PENDIDIKAN KEWARGANEGARAAN

CRITICAL BOOK REPORT

INVERTEBRATE TAXONOMY

“ANNELIDA”

Disusun Oleh:

“KELOMPOK 4”

1. JOSEPHINE GLORIA (4193342001)

2. TERESIA OKARINA (4193342005)
3. FADILLAH RAHMA PURBA (4193342004)
4. GRESIA PALENTINA HUTAGAOL (4193342003)

PENDIDIKAN BIOLOGI

FAKULTAS MATEMATIKA DAN ILMU PENGETAHUAN ALAM

UNIVERSITAS NEGERI MEDAN

2020
 FOREWORD

Praise and gratitude to God Almighty because thanks to His blessings and grace the
writer can complete this task properly and on time. The author also thanks Mrs. Dra. Katrina
Samosir, M.Pd. for guiding us through this task.

The author hopes that this critical book report paper can be useful for readers. The
author also realizes that in this critical book report paper there are still errors. Therefore, we
hope that there will be constructive criticism and suggestions as evaluation materials in
making this assignment.

The author also apologizes if there are mistakes in the words of this critical book report paper
which are not pleasing to the readers.

Medan, October 2020

Author

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 TABLE OF CONTENTS

Foreword ...................................................................................................ii

Table of contents ...................................................................................................iii

Chapter I Introduction ...................................................................................................4

Background ………………...........................................................................4

Destination ………………...........................................................................4

Benefits ………………...........................................................................4

Chapter II. Discussion ...................................................................................................5

Book Identity ………………...........................................................................5

Summary of Book Contents ...........................................................................6

Strengths and Weaknesses of Book Contents..............................................................25

Chapter III. Closing ..................................................................................................26

Conclusion ..........................................................................26

Suggestion ..........................................................................26

Bibliography ..................................................................................................27

iii
 CHAPTER I

PRELIMINARY

A. Background

One of the learning strategies applied to students is the Critical Book Report. Literally,
Critical Book Report is an activity to criticize a book. However, Critical Book Report is not just
a report or writing about the contents of a book, but focuses more on evaluation (explanation,
interpretation and analysis) of the strengths and weaknesses of a book, highlighting the
interesting points of the book's discussion. In other words, through the Critical Book Report we
test someone's critical thinking ability and then rewrite it based on the perspectives, knowledge,
and experiences we have.

B. Purpose

The objectives of the preparation of the Critical Book Report include:

1. Knowing the strengths and weaknesses of a book

2. Adding broad insight into knowledge, especially in the evaluation section.

C. Benefits

The benefits of the Critical Book Report are:

1. Provide information about the identity of a book

2. Adding insight in analyzing a book
3. To add broad knowledge, especially about Annelida material
4. The writer can think more critically than he knows.
5. Readers can know that there are shortcomings and advantages of the book
criticized by the author.

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 CHAPTER II

DISCUSSION

I. BOOK IDENTITY

1) MAIN BOOK
BOOK TITLE INVERTEBRATES Third Edition
AUTHOR Brusca, Richard C. I Moore, Wendy. I
Shuster, Stephen M.
PUBLISHER Sinauer Associates, Inc.
CITY ISSUED Sunder and, Massachusetts USA
PUBLICATION 2016
YEAR
ISBN 978-160-535-375-3

2) BOOK OF COMPARISON

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 BOOK TITLE An Introduction to the Invertebrates
AUTHOR Janet Moore
PUBLISHER Cambridge University Press
CITY ISSUED New York
PUBLICATION 2006
YEAR
ISBN 978-0-511-24942-6

II. SUMMARY OF BOOK CONTENT

1) MAIN BOOK
BAB XIV.ANNELIDA
Characteristics of the Phylum Annelida 1. Blaterally symmetrica . vermiform,
protostomes. Terrestria , fresh water. and marine. 2. Segmented (segmentation secondarily lost
in some groups, e.g., S puncula, Echiuridae): segments arise by teob ast c growth. 3. Deve
opment typ cally protostomous. with hooblastic, spiral, 4d, sch zocoeous embryogeny. 4. D
gestve tract complete, usually with regonal special i zation (secondarily lost in some groups). 5.
With a closed c rcu atory system (secondarily lost in some groups); resp ratory pgments may

6
 incude hemog obin. chorocruorin and hemerythrn 6. Most adu ts possess metanephrid a or,
less commonly, protonephrid a. 7. Nervous system well developed, with a dorsa cerebral gangl i
on, c rcumenteric connectives. and ventral gangl onated nerve cord(s) . 8. With lateral,
segmentally arranged epidermal chaetae (secondarily lost in some, e.g., Spuncula). 9. In most.
head composed of presegmental prostomium and peristomium (often wth body segments
fused to it; peristomium may be reduced). Echuridae and S puncula without obvious
prostomum/peristomoium. 10. Gonochoristic or hermaphroditic; deve opment indirect or d
rect; often wtth trochophore arva.
Although now known to be highly modified annelids, these groups are so distinct that
they are given detailed treatment at the end of this chapter. The overall classification with.in
Annelida is also currently undergoing significant revisions and has yet to stabilize. Phylogenomic
and other nolecular phylogenetic analyses have shown that taxonomic groupings based on
morphology are, in many cases, invalid. Annelida was traditionally divided into three classes.
Polychaeta was the largest and most diverse group, the other two being Oligoghaeta
(earthworms and their relatives) and the leeches (Hirudinoidea or Hirudinea). It is now dear
that the closest relative of Hirudinoidea is the freshwater group Lumbriculidae, a taxon that is
inside the Oligochaeta, and these two former classes (Hirudinoidea and Oligochaeta) are now
referred to as Clitellata, a taxon that actually dates back to 1919. Furthermore, Clitellata has
been shown to have arisen deep within Polychaeta, n1aking "Polychaeta" synonyn1ous with
Annelida. Hence, all three of the former classes are no longer ranked as such, the Oligochaeta
and Polychaeta are recognized as paraphyletic groups and no longer hold systen,atic ranking,
and "oligod,aete" and "polycl,aete" are inforn1al names only. Owing to their novel morphology
and lifestyle, Hirudinoidea are also given some detailed treatment at the end of this chapter.

