Annelids and Allied Taxa: Phylum Annelida, Including Pogonophorans (Siboglinids) Phylum Echiura Phylum Sipuncula

C H A P T E R
17
Annelids and Allied Taxa
• PHYLUM ANNELIDA, INCLUDING
POGONOPHORANS (SIBOGLINIDS)
• PHYLUM ECHIURA
• PHYLUM SIPUNCULA
Annelida
Echiura
Sipuncula
Chloeia sp., a polychaete.
Dividing the Body

Although a fluid-filled coelom provided an efficient hydrostatic skele- The evolutionary advent of metamerism was significant because
ton for burrowing, precise control of body movements was probably it made possible much greater complexity in structure and function.
difficult for the earliest coelomates. The force of muscle contraction Metamerism increased burrowing efficiency by permitting the inde-
in one area was carried throughout the body by the fluid in the undi- pendent movement of separate segments. Fine control of movements
vided coelom. In contrast, there were distinct coelomic compartments allowed, in turn, the evolution of a more sophisticated nervous sys-
within the bodies of ancestral annelids. Compartments, known as tem. Moreover, repetition of body parts gave the organisms a built-
segments or metameres, were separated from neighbors by partitions in redundancy that provided a safety factor: if one segment should
called septa. Septa permitted each fluid-filled segment to respond fail, others could still function. Thus an injury to one part would not
individually to local muscle contraction—one segment could be long necessarily be fatal.
and thin and another short and round. Annelids illustrate segmenta- The evolutionary potential of the metameric body plan is amply
tion, or metamerism; their bodies are composed of serially repeated demonstrated by the large and diverse phyla Annelida, Arthropoda
units. Each unit contains components of most organ systems, such as and Chordata, which likely represent three separate evolutionary
circulatory, nervous, and excretory systems. origins of metamerism.
hic70049_ch17_362-383.indd 362 7/14/07 3:18:00 PM

www.mhhe.com/hickmanipz14e CHAPTER 17 Annelids and Allied Taka 363
T
he wormlike animal phyla described in this chapter are Pogonophora, or placed in distinct phyla: Pogonophora and
coelomate protostomes belonging to subgroup Lophotro- Vestimentifera. These deep-ocean worms belong in clade Sibo-
chozoa. They develop by spiral mosaic cleavage, form glinidae within class Polychaeta.
mesoderm from derivatives of the 4d cell, make a coelom by Worms in phylum Echiura and phylum Sipuncula are benthic
schizocoely, and share a trochophore as the ancestral larval form. marine animals with unsegmented bodies. Several phylogenetic
Three phyla are discussed: Annelida, Echiura, and Sipuncula. studies using molecular sequence data place echiurans within
Members of phylum Annelida are segmented worms liv- phylum Annelida as a derived group of polychaetes where seg-
ing in marine, freshwater, and moist terrestrial habitats. Marine mentation has been lost, but this placement is not universally
bristle worms, leeches, and the familiar earthworms belong to accepted. We depict echiurans as the sister taxon to Annelida,
this group. Annelida also now includes pogonophoran and ves- and sipunculans as the sister taxon to a clade composed of
timentiferan worms, formerly either placed together in phylum Annelida and Echiura (Figure 17.1).
Lophotrochozoa (in part)

Annelida
Clitellata
Polychaeta Oligochaeta Hirudinida Echiura Sipuncula
(in part) (in part)
Acanthobdellida Branchiobdellida Hirudinea
15 segments 34 segments Proboscis Anterior

in front of retractable
27 segments mouth introvert
Loss of setae
Anterior body sucker
Superficial annuli
Posterior body sucker
Reduced septal walls
Reduced number of
setae
Distinct, fixed
reproductive system
Clitellum
Direct development
Hermaphroditism
Loss of parapodia
Figure 17.1
Cladogram of annelids, showing the appearance of shared
derived characters that specify the five monophyletic groups
Annelid head (based on Brusca and Brusca, 1990). The Acanthobdellida and the
Parapodia Branchiobdellida are two small groups that Brusca and Brusca
place with the Hirudinea (“true” leeches), within a single taxon,
Metameric body
the Hirudinida. This clade has several synapomorphies: tendency
toward reduction of septal walls, the appearance of a posterior
sucker, and the subdivision of body segments by superficial annuli.
Note also that, according to this scheme, the Oligochaeta have
Paired epidermal setae
no defining synapomorphies; that is, they are defined solely by
retention of plesiomorphies (retained primitive characters, p. 205),
Coelom by schizocoely and thus might be paraphyletic.
Source: Modified from R. C. Brusca and G. J. Brusca, Invertebrates.
Sinauer Associates, Inc., Sinderland, MA, 1990.
hic70049_ch17_362-383.indd 363 7/30/07 9:23:06 AM

364 PART THREE Diversity of Animal Life
PHYLUM ANNELIDA, INCLUDING meet to form septa, which are perforated by the gut and longitu-
dinal blood vessels. The body wall surrounding the peritoneum
POGONOPHORANS (SIBOGLINIDS) and coelom contains strong circular and longitudinal muscles
Phylum Annelida (an-neli-da) (L. annelus, little ring, ida, pl. adapted for swimming, crawling, and burrowing (Figure 17.3).
suffix) consists of the segmented worms. It is a diverse phylum, Except in leeches, the coelom of most annelids is filled with
numbering approximately 15,000 species, the most familiar of fluid and serves as a hydrostatic skeleton. Because the volume
which are earthworms and freshwater worms (class Oligochaeta) of fluid in a coelomic compartment is essentially constant, con-
and leeches (class Hirudinida). However, approximately two- traction of the longitudinal body-wall muscles causes a segment
thirds of the phylum comprises marine worms (class Polychaeta), to shorten and to become larger in diameter, whereas contrac-
which are less familiar to most people. Some polychaetes are tion of the circular muscles causes it to lengthen and become
grotesque in appearance whereas others are graceful and beauti- thinner. The presence of septa means that widening and elon-
ful. They include clamworms, plumed worms, parchment worms, gation occur in restricted areas; crawling motions are produced
scaleworms, lugworms, and many others. by alternating waves of contraction by longitudinal and circular
Annelida are worms whose bodies are divided into similar muscles passing down the body (peristaltic contractions). Seg-
segments, (also called metameres) arranged in linear series and ments in which longitudinal muscles are contracted widen and
externally marked by circular rings called annuli (the name of the anchor themselves against the substrate while other segments,
phylum refers to this characteristic). Body segmentation (metam- in which circular muscles are contracted, elongate and stretch
erism) is a division of the body into a series of segments, each of forward. Forces powerful enough for rapid burrowing as well as
which contains similar components of all major organ systems. In locomotion can thus be generated. Swimming forms use undula-
annelids the segments are delimited internally by septa. tory rather then peristaltic movements in locomotion.
Annelids are sometimes called “bristle worms” because, with An annelid body has a thin outer layer of nonchitinous cuticle
the exception of leeches, most annelids bear tiny chitinous bris- surrounding the epidermis (Figure 17.3). Paired epidermal setae
tles called setae (L. seta, hair or bristle). Short needlelike setae (Figures 17.2 and 17.17) are ancestral for annelids, although they
help anchor segments during locomotion and long, hairlike setae have been reduced or lost in some. The annelid digestive system
aid aquatic forms in swimming. Since many annelids burrow or is not segmented: the gut runs the length of the body perforating
live in secreted tubes, stiff setae also aid in preventing the worm each septum (Figure 17.3). Longitudinal dorsal and ventral blood
from being pulled out or washed out of its home. Robins know vessels follow the same path, as does the ventral nerve cord.
from experience how effective earthworms’ setae are. Traditionally, annelids are divided among three classes:
Annelids have a worldwide distribution, and a few species Polychaeta, Oligochaeta, and Hirudinida. Polychaeta is a para-
are cosmopolitan. Polychaetes are chiefly marine forms. Most phyletic class because ancestors of oligochaetes and hirudine-
are benthic, but some live pelagic lives in the open seas. Oligo- ans (leeches) arose from within polychaetes. Oligochaetes and
chaetes and leeches occur predominantly in freshwater or terres- leeches together form a monophyletic group called Clitellata (see
trial soils. Some freshwater species burrow in mud and sand and Figure 17.1), characterized by presence of a reproductive struc-
others among submerged vegetation. Many leeches are preda- ture called a clitellum (see p. 371). Some authorities now con-
tors, specialized for piercing their prey and feeding on blood or sider Clitellata to be an annelid class containing oligochaetes and
soft tissues. A few leeches are marine, but most live in freshwater leeches as orders, but we retain the three classes and consider
or in damp regions. Suckers are typically found at both ends of Clitellata a clade whose members are class Oligochaeta and class
the body for attachment to the substratum or to their prey. Hirudinida. Class Oligochaeta is a paraphyletic group because
ancestors of leeches arose from within it.
Body Plan
The annelid body typically has a two-part head, composed of Class Polychaeta
a prostomium and a peristomium followed by a segmented The largest class of annelids is the Polychaeta (Gr. polys, many,
body and a terminal portion called the pygidium bearing an chaitē, long hair) with more than 10,000 species, most of them
anus (Figure 17.2). The head and pygidium are not considered marine. Although most polychaetes are 5 to 10 cm long, some
to be segments. New segments differentiate during development are less than 1 mm, and others may be as long as 3 m. They may
just in front of the pygidium; thus the oldest segments are at the be brightly colored in reds and greens, iridescent, or dull.
anterior end and the youngest segments are at the posterior end. Many polychaetes are euryhaline and can tolerate a wide
Each segment typically contains circulatory, respiratory, nervous, range of environmental salinity. The freshwater polychaete fauna
and excretory structures, as well as a coelom. is more diversified in warmer regions than in temperate zones.
In most annelids the coelom develops embryonically as a Many polychaetes live under rocks, in coral crevices, or in
split in the mesoderm on each side of the gut (schizocoel), abandoned shells. A number of species burrow into mud or sand
forming a pair of coelomic compartments in each segment. Peri- and build their own tubes on submerged objects or in bottom
toneum (a layer of mesodermal epithelium) lines the body wall sediment. Others adopt the tubes or homes of other animals, and
of each compartment, forming dorsal and ventral mesenteries some are planktonic. They are extremely abundant in some areas;
that cover all organs (Figure 17.3). Peritonea of adjacent segments for example, a square meter of mudflat may contain thousands of
hic70049_ch17_362-383.indd 364 7/14/07 3:18:06 PM

