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15.4: Mollusks and Annelids - Biology

15.4: Mollusks and Annelids - Biology



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The mollusks are a diverse group (85,000 described species) of mostly marine species. They have a variety of forms, ranging from large predatory squid and octopus, some of which show a high degree of intelligence, to small grazing forms with elaborately sculpted and colored shells. The annelids traditionally include the oligochaetes, which include the earthworms and leeches, the polychaetes, which are a marine group, and two other smaller classes.

The phyla Mollusca and Annelida belong to a clade called the Lophotrochozoa, which also includes the phylum Nemertea, or ribbon worms ([link]). They are distinct from the Ecdysozoa (nematodes and arthropods) based on evidence from analysis of their DNA, which has changed our views of the relationships among invertebrates.

Phylum Mollusca

Mollusca is the predominant phylum in marine environments, where it is estimated that 23 percent of all known marine species belong to this phylum. It is the second most diverse phylum of animals with over 75,000 described species. The name “mollusca” signifies a soft body, as the earliest descriptions of mollusks came from observations of unshelled, soft-bodied cuttlefish (squid relatives). Although mollusk body forms vary, they share key characteristics, such as a ventral, muscular foot that is typically used for locomotion; the visceral mass, which contains most of the internal organs of the animal; and a dorsal mantle, which is a flap of tissue over the visceral mass that creates a space called the mantle cavity. The mantle may or may not secrete a shell of calcium carbonate. In addition, many mollusks have a scraping structure at the mouth, called a radula (Figure 15.4.1).

The muscular foot varies in shape and function, depending on the type of mollusk (described below in the section on mollusk diversity). It is a retractable as well as extendable organ, used for locomotion and anchorage. Mollusks are eucoelomates, but the coelomic cavity is restricted to a cavity around the heart in adult animals. The mantle cavity, formed inside the mantle, develops independently of the coelomic cavity. It is a multi-purpose space, housing the gills, the anus, organs for sensing food particles in the water, and an outlet for gametes. Most mollusks have an open circulatory system with a heart that circulates the hemolymph in open spaces around the organs. The octopuses and squid are an exception to this and have a closed circulatory system with two hearts that move blood through the gills and a third, systemic heart that pumps blood through the rest of the body.

ART CONNECTION

Which of the following statements about the anatomy of a mollusk is false?

  1. Mollusks have a radula for scraping food.
  2. Mollusks have ventral nerve cords.
  3. The tissue beneath the shell is called the mantle.
  4. The mantle cavity contains hemolymph.

Mollusk Diversity

This phylum is comprised of seven classes: Aplacophora, Monoplacophora, Polyplacophora, Bivalvia, Gastropoda, Cephalopoda, and Scaphopoda.

Class Aplacophora (“bearing no plates”) includes worm-like animals living mostly on deep ocean bottoms. These animals lack a shell but have aragonite spicules on their skin. Members of class Monoplacophora (“bearing one plate”) have a single, cap-like shell enclosing the body. The monoplacophorans were believed extinct and only known as fossils until the discovery of Neopilina galatheae in 1952. Today, scientists have identified nearly two dozen living species.

Animals in the class Polyplacophora (“bearing many plates”) are commonly known as “chitons” and bear an armor-like, eight-plated shell (Figure 15.4.2). These animals have a broad, ventral foot that is adapted for attachment to rocks and a mantle that extends beyond the shell in the form of a girdle. They breathe with ctenidia (gills) present ventrally. These animals have a radula modified for scraping. A single pair of nephridia for excretion is present.


Figure 15.4.2: This chiton from the class Polyplacophora has the eight-plated shell indicative of its class. (credit: Jerry Kirkhart)

Class Bivalvia (“two shells”) includes clams, oysters, mussels, scallops, and geoducks. They are found in marine and freshwater habitats. As the name suggests, bivalves are enclosed in a pair of shells (or valves) that are hinged at the dorsal side. The body is flattened on the sides. They feed by filtering particles from water and a radula is absent. They exchange gases using a pair of ctenidia, and excretion and osmoregulation are carried out by a pair of nephridia. In some species, the posterior edges of the mantle may fuse to form two siphons that inhale and exhale water. Some bivalves like oysters and mussels have the unique ability to secrete and deposit a calcareous nacre or “mother of pearl” around foreign particles that enter the mantle cavity. This property is commercially exploited to produce pearls.

CONCEPT IN ACTION

Watch animations of clams and mussels feeding to understand more about bivalves.

Gastropods (“stomach foot”) include well-known mollusks like snails, slugs, conchs, sea hares, and sea butterflies. Gastropods include shell-bearing species as well as species with a reduced shell. These animals are asymmetrical and usually present a coiled shell (Figure 15.4.3).

The visceral mass in the shelled species is characteristically twisted and the foot is modified for crawling. Most gastropods bear a head with tentacles that support eyes. A complex radula is used to scrape food particles from the substrate. The mantle cavity encloses the ctenidia as well as a pair of nephridia.

The class Cephalopoda (“head foot” animals) includes octopuses, squids, cuttlefish, and nautilus. Cephalopods include shelled and reduced-shell groups. They display vivid coloration, typically seen in squids and octopuses, which is used for camouflage. The ability of some octopuses to rapidly adjust their colors to mimic a background pattern or to startle a predator is one of the more awe-inspiring feats of these animals. All animals in this class are predators and have beak-like jaws. All cephalopods have a well-developed nervous system, complex eyes, and a closed circulatory system. The foot is lobed and developed into tentacles and a funnel, which is used for locomotion. Suckers are present on the tentacles in octopuses and squid. Ctenidia are enclosed in a large mantle cavity and are serviced by large blood vessels, each with its own heart.

Cephalopods (Figure 15.4.4) are able to move quickly via jet propulsion by contracting the mantle cavity to forcefully eject a stream of water. Cephalopods have separate sexes, and the females of some species care for the eggs for an extended period of time. Although the shell is much reduced and internal in squid and cuttlefish, and absent altogether in octopus, nautilus live inside a spiral, multi-chambered shell that is filled with gas or water to regulate buoyancy.

Members of the class Scaphopoda (“boat feet”) are known colloquially as “tusk shells” or “tooth shells.” Tooth shells are open at both ends and usually lie buried in sand with the front opening exposed to water and the reduced head end projecting from the back of the shell. Tooth shells have a radula and a foot modified into tentacles, each with a bulbous end that catches and manipulates prey (Figure 15.4.5).

Annelida

Phylum Annelida are segmented worms found in marine, terrestrial, and freshwater habitats, but the presence of water or humidity is a critical factor for their survival in terrestrial habitats. The name of the phylum is derived from the Latin word annellus, which means a small ring. Approximately 16,500 species have been described. The phylum includes earthworms, polychaete worms, and leeches. Like mollusks, annelids exhibit protostomic development.

