0ther lyPeS 0f Signals for commun'cati0n. Much the communication done by nonsocial insects involves find mg a mate. To attract females, male crickets chirp by rubbing eir fore wings together, and male cicadas buzz by vibratin? special membranes on the abdomen. Male fireflies turn a light producmg organ in their abdomens on and off, producing a disct series of flashes. When female fireflies (which are wingless and are known as glowworms) see the correct signal ey flash back a signal of their own and the males fly to them' view ^th ^ 3 g00d thing ^ the maIe firefly,s Point of v.ew the carnivorous females of one genus of fireflies can deX L S1fnal 0f anoth€r Senus and lure males to their death. Many female moths release chemicals that attract disant males to them. These chemicals are a type of pheromone and / SP. C chem,(:al messenger that affects the behavior and/or development of other individuals of the same species Communication in social insects is generally more complex an in nonsocial insects. A sophisticated system of communiFigure 28-31 A male tana moth's feathery antennae can detect pheromones released by a female several kilometers away. insert a society. Each species of social insect has its own language" of visual, touch, sound and the colony.18113 that information among members of Certafn^nhT"68 are particularly iniPortant in insect societies. ^ n Tm0T a nCti0n aS rapid messages at signal alarm, the death of a member of the colony or the she'head l ^ eXample' when a worker ant finds food, heads back to the nest, dragging her abdomen along the g ound. As she does so, she leaves behind a trail of a special m of Pheromone. Her nestmates can detect her trail by using Other'nh ' 0n ^ antennae and fol,ow ^ back to the food, er pheromones act as long-term controls over the colony or example, a queen honeybee produces a pheromone, called queen substance, that prevents the development of rival eggs lt ahnT" SUbtShtanCe makes worker bees unable to lay afQl]' a. H aUSeS T t0 raiSe fema,e larvae as ^rkers, not fhe hivp k , WeVe,i' When the amOUnt 0f q1166" stance in diet This T f 665 feed a feW female ,arvae a sPecial diet. This causes the larvae to develop into queens Honeybees communicate with sound and movement as well as with pheromones. Worker bees are able to convey infor¬ mation about the type, quality, direction, and distance of a food coded6bv^A Jhe ,anguage of the bee's dance was deered th/t K T b,0!0glSt KarI VOn FriSCh- Von Fr'sch discov- wTggle danc^05 ^ ^ danCeS; a ^d da- and a In the round dance, the bee that has found food circles first tells The othP^h11 th?H0thlr' 0Ver and 0Ver again' This dance other bees that there is a source of food within 50 meters of the hive. The frequency with which the dancing bee Round Dance Waggle Dance Figure 28-32 Bees communicate information about food sources by using a language of movement. The round dance tells members of the hive that a source of food is nearby. The waggle dance gives information about a more distant food source. changes direction indicates the quality of the food source—the more frequent the changes in direction, the greater the energy value of the food. By smelling the dancer with the chemical re¬ ceptors on their antennae, the other bees can determine what kind of flowers she has found. In the waggle dance, the bee that has found food runs for¬ ward in a straight line while waggling her abdomen, circles around one way, runs in a straight line again, and circles around the other way. See Figure 28-32. The waggle dance tells the other bees that the food is more than 50 meters away. Most of the information about the food source is conveyed by the part of the waggle dance called the straight run. The longer the bee takes to perform the straight run and the more she wag¬ gles, the farther away the food. The direction of the straight run indicates in which direction the food is to be found. For exam¬ ple, if the dancer runs straight up the vertical honeycomb, the food source is in the same direction as the sun. SECTION REVIEW _ 1. Compare the body plans and feeding habits of millipedes and centipedes. 2. Describe the basic body plan of an insect. 3. Give three specific examples of why and how insects communicate. 4. Explain how the mouthparts of bees, mosquitoes, and butterflies are adapted to different food sources. 5. How does the waggle dance of honeybees convey information about the location of a food source? 6. Critical Thinking—Relating Cause and Effect If all worker bees are females, why is the queen the only egg-layer in the colony? 