Jaw Everted
pharynx
Prostomial
tentacles Dorsal Oblique Dorsal Intestine Coelomic
cirrus muscle vessel epithelium
Palp
Longitudinal
Prostomium Respiratory Eggs muscle
Eyes capillaries
Circular muscle
Peristomium
Tentacles
(cirri) Notopodium
Parapodia
Parapodium
A
Setae
Neuropodium
Aciculum Ventral Nephridium Nerve Ventral Epidermis

B cirrus cord vessel
D
Figure 17.2
Nereis virens (A–D) and
Nereis diversicolor (E)
are errant polychaetes.
A, Anterior end, with
pharynx everted. B,
External structure.
C, Posterior end. D,
Generalized transverse Parapodia
section through region
of the intestine. E, In Pygidium
this photo of a live N.
diversicolor, note the Anus
well-defined segments,
the lobed parapodia, Cirrus
and the prostomium
with tentacles. C E
Longitudinal Dorsal blood Segment Intestine

muscle vessel
polychaetes. They play a significant part in marine food chains
because they are eaten by fish, crustaceans, hydroids, and many
Septum other predators.
Circular
muscle
Polychaetes differ from other annelids in having a well-
Ventral differentiated head with specialized sense organs; paired append-
mesentery ages, called parapodia, on most segments; and no clitellum (Fig-
Dorsal
mesentery ure 17.2). As their name implies, they have many setae, usually
Ventral arranged in bundles on the parapodia. They exhibit the most pro-
blood nounced differentiation of body segments and specialization of
vessel
sensory organs found in annelids (see p. 367).
Septum Polychaetes are often divided into two morphological groups
Parietal Circular Longitudinal based on their activity: sedentary polychaetes and errant (free-
peritoneum muscle muscle moving) polychaetes. Sedentary polychaetes spend much or all of
Visceral
peritoneum their time in tubes or permanent burrows. Many of them, especially
Intestine
those that live in tubes, have elaborate devices for feeding and
respiration (Figure 17.4). Errant polychaetes (L. errare, to wander),
Ventral Cuticle Ventral blood Epidermis
mesentery vessel
include free-swimming pelagic forms, active burrowers, crawlers,
and tubeworms that only leave their tubes for feeding or breed-
Figure 17.3 ing. Most of these, like clam worms in the genus Nereis (Gr. name
Annelid body plan. of a sea nymph) (see Figure 17.2), are predatory and equipped
hic70049_ch17_362-383.indd 365 7/14/07 3:18:06 PM

Figure 17.4
Tube-dwelling sedentary polychaetes. A, Christmas-tree worm, Spirobranchus giganteus, live in a calcareous tube. On its head are two whorls of
modified tentacles (radioles) used to collect suspended food particles from the surrounding water. Notice the finely branched filters visible on the
edge of one radiole. B, Sabellid polychaetes, Bispira brunnea, live in leathery tubes.
with jaws or teeth. They have an eversible, muscular pharynx

Tentacle
armed with teeth that can be thrust out with surprising speed to A
capture prey.
Gills B
Form and Function
A polychaete typically has a prostomium, which may or may C
not be retractile and which often bears eyes, tentacles, and sen-
sory palps (see Figure 17.2). The peristomium surrounds the
mouth and may bear setae, palps, or, in predatory forms, chitin-
D
ous jaws. Ciliary feeders may bear a crown of tentacles that can
be opened like a fan or withdrawn into the tube (Figure 17.4). Notopodia
Neuropodia
The polychaete trunk is segmented, and most segments bear
parapodia, which may have lobes, cirri, setae, and other parts on
them (see Figure 17.2). Parapodia are used in crawling, swim-
ming, or for anchoring the animal in its tube. They usually serve
as the chief respiratory organs, although some polychaetes also
have gills. Amphitrite (Gr. a mythical sea nymph), for example,
has three pairs of branched gills and long extensible tentacles (Fig-
ure 17.5). Arenicola (L. arena, sand, colo, inhabit), the burrow-
ing lugworm (Figure 17.6), has paired gills on certain segments.
Figure 17.5
Nutrition Amphitrite, which builds its tubes in mud or sand, extends long
grooved tentacles out over the mud to pick up bits of organic matter.
A polychaete’s digestive system consists of a foregut, a midgut, The smallest particles are moved along food grooves by cilia, larger
and a hindgut. The foregut includes a stomodeum, a phar- particles by peristaltic movement. Its plumelike gills are blood red.
ynx, and an anterior esophagus. It is lined with cuticle, and A, Section through exploratory end of tentacle. B, Section through
the jaws, where present, are constructed of cuticular protein. tentacle in area adhering to substratum. C, Section showing ciliary
groove. D, Particle being carried toward mouth.
The more anterior portions of the midgut secrete digestive
enzymes but absorption takes place toward the posterior end.
A short hindgut connects the midgut to the exterior via the
anus, which is on the pygidium. Circulation and Respiration
Errant polychaetes are typically predators and scavengers. Polychaetes show considerable diversity in both circulatory and
Sedentary polychaetes feed on suspended particles, or they may respiratory structures. As previously mentioned, parapodia and
be deposit feeders, consuming particles on or in the sediment. gills serve for gaseous exchange in various species. However,
hic70049_ch17_362-383.indd 366 7/14/07 3:18:09 PM

Sand falling into shaft Water movement lumen of the cup. The highest degree of eye development occurs
in the family Alciopidae, which has large, image-resolving eyes sim-
ilar in structure to those of some cephalopod molluscs (see Figure
16.40, p. 355), with cornea, lens, retina, and retinal pigments. Alci-
opid eyes also have accessory retinas, a characteristic independently
evolved by deep-sea fishes and some deep-sea cephalopods. The
accessory retinas of alciopids are sensitive to different wavelengths.
The eyes of these pelagic animals may be well adapted to function
because penetration by the different wavelengths of light varies
with depth. Studies with electroencephalograms show that they are
sensitive to dim light of the deep sea. Nuchal organs are ciliated
sensory pits or slits that appear to be chemoreceptive, an important
factor in food gathering. Some burrowing and tube-building poly-
chaetes have statocysts that function in body orientation.
Proboscis Gills
Figure 17.6 Reproduction and Development

Arenicola, the lugworm, lives in a U-shaped burrow in intertidal
mudflats. It burrows by successive eversions and retractions of its
Polychaetes have no permanent sex organs, and they usually
proboscis. By peristaltic movements it keeps water filtering through have separate sexes. Reproductive systems are simple: Gonads
the sand. The worm then ingests the food-laden sand. appear as temporary swellings of the peritoneum and shed their
gametes into the coelom. The gametes are then carried to the
in some polychaetes there are no special organs for respiration, outside through gonoducts, through the metanephridia, or by
and gaseous exchange takes place across the body surface. rupture of the body wall. Fertilization is external, and the early
The circulatory pattern varies greatly. In Nereis a dorsal lon- larva is a trochophore (see Figure 16.7).
gitudinal vessel carries blood anteriorly, and a ventral longitudi-
nal vessel conducts it posteriorly (see Figure 17.2D). Blood flows Some polychaetes live most of the year as sexually immature animals
between these two vessels via segmental networks in the para- called atokes, but during the breeding season a portion of the body
podia, septa, and around the intestine. In the burrowing preda- becomes sexually mature and swollen with gametes (Figure 17.7).
tory worm Glycera (Gr. Glykera, a feminine proper name) the An example is the palolo worm, which lives in burrows among coral
circulatory system is reduced and joins directly with the coelom. reefs. During the swarming period, the sexually mature portions,
Septa are incomplete, and thus the coelomic fluid assumes the now called epitokes, break off and swim to the surface. Just before
function of circulation. sunrise, the sea is literally covered with them, and at sunrise they
Many polychaetes have respiratory pigments such as hemo- burst, freeing eggs and sperm for fertilization. Anterior portions of
globin, chlorocruorin, or hemerythrin (p. 704). the worms regenerate new posterior sections. Swarming is of great
adaptive value because the synchronous maturation of all the epit-
Excretion okes ensures the maximum number of fertilized eggs. However, this
reproductive strategy is very hazardous; many types of predators
Excretory organs consist of protonephridia and mixed proto- and have a feast on the swarming worms. In the meantime, the atoke
metanephridia in some, but most polychaetes have metanephridia remains safely in its burrow to produce another epitoke at the next
(see Figure 17.2). There is one pair per segment, with the inner cycle. In some polychaetes, epitokes arise from atokes by asexual
end of each (nephrostome) opening into a coelomic compart- budding (Figure 17.8) and become complete worms.
ment. Coelomic fluid passes into the nephrostome, and selective
resorption occurs along the nephridial duct (see Figure 17.18).
Nervous System and Sense Organs Representative Polychaetes

Organization of the central nervous system in polychaetes fol- Clam Worms: Nereis Clam worms (see Figure 17.2), or sand
lows the basic annelid plan (see Figure 17.19). Dorsal cerebral worms as they are sometimes called, are errant polychaetes that
ganglia connect with a subpharyngeal ganglion via a circum- live in mucous-lined burrows in or near low tide. Sometimes they
pharyngeal connective. A double ventral nerve cord courses the are found in temporary hiding places, such as under stones, where
length of the worm, with metamerically arranged ganglia. they stay with their bodies covered and their heads protruding.
Sense organs are highly developed in polychaetes and include They are most active at night, when they wiggle out of their hiding
eyes, nuchal organs, and statocysts. Eyes, when present, may range places and swim or crawl over the sand in search of food.
from simple eyespots to well-developed organs. Eyes are most con- The body, containing about 200 segments, may grow to 30
spicuous in errant worms. Usually the eyes are retinal cups, with or 40 cm in length. The head is composed of a prostomium and
rodlike photoreceptor cells (lining the cup wall) directed toward the a peristomium. The prostomium bears a pair of stubby palps,
hic70049_ch17_362-383.indd 367 7/14/07 3:18:13 PM