SYNOPSES OF MAJOR GROUPS

OWENIIDAE, Fewer than 50 species generally small-bodied (some reach ng 10 cm), tubedwelling
annelids with numerous fine parapodial hooks. M.AGELONIDAE Around 70 speces, most in the genus
Magelona,Reatvely similar in appearance, with shovel-shaped anter or ends and a pair of papillated
palps. Thin, cylindr ca l worms, less than 1 mm wide, but reaching 15 cm length. CHAETOPTERIOAE
Around 70 species, p aced mostly in the genera Chaetopterus, Mesochaetopterus, Phyllochaetopterus
or Spiochaetopterus ),adults range in size from less than 1 cm to more than 40 cm, though there are
fewer than 60 segments in most taxa. AMPHINOMIDA, Includes Amph nomidae and Euphrosin dae over
200 species,the more motile forms. amphinom ds. are often qu te large (e.g., Chloeia. Eurythoe,

7
Hermodice) and commony referred to as fire worms. SIPUNCULA Peanut worms. About 150 species, al l
marne ,former y w th the rank of phylum, phylogenetic studies have shown th s group beongs w thin
Annelida; with six famil es. Sipuncu ans range in ength from less than 1 cm to about 50 cm. ERRANTIA,
This is an o d taxonom c grouping that was recently resurrected by Torsten Struck and colleagues. It
contains more than a quarter of all descrbed anne l i d spec es diversity . PROTODRILIDA, Over 60
species, mainly l i ving interstit ally in sediments ,includes Protodrilidae, Protodr i l od dae, and Saccocirr
dae, best known for being members of the now defunct taxon Arch annel da. Taxa such as Protodrilus
and Saccocirrus are found in medium to coarse sed ments in shallow waters. EUNICIDAE uncdas a
welldel neated group of over 1,000 spec es that have a ventral muscu arized pharynx with comp ex jaws
that ncude e ements such as ventra mandibles and dorsal maxllae. Their parapod a are supported by
rodl ke chaetae called acicu ae and often bear compound chaetae. DORVILLEIDAE,178 species, Includes
some of the smallest anne ids (Neotenotrocha). whil e others may reach severa l cent meters and have a
arge number of segments (e.g., Dorvil/ea). The head often has three antennae and there can be a pair of
palps that may either resemble or d ffer from the shape of the antennae. LUMBRINERIDAE 275 spec ies
thin and eongate and, except for Lysarete and Kuwaita; lack ng head appendages. Most crawl about in a
gal mats and ho dfasts, and small cracks in hard substrata; some burrow in sand or mud. OENONIDAE 87
species. Elongate, with small parapod a; lack ng head appendages. or w th three small antennae.
Resembling Lumbr neridae. though their jaws and chaetae d ffer. Often found n soft substrata.
Phyllodocida. with more than 4,600 species, is dist nguished by its members having an ax al muscular
probosc s (often armed w th two or more Jaws). They tend to show fus on of some anter or segments w
th the head. often on y identifiable by the retenton of anterior en arged c rri. and the head usually has
two or three antennae and a pa r of sensory palps. APHRODITIFORMA, Scale worms compr sea d everse
group of over 1,000 spec es in seven (or eight) fam lyranked taxa,acoetidae, Aphrodtidae. Eulepethdae,
Po ynodae (conta ns most spec es), lph ondae, Sgalion dae, and poss bly Pholo dae. Aphrod tforma a so
ncludes the scaeless scale worms of the former fam ly Ps ondae. Most are re at vely short and somewhat
flattened dorsoventrally. CHRYSOPET ALIDAE 135 species,the name (Lat n for "goden petals") refers to
the shape and co or of the go den, flattened notochaetae (called pa eae) that cover the dorsal surface in
many spec es (e.g., Chrysopeta/um). These are small to moderately s zed worms, wth adu ts varying in
length from 1 to 50 mm, and with as few as 10 to over 300 segments. GLYCERIDAE , 89 species,Cyindrica
tapered, homonomous body, usually red or p nk, reaching 30 cm n length. Enormous evers b e pharynx
that can be retracted into 1 /3 of the body length, armed with four hook-like jaws used in prey capture.
HESIONIDAE, 172 species,generally beaut ful worms, adults measuring from a few millimeters length to
more than 10 cm. Subtida , especially on rocky and mixed bottoms, and increasing y known from
hydrothermal vents and methane seeps. NEPHTYIDAE, 142 specis,usually long and sender, with well-
deve oped parapodia but s mp e heads. They can be very abundant in shallowwater sedments, and there
are often a number of d fferent taxa in a sngle sedment samp e. Wh le they are easly dentif ed to this
family eve. spec es ident ficat on is difficu t. Adu ts range from a few millimeters to 30-40 cm and up to
150 segments. NEREIDIDAE, 691 species ,among the best-known marine annelids and w dely used in
teach ng, aboratories, and as fishing ba t. Most nereid ds are found in shallow waters, though some are
found at hydrothermal vents. Small to very arge (over 100 cm) w th homonomous segments.
PHYLLODOCIDAE, 417 species,then, often elongate (over 50 cm) bodies of up to 700 homonomous

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segments; common y active ep benthc predators on sol i d substrata; a few burrow n mud (e.g., Eteone.
Eulalia, Notophyllum, Phyllodoce).

The Annelid Body Plan

The annelid body is comprised of four regions: a presegmental region derived from the larval
episphere, the prototroch region around the mouth, the serially repeated body segn1ents, and the
posterior pygid.iun1. The episphere (see the section on Reproduction and Development) becon1es the
presegmental prostomium, and the prototroch and buccaJ region give rise to the peristomium, the
region surrounding the mouth. The body segmentation of annelids is referred to as metamerism. The
extreme posterior end of the body is the pygidium and it bears the anus and often some cirri. As with
the prostomium, the pygid.ium is non-seginental and may well contain renu1ants of the larval body.
Fron1 this basic scheme, the tremendous diversity of body forms seen in annelids is built. The majority
of annelids show metamerism and a greatly elongate cylindrical body. They are triploblastic, have a
through (complete) gut, and a spacious coelom. The sin1plest annelid head is composed of a
prostonlitlll1 and a peristomium. The segmentation of the main body is usually v isible externally as
rings, or annuli, and is reflected internally by the serial arrangement of coelomjc compartments
separated from one another by intersegmental septa. Body Forms Most annelids are marine, living in
habitats ranging from the intertidal zone to extreme depths. But quite a few inhabit brackish or fresh
later, and the majority (C) Musculature of intestine Longitudinal muscles Dorsal vessel of Clitellata
(thousands of species) live on land. Annelids range in length from less than 0.5 mm as adults for some
interstitial species to over 3 m for sone giant eunicids and megascolecid clitellates (earthworms). The
myriad variations ill body form among aimelids can best be described relative to the basic annelid
regions of a head, segmented trunk, and pygidium. Note though, that sonle groups such as Echiuridae,
Hirudinea, Siboglinidae, and Sipuncula are so transformed from this condition, each it their own
interesting ways, that they are treated as special sections near the end of this chapter. Most annelids
are gonochoristic and proliferate the gametes from the peritoneum .into the coeloin here they develop.
Many annelids have broadcast spawning and produce unming larvae, but there is a great variety of life
history strategies in this phylum, with a wide range of kinds of parental care also present. Notably, any
annelids are hermaphrodites, induding all members of Clitellata, which typically exchange sperm and
produce brooded or encapsulated embryos that develop directly to juveniles.

