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The Segmented Worms Annelids and Allied Taxa • Phylum Annelida – Class Polychaeta – Class Oligochaeta – Class Hirudinea • Phylum Echiura • Phylum Sipuncula Phylum Annelida • Annelids are protostome coelomates in superphylum Lophotrochozoa. – Spiral, determinate cleavage. • Nervous system more centralized & circulatory system more complex than in previous phyla. Phylum Annelida • Annelids are segmented worms. • They have bodies composed of a series of fused rings. • Earthworms, leeches, clam worms. Phylum Annelida • The evolutionary innovation shown by annelids is segmentation (metamerism). – Segmentation evolved separately in annelids, arthropods, and chordates. • The body is divided into a series of segments, each having similar components of all major organ systems. – Built in fail-safe. – Allows for specialization. Phylum Annelida • Many annelids have chitinous bristles called setae. – Help in locomotion – Anchor worm in place – Deter predators Phylum Annelida – Body Plan • Prostomium – anterior part followed by segmented body. • Pygidium – terminal portion. Phylum Annelida – Body Plan • Peritonia (layers of mesodermal epithelium) of adjacent segments meet to form septa. • Fluid-filled coelom acts as a hydrostatic skeleton. Phylum Annelida • Tissues and organs differentiated • Segmentation highly developed – Mesodermal “blocks,” each with own coelom – Septa between segments Polychaetes • Members of class Polychaeta – Possess paddlelike parapodia that function as gills and aid in locomotion Parapodia Polychaetes • Resembling giant lipsticks, tubeworms (Riftia pachyptila) live over a mile deep on the Pacific Ocean floor near hydrothermal vents. • They may grow to about 3 meters (8 ft) long. • The worms’ white tube home is made of a tough, natural material called chitin (pronounced “kitein”). Class Polychaeta • Polychaetes have some features other annelids do not: – A well developed head. – Paired appendages, parapodia, that function as gills and aid in locomotion. – No clitellum. • Many setae Class Polychaeta • Polychaetes are mostly marine and mostly benthic. – May live under rocks, burrow into sediment, or build their own tubes. – Some are planktonic. Class Polychaeta • Sedentary and errant (freemoving) forms. • Sedentary forms often have elaborate devices for feeding and respiration. – Filter or deposit feeders. Class Polychaeta • Errant forms include pelagic and benthic types and are often predators or scavengers. Class Polychaeta Reproduction • • • • Gonads are temporary structures in polychaetes. Sexes usually separate. Fertilization is external. Early larva is a trochophore. Circulation and Respiration • Most have parapodia and gills for gaseous exchange. – Others use the body surface. • Circulation varies. – In Nereis a dorsal vessel carries blood forward and a ventral vessel carries blood posteriorly. - Blood flows across between these major vessels in networks around the parapodia and intestine. • In some, septa are incomplete and coelomic fluid serves circulatory function. • Many polychaetes have respiratory pigments Hemoglobin, chlorocruorin or hemerythrin. Excretion • Excretory organs vary, from protonephridia to metanephridia, and mixed forms. • One pair per metamere. • Inner end (nephrostome) opens into the coelomic cavity. • Coelomic fluid enters the nephrostome. • Selective resorption occurs along the nephridial duct. Nervous System and Sense Organs • Double ventral nerve cord runs length of the worm with ganglia in each metamere. • Sense organs include: – Eyes, nuchal organs and statocysts. – Eyes vary from simple eyespots to well-developed image-resolving eyes similar to mollusc eyes. – Nuchal organs are ciliated sensory pits that are probably chemoreceptive. – Some burrowing and tube-building polychaetes use statocysts to orient their body. Clade Siboglinidae (Pogonophorans) • Formerly members of phylum Pogonophora (beardworms). • Discovered in 1900. • 150 species described. • Most are small, less than 1 mm in diameter. • Giant beardworms that live in deepwater hydrothermal vents are 3 m long and 5 cm in diameter. Clade Siboglinidae (Pogonophorans) • Most live in mud on ocean floor at depths of 100 to 10,000 m. • Sessile animals that secrete and live in long chitinous tubes. • Tubes have general upright orientation in bottom sediments. • Tubes are generally three or four times the length of the animal. Clade Siboglinidae (Pogonophorans) • Long cylindrical body covered with cuticle. • Divided into a short anterior forepart, a long slender trunk, and a small, segmented opisthosoma. • Tentacles are hollow extensions of the coelom and bear minute pinnules. Clade Siboglinidae (Pogonophorans) • No mouth or digestive tract. • Nutrients such as glucose and amino acids absorbed from seawater through pinnules and microvilli of tentacles. Clade Siboglinidae (Pogonophorans) • Most energy derived from a mutualistic relationship with chemoautrophic bacteria that oxidizes hydrogen sulfide. – Trophosome, derived embryonically from midgut, houses the bacteria. Clade Siboglinidae (Pogonophorans) • Sexes are separate. • Research suggests that cleavage is unequal and atypical. – Appears to be spiral. • Coelom formed by schizocoely. • Embryo – Worm-shaped and ciliated. – Poor swimmer. – Probably carried by water currents until it settles. Class Oligochaeta • Earthworms are the most familiar oligochaetes, found in moist, rich soil. – They