28.3: Superphylum Lophotrochozoa

28.3.1: Superphylum Lophotrochozoa

Learning Objective

Describe the phylogenetic position and basic features of lophotrochozoa

Key Points

• Lophotrochozoa have a blastopore, which is an involution of the ectoderm that forms a rudimentary mouth opening to the alimentary canal, a condition called protostomy or "first mouth".
• The Lophotrochozoa are comprised of the trochozoans and the lophophorata, although the exact relationships between the different phyla are not clearly determined.
• Lophophores are characterized by the presence of the lophophore, a set of ciliated tentacles surrounding the mouth; they include the flatworms and several other phyla whose relationships are upheld by genetic evidence.
• Trochophore larvae are distinguished from the lophophores by two bands of cilia around the body; they include the Nemertea, Mollusca, Sipuncula, and Annelida.
• The lophotrochozoans have a mesoderm layer positioned between the ectoderm and endoderm and are bilaterally symmetrical, which signals the beginning of cephalization, the concentration of nervous tissues and sensory organs in the head of the organism.

Key Terms

blastopore

the opening into the archenteron

lophophore

a feeding organ of brachiopods, bryozoans, and phoronids

cephalization

an evolutionary trend in which the neural and sense organs become centralized at one end (the head) of an animal

Lophotrochozoans

Animals belonging to superphylum Lophotrochozoa are protostomes: the blastopore (or the point of involution of the ectoderm or outer germ layer) becomes the mouth opening to the alimentary canal. This is called protostomy or "first mouth." In protostomy, solid groups of cells split from the endoderm or inner germ layer to form a central mesodermal layer of cells. This layer multiplies into a band which then splits internally to form the coelom; this protostomic coelom is termed schizocoelom.

As lophotrochozoans, the organisms in this superphylum possess either lophophore or trochophore larvae. The exact relationships between the different phyla are not entirely certain. The lophophores include groups that are united by the presence of the lophophore, a set of ciliated tentacles surrounding the mouth. Lophophorata include the flatworms and several other phyla, including the Bryozoa, Entoprocta, Phoronida, and Brachiopoda. These clades are upheld when RNA sequences are compared. Trochophore larvae are characterized by two bands of cilia around the body. Previously, these were treated together as the Trochozoa, together with the arthropods, which do not produce trochophore larvae, but were considered close relatives of the annelids because they are both segmented. However, they show a number of important differences. Arthropods are now placed separately among the Ecdysozoa. The Trochozoa include the Nemertea, Mollusca, Sipuncula, and Annelida.

The lophotrochozoans are triploblastic, possessing an embryonic mesoderm sandwiched between the ectoderm and endoderm found in the diploblastic cnidarians. These phyla are also bilaterally symmetrical: a longitudinal section will divide them into right and left sides that are symmetrical . They also show the beginning of cephalization: the evolution of a concentration of nervous tissues and sensory organs in the head of the organism, which is where it first encounters its environment.

Lophotrochozoans

The Caribbean Reef Squid or Sepioteuthis sepioidea is a complex lophotrochozoan. Species in this group have bilateral symmetry.

28.3.2: Phylum Platyhelminthes

Learning Objective

Differentiate among the classes of platyhelminthes

Key Points

• The Platyhelminthes are acoelomate flatworms: their bodies are solid between the outer surface and the cavity of the digestive system.
• Most flatworms have a gastrovascular cavity rather than a complete digestive system; the same cavity used to bring in food is used to expel waste materials.
• Platyhelminthes are either predators or scavengers; many are parasites that feed on the tissues of their hosts.
• Flatworms posses a simple nervous system, no circulatory or respiratory system, and most produce both eggs and sperm, with internal fertilization.
• Platyhelminthes are divided into four classes: Turbellaria, free-living marine species; Monogenea, ectoparasites of fish; Trematoda, internal parasites of humans and other species; and Cestoda (tapeworms), which are internal parasites of many vertebrates.
• In flatworms, digested materials are taken into the cells of the gut lining by phagocytosis, rather than being processed internally.