Body Wall and Coelomic Arrangement

The annelid body is covered by a thin cuticle, which is composed of scleroprotein and
mucopolysaccharide fibers deposited by epidermal cellular n1icrovilli. The epidermis is a colunmar
epitheliwn that is often ciliated on certain parts of tl1e body. Beneath the epidermis lies a layer of
connective tissue, circular muscles (sometimes absent) and thick longitudinal muscles, the latter often
arranged as four bands. The circular muscles do not form a continuous sheath, but are interrupted at
least at the positions of the parapodia. The inner lining of the body wall is the peritoneum, which
surrounds the coelomic spaces and lines the surfaces of internal organs. The coelom of annelids with
ho1nonomous bodies is generally arranged as laterally paired. In addition to the main body wall and
septa I muscles, other muscles function to retract protrusible or eversible body parts (e.g., branchiae,

9
pharynx) and to operate the parapodi. Each parapodium is an evagination of the body v-,au and contains
a variety of muscles. Movable parapodia are operated primarily by sets of diagonal (oblique) muscles,
which have their origin near the ventral body midline.

Support and Locomotion

Annelids provide a great example of the employment of coelomic spaces as a hydrostatic

skeleton for body support. Coupled with the well-developed musculature, the metameric body, and the
parapodia, this hydrostatic quality provides the basis for understanding locon1otion in these vvorms.
The genera Nereis and Plntynereis (Phyllodocida, Nereididae) are errant, homonomous annelids that
show a variety of locomotory patterns that are worth describing. ln such annelids the intersegmental
septa are functionally complete, and thus the coelomic spaces in each segment can be effectively
isolated hydraulically from each other. Modifications on this fundamental arrangement are discussed
later. In addition to burrowing (see below), Nereis can engage in three basic epibenthic locomotory
patterns: slow crawling, rapid crawling, and swimming. The circular muscles are relatively thin and serve
primarily to maintain adequate hydrostatic pressure within the coelomic compartments. Each method of
locomotion in Nereis (and similar forms) in· volves the antagonistic action of the longitudinal muscles on
opposite sides of the body in each segment. During movement, the longitudinal muscles on one side of
any given segn1ent alternately contract and relax (and are stretched) in opposing synchrony with the
action of the muscles on the other side of the seg· ment. Thus, the body is thrown into undulations that
move in metachronal waves from posterior to anterior. Variations in the length and amplitude of these
waves combine with parapodial movements to produce the different patterns of locomotion. The
parapodia and their chaetae are extended maximally in a power stroke as they pass along the crest of
each metachronal wave. Conversely, the parapodia and chaetae retract in the wave troughs during their
recovery stroke. Thus, the parapodia on opposite sides of any given segment are exactly out of phase
with one another. In contrast to the above descriptions, scale worms (Polynoidae; cover photo on book)
have capitalized on the use of their muscular parapodia as efficient walking devices. The body undulates
little if at all, and tllere is a corresponding reduction in the size of the longitudinal muscle bands and
their importance in locomotion. These worms depend almost entirely on the action of the parapodia for
walking and most adult polynoids cannot swim, except for very short bursts. Nereis is also a burrower
and studies on N. virens showed that they can extend their burrows in muddy sediments by using
mechanically efficient fracturing style of locomotion called "crack propagation." This is an efficient use of
energy as the worm forces its way through the mud, and it involves everting the pharynx to apply
dorsoventral forces to burrow ,-valls that are amplified at the burrow tip making a crack in a forward
direction. The ·1-vonn then anchors itself by expanding the body laterally and pushing its narrow head
into the crack, whereupon it repeats the process. Most tube-dwelling annelids are heteronomous and
many, such as Terebelliformia, have rather soft bodies and relatively weak muscles. The parapodia are
reduced, so the chaetae are used to position and anchor the animal in its tube. Movement within the
tube is usually accomplished by slow peristaltic action of the body or by chaetae n1oven1ents. When the
anterior end is extended for feeding, it may be quickly withdrawn by special retractor n1uscles while the
unexposed portion of the body is anchored in the tube. Annelid tubes provide protection as well as
support for these soft-bodied worms, and also keep the animal oriented properly in relation to the

10
substratum. Some annelids build tubes composed entirely of their ovvn secretions. Most notable
an1ong these tube builders are the serpulids, which construct their tubes of calcium carbonate secreted
by a pair of large glands near a fold of the peristomium called the collar. The crystals of calcium
carbonate are added to an organic matrix; the mixture is molded to the top of the tube by the collar fold
and held in place until it hardens. Sabellids, close relatives of serpulids, produce parchn1ent-like or
membranous tubes of organic secretions n1olded by the collar. Some, such as Snbelln, nl.ix n1ucous
secretions with size-selected particles extracted from feeding currents, then lay down the tube with this
material. A few annelids are able to excavate burrows by boring into calcareous substrata, such as rocks,
coral skeletons, or mollusc shells (e.g., certain members of the families Cirratulidae, Eunicidae,
Spionidae, Sabellidae). In extreme situations, the activity of the annelids may have deleterious effects on
the "host." For example, a boring sabellid, Terebrnsabella, which lives in abalone shells, can cause
fatalities by making them deform their shells. Species of Polydora (Spionidae) often excavate galleries in
various calcareous substrata (e.g., shells) and have been responsible for killing oysters in comn,ercially
ha.tvested areas of Europe, Australia, and North America.

Feeding and Digestion

Feeding The great diversity of form and function an1ong annelids has allo,,ved them to exploit
nearly all marine food resources in one way or another, and to be critical ecosystem components of
most terrestrial soils. For convenience we have categorized annelids as raptorial, deposit, and
suspension feeders (see Chapter 4). However, there are several feeding methods and dietary
preferences ,,vi thin each of these basic designations. Following a discussion of selected exan1ples of
these feeding types, we mention a few of the sy1nbiotic annelids. The most familiar raptorial annelids
are hunting predators belonging to the clade Errantia (e.g., many phyllodocids, syUids, nereidids and
eunic.ids, all part of Aciculata). These animals tend toward homonomy and are capable of rapid
movement across the substratum. For the most part they feed on small invertebrates. When prey is
located by chemical or n1echanical means, the worm everts its pharynx by quick contractions of the
body wall n1uscles in the anterior segments, increasing the hydrostatic pressure in the coelomic spaces
and causing the eversion. As a result of the design of the pharynx, the jaws (if present) gape at the
anteriorn1ost end when the pharynx is everted Raptorial feeding also occurs in some freshwater
clitellates such as Lumbriculidae (e.g., Phagodrilus), which capture prey (often other clitellates!) with
their muscular pharynx. A number of annelids are deposit feeders that are relatively unselective, simply
ingesting the substratum and digesting the organic matter contained therein (e.g., members of
Arenicolidae, Opheliidae, Maldanidae, and n1any clitellates). Lug,vorms, such as Arenicola, excavate an
L-shaped burrow, which they irrigate with water drawn into the open end by peristaltic movements of
the worm's body. Selective deposit feeders are defined by their ability to effectively sort the organic
material from the sediment prior to ingestion (e.g., many species of Terebellidae, Spionidae, and
Pectinariidae). However, (G) everted. (G) The giant (1 to 3 m) tropical Bobbit worm, Eunice aphroditois,
with its head extended from the sediment at night. The jaws are locked open while it awaits a pass ng
victim, typically a fish. When the antennae or paps detect the prey, the jaws snap shut and it draws the
victim into its burrow. the methods used to sort food ruffers among these groups. Most terebellids (e.g.,
Aphitrite, Pistn, Terebelln) establish themselves in shallow burrows or permanent tubes.

