Dictionary:
an·ne·lid (ăn'ə-lĭd)  also an·nel·i·dan
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[From New Latin Annelida, phylum name, from French annelés, pl. past participle of anneler, to ring, from Old French anel, ring, from Latin ānellus, diminutive of ānus, ring.]
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Annelid Characteristics
Segmented Bodies
The fundamental characteristic of the phylum is the division of the body into a linear series of cylindrical segments, or metameres. Each metamere consists of a section of the body wall and a compartment of the body cavity with its internal organs. The external divisions, which may be seen in the common earthworm, correspond to the internal divisions. The annelid body consists of a head region; a trunk, made up of metameres; and an unsegmented terminal region called the pygidium. In some primitive members of the phylum the metameres are identical, or very similar to one another, each containing the same structures; in more advanced forms there is a tendency toward a consolidation of some segments and a restriction of certain organs to particular segments. Because of the soft nature of the annelid body, fossils are not common. Fossils of tube-dwelling polychaetes have been found, but there is scarcely any fossil record for earthworms and none for leeches.
The Body Wall
The body wall is covered with epidermis overlaid with a thin, pliant cuticle secreted by the epidermal cells. The body wall consists of well-developed, segmentally arranged muscles used for crawling and swimming movements. Most annelids possess short external bristles called setae, or chaetae, composed of chitin. Setae are used to grip the soil, to hold the animal in a tube, or to increase the surface areas of appendages for swimming.
Digestion
The digestive system of annelids consists of an unsegmented gut that runs through the middle of the body from the mouth, located on the underside of the head, to the anus, which is on the pygidium. The gut is separated from the body wall by the body cavity, called the coelom. The segmented compartments of the coelom are usually separated from each other by thin sheets of tissue, called septa, which are perforated by the gut and by blood vessels. Except in the leeches, the coelom is fluid filled and functions as a skeleton, providing the animal with rigidity and the resistance necessary for muscular movement. If the worm is punctured, it loses its ability to move properly, since functioning of the body muscles is dependent on the maintenance of the fluid volume in the coelom. In primitive annelids each compartment of the coelom is connected to the outside by ducts for the release of sex cells, and by paired excretory organs, or nephridia. These openings are closed except when functioning, thus preventing the loss of coelomic fluid. In more advanced species both excretory and reproductive functions are sometimes served by a single type of duct, and ducts may be absent in certain segments.
Circulation
Characteristics of the circulatory system vary within the phylum. The blood usually contains hemoglobin, a red oxygen-carrying pigment; some annelids have a green oxygen-carrying pigment, and others have unpigmented blood. The circulatory system is usually closed, i.e., confined within well-developed blood vessels; in some polychaetes and leeches the circulatory system is partly open, with blood and coelomic fluid mixing directly in the sinuses of the body cavity. Blood flows toward the head through a contractile vessel above the gut and returns to the terminal region through vessels below the gut; it is distributed to each body compartment by lateral vessels. Some of the lateral vessels are contractile and serve as hearts, i.e., pumping organs for driving the blood.
Respiration
Some aquatic annelids have thin-walled, feathery gills through which gases are exchanged between the blood and the environment. However, most annelids have no special organs for gas exchange, and respiration occurs directly through the body wall.
The Nervous System
The nervous system typically consists of a primitive brain, or ganglionic mass, located in the head region, connected by a ring of nerves to a ventral nerve cord that runs the length of the body; the cord gives rise to lateral nerves and ganglia in each segment. Sense organs of annelids generally include eyes, taste buds, tactile tentacles, and organs of equilibrium called statocysts.
Reproduction
Reproduction is sexual or asexual. Asexual reproduction is by fragmentation, budding, or fission. Among sexually reproducing annelids hermaphrodites are common, but most species have separate sexes. Fertilized eggs of marine annelids usually develop into free-swimming larvae. Eggs of terrestrial forms are enclosed in cocoons and hatch as miniature versions of the adults. The ability to regenerate lost body parts is highly developed in many polychaetes and digochaetes.
Class Polychaeta
The vast majority of the more than 8,000 known species of polychaete worms are marine; some, however, are found in fresh or brackish water. They are abundant from the intertidal zone to depths of over 16,405 ft (5,000 m). The polychaetes, so named because of the numerous setae (chaetae) they bear, range in length from less than 1/8 in. to more than 9 ft (2 mm to 3 m), but most are from 2 to 4 in. (5-10 cm) long. Their colors are often brilliant, and some species are iridescent. The class has usually been divided on the basis of mode of existence into two groups, the errantia and the sedentaria.
Errant Polychaetes
Errant polychaetes include actively crawling or swimming forms which may, however, also spend time in burrows or crevices, or under rocks on the seashore. A familiar errant polychaete is the clamworm, Nereis, widely used as bait. Errant polychaetes swim, crawl over the ocean bottom, or tunnel through surface sediments. Many are predators on small invertebrates; some are scavengers. In most the first few body segments bear sensory projections called cirri, while the remaining body segments bear conspicuous leglike appendages called parapodia. The parapodia, along with undulations of the body, propel the worm in crawling and swimming; parapodia are tipped with bundles of setae, usually made of chitin. Most errant polychaetes have well-developed head regions, which bear eyes, sensory tentacles, and a specialized organ, the nuchal organ, thought to detect chemicals. The anterior end of the gut often forms a protrusible structure, the proboscis, sometimes equipped with strong chitinous jaws and used in feeding. The setae of some polychaetes, e.g., the tropical fireworm, are composed of calcium carbonate rather than chitin and are hollow. These brittle setae are easily broken off and contain a toxin that produces a painful reaction in humans. In the scaleworms, a series of overlapping scales form a covering over the animal's upper surface. In the sea mouse these scales are completely covered by long, slender, feltlike setae projecting from the parapodia.