Key Terms

proglottid

any of the segments of a tapeworm; they contain both male and female reproductive organs

scolex

the structure at the rear end of a tapeworm which, in the adult, has suckers and hooks by which it attaches itself to a host

ectoparasite

a parasite that lives on the surface of a host organism

acoelomate

any animal without a coelom, or body cavity

Phylum Platyhelminthes

Phylum Platyhelminthes is composed of the flatworms: acoelomate organisms that include many free-living and parasitic forms. Most of the flatworms are classified in the superphylum Lophotrochozoa, which also includes the mollusks and annelids. The Platyhelminthes consist of two lineages: the Catenulida and the Rhabditophora. The Catenulida, or "chain worms" is a small clade of just over 100 species. These worms typically reproduce asexually by budding. However, the offspring do not fully detach from the parents; therefore, they resemble a chain. The remaining flatworms discussed here are part of the Rhabditophora.

Many flatworms are parasitic, including important parasites of humans. Flatworms have three embryonic tissue layers that give rise to surfaces that cover tissues (from ectoderm), internal tissues (from mesoderm), and line the digestive system (from endoderm). The epidermal tissue is a single layer cells or a layer of fused cells (syncytium) that covers a layer of circular muscle above a layer of longitudinal muscle. The mesodermal tissues include mesenchymal cells that contain collagen and support secretory cells that secrete mucus and other materials at the surface. The flatworms are acoelomates: their bodies are solid between the outer surface and the cavity of the digestive system.

Physiological Processes of Flatworms

The free-living species of flatworms are predators or scavengers. Parasitic forms feed on the tissues of their hosts. Most flatworms have a gastrovascular cavity rather than a complete digestive system; in such animals, the "mouth" is also used to expel waste materials from the digestive system. Some species also have an anal opening. The gut may be a simple sac or highly branched. Digestion is extracellular, with digested materials taken in to the cells of the gut lining by phagocytosis. One group, the cestodes, lacks a digestive system. Flatworms have an excretory system with a network of tubules throughout the body with openings to the environment and nearby flame cells, whose cilia beat to direct waste fluids concentrated in the tubules out of the body. The system is responsible for the regulation of dissolved salts and the excretion of nitrogenous wastes. The nervous system consists of a pair of nerve cords running the length of the body with connections between them and a large ganglion or concentration of nerves at the anterior end of the worm, where there may also be a concentration of photosensory and chemosensory cells.

There is neither a circulatory nor respiratory system, with gas and nutrient exchange dependent on diffusion and cell-cell junctions. This necessarily limits the thickness of the body in these organisms, constraining them to be "flat" worms. In addition, most flatworm species are monoecious; typically, fertilization is internal. Asexual reproduction is common in some groups.

Diversity of Flatworms

Platyhelminthes are traditionally divided into four classes: Turbellaria, Monogenea, Trematoda, and Cestoda. The class Turbellaria includes mainly free-living, marine species, although some species live in freshwater or moist terrestrial environments . The ventral epidermis of turbellarians is ciliated which facilitates their locomotion. Some turbellarians are capable of remarkable feats of regeneration: they may regrow the entire body from a small fragment.

Turbellaria

Pseudobiceros bedfordi (Bedford's Flatworm), a member of the Turbellaria, is a marine species which uses the epidermis of its belly for locomotion.

The monogeneans are ectoparasites, mostly of fish, with simple life cycles that consist of a free-swimming larva that attaches to a fish to begin transformation to the parasitic adult form. The worms may produce enzymes that digest the host tissues or simply graze on surface mucus and skin particles.

The trematodes, or flukes, are internal parasites of mollusks and many other groups, including humans . Trematodes have complex life cycles that involve a primary host in which sexual reproduction occurs and one or more secondary hosts in which asexual reproduction occurs. The primary host is almost always a mollusk. Trematodes are responsible for serious human diseases including schistosomiasis, a blood fluke.

Trematodes

Botulus microporus is a trematode that lives only in the intestinal tract of Lancetfish, Alepisaurus ferox.

The cestodes, or tapeworms, are also internal parasites, mainly of vertebrates . Tapeworms live in the intestinal tract of the primary host, remaining fixed by using a sucker on the anterior end, or scolex, of the tapeworm body. The remainder of the tapeworm is composed of a long series of units called proglottids. Each may contain an excretory system with flame cells and both female and male reproductive structures. Tapeworms do not possess a digestive system; instead, they absorb nutrients from the food matter passing through them in the host's intestine.

Cestodes

Taenia saginata, also known as Taeniarhynchus saginata or the beef tapeworm, is a parasite of both cattle and humans. It displays the long series of proglottid subunits characteristic of the species.

