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Worms, Animals

Worms, animals

A worm is an elongated soft-bodied invertebrate animal. The most famous is the earthworm, a member of phylum Annelida, but there are hundreds of thousands of different species that live in a wide variety of habitats other than soil. Originally, the word referred to any creeping or a crawling animal of any kind or size, such as a serpent, caterpillar, snail, or the like. Later this definition was narrowed to the modern definition which still includes several different animal groups. Major phyla include:
- Acanthocephala (spiny-headed worms)
- Annelida (segmented worms)
- Chaetognatha (arrow worms)
- Gnathostomulida (jaw worms)
- Nematoda (roundworms)
- Nematomorpha (horsehair worms)
- Nemertea (ribbonworms)
- Onychophora (velvet worms)
- Platyhelminthes (flatworms)
- Sipuncula (peanut worms) Worms may also be called helminths, especially in medical or zoological terminology. Some other invertebrate groups may be called worms, especially colloquially. Many insect larvae are called worms, such as the railroad worm, woodworm, glowworm, or bloodworms. When an animal, such as a dog, is said to have worms, it means that the dog is infested with parasitic worms, typically roundworm or tapeworm. Worm species differ in their abilities to move about on their own. Many species have bodies with no major muscles, and cannot move on their own. They must be moved by forces or other animals in their environment. Many species have bodies with major muscles, that let them move on their own. They are a type of muscular hydrostat. category:animals

Invertebrate

Invertebrate is a term coined by Jean-Baptiste Lamarck to describe any animal without a spinal column. It therefore includes all animals except vertebrates (fish, reptiles, amphibians, birds and mammals). Lamarck divided these animals into two groups, the Insecta and the Vermes, but nowadays, they are classified into over 30 phyla, from simple organisms such as sponges and flatworms to complex animals such as arthropods and mollusks. Since invertebrates include all animals except a certain group, invertebrates form a paraphyletic group, but, despite not forming a "natural group" (that is, monophyletic), "invertebrate" is still a widely used term. It is not uncommon for books entitled "Invertebrate Zoology" to be found. This reflects the bias in society and also in zoology towards larger, more complex animals that are more closely related to humans. Thus, there are relatively many scientists studying (and relatively much funding available for the study of) birds, mammals, reptiles, and so on, but far fewer scientists studying invertebrates, even though invertebrates include 97% of all animal species. For a full list of animals considered to be invertebrates, see animal. All the listed phyla are invertebrates along with two of the three subphyla in Phylum Chordata: Urochordata and Cephalochordata. These two, plus all the other known invertebrates, have only one cluster of Hox genes, while the vertebrates have duplicated their original cluster more than once.

External links

- [http://reference.allrefer.com/encyclopedia/categories/invertz.html Invertebrate Zoology]
- [http://digitalcommons.unl.edu/onlinedictinvertzoology/ Online Dictionary of Invertebrate Zoology]
- [http://www.goliathus.cz/en/museum-homepage-0.html Online museum] of many invertebrates, provided by [http://www.goliathus.cz/ goliathus.cz]. Category:Animals ms:Invertebrata ja:無脊椎動物

Animal

:For the Muppet Show character, see Animal (Muppet). For the professional wrestler, see Joseph Laurinaitis.

- Porifera (sponges)
- Ctenophora (comb jellies)
- Cnidaria (coral, jellyfish, anenomes)
- Placozoa (trichoplax)
- Subregnum Bilateria (bilateral symmetry)
- Acoelomorpha (basal)
- Orthonectida (flatworms, echinoderms, etc.)
- Rhombozoa (dicyemids)
- Myxozoa (slime animals)
- Superphylum Deuterostomia (blastopore becomes anus)
- Chordata (vertebrates, etc.)
- Hemichordata (acorn worms)
- Echinodermata (starfish, urchins)
- Chaetognatha (arrow worms)
- Superphylum Ecdysozoa (shed exoskeleton)
- Kinorhyncha (mud dragons)
- Loricifera
- Priapulida (priapulid worms)
- Nematoda (roundworms)
- Nematomorpha (horsehair worms)
- Onychophora (velvet worms)
- Tardigrada (water bears)
- Arthropoda (insects, etc.)
- Superphylum Platyzoa
- Platyhelminthes (flatworms)
- Gastrotricha (gastrotrichs)
- Rotifera (rotifers)
- Acanthocephala (acanthocephalans)
- Gnathostomulida (jaw worms)
- Micrognathozoa (limnognathia)
- Cycliophora (pandora)
- Superphylum Lophotrochozoa (trochophore larvae / lophophores)
- Sipuncula (peanut worms)
- Nemertea (ribbon worms)
- Phoronida (horseshoe worms)
- Ectoprocta (moss animals)
- Entoprocta (goblet worms)
- Brachiopoda (brachipods)
- Mollusca (mollusks)
- Annelida (segmented worms) Animals are a major group of organisms, classified as the kingdom Animalia or Metazoa. In general they are multicellular, capable of locomotion and responsive to their environment, and feed by consuming other organisms. Their body plan becomes fixed as they develop, usually early on in their development as embryos, although some undergo a process of metamorphosis later on. Along with sponges, gastropods, emus, dolphins and all other animals, Homo sapiens sapiens meet all the criteria above for membership in the group of organisms known as animals and they do not meet the criteria of the other groups. Some humans often consider themselves separate from animals, not on the grounds of biology, but through the use of "other contexts". Whilst self-delusion may be a unique characteristic of the human species it is not cause for exclusion from the Kingdom Animalia. The name animal comes from the Latin word animal, of which animalia is the plural, and ultimately from anima, meaning vital breath or soul.

Characteristics

Aristotle divided the living world between animals and plants, and this was followed by Carolus Linnaeus in the first hierarchical classification. Since then biologists have begun emphasizing evolutionary relationships, and so these groups have been restricted somewhat. For instance, microscopic protozoa were originally considered animals because they move, but are now treated separately. Kingdom Animalia has several characteristics that set it apart from other living things. First, animals are eukaryotic. This separates them from the Kingdom Monera. Second, animals are multicellular, which separates them from Kingdom Protista. Third, they are heterotrophic, setting them apart from Kingdom Plantae and several plant-like protists. Finally, Kingdom Animalia consists of organisms without cell walls, which makes it unique compared to Kingdom Plantae, algae, and Kingdom Fungi.

Structure

With a few exceptions, most notably the sponges (Phylum Porifera), animals have bodies differentiated into separate tissues. These include muscles, which are able to contract and control locomotion, and a nervous system, which sends and processes signals. There is also typically an internal digestive chamber, with one or two openings. Animals with this sort of organization are called metazoans, or eumetazoans when the former is used for animals in general. All animals have eukaryotic cells, surrounded by a characteristic extracellular matrix composed of collagen and elastic glycoproteins. This may be calcified to form structures like shells, bones, and spicules. During development it forms a relatively flexible framework upon which cells can move about and be reorganized, making complex structures possible. In contrast, other multicellular organisms like plants and fungi have cells held in place by cell walls, so develop by progressive growth. Also, unique to animal cells are the following intercellular junctions: tight junctions, gap junctions, and desmosomes.

