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Trilobites, Chelicerates, and Myriapods Chapter 19 Phylum Arthropoda Two out of every three known species of animals are arthropods. Members of the phylum Arthropoda are found in nearly all habitats of the biosphere. Phylum Arthropoda Arthropods are: Multicellular Bilaterally symmetrical Triploblastic Have a true coelom (protostomes) Segmented General Characteristics of Arthropods The diversity and success of arthropods are largely related to their segmentation, hard exoskeleton (made of chitin), and jointed appendages. General Characteristics of Arthropods Segments have combined into functional groups called tagmata. Tagmata have specialized purposes. General Characteristics of Arthropods As arthropods evolved, the segments fused, and the appendages became more specialized. The appendages of some living arthropods are modified for many different functions. General Characteristics of Arthropods Arthropods have an open circulatory system in which fluid called hemolymph is circulated into the spaces surrounding the tissues and organs. A variety of organs specialized for gas exchange have evolved in arthropods. A Versatile Exoskeleton The exoskeleton of arthropods is very protective, but still flexible. The exoskeleton is made of chitin. Prevents desiccation. Provides places for muscle attachment. Does not allow for growth, the outer covering must be molted – ecdysis. More Efficient Locomotion Usually, each segment bears a pair of jointed appendages. The appendages have sensory hairs and may be modified for sensory functions, food handling, or walking & swimming. Air Piped Directly to Cells Most terrestrial arthropods have an efficient tracheal system of air tubes, which delivers oxygen directly to the tissues and cells. Limits body size. Aquatic arthropods breathe using internal or external gills. Highly Developed Sensory Organs Arthropods have a variety of sensory organs. Complex Behavior Patterns Arthropods show complex behavior patterns. Mostly innate behaviors. Some learned. Metamorphosis Intraspecific competition (between members of one species) is reduced because of metamorphosis. Larval forms may be quite different from adults. Relationships Among Arthropod Subgroups Clade Panarthropoda, Phylum Arthropoda Divided into subphyla based on relationships between subgroups based on molecular data. Relationships Among Arthropod Subgroups Centipedes, millipedes, pauropods, and symphylans are placed in subphylum Myriapoda. Insects are placed in subphylum Hexapoda. Spiders, ticks, horseshoe crabs and their relatives form subphylum Chelicerata. Lobsters, crabs, barnacles, and others form subphylum Crustacea. Relationships Among Arthropod Subgroups Formerly, insects and myriapods were placed together in uniramia. Uniramous appendages. The “mandibulate hypothesis” suggests all arthropods with mandibles are more closely related to each other than to arthropods with chelicerae. Mandibles in each group may or may not be homologous. mtDNA data support this hypothesis. Subphylum Trilobita Early arthropods, such as trilobites showed little variation from segment to segment. Subphylum Trilobita Trilobites arose during the Cambrian – maybe earlier and lasted for 300 million years. Subphylum Trilobita Trilobites had a trilobed shape. Three tagmata: Head (cephalon) with a mouth, compound eyes, antennae, and 4 pairs of leglike appendages. Trunk with a variable number of segments each with a pair of biramous appendages. One of the branches of biramous appendage was fringed and may have been a gill. Pygidium – segments fused into a plate. Subphylum Trilobita Most could roll up like pill bugs. Probably benthic scavengers. Many (especially later species) had large, complex, many-faceted eyes. Subphylum Chelicerata Chelicerate arthropods include eurypterids, horseshoe crabs, spiders, ticks, mites, scorpions, & sea spiders. Subphylum Chelicerata They have 6 pairs of cephalothoracic appendages: Chelicerae (mouthparts) Pedipalps 4 pairs of walking legs Lack mandibles and antennae. Class Merostomata Class Merostomata includes the eurypterids and horseshoe crabs. Eurypterids were giant water scorpions up to 3 m in length. Cambrian through Permian. Predators, some with large crushing claws. Class Merostomata Three genera of horseshoe crabs live today. Limulus, found