28.3.3: Phylum Rotifera

Learning Objective

Identify the features of rotifers involved in movement and feeding

Key Points

• The rotifer body form consists of a head (containing the sensory organs in the form of a bi-lobed brain and small eyespots near the corona), the trunk (containing organs), and the foot (which can hold fast).
• The foot of the rotifer secretes a sticky material to help it adhere to surfaces.
• Rotifers are filter feeders that generate a current using the corona to pass food into the mouth, which then passes by digestive and salivary glands into the stomach and intestines.
• Rotifers exhibit sexual dimorphism; the gender of many species is determined by whether the egg is fertilized (and develops into a female) or unfertilized (and develops into a male).

Key Terms

pseudocoelomate

any invertebrate animal with a three-layered body and a pseudocoel

mastax

the pharynx of a rotifer which usually contains four horny pieces that work to crush the food

Phylum Rotifera

The rotifers are a microscopic (about 100 µm to 30 mm) group of mostly-aquatic organisms that get their name from the corona: a rotating, wheel-like structure that is covered with cilia at their anterior end. Although their taxonomy is currently in flux, one treatment places the rotifers in three classes: Bdelloidea, Monogononta, and Seisonidea. The classification of the group is currently under revision, however, as more phylogenetic evidence becomes available. It is possible that the "spiny headed worms" currently in phylum Acanthocephala will be incorporated into this group in the future.

Rotifers

A bdelloid rotifer is a member of a class of rotifers found in fresh water and moist soil. The rotifer body consists of a head, a truck, and a foot. They eat by filtering food into the mouth by creating currents with the corona.

The rotifer body form consists of a head (which contains the corona), a trunk (which contains the organs), and the foot. Rotifers are typically free-swimming and truly planktonic organisms, but the toes or extensions of the foot can secrete a sticky material forming a holdfast to help them adhere to surfaces. The head contains sensory organs in the form of a bi-lobed brain and small eyespots near the corona.

The rotifers are filter feeders that will eat dead material, algae, and other microscopic living organisms. Therefore, they are very important components of aquatic food webs. Rotifers obtain food that is directed toward the mouth by the current created from the movement of the corona. The food particles enter the mouth and travel to the mastax (pharynx with jaw-like structures). Food passes by digestive and salivary glands into the stomach and then into the intestines. Digestive and excretory wastes are collected in a cloacal bladder before being released out the anus.

Rotifers are pseudocoelomates commonly found in fresh water and some salt water environments throughout the world. About 2,200 species of rotifers have been identified. Rotifers are dioecious organisms (having either male or female genitalia) and exhibit sexual dimorphism (males and females have different forms). Many species are parthenogenic and exhibit haplodiploidy, a method of gender determination in which a fertilized egg develops into a female and an unfertilized egg develops into a male. In many dioecious species, males are short-lived and smaller, with no digestive system and a single testis. Females can produce eggs that are capable of dormancy, which protects eggs during harsh environmental conditions.

28.3.4: Phylum Nemertea

Learning Objective

Identify the key features of the Phylum Nemertea

Key Points

• The Nemertini are mostly bottom-dwelling marine organisms, although some are found in freshwater and terrestrial habitats.
• Most nemerteans are carnivores, some are scavengers, and others have evolved relationships with some mollusks that are benefit the Nemertean but do not harm the mollusk.
• Nemerteans vary greatly in size and are bilaterally symmetrical; they are unsegmented and resemble a flat tube which can change morphological presentation in response to environmental cues.
• Nemertini have a simple nervous system comprised of a ring of four nerve masses called "ganglia" at the anterior end between the mouth and the foregut from which paired longitudinal nerve cords emerge and extend to the posterior end.
• Nemertini are mostly sexually dimorphic, fertilizing eggs externally by releasing both eggs and sperm into the water; a larva may develop inside the resulting young worm and devour its tissues before metamorphosing into the adult.

Key Terms

proboscis

an elongated tube from the head or connected to the mouth, of an animal

rhynchocoel

a cavity which mostly runs above the midline and ends a little short of the rear of the body of a nemertean and extends or retracts the proboscis

protonephridia

an invertebrate organ which occurs in pairs and removes metabolic wastes from an animal's body

Phylum Nemertea

The Nemertea are colloquially known as ribbon worms. Most species of phylum Nemertea are marine (predominantly benthic or bottom dwellers) with an estimated 900 species known. However, nemertini have been recorded in freshwater and terrestrial habitats as well. Most nemerteans are carnivores, feeding on worms, clams, and crustaceans. Some species are scavengers, while other nemertini species, such as Malacobdella grossa, have also evolved commensalistic relationships with some mollusks. Interestingly, nemerteans have almost no predators, two species are sold as fish bait, and some species have devastated commercial fishing of clams and crabs.