can burrow deep underground and remain dormant in a slime chamber during dry weather. – Setae help prevent slipping while burrowing. Oligochaetes • The largest earthworm ever found was in South Africa and was 22 feet long! The Earthworm Giant Earthworm – Amazon Basin Class Oligochaeta - Feeding – Food is stored in a thin-walled crop. – Muscular gizzard grinds food into small pieces. – Digestion and absorption occur in intestine. Class Oligochaeta - Excretion • Each somite, except the 1st three and terminal one, have a pair of metanephridia. • A ciliated funnel, the nephrostome, draws in wastes and leads through the septum. • These coil until the nephridial duct ends at a bladder that empties outside at nephridiopore. • Wastes from both the coelom and the blood capillary beds are discharged. • Aquatic oligochaetes excrete toxic ammonia. Class Oligochaeta - Circulation and Respiration • Coelomic fluid and blood transport food, wastes, and respiratory gases. • Blood circulates in a closed system with five main trunks running lengthwise in the body. • Dorsal vessel contains valves and functions as a true heart. – Pumps blood anteriorly into 5 pairs of aortic arches. – Aortic arches ensure steady pressure in ventral vessel. Class Oligochaeta - Nervous System and Sense Organs • Central nervous system and peripheral nerves. • Pair of cerebral ganglia connect around the pharynx to the ganglia of the ventral nerve cord. • Neurosecretory cells in brain and ganglia secrete neurohormones. – Regulate reproduction, secondary sex characteristics, and regeneration. • Lack eyes but have many photoreceptors in the epidermis. • Free nerve endings in tegument are probably tactile structures. Class Oligochaeta - Reproduction • Earthworms are hermaphroditic – male and female organs in the same animal. • When mating, two worms are held together by mucus secreted by the clitellum. Class Oligochaeta - Reproduction • After mating, a cocoon forms around the clitellum, as it passes forward it gathers both gametes, and fertilization occurs inside. Class Oligochaeta - Reproduction • Development occurs inside the cocoon and young worms hatch out. – Development is direct, no larval stage. Class Oligochaeta - General Behavior • Avoid bright light (negative phototaxis). • Chemical stimuli are important in locating food. • Limited learning ability - primarily trialand-error learning. 3 ecological groups of earthworms Endogeic example: Aporrectodea caliginosa (angle worms) Epigeic example: Lumbricus rubellus (leaf worms) Anecic example: Lumbricus terrestris (nightcrawlers) Earthworm Cross Section Dorsal Blood Vessel Intestine Coelom Nephridia Epidermis Muscle Layer Ventral Nerve Cord Ventral Blood Vessel Class Hirudinea (leeches) • Most inhabit fresh water, or moist terrestrial environments • Feed on small invertebrates or are parasitic • Have significant medicinal value in treating bruising, and as a source of Pharmaceuticals Hirudinea Class Hirudinea • Many leeches live as carnivores on small invertebrates. • Some are temporary parasites. • Some are permanent parasites – they never leave their host. • Can be a benefit in medicine. Class Hirudinea • Leeches are hermaphroditic and have a clitellum (only appears during breeding season), like oligochaetes. • Leeches do not have setae. • They’ve developed suckers for attachment and a specialized gut for storing large amounts of blood. Class Hirudinea - Respiration and Excretion • Some fish leeches have gills. • All other leeches exchange gases across epidermis. • 10 to 17 pairs of nephridia. • Coelomocytes and other special cells may assist in excretion. Class Hirudinea - Nervous and Sensory Systems • Two “brains” – Anterior fused ganglia form a ring around the pharynx. – Seven pairs of posterior fused ganglia. • 21 pairs of segmental ganglia in between along a double nerve cord. • Epidermis contains free sensory nerve endings and photoreceptor cells. • Pigment-cup ocelli are present. Class Hirudinea - Circulation • Coelom reduced by invasion of connective tissue. – Forms system of coelomic sinuses and channels. • Some have a typical oligochaete circulatory system. – Coelomic system is auxiliary. • Some lack blood vessels and coelomic sinuses serve as only vascular system. Phylum Echiura • Approximately 140 species of marine worms that burrow into mud or sand. • Live in empty snail shells or sand-dollar tests, or rocky crevices. • Found in all oceans. • Length varies from a few millimeters to 40 or 50 cm. Phylum Echiura – Form and Function • Sausage-shaped. • Inextensible proboscis anterior to the mouth. • Often called spoon worms”. • Simple nervous system with a ventral nerve running length of the body. • Ciliated groove on the proboscis allows them to gather detritus over the mud while lying buried. • Muscular body wall is covered by a cuticle and epidermis which may be smooth or covered by papillae. Phylum Echiura • Large coelom. • Digestive tract long and coiled. • Pair of anal sacs may serve an excretory and osmoregulatory function. • Most have a closed circulatory system with colorless blood. – Hemoglobin found in certain cells and in coelomic corpuscles. • Respiration probably occurs in hindgut which is continually filled and emptied by cloacal irrigation. Phylum Echiura - Reproduction • Sexes are separate. • Gonads produced by special regions in peritoneum in each sex. • Fertilization usually external. • Early cleavage and trochophore stages similar to annelids.