in North America, has existed on earth almost unchanged since the Triassic period. Class Merostomata Horseshoe crabs have an unsegmented carapace (hard dorsal shield), a broad abdomen, and a long telson (tail piece). Cephalothorax Chelicerae Pedipalps 4 pairs walking legs Abdomen 6 pairs of thin appendages Book gills found on 5. Class Merostomata Horseshoe crabs have simple and compound eyes. Feed at night on worms and small molluscs. Come to shore in large numbers to mate at high tide. Trilobite larvae resemble trilobites. Class Pycnogonida Sea spiders, class Pycnogonida, have small, thin bodies and usually 4 pairs of walking legs. Found in all oceans, most common in polar seas. Some have chelicerae and pedipalps. Class Arachnida Class Arachnida includes spiders, scorpions, mites, and ticks. 50 µm Class Arachnida Two tagmata: Cephalothorax Chelicerae Pedipalps 4 pairs walking legs Abdomen Class Arachnida - Order Araneae Most spiders – order Araneae – have 8 simple eyes that can detect light and motion. Some hunting & jumping spiders may form images. Sensory setae detect air currents, web vibrations, and other stimuli. Spider’s vision usually poor, so awareness of environment depends largely on cuticular mechanoreceptors such as sensory setae. Class Arachnida - Order Araneae All are predaceous, mostly on insects. Many spin a web used for prey capture. Some chase & catch prey. Injected venom liquefies and digests the tissues which is sucked into spider’s stomach. Class Arachnida - Order Araneae Two or three pairs of spinnerets contain microscopic tubes that run to silk glands. Liquid scleroprotein secretion hardens as it is extruded from spinnerets. Silk threads are very strong and will stretch considerably before breaking. Silk is used for orb webs, lining burrows, forming egg sacs, and wrapping prey. Class Arachnida - Order Araneae Breathe by book lungs and/or tracheae. Book lungs unique to spiders - parallel air pockets extend into blood-filled chamber. Air enters chamber through a slit in body wall. Tracheae system is less extensive than in insects. Transports air directly to tissues. Tracheal systems of arthropods represent a case of evolutionary convergence. Class Arachnida - Order Araneae In spiders and insects, Malpighian tubules serve as excretory structures. Potassium, other solutes, and waste are secreted into tubules. Rectal glands reabsorb the potassium and water, leaving wastes and uric acid for excretion. Conserves water and allows the organisms to live in dry environments. Many spiders have coxal glands, modified nephridia, at the base of legs. Class Arachnida - Order Araneae Reproduction - before mating, male stores sperm in pedipalps. A courtship ritual is often required before the female will allow mating. Eggs may develop in a cocoon in the web or may be carried by female. Young hatch in about two weeks and may molt before leaving the egg cocoon. Class Arachnida - Order Araneae Are spiders really dangerous? Spiders are allies of humans in our battle with insects. American tarantulas rarely bite, and bite is not dangerous. Species of black widow spiders are dangerous. Venom is neurotoxic. Brown recluse spider has hemolytic venom that destroys tissue around the bite. Some Australian and South American spiders are the most dangerous and aggressive. Class Arachnida - Order Scorpiones Scorpions – order Scorpiones – feed on insects & spiders which they seize with their pedipalps. The last segment contains a bulbous base and a curved barb that injects venom. Scorpions are viviparous or ovoviviparous – females brood young within their reproductive tract. Class Arachnida - Order Opiliones Harvestmen – order Opiliones – differ from spiders in that the abdomen and cephalothorax are broadly joined rather than constricted. Only two eyes Abdomen shows segmentation Long legs end in tiny claws. Class Arachnida - Order Acari Mites and ticks – order Acari – have a fused cephalothorax & abdomen. Mites are tiny – 1mm or less. Some feed on plant juices and can be major pests. Several species of ticks carry diseases such as Lyme disease. Class Arachnida - Order Acari House dust mites free-living and often cause allergies. Spider mites - one of many important agricultural pest mites that suck out plant nutrients. Class Arachnida - Order Acari Hair follicle mite Demodex - harmless but