Reproduction and development

Nearly all animals undergo some form of sexual reproduction. Adults are diploid or occasionally polyploid. They have a few specialized reproductive cells, which undergo meiosis to produce smaller motile spermatozoa or larger non-motile ova. These fuse to form zygotes, which develop into new individuals. Many animals are also capable of asexual reproduction. This may take place through parthenogenesis, where fertile eggs are produced without mating, or in some cases through fragmentation. A zygote initially develops into a hollow sphere, called a blastula, which undergoes rearrangement and differentiation. In sponges, blastula larvae swim to a new location and develop into a new sponge. In most other groups, the blastula undergoes more complicated rearrangement. It first invaginates to form a gastrula with a digestive chamber, and two separate germ layers - an external ectoderm and an internal endoderm. In most cases, a mesoderm also develops between them. These germ layers then differentiate to form tissues and organs. Animals grow by indirectly using the energy of sunlight. Plants use this energy to turn air into simple sugars using a process known as photosynthesis. These sugars are then used as the building blocks which allow the plant to grow. When animals eat these plants (or eat other animals which have eaten plants), the sugars produced by the plant are used by the animal. They are either used directly to help the animal grow, or broken down, releasing stored solar energy, and giving the animal the energy required for motion. This process is known as glycolysis.

Origin and fossil record

Animals are generally considered to have evolved from flagellate protozoa. Their closest living relatives are the choanoflagellates, collared flagellates that have the same structure as certain sponge cells do. Molecular studies place them in a supergroup called the opisthokonts, which also include the fungi and a few small parasitic protists. The name comes from the posterior location of the flagellum in motile cells, such as most animal sperm, whereas other eukaryotes tend to have anterior flagella. The first fossils that might represent animals appear towards the end of the Precambrian, around 600 million years ago, and are known as the Vendian biota. These are difficult to relate to later fossils, however. Some may represent precursors of modern phyla, but they may be separate groups, and it is possible they are not really animals at all. Aside from them, most animal phyla with known phyla make a more or less simultaneous appearance during the Cambrian period, about 570 million years ago. It is still disputed whether this event, called the Cambrian explosion, represents a rapid divergence between different groups or a change in conditions that made fossilization possible.

Groups of animals

The sponges (Porifera) diverged from other animals early. As mentioned, they lack the complex organization found in most other phyla. Their cells are differentiated, but not organized into distinct tissues. Sponges are sessile and typically feed by drawing in water through pores all over the body, which is supported by a skeleton typically divided into spicules. The extinct Archaeocyatha, which have fused skeletons, may represent sponges or a separate phylum. Among the eumetazoan phyla, two are radially symmetric and have digestive chambers with a single opening, which serves as both the mouth and the anus. These are the Cnidaria, which include anemones, corals, and jellyfish, and the Ctenophora or comb jellies. Both have distinct tissues, but they are not organized into organs. There are only two main germ layers, the ectoderm and endoderm, with only scattered cells between them. As such, these animals are sometimes called diploblastic. The tiny phylum Placozoa is similar, but individuals do not have a permanent digestive chamber. The remaining animals form a monophyletic group called the Bilateria. For the most part, they are bilaterally symmetric, and often have a specialized head with feeding and sensory organs. The body is triploblastic, i.e. all three germ layers are well-developed, and tissues form distinct organs. The digestive chamber has two openings, a mouth and an anus, and there is also an internal body cavity called a coelom or pseudocoelom. There are exceptions to each of these characteristics, however - for instance adult echinoderms are radially symmetric, and certain parasitic worms have extremely simplified body structures. Genetic studies have considerably changed our understanding of the relationships within the Bilateria. Most appear to belong to four major lineages: # Deuterostomes # Ecdysozoa # Platyzoa # Lophotrochozoa In addition to these, there are a few small groups of bilaterians with relatively similar structure that appear to have diverged before these major groups. These include the Acoelomorpha, Rhombozoa, and Orthonectida. The Myxozoa, single-celled parasites that were originally considered Protozoa, are now believed to have developed from the Bilateria as well.

Deuterostomes

Deuterostomes differ from the other Bilateria, called protostomes, in several ways. In both cases there is a complete digestive tract. However, in protostomes the initial opening (the archenteron) develops into the mouth, and an anus forms separately. In deuterostomes this is reversed. In most protostomes cells simply fill in the interior of the gastrula to form the mesoderm, called schizocoelous development, but in deuterostomes it forms through evagination of the endoderm, called enterocoelic pouching. Deuterostomes also have a dorsal, rather than a ventral, nerve chord and their embryos undergo different cleavage. All this suggests the deuterostomes and protostomes are separate, monophyletic lineages. The main phyla of deuterostomes are the Echinodermata and Chordata. The former are radially symmetric and exclusively marine, such as sea stars, sea urchins, and sea cucumbers. The latter are dominated by the vertebrates, animals with backbones. These include fish, amphibians, reptiles, birds, and mammals. In addition to these, the deuterostomes also include the Hemichordata or acorn worms. Although they are not especially prominent today, the important fossil graptolites may belong to this group. The Chaetognatha or arrow worms may also be deuterostomes, but this is less certain.

Ecdysozoa

The Ecdysozoa are protostomes, named after the common trait of growth by moulting or ecdysis. The largest animal phylum belongs here, the Arthropoda, including insects, spiders, crabs, and their kin. All these organisms have a body divided into repeating segments, typically with paired appendages. Two smaller phyla, the Onychophora and Tardigrada, are close relatives of the arthropods and share these traits. The ecdysozoans also include the Nematoda or roundworms, the second largest animal phylum. Roundworms are typically microscopic, and occur in nearly every environment where there is water. A number are important parasites. Smaller phyla related to them are the Nematomorpha or horsehair worms, which are visible to the unaided eye, and the Kinorhyncha, Priapulida, and Loricifera, which are all microscopic. These groups have a reduced coelom, called a pseudocoelom. The remaining two groups of protostomes are sometimes grouped together as the Spiralia, since in both embryos develop with spiral cleavage.

Platyzoa

The Platyzoa include the phylum Platyhelminthes, the flatworms. These were originally considered some of the most primitive Bilateria, but it now appears they developed from more complex ancestors. A number of parasites are included in this group, such as the flukes and tapeworms. Flatworms lack a coelom, as do their closest relatives, the microscopic Gastrotricha. The other platyzoan phyla are microscopic and pseudocoelomate. The most prominent are the Rotifera or rotifers, which are common in aqueous environments. They also include the Acanthocephala or spiny-headed worms, the Gnathostomulida, Micrognathozoa, and possibly the Cycliophora. These groups share the presence of complex jaws, from which they are called the Gnathifera.

