Animal Diversity 7th Edition Hickman Roberts

Keen Larson Eisenhour 0073524255

9780073524252
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CHAPTER 11 ANNELIDS AND ALLIED TAXA

CHAPTER OUTLINE
11.1 Dividing the Body
A. A spacious, fluid-filled coelom provides an efficient hydrostatic skeleton.
B. When septa divided this coelom into a series of compartments, metamerism first arose.
C. Metamerism may have arisen independently in the deuterostome line, which includes the vertebrates.
D. If one segment should fail, another could still function.
E. Metamerism makes independent segmental movements possible.
11.2 Characteristics
A. General
1. Annelids and its allies are eucoelomate protostomes within the subgroup Lophotrochozoa
2. They develop by spiral mosaic cleavage, form a coelom by schizocoely, and have a trochophore
larva.
3. The three phyla discussed are Annelida (marine worms, leeches, earthworms), Echiura, and
Sipuncula (both benthic marine animals without segmented bodies)
5. Phylogenetic studies place echiurans within Annelida as a derived group of polycheates where
segmentation has been lost.
6. Sipuncula a sister taxon to Annelida (Figure 11.1).
B. Annelids
1. About 15,000 species of segmented worms are classified in the phylum Annelida.
2. About two-thirds are more obscure marine worms.
3. The nervous system is more centralized and the circulatory system is more complex than in other
worms.
4. Except for leeches, annelids have tiny chitinous bristles called setae.
5. Short setae anchor a segment in an earthworm so it prevents slipping backward.
6. Long setae help aquatic worms swim.
C. Ecology Relationships
1. Annelids occur in the sea, freshwater, and on land.
2. Marine annelids borrow in the mud or live in tubes.
3. Some feed on organic matter in the mud, some are suspension feeders, many are predators.
4. Freshwater annelids burrow in the mud, live among vegetation, or swim freely.

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 5. Leeches are bloodsuckers or are carnivorous.
D. Economic Importance:
1. Many annelids are grazers and consumers of detritus in food chains; many are preyed upon by
fish.
2. Burrowing on land or in oceanic mud and sand, annelids are important for drainage, aeration,
mixing of soil and distribution of organic matter .
3. Blood-sucking leeches are used medicinally.
11.3 Body Plan
A. Body Wall
1. A two part head has an anterior tip containing the prostomium and peristomium.
2. The terminal portion bearing the anus is the pygidium (Figure 11.2).
3. The prostomium and pygidium are not considered metameres like the other segments
4. New metameres form just in front of the pygidium; thus the newest segments are at the posterior.
5. The coelom develops embryonically as a split in mesoderm on each side of the gut (schizocoel).
6. Peritoneum (mesodermal epithelium) lines the body wall and forms dorsal and ventral
mesenteries (Figure 11.3).
7. Peritonea of adjacent segments meet to form the septa.
8. Except in leeches, the coelom is filled with fluid and serves as a hydrostatic skeleton.
a. The fluid volume remains constant.
b. Therefore contraction of longitudinal muscles causes the body to shorten and fatten.
c. Contraction of circular muscles causes the body to narrow and lengthen.
d. By separating this force into sections, widening and elongation move the whole animal.
e. Alternate waves of contraction, or peristalsis, allow efficient burrowing.
f. Swimming annelids use undulatory movements.
9. The surface is covered with an epidermis and a thin outer layer of non-chitinous cuticle (Figure
11.3).
10. Paired epidermal setae are ancestral for annelids; some are reduced or altogether lost (Figure
11.2D).
11. The gut is not segmented and longitudinal blood vessels extend through the septa. (Figure 11.3)
12. Annelids were originally divided into three classes: Polychaeta, Oligochaeta, Hirudinea.
13. Pogonophorans and euchiurans are within class Polychaeta due to molecular analysis.
14. Polychaeta is a paraphyletic class and recently separated; ancestors to oligochaetes and
hirudineans arose from within the polychaetes.
15. Oligocheates and leeches form the monophyletic group Clitellata. (Figure 11.1)
16. The clitellum is a reproductive structure present in leeches only during the reproductive season.
C. Characteristics of Phylum Annelida (see Inset).
11.4 Polychaetes
A. Characteristics
1. Polychaets is the largest groups of annelids with more than 10,000 species, mostly marine. They
have been separated into two classes: Errantia and Sedentaria (this also includes other annelids)
(Figure 11.4)
2. They vary from 1 mm to 3 meters long; most are 5 to 10 cm long.
3. Some live in crevices; others inhabit tubes or are pelagic.
4. Polychaetes are an important part of marine food chains.
5. Polychaetes have a well-differentiated head with sense organs.
7. Paired appendages called parapodia are on most segments (Figure 11.2 and p. 224).
8. They have no clitellum.
9. Many setae are arranged in bundles on the parapodia.
10. A head or prostomium may or may not be retractile; it often bears eyes, tentacles and sensory
palps. (Figure 11.2)
a. The first segment or peristomium surrounds the mouth and may have setae, palps or
chitinous jaws. (Figure 11.4)
b. Suspension feeders may bear a tentacular crown that opens like a fan but can be withdrawn in
to the tube.(Figure 11.4)
11. Most segments of the trunk bear parapodia with lobes, cirri, setae and other parts
(Figure 11.2D).