Figure 17.7
Eunice viridis, the Samoan palolo worm. Characteristics of Phylum Annelida
The posterior segments make up the
Atoke
epitokal region, consisting of segments 1. Unique annelid head and paired epidermal setae present
packed with gametes. Each segment has (lost in leeches); parapodia present in the ancestral condition
an eyespot on the ventral side. Once a 2. Marine, freshwater, and terrestrial
year the worms swarm, and the epitokes 3. Most free-living, some symbiotic, some ectoparasitic
detach, rise to the surface, and discharge 4. Body bilaterally symmetrical, metameric, often with distinct
their ripe gametes, leaving the water head
milky. By the next breeding season, the
Epitoke 5. Triploblastic body
epitokes are regenerated.
6. Coelom (schizocoel) well developed and divided by septa,
except in leeches; coelomic fluid functions as hydrostatic
sensitive to touch and taste; a pair of short skeleton
sensory tentacles; and two pairs of small 7. Epithelium secretes outer transparent moist cuticle
dorsal eyes that are light sensitive. The 8. Digestive system complete and not segmentally arranged
peristomium bears the ventral mouth, a 9. Body wall with outer circular and inner longitudinal muscle
pair of chitinous jaws, and four pairs of layers
sensory tentacles (see Figure 17.2A). 10. Nervous system with a double ventral nerve cord and a pair
Each parapodium has two lobes: a of ganglia with lateral nerves in each segment; brain a pair of
dorsal cerebral ganglia with connectives to ventral nerve cord
dorsal notopodium and a ventral neu-
11. Sensory system of tactile organs, taste buds, statocysts (in
ropodium (see Figure 17.2D) that bear
some), photoreceptor cells, and eyes with lenses (in some);
setae with many blood vessels. Para- specialization of head region into differentiated organs, such
podia are used for both creeping and as tentacles, palps, and eyespots of polychaetes
swimming and are controlled by oblique 12. Asexual reproduction by fission and fragmentation; capable
muscles that run from the midventral line of complete regeneration
to the parapodia in each segment. The 13. Hermaphroditic or separate sexes; larvae, if present, are
worm swims by lateral undulatory move- trochophore type; asexual reproduction by budding in some;
ment of the body. It can dart through the spiral cleavage and mosaic development
water with considerable speed. These undulatory movements can 14. Excretory system typically a pair of nephridia for each
also be used to suck water into or pump it out of the burrow. segment; nephridia remove waste from blood as well as
from coelom
Clam worms feed on small animals, other worms, and a
15. Respiratory gas exchange through skin, gills, or parapodia
variety of larval forms. They seize food with their chitinous jaws,
16. Circulatory system closed with muscular blood vessels
which they protrude through the mouth when they evert their and aortic arches (“hearts”) for pumping blood, segmentally
pharynx. Food is swallowed as the worm withdraws its pharynx. arranged; respiratory pigments (hemoglobin, hemerythrin, or
Movement of food through the alimentary canal is by peristalsis. chlorocruorin) often present; amebocytes in blood plasma
Scale Worms Scale worms (Figure 17.9) are members of the

family Polynoidae (Gr. Polynoē, daughter of Nereus and Doris,
Atoke
a sea god and goddess), one of the most diverse, abundant, and
widespread of polychaete families. Their flattened bodies are cov-
ered with broad scales, modified from dorsal parts of the para-
podia. Most species are of modest size, but some are enormous
(up to 190 mm long and 100 mm wide). They are carnivorous
and eat a wide variety of animals. Many are commensal, living in
burrows of other polychaetes or in association with cnidarians,
molluscs, or echinoderms.
Fireworms Hermodice carunculata (Gr. herma, reef, dex,

Epitokes
a worm found in wood) (Figure 17.10) and related species are
called fireworms because their hollow, brittle setae contain a
poisonous secretion. The setae puncture a hand that touches
Figure 17.8 them, and then break off in the wound to cause skin irritation.
Rather than transforming a portion of its body into Fireworms feed on corals, gorgonians, and other cnidarians.
an epitoke, Autolytus prolifer asexually buds off
complete worms from its posterior end that become Tubeworms Polychaete tube-dwellers secrete many types of
sexual epitokes. tubes. Some are parchmentlike or leathery (see Figure 17.4B);
hic70049_ch17_362-383.indd 368 7/14/07 3:18:18 PM

Cilia
Direction of
Radiole water flow
Pinnule section
Direction of
food movement
Pinnule
Mouth
Ventral sac Food groove
(sand storage)
B
Tube
Figure 17.9 Various sizes
A scale worm, Hesperonoe adventor, normally lives as a commensal in of particles
the tubes of Urechis (phylum Echiura, p. 379). sorted
Proximal radiole sorting

A mechanism
Figure 17.11
Sabella, a polychaete ciliary feeder, extends its crown of feeding
radioles from its leathery secreted tube, reinforced with sand and debris.
A, Anterior view of the crown. Cilia direct small food particles along
grooved radioles to mouth and discard larger particles. Sand grains are
directed to storage sacs and later are used in tube building. B, Distal
portion of radiole showing ciliary tracts of pinnules and food grooves.
deposit feeding. They live in a U-shaped burrow through which, by

peristaltic movements, they cause water to flow. Food particles are
trapped by the sand at the front of the burrow, and Arenicola then
ingests the food-laden sand (see Figure 17.6).
Fanworms, or “featherduster” worms, are beautiful tubeworms,
fascinating to watch as they emerge from their secreted tubes and
unfurl their lovely tentacular crowns to feed (see Figure 17.4). A
Figure 17.10 slight disturbance, sometimes even a passing shadow, causes them
A fireworm, Hermodice carunculata, feeds on gorgonians and stony
to duck back quickly into the safety of the homes. Food drawn
corals. Its setae are like tiny glass fibers and serve to ward off predators.
to the feathery arms, or radioles, by ciliary action is trapped in
mucus and carried down ciliated food grooves to the mouth (Fig-
some are firm, calcareous tubes attached to rocks or other surfaces ure 17.11). Particles too large for the food grooves pass along the
(see Figure 17.4A); and some are simply grains of sand or bits margins of the food grooves and fall away before they reach the
of shell or seaweed cemented together with mucous secretions. mouth. Only small particles of food enter the mouth; sand grains
Many species burrow in sand or mud, lining their burrows with are stored in a sac to be used later in enlarging the tube.
mucus (see Figure 17.6). The parchment worm, Chaetopterus (Gr. chaite¯, long hair,
Most sedentary tube and burrow dwellers are particle feeders, pteron, wing), feeds on suspended particles by an entirely
using cilia or mucus to obtain food, typically plankton and detritus. different mechanism (Figure 17.12). It lives in a U-shaped, parch-
Some deposit feeders, like Amphitrite (see Figure 17.5), protrude mentlike tube buried, except for the tapered ends, in sand or
their heads above the mud and extend long tentacles over the sur- mud along the shore. The worm attaches to the side of the
face to find food. Cilia and mucus on the tentacles entrap particles tube by ventral suckers. Fans (modified parapodia on segments
found on the sea bottom and move them toward the mouth. Lug- 14 to 16) pump water through the tube by rhythmical move-
worms, Arenicola, use an interesting combination of suspension and ments. A pair of enlarged parapodia on segment 12 secretes a
hic70049_ch17_362-383.indd 369 7/14/07 3:18:19 PM

Water movement Clade Siboglinidae (Pogonophorans) Members of for-

mer phylum Pogonophora (pogo-nofe-ra) (Gr. pōgōn, beard,
phora, bearing), or beardworms, were entirely unknown before
the twentieth century. The first specimens to be described were
collected from deep-sea dredgings off the coast of Indonesia in
Parchmentlike 1900. They have since been discovered in several seas, including
tube the western Atlantic off the U.S. eastern coast. Some 150 species
have been described so far. Most species are less than 1 mm in
diameter but can be 10 to 75 cm in length.
Most siboglinids live in mud on the ocean floor, at depths of
Mouth
100 to 10,000 m. This location accounts for their delayed discovery,
"Wing" (12th notopodia) for they are obtained only by dredging. They are sessile animals
that secrete very long chitinous tubes in which they live, and prob-
Mucous net ably only extend the anterior end of their body for absorbing nutri-
"Fans" ents. The tubes are generally oriented upright in bottom sediments.
A tube can be three to four times the length of the animal, which
can move up or down inside its tube but cannot turn around.
Beardworms have a long, cylindrical body covered with
Food cup
cuticle. Cuticle, epidermis, and circular and longitudinal muscles
compose the body wall. The body is divided into a short anterior
forepart; a long, very slender trunk; and a small, segmented
Figure 17.12 opisthosoma (Figure 17.13). Paired epidermal setae are present
Chaetopterus, a sedentary polychaete, lives in a U-shaped tube in the on the trunk and opisthosoma. At the anterior end of the body, a
sea bottom. It pumps water through the parchmentlike tube (of which cephalic lobe bears from 1 to 260 long tentacles (the “beard” that
one-half has been cut away here) with its three pistonlike fans. The gives this phylum its name), depending on species. Tentacles are
fans beat 60 times per minute to keep water currents moving. The
winglike notopodia of the twelfth segment continuously secrete a
hollow extensions of the coelom and bear minute pinnules. For
mucous net that strains out food particles. As the net fills with food, a part or all of their length, tentacles lie parallel with each other,
the food cup rolls it into a ball, and when the ball is large enough enclosing a cylindrical intertentacular space into which the pin-
(about 3 mm), the food cup bends forward and deposits the ball in a nules project (Figure 17.14).
ciliated groove to be carried to the mouth and swallowed. Siboglinids are remarkable in having no mouth or digestive
tract, making their mode of nutrition a puzzling matter. They
long mucous net that reaches back to a small food cup just in absorb some nutrients dissolved in seawater, such as glucose,
front of the fans. All water passing through the tube is filtered amino acids, and fatty acids, through the pinnules and microvilli
through this mucous net, the end of which is rolled into a ball of their tentacles. Most of their energy, however, apparently is
by cilia in the cup. When the ball is about the size of a BB shot derived from a mutualistic association with chemoautotrophic
(about 3 mm diameter), the fans stop beating and the ball of bacteria. These bacteria oxidize hydrogen sulfide to provide
food and mucus is rolled forward by ciliary action to the mouth energy to produce organic compounds from carbon dioxide.
and swallowed. Siboglinids bear the bacteria in an organ called a trophosome,
Tentacles Ciliated band Papilla
Cephalic lobe
Papillae Opisthosoma
Forepart
Bridle Surface
Band of Glandular
cilia shield Secreted
tube
Paired
Trunk
papillae
A B
Figure 17.13
Diagram of a typical siboglinid. A, External features. In life, the body is much more elongated than shown in this diagram. B, Position in tube.
hic70049_ch17_362-383.indd 370 7/14/07 3:18:24 PM