11
 Symbiotic relationships with other animals occur among several groups of annelids. There are
some interesting cases that reflect, again, the adaptive diversity of these vorms. Many symbiotic
annelids are hardly modified from thejr free-living counterparts and do not show the drastic adaptive
cl1aracteristics often associated with this sort of life. For many, the relationship with their host is a loose
one, the annelid often using the host merely as a protective refuge. We have already mentioned
annelids that burrow into the shells of other invertebrates that are quite similar to their free-living
relatives. Among the most commonly found commensaJ annelids are polynoid scale worms, especiaJJy
n,embers of the genera Arctonoe, Halosydnn, and Mnllicephnla, which live on the bodies of various
molluscs, echinoderms, and cnidarians. A most unusual symbiotic relationship exists between the
strange ampharetid Pompeii worm (Alvine/In pompejnnn; named after the deep-sea submersible Alvin)
and a variety of n,arine chen,oautotrophic sulfur bacteria. Alvine/la is a notable member of deep
hydrothermal vent communities of the East Pacific Rise.

Circulation and Gas Exchange

Given the relatively large size of many anneJjds, the compartmentalization of their coelomic
chambers, and the fact that only certain portions of tl,eir gut absorb digested food products, it is
essential that a circulatory mechanism be present for internal transport and distribution of nutrients.
Furthermore, many annelids have their gas exchange structures limited to particular body regions; thus
they depend on the circulatory system for internal transport of gases. It is easiest to understand the
circulatory system of annelids by considering it in concert with their gas exchange structures, which are
remarkably varied. In many annelids that lack appendages, the entire body surface functions in gas
exchange (e.g., lumbrinerids, oenonids). Some of the active epibenthic forms utiJjze hjghly vascul.arized
portions of the parapodia as gills. Special gas exchange structures, or branchiae, are fow1d in the form
of trunk filrunents (cirratulids, orbiniids), anterior gills (ampharetids, terebellids), and tentacuJar, or
branchial, crowns on the head (sabellids, serpulids, and siboglinids). Since the blood generally carries
respiratory pigments, the anaton1y of the circulatory system has evolved along with the structure and
location of these gas exchange structures. We again begin our examination witl, a hon1onon1ous
a1melid, such as Nereis, in which the parapodia are more or less similar to one another and the notopod
ia function as gills. The major blood vessels include a middorsal longitudinal vessel, which carries blood
anteriorly, and a rnidventral vessel, which carries blood posteriorly. Exchange of blood between these
vessels occurs through posterior and anterior vascular networks and serially arranged segmental vessels.

The movement of blood in Nereis depends on the action of the body wall muscles and on
intrinsic muscles in the walls of the blood vessels, especially the large dorsal vessel. There are no special
hearts or other pun1ping organs. The blood passing through the various segmental vessels supplies the
body wall muscles, gut, nephridia, and parapodia. Note that the oxygenated blood is being returned to
the dorsal vessel, thus maintaining a primary supply of oxygen to the anterior end of the ani mal,
including the feeding apparatus and cerebral ganglion. In many clitellates, three main longitudinal blood
vessels extend most of the body length and are connected to one another in each segment by additional
segmentally arranged vessels.

Reproduction and Development

12
 Regeneration and asexual reproduction Annelids show various degrees of regenerative
capabilities. Nearly all of them are capable of regenerating lost appendages such as palps, tentacles,
cirri, and parapodia. Most of them can also regenerate posterior body segments if the trunk is severed.
There are numerous exceptional cases of the regenerative powers of annelids. While regeneration of
the posterior end is common, most carmot regenerate lost heads. However, sabellids, syllids, and so me
others can regrow the anterior end. The most dramatic regenerative powers among the annelids occur,
oddly, in a few forms with highly specialized and heteronomous bodies. In Cliaetopterns, for example,
the anterior end will regenerate a nonnal posterior end as long as the regenerating part (the anterior
end) includes not more than fourteen segments; if the anin1al is cut behind the fourteenth segment,
regeneration does not occur. Furthermore, any single segment from among the first fourteen can
regenerate anteriorly and posteriorly to produce a complete worm. An even more dran1atic example of
regenerative power is known among certain species of Dodecaceria (Ci.rratulidae), ,,vhich are capable of
fragmenting their bodies into individual segments, each of which can regenerate a complete individual!
A similar situation is seen in another cirratulid, Ctenodrins, where segments transform into a series of
heads that then proliferate further segments behind, resulting in a chain of individuals that eventually
separates. Such clonal reproduction has been maintained by Ctemmodrins in culture for decades
without any instances of sexual reproduction. Regeneration appears to be controlled by neuroendocrine
secretions released by the central nervous syste1n at sites of regrowth. It is initiated by severing the
elements of the nervous system. Initiation has been demonstrated experimentally by cutting the ventral
nerve cord while leaving the body intact; the result is the formation of an extra part at the site of cutting
(e.g., two "tails"). The actual med1ruusm of regeneration has been studied in a variety of annelids and,
although the results are not entirely consistent, a general scenario can be outUned. Normal growth and
addition of segments (in young worms) take place immediately anterior to the pygidium, in a region
known as the growth zone. However, this growth zone is obviously not involved in regeneration. Rather,
when the trunk is severed, the cut region heals over and then a patch of generative tissue, or blastema,
forms. The blastema comprises an inner mass of cells originating from nearby tissues that were derived
originally from me soderm, and an outer covering of cells from ectodern1ally derived tissues such as the
epidermis. These two cell masses act somewhat as a growth zone analogue, proliferating new body
parts according to their tissue origin.

Sipuncula: The Peanut Worms

In the past, the coelomate worm phyla Sipuncula and Echiura Viere often dismissed in short
fashion as "minor" or "lesser" groups. However, thanks to recent molecular phylogenetic research, we
now kno"' that Caudal cirrus these two groups are remarkably modified spiralian clades embedded
within the phylu.n1 Annelida. Sipunculans and eclliurans resemble one another in several respects,
although they are not closely related, and they are often found in similar habitats. For these reasons,
biologists who find interest in one of these groups often study both. Sipunculans are never as abundant
or import.ant ecologically as some other wonns, especial!y the polycllaetes and nematodes.
Nonetheless, they display body plans that are different from any we have discussed so far and provide
important lessons in functional morphology and thus deserve special attention. The clade Sipuncula

13
(Greek, "little tube") includes about 150 species in 16 genera and 6 families. Usually called "peanut
worms," adult sipunculans show no evidence of segmentation or chaetae (two features viewed as
characteristic of annelids). The body is sausage-shaped and divisible into a retractable introvert and a
thicker trunk.