Sedentary Polychaetes
Sedentary polychaetes are usually adapted to living permanently in tubes or burrows; some attach themselves to rocks or piers. Many sedentary polychaetes, like the lugworm, Arenicola, live in burrows in sand or mud. The majority, however, are tube builders. Tubes of different species vary greatly in their composition and structure. They may be composed of sand, shell, or other particles held together with mucus, or made entirely of organic substances secreted by the worm that harden on contact with water. The tubes may be straight, branched, spiraled, or U-shaped. Most are permanently attached to a substrate, and the worm seldom or never ventures outside; however, the tube worm Cistenides moves about the seafloor, dragging along its delicate tube of sand grains. Sedentary polychaetes have greatly modified head regions for specialized feeding habits. Many are adapted for feeding on organic matter deposited on the ocean floor. For example, the lugworms have a simple, thin-walled, jawless proboscis, which is used to draw sand into the gut, where organic matter is removed. Other worms have feeding tentacles that extend from the tube opening and creep along the mud or sand, picking up organic deposits. Still others of the Sedentaria are filter feeders: the beautiful feather-duster worms have a crown of feathery, ciliated tentacles that extend from the tube opening to sweep small planktonic organisms from the water. The tentacles are quickly withdrawn if the animal is startled. The parapodia are reduced in the sedentary polychaetes, and the setae of many tube-dwelling forms are hooked to help the worm hold itself to the wall of its tube.
Polychaete Anatomy
The structure of the digestive tract of polychaetes is variable, reflecting the diversity of feeding types. Respiration is entirely through the body wall in some polychaetes, and partially so in most. Many species have thin-walled extensions of the body surface, i.e., gills, used for gas exchange; most commonly the gills are extensions of the parapodia. The tentacles of feather-duster worms are used for respiratory exchange as well as for feeding. A polychaete may have a single pair of excretory tubes or a pair in each segment. Sedentary polychaetes have various modifications to insure that wastes will be deposited near the mouth of the tube or burrow, where they are washed away.
Polychaete Reproduction
Most polychaetes reproduce sexually, and the sexes are separate. Sex cells develop from masses of tissue in the metameres and leave by way of tubules or by rupture of the body wall. In most cases fertilization of the eggs by sperm occurs externally in seawater and results in the formation of free-swimming larvae. Variations include internal fertilization, laying of egg masses that are attached to objects with mucus, and brooding of developing eggs in the worm's body. Some errant polychaetes, including the clamworm, undergo extreme changes in appearance and become active swimmers at the time of year that the sex cells mature; males and females swarm to the surface of the sea to spawn. In some of these species the portion of the body containing the sex cells breaks free and engages in swarming and spawning, leaving the asexual portion behind to regenerate its lost parts. Swarming generally occurs at night and is correlated with particular phases of the moon. Some species perform a kind of nuptial dance, swimming in circles as they spawn. In some species the worms liberate a luminous secretion, which produces circles of light on the ocean surface as they dance. The most famous swarming polychaete is the tropical palolo worm, a name sometimes applied to all swarming polychaetes.
Archiannelida and Myzostomaria
Two groups of polychaetes that are sometimes regarded as separate classes are the Archiannelida and the Myzostomaria. The former group includes a variety of minute marine worms living in surface mud, in tidepools near the high-tide line, and in the interstitial spaces of mud and sand in some subtidal areas. All archiannelids are scavengers. They have a ciliated epidermis and only a few body segments; many resemble the larvae of other polychaetes. The Myzostomaria are a small group of marine worms parasitic on certain echinoderms (crinoids, starfish, and brittlestars). They are disk-shaped and flattened, with a series of reduced parapodia with hooked setae; they often match the color pattern of the host.
Class Oligochaeta
This class includes about 3,500 species of earthworms and freshwater worms. The members of the class range in length from about 1/32 in. to 10 ft (0.5 mm-3 m), but most are comparable to the polychaetes in size. Oligochaetes occur in a variety of habitats throughout the world. Most are burrowers in the soil, but the class also includes worms that inhabit wells, marshes, and swamps. Other species live under rocks on the seashore, in the leaves of tropical trees and vines, on the surface of glaciers, or on the gills of freshwater crayfish.
Oligochaete Anatomy
Like the polychaetes, oligochaetes have bodies divided into segments. However, they lack parapodia and, with a few exceptions, have relatively few and inconspicuous setae. The setae are usually arranged in four bundles on each segment; those of aquatic forms are longer than those of land forms. The setae of an earthworm may be felt as a roughness if one rubs a finger along its side.
Oligochaetes are less varied in their external form than the polychaetes, but are much more numerous. As many as 4,000 oligochaetes have been counted in 1 square meter of lake bottom, and about 9,000 in 1 square meter of meadow soil. In almost all oligochaetes, the head is a simple cone-shaped structure without sensory appendages. Light is detected by photoreceptor cells in the skin, usually concentrated toward the front of the animal.
Oligochaete Digestion
The mouth, located under the head, leads to a relatively simple, straight digestive tract consisting of a pharynx, an esophagus, and an intestine, terminating in an anal opening. Terrestrial oligochaetes tunnel through the ground, swallowing soil as they go. The digestive tract of such a worm is specially modified for this rough diet. Typically it has a thin-walled storage area, or crop, and a muscular gizzard for grinding the soil to remove the organic matter that is the actual food of the worm. Specialized calciferous glands remove excess calcium, magnesium, strontium, and phosphate and regulate the level of these ions in the blood. Solid wastes are egested and plastered against the burrow wall, or ejected from the mouth of the burrow; the ejected material is called castings. Earthworms, through their burrowing and digestive processes, are largely responsible for the mixing and aeration of the soil. Not all oligochaetes have soil diets; some of the small aquatic worms are active predators on other small invertebrates. Excretion is typically carried out by a pair of tubes in each segment.
Oligochaete Circulation and Respiration
The circulatory system is that typical of the annelids and has many contractile vessels, or hearts. Although a few aquatic forms have gills for respiration, most oligochaetes lack such specialized structures and use the capillaries of their body walls for respiratory exchange. Oxygen dissolved in the soil water diffuses through the moist epidermis of the worm. If earthworms are forced to the surface, as when their burrows are filled with rainwater, they suffocate as a result of desiccation.