Morphology

Ribbon worms vary in size from 1 cm to several meters. They show bilateral symmetry and remarkable contractile properties. Because of their contractility, they can change their morphological presentation in response to environmental cues. Animals in phylum Nemertea also show a flattened morphology: they are flat from front to back, like a flattened tube. In addition, nemertea are soft, unsegmented animals .

Morphology of Nemertea

A terrestrial Geonemertes, a Nemertean, displaying the flat, ribbon-like body of the organism that is unsegmented.

A unique characteristic of this phylum is the presence of a proboscis enclosed in a rhynchocoel. The proboscis serves to capture food and may be ornamented with barbs in some species. The rhynchocoel is a fluid-filled cavity that extends from the head to nearly two-thirds of the length of the gut in these animals . The proboscis may be extended or retracted by the retractor muscle attached to the wall of the rhynchocoel.

Internal structures of the Nemertini

This image shows the internal structures of a basic nemertean, including the proboscis and the rhynchocoel: 1: Proboscis 2: Rhynchocoel 3: Dorsal commissure of brain 4: Rhynchodeum 5: Proboscis pore 6: Ventral commissure of brain 7: Mouth 8: Foregut 9: Stomach

Metabolism

The nemertini show a very well-developed digestive system. A mouth opening that is ventral to the rhynchocoel leads into the foregut, followed by the intestine. The intestine is present in the form of diverticular pouches which ends in a rectum that opens via an anus. Gonads are interspersed with the intestinal diverticular pouches, opening outwards via genital pores. A circulatory system consists of a closed loop of a pair of lateral blood vessels. The circulatory system is derived from the coelomic cavity of the embryo. Some animals may also have cross-connecting vessels in addition to lateral ones. Although these are called blood vessels, since they are of coelomic origin, the circulatory fluid is colorless. Some species bear hemoglobin as well as yellow or green pigments. The blood vessels are connected to the rhynchocoel. The flow of fluid in these vessels is facilitated by the contraction of muscles in the body wall. A pair of protonephridia, or primitive kidneys, is present in these animals to facilitate osmoregulation. Gaseous exchange occurs through the skin in the nemertini.

Nervous System

Nemertini have a ganglion or "brain" situated at the anterior end between the mouth and the foregut, surrounding the digestive system as well as the rhynchocoel. A ring of four nerve masses called "ganglia" comprises the brain in these animals. Paired longitudinal nerve cords emerge from the brain ganglia, extending to the posterior end. Ocelli or eyespots are present in pairs, in multiples of two in the anterior portion of the body. It is speculated that the eyespots originate from neural tissue and not from the epidermis.

Reproduction

Animals in phylum Nemertea show sexual dimorphism, although freshwater species may be hermaphroditic. Eggs and sperm are released into the water; fertilization occurs externally. The zygote develops into a special kind of nemertean larvae called a planuliform larva. In some nemertine species, another larva specific to the nemertinis, a pilidium, may develop inside the young worm from a series of imaginal discs. This larval form, characteristically shaped like a deerstalker cap, devours tissues from the young worm for survival before metamorphosing into the adult-like morphology.

28.3.5: Phylum Mollusca

Learning Objective

Describe the unique anatomical and morphological features of molluscs

Key Points

• A mollusk's muscular foot is used for locomotion and anchorage, varies in shape and function, and can both extend and retract.
• The visceral mass inside the mollusk includes digestive, nervous, excretory, reproductive, and respiratory systems.
• Most mollusks possess a radula, which is similar to a tongue with teeth-like projections, serving to shred or scrape food.
• The mantle is the dorsal epidermis in mollusks; in some mollusks it secretes a chitinous and hard calcareous shell.

Key Terms

visceral mass

the soft, non-muscular metabolic region of the mollusc that contains the body organs

mantle

the body wall of a mollusc, from which the shell is secreted

radula

the rasping tongue of snails and most other mollusks

Phylum Mollusca

Phylum Mollusca is the predominant phylum in marine environments. It is estimated that 23 percent of all known marine species are mollusks; there are around 85,000 described species, making them the second most diverse phylum of animals. The name "mollusca" signifies a soft body; the earliest descriptions of mollusks came from observations of unshelled cuttlefish. Mollusks are predominantly a marine group of animals; however, they are known to inhabit freshwater as well as terrestrial habitats. Mollusks display a wide range of morphologies in each class and subclass. They range from large predatory squids and octopus, some of which show a high degree of intelligence, to grazing forms with elaborately-sculpted and colored shells. In spite of their tremendous diversity, however, they also share a few key characteristics, including a muscular foot, a visceral mass containing internal organs, and a mantle that may or may not secrete a shell of calcium carbonate .