Annelids are bilaterally symmetrical and have a worm-like appearance. Their particular segmented body plan results in repetition of internal and external features in each body segment. This type of body plan is called metamerism. The evolutionary benefit of such a body plan is thought to be the capacity it allows for the evolution of independent modifications in different segments that perform different functions. The overall body can then be divided into head, body, and tail.

Physiological Processes of Annelida

The skin of annelids is protected by a cuticle that is thinner than the cuticle of the ecdysozoans and does not need to be molted for growth. Chitinous hairlike extensions, anchored in the skin and projecting from the cuticle, called chaetae, are present in every segment in most groups. The chaetae are a defining character of annelids. Polychaete worms have paired, unjointed limbs called parapodia on each segment used for locomotion and breathing. Beneath the cuticle there are two layers of muscle, one running around its circumference (circular) and one running the length of the worm (longitudinal). Annelids have a true coelom in which organs are distributed and bathed in coelomic fluid. Annelids possess a well-developed complete digestive system with specialized organs: mouth, muscular pharynx, esophagus, and crop. A cross-sectional view of a body segment of an earthworm is shown in Figure 15.4.6; each segment is limited by a membrane that divides the body cavity into compartments.

Annelids have a closed circulatory system with muscular pumping “hearts” in the anterior segments, dorsal and ventral blood vessels that run the length of the body with connections in each segment, and capillaries that service individual tissues. Gas exchange occurs across the moist body surface. Excretion is carried out by pairs of primitive “kidneys” called metanephridia that consist of a convoluted tubule and an open, ciliated funnel present in every segment. Annelids have a well-developed nervous system with two ventral nerve cords and a nerve ring of fused ganglia present around the pharynx.

Annelids may be either monoecious with permanent gonads (as in earthworms and leeches) or dioecious with temporary or seasonal gonads (as in polychaetes).

CONCEPT IN ACTION

This video and animation provides a close-up look at annelid anatomy.

Annelid Diversity

Phylum Annelida includes the classes Polychaeta and Clitellata (Figure 15.4.7); the latter contains subclasses Oligochaeta, Hirudinoidea, and Branchiobdellida.

Earthworms are the most abundant members of the subclass Oligochaeta, distinguished by the presence of the clitellum, a ring structure in the skin that secretes mucus to bind mating individuals and forms a protective cocoon for the eggs. They also have a few, reduced chaetae (oligo- = “few”; -chaetae = “hairs”). The number and size of chaetae is greatly diminished in oligochaetes as compared to the polychaetes (poly- = “many”; -chaetae = “hairs”). The chaetae of polychaetes are also arranged within fleshy, flat, paired appendages on each segment called parapodia.

The subclass Hirudinoidea includes leeches. Significant differences between leeches and other annelids include the development of suckers at the anterior and posterior ends, and the absence of chaetae. Additionally, the segmentation of the body wall may not correspond to internal segmentation of the coelomic cavity. This adaptation may allow leeches to swell when ingesting blood from host vertebrates. The subclass Branchiobdellida includes about 150 species that show similarity to leeches as well as oligochaetes. All species are obligate symbionts, meaning that they can only survive associated with their host, mainly with freshwater crayfish. They feed on the algae that grows on the carapace of the crayfish.

Section Summary

The phylum Mollusca is a large, mainly marine group of invertebrates. Many mollusks secrete a calcareous shell for protection, but in other species, the shell is reduced or absent. Mollusks are protostomes. The dorsal epidermis in mollusks is modified to form the mantle, which encloses the mantle cavity and visceral organs. This cavity is distinct from the coelomic cavity, which the adult animal retains, surrounding the heart. Respiration is facilitated by gills known as ctenidia. A chitinous scraper called the radula is present in most mollusks. Mollusks are mostly dioecious and are divided into seven classes.

The phylum Annelida includes worm-like, segmented animals. Segmentation is both external and internal, which is called metamerism. Annelids are protostomes. The presence of chitinous hairs called chaetae is characteristic of most members. These animals have well-developed nervous and digestive systems. Polychaete annelids have parapodia that participate in locomotion and respiration. Suckers are seen in the order Hirudinea. Breeding systems include separate sexes and hermaphroditism.

Art Connections

Figure 15.4.1 Which of the following statements about the anatomy of a mollusk is false?

A. Mollusks have a radula for scraping food.
B. Mollusks have ventral nerve cords.
C. The tissue beneath the shell is called the mantle.
D. The mantle cavity contains hemolymph.

Figure 15.4.1 D

Review Questions

A mantle and mantle cavity are present in _____.

A. class Oligochaeta
B. class Bivalvia
C. class Polychaeta
D. class Hirudinea

C

Annelids have a _____.

A. pseudocoelom
B. a true coelom
C. no coelom
D. none of the above

B

Free Response

Describe the morphology and anatomy of mollusks.

Mollusks have a large muscular foot that may be modified in various ways, such as into tentacles, but it functions in locomotion. They have a mantle, a structure of tissue that covers and encloses the dorsal portion of the animal and secretes the shell when it is present. The mantle encloses the mantle cavity, which houses the gills (when present), excretory pores, anus, and gonadopores. The coelom of mollusks is restricted to the region around the systemic heart. The main body cavity is a hemocoel. Many mollusks have a radula near the mouth that is used for scraping food.

Glossary

Annelida
a phylum of worm-like animals with metamerism
chaeta
a chitinous projection from the cuticle found in annelids
clitellum
a specialized band of fused segments in some annelids, which aids in reproduction
ctenidia
specialized gills in mollusks
Lophotrochozoa
a clade of invertebrate organisms that is a sister group to the Ecdysozoa
mantle
a specialized epidermis that encloses all visceral organs and secretes shells in mollusks
metamerism
having a series of body structures that are similar internally and externally, such as segments
Mollusca
a phylum of protostomes with soft bodies and no segmentation
nacre
a calcareous secretion produced by bivalve mollusks to line the inner side of shells as well as to coat foreign particulate matter

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15 Cards in this Set

Charactersitic's of Mollusks

Have a three part body plan

Ex. Snails, oysters, clams, octopus and squid

Mollusks Body Plan & Organ System

The three parts that make up the mollusk are the mantle, visceral mass and the foot

Mollusk Feeding & Digestion

All mollusks (Except bivalves) have a two shelled radula which is a tongue like organ covered with thousands of pointing curled teeth.

Food is digested in the stomach and intestine of the one way digestive tract.

Waste passes out the anus (Tube within a tube)

Animals with true coeloms

The gut and other internal organs are suspended from the body wall and are cushioned by the fluid within the coelom.

The visceral mass is the central section of the mollusk body and contains the mollsck's organs.