627 TECHNOLOGY, AND SOCIETY Controlling Agricultural Pests Many plant-eating insects are enemies of both crops and plants in nature. Yet although cornfields can be destroyed by insects, forests and grasslands are never wiped out. Why not? And how can the answer to that question help us protect food crops? Pests rarely get out of control in nature for three main reasons. First, they are eaten by such natural enemies as ladybugs, spiders parasitic wasps, and birds, and suffer from dis¬ eases caused by bacteria and fungi. Second many plants produce compounds that taste bad to insects, poison them, or mimic the ac¬ tion of insect juvenile hormones. Larvae that eat these hormone mimics cannot pupate never mature, and cannot reproduce Third natural environments contain several plant species mixed together, so insect pests have to work harder to find their favorite foods. For many years now. farmers have taken their cues from poisonous plants, and have re- hed on toxic sprays such as DDT to kill pests It is clear, however, that widespread, constant use of poison sprays has created more prob¬ lems than it solved. Many pesticides are deadly to humans, livestock, wildlife, and the natural enemies of pests. Some chemicals stay m the environment for a long time, poisoning rivers and water supplies. Furthermore, many insecticides are no longer useful because pests have evolved resistance to them. In 1938, there were only 7 pests that were resistant to chemi¬ cals; by 1985 there were 447. Clearly, different insect control methods are needed. Researchers have therefore fo¬ cused their attention on pests' natural enemies and on ancient agricultural practices. A great deal has been learned in recent years. Insect biologists are learning to breed and collect pest-eating insects, and to release them in fields at the right time to control pests. Biotechnology companies are learning to manufacture hormone mimics economi- Mult ladybugs and their larvae feed on crop- leT? /n ^ /0ng run' the use of natural pest controls is safer and more efficient and pesiHides USe ofarm^ chemical cally. Farmers are learning to use these non¬ toxic sprays, which affect only the insects at which they are targeted and remain useful for a long time. (Pests cannot usually evolve resis¬ tance against hormone mimics, because that would involve extensive modifications to its complex internal control system.) Finally, ecologists and anthropologists have discovered that certain traditional agri¬ cultural practices control plant pests and dis¬ eases remarkably well. Some practices en¬ courage animals that prey on pests to live in farmers fields. Others control harmful insects by starving them—through planting different crops in different years and leaving some fields unplanted every few seasons. 28-5 How Arthropods Fit into the World As you might expect from such a large, diverse group of an¬ imals, arthropods play many roles in the natural world. They are a direct source of food for many carnivorous organisms— from protists such as radiolarians to plants such as Venus' fly¬ traps to animals such as sea anemones, fishes, frogs, turtles, birds, whales, and humans. And they are also an indirect source of food for many other organisms. The interactions of arthropods with other organisms are not limited to eating or being eaten. Two thirds of the world's flowering plants depend on insects to pollinate them. Some plants live even more intimately with arthropods. The bull's horn acacia tree has hollow swellings at the base of some of its thorns that house symbiotic ants and special structures that feed the ants. The ants protect the acacia by eating herbivorous insects and by driving away larger herbivores with their painful bites and stings. Animals are also involved in symbiotic rela¬ tionships with arthropods. It is amazing to see a large fish allow a bite-sized cleaner shrimp to crawl on its body and even into its mouth. But by allowing the cleaner shrimp to go un¬ harmed, the fish is cleaned of annoying parasites and bits of dead tissue, and the cleaner shrimp gets a meal. Figure 28-34 The cleaner shrimp and fish such as the queen angelhsh, shown here, engage in a symbiotic relationship that benefits both organisms. The cleaner shrimp gets a meal by eating parasites on the hsh and the fish gets rid of annoying pests. A number of fishes can sometimes be observed congregating around a shrimp's cleaning station, waiting for their turn to be cleaned! Guide For Reading How do arthropods interact with other organisms in nature? In what ways do arthropods affect humans? Figure 28-33 The damsel fly has been trapped by a carnivorous sundew plant. Although sundews and other carnivorous plants are photosynthetic, they need to "eat" insects and other small animals to obtain the nitrogen compounds they need to survive. Today, agricultural experts favor combin¬ ing all these methods into a strategy called In¬ tegrated Pest Management or