other species cause mange in domestic animals. Human itch mite causes intense itching. Class Arachnida - Order Acari Tick species of Ixodes carry Lyme disease. Tick species of Dermacentor transmit Rocky Mountain spotted fever. Cattle tick transmits Texas cattle fever. Subphylum Myriapoda Subphylum Myriapoda includes these classes: Chilopoda (centipedes) Diplopoda (millipedes) Pauropoda (pauropods) Symphyla (symphylans) Use trachea to transport air. Excretion usually by Malpighian tubules. Class Chilopoda Centipedes – class Chilopoda – contain a few or many segments each (except the first behind the head and the last two) with a pair of jointed legs. Last pair of legs has a sensory function. Class Chilopoda Head appendages: One pair antennae One pair mandibles One or two pairs of maxillae. Dorsoventrally flattened. Class Chilopoda Sexes are separate. Some lay eggs (oviparous). Some have live young (viviparous). Young like little adults – no metamorphosis. Class Chilopoda Centipedes live in moist environments. They are carnivores, feeding on insects & worms. Prey is killed with poison claws on the first segment. Class Diplopoda Millipedes (Class Diplopoda) have two pairs of legs on each segment. Head has one pair each of antennae, mandibles, & maxillae. Body is more cylindrical. Class Diplopoda Millipedes live in dark, moist places – under rocks or logs. Most are herbivores, feeding on decayed plant matter or occasionally living plants. Slow moving, coil up when disturbed. Toxic or repellent fluids secreted when disturbed. Class Diplopoda After copulation, female lays eggs in a nest and guards them. Larvae have only one pair of legs per segment. Class Pauropoda Live in moist soil, leaf litter, decaying vegetation, or under bark and debris. Least well known of myriapods. Soft-bodied, small (2 mm or less). Approximately 500 species. Head lacks true eyes, has branched antennae, and a pair of sense organs. Class Pauropoda 12 trunks segments bear 9 pairs of legs but none on the first or last 2 segments. One tergal plate covers two segments. Lack tracheae, spiracles, and circulatory system. Probably most closely related to diplopods. Class Symphyla Live in humus, leaf mold, and debris. Male Scutigerella places a spermatophore at end of a stalk. Female stores the sperm in special pouches. Removes and smears eggs with sperm before attaching them to moss or lichen. Young hatch with only 6 or 7 pairs of legs. Class Symphyla Small (2–10 mm) with centipede-like bodies. Soft-bodied with 14 segments - 12 segments bear legs and one bears a pair of spinnerets. Antennae are long and unbranched. About 160 species are known. Eyeless with sensory pits at base of antennae. Phylogeny Relationships between subphyla are debated. Taxon of Pancrustacea, which includes hexapods and crustaceans, is well-supported. Phylogenies using molecular data rarely support grouping Myriapoda with Pancrustacea. There is support for placement of Myriapoda as the sister taxon for Cheliceratae. Phylogeny Biologists assume that the ancestral arthropod had a segmented body with one pair of legs per segment. Evolution caused adjacent segments to fuse and to make body regions. Hox gene studies indicate that the first five segments fused to form the head tagma in all four extant subphyla. In spiders, Hox gene studies indicate that the entire prosoma corresponds to the head of other arthropods. Phylogeny Genetic studies have been helpful in understanding the evolution of uniramous and biramous appendages. Molecular evidence repeatedly places hexapods with crustaceans even though hexapods have uniramous appendages and crustaceans have biramous appendages. Leads to the question: Did uniramous appendage development evolve more than once? Adaptive Diversification In contrast to annelids, arthropods have pronounced tagmatization by fusion of somites. Those with primitive characters have appendages on each somite. Derived forms are specialized. Modification of exoskeleton and appendages allowed variation in feeding and movement. Adaptations made possible by cuticular exoskeleton and small size fostered high diversity. Phylogeny Subphylum Trilobita Subphylum Chelicerata Class Merostomata Class Pycnogonida Class Arachnida Subphylum Myriapoda Class Diplopoda Class Chilopoda Class Pauropoda Class Symphyla Subphylum Crustacea Subphylum Hexapoda