Lophotrochozoa

The Lophotrochozoa include two of the most successful animal phyla, the Mollusca and Annelida. The former includes animals such as snails, clams, and squids, and the latter comprises the segmented worms, such as earthworms and leeches. These two groups have long been considered close relatives because of the common presence of trochophore larvae, but the annelids were considered closer to the arthropods, because they are both segmented. Now this is generally considered convergent evolution, owing to many morphological and genetic differences between the two phyla. The Lophotrochozoa also include the Nemertea or ribbon worms, the Sipuncula, and several phyla that have a fan of cilia around the mouth, called a lophophore. These were traditionally grouped together as the lophophorates, but it now appears they are paraphyletic, some closer to the Nemertea and some to the Mollusca and Annelida. They include the Brachiopoda or lamp shells, which are prominent in the fossil record, the Entoprocta, the Phoronida, and possibly the Ectoprocta or moss animals.

History of classification

In Linnaeus' original scheme, the animals were one of three kingdoms, divided into the classes of Vermes, Insecta, Pisces, Amphibia, Aves, and Mammalia. Since then the last four have all been subsumed into a single phylum, the Chordata, whereas the various other forms have been separated out. The above lists represent our current understanding of the group, though there is some variation from source to source.

Usage of the word animal

In everyday usage animal refers to any member of the animal kingdom that is not a human being, and sometimes excludes insects (although including such arthropods as crabs). This confusion stems primarily from the familiarity with zoo animals, farm animals and pets, not from an analytical distinction between insects, humans and the rest of the animal kingdom.

Examples

Some well-known types of animals, listed by their common names:
- alpaca, ant, antelope, badger, bat, bear, bee, beetle, bird, bison, butterfly, cat, chicken, cockroach, coral, cow, deer, dinosaur, dog, dolphin, earthworm, elephant, elk, fish, fly, fox, frog, giraffe, goat, gorilla, hippopotamus, horse, human, iguana, jellyfish, kangaroo, lion, lizard, llama, lynx, monkey, mouse, nightingale, octopus, owl, ox, parrot, penguin, pig, quail, rabbit, rat, rhinoceros, salamander, scorpion, seahorse, shark, sheep, sloth, snake, spider, squid, starfish, tiger, turtle, urial, vole, whale, wolf, yak, zebra

References

External links

- [http://www.animool.com/animals/index.jsp Animals Search Engine]
- [http://www.wikianimals.com wikianimals.com] - Documenting the animal kingdom
- [http://tolweb.org/tree?group=Animals&contgroup=Eukaryotes Tree of Life]
- [http://www.arkive.org A Multimedia Database of Various UK or Endangered Species]
- [http://freepages.genealogy.rootsweb.com/~wakefield/animals.html Animals and Birds Names] - Large table of words: animal, collective, male, female, young, & home
- [http://www273.pair.com/med/words/animal_adjectives.htm English Animal Adjectives]
- [http://www.georgetown.edu/faculty/ballc/animals/animals.html Sounds of the World's Animals] - animal sounds in many languages
- [http://www.findsounds.com/ FindSounds - Search the Web for Sounds] - sound files including animal sound files
- [http://www.australianfauna.com/ Australian Animals]
- [http://www.animalreviews.com AnimalReviews] - animals reviewed and evaluated
- [http://animals.timduru.org/ The animal photo archive] - Photos of animals
- [http://www.wildlife-photo.org Photo gallery of animals pictures from the entire world.]
- [http://www.wildlife-photo.org/birds_list.htm Birds Name Check List in Latin, English, Russian and Hebrew.]
- [http://www.wildanimalsonline.com Wild Animals Online] - an online encyclopedia of wild animals - facts, photos Category:Animals zh-min-nan:Tōng-bu̍t ko:동물 ms:Haiwan ja:動物 simple:Animal th:สัตว์

Earthworm

Image:Earth-worm_1.jpg Earthworm is the common reference for the larger members of the Oligochaeta (which is either a class or subclass depending on the author) in the phylum Annelida. In classical systems they were placed in the order Opisthopora, on the basis of the male pores opening to the outside of body posterior to the female pores, even though the male segments are anterior to the female. Cladistic studies have supported placing them instead in the Haplotaxida, which also includes the family Haplotaxidae. Folk names for earthworm include "dew-worm"，"night crawler" and "angleworm." Earthworms are also called megadriles (or big worms), as opposed to the microdriles, which include the families Tubificidae, Lumbriculidae, and Enchytraeidae, among others. The haplotaxids have been traditionally considered microdriles. The megadriles are characterized by having a multilayered clitellum (which is much more obvious than the single-layered one of the microdriles), a vascular system with true capillaries, and male pores behind the female pores.

Overview

There are over 2,200 species known worldwide, existing everywhere but Arctic and arid climates. They range in size from two centimeters (less than one inch) to over three meters (almost ten feet) in the Giant Gippsland Earthworm. Amongst the main earthworm species commonly found in the soil are the red coloured Lumbricus terrestris, which dwells close to and leaves its deposits on the surface, whilst the greyish blue Allolobophora caliginosa is deeper burrowing. In temperate zone areas, the most commonly seen earthworms are lumbricids (Lumbricidae), mostly due to the recent rapid spread of a relatively small number of European species, but there are several other families, e.g. Megascolecidae, Sparganophilidae, Glossoscolecidae, Haplotaxidae, and others. These other families are often very different from the lumbricids in behavior, physiology and habitat.

Anatomy

Earthworms have a closed circulatory system. They have two main blood vessels that extend through the length of their body: a ventral blood vessel which leads the blood to the posterior end, and a dorsal blood vessel which leads to the anterior end. The dorsal vessel is contractile and pumps blood forward, where it is pumped into the ventral vessel by a series of "hearts" which vary in number in the different taxa. A typical lumbricid will have 5 pairs of hearts. The blood is distributed from the ventral vessel into capillaries on the body wall and other organs and into a vascular sinus in the gut wall where gases and nutrients are exchanged. This arrangement may be complicated in the various groups by suboesophageal, supraoesophageal, parietal and neural vessels, but the basic arrangement holds in all earthworms.

Reproduction

Earthworms are hermaphrodites (both female and male organs within the same individual) but cannot fertilize their own eggs. They have testes, seminal vesicles and male pores which produce, store and release the sperm, and ovaries and ovipores. However, they also have one or more pairs of spermathecae (depending on the species) that are internal sacs which receive and store sperm from the other worm in copulation. Copulation and reproduction are separate processes in earthworms. The mating pair overlap front ends ventrally and each exchanges sperm with the other. The cocoon, or egg case, is secreted by the clitellum, the external glandular band which is near the front of the worm, but behind the spermathecae. Some indefinite time after copulation, long after the worms have separated, the clitellum secretes the cocoon which forms a ring around the worm. The worm then backs out of the ring, and as it does so, injects its own eggs and the other worm's sperm into it. As the worm slips out, the ends of the cocoon seal to form a vaguely lemon-shaped incubator (cocoon) in which the embryonic worms develop. They emerge as small, but fully formed earthworms, except for lacking the sexual structures, which develop later. Some earthworm species are mostly parthenogenetic, in which case the male structures and spermathecae may become abnormal, or missing.