Mollusk shells

Helix aspersa, a common land snail, has a calcium carbonate shell.

Mollusks have a muscular foot used for locomotion and anchorage that varies in shape and function, depending on the type of mollusk under study. In shelled mollusks, this foot is usually the same size as the opening of the shell. The foot is a retractable as well as an extendable organ. It is the ventral-most organ, whereas the mantle is the limiting dorsal organ. Mollusks are eucoelomate, but the cavity is restricted to a region around the heart in adult animals. The mantle cavity develops independently of the coelomic cavity.

The visceral mass is present above the foot in the visceral hump. This includes digestive, nervous, excretory, reproductive, and respiratory systems. Mollusk species that are exclusively aquatic have gills for respiration, whereas some terrestrial species have lungs for respiration. Additionally, a tongue-like organ called a radula, which bears chitinous tooth-like ornamentation, is present in many species, serving to shred or scrape food. The mantle (also known as the pallium) is the dorsal epidermis in mollusks; shelled mollusks are specialized to secrete a chitinous and hard calcareous shell.

A "generalized mollusk"

An anatomical diagram of a hypothetical ancestral mollusk, showing features common to many mollusk types.

Most mollusks are dioecious animals where fertilization occurs externally, although this is not the case in terrestrial mollusks, such as snails and slugs, or in cephalopods. In some mollusks, the zygote hatches and undergoes two larval stages, trochophore and veliger, before becoming a young adult; bivalves may exhibit a third larval stage, glochidia.

28.3.6: Classification of Phylum Mollusca

Learning Objective

Differentiate among the classes in the phylum mollusca

Key Points

• Mollusks can be segregated into seven classes: Aplacophora, Monoplacophora, Polyplacophora, Bivalvia, Gastropoda, Cephalopoda, and Scaphopoda. These classes are distinguished by, among other criteria, the presence and types of shells they possess.
• Class Aplacophora includes worm-like animals with no shell and a rudimentary body structure.
• Members of class Monoplacophora have a single shell that encloses the body.
• Members of class Polyplacophora are better known as "chitons;" these molluscs have a large foot on the ventral side and a shell composed of eight hard plates on the dorsal side.
• Class Bivalvia consists of mollusks with two shells held together by a muscle; these include oysters, clams, and mussels.
• Members of class Gastropoda have an asymmetrical body plan and usually have a shell, which can be planospiral or conispiral. Their key characteristic is the torsion around the perpendicular axis on the center of the foot that is modified for crawling.
• Class Scaphopoda consists of mollusks with a single conical shell through which the head protrudes, and a foot modified into tentacles known as captaculae that are used to catch and manipulate prey.

Key Terms

ctenidium

a respiratory system, in the form of a comb, in some molluscs

captacula

the foot of a Scaphalopod, modified into tentacles for capturing prey

nephridium

a tubular excretory organ in some invertebrates

Classes in Phylum Mollusca

Phylum Mollusca is a very diverse (85,000 species) group of mostly marine species, with a dramatic variety of form. This phylum can be segregated into seven classes: Aplacophora, Monoplacophora, Polyplacophora, Bivalvia, Gastropoda, Cephalopoda, and Scaphopoda.

Class Aplacophora

Class Aplacophora ("bearing no plates") includes worm-like animals primarily found in benthic marine habitats. These animals lack a calcareous shell, but possess aragonite spicules on their epidermis. They have a rudimentary mantle cavity and lack eyes, tentacles, and nephridia (excretory organs).

Class Monoplacophora

Members of class Monoplacophora ("bearing one plate") posses a single, cap-like shell that encloses the body. The morphology of the shell and the underlying animal can vary from circular to ovate. A looped digestive system, multiple pairs of excretory organs, many gills, and a pair of gonads are present in these animals. The monoplacophorans were believed extinct and only known via fossil records until the discovery of Neopilina galathaea in 1952. Today, scientists have identified nearly two dozen extant species.