Sexes are separate, with a pair of

gonads and a pair of gonoducts in the
Nerve trunk section. Little developmental work
Tentacles
Coelom has been done on these deep-sea worms,
Pinnule Tentacle but research suggests that cleavage is
Cuticle
Efferent blood unequal and atypical. It seems to be closer
vessel to spiral than to radial. Development of
Coelomic the apparent coelom is schizocoelic, not
Afferent blood
canal
vessel enterocoelic as was originally described.
The worm-shaped embryo is ciliated but
Cephalic Pinnule containing a poor swimmer. It is probably carried by
lobe Intertentacular
cavity afferent and water currents until it settles.
Cross section of efferent branches
tentacular crown
Enlargement of
Clade Clitellata
Anterior portion two tentacles Clade Clitellata contains earthworms,
of Lamellisabella in cross section
and their relatives, in class Oligochaeta
Figure 17.14 and leeches in class Hirudinida. Mem-
Cross section of tentacular crown of siboglinid Lamellisabella. Tentacles arise from ventral side of bers of this clade share a unique repro-
forepart at base of cephalic lobe. Tentacles (which vary in number in different species) enclose a ductive structure called a clitellum. The
cylindrical space, with the pinnules forming a kind of nutrient uptake network. Food molecules may clitellum is a ring of secretory cells in the
be absorbed into the blood supply of tentacles and pinnules. epidermis that appears on the worm’s
exterior as a fat band around the body
which is derived embryonically from the midgut (all traces of about one-third of the body length from the anterior end. The cli-
foregut and hindgut are absent in adults). tellum is always visible in oligochaetes, but it appears only dur-
There is a well-developed closed circulatory system. Photo- ing the reproductive season in leeches. Members of Clitellata are
receptors are similar to those of other annelids. derived annelids that lack parapodia, presumably an evolutionary
loss from a polychaete ancestor. Clitellates are all hermaphroditic
Among the most amazing animals found in deep-water, Pacific (monoecious) animals that exhibit direct development: Young
rift communities (see Chapter 38, p. 839) are vestimentiferans, develop inside a cocoon secreted by the clitellum, so no trocho-
Riftia pachyptila. These giant beardworms live around deep-water phore larva is visible. Small worms emerge from cocoons.
hydrothermal vents and grow up to 3 m long and 5 cm in diameter
(Figure 17.15). The trophosome of other siboglinids is confined Class Oligochaeta
to the posterior part of the trunk, which is buried in sulfide-rich More than 3000 species of oligochaetes are found in a great vari-
sediments, but the trophosome of Riftia occupies most of its large ety of sizes and habitats. They include the familiar earthworms
trunk. It has a much larger supply of hydrogen sulfide, enough to and many species that live in freshwater. Most are terrestrial
nourish its large body, in the effluent of the hydrothermal vents. or freshwater forms, but some are parasitic, and a few live in
marine or brackish water.
With few exceptions, oligochaetes bear setae, which may be
long or short, straight or curved, blunt or needlelike, or arranged
singly or in bundles. Whatever the type, setae are less numerous
in oligochaetes than in polychaetes, as is implied by the class
name, which means “few long hairs.” Aquatic forms usually have
longer setae than do earthworms.
Form and Function The main features of an oligochaete
body are described with reference to the familiar earthworm.
The circulatory system and excretory structures described in
earthworms are typical of annelids in general, but the digestive
and nervous systems have aspects specific to oligochaetes.
Earthworms, sometimes called “night crawlers,” burrow in
moist rich soil, and usually live in branched, interconnected tun-
nels. The species commonly studied in laboratories is Lumbri-
Figure 17.15 cus terrestris (L. lubricum, earthworm). It ranges in size from
A colony of giant beardworms (vestimentiferans, clade Siboglinidae) 12 to 30 cm long (Figure 17.16), but is small in comparison to
at great depth near a hydrothermal vent along the Galápagos Trench, giant tropical forms whose 4-m-long bodies may comprise 150 to
eastern Pacific Ocean. upward of 250 segments.
hic70049_ch17_362-383.indd 371 7/14/07 3:18:31 PM

To Part C below
Prostomium
Seminal receptacle Genital openings Clitellum Setae

openings
A
Anus
Pharynx Lateral Second left Seminal Esophagus Epidermis Gizzard Circular Longitudinal
nerve aortic arch vesicles muscle muscle
Brain Dorsal
vessel
Buccal cavity
Crop
Prostomium
Intestine
Mouth
Testes Ovary
B Left circumpharyngeal Seminal Sperm Egg funnel Nerve Subneural Ventral

connective receptacle funnel and oviduct cord vessel Nephridium vessel
Circular Epidermis Cuticle Dorsal vessel Mucous

muscle gland cell Pores of gland
Longitudinal Sensory (receptor) cell
muscle of sense organ
Peritoneum Epithelial cell
Cuticle
Setae
Typhlosole Photoreceptor cell
Intestinal lumen Sensory fibers

Setal retractor D
Chloragogen cells muscle
Figure 17.16
Intestinal Earthworm anatomy. A, External features, lateral
epithelium Nephridium view. B, Internal structure of anterior portion of
worm. C, Generalized transverse section through
Lateroneural Ventral Subneural Ventral region posterior to clitellum. D, Portion of epidermis
C vessel vessel vessel nerve cord showing sensory, glandular, and epithelial cells.
Earthworms normally emerge at night, but in damp rainy against the sides of the burrow. As they do so, bristlelike rods
weather they stay near the surface, often with mouth or anus protrud- called setae project outward through small pores in the cuticle.
ing from the burrow. In very dry weather they may burrow several Setae dig into the walls of the burrow to anchor the forward seg-
feet underground, coil in a slime chamber and become dormant. ments; contractions of longitudinal muscles then shorten the rest
Earthworms use peristaltic movement: Contractions of cir- of the body, pulling the posterior end up behind the anchored
cular muscles in the anterior end lengthen the body, pushing anterior region. As waves of extension and contraction pass
the anterior end forward where it anchors. Anchoring is accom- along the entire body, it gradually moves forward.
plished by contraction of the longitudinal muscles in forward The paired epidermal setae of oligochaetes are set in a sac
segments—these segments become short and wide, pushing within the body wall and moved by muscles (Figure 17.17), as
hic70049_ch17_362-383.indd 372 7/14/07 3:18:33 PM

Figure 17.17 Longitudinal as a center for synthesis of glycogen and fat, a func-
muscle tion roughly equivalent to that of liver cells. When
Seta with its muscle
attachments Formative cell Circular full of fat, chloragogen cells are released into the coe-
showing relation to muscle lom where they float freely as cells called eleocytes
adjacent structures.
Seta
(Gr. elaio, oil, kytos, hollow vessel [cell]), which
Setae lost by wear
transport materials to the body tissues. Eleocytes can
are replaced by new
ones, which develop pass from segment to segment and may accumu-
from formative cells. Epidermis
late around wounds and regenerating areas, where
they break down and release their contents into the
Retractor coelom. Chloragogen cells also function in excretion.
muscle Cuticle
Peritoneum Protractor Circulation and Respiration Annelids have a
muscle
double transport system: coelomic fluid and a closed
they are in polychaetes. However, oligochaetes do not have circulatory system. Food, wastes, and respiratory
parapodia; instead the setae extend directly out of the body wall gases are carried by both coelomic fluid and blood in varying
on each segment. In most earthworms each segment bears four degrees. Blood circulates in a closed system of vessels, which
pairs of chitinous setae (Figure 17.16C), although there may be includes capillary systems in the tissues. Five main blood trunks
more than 100 such setae per segment in some oligochaetes. run lengthwise through the body.
A single dorsal vessel runs above the alimentary canal from
the pharynx to the anus. It is a pumping organ, provided with
Aristotle called earthworms the “intestines of the soil.” Some 22 valves, and it functions as a true heart. This vessel receives blood
centuries later Charles Darwin published his observations in his from vessels of the body wall and digestive tract and pumps it ante-
classic The Formation of Vegetable Mould Through the Action of riorly into five pairs of aortic arches. The function of aortic arches
Worms. He showed how worms enrich soil by bringing subsoil to is to maintain a steady pressure of blood in the ventral vessel.
the surface and mixing it with topsoil. An earthworm can ingest A single ventral vessel serves as an aorta. It receives blood
its own weight in soil every 24 hours, and Darwin estimated that from the aortic arches and delivers it to the brain and rest of the
from 10 to 18 tons of dry earth per acre pass through their intestine body, providing segmental vessels to the walls, nephridia, and
annually, thus bringing potassium and phosphorus from the subsoil digestive tract.
and also adding nitrogenous products to the soil from their own Their blood contains colorless ameboid cells and a dissolved
metabolism. They also drag leaves, twigs, and organic substances respiratory pigment, hemoglobin (p. 704). The blood of some
into their burrows closer to the roots of plants. Their activities are annelids may have respiratory pigments other than hemoglobin,
vitally important in aerating soil. Darwin’s views were at odds with as noted on page 367.
his contemporaries, who thought earthworms were harmful to Earthworms have no special respiratory organs, but gaseous
plants. But recent research has amply confirmed Darwin’s findings, exchange occurs across their moist skin.
and earthworm management is now practiced in many countries.
Excretion Each segment (except the first three and the last
one) bears a pair of metanephridia. Each metanephridium
Nutrition Most oligochaetes are scavengers. Earthworms feed occupies parts of two successive segments (Figure 17.18). A cili-
mainly on decaying organic matter, bits of leaves and vegetation, ated funnel, the nephrostome, lies just anterior to an interseg-
refuse, and animal matter. After being moistened by secretions from mental septum and leads by a small ciliated tubule through the
the mouth, food is drawn inward by the sucking action of their septum into the segment behind, where it connects with the
muscular pharynx. The liplike prostomium aids in manipulating main part of the nephridium. Several complex loops of increas-
food into position. Calcium from soil swallowed with food tends ing size compose the nephridial duct, which terminates in a blad-
to produce a high blood calcium level. Calciferous glands along derlike structure leading to an opening, the nephridiopore.
the esophagus secrete calcium ions into the gut and so reduce the The nephridiopore opens to the outside near the ventral row of
calcium ion concentration of their blood. Calciferous glands also setae. By means of cilia, wastes from the coelom are drawn into
function in regulating acid-base balance of body fluids. the nephrostome and tubule, where they are joined by salts and
Leaving the esophagus, food is stored temporarily in the organic wastes transported from blood capillaries in the glandu-
thin-walled crop before being passed on to the gizzard, which lar part of the nephridium. Waste is discharged to the outside
grinds food into small pieces. Digestion and absorption occur in through a nephridiopore.
the intestine. The wall of the intestine is infolded dorsally to Aquatic oligochaetes excrete ammonia; terrestrial oligo-
form a typhlosole, which greatly increases the absorptive and chaetes usually excrete the much less toxic urea. Lumbri-
digestive surface (Figure 17.16C). cus produces both, the level of urea depending somewhat
Surrounding the intestine and dorsal vessel and filling much on environmental conditions. Both urea and ammonia are
of the typhlosole is a layer of yellowish chloragogen tissue produced by chloragogen cells, which may break off and
(Gr. chlōros, green, agōgē, a carrying away). This tissue serves enter the metanephridia directly, or their products may be
hic70049_ch17_362-383.indd 373 7/14/07 3:18:36 PM