Sipunculan Classification

The first published illustrations of sipunculans were produced from woodcuts made in the mid-
sixteenth century. Linnaeus included these animals in the twelfth edition of his Systemn Nnturne (1767)
and placed them in the Vermes, along with so many other odds and ends. In the nineteenth century,
Lamarck and Cuvier considered the sipunculans to be relatives of holothuroid echinoderms (sea
cucumbers). No separate taxon was established for these worms until 1828, when Henri Marie Ducrotay
de Blainville introduced the name Sipunculida and allied the group with certain parasitic helnlinths. Tn
1847 Jean Louis Armand de Quatrefages invented the group Gephyrea to include sipunculans, echiurans,
and priapulans. The Greek root gephyra means "bridge," as Quatrefages regarded these animals as
intermediate between annelids and echinoderms. Antillesomatidae Phascolosomatidae
Aspidosiphonidae Phylogenetic relationships of the sipunculan families. The gephyrean concept was
founded on superficial characteristics, but it persisted well into the twentieth century even though many
authors attempted to raise the constituent groups to individual phylum status. Finally, Libbie Hyman
(1959), recognizing the polyphyletic nature of the Gephyrea, elevated the sipunculans to a separate
phylum rank; her view was quickly accepted. At that time, however, no classes, orders, or families v.•ere
recognized, and the phylum was divided into only genera and species. The Herculean effort by
Alexander Stephen and Stanley Edmonds (1972) and subsequent modifications by other 1-vorkers (e.g.,
Mary Rice 1982) led to a classification comprising 4 families and 16 genera. Later, Edward Cutler and
Peter Gibbs (1985) applied modern phylogenetic methods to the sipunculans and produced a
classification scheme of 2 classes, 4 orders, 6 families, and 17 genera, which has been used until
recently. However, with the use of molecular phylogenetics, this classification was challenged, the
phylum was subsumed within Annelida, and a new system, consisting of 6 families and 16 genera has
been proposed.

The Sipunculan Body Plan Body Wall, Coelom, Circulation, and Gas Exchange

The sipunculan body s urface is covered by a well-developed cuticle that varies fron1 thin on
the tentacles to quite thick and layered over much of the trunk. The cuticle often bears papillae, yarts, or
spines of various shapes. Beneath the cuticle lies the epiderntls, the cells of which are cuboidaJ over
most of the body but grade to columnar and ciliated on the tentacles. The epidermis contains a variety
of unicellular and multicellular glands called epidermal organs, some of wltlch project into the cuticle
and produce some of the surface papillae or knobs. Some of these glands are associated with sensory
nerve endings; others are responsible for producing the cuticle or for mucus secretion. In both larval and
adult sipunculans, epidermal organs are externally encapsulated by a c uticle and internally delimited by

14
regular epidermal cells. If pocketed deeply into the body ball, epidermal orgai1s are also delin1.ited by
subepidermal musculature.

The fluid of the body cavity contains a variety of cells and other inclusions. There are both
granular and agranular amebocytes of uncertain function, and red blood cells containing hemerythrin.
Also contained in the coelomic fluid are unique and fascinating multicellular structures called urns, some
of which are fixed to the peritoneum and some of which swim free in the fluid. The urns accu1nulate
waste materials and dead cells by trapping them with cilia and mucus. Gas exchange apparently varies
an1ong species. It has been suggested that rock borers and those that burrow in sediments of low
oxygen content (e.g., Themiste) exchange gases largely across the tentacles, which extend into the
overlying water. Other burrowers with long introverts may use the body surface of both the tentacles
and the introvert for gas exchange, whereas others, such as the large-bodied species of the genus, may
use the entire body surface. The coelomic fluid in the body cavity and in body wall channels provides a
circulatory medium, aided by diffusion and body movements. The erythrocytes that circulate in the
coelomic fluid of sipunculans contain . intracellular polymeric hemerythrin for storing and transporting
oxygen.

Support and Locomotion

Sipunculans are sedentary creatures. The general body shape is maintained by the muscles of
the body wall and the hydrostatic skeleton established by the large coelom. The body is essentially a
fluid-filled bag of constant volume, so any constriction at one point is accompanied by an expansion at
another. Burrowing in soft substrata is accomplished by peristalsis, driven by the circular and
longitudinal muscles of the body wall and by the action of the introvert. Movement through algal
holdfasts and bottom rubble occurs in a similar manner. Some species with an anterior cuticular shield
burrow into hard substrata and use the shield as a functional operculum to close the burrow entrance.
Burrowing into hard substrata is probably accomplished by both mechanical and chemical means, the
former using cuticular structures (such as spines and the posterior shield) as rasps, the latter facilitated
by the secretions of epidermal glands. The introvert is extended when the coelomic pressure is
increased by contracting the circular muscles of the body wall. Withdrawal is accomplished by the
retractor muscles, which pull from the mouth end, turning the introvert inward as the body wall muscles
relax. When the mtrovert is fully extended, the tentacles are erected by increasing the presslLre on the
compensation sacs. Sipunculans are highly tactile and strongly thigmotactic, requiring contact with their
surroundings. Placed alone in a glass dish, they are rather inactive except for rolling the introvert in and
out. However, if several sipunculans are placed together or with small stones or shell fragments, they
soon respond by makmg contact with eacli other or surrounding objects.

Feeding and Digestion

There is surprisingly little information on the details of sipw1culan feeding n1echanisms. Indirect
evidence from anatomy, gut contents, and general behavior suggests that these animals use different
feeding methods in different habitats. Most of the sipunculans that can place their tentacles at a
substratum-water interface are selective or nonselective detritivores (e.g., shallow burrowers, algal

15
holdfast dwellers); they use the mucus and cilia on the tentacles to obtain food. Deeper burrowers in
sand are direct deposit feeders. Some appear to be ciliary-mucus suspension feeders, using the
tentacles to extract organic material £ron1 the water. Sipw1culans that burrow in calcareous substrata
use spines or hooks on their introverts to retrieve organic detritus within reach and ingest the material
by retracting the introvert. Limited data suggest that at least some sipunculans take up dissolved organic
compounds directly across the body wall. Some workers have speculated that up to 10% of these
animals' nutritional requirements 1nay be met in this fashion.

Excretion and Osmoregulation

Most sipunculans possess one pair of elongate, saclike, tubular metanephridia located
ventrolaterally at the anterior end of the trunk. Two genera (Onchnesonm and Phnscolion) have but a
single nephridium. Species in these genera tend to be asymmetrically coiled. The nephridiopores are
located ventrally on the anterior region of the h·unk. The nephrostome Lies close to the body wall, near
the pore, and leads to a large nephridial sac that extends posteriorly in the trunk. Sipunculans are
basically os1noconformers and they are unknown from fresh and brackish-water habitats. Under normal
conditions, the coelomic fluid is nearly isotonic to the surrounding seawater. However, when placed in
hypotonic or hypertonic environments, the body volume increases or decreases, respectively.
Interestingly, the rates of volume change differ when the animal is exposed to these opposing
environments, suggesting that sipunculans are better at preventing water loss than at preventing water
gain. This situation may be due to a differential permeability of the cuticle, or perhaps to some active
mechanism of the nephridia. In any case, sipunculans rarely face severe osmotic problems in their usual
environments, and even in laboratory experiments they are able to recover nicely from most conditions
of osmotic stress.