The heavy fold of tissue outside of the visceral mass which forms the outer layer of the body and secretes the mollusk's shell.

A muscular mass found in every mollusk which is used primarily for locomotion

The coelom of the mollusk collects waste body fluids

The cilia then pull the fluid from the coelom into a tubelike sructure called the nepheridia which recovers an useful materials (Ie. salt, sugar)

The remaining fluid leave via the anal pore

Most mollusks have a 3 chambered heart and an open circulatory system

Optopi and their relatives are exceptions as they have closed circulatory systems where blood remains inside of the vessels

Most mollusk respire with gills found in the mantle cavity

Mollusk gills extract 50% or more of the dissolved oxygen from the water that passes over them

Most terrestrial snails have no gills and the thin membrane that lines the snail's empty mantle cavity works as a primitive lung

Space located between the visceral mass and the mantle

Distinct female and male invididuals (some snails and slugs are hermaphrodites)

Fertilization of eggs occurs externally in most aquatic mollusks and internally in terrestrial mollusks and octopuses and their relatives.

The fertilized eggs develop into trochophore larave

Gastropods, cepholopods and bivavles share the same basic organ systems and tissue layers but different feeding and body plans

snails and slugs that live primarily marine but some in terrestrial and freshwater

Most gastropods have a pair of tentacles on their heads that have eyes on their tips

Most have a single shell but slugs and nudibranches have no shell

The foot of gastropods is adapted for locomotion. In terrestrials, the base secretes mucus from the base of the foot to provide a slimy path for them to glide along


Phylum Mollusca

Mollusca is the predominant phylum in marine environments, where it is estimated that 23 percent of all known marine species belong to this phylum. It is the second most diverse phylum of animals with over 75,000 described species. The name “mollusca” signifies a soft body, as the earliest descriptions of mollusks came from observations of unshelled, soft-bodied cuttlefish (squid relatives). Although mollusk body forms vary, they share key characteristics, such as a ventral, muscular foot that is typically used for locomotion the visceral mass, which contains most of the internal organs of the animal and a dorsal mantle, which is a flap of tissue over the visceral mass that creates a space called the mantle cavity. The mantle may or may not secrete a shell of calcium carbonate. In addition, many mollusks have a scraping structure at the mouth, called a radula ([Figure 1]).

The muscular foot varies in shape and function, depending on the type of mollusk (described below in the section on mollusk diversity). It is a retractable as well as extendable organ, used for locomotion and anchorage. Mollusks are eucoelomates, but the coelomic cavity is restricted to a cavity around the heart in adult animals. The mantle cavity, formed inside the mantle , develops independently of the coelomic cavity. It is a multi-purpose space, housing the gills, the anus, organs for sensing food particles in the water, and an outlet for gametes. Most mollusks have an open circulatory system with a heart that circulates the hemolymph in open spaces around the organs. The octopuses and squid are an exception to this and have a closed circulatory system with two hearts that move blood through the gills and a third, systemic heart that pumps blood through the rest of the body.

Art Connection

Figure 1: There are many species and variations of mollusks the gastropod mollusk anatomy is shown here, which shares many characteristics common with other groups.

Which of the following statements about the anatomy of a mollusk is false?

  1. Mollusks have a radula for scraping food.
  2. Mollusks have ventral nerve cords.
  3. The tissue beneath the shell is called the mantle.
  4. The mantle cavity contains hemolymph.
    [reveal-answer q=�″]Show Answer[/reveal-answer]
    [hidden-answer a=�″]4[/hidden-answer]

This phylum is comprised of seven classes: Aplacophora, Monoplacophora, Polyplacophora, Bivalvia, Gastropoda, Cephalopoda, and Scaphopoda.

Class Aplacophora (“bearing no plates”) includes worm-like animals living mostly on deep ocean bottoms. These animals lack a shell but have aragonite spicules on their skin. Members of class Monoplacophora (“bearing one plate”) have a single, cap-like shell enclosing the body. The monoplacophorans were believed extinct and only known as fossils until the discovery of Neopilina galatheae in 1952. Today, scientists have identified nearly two dozen living species.

Animals in the class Polyplacophora (“bearing many plates”) are commonly known as “chitons” and bear an armor-like, eight-plated shell ([Figure 2]). These animals have a broad, ventral foot that is adapted for attachment to rocks and a mantle that extends beyond the shell in the form of a girdle. They breathe with ctenidia (gills) present ventrally. These animals have a radula modified for scraping. A single pair of nephridia for excretion is present.

Figure 2: This chiton from the class Polyplacophora has the eight-plated shell indicative of its class. (credit: Jerry Kirkhart)

Class Bivalvia (“two shells”) includes clams, oysters, mussels, scallops, and geoducks. They are found in marine and freshwater habitats. As the name suggests, bivalves are enclosed in a pair of shells (or valves) that are hinged at the dorsal side. The body is flattened on the sides. They feed by filtering particles from water and a radula is absent. They exchange gases using a pair of ctenidia, and excretion and osmoregulation are carried out by a pair of nephridia. In some species, the posterior edges of the mantle may fuse to form two siphons that inhale and exhale water. Some bivalves like oysters and mussels have the unique ability to secrete and deposit a calcareous nacre or “mother of pearl” around foreign particles that enter the mantle cavity. This property is commercially exploited to produce pearls.

Watch animations of clams and mussels feeding to understand more about bivalves.

Gastropods (“stomach foot”) include well-known mollusks like snails, slugs, conchs, sea hares, and sea butterflies. Gastropods include shell-bearing species as well as species with a reduced shell. These animals are asymmetrical and usually present a coiled shell ([Figure 3]).

Figure 3: (a) Like many gastropods, this snail has a stomach foot and a coiled shell. (b) This slug, which is also a gastropod, lacks a shell. (credit a: modification of work by Murray Stevenson credit b: modification of work by Rosendahl)

The visceral mass in the shelled species is characteristically twisted and the foot is modified for crawling. Most gastropods bear a head with tentacles that support eyes. A complex radula is used to scrape food particles from the substrate. The mantle cavity encloses the ctenidia as well as a pair of nephridia.

The class Cephalopoda (“head foot” animals) includes octopuses, squids, cuttlefish, and nautilus. Cephalopods include shelled and reduced-shell groups. They display vivid coloration, typically seen in squids and octopuses, which is used for camouflage. The ability of some octopuses to rapidly adjust their colors to mimic a background pattern or to startle a predator is one of the more awe-inspiring feats of these animals. All animals in this class are predators and have beak-like jaws. All cephalopods have a well-developed nervous system, complex eyes, and a closed circulatory system. The foot is lobed and developed into tentacles and a funnel, which is used for locomotion. Suckers are present on the tentacles in octopuses and squid. Ctenidia are enclosed in a large mantle cavity and are serviced by large blood vessels, each with its own heart.