Posterior Lateral nerves Pharynx Cerebral ganglia

Nephridia
Buccal cavity
Anterior
Prostomium
Nephric
tubule
Gut
Capillary
network
Bladder
Sensory
Septum endings
Ciliated funnel
Mouth
(nephrostome)
Nerve cord Subpharyngeal Circumpharyngeal
ganglion connective
Figure 17.19
Nephridiopore Anterior portion of earthworm and its nervous system. Note
concentration of sensory endings in this region.
Figure 17.18
Nephridium of earthworm. Wastes are drawn into the ciliated
nephrostome in one segment, then passed through the loops of
Neurosecretory cells have been found in the brain and
the nephridium, and expelled through the nephridiopore of the
next segment. ganglia of both oligochaetes and polychaetes. They are endo-
crine in function and secrete neurohormones concerned with
the regulation of reproduction, secondary sex characteristics,
carried by the blood. Some nitrogenous waste is eliminated
and regeneration.
through the body surface.
For rapid escape movements most annelids have from
Oligochaetes are largely freshwater animals, and even such
one to several very large axons commonly called giant
terrestrial forms as earthworms must exist in a moist environ-
axons ( Figure 17.20 ), or giant fi bers, located in the ventral
ment. Osmoregulation is a function of the body surface and the
nerve cord. Their large diameter increases rate of conduction
nephridia, as well as the gut and dorsal pores. Lumbricus will
(see p. 730) and makes possible simultaneous contractions of
gain weight when placed in tap water and lose it when returned
muscles in many segments.
to soil. Salts as well as water can pass across the integument,
salts apparently being actively transported.
Nervous System and Sense Organs

The nervous system in earthworms (Fig-
ure 17.19 ) consists of a central system
and peripheral nerves. The central system
reflects the typical annelid pattern: a pair Lateral giant
of cerebral ganglia (the “brain”) above fiber connections
the pharynx, a pair of connectives passing Median giant
fiber Nerve sheath
around the pharynx connecting the brain
with the first pair of ganglia in the nerve Lateral giant
Lateral nerve
cord; a solid ventral nerve cord, really fiber
double, running along the floor of the coe-
lom to the last segment; and a pair of fused Sensory neuron
ganglia on the nerve cord in each segment.
Each pair of fused ganglia provides nerves
to the body structures, which contain both Motor neuron
sensory and motor fibers. Ventral giant
nerve cells Association neuron
Figure 17.20
Portion of nerve cord of earthworm showing arrangement of simple reflex arc (in foreground; see also Sensory cell
p. 734) and the three dorsal giant fibers that are adapted for rapid reflexes and escape movements. (receptor)
Ordinary crawling involves a succession of reflex acts, the stretching of one segment stimulating the next to
stretch. Impulses are transmitted much faster in giant fibers than in regular nerves so that all segments can Muscle
contract simultaneously when quick withdrawal into a burrow is necessary. (effector)
hic70049_ch17_362-383.indd 374 7/30/07 9:24:14 AM

When mating, worms extend their anterior ends from their bur-
In the dorsal median giant fiber of Lumbricus, which is 90 to rows and bring their ventral surfaces together (Figure 17.21).
160 m in diameter, speed of conduction has been estimated Their surfaces are held together by mucus secreted by the
at 20 to 45 m/second, several times faster than in ordinary neu- clitellum (L. clitellae, packsaddle) and by special ventral setae,
rons of this species. This is also much faster than in polychaete which penetrate each other’s bodies in the regions of contact.
giant fibers, probably because in earthworms the giant fibers are After discharge, sperm travel to seminal receptacles of the other
enclosed in myelinated sheaths, which insulate them. worm via its seminal grooves. After copulation each worm
secretes first a mucous tube and then a tough, chitinlike band
Simple sense organs are distributed all over the body. Earth- that forms a cocoon around its clitellum. As the cocoon passes
worms have no eyes but do have many lens-shaped photo- forward, eggs from the oviducts, albumin from skin glands, and
receptors in their epidermis. Most oligochaetes are negatively sperm from the mate (stored in the seminal receptacles) pour into
phototactic to strong light but positively phototactic to weak it. Fertilization of eggs then occurs within the cocoon. When the
light. Many single-celled sense organs are widely distributed in cocoon slips past the anterior end of the worm, its ends close,
the epidermis. What are presumably chemoreceptors are most producing a sealed, lemon-shaped body. Embryogenesis occurs
numerous on the prostomium. In the integument are many free within the cocoon, and the form that hatches from the egg is
nerve endings which are probably tactile in nature. a young worm similar to the adult. Thus development is direct
with no metamorphosis. Juveniles do not develop a clitellum
General Behavior Earthworms are among the most defense- until they are sexually mature.
less of creatures, yet their abundance and wide distribution indi- Representative Oligochaetes Freshwater oligochaetes
cate their ability to thrive. Although they have no specialized usually are smaller and have more conspicuous setae than earth-
sense organs, they are sensitive to many stimuli. They react worms. They are more mobile than earthworms and tend to have
positively to mechanical stimuli when such stimuli are moderate better-developed sense organs. Most are benthic forms that crawl
and negatively to a strong stimulus (such as footfall near them), on the substrate or burrow in soft mud. Aquatic oligochaetes are
which causes them to retire quickly into their burrows. They an important food source for fishes. A few are ectoparasitic.
react to light, which they avoid unless it is very weak. Chemical Some of the more common freshwater oligochaetes are the
responses aid them in the choice of food. 1 mm long Aeolosoma (Gr. aiolos, quick-moving, soma, body)
Chemical as well as tactile responses are very important to (Figure 17.22B); the 10 to 25 mm long Stylaria (Gr. stylos, pillar)
earthworms. They not only must sample the organic content of (Figure 17.22A); the 5 to 10 mm long Dero (Gr. dere, neck or
soil to find food, but also must sense its texture, acidity, and throat) (Figure 17.22D). The common 30 to 40 mm long Tubifex
calcium content. (L. tubus, tube, faciens, to make or do) (Figure 17.22C) is
Experiments show that earthworms have some learning abil- reddish and lives with its head in mud at the bottom of ponds
ity. They can be taught to avoid an electric shock, and thus can and its tail waving in the water. Tubifex is an alternate host nec-
develop an association reflex. Darwin credited earthworms with a essary in the life cycle of Myxobolus cerebralis, a parasite that
great deal of intelligence because they pulled leaves into their bur- causes a very serious condition called whirling disease in rain-
rows by the narrow end, the easiest way for drawing a leaf-shaped bow trout in North America. Some oligochaetes, such as Aeolo-
object into a small hole. Darwin assumed that seizure of leaves by soma, may asexually form chains of zooids by transverse fission
worms did not result from random handling or from chance but (Figure 17.22B).
was deliberate. However, investigations since Darwin’s time have
shown that the process is mainly one of trial and error, for earth-
worms often seize a leaf several times before getting it right. Class Hirudinida: Leeches
Class Hirudinida is divided into three orders, Hirudinea, the “true”
Reproduction and Development Earthworms are monoe- leeches, and two others that merit mention here because their
cious (hermaphroditic); both male and female organs are found in members are morphological intermediates between oligochaetes
the same animal (see Figure 17.16B). In Lumbricus reproductive and true leeches (see Figure 17.1). Oligochaetes have variable
systems are found in segments 9 to 15. Two pairs of small testes numbers of segments, segments bear setae, and there are no
and two pairs of sperm funnels are surrounded by three pairs of suckers on the body. True leeches have 34 segments, entirely
large seminal vesicles. Immature sperm from the testes mature in lack setae, and possess anterior and posterior suckers. Members
seminal vesicles, then pass into sperm funnels and down sperm of order Acanthobdellida have 27 segments, bear setae on the
ducts to the male genital pores in segment 15, where they are first five segments, and have a posterior sucker. Members of
expelled during copulation. Eggs are discharged by a pair of order Branchiobdellida have 14 or 15 segments, no setae, and an
small ovaries into the coelomic cavity, where ciliated funnels of anterior sucker. Branchiobdellids are commensal or parasitic on
the oviducts carry them outside through female genital pores on crayfish. Hereafter, leech refers to members of order Hirudinea.
segment 14. Two pairs of seminal receptacles in segments 9 and Leeches occur predominantly in freshwater habitats, but a
10 receive and store sperm from the mate during copulation. few are marine, and some have even adapted to terrestrial life in
Reproduction in earthworms may occur throughout the year warm, moist places. They are more abundant in tropical coun-
as long as warm, moist weather prevails at night (Figure 17.21). tries than in temperate zones.
hic70049_ch17_362-383.indd 375 7/14/07 3:18:42 PM

Seminal Testis Sperm Seminal

vesicles (in red) receptacle
Clitellum Sperm Ovary
duct
A Sperm exchange (copulation)

in earthworms
Egg sac Eggs Oviduct Mating earthworms
MATING AND REPRODUCTION

B Deposition of eggs in a tough Clitellum IN EARTHWORMS
band that becomes a cocoon
Sperm are added
Seminal
receptacle
(with sperm)
C Fertilization
Fertilized eggs
D Cocoon slipping off

Worm
emerging
Figure 17.21 E Cocoon F

Earthworm copulation and formation of egg cocoons. A, Mutual insemination; sperm from genital pore (segment 15) pass along seminal grooves
to seminal receptacles (segments 9 and 10) of each mate. B and C, After worms separate, the clitellum secretes first a mucous tube and then a
tough band that forms a cocoon. The developing cocoon passes forward to receive eggs from oviducts and sperm from seminal receptacles. D, As
cocoon slips off over anterior end, its ends close and seal. E, Cocoon is deposited near burrow entrance. F, Young worms emerge in 2 to 3 weeks.
G, Two earthworms in copulation. Their anterior ends point in opposite directions as their ventral surfaces are held together by mucous bands
secreted by the clitella.
Most leeches are between 2 and 6 cm in length, but some, Form and Function Unlike other annelids, leeches have a
including “medicinal” leeches, reach 20 cm. The giant of all is the fixed number of segments but they appear to have many more
Amazonian Haementeria (Gr. haimateros, bloody) (Figure 17.23), because each segment is marked by transverse grooves to form
which reaches 30 cm. superficial rings (Figure 17.24).
Leeches are usually flattened dorsoventrally and exhibit a Unlike other annelids, leeches lack distinct coelomic com-
variety of patterns and colors: black, brown, red, or olive green. partments. In all but one species the septa have disappeared,
Many leeches live as carnivores on small invertebrates; some are and the coelomic cavity is filled with connective tissue and a
temporary parasites; and some are permanent parasites, never system of spaces called lacunae. The coelomic lacunae form a
leaving their host. Some leeches attack human beings and are a regular system of channels filled with coelomic fluid, which in
nuisance to outdoor enthusiasts. some leeches serves as an auxiliary circulatory system.
Like oligochaetes, leeches are hermaphroditic and have a Leeches are more highly specialized than oligochaetes.
clitellum, which appears only during breeding season. The clitel- They have lost the setae used by oligochaetes in locomotion
lum secretes a cocoon for reception of eggs. and have developed suckers for attachment while sucking
hic70049_ch17_362-383.indd 376 7/14/07 3:18:43 PM