Oligochaete Reproduction
All oligochaetes are hermaphroditic, and nearly all cross-fertilize by copulation. Male and female reproductive organs are located in separate segments. The copulating pair exchange sperm, which are stored in the body of the recipient worm until its eggs are mature. The worm then secretes a cocoon into which it deposits the eggs and the sperm; fertilization and development of the eggs occur in the cocoon. When the young emerge they are miniatures of the adults. The cocoon is secreted by a glandular region, the clitellum, consisting of several thickened segments. The clitellum of an earthworm is a conspicuous saddle-shaped region near its front end.
Class Hirudinea
This class includes the 500 species of leeches, flattened, predacious or parasitic annelids equipped with suckers used for creeping. Leeches range in length from about 1/2 in. to 8 in. (1 cm-20 cm); most are under 2 in. (5 cm) long. They are commonly black, brown, green, or red, and may have stripes or spots. Leeches are primarily freshwater annelids, but some live in the ocean and some in moist soil or vegetation. The majority of leeches are predators on small invertebrates; most swallow their prey whole, but some suck the soft parts from their victims. Some leeches are parasites rather than predators, and suck the body fluids of their victims without killing them. The distinction is not sharp, as many predatory leeches take blood meals on occasion.
Leech Anatomy
Leeches are the only annelids with a fixed number (34) of body segments; each segment has secondary subdivisions known as annuli. A clitellum, less conspicuous than that of oligochaetes, is present; there are no parapodia. A leech has a small anterior sucker and a larger posterior one; the leech crawls by moving the anterior sucker forward, attaching it, and drawing up the posterior sucker. Most leeches can swim by rapid undulations of the body, using well-developed muscles of the body wall.
The coelom differs from that of other annelids in that it is largely filled in with tissue. Coelomic fluid is contained in a system of sinuses, which in some leeches functions as a circulatory system; there is a tendency in this group toward the loss of true blood vessels. The blood of some leeches is red. In others the blood lacks oxygen-carrying pigments and is therefore colorless; the oxygen dissolved directly in the blood is sufficient for respiration. Gas exchange occurs through the body surface of most leeches, although many fish-parasitizing leeches have gills.
The sense organs consist of sensory cells of various types, including photoreceptor cells, scattered over the body surface. There are also from 2 to 10 eyes, consisting of clusters of photoreceptor cells and located toward the front of the body.
Leech Predation and Digestion
Many leeches have a proboscis used for swallowing the prey or for sucking its fluids; others have jaws for biting. Many parasitic leeches are able to parasitize a wide variety of hosts. Most of the marine and some of the freshwater leeches are fish parasites. The medicinal leech, Hirudo medicinalis, is one of a group of aquatic bloodsucking leeches with jaws. Another group of jawed bloodsuckers is terrestrial; these leeches live in damp tropical vegetation and drop onto their mammalian prey. Most parasitic leeches attach to the host only while feeding; a single meal may be 5 or 10 times the weight of the leech and provide it with food for several months. The digestive tract of bloodsuckers produces an anticoagulant, hirudin, which keeps the engorged blood from clotting. A few leeches attach permanently to the host, leaving only to reproduce. Predatory leeches are active at night and hide by day.
Leech Reproduction
Like the oligochaetes, leeches are hermaphroditic and cross-fertilizing, although fertilization is internal. In some species the sperm are enclosed in sacs, called spermatophores, that are attached to the outside of the partner; the sperm pass through the body wall to the ovaries, where the eggs are fertilized. In other species the sperm are not enclosed and are transferred directly into the body of the partner by copulation. A courtship display is seen among some leeches at the time of mating. The fertilized eggs are deposited in a cocoon, secreted by the clitellum; the cocoon is buried in mud or affixed to submerged objects. The young emerge as small copies of the adults.
Bibliography
See R. O. Brinkhurst and B. G. Jamieson, Aquatic Oligochaeta of the World (1972); K. Fauchald, The Polychaete Worms (1977); R. W. Pennak, Fresh-water Invertebrates of the United States (3d ed. 1989).
| Veterinary Dictionary: annelid |
A member of the phylum annelida.
| Wikipedia: Annelid |
| Annelids Fossil range: 518–0 Ma Cambrian - Recent |
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| Glycera sp. | |
| Scientific classification | |
| Kingdom: | Animalia |
| Superphylum: | Lophotrochozoa |
| Phylum: | Annelida Lamarck, 1809 |
| Classes and subclasses | |
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Class Polychaeta (paraphyletic?) |
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The annelids, collectively called Annelida (from French annelés "ringed ones", ultimately from Latin anellus "little ring"[1]), are a large phylum of segmented worms, with over 17,000 modern species including ragworms, earthworms and leeches. They are found in marine environments from tidal zones to hydrothermal vents, in freshwater, and in moist terrestrial environments. Although most textbooks still use the traditional division into polychaetes (almost all marine), oligochaetes (which include earthworms) and leech-like species, research since 1997 has radically changed this scheme, viewing leeches as a sub-group of oligochaetes and oligochaetes as a sub-group of polychaetes. In addition, the Pogonophora, Echiura and Sipuncula, previously regarded as separate phyla, are now regarded as sub-groups of polychaetes. Annelids are considered members of the Lophotrochozoa, a "super-phylum" of protostomes that also includes molluscs, brachiopods, flatworms and nemerteans.
The basic annelid form consists of multiple segments, each of which has the same sets of organs and, in most polychaetes, a pair of parapodia that many species use for locomotion. Septa separate the segments of many species, but are poorly-defined or absent in some, and Echiura and Sipuncula show no obvious signs of segmentation. In species with well-developed septa, the blood circulates entirely within blood vessels, and the vessels in segments near the front ends of these species are often built up with muscles to act as hearts. The septa of these species also enable them to change the shapes of individual segments, which facilitates movement by peristalsis ("ripples" that pass along the body) or by undulations that improve the effectiveness of the parapodia. In species with incomplete septa or none, the blood circulates through the main body cavity without any kind of pump, and there is a wide range of locomotory techniques – some burrowing species turn their pharynges inside out to drag themselves through the sediment.