Cephalopods ([Figure 4]) are able to move quickly via jet propulsion by contracting the mantle cavity to forcefully eject a stream of water. Cephalopods have separate sexes, and the females of some species care for the eggs for an extended period of time. Although the shell is much reduced and internal in squid and cuttlefish, and absent altogether in octopus, nautilus live inside a spiral, multi-chambered shell that is filled with gas or water to regulate buoyancy.

Figure 4: The (a) nautilus, (b) giant cuttlefish, (c) reef squid, and (d) blue-ring octopus are all members of the class Cephalopoda. (credit a: modification of work by J. Baecker credit b: modification of work by Adrian Mohedano credit c: modification of work by Silke Baron credit d: modification of work by Angell Williams)

Members of the class Scaphopoda (“boat feet”) are known colloquially as “tusk shells” or “tooth shells.” Tooth shells are open at both ends and usually lie buried in sand with the front opening exposed to water and the reduced head end projecting from the back of the shell. Tooth shells have a radula and a foot modified into tentacles, each with a bulbous end that catches and manipulates prey ([Figure 5]).

Figure 5: Antalis vulgaris shows the classic Dentaliidae shape that gives these animals their common name of “tusk shell.” (credit: Georges Jansoone)


Phylogenetic relationships of annelids, molluscs, and arthropods evidenced from molecules and morphology

Annelids and arthropods have long been considered each other's closest relatives, as evidenced by similarities in their segmented body plans. An alternative view, more recently advocated by investigators who have examined partial 18S ribosomal RNA data, proposes that annelids, molluscs, and certain other minor phyla with trochophore larva stages share a more recent common ancestor with one another than any do with arthropods. The two hypotheses are mutually exclusive in explaining spiralian relationships. Cladistic analysis of morphological data does not reveal phylogentic relationships among major spiralian taxa but does suggest monophyly for both the annelids and molluscs. Distance and maximum-likelihood analyses of 18S rRNA gene sequences from major spiralian taxa suggest a sister relationship between annelids and molluscs and provide a clear resolution within the major groups of the spiralians. The parsimonious tree based on molecular data, however, indicates a sister relationship of the Annelida and Bivalvia, and an earlier divergence of the Gastropoda than the Annelida-Bivalvia clade. To test further hypotheses on the phylogenetic relationships among annelids, molluscs, and arthropods, and the ingroup relationships within the major spiralian taxa, we combine the molecular and morphological data sets and subject the combined data matrix to parsimony analysis. The resulting tree suggests that the molluscs and annelids form a monophyletic lineage and unites the molluscan taxa to a monophyletic group. Therefore, the result supports the Eutrochozoa hypothesis and the monophyly of molluscs, and indicates early acquisition of segmented body plans in arthropods.


Molluscs

This is a description of their main characteristic -all molluscs ave a soft body, which ( with the exceptions of species from certain families) is generally protected by an external, calcium-containing shell. In the octopuses and sea-slugs the shells is completely absent where as in squids it is located internally.

Main characteristics of Mollusca :

A ll modern mollusks have the same fundamental body plan. There are three distinct body zones: a head-foot, a visceral mass and a mantle. Some common features are :

  1. Unsegmented soft body with bilateral symmetry
  2. Presence of an internal or external shell
  3. A toothed tongue (made mostly of chitin) called the radula
  4. A mantle which is a fold in the body wall that lines the shell
  5. Muscular foot (and/or tentacles in some).
  6. Possession of a fluid-filled cavity called coelom

Diagram of body plan. – Shall soon appear here

The Head-foot region contains the sensory and motor organs. The foot is muscular and is used for a variety of purposes namely locomotion, capturing prey for food and attachment to the substratum.
The visceral mass contains the internal organs, namely the organs of digestive excretory and reproductive systems.
The mantle is a specialized tissue that surrounds the visceral mass but does not cover it completely it also secretes the shell and contributes to the development of breathing organs. The mantle cavity is the space between mantle and the visceral mass. The gills are located here and the digestive, excretory and reproductive organs discharge their contents into this cavity.
Radula is a feature that is seen only in molluscs. Its structure ranges from simple tongue like ones to scrape algae off the rocks to complex harpoon-like ones seen in the cone snails that eject chemicals capable of killing humans. Most of the molluscs have gills to obtain the oxygen from water. The coelom is a haemocoel – quite reduced and found near and around the heart and as cavities around the excretory and reproductive organs

Some anatomical features :
All molluscs have a complete digestive tract starting with the mouth and ending with anus. The digestive tract is ciliated and stomach structures vary in molluscs depending upon their food intake. The undigested food is compressed and packaged into solid wastes that are sent from anus to mantle to be released outside. Many of them are filter-feeders while others are predatory.
With the exception of cephalopods all molluscs have an open circulatory system.
The blood circulates between gills and heart via the blood-filled space haemocoel.
Excretion is carried out by a pair of nephridia, who empty their contents into the mantle cavity.
Locomotion is accomplished with the help of the muscular foot. The herbivores glide where as the bivalves use it to burrow in the sand,the cephalopods swim actively creating a kind of jet propulsion by expelling water from their mantle cavity. The cuttle fishes use their lateral fins for greater maneuverability.
Molluscs have a complex nervous system made up of ganglia and nerve cords, with the octopus having the most highly evolved one. It is also considered to be a very intelligent animal with supposedly an IQ equivalent to that of a domestic cat. The giant squids have very large eyes almost the size of a quarter plate.
The basic pattern of reproduction is one of having separate sexes but several snails are hermaphrodites. Sperm and eggs are spawned into the water, with fertilization being external, but internal fertilization can also be seen in several species. The development stages suggest a close relationship between molluscs and annelids( earthworm is an example of annelid). Some Oysters exhibit parental care.


15.4: Mollusks and Annelids - Biology

By the end of this section, you will be able to do the following:

  • Describe the unique anatomical and morphological features of molluscs and annelids
  • Describe the formation of the coelom
  • Identify an important extracoelomic cavity in molluscs
  • Describe the major body regions of Mollusca and how they vary in different molluscan classes
  • Discuss the advantages of true body segmentation
  • Describe the features of animals classified in phylum Annelida

The annelids and the mollusks are the most familiar of the lophotrochozoan protostomes. They are also more “typical” lophotrochozoans, since both groups include aquatic species with trochophore larvae, which unite both taxa in common ancestry. These phyla show how a flexible body plan can lead to biological success in terms of abundance and species diversity. The phylum Mollusca has the second greatest number of species of all animal phyla with nearly 100,000 described extant species, and about 80,000 described extinct species. In fact, it is estimated that about 25 percent of all known marine species are mollusks! The annelids and mollusca are both bilaterally symmetrical, cephalized, triploblastic, schizocoelous eucoeolomates They include animals you are likely to see in your backyard or on your dinner plate!

