Annelid

? Annelids
230px Glycera sp.
Scientific classification
Kingdom: Animalia Phylum: Annelida Lamarck, 1809
Classes and subclasses
Class Polychaeta Class Clitellata    Oligochaeta - Earthworms and others    Acanthobdellida    Branchiobdellida    Hirudinea - Leeches Class Myzostomida Class Archiannelida *Some authors consider the subclasses under Clitellata to be classes

The annelids (from Latin anellus "little ring") are a large phylum (Annelida) of invertebrate animals, comprising the segmented worms, including the well-known earthworms and leeches. There are about 15,000 known modern species of annelids.

Annelids are found in most wet environments and include many terrestrial, freshwater, and especially marine species (such as the polychaetes), as well as some which are parasitic or mutualistic. They range in length from under a millimeter to over 3 meters (the seep tube worm Lamellibrachia luymesi).

Anatomy

Annelids are triploblastic protostomes with a coelom (at least historically), closed circulatory system, and true segmentation. Protosomes are animals with bilaterial symmetry where the first opening in development, the blastophore, becomes its mouth. Triploblastic means that they have three primary tissue areas formed during embryogenesis. A coelom is a fluid-filled body cavity.

Oligochaetes and polychaetes typically have spacious coeloms; in leeches, the coelom is largely filled in with tissue and reduced to a system of narrow canals; archiannelids may lack the coelom entirely. The coleom is divided into a sequence of compartments by walls called septa. In the most general forms, each compartment corresponds to a single segment of the body, which also includes a portion of the nervous and (closed) circulatory systems, allowing it to function relatively independently. Each segment is marked externally by one or more rings, called annuli. Each segment also has an outer layer of circular muscle underneath a thin cuticle and epidermis, and a system of longitudinal muscles. In earthworms, the longitudinal muscles are strengthened by collagenous lamellae; the leeches have a double layer of muscles between the outer circulars and inner longitudinals. In most forms, they also carry a varying number of bristles, called setae, and among the polychaetes a pair of appendages, called parapodia.

Polychaeta: "A variety of marine worms" plate from Das Meer by M. J. Schleiden (1804–1881)

Anterior to the true segments lies the prostomium and peristomium, which carries the mouth, and posterior to them lies the pygidium, where the anus is located. The digestive tract is quite variable but is usually specialized. For example, in some groups (notably most earthworms) it has a typhlosole (internal fold of the intestine or intestine inner wall), to increase surface area, along much of its length.

Different species of annelids have a wide variety of diets, including active and passive hunters, scavengers, filter feeders, direct deposit feeders that simply ingest the sediments, and blood-suckers.

The vascular system and the nervous system are separate from the digestive tract. The vascular system includes a dorsal vessel conveying the blood toward the front of the worm, and a ventral longitudinal vessel that conveys the blood in the opposite direction. The two systems are connected by a vascular sinus and by lateral vessels of various kinds, including in the true earthworms, capillaries on the body wall.

The nervous system has a solid, ventral nerve cord from which lateral nerves arise in each segment. Every segment has an autonomy; however, they unite to perform as a single body for functions such as locomotion. Growth in many groups occurs by replication of individual segmental units; in others, the number of segments is fixed in early development.

Reproduction

Depending upon the species, annelids can reproduce both sexually and asexually.

Asexual reproduction

Asexual reproduction by fission is a method used by some annelids and allows them to reproduce quickly. The posterior part of the body breaks off and forms a new individual. The position of the break is usually determined by an epidermal growth. Lumbriculus and Aulophorus, for example, are known to reproduce by the body breaking into such fragments. Many other taxa (such as most earthworms) cannot reproduce this way, though they have varying abilities to regrow amputated segments.

Sexual reproduction

Sexual reproduction allows a species to better adapt to its environment. Some annelida species are hermaphroditic, while others have distinct sexes.

Most polychaete worms have separate males and females and external fertilization. The earliest larval stage, which is lost in some groups, is a ciliated trochophore, similar to those found in other phyla. The animal then begins to develop its segments, one after another, until it reaches its adult size.

Earthworms and other oligochaetes, as well as the leeches, are hermaphroditic and mate periodically throughout the year in favored environmental conditions. They mate by copulation. Two worms, which are attracted by each other's secretions, lay their bodies together with their heads pointing in opposite directions. The fluid is transferred from the male pore to the other worm. Different methods of sperm transference have been observed in different genera, and may involve internal spermathecae (sperm storing chambers) or spermatophores that are attached to the outside of the other worm's body. The clitellata lack the free-living ciliated trochophore larvae present in the polychaetes, the embryonic worms developing in a fluid-filled "cocoon" secreted by the clitellum.

Fossil record

The annelid fossil record is sparse, but a few definite forms are known as early as the Cambrian. There are some signs they may have been around in the later Precambrian. Because the creatures have soft bodies, fossilization is an especially rare event. The best-preserved and oldest annelid fossils come from Cambrian Lagerstätten, such as the Burgess Shale of Canada, and the Middle Cambrian strata of the House Range in Utah. The annelids are also diversely represented in the Pennsylvanian-age Mazon Creek fauna of Illinois.

Relationships

The arthropods and their kin have long been considered the closest relatives of the annelids on account of their common segmented structure. However, a number of differences between the two groups suggest this may be convergent evolution rather than a feature passed on by common descent.

The other major phylum that is of definite relation to the annelids are the mollusks, which share with them the presence of trochophore larvae. Annelids and Mollusks are thus united as the Trochozoa, a taxon more strongly supported by molecular evidence.

References

ISBN links support NWE through referral fees

• Annelid Fossils (Web page). The Virtual Fossil Museum (2006). Retrieved May 20, 2006. – Descriptions and images of annelid fossils from Mazon Creek and the Utah House Range.

References

Annelid Fossils.[[1]] (web page) The Virtual Fossil Museum: (2006). Retrieved on May 20, 2006. – Descriptions and images of annelid fossils from Mazon Creek and the Utah House Range.

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