Although many species can reproduce asexually and use similar mechanisms to regenerate after severe injuries, sexual reproduction is the normal method in species whose reproduction has been studied. The minority of living polychaetes whose reproduction and lifecycles are known produce trochophore larvae, which live as plankton and then sink and metamorphose into miniature adults. Oligochaetes are full hermaphrodites and produce a ring-like cocoon round their bodies, in which the eggs and hatchlings are nourished until they are ready to emerge.
Earthworms support terrestrial food chains both as prey and by aerating and enriching soil. The burrowing of marine polychaetes, which may constitute up to a third of all species in near-shore environments, encourages the development of ecosystems by enabling water and oxygen to penetrate the sea floor. In addition to improving soil fertility, annelids serve humans as food and as bait. Scientists observe annelids to monitor the quality of marine and fresh water. Although blood-letting is no longer in favor with doctors, some leech species are regarded as endangered species because they have been over-harvested for this purpose in the last few centuries. Ragworms' jaws are now being studied by engineers as they offer an exceptional combination of lightness and strength.
Since annelids are soft-bodied, their fossils are rare – mostly jaws and the mineralized tubes that some of the species secreted. Although some late Ediacaran fossils may represent annelids, the oldest known fossil that is identified with confidence comes from about in the early Cambrian period. Fossils of most modern mobile polychaete groups appeared by the end of the Carboniferous, about . Scientists disagree about whether some body fossils from the mid Ordovician, about , are the remains of oligochaetes, and the earliest certain fossils of the group appear in the Tertiary period, which began .
Contents |
There are over 17,000 living annelid species,[2] ranging in size from microscopic to the Australian giant Gippsland earthworm, which can grow up to 3 metres (9.8 ft) long.[3][4] Although research since 1997 has radically changed scientists' views about the evolutionary family tree of the annelids,[5][6] most textbooks use the traditional classification into the following sub-groups:[3][7]
The Archiannelida, minute annelids that live in the spaces between grains of sediment, were treated as a separate class because of their simple body structure, but are now regarded as polychaetes.[7] Some other groups of animals have been classified in various ways, but are now widely regarded as annelids:
No single feature distinguishes Annelids from other invertebrate phyla, but they have a distinctive combination of features. Their bodies are long, with segments that are divided externally by shallow ring-like constrictions called annuli and internally by septa ("partitions") at the same points, although in some species the septa are incomplete and in a few cases missing. Most of the segments contain the same sets of organs, although sharing a common gut, circulatory system and nervous system makes them inter-dependent.[3][7] Their bodies are covered by a cuticle (outer covering) that does not contain cells but is secreted by cells in the skin underneath, is made of tough but flexible collagen[3] and does not molt[13] – on the other hand arthropods' cuticles are made of the more rigid α-chitin,[3][14] and molt until the arthropods reach their full size.[15] Most annelids have closed circulatory systems, where the blood makes its entire circuit via blood vessels.[13]
| Annelida[3] | Recently merged into Annelida[5] | Closely-related | Similar-looking phyla | |||
|---|---|---|---|---|---|---|
| Echiura[16] | Sipuncula[17] | Nemertea[18] | Arthropoda[19] | Onychophora[20] | ||
| External segmentation | Yes | no | no | Only in a few species | Yes, except in mites | no |
| Repetition of internal organs | Yes | no | no | Yes | In primitive forms | Yes |
| Septa between segments | In most species | no | no | No | No | No |
| Cuticle material | collagen | collagen | collagen | none | α-chitin | α-chitin |
| Molting | Generally no;[13] but some polychaetes molt their jaws, and leeches molt their skins[21] | no[22] | no[22] | no[22] | Yes[15] | Yes |
| Body cavity | Coelom; but this is reduced or missing in many leeches and some small polychaetes[13] | 2 coeloms, main and in proboscis | 2 coeloms, main and in tentacles | Coelom only in proboscis | Hemocoel | Hemocoel |
| Circulatory system | Closed in most species | Open outflow, return via branched vein | Open | Closed | Open | Open |
Most of an annelid's body consists of segments that are practically identical, having the same sets of internal organs and external chetae (Greek χαιτα, meaning "hair") and, in some species, appendages. However, the frontmost and rearmost sections are not regarded as true segments as they do not contain the standard sets of organs and do not develop in the same way as the true segments. The frontmost section, called the prostomium (Greek προ- meaning "in front of" and στομα meaning "mouth") contains the brain and sense organs, while the rearmost, called the pygidium (Greek πυγιδιον, meaning "little tail") contains the anus, generally on the underside. The first section behind the prostomium, called the peristomium (Greek περι- meaning "around" and στομα meaning "mouth"), is regarded by some zoologists as not a true segment, but in some polychaetes the peristomium has chetae and appendages like those of other segments.[3]
The segments develop one at a time from a growth zone just ahead of the pygidium, so that an annelid's youngest segment is just in front of the growth zone while the peristomium is the oldest. This pattern is called teloblastic growth.[3] Some groups of annelids, including all leeches,[10] have fixed maximum numbers of segments, while others add segments throughout their lives.[7]
The phylum's name is derived from the Latin word annelus, meaning "little ring".[2]
Annelids' cuticles are made of collagen fibers, usually in layers that spiral in alternating directions so that the fibers cross each other. These are secreted by the one-cell deep epidermis (outermost skin layer). A few marine annelids that live in tubes lack cuticles, but their tubes have a similar structure, and mucus-secreting glands in the epidermis protect their skins.[3] Under the epidermis is the dermis, which is made of connective tissue, in other words a combination of cells and non-cellular materials such as collagen. Below this are two layers of muscles, which develop from the lining of the coelom (body cavity): circular muscles make a segment longer and slimmer when they contract, while under them are longitudinal muscles, usually four distinct strips,[13] whose contractions make the segment shorter and fatter.[3] Some annelids also have oblique internal muscles that connect the underside of the body to each side.[13]