Stylaria
B
C
Aeolosoma
Figure 17.22
Some freshwater oligochaetes. Tubifex
A, Stylaria has the prostomium
drawn out into a long snout. B,
Aeolosoma uses cilia around the
mouth to sweep in food particles,
and it buds off new individuals
D
asexually. C, Tubifex lives head
down in long tubes. D, Dero has
ciliated anal gills. Dero
blood (their gut is specialized for storage of large quantities

of blood). Most leeches crawl with looping movements of the
body, by attaching first one sucker and then the other and pull- Figure 17.23
ing the body along the surface. Aquatic leeches swim with a The world’s largest leech, Haementeria ghilianii, on the arm of Dr. Roy
graceful undulatory movement. K. Sawyer, who found it in French Guiana, South America.
Nutrition Leeches are popularly considered parasitic, but

many are predaceous. Most freshwater leeches are active pred-
For centuries “medicinal leeches” (Hirudo medicinalis) were used
ators or scavengers equipped with a proboscis that can be
for bloodletting because of the mistaken idea that a host of bodily
extended to ingest small invertebrates or to take blood from
disorders and fevers were caused by an excess of blood. A 10- to
cold-blooded vertebrates. Some can force their pharynx or
12-cm-long leech can extend to a much greater length when dis-
proboscis into soft tissues such as the gills of fish. Some ter-
tended with blood, and the amount of blood it can suck is consid-
restrial leeches feed on insect larvae, earthworms, and slugs,
erable. Leech collecting and leech culture in ponds were practiced
which they hold by an oral sucker while using a strong sucking
in Europe on a commercial scale during the nineteenth century.
pharynx to ingest food. Other terrestrial forms climb bushes
Wordsworth’s poem “The Leech-Gatherer” was based on this use
or trees to reach warm-blooded vertebrates such as birds or
of leeches.
mammals.
Leeches are once again being used medically. When fingers,
Most leeches are fluid feeders. Many prefer to feed on tis-
toes, or ears are severed, microsurgeons can reconnect arteries but
sue fluids and blood pumped from open wounds. Some fresh-
not all the more delicate veins. Leeches are used to relieve conges-
water leeches are true bloodsuckers, preying on cattle, horses,
tion until the veins can grow back into the healing appendage.
humans, and other mammals. True bloodsuckers, which include
the so-called medicinal leech, Hirudo medicinalis (L. hirudo, a
leech) (Figure 17.25), have cutting plates, or chitinous “jaws,” for
cutting through tough skin. Some parasitic leeches leave their Respiration and Excretion Gas exchange occurs only
hosts only during the breeding season, and certain fish parasites through the skin except in some fish leeches, which have gills.
are permanently parasitic, depositing their cocoons on their host There are 10 to 17 pairs of nephridia, in addition to coelomocytes
fish. However, even the true bloodsuckers rarely remain on the and certain other specialized cells that also may be involved in
host for a long period of time. excretory functions.
hic70049_ch17_362-383.indd 377 7/14/07 3:18:46 PM

Anterior sucker
Eyes
Mouth
Classification of Phylum Annelida
Nephridiopore
Classification of annelids is based primarily on the presence or
Proboscis absence of parapodia, setae, and other morphological features.
Because both oligochaetes and hirudineans (leeches) bear a
Salivary gland
clitellum, these two groups are often placed under the head-
Segment Male gonopore ing Clitellata (cli-tel-lata) and members are called clitellates.
Seminal vesicle Alternatively, because both Oligochaeta and Polychaeta possess
setae, some authorities place them together in a group called
Female gonopore
Chaetopoda (ke-topo-da) (N.L. chaeta, bristle, from Gr. chaitē,
Testis long hair, pous, podos, foot).
Ovary Class Polychaeta (pole-keta) (Gr. polys, many, chaitē,
long hair). Mostly marine; head distinct and bearing eyes and
Crop
tentacles; most segments with parapodia (lateral appendages)
bearing tufts of many setae; clitellum absent; sexes usually
Sperm duct separate; gonads transitory; asexual budding in some;
trochophore larva usually present. Examples: Nereis, Aphrodita,
Intestine Glycera, Arenicola, Chaetopterus, Amphitrite, Riftia.
Ceca Class Oligochaeta (oli-go-keta) (Gr. oligos, few, chaitē, long
Sensillae hair). Body with conspicuous segmentation; number of segments
variable; setae few per segment; no parapodia; head absent;
Anus
coelom spacious and usually divided by intersegmental septa;
Posterior Posterior hermaphroditic; development direct, no larva; chiefly terrestrial and
sucker sucker freshwater. Examples: Lumbricus, Stylaria, Aeolosoma, Tubifex.
A B Class Hirudinida (hiru-dini-da) (L. hirudo, leech, ida,
pl. suffix): leeches. Body with fixed number of segments
Figure 17.24 (normally 34; 15 or 27 in some groups) with many annuli; oral
Structure of a leech, Placobdella. A, External appearance, dorsal view. and posterior suckers usually present; clitellum present; no
B, Internal structure, ventral view. parapodia; setae absent (except in Acanthobdellida); coelom
closely packed with connective tissue and muscle; development
direct; hermaphroditic; terrestrial, freshwater, and marine.
Examples: Hirudo, Placobdella, Macrobdella.
Leeches are highly sensitive to stimuli associated with the presence

of a prey or host. They are attracted by and will attempt to attach
to an object smeared with appropriate host substances, such as fish
scales, oil secretions, or sweat. Those that feed on the blood of
mammals are attracted by warmth; terrestrial haemadipsids of the
tropics will converge on a person standing in one place.
Reproduction Leeches are hermaphroditic but cross-fertilize

during copulation. Sperm are transferred by a penis or by hypo-
dermic impregnation (a spermatophore is expelled from one
Figure 17.25 worm and penetrates the integument of the other). After copu-
Hirudo medicinalis feeding on blood from human arm.
lation their clitellum secretes a cocoon that receives eggs and
sperm. Leeches may bury their cocoons in mud, attach them to
Nervous and Sensory Systems Leeches have two “brains”: submerged objects, or, in terrestrial species, place them in damp
one is anterior and composed of six pairs of fused ganglia (form- soil. Development is similar to that of oligochaetes.
ing a ring around the pharynx), the other is posterior and com-
posed of seven pairs of fused ganglia. An additional 21 pairs of Circulation The coelom of leeches has been reduced by the
segmental ganglia occur along the double nerve cord. In addi- invasion of connective tissue and, in some, by a proliferation of
tion to free sensory nerve endings and photoreceptor cells in the chloragogen tissue, to a system of coelomic sinuses and channels.
epidermis, there is a row of sense organs, called sensillae, in Some orders of leeches retain a typical oligochaete circulatory sys-
the central annulus of each segment. Pigment-cup ocelli also are tem, and in these the coelomic sinuses act as an auxiliary blood-
present in many species. vascular system. In other orders the traditional blood vessels are
hic70049_ch17_362-383.indd 378 7/14/07 3:18:55 PM

lacking and the system of coelomic sinuses forms the only blood-
vascular system. In those orders contractions of certain longitudi-
nal channels provide propulsion for the blood (the equivalent of
coelomic fluid). Ciliated groove Proboscis
PHYLUM ECHIURA
Phylum Echiura (ek-ee-yur a) (Gr. echis, viper, serpent, Body
oura tail, ida, pl. suffix) consists of about 140 species of
marine worms that burrow into mud or sand, live in empty
snail shells or sand-dollar tests, or rocky crevices. They are
found in all oceans—most commonly in littoral zones of warm
waters—but some are found in polar waters or dredged from
depths of up to 10,000 m. They vary in length from a few mil-
limeters to 40 or 50 cm.
Echiurans are cylindrical and somewhat sausage-shaped
(Figure 17.26). Anterior to the mouth is a flattened, extensible Figure 17.27
proboscis, which cannot be retracted into the trunk. Echiurans Bonellia (phylum Echiura) is a detritus feeder. Lying in its burrow, it
are often called “spoon worms” because of the shape of the con- explores the surface with its long proboscis, which picks up organic
tracted proboscis in some species. The nervous system of echiu- particles and carries them along a ciliated groove to the mouth.
rans is fairly simple with a ventral nerve cord that runs the length
of the trunk and continues dorsally into the proboscis. The pro-
boscis has a ciliated groove leading to the mouth. While they lie short in some forms and long in others. Bonellia, which is only
buried, the proboscis can extend out over the mud for explora- 8 cm long, can extend its proboscis up to 2 m.
tion and deposit-feeding (Figure 17.27). Most species gather very One common form, Urechis (Gr. oura, tail, echis, viper,
small particles of detritus and move them along the proboscis serpent), has a very short proboscis and lives in a U-shaped bur-
by cilia; larger particles are moved by a combination of cilia and row in which it secretes a funnel-shaped mucous net. It pumps
muscular action or by muscular action alone. Unwanted particles water through the net, capturing bacteria and fine particulate
can be rejected along the route to the mouth. The proboscis is material in it. Urechis periodically swallows the food-laden net.
Lissomyema (Gr. lissos, smooth, mys, muscle) lives in empty
gastropod shells in which it constructs galleries irrigated by
Proboscis
Ciliated groove
("gutter")
Proboscis
Mouth
Seta Mouth
Pharynx
Nephridium
Anterior setae Dorsal
vessel
Ring
vessel
Ventral
nerve cord
Ventral
vessel Intestine
Posterior setae
Anal Gonad
A B vesicle
Figure 17.26 Cloaca