Behavior

One often sees earthworms come to the surface in large numbers after a rainstorm. There are three theories for this behavior. The first is that the waterlogged soil has insufficient oxygen for the worms, therefore, earthworms come to the surface to get the oxygen they need and breathe more easily. Secondly, some species (notably Lumbricus terrestris) come to the surface to mate. This behavior is, however, limited to a few species. Thirdly, the worms may be using the moist conditions on the surface to travel more quickly than they can underground, thus colonizing new areas more quickly. This is in any event a dangerous activity in the daytime, since earthworms die quickly when exposed to direct sunlight with its strong UV content. UV

Locomotion and importance to soil

Earthworms travel underground by the means of waves of muscular contractions which alternately shorten and lengthen the body. The shortened part is anchored to the surrounding soil by tiny claw-like bristles (setae) set along its segmented length. The whole process is aided by the secretion of a slimy lubricating mucous. In more compacted soils the earthworm actually eats its way through the soil, cutting a passage with its muscular pharynx and dragging the rest of the body along. The ingested soil is ground up, digested, and the waste deposited behind the worm. This process aerates and mixes the soil, and is often considered greatly helpful by gardeners and farmers. In addition, many earthworms will come to the surface and graze on the higher concentrations of organic matter there, mixing it with the mineral soil. Because a high level of organic matter is associated with soil fertility, an abundance of earthworms is a happy sight for the organic gardener. In fact as long ago as 1881 Charles Darwin wrote:

"It may be doubted whether there are any other animals which have played so important a part in the history of the world, as have these lowly creatures"

:(The Formation Of Vegetable Mould Through The Action Of Worms, Charles Darwin)

Benefits

The major benefits of earthworm activities to soil fertility can be summarised as:
- Biological. The earthworm is essential to composting; the process of converting dead organic matter into rich humus, a medium vital to the growth of healthy plants, and thus ensuring the continuance of the cycle of fertility. This is achieved by the worm's actions of pulling down below any organic matter deposited on the soil surface (eg, leaf fall, manure, etc) either for food or when it needs to plug its burrow. Once in the burrow, the worm will shred the leaf and partially digest it, then mingle it with the earth by saturating it with intestinal secretions. Worm casts (see below) can contain 40% more humus than the top 6" of soil in which the worm is living.
- Chemical. As well as dead organic matter, the earthworm also ingests any other soil particles that are small enough (including stones up to 1/20 of an inch across) into its 'crop' wherein minute fragments of grit grind everything into a fine paste which is then digested in the stomach. When the worm excretes this in the form of casts which are deposited on the surface or deeper in the soil, a perfectly balanced selection of minerals and plant nutrients is made available in an accessible form. Investigations in the US show that fresh earthworm casts are 5 times richer in available nitrogen, 7 times richer in available phosphates and 11 times richer in available potash than the surrounding upper 6 inches (150 mm) of soil. In conditions where there is plenty of available humus, the weight of casts produced may be greater than 4.5 kg (10 lb) per worm per year, in itself an indicator of why it pays the gardener or farmer to keep worm populations high.
- Physical. By its burrowing actions, the earthworm is of great value in keeping the soil structure open, creating a multitude of channels which allow the processes of both aeration and drainage to occur. Permaculture co-founder Bill Mollison points out that by sliding in their tunnels, earthworms "act as an innumerable army of pistons pumping air in and out of the soils on a 24 hour cycle (more rapidly at night)" (Permaculture- A Designer's Manual, Tagari Press, 1988). Thus the earthworm not only creates passages for air and water to traverse, but is itself a vital component in the living biosystem that is healthy soil. It is important that we do not take the humble earthworm for granted. Dr. W. E. Shewell Cooper observed "tremendous numerical differences between adjacent gardens" (Soil, Humus And Health), and worm populations are affected by a host of environmental factors, many of which can be influenced by good management practices on the part of the gardener or farmer. Darwin estimated that arable land contains up to 53,000 worms per acre (13/m²), but more recent research from Rothamsted Experimental Station has produced figures suggesting that even poor soil may support 250,000/acre (62/m²), whilst rich fertile farmland may have up to 1,750,000/acre (432/m²). Professor I. L. Heiberg of State University of New York College of Environmental Science and Forestry has stated that in optimum conditions, the worm population may even reach 250,000,000 per acre (62,000/m²), meaning that the weight of earthworms beneath the farmer's soil could be greater than that of his livestock upon its surface. One thing is certain however: rich, fertile soil that is cared for organically and well-fed and husbanded by its steward will reap its reward in a healthy worm population, whilst denuded, overworked, and eroded land will almost certainly contain fewer, scrawny, undernourished specimens.

Earthworms as invasives

Lumbricid earthworms are invasive to North America and not only have displaced native earthworms in much of the continent, but have invaded areas where earthworms did not formerly exist. There are no native earthworms in much of North America, especially in the north, and the forests there developed relying on a large amount of undecayed leaf matter. The worms decompose that leaf layer, making the habitat unsurvivable for young native trees, ferns and wildflowers. Currently there is no economically feasible method for controlling earthworms in forests, besides preventing introductions. Earthworms normally spread slowly, but can be widely introduced by human activities such as construction earthmoving, or by fishermen releasing bait, or by plantings from other areas.

Special habitats

While, as the name earthworm suggests, the main habitat of earthworms is in soil, the situation is more complicated than that. The brandling worm Eisenia foetida (or fetida) lives in decaying plant matter and manure. Arctiostrotus vancouverensis from Vancouver Island and the Olympic Peninsula is generally found in decaying conifer logs or in extremely acid humus. Alollobophora limicola and Sparganophilus and several others are found in mud in streams. Even in the soil species, there are special habitats, such as soils derived from serpentine which have an earthworm fauna of their own.

Ecology

Earthworm populations depend on both physical and chemical properties of the soil (such as soil temperature, moisture, pH, salts, aeration and texture), as well as available food, and the ability of the species to reproduce and disperse. One of the most important environmental factors is pH, but earthworms vary in their preferences. Most earthworms favor neutral to slightly acid soil. However, Lumbricus terrestris are still present in pH of 5.4 and Dendrobaena octaedra at pH of 4.3 and some Megascolecidae are present in extremely acid humic soils. Soil pH may also influence the numbers of worms that go into diapause. The more acid the soil, the sooner worms went into diapause, and remain in diapause the longest time at pH of 6.4. Earthworms form the base of many food chains. They are preyed upon by many species of birds, e.g. starlings, thrushes, gulls, crows, and robins. Mammals such as hedgehogs and moles eat many earthworms as well. Earthworms are also eaten by many invertebrates such as Ground beetles and other beetles, snails, slugs and flatworms. Earthworms have many internal parasites including Protozoa, Platyhelminthes, nematodes. They are found in many part of earthworms' bodies like the blood, seminal vesicles, coelom, intestine, or in the cocoons.