Nervous System and Sense Organs

The general struchue of the sipunculan nervous system is similar in many respects to that in
other annelids. A bilobed cerebral ganglion lies dorsally in the introvert, just behind the mouth.
Circumenteric connectives extend fron1 the cerebral ganglion to a ventral nerve cord running along the
body wall th.rough the introvert and truk. The adult ventral nerve cord is single, and there is no
evidence of segmental ganglia. In Plznscolosoma ngassizii, a double ventral nerve cord forms initially,
but later fuses into a single cord, and neurogenesis initially follows a segmental pattern similar to that of
armelids. Starting out with paired FMRFamidergic and serotonergic axons, four pairs of associated
serotonergic perikarya and interco1u1ecti.ng commissures form one after an other in an anterior-
posterior progression. In late-stage larvae, the two serotonergic axons of the VNCs fuse, the
comm.issures disappear, and one additional pair of perikarya is formed. These cells (ten in total) migrate
toward one another, eventually forming two clusters of five ceUs each. These neural-remodeling
processes result in the single nonmetameric central nervous system of the adult sipunculan. This
ontogenetic example suggests that the ancestral sipunculan condition may have been a double ventral
nerve cord like that found in some primitive annelids. Lateral nerves arise fron1 the nerve cord and
extend to the body "'all muscles and sensory receptors in the epidermis.

16
Reproduction and Development

Sipunculans possess reasonable powers of regeneration. Most species are able to regrow lost
parts of the tentacles and even the introvert, and some can regenerate portions of the trunk and the
digestive tract. It "'as long believed that sipunculans could not reproduce asexually. However, in the
1970s it was discovered that at least some species do possess this capacity. The process takes place by
transverse fission of the body, whereby the worm divides into a small posterior fragment and a larger
anterior portion. Both portions then regrow the missing parts. Regeneration from the small posterior
part is quite remarkable, smce most of the trunk, anterior gut, retractor muscles, nephridia, introvert,
and so on must be regrown. Except for Golftngin mi,zuta, sipunculans are gonocl1oristic. (Facultative
parthenogenesis has been reported in one species, The,niste lngenifonnis.) The gametes arise from the
coelonlic Uning, often near the origins of the retractor muscles. Gametes are released into the coelom,
where they mature. Ripe eggs and sperm are picked up selectively by the nephridia and stored in the
sacs until released. The eggs are encased in a layered, porous covering. Males spawn first, probably il1
response to some environmental cue, and the presence of sperm in the water stimulates females to
spawn. Following external fertilization, the zygotes pass through typical spiralian development. In
Golfi11gin cleavage is spiral and holoblastic, but the relative sizes of micromeres and macromeres differ
among species, depending on the amount of yolk in the egg. Although traditionally sipunculan
embryos ,vere described as having a "molluscan cross," whlch suggested a close relationship with the
Mollusca, this character is now known to have been misinterpreted, and sipunculans are now
considered to be an early but highly derived offshoot of Annelida. The cell fates are the same as those il l
most typical spiralians. The first three quartets of micromeres become ectoderm and ectomesoderm;
the 4d cell produces endomesoderm; and form the endoderm. The mouth opens at the site of the
blastopore, and the surrounding ectodermal cells grow inward as a stomodeum. The anus breaks
through secondarily on the dorsal surface. The 4d mesoderm proliferates as two bands, as it does in
other annelids, but yields the major trunk coelom without segmentation. Four different developmental
sequences have been recognized among the sipunculans, includmg direct (e.g., Golfingin, Theiste
pyroides) and mdirect development. ln direct development, the eggs are covered by an adhesive jelly
and attached to the substratum after fertilization. The embryo develops directly to a vermiform
individual that hatches as a mmute juvenile sipunculan. The other three developmental patterns are
mdirect, involving various combinations of larval stages. In some species (e.g., Phnscolion strombi), a
free -living lecithotrophlc trochophore larva develops and metamorphoses into a juvenjle worm. The
other two developmental patterns involve a second larval stage, the pelagosphera larva, that forms after
a metamorphosis of the trodlophore. In some species, both the troc11ophore and the pelagosphera
forms are lecithotrophjc and relatively short-lived (e.g., son1e species of Golftngin and Themiste), while
in others the pelagosphera larva is planktotrophic and may live for extended periods of time in the
plankton (e.g., Aspidosiphon parvulus, members of the genus Phascoloson,n). The transformation of the
trochophore to the pelagosphera larva involves a reduction or loss of the prototrochal ciliary band and
the formation of a single metatrochal band for locomotion. The pelagosphera eventually elongates,
settles, and becomes a juvenile sipunculan. Si.nee the pelagosphera larva of some species are estimated
to spend several months in the plankton and sipunculans display little morphological variation, a
disproportionate number of species, when compared to any other marme worm groups, have been

17
suggested to be cosmopolitan. However detailed developmental and molecular analyses of several
supposed cosmopolitan species have shown that cosmopolitanism cannot be the norm, and that instead
sipunculans have many cryptic or pseudocryptic species.

Annelid Phylogeny

There are undoubted annelid fossils that date back to the Cambrian period, with well-known
examples being Cnnndin and Biirgessochnetn. These show dear evidence of segmentation and chaetae,
though it is a subject of debate as to whether they could be classified very closely with any extant
lineage of annelids. But these were quite complex annelids and the early origins of the group are
obscure. As outlined in the Introduction, the traditionally recognized classes were Polychaeta,
Oligochaeta, and Hirudinea. The latter two are novv viewed as the taxon Clitellata, sin.ce recognizing
Hirudinea with class rartk renders Oligochaeta paraphyletic. Furthennore, we now know that Polychaeta
contains Clitellata, making it synonymous with Annelida and so the name "Polychaeta" is no longer used
as a forn1al taxon. In addition to the recognition that the traditional classification had to change, the
past few decades has seen a series of major changes in annelid membership as well, mainly due to
molecular-based phylogenetic studies. Echiura, Sipuncula, Pogonophora, and Vestimentifera, all of
which were once considered separate phyla, are now clearly seen as part of Annelida. Hence, Annelida
now encompasses many ani1nals that do not conform to the traditional wuque features of the group
such as possession of body segmentation and chaetae. Even more problematic has been resolving the
relationships among the groups that have long been regarded as part of the annelid radiation, a work
that must still be regarded as very much in progress. While a deep understanding of the origin and
evolution of annelids still eludes us, considerable progress has been recently made with the use of Next
Generation sequencing technology, which holds great promise. This means we will likely see a stable
annelid phylogenetic tree in the coming years. Our current knov.r ledge of annelid relationships, as used
in this chapter, is based on the recent phylogenonuc studies of Sonia Andrade and Anne Weigert and
their colleagues, and these were used to generate the sun,mary tree and classification used in this
chapter. As future phylogenomic analyses are integrated with increasing knowledge of fossil aru1elids,
we will be able to make better inferences as to the origin and evolution of this amazingly diverse group
of animals.

2) SUMMARY OF COMPARATIVE BOOK

Annelida

Most of the 15 000 known species of annelid worms live in the sea, crawling under rocks on
the shore and sea bottom, freely swimming or seeking protection from predators in burrows or
tubes. Freshwater and terrestrial annelid radiation has been small except for the earthworms,
which are outstandingly successful, and the leeches, which are widespread specialized suctorial

18
predators. Annelids vary in length from 1mm between sand grains to 3 meters in some Australasian
earthworms.