Class Polyplacophora

Animals in the class Polyplacophora ("bearing many plates") are commonly known as "chitons" and bear an armor-like, eight-plated dorsal shell. These animals have a broad, ventral foot that is adapted for suction to rocks and other substrates, and a mantle that extends beyond the shell in the form of a girdle. Calcareous spines may be present on the girdle to offer protection from predators. Chitons live worldwide, in cold water, warm water, and the tropics. Most chiton species inhabit intertidal or subtidal zones, and do not extend beyond the photic zone. Some species live quite high in the intertidal zone and are exposed to the air and light for long periods.

Chiton morphology

The underside of the gumboot chiton, Cryptochiton stellari, showing the foot in the center, surrounded by the gills and mantle. The mouth is visible to the left in this image.

Class Bivalvia

Bivalvia is a class of marine and freshwater molluscs with laterally compressed bodies enclosed by a shell in two hinged parts. Bivalves include clams, oysters, mussels, scallops, and numerous other families of shells. The majority are filter feeders and have no head or radula. The gills have evolved into ctenidia, specialised organs for feeding and breathing. Most bivalves bury themselves in sediment on the seabed, while others lie on the sea floor or attach themselves to rocks or other hard surfaces.

The shell of a bivalve is composed of calcium carbonate, and consists of two, usually similar, parts called valves. These are joined together along one edge by a flexible ligament that, in conjunction with interlocking "teeth" on each of the valves, forms the hinge.

Empty shell of a bivalve

The empty shell of the giant clam, Tridacna gigas. Note the pair of shells that are hinged together, a characteristic of members of the class Bivalvia.

Class Gastropoda

Animals in class Gastropoda ("stomach foot") include well-known mollusks like snails, slugs, conchs, sea hares, and sea butterflies. Gastropoda includes shell-bearing species as well as species with a reduced shell. These animals are asymmetrical and usually present a coiled shell. 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.

The visceral mass in the shelled species displays torsion around the perpendicular axis on the center of the foot, which is the key characteristic of this group, along with a foot that is modified for crawling . Most gastropods bear a head with tentacles, eyes, and a style. A complex radula is used by the digestive system and aids in the ingestion of food. Eyes may be absent in some gastropods species. The mantle cavity encloses the ctenidia (singluar: ctenidium) as well as a pair of nephridia (singular: nephridium).

Gastropod foot

Gastropods, such as this Roman snail, have a large foot that is modified for crawling.

Class Cephalopoda

Class Cephalopoda ("head foot" animals) includes octopi, squids, cuttlefish, and nautilus. Cephalopods are a class of shell-bearing animals as well as mollusks with a reduced shell. They display vivid coloration, typically seen in squids and octopi, which is used for camouflage. All animals in this class are carnivorous predators and have beak-like jaws at the anterior end. All cephalopods show the presence of a very well-developed nervous system along with eyes, as well as a closed circulatory system. The foot is lobed and developed into tentacles and a funnel, which is used as the mode of locomotion. Locomotion in cephalopods is facilitated by ejecting a stream of water for propulsion ("jet" propulsion) . Cephalopods, such as squids and octopi, also produce sepia or a dark ink, which is squirted upon a predator to assist in a quick getaway. Suckers are present on the tentacles in octopi and squid. Ctenidia are enclosed in a large mantle cavity serviced by blood vessels, each with its own associated heart. The mantle has siphonophores that facilitate exchange of water.

Cephalopods

Cephalopods ("head foot") include this octopus, which ejects a stream of water from a funnel in its body to propel itself through the water.

A pair of nephridia is present within the mantle cavity. Sexual dimorphism is seen in this class of animals. Members of a species mate, then the female 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. Reproduction in cephalopods is different from other mollusks in that the egg hatches to produce a juvenile adult without undergoing the trochophore and veliger larval stages.

Class Scaphopoda

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. Scaphopods are usually buried in sand with the anterior opening exposed to water. These animals bear a single conical shell, which has both ends open . The head is rudimentary and protrudes out of the posterior end of the shell. These animals do not possess eyes, but they have a radula, as well as a foot modified into tentacles with a bulbous end, known as captaculae. Captaculae serve to catch and manipulate prey. Ctenidia are absent in these animals.

The Scaphopods

The Scaphopods ("boat feet") include the Antalis vulgaris, the shell of which is depicted here.