The chetae ("hairs") of annelids project out from the epidermis to provide traction and other capabilities. The simplest are unjointed and form paired bundles near the top and bottom of each side of each segment. The parapodia ("limbs") of annelids that have them often bear more complex chetae at their tips – for example jointed, comb-like or hooked.[3] Chetae are made of moderately flexible β-chitin and are formed by follicles, each of which has a chaetoblast ("hair-forming") cell at the bottom and muscles that can extend or retract the cheta. The chetoblasts produce chetae by forming microvilli, fine hair-like extensions that increase the area available for secreting the cheta. When the cheta is complete, the microvilli withdraw into the chetoblast, leaving parallel tunnels that run almost the full length of the cheta.[3] Hence annelids' chetae are structurally different from the setae ("bristles") of arthropods, which are made of the more rigid α-chitin, have a single internal cavity, and are mounted on flexible joints in shallow pits in the cuticle.[3]
Nearly all polychaetes have parapodia that function as limbs, while other major annelid groups lack them. Parapodia are unjointed paired extensions of the body wall, and their muscles are derived from the circular muscles of the body. They are often supported internally by one or more large, thick chetae. The parapodia of burrowing and tube-dwelling polychaetes are often just ridges whose tips bear hooked chetae. In active crawlers and swimmers the parapodia are often divided into large upper and lower paddles on a very short trunk, and the paddles are generally fringed with chetae and sometimes with cirri (fused bundles of cilia) and gills.[13]
The brain generally forms a ring round the pharynx (throat), consisting of a pair of ganglia (local control centers) above and in front of the pharynx, linked by nerve cords either side of the pharynx to another pair of ganglia just below and behind it.[3] The brains of polychaetes are generally in the prostomium, while those of clitellates are in the peristomium or sometimes the first segment behind the peristomium.[23] In some very mobile and active polychaetes the brain is enlarged and more complex, with visible hindbrain, midbrain and forebrain sections.[13] The rest of the central nervous system is generally "ladder-like", consisting of a pair of nerve cords that run through the bottom part of the body and have in each segment paired ganglia linked by a transverse connection. From each segmental ganglion a branching system of local nerves runs into the body wall and then encircles the body.[3] However, in most polychaetes the two main nerve cords are fused, and in the tube-dwelling genus Owenia the single nerve chord has no ganglia and is located in the epidermis.[7][24]
As in arthropods, each muscle fiber (cell) is controlled by more than one neuron, and the speed and power of the fiber's contractions depends on the combined effects of all its neurons. Vertebrates have a different system, in which one neuron controls a group of muscle fibers.[3] Most annelids' longitudinal nerve trunks include giant axons (the output signal lines of nerve cells). Their large diameter decreases their resistance, which allows them to transmit signals exceptionally fast. This enables these worms to withdraw rapidly from danger by shortening their bodies. Experiments have shown that cutting the giant axons prevents this escape response but does not affect normal movement.[3]
The sensors are primarily single cells that detect light, chemicals, pressure waves and contact, and are present on the head, appendages (if any) and other parts of the body.[3] Nuchal ("on the neck") organs are paired, ciliated structures found only in polychaetes, and are thought to be chemosensors.[13] Some polychaetes also have various combinations of ocelli ("little eyes") that detect the direction from which light is coming and camera eyes or compound eyes that can probably form images.[24] The compound eyes probably evolved independently of arthropods' eyes.[13] Some tube-worms use ocelli widely spread over their bodies to detect the shadows of fish, so that they can quickly withraw into their tubes.[24] Some burrowing and tube-dwelling polychaetes have statocysts (tilt and balance sensors) that tell them which way is down.[24] A few polychaete genera have on the undersides of their heads palps that are used both in feeding and as "feelers", and some of these also have antennae that are structurally similar but probably are used mainly as "feelers".[13]
Most annelids have a pair of coeloms (body cavities) in each segment, separated from other segments by septa and from each other by vertical mesenteries. Each septum forms a sandwich with connective tissue in the middle and mesothelium (membrane that serves as a lining) from the preceding and following segments on either side. Each mesentery is similar except that the mesothelium is the lining of each of the pair of coeloms, and the blood vessels and, in polychaetes, the main nerve cords are embedded in it.[3] The mesothelium is made of modified epitheliomuscular cells,[3] in other words their bodies form part of the epithelium but their bases extend to form muscle fibers in the body wall.[25] The mesothelium may also form radial and circular muscles on the septa, and circular muscles around the blood vessels and gut. Parts of the mesothelium, especially on the outside of the gut, may also form chloragogen cells that perform similar functions to the livers of vertebrates: producing and storing glycogen and fat; producing the oxygen-carrier hemoglobin; breaking down proteins; and turning nitrogenous waste products into ammonia and urea to be excreted.[3]
Many annelids move by peristalsis (waves of contraction and expansion that sweep along the body),[3] or flex the body while using parapodia to crawl or swim.[26] In these animals the septa enable the circular and longitudinal muscles to change the shape of individual segments, by making each segment a separate fluid-filled "balloon".[3] However, the septa are often incomplete in annelids that are semi-sessile or that do not move by peristalsis or by movements of parapodia – for example some move by whipping movements of the body, some small marine species move by means of cilia (fine muscle-powered hairs) and some burrowers turn their pharynges (throats) inside out to penetrate the sea-floor and drag themselves into it.[3]
The fluid in the coeloms contains coelomocyte cells that defend the animals against parasites and infections. In some species coelomocytes may also contain a respiratory pigment – red hemoglobin in some species, green chlorocruorin in others[13] – and provide oxygen transport within their segments. Respiratory pigment is also dissolved in the blood plasma. Species with well-developed septa generally also have blood vessels running all long their bodies above and below the gut, the upper one carrying blood forwards while the lower one carries it backwards. Networks of capillaries (fine blood vessels) in the body wall and around the gut transfer blood between the main blood vessels and to parts of the segment that need oxygen and nutrients. Both of the major vessels, especially the upper one, can pump blood by contracting. In some annelids the forward end of the upper blood vessel is enlarged with muscles to form a heart, while in the forward ends of many earthworms some of the vessels that connect the upper and lower main vessels function as hearts. Species with poorly-developed or no septa generally have no blood vessels and rely on the circulation within the coelom for delivering nutrients and oxygen.[3]