Phylum Mollusca

The name “Mollusca” means “soft” body, since the earliest descriptions of molluscs came from observations of “squishy,” unshelled cuttlefish. Molluscs are predominantly a marine group of animals however, they are also known to inhabit freshwater as well as terrestrial habitats. This enormous phylum includes chitons, tusk shells, snails, slugs, nudibranchs, sea butterflies, clams, mussels, oysters, squids, octopuses, and nautiluses. Molluscs display a wide range of morphologies in each class and subclass, but share a few key characteristics ((Figure)). The chief locomotor structure is usually a muscular foot. Most internal organs are contained in a region called the visceral mass. Overlying the visceral mass is a fold of tissue called the mantle within the cavity formed by the mantle are respiratory structures called gills, that typically fold over the visceral mass. The mouths of most mollusks, except bivalves (e.g., clams) contain a specialized feeding organ called a radula, an abrasive tonguelike structure. Finally, the mantle secretes a calcium-carbonate-hardened shell in most molluscs, although this is greatly reduced in the class Cephalopoda, which contains the octopuses and squids.

Art Connection

Figure 1. Molluscan body regions. There are many species and variations of molluscs this illustration shows the anatomy of an aquatic gastropod. In a terrestrial gastropod, the mantle cavity itself would serve as a respiratory organ.

Which of the following statements about the anatomy of a mollusc is false?

  1. Most molluscs have a radula for grinding food.
  2. A digestive gland is connected to the stomach.
  3. The tissue beneath the shell is called the mantle.
  4. The digestive system includes a gizzard, a stomach, a digestive gland, and the intestine.

The muscular foot is the ventral-most organ, whereas the mantle is the limiting dorsal organ that folds over the visceral mass. The foot, which is used for locomotion and anchorage, varies in shape and function, depending on the type of mollusk under study. In shelled mollusks, the foot is usually the same size as the opening of the shell. The foot is both retractable and extendable. In the class Cephalopoda (“head-foot”), the foot takes the form of a funnel for expelling water at high velocity from the mantle cavity and the anterior margin of the foot has been modified into a circle of arms and tentacles.

The visceral mass is present above the foot, in the visceral hump. This mass contains digestive, nervous, excretory, reproductive, and respiratory systems. Molluscan species that are exclusively aquatic have gills that extend into the mantle cavity, whereas some terrestrial species have “lungs” formed from the lining of the mantle cavity. Mollusks are schizocoelous eucoelomates, but the coelomic cavity in adult animals has been largely reduced to a cavity around the heart. However, a reduced coelom sometimes surrounds the gonads, part of the kidneys, and intestine as well. This overall coelomic reduction makes the mantle cavity the major internal body chamber.

Most mollusks have a special rasp-like organ, the radula, which bears chitinous filelike teeth. The radula is present in all groups except the bivalves, and serves to shred or scrape food before it enters the digestive tract. The mantle (also known as the pallium) is the dorsal epidermis in mollusks all mollusks except some cephalopods are specialized to secrete a calcareous shell that protects the animal’s soft body.

Most mollusks are dioecious animals and fertilization occurs externally, although this is not the case in terrestrial mollusks, such as snails and slugs, or in cephalopods. In most aquatic mollusks, the zygote hatches and produces a trochophore larva, with several bands of cilia around a toplike body, and an additional apical tuft of cilia. In some species, the trochophore may be followed by additional larval stages, such as a veliger larvae, before the final metamorphosis to the adult form. Most cephalopods develop directly into small versions of their adult form.

Classification of Phylum Mollusca

Phylum Mollusca comprises a very diverse group of organisms that exhibits a dramatic variety of forms, ranging from chitons to snails to squids, the latter of which typically show a high degree of intelligence. This variability is a consequence of modification of the basic body regions, especially the foot and mantle. The phylum is organized into eight classes: Caudofoveata, Solenogastres, Monoplacophora, Polyplacophora, Gastropoda, Cephalopoda, Bivalvia, and Scaphopoda. Although each molluscan class appears to be monophyletic, their relationship to one another is unclear and still being reviewed.

Both the Caudofoveata and the Solenogastres include shell-less, worm-like animals primarily found in benthic marine habitats. Although these animals lack a calcareous shell, they get some protection from calcareous spicules embedded in a cuticle that covers their epidermis. The mantle cavity is reduced, and both groups lack eyes, tentacles, and nephridia (excretory organs). The Caudofoveata possess a radula, but the Solenogastres do not have a radula or gills. The foot is also reduced in the Solenogastres and absent from the Caudofoveata.

Long thought to be extinct, the first living specimens of Monoplacophora, Neopilina galatheae, were discovered in 1952 on the ocean bottom near the west coast of Costa Rica. Today there are over 25 described species. Members of class Monoplacophora (“bearing one plate”) possess a single, cap-like shell that covers the dorsal body. The morphology of the shell and the underlying animal can vary from circular to ovate. They have a simple radula, a looped digestive system, multiple pairs of excretory organs, and a pair of gonads. Multiple gills are located between the foot and the edge of the mantle.

Animals in class Polyplacophora (“bearing many plates”) are commonly known as “chitons” and bear eight limy plates that make up the dorsal shell ((Figure)). These animals have a broad, ventral foot that is adapted for suction onto rocks and other substrates, and a mantle that extends beyond the edge of the shell. Calcareous spines on the exposed mantle edge provide protection from predators. Respiration is facilitated by multiple pairs of gills in the mantle cavity. Blood from the gills is collected in a posterior heart, and then sent to the rest of the body in a hemocoel—an open circulation system in which the blood is contained in connected chambers surrounding various organs rather than within individual blood vessels. The radula, which has teeth composed of an ultra-hard magnetite, is used to scrape food organisms off rocky surfaces. Chiton teeth have been shown to exhibit the greatest hardness and stiffness of any biomineral material reported to date, being as much as three-times harder than human enamel and the calcium carbonate-based shells of mollusks.

The nervous system is rudimentary with only buccal or “cheek” ganglia present at the anterior end. Multiple tiny sensory structures, including photosensors, extend from the mantle into channels in the upper layer of the shell. These structures are called esthetes and are unique to the chitons. Another sensory structure under the radula is used to sample the feeding environment. A single pair of nephridia is used for the excretion of nitrogenous wastes.

