segmented worms chapter 17. annelids and allied taxa phylum annelida class polychaeta class...

Segmented Worms

Chapter 17

Annelids and Allied Taxa

Phylum Annelida Class

Polychaeta Class

Oligochaeta Class Hirudinida

Phylum Echiura Phylum

Sipuncula

Annelids and Allied Taxa

Members of phyla Echiura and Sipuncula are benthic marine animals with unsegmented bodies.

Molecular sequence data place echiurans within phylum Annelida.

Echiurans - Sister taxon to Annelida.Sipunculans - Sister taxon to a clade

composed of Annelida and Echiura.

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

Annelids can be found worldwide in marine, freshwater, and terrestrial habitats.

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.

Phylogeny

Traditionally, annelids are divided among 3 classes: Class Polychaeta Class Oligochaeta Class Hirudinida

Polychaeta is a paraphyletic class because ancestors of the clitellates arose from within it.

Oligochaeta and Hirudinida form a monophyletic group called Clitellata. Characterized by reproductive structure called a clitellum.

Class Oligochaeta is a paraphyletic group because ancestors of leeches arose from within it.

Class Polychaeta

Polychaeta is the largest, most diverse class. May be

brightly colored, variable shape.

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 (free-moving) 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.

Representative Polychaetes

Clam Worms: Nereis Errant polychaetes Live in mucus-lined burrows

near low tide level. Come out of hiding places at

night to search for food. Prostomium bears a pair of

palps sensitive to touch and taste, a pair of short sensory tentacles, and two small dorsal eyes sensitive to light.

Peristomium has a ventral mouth, a pair of jaws, and four pairs of sensory tentacles.

Representative Polychaetes

Scale worms Flattened bodies

are covered with broad scales.

Some are large, all are carnivores and some are commensals in burrows of other organisms.

Representative Polychaetes

Fireworms Have hollow, brittle

setae that contain poisonous secretions.

Feed on cnidarians.

Representative Polychaetes

Tubeworms Tube-dwellers May line their

burrows with mucus Use cilia or mucus

to obtain food

Representative Polychaetes

Fanworms or Featherduster worms Unfurl tentacular

crowns to feed. Food moved from

radioles to mouth by ciliary action.

Representative Polychaetes

Parchment Worms

Lives in a U-shaped tube.

Modified segments pump water through tube.

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.

http://www.youtube.com/watch?v=2FFnrW_SUdM

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.

Osedax – bone eating worms

http://www.youtube.com/watch?v=URi8KccVkks

Clade Clitellata

Class Oligochaeta and Class Hirudinida Form reproductive structure called a clitellum.

Ring of secretory cells found in a band around the body. Permanent in oligochaetes but visible only during

reproductive season in leeches. Members are derived annelids that lack parapodia. Hermaphroditic (monoecious) animals that exhibit

direct development. Young develop inside a cocoon secreted by the

clitellum, and emerge as small worms.

Class Oligochaeta

Class Oligochaeta includes earthworms and many freshwater worms. They possess setae, but not as much as

polychaetes.

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.

Class Oligochaeta

Darwin wrote about earthworms in The Formation of Vegetable Mould Through the Action of Worms.He noted the beneficial activities of worms,

aeration, moving nutrients up from subsoil, adding nitrogenous products, breakdown of organic matter in dead leaves etc.

An earthworm can ingest its own weight in soil every 24 hours.

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 - 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 segment, 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 - General Behavior

Avoid bright light (negative phototaxis).Chemical stimuli are important in

locating food.Limited learning ability - primarily trial-

and-error learning.

Class Hirudinida

Class Hirudinida includes the leeches. Primarily

freshwater, a few marine & terrestrial.

More common in tropical climates.

Class Hirudinida

Many leeches live as carnivores on small invertebrates.

Some are temporary parasites. Some are permanent parasites – they never

leave their host.

Class Hirudinida

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 Hirudinida - 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 Hirudinida - 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 Hirudinida - 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.

Class Hirudinida

Leeches are highly sensitive to stimuli associated with the presence of prey.Those that feed on mammals are attracted

by warmth.

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.

Phylum Sipuncula

Approximately 250 species of benthic marine worms.

Sedentary, living in burrows of mud or sand, snail shells, coral crevices, or among vegetation.

More than ½ restricted to tropical zones.

Some are tiny, slender worms, but most range from 3 to 10 cm in length.

Some are known as “peanut worms” because when disturbed, they contract to a peanut shape.

Phylum Sipuncula - Form and Function

No segmentation or setae.Slender, retractable introvert or

proboscis at anterior end.Walls of the trunk are muscular.

Phylum Sipuncula - Nutrition

Some appear to be detritivores and others suspension feeders.

Some nutrition may come from dissolved organic matter in the surrounding water.

From burrow or hiding place, they extend tentacles to explore and feed.

Collected organic matter moved from mucus on tentacles to mouth by ciliary action.

Large fluid-filled coelom. Digestive tract is U-shaped.

Phylum Sipuncula - Respiration

Lack a circulatory and respiratory system.

Gas exchange appears to occur across the introvert and tentacles.

Phylum Sipuncula - Nervous and Sensory Systems

Bilobed cerebral ganglion behind tentacles.

Ventral cord extends the length of body.

Phylum Sipuncula - Reproduction

Sexes are separate.Sex organs develop seasonally within

the connective tissue covering the origins of the retractor muscles.

Sex cells are released through the nephridia.

Asexual reproduction occurs by transverse fission.

Phylogeny

Similarities in the early development of molluscs, annelids, and some primitive arthropods indicate that these three groups are probably closely related.Trochophore larvaSpiral cleavageSchizocoelous coelom formation

Evolutionary Significance of Metamerism

No satisfactory explanation for origins of metamerism and coelom has gained acceptance.

Coelom may have been advantageous as a hydrostatic skeleton.

Coelomic fluid would have acted as a circulatory fluid and reduce need for flame cells everywhere.

Coelom could store gametes for timed release. Would require nervous and endocrine control.

Unlikely that segmentation is homologous among annelids, arthropods, and chordates.

Current evidence supports the hypothesis that segmentation arose independently multiple times.

Evolutionary Significance of Metamerism

Selective advantage of a segmented body for annelids appears to lie in the efficiency of burrowing.

However, does not explain segmentation in arthropods given the rigidity of the exoskeleton.

Phylogeny

Molluscs and annelids share many developmental features so are presumed to be closely related.

However, shared features are likely to be a retained ancestral feature for lophotrochozoan protostomes.

Pogonophorans have been included within the annelids as derived polychaetes.

Molecular analyses place sipunculids and echiurans closely related to the annelids.

Phylogeny

With more studies, Echiura, like Pogonophora, may no longer be a valid phylum.

Placement of Sipuncula is contentious. Members are not metameric and lack setae. Larval development similar to annelids, molluscs,

and echiurans. Molecular data may clarify position within

Lophotrochozoa. Presently, depicted as the sister taxon to a clade of

annelids and echiurans. Polychaeta is a paraphyletic group.