However, leeches and their closest relatives have a body structure that is very uniform within the group but significantly different from that of other annelids, including other members of the Clitellata.[10] In leeches there are no septa, the connective tissue layer of the body wall is so thick that it occupies much of the body, and the two coeloms are widely separated and run the length of the body. They function as the main blood vessels, although they are side-by-side rather than upper and lower. However, they are lined with mesothelium, like the coeloms and unlike the blood vessels of other annelids. Leeches generally use suckers at their front and rear ends to move like inchworms. The anus is on the upper surface of the pygidium.[10]
In some annelids, including earthworms, all respiration is via the skin. However, many polychaetes and some clitellates (the group to which earthworms belong) have gills associated with most segments, often as extensions of the parapodia in polychaetes. The gills of tube-dwellers and burrowers usually cluster around whichever end has the stronger water flow.[13]
Feeding structures in the mouth region vary widely, and have little correlation with the animals' diets. Many polychaetes have a muscular pharynx that can be everted (turned inside out to extend it). In these animals the foremost few segments often lack septa so that, when the muscles in these segments contract, the sharp increase in fluid pressure from all these segments everts the pharynx very quickly. Two families, the Eunicidae and Phyllodocidae, have evolved jaws, which can be used for seizing prey, biting off pieces of vegetation, or grasping dead and decaying matter. On the other hand some predatory polychaetes have neither jaws nor eversible pharynges. Selective deposit feeders generally live in tubes on the sea-floor and use palps to find food particles in the sediment and them wipe them into their mouths. Filter feeders use "crowns" of palps covered in cilia that wash food particles towards their mouths. Non-selective deposit feeders ingest soil or marine sediments via mouths that are generally unspecialized. Some clitellates have sticky pads in the roofs of their mouths, and some of these can evert the pads to capture prey. Leeches often have an eversible proboscis, or a muscular pharynx with two or three teeth.[13]
The gut is generally an almost straight tube supported by the mesenteries (vertical partitions within segments), and ends with the anus on the underside of the pygidium.[3] However, in members of the tube-dwelling family Siboglinidae the gut is blocked by a swollen lining that houses symbiotic bacteria, which can make up 15% of the worms' total weight. The bacteria convert inorganic matter – such as hydrogen sulfide and carbon dioxide from hydrothermal vents, or methane from seeps – to organic matter that feeds themselves and their hosts, while the worms extend their palps into the gas flows to absorb the gases needed by the bacteria.[13]
Annelids with blood vessels use metanephridia to remove soluble waste products, while those without use protonephridia.[3] Both of these systems use a two-stage filtration process, in which fluid and waste products are first extracted and these are filtered again to re-absorb any re-usable materials while dumping toxic and spent materials as urine. The difference is that protonephridia combine both filtration stages in the same organ, while metanephridia perform only the second filtration and rely on other mechanisms for the first - in annelids special filter cells in the walls of the blood vessels let fluids and other small molecules pass into the coelomic fluid, where it circulates to the metanephridia.[27] In annelids the points at which fluid enters the protonephridia or metanephridia are on the forward side of a septum while the second-stage filter and the nephridiopore (exit opening in the body wall) are in the following segment. As a result the hindmost segment (before the growth zone and pygidium) has no structure that extracts its wastes, as there is no following segment to filter and discharge them, while the first segment contains an extraction structure that passes wastes to the second, but does not contain the structures that re-filter and discharge urine.[3]
Polychaetes can reproduce asexually, by dividing into two or more pieces or by budding off a new individual while the parent remains a complete organism.[3][28] Some oligochaetes, such as Aulophorus furcatus, seem to reproduce entirely asexually, while others reproduce asexually in summer and sexually in autumn. Asexual reproduction in oligochaetes is always by dividing into two or more pieces, rather than by budding.[7][29] However, leeches have never been seen reproducing asexually.[7][30]
Most polychaetes and oligochaetes also use similar mechanisms to regenerate after suffering damage. Two polychaete genera, Chaetopterus and Dodecaceria, can regenerate from a single segment, and others can regenerate even if their heads are removed.[7][28] Annelids are the most complex animals that can regenerate after such severe damage.[31] On the other hand leeches cannot regenerate.[30]
It is thought that annelids were originally animals with two separate sexes, which released ova and sperm into the water via their nephridia.[3] The fertilized eggs develop into trochophore larvae, which live as plankton.[33] Later they sink to the sea-floor and metamorphose into miniature adults: the part of the trochophore between the apical tuft and the prototroch becomes the prostomium (head); a small area round the trochophore's anus becomes the pygidium (tail-piece); a narrow band immediately in front of that becomes the growth zone that produces new segments; and the rest of the trochophore becomes the peristomium (the segment that contains the mouth).[3]
However, the lifecycles of most living polychaetes, which are almost all marine animals, are unknown, and only about 25% of the 300+ species whose lifecycles are known follow this pattern. About 14% use a similar external fertilization but produce yolk-rich eggs, which reduce the time the larva needs to spend among the plankton, or eggs from which miniature adults emerge rather than larvae. The rest care for the fertilized eggs until they hatch – some by producing jelly-covered masses of eggs which they tend, some by attaching the eggs to their bodies and a few species by keeping the eggs within their bodies until they hatch. These species use a variety of methods for sperm transfer; for example, in some the females collect sperm released into the water, while in others the males have penes that inject sperm into the female.[33] There is no guarantee that this is a representative sample of polychaetes' reproductive patterns, and it simply reflects scientists' current knowledge.[33]