Figure 2. A chiton. This chiton from the class Polyplacaphora has the eight-plated shell for which its class is named. (credit: Jerry Kirkhart)

Class Bivalvia (“two-valves”) includes clams, oysters, mussels, scallops, geoducks, and shipworms. Some bivalves are almost microscopic, while others, in the genus Tridacna, may be one meter in length and weigh 225 kilograms. Members of this class are found in marine as well as freshwater habitats. As the name suggests, bivalves are enclosed in two-part valves or shells ((Figure)a) fused on the dorsal side by hinge ligaments as well as shell teeth on the ventral side that keep the two halves aligned. The two shells, which consist of an outer organic layer, a middle prismatic layer, and a very smooth nacreous layer, are joined at the oldest part of the shell called the umbo. Anterior and posterior adductor and abductor muscles close and open the shell respectively.

The overall body of the bivalve is laterally flattened the foot is wedge-shaped and the head region is poorly developed (with no obvious mouth). Bivalves are filter-feeders, and a radula is absent in this class of mollusks. The mantle cavity is fused along the edges except for openings for the foot and for the intake and expulsion of water, which is circulated through the mantle cavity by the actions of the incurrent and excurrent siphons. During water intake by the incurrent siphon, food particles are captured by the paired posterior gills (ctenidia) and then carried by the movement of cilia forward to the mouth. Excretion and osmoregulation are performed by a pair of nephridia. Eyespots and other sensory structures are located along the edge of the mantle in some species. The “eyes” are especially conspicuous in scallops ((Figure)b). Three pairs of connected ganglia regulate activity of different body structures.

Figure 3. Bivalves. These mussels (a), found in the intertidal zone in Cornwall, England, show the bivalve shell. The scallop Argopecten irradians (b) has a fluted shell and conspicuous eyespots. (credit (a): Mark A. Wilson. credit (b) Rachael Norris and Marina Freudzon. https://commons.wikimedia.org/w/index.php?curid=17251065)

One of the functions of the mantle is to secrete the shell. Some bivalves, like oysters and mussels, possess the unique ability to secrete and deposit a calcareous nacre or “mother of pearl” around foreign particles that may enter the mantle cavity. This property has been commercially exploited to produce pearls.

Link to Learning

Watch the animations of bivalves feeding: View the process in clams and mussels at these sites.

More than half of molluscan species are in the class Gastropoda (“stomach foot”), which includes well-known mollusks like snails, slugs, conchs, cowries, limpets, and whelks. Aquatic gastropods include both marine and freshwater species, and all terrestrial mollusks are gastropods. Gastropoda includes shell-bearing species as well as species without shells. Gastropod bodies are asymmetrical and usually present a coiled shell ((Figure)a). Shells may be planospiral (like a garden hose wound up), commonly seen in garden snails, or conispiral, (like a spiral staircase), commonly seen in marine conches. Cowrie shells have a polished surface because the mantle extends up over the top of the shell as it is secreted.

Figure 4. Gastropods. Snails(a) and slugs(b) are both gastropods, but slugs lack a shell. (credit a: modification of work by Murray Stevenson credit b: modification of work by Rosendahl)

A key characteristic of some gastropods is the embryonic development of torsion. During this process, the mantle and visceral mass are rotated around the perpendicular axis over the center of the foot to bring the anal opening forward just behind the head ((Figure)), creating a very peculiar situation. The left gill, kidney, and heart atrium are now on the right side, whereas the original right gill, kidney, and heart atrium are on the left side. Even stranger, the nerve cords have been twisted and contorted into a figure-eight pattern. Because of the space made available by torsion in the mantle cavity, the animal’s sensitive head end can now be withdrawn into the protection of the shell, and the tougher foot (and sometimes the protective covering or operculum) forms a barrier to the outside. The strange arrangement that results from torsion poses a serious sanitation problem by creating the possibility of wastes being washed back over the gills, causing fouling. There is actually no really perfect explanation for the embryonic development of torsion, and some groups that formerly exhibited torsion in their ancestral groups are now known to have reversed the process.

Gastropods also have a foot that is modified for crawling. Most gastropods have a well-defined head with tentacles and eyes. A complex radula is used to scrape up food particles. In aquatic gastropods, the mantle cavity encloses the gills (ctenidia), but in land gastropods, the mantle itself is the major respiratory structure, acting as a kind of lung. Nephridia (“kidneys”) are also found in the mantle cavity.

Figure 5. Torsion in gastropods. During embryonic development of some gastropods, the visceral mass undergoes torsion, or counterclockwise rotation of the visceral anatomical features. As a result, the anus of the adult animal is located over the head. Although torsion is always counterclockwise, the shell may coil in either direction thus coiling of a shell is not the same as torsion of the visceral mass.

Everyday Connection

Can Snail Venom Be Used as a Pharmacological Painkiller?

Marine snails of the genus Conus ((Figure)) attack prey with a venomous stinger, modified from the radula. The toxin released, known as conotoxin, is a peptide with internal disulfide linkages. Conotoxins can bring about paralysis in humans, indicating that this toxin attacks neurological targets. Some conotoxins have been shown to block neuronal ion channels. These findings have led researchers to study conotoxins for possible medical applications.

Conotoxins are an exciting area of potential pharmacological development, since these peptides may be possibly modified and used in specific medical conditions to inhibit the activity of specific neurons. For example, conotoxins or modifications of them may be used to induce paralysis in muscles in specific health applications, similar to the use of botulinum toxin. Since the entire spectrum of conotoxins, as well as their mechanisms of action, is not completely known, the study of their potential applications is still in its infancy. Most research to date has focused on their use to treat neurological diseases. They have also shown some efficacy in relieving chronic pain, and the pain associated with conditions like sciatica and shingles. The study and use of biotoxins—toxins derived from living organisms—are an excellent example of the application of biological science to modern medicine.

Figure 6. Conus. Members of the genus Conus produce neurotoxins that may one day have medical uses. The tube above the eyes is a siphon used both to circulate water over the gills and to sample the water for chemical evidence of prey nearby. Note the eyes below the siphon. The proboscis, through which the venomous harpoon is projected, is located between the eyes. (credit: David Burdick, NOAA)

Class Cephalopoda (“head foot” animals), includes octopuses, squids, cuttlefish, and nautiluses. Cephalopods include both animals with shells as well as animals in which the shell is reduced or absent. In the shell-bearing Nautilus, the spiral shell is multi-chambered. These chambers are filled with gas or water to regulate buoyancy. A siphuncle runs through the chambers, and it is this tube that regulates the amount of water and gases (nitrogen, carbon dioxide, and oxygen mixture) present in the chambers. Ammonites and other nautiloid shells are commonly seen in the fossil record. The shell structure in squids and cuttlefish is reduced and is present internally in the form of a squid pen and cuttlefish bone, respectively. Cuttle bone is sold in pet stores to help smooth the beaks of birds and also to provide birds such as egg-laying chickens and quail with an inexpensive natural source of calcium carbonate. Examples of cephalopods are shown in (Figure).