IPM. IPM strives to control pests more naturally, while using dangerous poison sprays only in emergencies This tactic has dramatically reduced the use of pesticides in many places around the world. Importantly, farmers are discovering that 1PM •s often cheaper in the long run. This new technology thus has the potential to improve our environment while saving farmers world¬ wide many millions of dollars each year. 629 P IN BIOLOGY T~^\ Humans encounter arthropods almost everywhere Th pores of our skin are home to thousands of harmless micr 6 scopic mites that feed on dead skin and oil. And no matter how zl neat and clean we are, our homes, even our beds, contain mil Figure 28-35 Millions of microscopic dust mites live in human homes. The mites feed on the tiny flakes of dead skin that are constantly being shed from the bodies of humans. lions of microscopic dust mites. For the most part, these tin harmless mites are of little consequence. But many of the more visible arthropods are of significance because they are either useful to us or a great nuisance. Arthropods contribute enormously to the richness of human life. Agriculture would be impossible without the bees butterflies, wasps, moths, and flies that pollinate crops. Bees manufacture honey, and silkworms produce silk. In Southeast Asia and Japan, whole shrimp and shrimp paste are important sources of protein and major ingredients in cooking. In the United States, shrimp, crab, crayfish, and lobster are consid¬ ered delicacies. In Africa and Asia, many people eat insects such as grasshoppers and termites. (These insects, which are quite nutritious, are said to taste rather good.) And many in¬ sects and spiders are predators or parasites that prey upon harmful species. There are many useful chemicals that may be obtained from arthropods—far too many to list here. An extract of horseshoe-crab blood, for example, is used to test the purity of medications. Chitin extracted from crustacean shells is used to dress wounds and to make thread for surgical stitches. The chemical that makes fireflies glow is used in medical tests and as a marker in genetic engineering. Many new applications of arthropod chemicals are cur¬ rently being investigated. For example, chitin could be sprayed onto fruit and frozen food to prevent spoilage and to preserve flavor. The adhesive that barnacles use to attach themselves to rocks, which sets quickly and hardens under water into a per¬ manent bond, could be useful in applications ranging from den¬ tistry to underwater construction and repair. Chemicals in spider venom are being tested for potential applications as pesticides. And scientists are currently trying to produce ge¬ netically engineered spider silk that could be used in making products as diverse as aircraft, helmets, bulletproof vests, and surgical thread. Not all arthropods are beneficial to humans, however. In¬ sects (such as locusts and "medflies") and arachnids (such as mites and ticks) cause billions of dollars in damage each year to livestock and crops around the world. Mosquitoes inflict n identifying arthropods Having heard that you are now an expert on arthropods, some of your friends bring you the rreatures pictured below and ask you to iden¬ tify them. Can you place each in the correct arthropod group? Be as specific as possible. (Hint: Check the following important features, number of body segments, walking legs, and pairs of antennae; presence of wings; presence of claws. Compare the animals with the de¬ scriptions and photos in the text.) annoying bites, and some species carry malaria and yellow fever Biting flies carry diseases such as sleeping sickness and rC blindness, and fleas can carry bubonic plague. Termites cause extensive damage to wooden structures. Locusts have destroyed crops from the time humans first began to farm. Boll weevils are notorious for the trouble they cause cotton fanners in the South. For years, farmers have spent billions of dollars on poisonous chemicals to save their crops from these pests. mm section 1.' : R E V 1 E W 1. Why are certain insects essential to agriculture? Figure 28-36 Some species of grasshoppers exist in two distinctly different forms: a dull-colored solitary grasshopper or a brightly colored gregarious locust (shown here). Locusts travel in immense swarms that may contain as many as 50 thousand million individuals. A swarm can devastate huge areas of crops. One swarm destroyed 167,000 tons of growing grain—enough to feed 1 million people for a year. 2. How are arthropods beneficial to other living things? Give specific examples. 