Some polychaetes breed only once in their lives, while others breed almost continuously or through several breeding seasons. While most polychaetes remain of one sex all their lives, a significant percentage of species are full hermaphrodites or change sex during their lives. Most polychaetes whose reproduction has been studied lack permanent gonads, and it is uncertain how they produce ova and sperm. In a few species the rear of the body splits off and becomes a separate individual that lives just long enough to swim to a suitable environment, usually near the surface, and spawn.[33]
Most mature clitellates (the group that includes earthworms and leeches) are full hermaphrodites, although in a few leech species younger adults function as males and become female at maturity. All have well-developed gonads, and all copulate. Earthworms store their partners' sperm in spermathecae ("sperm stores") and then the clitellum produces a cocoon that collects ova from the ovaries and then sperm from the spermathecae. Fertilization and development of earthworm eggs takes place in the cocoon. Leeches' eggs are fertilized in the ovaries, and then transferred to the cocoon. In all clitellates the cocoon also either produces yolk when the eggs are fertilized or nutrients while they are developing. All clitellates hatch as miniature adults rather than larvae.[33]
Charles Darwin's book The Formation of Vegetable Mould through the Action of Worms (1881) presented the first scientific analysis of earthworms' contributions to soil fertility.[34] Some burrow while others live entirely on the surface, generally in moist leaf litter. The burrowers loosen the soil so that oxygen and water can penetrate it, and both surface and burrowing worms help to produce soil by mixing organic and mineral matter, by accelerating the decomposition of organic matter and thus making it more quickly available to other organisms, and by concentrating minerals and converting them to forms that plants can use more easily.[35][36] Earthworms are also important prey for birds ranging in size from robins to storks, and for mammals ranging from shrews to badgers, and in some cases conserving earthworms may be essential for conserving endangered birds.[37]
Marine annelids may account for over one-third of bottom-dwelling animal species round coral reefs and in tidal zones.[34] Burrowing species increase the penetration of water and oxygen and water into the sea-floor sediment, which encourages the growth of populations of bacteria and small animals alongside their burrows.[38]
Although blood-sucking leeches do little direct harm to their victims, some transmit flagellates that can be very dangerous to their hosts. Some small tube-dwelling oligochaetes transmit myxosporean parasites that cause whirling disease in fish.[34]
Earthworms make a significant contribution to soil fertility.[34] The rear end of the Palolo worm, a marine polychaete that tunnels through coral, detaches in order to spawn at the surface, and the people of Samoa regard these spawning modules as a delicacy.[34] Anglers sometimes find that worms are more effective bait than artificial flies, and worms can be kept for several days in a tin lined with damp moss.[39] Ragworms are commercially important as bait and as food sources for aquaculture, and there have been proposals to farm them in order to reduce over-fishing of their natural populations.[38] Some marine polychaetes' predation on molluscs causes serious losses to fishery and aquaculture operations.[34]
Scientists study aquatic annelids to monitor the oxygen content, salinity and pollution levels in fresh and marine water.[34]
Accounts of the use of leeches for the medically dubious practise of blood-letting have come from China around 30 AD, India around 200 AD, ancient Rome around 50 AD and later throughout Europe. In the 19th century medical demand for leeches was so high that some areas' stocks were exhausted and other regions imposed restrictions or bans on exports, and Hirudo medicinalis is treated as an endangered species by both IUCN and CITES. More recently leeches have been used to assist in microsurgery, and their saliva has provided anti-inflammatory compounds and several important anticoagulants, one of which also prevents tumors from spreading.[34]
Ragworms' jaws are strong but much lighter than the hard parts of many other organisms, which are biomineralized with calcium salts. These advantages have attracted the attention of engineers. Investigations showed that ragworm jaws are made of unusual proteins that bind strongly to zinc.[40]
Since annelids are soft-bodied, their fossils are rare.[41] Polychaetes' fossil record consists mainly of the jaws that some species had and the mineralized tubes that some secreted.[42] Some Ediacaran fossils such as Dickinsonia in some ways resemble polychaetes, but the similarities are too vague for these fossils to be classified with confidence.[43] The small shelly fossil Cloudina, from , has been classified by some authors as an annelid, but by others as a cnidarian (i.e. in the phylum to which jellyfish and sea anemones belong).[44] Until 2008 the earliest fossils widely accepted as annelids were the polychaetes Canadia and Burgessochaeta, both from Canada's Burgess Shale, formed about in the early Cambrian.[45] Myoscolex, found in Australia and a little older than the Burgess Shale, was possibly an annelid. However, it lacks some typical annelid features and has features which are not usually found in annelids and some of which are associated with other phyla.[45] Then Simon Conway Morris and John Peel reported Phragmochaeta from Sirius Passet, about , and concluded that it was the oldest annelid known to date.[43] There has been vigorous debate about whether the Burgess Shale fossil Wiwaxia was a mollusc or an annelid.[45] Polychaetes diversified in the early Ordovician, about . By the end of the Carboniferous, about , fossils of most of the modern mobile polychaete groups had appeared.[45] Many fossil tubes look like those made by modern sessile polychaetes, but the first tubes clearly produced by polychaetes date from the Jurassic, less than .[45]
The earliest good evidence for oligochaetes occurs in the Tertiary period, which began , and it has been suggested that these animals evolved around the same time as flowering plants in the early Cretaceous, from .[46] A trace fossil consisting of a convoluted burrow partly filled with small fecal pellets may be evidence that earthworms were present in the early Triassic period from .[46][47] Body fossils going back to the mid Ordovician, from , have been tentatively classified as oligochaetes, but these identifications are uncertain and some have been disputed.[46][48]
| Annelida |