Annelid worms are coelomates with metameric segmentation. The annelid coelom is a large
fluid-filled body cavity surrounded by mesoderm, providing an efficient hydrostatic skeleton.
Metamerism is the serial repetition of similar parts along the length of an animal, manifested
primarily in the separation of the mesoderm into segmental blocks of muscle. In annelids there are
usually internal partitions (septa) between the segments. Both the coelom and metamerism
improve the effectiveness of muscle contractions, so that active locomotion can be much faster than
in acoelomate animals. At the same time these advances demand greater complexity: when an
extensive coelom separates the inner and outer tissues, a transport system is required and more
elaborate respiratory and excretory organs may need to develop. This complexity then allows more
structural differentiation and an increase in size.

What is an Annelid?

An annelid is a protostome worm with three cell layers, a gut with a mouth and an anus and
a body wall with both circular and longitudinal muscle. The coelomic body cavity is formed by the
splitting of embryonic mesoderm. The outer epidermis is covered by a thin cuticle, typically bearing
chitinous bristles ('chaetae' or 'Setae'). Metameric segmentation, always shown in the muscular and
nervous systems, is characteristically also evident externally. The nervous system has a supra-
oesophageal ganglion (i.e. group of the main bodies of nerve cells), which is called a brain although
it may be little more than a sensory relay, and a ventral nerve cord bearing segmental ganglia giving
off segmental nerves. There is a closed blood system with the blood moving forward in the dorsal
longitudinal vessel. Segmental ducts between the coelom and the outside are used for excretion and
reproduction.

What annelids are there?

The three main classes are:

1. Polychaeta
This large class of marine annelids shows great diversity. Characteristically annelids bear
'parapodia', paired lobes on the sides of most segments forming paddle-like appendages
with bundles of chaetae.
2. Oligochaeta

19
 Mainly freshwater or terrestrial annelids, without parapodia and with a few unjointed
chaetae. The coelom is large and spacious. Feeding is suctorial; there are no jaws.
Oligochaetes are characterised by a saddlelike 'Clitellum', an epidermal thickening that
secretes a cocoon.
3. Hirudinea (leeches).
Leeches have a clitellum, anterior and posterior suckers and no chaetae. They are
specialized predators, often ectoparasites, in fresh water, on land or sometimes in the sea.
Oligochaeta and Hirudinea are now usually united as Clitellata.

What are the advantages of the coelomand of metamerism?

 The coelom
A coelom is a body cavity surrounded by mesoderm and lined by a mesodermal epithelium,
Such a cavity is useful in many ways. Separation of the gut and body wall enables each to
move independently: the worm can bend without pushing food along the gut. A fluid-filled
body cavity makes transport more effective and excretory organs, gonads and their
products can be contained in the coelom. But probably the most important function of the
coelom is to serve as a hydrostatic skeleton against which muscles can contract.
 Metameric segmentation
Like the coelom, metameric segmentation is above all concerned with movement. The
primary metameric structures are the body wall muscles, innervated in annelids by
segmental nerves arising from a chain of segmental nerve ganglia. Segmentation allows
control of particular portions of muscle, enabling each to contract independently of its
neighbor. When each muscle contracts in succession, a wave of contraction passes along the
worm and can set up lateral undulations or peristaltic waves that can be used for
burrowing. First a 'penetration anchor' makes and enters the burrow, and then a 'terminal
anchor' takes hold and pulls in the rest of the worm. Such a burrowing sequence is common
in unsegmented animals also, but is very much faster and more efficient where fluid
localized by septa in a front coelomic compartment can build up considerable hydrostatic
pressure, as in earthworms. Burrowers in soft sand, such as lugworms, by contrast lose
some septa and move coelomic fluid forward in bulk.
 Methods of locomotion
The slow walking of a typical polychaete such as Nereis is achieved by stepping with
parapodia, their chaetae increasing traction with the substratum. Fast walking however

20
 uses the parapodia much less. The body is thrown into horizontal waves by the successive
contraction and relaxation of the main body wall muscles alternately on the two sides; the
lateral components of the force generated will cancel out and the worm moves forward by
pressing backward on its environment. When the polychaete changes from fast walking to
swimming, the very rapid wave illustrates the activity possible for a segmented coelomate
worm. Polychaete lateral undulation differs from wave propulsion in a nematode, eel or
snake in that there are projecting parapodia. They do not contribute much directly, but their
presence, spread out on the outside and clustered on the inside of each bend means that the
propulsive force is exerted on the outside convex surface rather than on the inside concave
one. The wave of contraction has then to progress from the back to the front of the worm,
unlike the front to back wave of a nematode, eel or snake, and the polychaete brain has to
initiate locomotion at the far end of the body. A polychaete with all parapodia removed
swims backwards. Oligochaetes use the alternating contraction of circular and longitudinal
muscles not to undulate but for peristalsis. Circular muscle contraction enables an
earthworm to extend a longer thinner anterior portion while chaetae grip the ground in the
portion just behind, made shorter and fatter by contraction of longitudinal muscles. The
anterior circular muscles then relax and the wave of circular contraction passes backward
along the worm. Leeches may 'loop' by attaching and detaching them suckers, or may swim
by up-and-down waves propagated from the anterior end. To do this they need to flatten
and stiffen their bodies by contraction of dorsoventral muscles, so that they can flex their
muscles without the body shortening. The control of locomotion is more advanced than in
other annelids, with a more condensed nervous system and more centralized functions.

How does a coelomintroduce complexity?

The coelom both allows increase in size and activity and introduces a barrier between the outer
body wall and the gut. A transport system therefore becomes essential, to supply the inner tissues
with oxygen and the outer tissues with food.

 The blood system

A small part of the primary body cavity or 'blastocoel' of an annelid persists during
development, becomes filled with blood and forms a closed system of vessels. General
contractility of the dorsal longitudinal vessel pushes the blood forward, and then through a
varying number of connecting vessels to flow posteriorly in the ventral vessel beneath the
gut. While there is no true heart, there may be a number of blind ending contractile vessels

21
 that assist blood flow. There may be further vessels, for example to the parapodia in
polychaetes. Such transverse vessels require the support of segmental septa (it is uncertain
which of these structures came first and necessitated the other). Leeches have very
different circulatory systems. The coelom becomes packed with connective ‘botryoidal’
tissue, leaving coelomic channels in which fluid circulates. The blood system then becomes
much reduced or is absent. lobes with a large surface area and internal blood supply. Tube-
living polychaetes may need separate gills if the tentacles collect food from sand.
Respiratory pigments are now necessary, either in the blood to increase its oxygen carrying
capacity or in tissues to facilitate diffusion. Hemoglobin is the commonest respiratory
pigment, occurring in a very wide range of animals. It occurs in many annelids, but so also
do two other rare iron-based pigments, haemerythrin and chlorocruorin, and the copper-
based haemocyanin: no other phylum has so great a range.
 Excretion
While most polychaetes are osmoconformers, osmotic regulation is an important aspect of
excretion in annelids able to colonize the dilute environments. For example, Nereis
diversicolor in estuaries and earthworms in wet soil can produce hypotonic urine. The
excretory organs in larvae and some adult polychaetes (and possibly in ancestral annelids)
are 'protonephridia', tubes with blind ends where cilia-mediated filtrationoccurs. Most
polychaetes have 'metanephridia', ducts opening from the coelom by ciliated funnels. The
tubes may lead separately to the outside but more often open into the mesodermal gonadal
ducts occurring in most segments.