Cephalopods can display vivid and rapidly changing coloration, almost like flashing neon signs. Typically these flashing displays are seen in squids and octopuses, where they may be used for camouflage and possibly as signals for mating displays. We should note, however, that researchers are not entirely sure if squid can actually see color, or see color in the same way as we do. We know that pigments in the skin are contained in special pigment cells (chromatophores), which can expand or contract to produce different color patterns. But chromatophores can only make yellow, red, brown, and black pigmentation however, underneath them is a whole different set of elements called iridophores and leucophores that reflect light and can make blue, green, and white. It is possible that squid skin might actually be able to detect some light on its own, without even needing its eyes!

All animals in this class are carnivorous predators and have beak-like jaws in addition to the radula. Cephalopods include the most intelligent of the mollusks, and have a well-developed nervous system along with image-forming eyes. Unlike other mollusks, they have a closed circulatory system, in which the blood is entirely contained in vessels rather than in a hemocoel.

The foot is lobed and subdivided into arms and tentacles. Suckers with chitinized rings are present on the arms and tentacles of octopuses and squid. Siphons are well developed and the expulsion of water is used as their primary mode of locomotion, which resembles jet propulsion. Gills (ctenidia) are attached to the wall of the mantle cavity and are serviced by large blood vessels, each with its own heart. A pair of nephridia is present within the mantle cavity for water balance and excretion of nitrogenous wastes. Cephalopods such as squids and octopuses also produce sepia or a dark ink, which contains melanin. The ink gland is located between the gills and can be released into the excurrent water stream. Ink clouds can be used either as a “smoke screen” to hide the animal from predators during a quick attempt at escape, or to create a fake image to distract predators.

Cephalopods are dioecious. Members of a species mate, and the female then lays the eggs in a secluded and protected niche. Females of some species care for the eggs for an extended period of time and may end up dying during that time period. While most other aquatic mollusks produce trochophore larvae, cephalopod eggs develop directly into a juvenile without an intervening larval stage.

Figure 7. Cephalopods. The (a) nautilus, (b) giant cuttlefish, (c) reef squid, and (d) blue-ring octopus are all members of the class Cephalopoda. (credit a: modification of work by J. Baecker credit b: modification of work by Adrian Mohedano credit c: modification of work by Silke Baron credit d: modification of work by Angell Williams)

Members of class Scaphopoda (“boat feet”) are known colloquially as “tusk shells” or “tooth shells,” as evident when examining Dentalium, one of the few remaining scaphopod genera ((Figure)). Scaphopods are usually buried in sand with the anterior opening exposed to water. These animals have a single conical shell, which is open on both ends. The head is not well developed, but the mouth, containing a radula, opens among a group of tentacles that terminate in ciliated bulbs used to catch and manipulate prey. Scaphopods also have a foot similar to that seen in bivalves. Ctenidia are absent in these animals the mantle cavity forms a tube open at both ends and serves as the respiratory structure in these animals.

Figure 8. Tooth shells. Antalis vulgaris shows the classic Dentaliidae shape that gives these animals their common name of “tusk shell.” (credit: Georges Jansoone)

Phylum Annelida

Phylum Annelida comprises the true, segmented worms. These animals are found in marine, terrestrial, and freshwater habitats, but the presence of water or humidity is a critical factor for their survival in terrestrial habitats. The annelids are often called “segmented worms” due to their key characteristic of metamerism, or true segmentation. Approximately 16,500 species have been described in phylum Annelida, which includes polychaete worms (marine annelids with multiple appendages), and oligochaetes (earthworms and leeches). Some animals in this phylum show parasitic and commensal symbioses with other species in their habitat.

Morphology

Annelids display bilateral symmetry and are worm-like in overall morphology. The name of the phylum is derived from the Latin word annullus, which means a small ring, an apt description of the ring-like segmentation of the body. Annelids have a body plan with metameric segmentation, in which several internal and external morphological features are repeated in each body segment. Metamerism allows animals to become bigger by adding “compartments,” while making their movement more efficient. The overall body can be divided into head, body, and pygidium (or tail). During development, the segments behind the head arise sequentially from a growth region anterior to the pygidium, a pattern called teloblastic growth. In the Oligochaetes, the clitellum is a reproductive structure that generates mucus to aid sperm transfer and also produces a “cocoon,” within which fertilization occurs it appears as a permanent, fused band located on the anterior third of the animal ((Figure)).

Figure 9. The clitellum of an earthworm. The clitellum, seen here as a protruding segment with different coloration than the rest of the body, is a structure that aids in oligochaete reproduction. (credit: Rob Hille)

Anatomy

The epidermis is protected by a collagenous, external cuticle, which is much thinner than the cuticle found in the ecdysozoans and does not require periodic shedding for growth. Circular as well as longitudinal muscles are located interior to the epidermis. Chitinous bristles called setae (or chaetae) are anchored in the epidermis, each with its own muscle. In the polychaetes, the setae are borne on paired appendages called parapodia.

Most annelids have a well-developed and complete digestive system. Feeding mechanisms vary widely across the phylum. Some polychaetes are filter-feeders that use feather-like appendages to collect small organisms. Others have tentacles, jaws, or an eversible pharynx to capture prey. Earthworms collect small organisms from soil as they burrow through it, and most leeches are blood-feeders armed with teeth or a muscular proboscis. In earthworms, the digestive tract includes a mouth, muscular pharynx, esophagus, crop, and muscular gizzard. The gizzard leads to the intestine, which ends in an anal opening in the terminal segment. A cross-sectional view of a body segment of an earthworm is shown in (Figure) each segment is limited by a membranous septum that divides the coelomic cavity into a series of compartments.

Most annelids possess a closed circulatory system of dorsal and ventral blood vessels that run parallel to the alimentary canal as well as capillaries that service individual tissues. In addition, the dorsal and ventral vessels are connected by transverse loops in every segment. Some polychaetes and leeches have an open system in which the major blood vessels open into a hemocoel. In many species, the blood contains hemoglobin, but not contained in cells. Annelids lack a well-developed respiratory system, and gas exchange occurs across the moist body surface. In the polychaetes, the parapodia are highly vascular and serve as respiratory structures. Excretion is facilitated by a pair of metanephridia (a type of primitive “kidney” that consists of a convoluted tubule and an open, ciliated funnel) that is present in every segment toward the ventral side. Annelids show well-developed nervous systems with a ring of fused ganglia present around the pharynx. The nerve cord is ventral in position and bears enlarged nodes or ganglia in each segment.

Figure 10. Segmental anatomy of an earthworm. This schematic drawing shows the basic anatomy of annelids in a cross-sectional view.