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Traditionally the annelids have been divided into two major groups, the polychaetes and clitellates. In turn the clitellates were divided into oligochaetes, which include earthworms, and hirudinomorphs, whose best-known members are leeches.[3] For many years there was no clear arrangement of the approximately 80 polychaete families into higher-level groups.[5] In 1997 Greg Rouse and Kristian Fauchald attempted a "first heuristic step in terms of bringing polychaete systematics to an acceptable level of rigour", based on anatomical structures, and divided polychaetes into:[50]
Also in 1997 Damhnait McHugh, using molecular phylogenetics to compare similarities and differences in one gene, presented a very different view, in which: the clitellates were an off-shoot of one branch of the polychaete family tree; the pogonophorans and echiurans, which for a few decades had been regarded as a separate phyla, were placed on other branches of the polychaete tree.[52] Subsequent molecular phylogenetics analyses on a similar scale presented similar conclusions.[53]
In 2007 Torsten Struck and colleagues compared 3 genes in 81 taxa, of which 9 were outgroups,[5] in other words not considered closely related to annelids but included to give an indication of where the organisms under study are placed on the larger tree of life.[54] For a cross-check the study used an analysis of 11 genes (including the original 3) in 10 taxa. This analysis agreed that clitellates, pogonophorans and echiurans were on various branches of the polychaete family tree. It also concluded that the classification of polychaetes into Scolecida, Canalipalpata and Aciculata was useless, as the members of these alleged groups were scattered all over the family tree derived from comparing the 81 taxa. In addition, it also placed sipunculans, generally regarded at the time as a separate phylum, on another branch of the polychaete tree, and concluded that leeches were a sub-group of oligochaetes rather than their sister-group among the clitellates.[5] Rouse accepted the analyses based on molecular phylogenetics,[7] and their main conclusions are now the scientific consensus, although the details of the annelid family tree remain uncertain.[6]
In addition to re-writing the classification of annelids and 3 previously independent phyla, the molecular phylogenetics analyses undermine the emphasis that decades of previous writings placed on the importance of segmentation in the classification of invertebrates. Polychaetes, which these analyses found to be the parent group, have completely segmented bodies, while polychaetes' echiurans and sipunculan offshoots are not segmented and pogonophores are segmented only in the rear parts of their bodies. It now seems that segmentation can appear and disappear much more easily in the course of evolution than was previously thought.[5][52] The 2007 study also noted that the ladder-like nervous system, which is associated with segmentation, is less universal than previously thought in both annelids and arthropods.[5]
| Bilateria |
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Annelids are members of the protostomes, one of the two major superphyla of bilaterian animals - the other is the deuterostomes, which includes vertebrates.[53] Within the protostomes, annelids used to be grouped with arthropods under the super-group Articulata ("jointed animals"), as segmentation is obvious in most members of both phyla. However, the genes that drive segmentation in arthropods do not appear to do the same in annelids. Arthropods and annelids both have close relatives that are unsegmented. It is at least as easy to assume that they evolved segmented bodies independently as it is to assume that the ancestral protostome or bilaterian was segmented and that segmentation disappeared in many descendant phyla.[53] The current view is that annelids are grouped with molluscs, brachiopods and several other phyla that have lophophores (fan-like feeding structures) and/or trochophore larvae as members of Lophotrochozoa.[55] Bryzoa may be the most basal phylum (the one that first became distinctive) within the Lophotrochozoa, and the relationships between the other members are not yet known.[53] Arthropods are now regarded as members of the Ecdysozoa ("animals that molt"), along with some phyla that are unsegmented.[53][56]
The "Lophotrochozoa" hypothesis is also supported by the fact that many phyla within this group, including annelids, molluscs, nemerteans and flatworms, follow a similar pattern in the fertilized egg's development. When their cells divide after the 4-cell stage, descendants of these 4 cells form a spiral pattern. In these phyla the "fates" of the embryo's cells, in other words the roles their descendants will play in the adult animal, are the same and can be predicted from a very early stage.[57] Hence this development pattern is often described as "spiral determinate cleavage".[58]
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| Translations: Annelid |
Dansk (Danish)
n. - ledorm
adj. - ledorms-
Français (French)
n. - annélide, (Zool) annélide
adj. - annélidien
Deutsch (German)
n. - Ringelwurm
adj. - zu den Ringelwürmern gehörig
Ελληνική (Greek)
n. - (ζωολ.) αννελίδης, ζωνοσκώληκας
Português (Portuguese)
n. - anelídeo (m) (Zool.)
Русский (Russian)
кольчатый червь
Español (Spanish)
n. - anélido
adj. - relacionado con los anélidos
Svenska (Swedish)
n. - ringmask
中文（简体）(Chinese (Simplified))
环节动物, 环节动物的
中文（繁體）(Chinese (Traditional))
n. - 環節動物
adj. - 環節動物的
한국어 (Korean)
n. - 환형동물
adj. - 환형동물의
日本語 (Japanese)
n. - 環形動物
adj. - 環形動物の
العربيه (Arabic)
(الاسم) الدوده الحلقيه, دوده يتكون جسمها من حلقات متتاليه
עברית (Hebrew)
n. - תולעת טבעות
adj. - של תולעת טבעות
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| Dinophilidae (invertebrate zoology) |
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Copyrights:
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 | Dictionary. The American Heritage® Dictionary of the English Language, Fourth Edition Copyright © 2007, 2000 by Houghton Mifflin Company. Updated in 2009. Published by Houghton Mifflin Company. All rights reserved. Read more |
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| Britannica Concise Encyclopedia. Britannica Concise Encyclopedia. © 2006 Encyclopædia Britannica, Inc. All rights reserved. Read more |
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| Columbia Encyclopedia. The Columbia Electronic Encyclopedia, Sixth Edition Copyright © 2003, Columbia University Press. Licensed from Columbia University Press. All rights reserved. www.cc.columbia.edu/cu/cup/. Read more |
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| Veterinary Dictionary. Saunders Comprehensive Veterinary Dictionary 3rd Edition. Copyright © 2007 by D.C. Blood, V.P. Studdert and C.C. Gay, Elsevier. All rights reserved. Read more |
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| Wikipedia. This article is licensed under the Creative Commons Attribution/Share-Alike License. It uses material from the Wikipedia article "Annelid". Read more |
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