How do annelids reproduce and feed?

Polychaetes are by far the largest class of annelids, showing remarkably great diversity, while
the oligochaetes (and still more the leeches) are more uniform. The division between polychaetes
and oligochaetes is at first sight extremely tidy, with a habitat difference typically related to
reproduction, as follows:

 Reproduction
Polychaetes are marine and have separate sexes, external sperm transfer and external
fertilisation. The eggs are shed through the segmental organs into the sea, where they
develop into planktonic 'Trochophore' larvae. Oligochaetes, in fresh water or in soil, are

22
 cross-fertilized hermaphrodites with no external sperm transfer: the yolky eggs are
fertilised within the worm or in the cocoon. The clitellum, situated near the female pores,
secretes the cocoon into which (before or after fertilisation) the eggs are shed, and where
they develop directly with no larval stage. Leeches in any habitat reproduce like
oligochaetes, with direct development in a cocoon. Generalization provides a useful
framework but (as so often in biology) breaks down on account of the versatility of natural
selection. Polychaetes are not all marine but have a few freshwater species, and rather more
living under damp rotten logs on land; oligochaetes have a number of marine species. While
the generalizations about reproduction and development in oligochaetes may hold,
polychaetes vary widely. The female may collect and store sperm, for example, or may be
viviparous; the trochophore larva may be modified or suppressed where development
proceeds in a yolky egg. Asexual reproduction (with corresponding regenerative powers)
may occur in polychaetes by budding, or may be combined with the sexual process as when
the back end of a Nereis breaks off and becomes transformed into a pelagic 'heteronereis':
like other pelagic polychaetes, it then floats due to its extended surface and has very large
eyes. Gamete release may be synchronized in a swarm of sexually mature polychaetes.
 Feeding
Polychaetes freely crawling or swimming in the sea are usually brightly colored active
predators, with biting jaws borne on an eversible pharynx. Most are carnivores, but
herbivores, detritivores and omnivores are known. There are very many genera, mostly
with names inspired by Greek nymphs and goddesses. Nereis is a familiar 'ragworm', as are
Nephtys and Phylloduce; the Syllidae are slender with feathery parapodia, the Glyceridae
have balloonlike pharynges, the Eunicidae have especially strong jaws and the scale worms
such as Aphrodite have gill covers all down their backs. There are some 25 families,
formerly grouped as 'Errantia', but they are not all closely related. The arrival of arthropod
predators was no doubt a stimulus for rapid evolution among relatively defenceless worms;
many times separately they took to living in burrows as sedimentary or detritus feeders.
External structures became reduced but muscles remained well developed and gills were
needed.

How are annelids related to each other?

23
 The relationship between the three main classes of annelid is clear only in that the Clitellata
are distinct, whatever their origin. Within this group, there can no longer be a clear demarcation
between classes: the highly specialized Hirudinea (leeches) arose, probably more than once, from
among the Oligochaetes. Relationships among the many orders of polychaetes are very hard to
elucidate: clearly the protection of burrows and tubes was achieved many times separately by
different ancestors. There has long been controversy about the size and nature of the annelid
ancestor; now the question is, did the polychaetes (and therefore the annelids as a whole) have a
common ancestor at all? To apply the language of cladistics, polychaetes may not be ‘Paraphyletic’
(an assemblage including some but not all members of a monophyletic group) but 'polyphyletic' (a
group of animals with more than one ancestor). Mercifully we have insufficient evidence to pursue
these findings to a conclusion, and we continue at present to talk about 'Polychaeta' and 'Annelida'
as we always have done.

How are annelids related to other phyla?

The first step towards an answer is clear: annelid early development is unambiguously
protostome. Cleavage is spiral, the blastopore becomes the mouth and the mesoderm is early set
aside, later becoming blocks that form segments and split to make the coelom. Annelids are
traditionally placed near the base of the protostome radiation and close to both the molluscs and
the arthropods; more recent morphological assessment has maintained the link with molluscs but
has denied that the common possession of metameric segmentation united annelids and
arthropods. Molecular evidence supports the complete separation of annelids and arthropods and
the placing of annelids in a group with other soft-bodied worms and with molluscs, but the
relationships within this group remain uncertain. Molecular evidence is an exciting new guide to
relationships, but it cannot in isolation tell us anything about the direction of evolution, nor can it
replace the need to study the animals themselves. Observing the rapid and well-controlled
locomotion of many annelids makes it hard to believe that secondary loss of a coelom could have
been advantageous for an active predator. Likewise, it seems most improbable that nemertines
evolved from animals with coelomic body cavities: it is far more likely that localized structures such
as the rhynchocoel evolved convergently. Annelids show us that metameric segmentation, like the
coelom, can be a great functional asset, and we should not be surprised that it almost certainly
evolved several times separately. Phylogenies uniting animals because they share such clearly

24
advantageous characters may be very misleading: the likelihood of convergence should not be
underestimated.

III. ADVANTAGES AND WEAKNESS OF BOOK CONTENTS

1) MAIN BOOK
A. ADVANTAGES
 The language used in the discussion of this book is easy for readers to understand
 This book has a more complete discussion than the comparison book.
 This book describes annelids in detail and in this book the classification of
annelids is presented so that it can help readers gain insight into annelids in detail.
 This book is also equipped with good pictures so that the reader is not too difficult
to read.
B. WEAKNESS

25
  The way of writing and the layout of this book is not good, such as the spacing is
too tight and there are some letters missing so that readers who do not understand
English will find it difficult to understand.

2) BOOK OF COMPARISON
A. ADVANTAGES
 Use language that is easy for readers to understand.
 Good writing and layout of letters so that readers are not difficult to understand
 In this book, the important parts are given in a table or box so that it helps the
author to find important things.
B. WEAKNESS
 This book does not contain pictures or images that are displayed are too few.
 In this book the material is not as complete as in the main book, this book only
focuses on annelids in general.

CHAPTER III

CLOSING

A. CONCLUSION
Based on the strengths and weaknesses of the book, it is concluded that the main
book is more suitable as a student handbook because it supports learning with clearer
material exposure than the comparison book even though both books use language that is
easily understood by readers.

B. ADVICE
According to our group, the main book pays more attention to the way of writing
or the spacing used because this is an ebook so that readers are more interested in reading
it and for the second book to add more material that is lacking or not included in the
material explanation and it is hoped that with add images to the material.

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 BIBLIOGRAPHY

Brusca, Richard C. I Moore, Wendy. I Shuster, Stephen M,2016, Invertebrate Third Edition,USA:
Sinauer Associates, Inc.

Janet Moore,2006, An Introduction to the Invertebrates,New York : Cambridge University Press.

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