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Biology 320 Invertebrate Zoology Fall 2005 Chapter 21 – Phylum Arthropoda, Superclass Hexapoda Class Insecta Majority of all animals 1,000,000 described species Estimated 10 to 30 million undescribed Do not live in oceans, with the exception of a few intertidal species Well adapted for terrestrial existence 2 – 250 Malpighian tubules Many other adaptations and behaviors Holometabolus (indirect) development has led to success on land by promoting resource partitioning Largest insect orders (Lepidoptera, Hymenooptera, and Diptera) exhibit indirect development Insect flight allows for: Dispersal Predator escape Access to food and optimum habitat Many have interesting coevolutionary relationships with flowering plants Some are vectors of disease Herbivory by insects does large amounts of damage to crops Body Form Three tagmata Head Two compound eyes Three unpaired ocelli One pair of antennae Preoral cavity with complex mouthparts Thorax Three pairs of legs Forelegs Middle legs Hind legs Thorax cont… Two pairs of wings in most adults, while juveniles have wing pads Some only have one pair Some lack altogether Some only possess wings during reproductive periods Abdomen Typically 9 – 11 segments Female gonopore on segment 8 Male gonopore on segment 9 Some have sensory appendages called cerci on last segment Wings and Flight Wings composed of two sheets of epidermis covered with cuticle Sclerotized areas called veins offer support Hollow House tracheae, nerves, and blood Permanently outstretched wings is a primitive trait The evolution of wing folding has allowed insects to exploit microhabitats where outstretched wings would be a handicap Flight muscles fill most of the thorax Striated fibers with huge mitochondria, and profuse tracheal innervation (to meet high oxygen demand) Wings move up, down, forwards, and backwards Wing angles are frequently changed Insects are probably the most skilled flyers Houseflies can outperform any bird Can turn in the distance of one body length Can fly upside down Hummingbird moths and botflies can fly 25 mph Display positive phototaxis, and often fly into lamps Nutrition Diverse nutritional strategies, and thus diversity in mouthpart structure and function Mandibulate mouthparts Grasshoppers and ants Large jaws for cutting and chewing Found in many herbivores and carnivores Sucking mouthparts Butterflies and moths Suck nectar and fruit juices with a long proboscis formed from maxillae Rolled and coiled when not feeding Diet is N and Na deficient so they drink other types of liquids (mud puddles, carrion, animal wastes, etc.) to supplement Piercing mouthparts Aphids drink plant juices Female mosquitoes require a blood meal to produce eggs Stylet pierces prey Has a salivary channel and a food channel Proteins in saliva produce itching associated with mosquito bites Surrounding structures support stylet Sponging mouthparts Nonbiting flies Enlarged labium is used to absorb liquids Liquefy food with enzymes Digestive system Salivary glands produce many types of secretions Pectinase for breaking down plant cell walls Typical secretions Some have silk glands in place of salivary glands Used for making cocoons Digestive system cont… Peritrophic membrane Protects non-cuticularized midgut Partitions areas where digestive enzymes function Membrane is perforated, so some enzymes pass through membrane and act on food bolus Hydrolyzed biomolecules pass from bolus side to lumen side Additional enzymes facilitate further digestion Midgut has two to six gastric ceca Hemocoel contains a fat body that synthesizes and stores biomolecules Hemal System Not many vessels Accessory hearts pump blood to important structures (antennae, mouthparts, legs, wings, cerci, etc.) Located at bases of extremities Many insects can survive subfreezing temperatures (-30°C) Compounds in blood, such as glycerol, function as antifreeze Respiratory System Elaborate tracheal system Branch and diverge into a network of smaller and smaller tubes Smallest tubes are called tracheoles (make direct contact with cells) 10 pairs of spiracles Two pairs on thorax Spiracles can be closed, which prevents: Water loss Dust from entering Parasites from entering Cartilagenous support rings Sensory Organs Most are derived from setae and are associated with appendages Tarsi of many have gustatory (taste) or tactile (touch) receptors Flies taste food by walking on it Eyes Form color images in some Some (especially pollinators) are sensitive to UV light Reproductive System Gonochoric with internal fertilization Female system Two ovaries produce eggs with a waterproof capsule Two oviducts converge and empty into a vagina Oviduct opens on segment 8 Tubular ovipositor for depositing eggs in appropriate substratum (i.e. soil, wood, plant, animal, etc.) Ovipositor is modified to form a sting with poison glands in some Hymenopteran females Male system Two testes with sperm ducts Accessory glands Median ejaculatory duct Penis is extensible / retractable Everts through gonopore on segment 9 Releases spermatophore or sperm Sperm plug (mating plug) May have claspers for holding female during copulation Sperm enter egg through a micropyle in egg capsule Insect Development Under hormonal control Three main stages: Egg Juvenile Many instars separated by molts Main role is to feed and grow Adult Technically termed an imago Sexually mature with functional wings May / may not continue to molt Main role is dispersal and reproduction Direct Development Three main types Paurometabolous development is the only type we will discuss Grasshopper, cockroaches, etc. Juveniles are morphologically similar to adults Adults don’t molt Does not promote resource partitioning Adults and nymphs in competition with one another Indirect Development Holometabolus development Hymenopterans, flies, beetles, and Lepidopterans Juveniles (larvae) are wormlike, and called grubs, maggots, or caterpillars Many larval instars Pupa phase between last instar and imago Undergoes metamorphosis Imago has wings and looks nothing like juvenile Different mouthparts, and therefore they feed on different types of foods Promotes resource partitioning Coevolution Definition - evolution of plants and insects in response to one another Plants and insects are vital to each other Plants provide food (nectar, pollen, and plant