3 Critical Thinking—Relating Concepts Name three dangerous or destructive arthropods and explain how they cause problems for humans. ^ vS* I w w * to i I p PROBLEM What changes occur as some insects grow and develop? The key concepts in each section of this chapter are listed below to help you review the chapter content. Make sure you understand each concept and its relationship to other concepts and to the theme of this chapter. MATERiALS (per group) 600-mL beaker cheesecloth corn flakes dissecting tray SUMMARIZING THE CONCEPTS hand lens 25 mealworms probe rubber band 4. Put the mealworm back into the beaker of corn flakes. Cover the beaker with a piece of cheesecloth and secure the cheesecloth in place with a rubber band. 5. Check the beaker every other day for a few months by moving the probe carefully through the corn flakes. Be careful not to injure the lar¬ vae. Look for any changes in the larvae with respect to size or shape. Do you find any lifeess shells that look like the exoskeletons of the larvae? 6. If you find a short, thick motionless footballshaped object among the corn flakes, carefully slide it mto the dissecting tray with a probe I his is the pupal stage of the Tenebrio beetle Carefully examine the pupa with a hand lens' noting the location and number of append¬ ages. Draw a diagram of the pupa. 7. Put the pupa back into the beaker. Continue beet'lef 0bServations until y™ find adult 28-4 Insects and Their Relatives • Arthropods are characterized by an exoskeleton of chitin, jointed appendages, and a segmented body. • Uniramians include centipedes, millipedes, and insects. • In order to grow, arthropods must periodi¬ cally shed their exoskeletons in a process called molting. PROCEDURE 1 B 1. Fill a 600-mL beaker halfway with corn flakes. ni 25 ™al— into the beaker of corn flakes. Observe their behavior. 3. Using a probe, slide one of the mealworms mto a dissecting tray. The mealworm is the arvai stage of the Tenebrio beetle. Carefully examine the larva with a hand lens, noting the location and number of appendages. Draw a diagram of the larva. 28-1 Introduction to Arthropods 8. Do not remove the beetles from the con flakes, as they may fly. Instead, try to keep ; beetle uncovered in the beaker and examine il with a hand lens. Note the location and adult" 0f appendages- Draw a diagram of the OBSERVATIONS 1. What did the mealworms do when you placed them in the corn flakes? ® Centipedes are carnivores that have poison claws and possess one pair of legs per body segment. Millipedes are herbivores that have two pairs of legs per body segment. ® The process of growth and development in arthropods often involves metamorphosis, • Insects have a body that is divided into three parts: head, thorax, and abdomen. They have three pairs of legs attached to the thorax. 2.8-2 Spiders and Their Relatives ® Members of insect societies are specialized for performing different functions. ® Chelicerates have a body that consists of two parts—cephalothorax and abdomen. Cheli¬ cerates have chelicerae and lack antennae. • Arachnids, such as spiders, scorpions, and mites, are typically carnivores that have four pairs of walking legs. ® Insects communicate. Some forms of commu¬ nication rely on pheromones. 28-5 How Arthropods Fit into the World ® Arthropods play many roles in the natural world, 28-3 Crustaceans ® Crustaceans, such as crabs and crayfish, are characterized by a stony exoskeleton, two pairs of antennae, and mandibles. ® Some arthropods are of little significance to humans; others are important because they are useful or a great nuisance. 2' """J 'egS d0es each of the stages have? What other appendages does each of the stages have? 3. What happened to the sizes of the larvae over timer 4. Compare the larval, pupal, and adult stages in terms of appearance and behavior. ANALYSIS AND CONCLUSIONS 1. What did the mealworms use for food during growth and development? 2. What evidence did you find of molting? Why is molting necessary? 3. What changes occur as Tenebrio beetles grow sm and develop? REVIEWING KEY TERMS Vocabulary terms are important to your understanding of biology. The key terms listed below are those you should be especially familiar with. Review these terms and their meanings. Then use each term in a complete sentence. If you are not sure of a term's meaning, return to the appropriate section and review its definition. 28-1 Introduction to uniramian exoskeleton trilobite chitin molt chelicerate crustacean pupa arthropod metamorphosis 28-2 Spiders and Their Relatives arachnid 2B-4 Insects and Their Relatives pheromone 28-3 Crustaceans mandible 632 633 c o N c E P "LM AST E R Y CONTENT REVIEW Choose the letter of the answer that best completes each statement. 1. Which of these is an arachnid? a. scorpion c. grasshopper b. horseshoe crab d. lobster 2. A free-living arthropod is certain to have a. antennae. c. jointed appendages. b. chelicerae. d. gills. 