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 a. Parapodia have two lobes, the dorsal notopodium and the ventral neuropodium.
b. Usually the parapodia are the chief respiratory organ although the worm may also possess
gills.
c. Parapodia help crawl, swim, and anchor the worm in a tube.
d. Along with gills, they serve is chief respiratory organs.
e. Like some, Amphritite has plumelike gills (Figure 11.5).
12. Sense organs include eyes and statocysts that are more developed than oligochaetes.
a. Eyes vary from simple eyespots to well-developed image-resolving eyes.
13. In contrast to clitellates, polychaetes have no permanent sex organs and have separate sexes.
a. Gonads appear as simple temporary swellings of the peritoneum.
b. Gametes are shed into the coelom and exit by gonoducts, metanephridia or rupturing of the
body.
c. Fertilization is external and the early larva is a trochophore
d. Most of the year polycheates live as sexually unripe worms called atokes; when swollen with
gametes they are called epitokes. During the swarming period, which occurs at the
beginning of the last quarter of the October-November moon, these epitokes break off and
swim to the surface; swarming provides for synchronous maturation of all epitokes (Figure
11.9)
11.5 Class Errantia (mobile polychaetes)
a. The clamworm Nereis is an example of a predatory polychaete with jaws. (Figure 11.2)
b. Scale Worms (Figure 11.6) are carnivores and some are commensals living in burrows of
other organisms. (Figure 11.7)
11.6 Class Sedentaria (sedentary polychates, oligochaetes, and leeches).
a. Amphitrite traps particles with extensible tentacles. (Figure 11.5)
b. Featherduster worms or fan worms live in tubes but unfurl tentacular crowns to feed.
c. Fanworms are polychaete ciliary feeders, directing small food balls along grooved
radioles to the mouth by ciliary action (Figure 11.8).
f. As the net fills with food, the food cup rolls it into a ball, which is passed into a groove
toward the mouth.
g. Arenicola, the lugworm, lives in an L-shaped burrow in intertidal mudflats and ingests food-
laden sand (Figure 11.11)
h. Tubes may be parchmentlike, attached to rocks, and composed of sand, shell, or seaweed.
(Figure 11.4A)
11.7 Class Sedentaria, Family Siboglinidae (Pogonophorans)
A. General Description (Figure 11.12)
1. Pogonophorans or beardworms have been found from Indonesia coasts to the Atlantic Ocean.
2. Some 150 species have been described from marine environments.
3. Molecular and morphological analysis indicates they are derived from polycheates.
4. Segmentation is present in only one body region.
5. They are sessile, bottom living animals found at depths of more than 200 m.
6. Body characteristics:
a. Most are 5 to85 cm in length with a diameter of less than a millimeter
b. They have a forepart, trunk, and opisthosoma. (Figure 11.12)
c. A cuticle, setae, coelomic compartments, and tentacles are present.
d. A mouth and digestive tract are absent.
7. Soboglinids absorb nutrients from seawater through pinnules and tentacles.
8. They form a mutual association with chemoautotrophic bacteria that oxidize hydrogen sulfide.
9. The trophosome of the midgut bears bacteria.
10. Sexes are separate.
11.8 Class Sedentaria, FamilyEchiuridae
1. Consists of marine worms that burrow in mud or sand, or live in empty snail shells.
2. Mostly littoral; vary in length from a few millimeters to 40 to 50 centimeters.
3. Echiurans are cylindrical and sausage-shaped.
4. Anterior to the mouth is a flattened, extensible proboscis for deposit feeding.
5. Often called “spoonworms (Figure 11.13).”
6. While buried the animal can extend its proboscis out for exploration and deposit feeding (Figure 11.14).