Caecum
A, Echiurus, an echiuran common on both Atlantic and Pacific coasts
of North America. B, Anelassorhynchus, an echiuran of the tropical
Pacific. The shape of their proboscis lends them the common name of Figure 17.28
“spoon worms.” Internal anatomy of an echiuran.
hic70049_ch17_362-383.indd 379 7/14/07 3:18:59 PM

rhythmical pumping of water and feeds on detritus and the Introvert

organic coating of sand and mud gathered by this process.
Cuticle and epithelium, which may be smooth or orna-
mented with papillae, cover the muscular body wall. There
may be a pair of anterior setae or a row of bristles around the
posterior end (see Figure 17.26). The coelom is large. The diges-
tive tract is long and coiled and terminates at the posterior end
(Figure 17.28). A pair of anal sacs may have an excretory and Trunk
osmoregulatory function. Most echiurans have a closed circula-
tory system with colorless blood but contain hemoglobin in coe-
lomic corpuscles and certain body cells. There are one to many
pairs of nephridia, which serve mainly as gonoducts in some
species. Gas exchange probably occurs primarily in the hindgut,
which is continually filled and emptied by cloacal irrigation.
In some species sexual dimorphism is pronounced, with the female

being much the larger of the two. Bonellia has an extreme sexual
dimorphism, and tiny males live on the body of the female or in A B
her nephridia. Determination of sex in Bonellia is very interesting.
Free-swimming larvae are sexually undifferentiated. Those that
Figure 17.29
Sipunculans. Themiste (A) and Phascolosoma (B) are both burrowing
settle on the proboscis of a female become males (1 to 3 mm long). genera of cosmopolitan distribution.
About 20 males are usually found in a single female. Larvae that
do not contact a female proboscis metamorphose into females. The
or detritivores, whereas others appear to be suspension feeders.
stimulus for development into males is apparently a hormone pro-
Some nutrition may also come from dissolved organic compounds
duced by a female’s proboscis.
directly from the water column. Undisturbed sipunculans usually
extend the anterior end from their burrow or hiding place and
Sexes are separate, with gonads being produced by special stretch out their tentacles to explore and to feed. Organic matter
regions of the peritoneum in each sex. Mature sex cells break collected in mucus on the tentacles is moved to the mouth by cili-
loose from these gonadal regions and leave the body cavity by ary action. The introvert is extended by hydrostatic pressure pro-
way of the nephridia. Fertilization is usually external. duced by contraction of body-wall muscles against the coelomic
Early cleavage and trochophore stages are very similar to fluid. The lumen of the hollow tentacles is not connected to the
those of annelids and sipunculans. The trochophore stage, which coelom but rather to one or two blind, tubular compensation sacs
may last from a few days to 3 months, according to species, is that lie along their esophagus (Figure 17.30). These sacs receive
followed by gradual metamorphosis to a wormlike adult. fluid from the tentacles when the introvert is retracted. Retraction
is effected by special retractor muscles. The surface of the intro-
vert is often rough because of surface spines, hooks, or papillae.
PHYLUM SIPUNCULA There is a large, fluid-filled coelom traversed by muscle and
Phylum Sipuncula (sigh-punkyu-la) (L. sipunculus, little siphon) connective-tissue fibers. Their digestive tract is a long tube that
consists of about 250 species of benthic marine worms, at depths doubles back on itself to form a U-shape and ends in an anus near
ranging from the intertidal to over 5000 m. They live seden- the base of the introvert (Figure 17.30). A pair of large nephridia
tary lives in burrows in mud or sand, occupy borrowed snail opens to the outside to expel waste-filled coelomic amebocytes;
shells, or live in coral crevices or among vegetation. Some spe- the nephridia also serve as gonoducts. Circulatory and respira-
cies construct their own rock burrows by chemical and perhaps tory systems are lacking, but coelomic fluid contains red cor-
mechanical means. More than half of the species are restricted puscles that have a respiratory pigment, hemerythrin, used in
to tropical zones. Some are tiny, slender worms, but the majority transportation of oxygen. Gas exchange appears to occur largely
range from 3 to 10 cm in length. Some are commonly known as across the tentacles and introvert. Their nervous system has a
“peanut worms” because, when disturbed, they can contract to a bilobed cerebral ganglion just behind the tentacles and a ventral
peanut shape (Figure 17.29). nerve cord extending the length of the body.
Sipunculans have no segmentation or setae. They are With only a few exceptions, sexes are separate. Permanent
most easily recognized by a slender retractile introvert, or gonads are lacking, and ovaries or testes develop seasonally in
proboscis, which is continually and rapidly being run in and out the connective tissue covering the origins of one or more of the
of the anterior end. Walls of the trunk are muscular. When the retractor muscles. Sex cells are released through the nephridia.
introvert is everted, the mouth can be seen at its tip surrounded The larval form is usually a trochophore. Asexual reproduction
by a crown of ciliated tentacles. Little is known about the details also occurs by transverse fission, the posterior one-fifth of the par-
of sipunculan feeding. Some species appear to be deposit feeders ent constricting off to become a new individual in some species.
hic70049_ch17_362-383.indd 380 7/14/07 3:19:03 PM

Mouth nervous and endocrine control. The coelom may have evolved
in response to different selective pressures in protostomes and
Tentacles deuterostomes.
Brain The origins of a metameric (segmented) body are at least
Esophagus
as puzzling as the origins of the coelom. True metamerism
occurs in annelids, arthropods, and chordates. The placement
Retractor muscle
of the annelids and arthropods in Protostomia and of the chor-
Compensation sac dates in Deuterostomia makes it unlikely that segmentation is
Nephridium homologous among these three taxa. Within the protostomes,
annelids are placed in clade Lophotrochozoa, whereas arthro-
Location of anus
pods are in clade Ecdysozoa. In both clades most phyla are
not segmented, again making it unlikely that members of these
Ventral
nerve cord two phyla inherited a segmented body plan from a common
ancestor. Annelids and molluscs have very similar develop-
Intestine
mental programs leading to a trochophore larva, but the anne-
lid trochophore develops a series of segments as it grows,
Longitudinal whereas the mollusc trochophore does not grow in this way
muscles (see discussion p. 336).
It is possible that all bilaterally symmetrical metazoans
shared a segmented ancestor and that segmentation genes
were suppressed in most lineages, but preliminary studies of
the details of how segments form (genetic control and chemi-
cal signaling) in different phyla do not support this hypothesis.2
Instead, current evidence supports the hypothesis that segmen-
Figure 17.30 tation arose independently multiple times.
Internal structure of Sipunculus.
The selective advantage of a segmented body for anne-
lids appears to lie in the efficiency of burrowing made pos-
sible by shape change in individual coelomic compartments of
EVOLUTIONARY SIGNIFICANCE the hydrostatic skeleton. However, this explanation cannot be
extended to the arthropods because, as Chapters 19, 20, and
OF METAMERISM 21 describe, the rigid exoskeleton of the arthropods prohib-
No truly satisfactory explanation has yet been given for the ori- its shape change among segments, and the coelom is small in
gins of segmentation and the coelom, although the subject has comparison to that of annelids. Clearly, there is much to learn
stimulated much speculation and debate. All classical explana- about metamerism.
tions have had important arguments leveled against them, and
more than one may be correct, or none, as suggested by R. B.
Clark.1 Clark stressed the functional and evolutionary signifi- PHYLOGENY AND ADAPTIVE
cance of these features to the earliest animals that possessed
them. He argued forcefully that the adaptive value of a coe- DIVERSIFICATION
lom was as a hydrostatic skeleton in a burrowing animal.
Thus contraction of muscles in one part of the animal could act
Phylogeny
antagonistically on muscles in another part by transmission of Annelids and molluscs share many developmental features,
the force of contraction through the enclosed constant volume so they were presumed by many biologists to be very closely
of fluid in the coelom. related, perhaps sister taxa. However, the shared developmental
Although the original function of the coelom may have features are likely to be retained ancestral features for lophotro-
served to facilitate burrowing in the substrate, certain other chozoan protostomes.
advantages accrued to its possessors. For example, coelomic Pogonophoran and vestimentiferan worms were once placed
fluid would have acted as a circulatory fluid for nutrients and outside phylum Annelida, but they have been reinterpreted as
wastes, making large numbers of flame cells distributed through- derived members of class Polychaeta and are now placed in
out the tissues unnecessary. Gametes could be stored in the clade Siboglinidae within this class. Only a small portion of the
spacious coelom for simultaneous release by all individuals in siboglinid body is segmented.
the population (thus enhancing chances of fertilization). Such a Two other groups of worms, sipunculids and echiu-
synchronous release of gametes would have selected for greater rans, are closely related to annelids according to phylogenies
using molecular characters. Some of these phylogenies place
1
Clark, R. B. 1964. Dynamics in metazoan evolution. The origin of the
2
coelom and segments. Oxford, U.K., Clarendon Press. Seaver, E. C. 2003. Int. J. Dev. Biol. 47:583–595.
hic70049_ch17_362-383.indd 381 7/14/07 3:19:05 PM

echiurans within Annelida, as derived polychaetes that have Adaptive Diversification