Threats to earthworms

The application of chemical fertilisers, sprays and dusts can have a disastrous effect on earthworm populations. Nitrogenous fertilisers tend to create acid conditions, which are fatal to the worms, and often dead specimens are to be found on the surface following the application of substances like DDT, lime sulphur and lead arsenate. In Australia, the use of superphosphate on pastures almost totally wiped out the giant Gippsland earthworm. In addition, as earthworms are processors of large amounts of plant and mineral materials, even if not killed themselves they can accumulate pollutants such as DDT, lead, cadmium, and dioxins at levels up to 20 times higher than in the soil, which in turn are passed on at lethal dosages to the wildlife which feed upon them such as foxes, moles or birds. Therefore, the most reliable way to maintain or increase the levels of worm population in the soil is to avoid the application of artificial chemicals, as well as adding organic matter, preferably as a surface mulch, on a regular basis. This will not only provide them with their food and nutrient requirements, but also creates the optimum conditions of heat (cooler in summer and warmer in winter) and moisture to stimulate their activity. A recent threat to earthworm populations in the UK is the New Zealand Flatworm (Artiposthia triangulata), which feeds upon the earthworm, but in this country has no natural predator itself. At present sightings of the NZFW have been mainly localised, but this is no reason for complacency as it has spread extensively since its introduction in 1960 through contaminated soil and plant pots. Any sightings of the flatworm should be reported to the Scottish Crop Research Institute, who are monitoring its spread.

Economic Impact

Various species of worms are used in vermiculture, the practice of feeding organic waste to earthworms to decompose (digest) it, a form of composting by the use of worms. These are usually Eisenia foetida or the Brandling worm, also known as the Tiger worm or Red Wriggler, and are distinct from soil-dwelling earthworms. Earthworms are sold all over the world. The earthworm market is sizeable. According to Doug Collicut (see "Nightcrawler" link below), "In 1980, 370 million worms were exported from Canada, with a Canadian export value of $13 million and an American retail value of $54 million."

External References

- [http://www.sarep.ucdavis.edu/worms/ Earthworm Information (UC Davis)]
- [http://www.encyclopedia.com/html/e1/earthworm.asp earthworm on Encyclopedia.com]
- [http://www.naturenorth.com/fall/ncrawler/ncrawlF.html Biology of the Night Crawler (Lumbricus terrestris)]
- [http://www.naturewatch.ca/english/wormwatch/about/guide/about_guide_redworms.html WormWatch - Field guide to earthworms]
- [http://flatworm.csl.gov.uk/ New Zealand flatworm page (UK Govt.)]
- [http://www.hdra.org.uk/factsheets/pc21.htm New Zealand flatworm page (HDRA)]
- [http://www.strayreality.com/plants2/earthworms.htm Earthworms Saving the Earth]
- [http://www.naturewatch.ca/english/wormwatch/index.html Worm Watch] Canadian worm awareness and appreciation site, with detailed worm anatomy.
- [http://www.dnr.state.mn.us/invasives/terrestrialanimals/earthworms/index.html Minnesota Invasive Earthworms] Minnesota DNR information on the negative impacts of earthworms Category:Annelids (worms) Category:cryptic animals ja:ミミズ

Annelida

Class Polychaeta (paraphyletic?)
Class Clitellata
   Oligochaeta - Earthworms and others
   Acanthobdellida
   Branchiobdellida
   Hirudinea - Leeches
Class Myzostomida
Class Archiannelida (polyphyletic)
Class Echiura

- Some authors consider the subclasses
under Clitellata to be classes The annelids, collectively called Annelida (from Latin annellus "little ring"), are a large phylum of animals, comprising the segmented worms, with about 15 000 modern species including the well-known earthworms and leeches. They are found in most wet environments, and include many terrestrial, freshwater, and especially marine species (such as the often gorgeous polychaetes), as well as some which are parasitic or mutualistic. They range in length from under a millimetre to over 3 metres.

Anatomy

Annelids are triploblastic protostomes. The body cavity is a coelom, a fluid-filled cavity in which the gut and other organs are suspended. 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. 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 usually specialized. Different species of annelids have a wide variety of diets, including active and passive hunters, scavengers, filter feeders, direct deposit feeders which 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 which 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 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 can regrow the posteriormost segments in most instances.

Sexual reproduction

Sexual reproduction allows a species to better adapt to its environment. Some annelid species are hermaphroditic, while others have distinct genders. Hermaphrodite annelids like earthworms mate periodically throughout the year in favored environmental conditions. Earthworms mate by copulation. Two worms which are attracted by each other's secretions lay their bodies together with their heads pointing opposite directions. The fluid is transferred from the male pore to the other worm. Different methods of sperm tranference 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. 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. The oligochaetes and leeches tend to be hermaphroditic and lack free-living larvae of this sort. While annelids have some regenerative abilities, sometimes to the point where each half of an adult divided cross-wise will survive, this is not universal, and especially does not occur among the earthworms as folklore would suggest.

Fossil record

The annelid fossil record is sparse, but a few definite forms are known as early as the Cambrian, and there are some signs they were around in the later Precambrian. A few small groups have been treated as separate phyla: the Pogonophora and Vestimentifera, now included in the family Siboglinidae, and the Echiura.

Relationships

The arthropods and their kin have long been considered the closest relatives of the annelids, on account of their common segmented structure, but a number of differences between the two groups suggest this may be convergent evolution. The other major phylum which is of definite relation to the annelids are the mollusks, which share with them the presence of trochophore larvae. These groups are united as the Trochozoa, and when the arthropods are included, they and the annelids are treated in a subgroup called the Articulata.

Classes and subclasses of Annelida

- Clitellata
- Oligochaeta - The class Oligochaeta includes the megadriles (earthworms), which are both aquatic and terrestrial, and the microdrile families such as tubificids, which include many marine members as well.
- Leeches (Hirudinea) - These include both bloodsucking external parasites and predators of small invertebrates.
- Polychaeta - This is the largest group of the Annelids and majority are marine. All segments are identical each consisting a parapodia. The parapodia are used for swimming, burrowing and feeding current. Category:Animals Category:Annelids (worms) ja:環形動物

Species

In biology, a species is the basic unit of biodiversity. In scientific classification, a species is assigned a two-part name in Latin. The genus is listed first (and capitalized), followed by a specific epithet. For example, humans belong to the genus Homo, and are in the species Homo sapiens. The name of the species is the whole binomial not just the second term (the specific epithet). The binomial, and most other purely formal aspects of the biological codes of nomenclature, were formalized by Carolus Linnaeus in the 1700's and as a result are called the "Linnaean system". At that time, species were thought to represent independent acts of creation by God, and were therefore considered objectively real and immutable. Since the advent of the theory of evolution, the conception of species has undergone vast changes in biology, however no consensus on the definition of the word has yet been reached. The most commonly cited definition of "species" was first coined by Ernst Mayr. By this definition, called the biological species concept or isolation species concept, species are "groups of actually or potentially interbreeding natural populations which are reproductively isolated from other such groups". However, many other species concepts are also used (see other definitions of species below). The scientific name of a species is properly typeset in italics. When an unknown species is being referred to this may be done by using the abbreviation "sp." in the singular or "spp." in the plural in the place of the second part of the scientific name. Note that the word "specie" is not the singular of "species". It refers to coined money.