tissues) and shelter to insects Insects pollinate plants Bees, wasps, butterflies and moths are the primary insect pollinators Insects pollinate 70% of all flowering plants Plants adapt to make improvements that minimize costs to both organisms, and maximize efficiency of pollination process Characteristic colors, odors, and nectars to attract insects Guide marks and UV-sensitive patterns on flowers Time that flowers bloom coincides with periods of insect activity Coevolution in response to herbivory Plants have evolved a suite of defenses against insect herbivory Mechanical – spines, bristles, and hairs Chemical – nicotine, caffeine, cyanide, and tannins In turn, insects develop counter offenses Enzymes that detoxify protected tissues These insects specialize on plants they can detoxify, and have little competition Defensive chemicals may become attractants Arms race continues Galls Another interesting aspect of insectplant relationships is gall formation Many herbivorous insects cause plants to form galls Swollen portion of plant tissue containing eggs or larvae Forms in response to compounds secreted by ovipositing female and larvae Gall shape is characteristic of the insect that laid eggs Gall protects larvae, larvae feed on gall tissue; eventually emerge Parasitism Fleas, lice and biting flies are parasitic Usually only one life phase is parasitic Human botfly (Dermatobia) Captures a mosquito or biting fly Deposits eggs on the captured fly and releases Recently captured insect bites a human or other animal Egg falls into wound Grub develops in a cavity called a warble Molts several times Crawls to surface, falls to ground and pupates Parasitoidism Insects that parasitize larger insects Typically deposit eggs in or on host using an ovipositor Eggs hatch and consume host Very common It has been hypothesized that each insect species has a parasitoid Parasitoids are important for controlling insect populations Example 1: Wasp secretes chemical that causes worker ants to fight each other While ants fight, wasp enters nest and oviposits on larvae Example 2: Apanteles (wasp) oviposits in Manduca (Sphinx moth) larvae Up to 500 larvae can grow inside caterpillar Rupture body wall and form cocoon on surface Communication Pheromones Chemicals used for: Attracting potential mates Marking territories and trails Locating and removing dead individuals from colony Defense Warning conspecifics of danger Locating food Usually dispersed by wind Effective in very small quantities Flashing Fireflies Species-specific flashing pattern Some female fireflies mimic flashing patterns of other species Attracts male of a different species Female preys upon male Sound Stridulation is produced by wings and leg files Other types of sound produced by wing beating, forcing air from tracheae, etc. Social Insects Isopterans (termites) and Hymenopterans (bees, wasps, and ants) All groups share several characteristics One queen that mates with other fertile individuals (drones) Physical labor is carried out by sterile individuals Caring for juveniles Foraging Soldiers No individual can exist independent of colony No individual can be accepted into a colony other than its own However, some ants raid nests of other species, kidnap pupae, raise juveniles, and force them to work as slaves Characteristics… Caste system Individuals with different morphologies Have different duties Examples: soldiers, workers, and drones Ants and termites usually inhabit elaborate colonies with many tunnels, chambers, entrances and exits Army ants are an exception Leaf cutter ants have chambers for growing fungus Colony formation In bees, new queens are produced when larvae eat a substance known as “royal jelly” that is produced by the hypopharyngeal glands of nursery workers Queen usually takes part in nuptial flight in order to start new colony In bees 20,000 to 60,000 individuals will swarm and search for a new colony location Diversity There are around 30 insect orders We will briefly cover 14 Order Odonata Dragonflies and damselflies Voracious predators found near water Nymphs are aquatic Dragonflies are heavy bodied with outstretched wings Damselflies are more slender and delicate, with wings held over abdomen while at rest Order Orthoptera Grasshoppers, crickets, and katydids Large insects with large compound eyes Large hind legs for jumping Stridulating and auditory organs Herbivores Order Phasmida Walking sticks Mimic sticks or vegetation Elongate and cylindrical bodies Usually lack wings Some up to 30cm long Order Dermaptera Earwigs Nocturnal and omnivorous Elongate body Large pincer-like cerci Order Isoptera Termites Soft bodied and pale in color Build large mounds or nests in trees Can infest buildings Have endosymbiotic microbes that produce cellulase Order Mantodea Mantids Large and slow moving Cryptic coloration Ambush predators with raptorial forelegs Large eyes on a freely movable head Order Blattaria Cockroaches Fast runners Nocturnal Omnivorous Endosymbionts similar to those of termites Order Phthiraptera Lice Ectoparasites of mammals and birds Prehensile legs for clinging to hair or feathers Wingless All life stages are spent on host Host to host physical contact is required for transmission Order Hemiptera True bugs Stink bugs, water striders, aphids, and cicadas Sucking mouthparts for ingestion of liquids Herbivory is of economic significance Some are vectors of plant or animal disease Order Coleoptera Beetles Largest order at approximately 600,000 species Extremely sclerotized Forewings are called elytra Most are herbivores, but ladybird beetles are carnivorous Order Hymenoptera Ants, wasps and bees Chewing mouthparts Most have wings, but worker ants lack In females, ovipositor may be a sting Maggot-like larvae Order Lepidoptera Butterflies and moths Adults have a coiled proboscis Drink nectar and fruit juices Larvae are herbivorous caterpillars Cocoon or chrysalis is constructed for pupa phase Order Siphonaptera Fleas Small and wingless Laterally compressed Ectoparasites of mammals and birds Excellent jumpers Larvae aren’t parasitic Some are vectors of disease like bubonic plague Order Diptera True flies Mosquitoes, houseflies, gnats, and craneflies Large compound eyes Feed on animal or plant fluids Some are vectors of disease Eggs deposited in wet substrata such as flesh, fruit, or water Larvae called maggots