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 7. Urechis often has commensals living with it in its burrow, such as crabs, fish, clams, and polychaete
annelids.
8. They have a closed circulatory system.
9. Nephridia are present; all have a nerve ring and ventral nerve cord.
10. A pair of anal sacs arises from the rectum and opens into the coelom.
11. A trochophore larva is present similar to those of annelids and sipunculans.
12. Sexual dimorphism is pronounced, with females being much larger than males.
11.9 Class Sedentaria, Order Clitellata: (Oligochaetes and Leeches)
A. Oligochaetes 1. Over 3000 species occur in habitats from soil to fresh water; a few are marine or
parasitic.
2. Nearly all bear setae; although highly varied, setae are fewer than in polychaetes.
3. Earthworms burrow in moist, rich soil; they emerge at night.
4. In wet weather they stay near the surface; in dry weather they burrow deep and become
dormant
5. Lumbricus terrestris is commonly studied in school laboratories. (Figure 11.16)
6. Darwin studied earthworms and estimated 10-18 tons [9-16 metric tonnes] of dry earth passed
through earthworm intestines per acre [0.4 hectares] annually.
7. Earthworms have an important role in churning the soil, mixing materials and adding
nutrients.
8. Giant tropical earthworms may reach 4 meters long and live in interconnected tunnels.
B. Form and Function
1. The prostomium overhangs the mouth at the anterior end (Figures 11.16B).
2. In most earthworms, each segment bears four pairs of chitinous setae; some may bear
over 100 (Figure 11.16C).
3. Each seta is a bristle-like rod set in a sac and moved by tiny muscles. (Figure 11.17)
4. Setae anchor segments during burrowing.
5. Earthworms move by peristalsis.
a) Circular muscles contract, the anterior end lengthens, setae anchor the forward end.
b) Longitudinal muscles contract, body shortens, posterior end is pulled forward.
6. Most oligochaetes are scavengers, feeding on decayed organic matter, leaves, refuse, etc.
a) Food is moistened by the mouth and drawn in by a sucking action of the muscular
pharynx. (Figure 11.16A)
b) Digestion and absorption occur in the intestine; an infolded typhlosole increases
surface
area.
c). Chloragogen tissue surrounds the intestine and synthesizes glycogen and fat; cells full of
fat float free in the coelom.
d). Chloragogen cells also function in excretion.
7. Both coelomic fluid and blood carry food, wastes and respiratory gases.
8. Blood circulates in a closed system with five main trunks running lengthwise in the body.
9. The dorsal blood vessel above the alimentary canal has valves and functions as a true heart
(Figure 11.16A)
10. The dorsal vessel pumps blood anteriorly into five pairs of aortic arches. (Figure 11.13A)
11. Blood contains colorless ameboid cells and dissolved hemoglobin.
12. Earthworms have no special gaseous exchange organs; the moist skin handles all exchanges.
13. Each somite except the first three and last one have a pair of nephridia (Figure 11.18).
14. Each nephridium occupies parts of two adjacent segments.
15. A ciliated funnel, the nephrostome, draws in wastes and leads through the septum.
16. These coil until the nephridial duct ends at a bladder that empties outside at the nephridiopore.
17. Wastes from the coelom are discharged.
18. A pair of cerebral ganglia connects around the pharynx to the ganglia of the ventral nerve cord.
(Figure 11.16A)
19. Fused ganglia in each somite contain both sensory and motor fibers.
20. One or more giant axons are located in the ventral nerve cord to increase the rate of conduction
and stimulate contractions of muscles in many segments.
21. Earthworms are hermaphroditic