3. insects are characterized by a. a stony exoskeleton containing calcium carbonate. b. chelicerae and pedipalps. c. three pairs of legs on the thorax. d. many body segments, each of which bears two pairs of legs. 4. In crustaceans, nitrogenous wastes are 5. A wormlike immature animal undergoes a resting stage during which it changes into an adult that has four wings and six legs. This animal is a (an) a. crustacean. c. chelicerate. b. insect. d. trilobite. 6. Most spiders breathe using a. mandibles. c. Malpighian tubules. b. tracheal tubes. d. book lungs. 7. Which is most likely to be a herbivore? a. spider c. tick b. centipede d. millipede 8. Trilobites a. are primarily terrestrial. excreted with the help of a. green glands c. Malpighian tubules. c. have highly specialized appendages. b. spiracles d. pheromones. d. communicate by "dancing." b. are extinct. Useyourunders,anding of the concerts «ed chaPter to ansuter eactt of the following in a brief paragraph. arthropods belong to the subphylum , Beetles undergo complete metamorphosis Mandibulata according to this classification ' and dragonflies undergo incomplete scheme? Explain why many experts do not metamorphosis, Describe the major events favor this method of grouping arthropods. Of the life cycles of beetles and drag°""ies3. Using a crayfish as your representative Be sure to include a comparison of their organism, discuss the distinguishing characteristics of arthropods. 9 Irfsom^ciassification schemes, arthropods 4 Certain chemicals bind with juvenile ' are divided into two subphyla—Chehcerata hormone and make it inactive. Describe and Mandibulata—based on the type of how exposure to such chemicals would mouthparts they possess (chelicera and affect the development of a moth. mandibles, respectively). Which groups of CRITICAL AND CREATIVE THINKING Discuss each of the following in a brief paragraph. 1. Applying concepts Explain why you will never see spiders three stories tall or ants big enough to eat New York (excep True or False Determine whether each statement is true or false. If it is true, write "true." If it is false, change the underlined word or words to make the statement true. 1. If the level of juvenile hormone in an insect's body is high, the insect is in the pupal stage. 2. Arthropods have a closed circulatory system. 3. Centipedes are herbivores that have two pairs of legs on each segment. 4. A spider uses fanglike pedipalps to inject venom into its prey. 5. All the members of the class Uniramia are now extinct. 6. Horseshoe crabs are classified as crustaceans. 7. Arthropods are characterized by an exoskeleton composed of calcium carbonate. 8. Arthropods must periodically molt, or shed, their exoskeletons. Word Relationships some crabs are soft-shelled. 3. Relating cause and effect ^ople who squash an annoying hornet are often unpleasantly surprised to find themselve suddenly under attack by dozens of hornets. Explain this phenomenon, 4. Making inferences Instead of spray.ng a field with chemicals, a plane disperses tens of thousands of tiny wasps oyer he In each of the following sets of terms, three of the terms are related. One term does not belong. Determine the characteristic common to three of the terms and then identify the term that does not belong. 1. tracheal tube, book lung, spiracle, pupa 2. green gland, gill, Malpighian tubule, chitin 2 Relating0'concepts Blue crabs usually have ha'rd, stony shells. However some blue crabs have thin, papery shells^ These crabs are called soft-shell crabs and are a popular food for some people, who eat them whole-shell and all! Explain why 3. mandible, chelicera, pedipalp, walking leg 4. chelicerate, uniramian, crustacean, insect growing plants. What is the most likefy reason for such an action? 3 Anplving concepts At the park one day, ¦ you observe a bee flying around an open can of soda. Soon after, you notice that there are a lot of bees buzzing around this can However, there are no bees on other open cans of soda a few meters away, a Explain how the bees probably found the first can of soda, b Explain why the bees do not seeni interested in the other cans of soda. 6. Assessing concepts Which do you think is a better arrangement for an insect society: having workers that can each perform all necessary tasks or having workers that are specialized for specific tasks? Explain your answer. 7 Using the writing process Certam crabs • have a peculiar symbiotic relationship with coral- They cause branches of coral to grow around them to form a P">tect've orison The imprisoned crab obtains fo and oxygen from the currents of water that flow through its coral cage. Write a short story or play in which one of these imprisoned crabs converses with a more typical crustacean.