Definitions of species

The definition of a species given above as taken from Mayr, is somewhat idealistic. Since it assumes sexual reproduction, it leaves the term undefined for a large class of organisms that reproduce asexually. Biologists frequently do not know whether two morphologically similar groups of organisms are "potentially" capable of interbreeding. Further, there is considerable variation in the degree to which hybridization may succeed under natural and experimental conditions, or even in the degree to which some organisms use sexual reproduction between individuals to breed. Consequently, several lines of thought in the definition of species exist: ; Typological species : A group of organisms in which individuals are members of the species if they sufficiently conform to certain fixed properties. The clusters of variations or phenotypes within specimens (ie: longer and shorter tails) would differentiate the species. This method was used as a "classical" method of determining species, such as with Linnaeus early in evolutionary theory. However, we now know that different phenotypes do not always constitute different species (e.g.: a 4-winged Drosophila born to a 2-winged mother is not a different species). Species named in this manner are called morphospecies. ; Morphological species : A population or group of populations that differs morphologically from other populations. For example, we can distinguish between a chicken and a duck because they have different shaped bills and the duck has webbed feet. Species have been defined in this way since well before the beginning of recorded history. This species concept is much criticised because more recent genetic data reveals that genetically distinct populations may look very similar and, contrarily, large morphological differences sometimes exist between very closely-related populations. Nonetheless, most species known have been described solely from morphology. ; Biological / Isolation species : A set of actually or potentially interbreeding populations. This is generally the most useful formulation for scientists working with living examples of the higher taxa like mammals, fish, and birds, but meaningless for organisms that do not reproduce sexually. It does not distinguish between the theoretical possibility of interbreeding and the actual likelihood of gene flow between populations and is thus impractical in instances of allopatric (geographically isolated) populations. The results of breeding experiments done in artificial conditions may or may not reflect what would happen if the same organisms encountered each other in the wild, making it difficult to gauge whether or not the results of such experiments are meaningful in reference to natural populations. ; Mate-recognition species : A group of organisms that are known to recognise one another as potential mates. Like the isolation species concept above, it applies only to organisms that reproduce sexually. Unlike the isolation species concept, it focuses specifically on pre-mating reproductive isolation. ; Phylogenetic / Evolutionary / Darwinian species : A group of organisms that shares an ancestor; a lineage that maintains its integrity with respect to other lineages through both time and space. At some point in the progress of such a group, members may diverge from one another: when such a divergence becomes sufficiently clear, the two populations are regarded as separate species. ; Microspecies : Species that reproduce without meiosis or mitosis so that each generation is genetically identical to the previous generation. See also apomixis. In practice, these definitions often coincide, and the differences between them are more a matter of emphasis than of outright contradiction. Nevertheless, no species concept yet proposed is entirely objective, or can be applied in all cases without resorting to judgement. Given the complexity of life, some have argued that such an objective definition is in all likelihood impossible, and biologists should settle for the most practical definition. For most vertebrates, this is the biological species concept, and to a lesser extent (or for different purposes) the phylogenetic species concept. Many BSC subspecies are considered species under the PSC; the difference between the BSC and the PSC can be summed up insofar as that the BSC defines a species as a consequence of manifest evolutionary history, while the PSC defines a species as a consequence of manifest evolutionary potential. Thus, a PSC species is "made" as soon as an evolutionary lineage has started to separate, while a BSC species starts to exist only when the lineage separation is complete.

Importance in biological classification

The idea of species has a long history. It is one of the most important levels of classification, for several reasons:
- It often corresponds to what lay people treat as the different basic kinds of organism - dogs are one species, cats another.
- It is the standard binomial nomenclature (or trinomial nomenclature) by which scientists typically refer to organisms.
- It is the only taxonomic level which has empirical content, in the sense that asserting that two animals are of different species is saying something more than classificatory about them. After thousands of years of use, the concept remains central to biology and a host of related fields, and yet also remains at times ill-defined and controversial.

Implications of assignment of species status

The naming of a particular species should be regarded as a hypothesis about the evolutionary relationships and distinguishability of that group of organisms. As further information comes to hand, the hypothesis may be confirmed or refuted. Sometimes, especially in the past when communication was more difficult, taxonomists working in isolation have given two distinct names to individual organisms later identified as the same species. When two named species are discovered to be of the same species, the older species name is usually retained, and the newer species name dropped, a process called synonymization, or convivially, as lumping. Dividing a taxon into multiple, often new, taxons is called splitting. Taxonomists are often referred to as "lumpers" or "splitters" by their colleagues, depending on their personal approach to recognizing differences or commonalities between organisms (see lumpers and splitters). Traditionally, researchers relied on observations of anatomical differences, and on observations of whether different populations were able to interbreed successfully, to distinguish species; both anatomy and breeding behavior are still important to assigning species status. As a result of the revolutionary (and still ongoing) advance in microbiological research techniques, including DNA analysis, in the last few decades, a great deal of additional knowledge about the differences and similarities between species has become available. Many populations which were formerly regarded as separate species are now considered to be a single taxon, and many formerly grouped populations have been split. Any taxonomic level (species, genus, family, etc.) can be synonymized or split, and at higher taxonomic levels, these revisions have been still more profound. From a taxonomical point of view, groups within a species can be defined as being of a taxon hierarchically lower than a species. In zoology only the subspecies is used, while in botany the variety, subvariety, and form are used as well.

The isolation species concept in more detail

In general, for large, complex, organisms that reproduce sexually (such as mammals and birds), one of several variations on the isolation or biological species concept is employed. Often, the distinction between different species, even quite closely related ones, is simple. Horses (Equus caballus) and donkeys (Equus asinus) are easily told apart even without study or training, and yet are so closely related that they can interbreed after a fashion. Because the result, a mule or hinny, is not usually fertile, they are clearly separate species. But many cases are more difficult to decide. This is where the isolation species concept diverges from the evolutionary species concept. Both agree that a species is a lineage that maintains its integrity over time, that is diagnosably different to other lineages (else we could not recognise it), is reproductively isolated (else the lineage would merge into others, given the chance to do so), and has a working intra-species recognition system (without which it could not continue). In practice, both also agree that a species must have its own independent evolutionary history—otherwise the characteristics just mentioned would not apply. The species concepts differ in that the evolutionary species concept does not make predictions about the future of the population: it simply records that which is already known. In contrast, the isolation species concept refuses to assign the rank of species to populations that, in the best judgement of the researcher, would recombine with other populations if given the chance to do so.