28.3.7: Phylum Annelida

Learning Objective

Describe the morphological and anatomical features of annelids

Key Points

• Annelids are often called "segmented worms" because they possess true segmentation of their bodies, with both internal and external morphological features repeated in each body segment.
• The clitellum is a structure on the anterior portion of the worm that generates mucus to aid in sperm transfer from one worm to another; it also forms a cocoon within which fertilization occurs.
• Most annelids have chitinous hairlike extensions in every segment called chaetae that are anchored in the epidermis, although the number and size of chaetae can vary in the different classes.
• Annelids possess a closed circulatory system, lack a well-developed respiratory system, but have well-developed nervous systems.
• Annelids can either have distinct male and female forms or be hermaphrodites (having both male and female reproductive organs). Earthworms are hermaphrodites and can self-fertilize, but prefer to cross-fertilize if possible.

Key Terms

metamerism

the segmentation of the body into similar discrete units

chaeta

a chitinous bristle of an annelid worm

clitellum

a glandular swelling in the epidermis of some annelid worms; it secretes a viscous fluid in which the eggs are deposited

Phylum Annelida

Phylum Annelida contains the class Polychaeta (the polychaetes) and the class Oligochaeta (the earthworms, leeches, and their relatives). These animals are found in marine, terrestrial, and freshwater habitats, but a presence of water or humidity is a critical factor for their survival, especially in terrestrial habitats. The name of the phylum is derived from the Latin word annellus, which means a small ring. Animals in this phylum show parasitic and commensal symbioses with other species in their habitat. Approximately 16,500 species have been described in phylum Annelida. The phylum includes earthworms, polychaete worms, and leeches. Annelids show protostomic development in embryonic stages and are often called "segmented worms" due to their key characteristic of metamerism, or true segmentation.

Morphology

Annelids display bilateral symmetry and are worm-like in overall morphology. They have a segmented body plan where the 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. This metamerism is thought to arise from identical teloblast cells in the embryonic stage, which develop into identical mesodermal structures. The overall body can be divided into head, body, and pygidium (or tail). The clitellum is a reproductive structure that generates mucus that aids in sperm transfer and gives rise to a cocoon within which fertilization occurs; it appears as a fused band in the anterior third of the animal .

Clitellum

The clitellum is the reproductive structure of an annelid. It creates mucus that aids in sperm transfer and gives rise to a cocoon within which fertilization occurs. It can be seen in this image as the enlarged band around the animal.

Anatomy

The epidermis is protected by an acellular, external cuticle, but this 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 hairlike extensions, anchored in the epidermis and projecting from the cuticle, called setae/chaetae are present in every segment. Annelids show the presence of a true coelom, derived from embryonic mesoderm and protostomy. Hence, they are the most advanced worms. A well-developed and complete digestive system is present in earthworms (oligochaetes) with a mouth, muscular pharynx, esophagus, crop, and gizzard being present. The gizzard leads to the intestine and ends in an anal opening. Each segment is limited by a membranous septum that divides the coelomic cavity into a series of compartments.

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, these vessels are connected by transverse loops in every segment. These animals lack a well-developed respiratory system; gas exchange occurs across the moist body surface. 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 towards the ventral side. Annelids show well-developed nervous systems with a nerve ring of fused ganglia present around the pharynx. The nerve cord is ventral in position, bearing enlarged nodes or ganglia in each segment.

Annelids may be either monoecious, with permanent gonads (as in earthworms and leeches), or dioecious, with temporary or seasonal gonads that develop (as in polychaetes). However, cross-fertilization is preferred in hermaphroditic animals. These animals may also show simultaneous hermaphroditism, participating in simultaneous sperm exchange when they are aligned for copulation.

Earthworms are the most abundant members of the class Oligochaeta, distinguished by the presence of the clitellum as well as few, reduced chaetae ("oligo- = "few"; -chaetae = "hairs"). The number and size of chaetae are greatly diminished in Oligochaeta compared to the polychaetes (poly=many, chaetae = hairs). The many chetae of polychaetes are also arranged within fleshy, flat, paired appendages that protrude from each segment. These parapodia may be specialized for different functions in the polychates. A significant difference between leeches and other annelids is the development of suckers at the anterior and posterior ends and an absence of chaetae . Additionally, 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.

Leeches

Unlike earthworms, leeches lack chaetae and have suckers at both ends of the body.