Annelids may be either monoecious with permanent gonads (as in earthworms and leeches) or dioecious with temporary or seasonal gonads (as in polychaetes). However, cross-fertilization is preferred even in hermaphroditic animals. Earthworms may show simultaneous mutual fertilization when they are aligned for copulation. Some leeches change their sex over their reproductive lifetimes. In most polychaetes, fertilization is external and development includes a trochophore larva, which then metamorphoizes to the adult form. In oligochaetes, fertilization is typically internal and the fertilized eggs develop in a cocoon produced by the clitellum development is direct. Polychaetes are excellent regenerators and some even reproduce asexually by budding or fragmentation.

Link to Learning

This combination video and animation provides a close-up look at annelid anatomy.

Classification of Phylum Annelida

Phylum Annelida contains the class Polychaeta (the polychaetes) and the class Oligochaeta (the earthworms, leeches, and their relatives). The earthworms and the leeches form a monophyletic clade within the polychaetes, which are therefore paraphyletic as a group.

There are more than 22,000 different species of annelids, and more than half of these are marine polychaetes (“many bristles”). In the polychaetes, bristles are arranged in clusters on their parapodia—fleshy, flat, paired appendages that protrude from each segment. Many polychaetes use their parapodia to crawl along the sea floor, but others are adapted for swimming or floating. Some are sessile, living in tubes. Some polychaetes live near hydrothermal vents. These deepwater tubeworms have no digestive tract, but have a symbiotic relationship with bacteria living in their bodies.

Earthworms are the most abundant members of the class Oligochaeta (“few bristles”), distinguished by the presence of a permanent clitellum as well as the small number of reduced chaetae on each segment. (Recall that oligochaetes do not have parapodia.) The oligochaete subclass Hirudinea, includes leeches such as the medicinal leech, Hirudo medicinalis, which is effective at increasing blood circulation and breaking up blood clots, and thus can be used to treat some circulatory disorders and cardiovascular diseases. Their use goes back thousands of years. These animals produce a seasonal clitellum, unlike the permanent clitellum of other oligochaetes. A significant difference between leeches and other annelids is the lack of setae and the development of suckers at the anterior and posterior ends, which are used to attach to the host animal. Additionally, in leeches, the segmentation of the body wall may not correspond to the internal segmentation of the coelomic cavity. This adaptation possibly helps the leeches to elongate when they ingest copious quantities of blood from host vertebrates, a condition in which they are said to be “engorged.” The subclass Brachiobdella includes tiny leechlike worms that attach themselves to the gills or body surface of crayfish.

Figure 11. Annelid groups. The (a) earthworm, (b) leech, and (c) featherduster are all annelids. The earthworm and leech are oligochaetes, while the featherduster worm is a tube-dwelling filter-feeding polychaete. (credit a: modification of work by S. Shepherd credit b: modification of work by “Sarah G…”/Flickr credit c: modification of work by Chris Gotschalk, NOAA)

Section Summary

Phylum Mollusca is a large, group of protostome schizocoelous invertebrates that occupy marine, freshwater, and terrestrial habitats. Mollusks can be divided into seven classes, each of which exhibits variations on the basic molluscan body plan. Two defining features are the mantle, which secretes a protective calcareous shell in many species, and the radula, a rasping feeding organ found in most classes. Some mollusks have evolved a reduced shell, and others have no radula. The mantle also covers the body and forms a mantle cavity, which is quite distinct from the coelomic cavity—typically reduced to the area surrounding the heart, kidneys, and intestine. In aquatic mollusks, respiration is facilitated by gills (ctenidia) in the mantle cavity. In terrestrial mollusks, the mantle cavity itself serves as an organ of gas exchange. Mollusks also have a muscular foot, which is modified in various ways for locomotion or food capture. Most mollusks have separate sexes. Early development in aquatic species occurs via one or more larval stages, including a trochophore larva, that precedes a veliger larva in some groups.

Phylum Annelida includes vermiform, segmented animals. Segmentation is metameric (i.e., each segment is partitioned internally as well as externally, with various structures repeated in each segment). These animals have well-developed neuronal, circulatory, and digestive systems. The two major groups of annelids are the polychaetes, which have parapodia with multiple bristles, and oligochaetes, which have no parapodia and fewer bristles or no bristles. Oligochaetes, which include earthworms and leeches, have a specialized band of segments known as a clitellum, which secretes a cocoon and protects gametes during reproduction. The leeches do not have full internal segmentation. Reproductive strategies include separate sexes, hermaphroditism, and serial hermaphroditism. Polychaetes typically have trochophore larvae, while the oligochaetes develop more directly.

Art Connections

(Figure) Which of the following statements about the anatomy of a mollusk is false?


Annelida

Phylum Annelida are segmented worms found in marine, terrestrial, and freshwater habitats, but the presence of water or humidity is a critical factor for their survival in terrestrial habitats. The name of the phylum is derived from the Latin word annellus, which means a small ring. Approximately 16,500 species have been described. The phylum includes earthworms, polychaete worms, and leeches. Like mollusks, annelids exhibit protostomic development.

Annelids are bilaterally symmetrical and have a worm-like appearance. Their particular segmented body plan results in repetition of internal and external features in each body segment. This type of body plan is called metamerism . The evolutionary benefit of such a body plan is thought to be the capacity it allows for the evolution of independent modifications in different segments that perform different functions. The overall body can then be divided into head, body, and tail.


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Phylum Annelida includes the classes Polychaeta and Clitellata ( [link] ) the latter contains subclasses Oligochaeta, Hirudinoidea, and Branchiobdellida.

Earthworms are the most abundant members of the subclass Oligochaeta, distinguished by the presence of the clitellum , a ring structure in the skin that secretes mucus to bind mating individuals and forms a protective cocoon for the eggs. They also have a few, reduced chaetae (oligo- = &ldquofew&rdquo -chaetae = &ldquohairs&rdquo). The number and size of chaetae is greatly diminished in oligochaetes as compared to the polychaetes (poly- = &ldquomany&rdquo -chaetae = &ldquohairs&rdquo). The chaetae of polychaetes are also arranged within fleshy, flat, paired appendages on each segment called parapodia.

The subclass Hirudinoidea includes leeches. Significant differences between leeches and other annelids include the development of suckers at the anterior and posterior ends, and the absence of chaetae. Additionally, the segmentation of the body wall may not correspond to internal segmentation of the coelomic cavity. This adaptation may allow leeches to swell when ingesting blood from host vertebrates. The subclass Branchiobdellida includes about 150 species that show similarity to leeches as well as oligochaetes. All species are obligate symbionts, meaning that they can only survive associated with their host, mainly with freshwater crayfish. They feed on the algae that grows on the carapace of the crayfish.

The (a) earthworm and (b) leech are both annelids. (credit a: modification of work by "schizoform"/Flickr credit b: modification of work by "Sarah G. "/Flickr)


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Watch the video: diversity of invertebrates 2: nematodes, annelids, mollusks (August 2022).