The isolation question

There are, essentially, two questions to resolve. First, is the proposed species consistently and reliably distinguishable from other species? Secondly, is it likely to remain so in the future? To take the second question first, there are several broad geographic possibilities.
- The proposed species are sympatric—they occupy the same habitat. Observation of many species over the years has failed to establish even a single instance of two diagnostically different populations that exist in sympatry and have then merged to form one united population. Without reproductive isolation, population differences cannot develop, and given reproductive isolation, gene flow between the populations cannot merge the differences. This is not to say that cross breeding does not take place at all, simply that it has become negligible. Generally, the hybrid individuals are less capable of successful breeding than pure-bred individuals of either species.
- The proposed species are allopatric—they occupy different geographical areas. Obviously, it is not possible to observe reproductive isolation in allopatric groups directly. Often it is not possible to achieve certainty by experimental means either: even if the two proposed species interbreed in captivity, this does not demonstrate that they would freely interbreed in the wild, nor does it always provide much information about the evolutionary fitness of hybrid individuals. A certain amount can be inferred from other experimental methods: for example, do the members of population A respond appropriately to playback of the recorded mating calls of population B? Sometimes, experiments can provide firm answers. For example, there are seven pairs of apparently almost identical marine snapping shrimp (Altheus) populations on either side of the Isthmus of Panama, which did not exist until about 3 million years ago. Until then, it is assumed, they were members of the same seven species. But when males and females from opposite sides of the isthmus are placed together, they fight instead of mating. Even if the isthmus were to sink under the waves again, the populations would remain genetically isolated: therefore they are now different species. In many cases, however, neither observation nor experiment can produce certain answers, and the determination of species rank must be made on a 'best guess' basis from a general knowledge of other related organisms.
- The proposed species are parapatric—they have breeding ranges that abut but do not overlap. This is fairly rare, particularly in temperate regions. The dividing line is often a sudden change in habitat (an ecotone) like the edge of a forest or the snow line on a mountain, but can sometimes be remarkably trivial. The parapatry itself indicates that the two populations occupy such similar ecological roles that they cannot coexist in the same area. Because they do not crossbreed, it is safe to assume that there is a mechanism, often behavioral, that is preventing gene flow between the populations, and that therefore they should be classified as separate species.
- There is a hybrid zone where the two populations mix. Typically, the hybrid zone will include representatives of one or both of the 'pure' populations, plus first-generation and back-crossing hybrids. The strength of the barrier to genetic transmission between the two pure groups can be assessed by the width of the hybrid zone relative to the typical dispersal distance of the organisms in question. The dispersal distance of oaks, for example, is the distance that a bird or squirrel can be expected to carry an acorn; the dispersal distance of Numbats is about 15 kilometres, as this is as far as young Numbats will normally travel in search of vacant territory to occupy after leaving the nest. The narrower the hybrid zone relative to the dispersal distance, the less gene flow there is between the population groups, and the more likely it is that they will continue on separate evolutionary paths. Nevertheless, it can be very difficult to predict the future course of a hybrid zone; the decision to define the two hybridizing populations as either the same species or as separate species is difficult and potentially controversial.
- The variation in the population is clinal; at either extreme of the population's geographic distribution, typical individuals are clearly different, but the transition between them is seamless and gradual. For example, the Koalas of northern Australia are clearly smaller and lighter in colour than those of the south, but there is no particular dividing line: the further south an individual Koala is found, the larger and darker it is likely to be; Koalas in intermediate regions are intermediate in weight and colour. In contrast, over the same geographic range, black-backed (northern) and white-backed (southern) Australian Magpies do not blend from one type to another: northern populations have black backs, southern populations white backs, and there is an extensive hybrid zone where both 'pure' types are common, as are crossbreeds. The variation in Koalas is clinal (a smooth transition from north to south, with populations in any given small area having a uniform appearance), but the variation in magpies is not clinal. In both cases, there is some uncertainty regarding correct classification, but the consensus view is that species rank is not justified in either. The gene flow between northern and southern magpie populations is judged to be sufficiently restricted to justify terming them subspecies (not full species); but the seamless way that local Koala populations blend one into another shows that there is substantial gene flow between north and south. As a result, experts tend to reject even subspecies rank in this case.

The difference question

Obviously, when defining a species, the geographic circumstances become meaningful only if the populations groups in question are clearly different: if they are not consistently and reliably distinguishable from one another, then we have no grounds for believing that they might be different species. The key question in this context, is "how different is different?" and the answer is usually "it all depends". In theory, it would be possible to recognise even the tiniest of differences as sufficient to delineate a separate species, provided only that the difference is clear and consistent (and that other criteria are met). There is no universal rule to state the smallest allowable difference between two species, but in general, very trivial differences are ignored on the twin grounds of simple practicality, and genetic similarity: if two population groups are so close that the distinction between them rests on an obscure and microscopic difference in morphology, or a single base substitution in a DNA sequence, then a demonstration of restricted gene flow between the populations will probably be difficult in any case. More typically, one or other of the following requirements must be met:
- It is possible to reliably measure a quantitative difference between the two groups that does not overlap. A population has, for example, thicker fur, rougher bark, longer ears, or larger seeds than another population, and although this characteristic may vary within each population, the two do not grade into one another, and given a reasonably large sample size, there is a definite discontinuity between them. Note that this applies to populations, not individual organisms, and that a small number of exceptional individuals within a population may 'break the rule' without invalidating it. The less a quantitative difference varies within a population and the more it varies between populations, the better the case for making a distinction. Nevertheless, borderline situations can only be resolved by making a 'best-guess' judgement.
- It is possible to distinguish a qualitative difference between the populations; a feature that does not vary continuously but is either entirely present or entirely absent. This might be a distinctively shaped seed pod, an extra primary feather, a particular courting behaviour, or a clearly different DNA sequence. Sometimes it is not possible to isolate a single difference between species, and several factors must be taken in combination. This is often the case with plants in particular. In eucalypts, for example, Corymbia ficifolia cannot be reliably distinguished from its close relative Corymbia calophylla by any single measure (and sometimes individual trees cannot be definitely assigned to either species), but populations of Corymbia can be clearly told apart by comparing the colour of flowers, bark, and buds, number of flowers for a given size of tree, and the shape of the leaves and fruit. When using a combination of characteristics to distinguish between populations, it is necessary to use a reasonably small number of factors (if more than a handful are needed, the genetic difference between the populations is likely to be insignificant and is unlikely to endure into the future), and to choose factors that are functionally independent (height and weight, for example, should usually be considered as one factor, not two).