Attributions

• Superphylum Lophotrochozoa
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  • "cephalization." http://en.wiktionary.org/wiki/cephalization. Wiktionary CC BY-SA 3.0.
  • "lophophore." http://en.wiktionary.org/wiki/lophophore. Wiktionary CC BY-SA 3.0.
  • "OpenStax College, Biology. October 23, 2013." http://cnx.org/content/m44665/latest/?collection=col11448/latest. OpenStax CNX CC BY 3.0.
  • "Caribbean reef squid." http://en.wikipedia.org/wiki/File:Caribbean_reef_squid.jpg. Wikipedia Public domain.
• Phylum Platyhelminthes
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  • "proglottid." http://en.wiktionary.org/wiki/proglottid. Wiktionary CC BY-SA 3.0.
  • "acoelomate." http://en.wiktionary.org/wiki/acoelomate. Wiktionary CC BY-SA 3.0.
  • "OpenStax College, Biology. October 17, 2013." http://cnx.org/content/m44665/latest/?collection=col11448/latest. OpenStax CNX CC BY 3.0.
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  • "Bedford's Flatworm." http://en.wikipedia.org/wiki/File:Bedford's_Flatworm.jpg. Wikipedia Public domain.
• Phylum Rotifera
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• Phylum Nemertea
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  • "protonephridia." http://en.wikipedia.org/wiki/protonephridia. Wikipedia CC BY-SA 3.0.
  • "rhynchocoel." http://en.wiktionary.org/wiki/rhynchocoel. Wiktionary CC BY-SA 3.0.
  • "OpenStax College, Biology. October 17, 2013." http://cnx.org/content/m44665/latest/?collection=col11448/latest. OpenStax CNX CC BY 3.0.
  • "proboscis." http://en.wiktionary.org/wiki/proboscis. Wiktionary CC BY-SA 3.0.
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  • "Nemertea Anopla n Enopla Head Sagittal." http://en.wikipedia.org/wiki/File:Nemertea_Anopla_n_Enopla_Head_Sagittal.png. Wikipedia CC BY-SA.
• Phylum Mollusca
  • "OpenStax College, Biology. October 26, 2013." http://openstaxcollege.org/textbooks/biology. OpenStax CNX CC BY-SA 3.0.
  • "visceral mass." http://en.wikipedia.org/wiki/visceral%20mass. Wikipedia CC BY-SA 3.0.
  • "Mollusca." http://en.wikipedia.org/wiki/Mollusca. Wikipedia CC BY-SA 3.0.
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  • "radula." http://en.wiktionary.org/wiki/radula. Wiktionary CC BY-SA 3.0.
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• Classification of Phylum Mollusca
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  • "Bivalvia." http://en.wikipedia.org/wiki/Bivalvia. WIKIPEDIA CC BY-SA 3.0.
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  • "Octopus3." http://en.wikipedia.org/wiki/File:Octopus3.jpg. Wikipedia CC BY-SA.
  • "Grapevinesnail 01a." http://en.wikipedia.org/wiki/File:Grapevinesnail_01a.jpg. Wikipedia CC BY-SA.
  • "Black Chiton." http://commons.wikimedia.org/wiki/File:Black_Chiton.jpg. Wikimedia Commons CC BY-SA 3.0.
  • "Black Chiton." http://commons.wikimedia.org/wiki/File:Black_Chiton.jpg. Wikimedia Commons CC BY-SA.
  • "File:Tridacna gigas.001 - Aquarium Finisterrae.JPG." http://en.wikipedia.org/wiki/File:Tridacna_gigas.001_-_Aquarium_Finisterrae.JPG. Wikipedia CC BY-SA 3.0.
  • "File:Cryptochiton stelleri underside.jpg." http://en.wikipedia.org/wiki/File:Cryptochiton_stelleri_underside.jpg. Wikipedia CC BY-SA 3.0.
  • "Black Chiton." http://commons.wikimedia.org/wiki/File:Black_Chiton.jpg. Wikimedia Commons CC BY-SA 3.0.
• Phylum Annelida
  • "Boundless." http://www.boundless.com/. Boundless Learning CC BY-SA 3.0.
  • "OpenStax College, Biology. October 17, 2013." http://cnx.org/content/m44665/latest/?collection=col11448/latest. OpenStax CNX CC BY 3.0.
  • "chaeta." http://en.wiktionary.org/wiki/chaeta. Wiktionary CC BY-SA 3.0.
  • "clitellum." http://en.wiktionary.org/wiki/clitellum. Wiktionary CC BY-SA 3.0.
  • "metamerism." http://en.wiktionary.org/wiki/metamerism. Wiktionary CC BY-SA 3.0.
  • "Egel als Schneckenparasit 04." http://en.wikipedia.org/wiki/File:Egel_als_Schneckenparasit_04.JPG. Wikipedia GNU FDL.
  • "Regenwurm1." http://en.wikipedia.org/wiki/File:Regenwurm1.jpg. Wikipedia GNU FDL.