lost segmentation. There are serially repeated structures, such
Annelids are an ancient group that has undergone extensive
as nerve-cord ganglia and mucous glands in echiuran larvae,
adaptive diversification. The basic body structure, particularly
and serially repeated nephridia in echiuran adults. Some biol-
of polychaetes, lends itself to almost endless modification. As
ogists interpret these repeated structures as remnants from a
marine worms, polychaetes have a wide range of habitats.
segmented ancestor and place echiurans within Annelida. The A basic adaptive feature in evolution of annelids is their sep-
presence of paired epidermal setae in some echiuran species tal arrangement, resulting in fluid-filled coelomic compartments.
provides strong support for placing echiurans within Annelida. Fluid pressure in these compartments is used to create a hydro-
One recently developed phylogenetic tree placed echiurans near static skeleton, which in turn permits precise movements such
capitellid polychaetes; both taxa dwell in sediments. If this result as burrowing and swimming. Powerful circular and longitudinal
is supported by more studies, Echiura, like Pogonophora, may muscles can flex, shorten, and lengthen the body.
no longer be a valid phylum. Feeding adaptations show great variation, from the sucking
The placement of Sipuncula is more contentious than that pharynx of oligochaetes and the chitinous jaws of carnivorous
of Echiura. Sipunculans are not metameric and they do not have polychaetes to the specialized tentacles and radioles of particle
setae. Larval development is similar to that of annelids, molluscs, feeders. The evolution of a trophosome to house the chemo-
and echiurans. Further study, especially where molecular char- autotrophic bacteria that provide nutrients to siboglinids is an
acters are used, may clarify the position of these worms within adaptation to deep-sea life.
Lophotrochozoa. For the present, we depict them as the sister In polychaetes the parapodia have been adapted in many ways
taxon to a clade of annelids and echiurans. and for a variety of functions, chiefly locomotion and respiration.
Within phylum Annelida, class Polychaeta is a paraphyletic In leeches many adaptations (such as suckers, cutting jaws,
group because we have evidence that ancestral clitellates arose pumping pharynx, and distensible gut) relate to their predatory
from within Polychaeta. and bloodsucking habits.
SUMMARY
Phylum Annelida is a large, cosmopolitan group containing marine (compared with Polychaeta) and no parapodia. They have a closed
polychaetes, earthworms and freshwater oligochaetes, and leeches. circulatory system, and the dorsal blood vessel is the main pump-
Certainly the most important structural innovation underlying diver- ing organ. Paired nephridia occur in most segments. Earthworms
sification of this group is metamerism (segmentation), a division of contain the typical annelid nervous system: dorsal cerebral ganglia
the body into a series of similar segments, each of which contains a connected to a double, ventral nerve cord with segmental ganglia
repeated arrangement of many organs and systems. The coelom also running the length of the worm. Oligochaetes are hermaphroditic
is highly developed in annelids, and this, together with the septal and practice cross-fertilization. The clitellum plays an important
arrangement of fluid-filled compartments and a well-developed body- role in reproduction, including secretion of mucus to surround the
wall musculature, is an effective hydrostatic skeleton for precise bur- worms during copulation and secretion of a cocoon to receive
rowing and swimming movements. Further segmented specialization eggs and sperm and in which embryonation occurs. A small, juve-
occurs in arthropods, the subjects of Chapters 19, 20, and 21. nile worm hatches from the cocoon.
Polychaetes, the largest class of annelids, are mostly marine. Leeches (class Hirudinida) are mostly freshwater, although a
On each segment they have many setae, which are borne on few are marine and a few are terrestrial. They feed mostly on flu-
paired parapodia. Parapodia show a wide variety of adaptations ids; many are predators, some are temporary parasites, and a few
among polychaetes, including specialization for swimming, res- are permanent parasites. The hermaphroditic leeches reproduce in
piration, crawling, maintaining position in a burrow, pumping a fashion similar to that of oligochaetes, with cross-fertilization and
water through a burrow, and accessory feeding. Some polychaetes cocoon formation by the clitellum.
are mostly predaceous and have an eversible pharynx with jaws. Echiurans are burrowing marine worms, and most are deposit
Other polychaetes rarely leave the burrows or tubes in which feeders, with a proboscis anterior to their mouth. Some species bear
they live. Several types of deposit- and filter-feeding are known epidermal setae. They lack segmentation. The validity of this group
among members of this group. Polychaetes are dioecious, have a as a phylum is a subject of debate.
reproductive system lacking a clitellum, external fertilization, and Sipunculans are small, burrowing marine worms with an eversi-
a trochophore larva. ble introvert at their anterior end. The introvert bears tentacles used
Siboglinids live in tubes on the deep-ocean floor, and they are for deposit feeding. Sipunculans are not segmented.
metameric. They have no mouth or digestive tract but apparently Embryological evidence places annelids with molluscs and
absorb some nutrient by the crown of tentacles at their anterior end. arthropods in the Protostomia. Recent molecular evidence sug-
Much of their energy is due to chemoautotrophy of bacteria in their gests that annelids and molluscs are more closely related to each
trophosome. other (in Lophotrochozoa) than either phylum is to arthropods
Clade Clitellata encompasses class Oligochaeta and class (in Ecdysozoa). Echiurans are closely related to annelids and may
Hirudinida. Class Oligochaeta contains earthworms and many have arisin within this phylum. Sipunculans are also allied to
freshwater forms; they have a small number of setae per segment annelids, but also share certain features with molluscs.
hic70049_ch17_362-383.indd 382 7/20/07 3:46:29 PM

SELECTED REFERENCES
Childress, J. J., H. Felbeck, and G. N. Somero. 1987. Symbiosis in the This curious echiuran has detoxification bodies in its gut cells
deep sea. Sci. Am. 256:114–120 (May). The amazing story of how the and epithelial cells that allow it to live in a highly toxic sulfide
animals around deep-sea vents, including Riftia pachyptila, absorb environment.
hydrogen sulfide and transport it to their mutualistic bacteria. For most Mirsky, S. 2000. When good hippos go bad. Sci. Am. 282:28 (Jan.).
animals, hydrogen sulfide is highly toxic. Placobdelloides jaegerskioeldi is a parasitic leech that breeds only in
Cutler, E. B. 1995. The Sipuncula. Their systematics, biology, and evolution. the rectum of hippopotomuses.
Ithaca, New York, Cornell University Press. The author tried to “bring Patel, N. H. 2003. The ancestry of segmentation. Dev. Cell 5:2–4. Explores
together everything known about” sipunculans. the idea that segmentation is an ancestral feature of all bilaterally
Davis, G. K., and N. H. Patel. 2000. The origin and evolution of symmetrical animals.
segmentation. Trends Genet. 15:M68–M72. Discussion of segmentation Pernet, B. 2000. A scaleworm’s setal snorkel. Invert. Biol. 119:147–151.
with a focus on arthropods. Sthenelais berkeleyi is an apparently rare but large (20 cm) polychaete
Fischer, A., and U. Fischer. 1995. On the life-style and life-cycle of the that buries its body in sediment and communicates with water above
luminescent polychaete Odontosyllis enopla (Annelida: Polychaeta). just by its anterior end. Ciliary movement on parapodia pumps water
Invert. Biol. 114:236–247. If epitokes of this species survive their into the burrow for ventilation. The worm remains immobile for long
spawning swarm, they can return to a benthic existence. periods, except when prey comes near; it then rapidly everts its pharynx
Halanych, K. M., T. D. Dahlgren, and D. McHugh. 2002. Unsegmented to capture prey.
annelids? Possible origins of four lophotrochozoan worm Rouse, G. W. 2001. A cladistic analysis of Siboglinidae Caullery, 1914
taxa. Integ. and Comp. Biol. 42:678–684. A nice summary of (Polychaeta: Annelida): Formerly the phyla Pogonophora and
current morphological and molecular studies on classification of Vestimentifera. Zool. J. Linn. Soc. 132:55–80. Diagnostic features of
pogonophorans, echiurids, myzostomids, and sipunculans. Siboglinidae and its subgroups are provided.
Lent, C. M., and M. H. Dickinson. 1988. The neurobiology of feeding in Seaver, E. C. 2003. Segmentation: mono- or polyphyletic. Int. J. Dev. Biol.
leeches. Sci. Am. 258:98–103 (June). Feeding behavior in leeches is 47:583–595. Preliminary comparisons of the segmentation process
controlled by a single neurotransmitter (serotonin). in annelids, arthropods, and chordates suggest that annelids and
McClintock, J. 2001. Blood suckers. Discover 22:56–61 (Dec.). Describes arthropods do not share mechanisms of segmentation, but vertebrates
modern medical uses for leeches. and arthropods may share some mechanisms.
McHugh, D. 2000. Molecular phylogeny of Annelida. Can. J. Zool. 78:1873– Winnepenninckx, B. M. H., Y. Van de Peer, and T. Backeljau. 1998.
1884. Descriptions of monophyletic groups within Annelida supported Metazoan relationships on the basis of 18S rRNA sequences: A few
by molecular data. years later . . . Am. Zool. 38:888–906. Their calculations and analysis
Menon, J., and A. J. Arp. 1998. Ultrastructural evidence of detoxification support monophyly of Clitellata but cast doubt on monophyly of
in the alimentary canal of Urechis caupo. Invert. Biol. 117:307–317. Polychaeta.
hic70049_ch17_362-383.indd 383 7/14/07 3:19:08 PM

Annelids and Allied Taxa: Phylum Annelida, Including Pogonophorans (Siboglinids) Phylum Echiura Phylum Sipuncula

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Annelids and Allied Taxa: Phylum Annelida, Including Pogonophorans (Siboglinids) Phylum Echiura Phylum Sipuncula

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C H A P T E R

Chloeia sp., a polychaete.

Dividing the Body

hic70049_ch17_362-383.indd 362 7/14/07 3:18:00 PM

Lophotrochozoa (in part)

Acanthobdellida Branchiobdellida Hirudinea

15 segments 34 segments Proboscis Anterior

hic70049_ch17_362-383.indd 363 7/30/07 9:23:06 AM

hic70049_ch17_362-383.indd 364 7/14/07 3:18:06 PM

Aciculum Ventral Nephridium Nerve Ventral Epidermis

Longitudinal Dorsal blood Segment Intestine

hic70049_ch17_362-383.indd 365 7/14/07 3:18:06 PM

with jaws or teeth. They have an eversible, muscular pharynx

hic70049_ch17_362-383.indd 366 7/14/07 3:18:09 PM

Figure 17.6 Reproduction and Development

Nervous System and Sense Organs Representative Polychaetes

hic70049_ch17_362-383.indd 367 7/14/07 3:18:13 PM

Scale Worms Scale worms (Figure 17.9) are members of the

Fireworms Hermodice carunculata (Gr. herma, reef, dex,

hic70049_ch17_362-383.indd 368 7/14/07 3:18:18 PM

Proximal radiole sorting

deposit feeding. They live in a U-shaped burrow through which, by

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Water movement Clade Siboglinidae (Pogonophorans) Members of for-

Tentacles Ciliated band Papilla

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Sexes are separate, with a pair of

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Seminal receptacle Genital openings Clitellum Setae

B Left circumpharyngeal Seminal Sperm Egg funnel Nerve Subneural Ventral

Circular Epidermis Cuticle Dorsal vessel Mucous

Intestinal lumen Sensory fibers

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Posterior Lateral nerves Pharynx Cerebral ganglia

Nervous System and Sense Organs

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Seminal Testis Sperm Seminal

A Sperm exchange (copulation)

Egg sac Eggs Oviduct Mating earthworms

MATING AND REPRODUCTION

D Cocoon slipping off

Figure 17.21 E Cocoon F

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blood (their gut is specialized for storage of large quantities

Nutrition Leeches are popularly considered parasitic, but

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Leeches are highly sensitive to stimuli associated with the presence

Reproduction Leeches are hermaphroditic but cross-fertilize

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Figure 17.26 Cloaca

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rhythmical pumping of water and feeds on detritus and the Introvert

In some species sexual dimorphism is pronounced, with the female

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echiurans within Annelida, as derived polychaetes that have Adaptive Diversification