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 22. Earthworms mate at night during warm, moist weather.
23. They mate by aligning in different directions with ventral surfaces together. (Figure 11.19)
24. Mucus secreted by the clitellum holds them together.
25. Sperm travel to the seminal receptacles of the other worm along seminal grooves.
26. After sperm exchange, each worm secretes a mucous tube and chitinous band to form a cocoon.
(Figure 11.19)
27. As the cocoon passes forward, eggs, albumin and sperm pour into it.
28. Fertilization and embryogenesis takes place in the cocoon; young worms emerge.
11.10 Class Sedentaria, Order Clitellata, Family Hirudinidae: Leeches
A. Diversity
1. Most leeches live in fresh water but a few are marine or in moist terrestrial environments.
2. About 500 species, they are more common in the tropics than temperate zones.
3. Leeches vary in color: black, brown, red and olive green.
4. Most are flattened; they range in size from 2 cm to 20 cm or more (Figures 11.20).
B. Form and Function
1. True leeches usually have 34 segments.
2. There are no setae and no parapodia but typically have both an anterior and posterior sucker
(Figure 11.21).
3. Most leeches lack distinct coelomic compartments and septa have disappeared.
4. The coelomic cavity is filled with connective tissue.
5. Some Characteristics
a. Some are carnivores on small invertebrates; others are temporary or permanent parasites.
b. Leeches feed on prey with a protrusible proboscis or muscular pharynx armed with teeth.
a. Most are feed on tissue fluids and blood pumped from open wounds.
b. Bloodsuckers secrete an anticoagulant in their salvia
c. They depend mostly on bacteria in their gut for digestion of their blood meal.
f. Leeches are hermaphroditic and have a clitellum during the breeding season.
g. The clitellum secretes a cocoon for reception of eggs.
h. The cocoons are buried in mud or damp soil, and development is similar to that oligochaetes.
6. Leeches are being used again medically to relieve congestion until veins grow back in severed
digits.
11.9 Phylum Sipuncula
A. Characteristics
1. Consist of 250 or so species of benthic marine worms, mostly littoral (also called “peanut worms”).
2. Burrow in mud or sand, occupy borrowed snail shells, etc. (Figure 11.23).
3. Sipunculans are not metameric; no setae.
4. Head is in the form of an introvert—crowned by ciliated tentacles surrounding the mouth. (Figure
11.23)
5. Cerebral ganglion, nerve cord, and pair of nephridia present.
6. Coelomic fluid contains red blood cells with respiratory pigment called hemerythrin.
7. Larvae are trochophores.
8. They have separate sexes and in most gonads develop seasonally.
11.10 Phylogeny and Adaptive Diversification
A. Phylogeny
1. Molecular and morphological analysis supports pognophorans and echiurans are within the
phylum Annelida.
2. Echiuran larvae have some serially repeated structures that could re remnant structures from a
metameric ancestor.
3. Origin of segmentation
a) True metamerism is seen in annelids, arthropods, and chordates
b) Placement of annelids and arthropods makes it unlikely segmentation is homologous in the
three taxa.
c) The clade Lophotrochozoa and Ecdysozoa contain most phyla that are unsegmented; it is
unlikely members inherited a segmented body plan from a common ancestor.
d) Annelids and mollusks have similar trochophore development; but annelid larva develop in
segments
e) Possibly all bilateral symmetrical metazoans share a segmented ancestor.

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 f) Studied of how segments form does not support this.
g) The advantage of segmentation for annelids is in burrowing efficiency; it allows the body to
change shape; the same cannot be applied to arthropods.
4. Class Polychaeta is paraphyletic even with siboglinids. This class has recently been dissolved.
B. Adaptive Diversification
1. Septal arrangement with fluid-filled compartments has been varied for precise movements.
2. Powerful circular and longitudinal muscles shorten and lengthen the body.
3. Oligochaetes are constrained by the terrestrial soil environment.
4. Polychaetes inhabit a wide range of habitats and are not constrained by the physical environment
to a great degree.
5. Feeding adaptations vary widely, from chitinous jaws to specialized tentacles.
6. Leeches have developed both parasitic and predatory adaptations.
C. Classification
Class Errantia
Class Sedentaria
Order Clitellata
Family Hirudinidae

Lecture Enrichment
1. Annelids occasionally are central to native life and customs. The epitokes of the marine palolo worms rise to
the surface in huge numbers during their reproductive cycle and natives wade out to collect them in tubs and
hold feasts on this egg-like food.
2. The problem of the evolutionary origin and function of segmentation provides an excellent illustration of how a
concept that may seem to have several very “obvious” advantages remains a “just so” story until researchers can
find fossil evidence or devise an experimental regime to actually demonstrate its advantages. “Proving” an
evolutionary advantage requires perceptive research.
3. Ask students to consider why a segmented animal might not bud segments from the tail end, rather than
somewhere between the first and last segments.

Commentary
Background: Coastal students may have encountered clam worms, etc. in beachcombing, but the wide array of
polychaetes will require visual illustrations or specimens. Only the avid fishermen among students may have
directly experienced leeches; their testimony can generate student interest. Others may remember leech scenes from
movies such as “Stand By Me” or “African Queen.” Students who put earthworms on hooks when fishing may have
made one insight into earthworm anatomy; they should be aware of the fact that earthworms have some blood
vessels and blood.
Misconceptions: The urban legend that various hamburger chains use earthworms instead of beef is still extant;
note that it is both impractical and highly uneconomical to substitute this puny animal for such use. Some have been
led to believe that the “comeback” of leeches in medicine in some way admits that the earlier abandonment of
leeching was premature and a medical mistake. Note that the original theory (Galen’s “body humors”) remains
completely disproved and that modern use centers around a very narrow application well within the modern medical
paradigm: reattaching appendages. Even biology majors often mentally envision all oligochaetes as terrestrial
earthworms and may even “rescue” a poor “earthworm” found drowning in a freshwater stream; use of Tubifex and
other aquatic oligochaetes will help rectify this misconception. Again, students often assume that evolution only
adds complexity; leeches are an example where quite a few features are reduced or lost from ancestors. The use of
the term “brain” for fused ganglia and “heart” for pumping tubes with valves may lead students to assume more
complexity and efficiency to these structures than is justified.

ADVANCED CLASS QUESTIONS:

1. How does the development of segmentation or metamerism lead to specialization of parts?

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2. Animals are adapted to the manner they acquire food. How does this explain so much of polychaete,
oligochaete and leech diversity? What keeps members of these groups from evolving back into the other’s
adaptive “territory?”
3. Peripatus is a living organism that resembles an earthworm with stubby legs and it has been considered a “link”
between annelids and arthropods. Peripatus uses hypodermic insemination. Does this method of reproduction
logically fit with its classical position as a surviving member of the common ancestor?
4. Describe a scenario for the evolution of leeches from ancestral oligochaetes. Would the ancestor be terrestrial
or aquatic? What features would gradually be gained or lost? What chemosensors would be needed and what
digestive enzymes would be selected? How is this scenario supported or not supported by modern leech
anatomy, physiology, behavior and environmental occurrence?