Historical development of the species concept

In the earliest works of science, a species was simply an individual organism that represented a group of similar or nearly identical organisms. No other relationships beyond that group were implied. Aristotle used the words genus and species to mean generic and specific categories. Aristotle and other pre-Darwinian scientists took the species to be distinct and unchanging, with an "essence", like the chemical elements. When early observers began to develop systems of organization for living things, they began to place formerly isolated species into a context. To the modern mind, many of the schemes delineated are whimsical at best, such as those that determined consanguinity based on color (all plants with yellow flowers) or behavior (snakes, scorpions and certain biting ants). In the 18th century Carolus Linnaeus classified organisms according to differences in the form of reproductive apparatus. Although his system of classification sorts organisms according to degrees of similarity, it made no claims about the relationship between similar species. At the time, it was still widely believed that there is no organic connection between species, no matter how similar they appear; every species was individually created by God, a view today called creationism. This approach also suggested a type of idealism: the notion that each species exists as an "ideal form". Although there are always differences (although sometimes minute) between individual organisms, Linnaeus considered such variation problematic. He strove to identify individual organisms that were exemplary of the species, and considered other non-exemplary organisms to be deviant and imperfect. By the 19th century most naturalists understood that species could change form over time, and that the history of the planet provided enough time for major changes. As such, the new emphasis was on determining how a species could change over time. Jean-Baptiste Lamarck suggested that an organism could pass on an acquired trait to its offspring, i.e., the giraffe's long neck was attributed to generations of giraffes stretching to reach the leaves of higher treetops (this well-known and simplistic example, however, does not do justice to the breadth and subtlety of Lamarck's ideas). Lamarck's most important insight may have been that species can be extraordinarily fluid; his 1809 Zoological Philosophy contained one of the first logical refutations of creationism. With the acceptance of the work of Charles Darwin in the 1860s, Lamarck's view of evolution was quickly eclipsed. It was not until the late 20th century that his work began to be reexamined, and took its place as a fundamental stepping stone to the modern theory of adaptive mutation. Lamarck's long-discarded ideas of the goal-oriented evolution of species, also known the teleological process, have also received renewed attention, particularly by proponents of artificial selection. Charles Darwin and Alfred Wallace provided what scientists now consider the most powerful and compelling theory of evolution. Basically, Darwin argued that it is populations that evolve, not individuals. His argument relies on a radical shift in perspective from Linnaeus: rather than defining species in ideal terms (and searching for an ideal representative and rejecting deviations), Darwin considered variation among individuals to be natural. He further argued that variation, far from being problematic, actually provides the explanation for the existence of distinct species. Darwin's work drew on Thomas Malthus' insight that the rate of growth of a biological population will always outpace the rate of growth of the resources in the environment, such as the food supply. As a result, Darwin argued, not all the members of a population will be able to survive and reproduce. Those that did will, on average, be the ones possessing variations—however slight—that make them slightly better adapted to the environment. If these variable traits are heritable, then the offspring of the survivors will also possess them. Thus, over many generations, adaptive variations will accumulate in the population, while counter-adaptive will be eliminated. It should be emphasized that whether a variation is adaptive or non-adaptive depends on the environment: different environments favor different traits. Since the environment effectively selects which organisms live to reproduce, it is the environment (the "fight for existence") that selects the traits to be passed on. This is the theory of evolution by natural selection. In this model, the length of a giraffe's neck would be explained by positing that proto-giraffes with longer necks would have had a significant reproductive advantage to those with shorter necks. Over many generations, the entire population would be a species of long-necked animals. In 1859, when Darwin published his theory of natural selection, the mechanism behind the inheritance of individual traits was unknown. Although Darwin made some speculations on how traits are inherited (pangenesis), his theory relies only on the fact that inheritable traits exist, and are variable (which makes his accomplishment even more remarkable.) Although Gregor Mendel's paper on genetics was published in 1866, its significance was not recognized. It was not until 1900 that his work was rediscovered by Hugo de Vries, Carl Correns and Erich von Tschermak, who realised that the "inheritable traits" in Darwin's theory are genes. The theory of the evolution of species through natural selection has two important implications for discussions of species -- consequences that fundamentally challenge the assumptions behind Linnaeus' taxonomy. First, it suggests that species are not just similar, they may actually be related. Some students of Darwin argue that all species are descended from a common ancestor. Second, it supposes that "species" are not homogeneous, fixed, permanent things; members of a species are all different, and over time species change. This suggests that species do not have any clear boundaries but are rather momentary statistical effects of constantly changing gene-frequencies. One may still use Linnaeus' taxonomy to identify individual plants and animals, but one can no longer think of species as independent and immutable. The rise of a new species from a parental line is called speciation. There is no clear line demarcating the ancestral species from the descendant species. Although the current scientific understanding of species suggests there is no rigorous and comprehensive way to distinguish between different species in all cases, biologists continue to seek concrete ways to operationalize the idea. One of the most popular biological definitions of species is in terms of reproductive isolation; if two creatures cannot reproduce to produce fertile offspring, then they are in different species. This definition captures a number of intuitive species boundaries, but nonetheless has some problems, however. It has nothing to say about species that reproduce asexually, for example, and it is very difficult to apply to extinct species. Moreover, boundaries between species are often fuzzy: there are examples where members of one population can produce fertile offspring with a second population, and members of the second population can produce fertile offspring with members of a third population, but members of the first and third population cannot produces fertile offspring. Consequently, some people reject this notion of species. In recent years we have witnessed the drastic reduction in the size of breeding populations and the geographical range of many physically large mammals. In earlier times it was assumed that every species existed in at least a few thousand living individuals, except very rare relic, isolated groups. In the present, many well know mammal & bird species are so stressed by habitat loss, and other effects of the modern world, that only a very few breeding males may contribute the genetic material to a small number of breeding females. In these highly stressed conditions, the likelihood of change is very much greater. Mammals may become smaller, have darker fur, more stripes, more cautious behavior, even over time learn to co-exist with the human world. Very likely, evolution is radically accelerated, and we are only beginning to notice it. Species in transition before our eyes. It is possible that this severe stress is essential to the creation of new species, and may have been a prime factor throughout biological history, from other population reducing influences. Richard Dawkins defines two organisms as conspecific if and only if they have the same number of chromosomes and, for each chromosome, both organisms have the same number of nucleotides (The Blind Watchmaker, p. 118). However, most if not all taxonomists would strongly disagree. For example, in many amphibians, most notably in New Zealand's Leiopelma frogs, the genome consists of "core" chromosomes which are mostly invariable and accessory chromosomes, of which exist a number of possible combinations. Even though the chromosome numbers are highly variable between populations, these can interbreed successfully and form a single evolutionary unit. In plants, polyploidy is extremely commonplace with few restrictions on interbreeding; as individuals with an odd number of chromosome sets are usually sterile, depending on the actual number of chromosome sets present, this results in the odd situation where some individuals of the same evolutionary unit can interbreed with certain others and some cannot, with all populations being eventually linked as to form a common gene pool. The classification of species has been profoundly affected by technological advances that have allowed researchers to determine relatedness based on molecular markers, starting with the comparatively crude blood plasma precipitation assays in the mid-20th century and coming into full swing with Charles Sibley's ground-breaking DNA-DNA hybridisation studies in the 1970s. The results of the technique caused revolutionary changes in the higher taxonomic categories (such as phyla and classes), resulting in the reordering of many branches of the phylogenetic tree (see also: molecular phylogeny). For taxonomic categories below genera, the results have been mixed so far; the pace of evolutionary change on the molecular level is rather slow, yielding clear differences only after considerable periods of reproductive separation. Instances of hybridization can result in misleading molecular data, the Pomarine Skua - Great Skua phenomenon being a famous example. Turtles have been determined to evolve with just one-eighth of the speed of other reptiles on the molecular level, and the rate of molecular evolution in albatrosses is half of what is found in the rather closely related storm-petrels. The hybridization technique is however no longer considered a good technique and more reliable computational techniques for sequence comparison are now used for. Molecular taxonomy does not directly measure the evolutionary processes, but rather the overall change brought upon by these processes. The processes that lead to the generation and maintenance of variation such as mutation, crossover and selection are not uniform (see also molecular clock). DNA is only extremely rarely a direct target of natural selection rather than changes in the DNA sequence enduring over generations being a result of the latter; for example, silent transition-transversion combinations would alter the melting point of the DNA sequence, but not the sequence of the encoded proteins and thus are a possible example where, for example in microorganisms, a mutation confers a change in fitness all by itself.

External links

- http://users.rcn.com/jkimball.ma.ultranet/BiologyPages/S/Speciation.html
- [http://www.sciencedaily.com/releases/2003/12/031231082553.htm 2003-12-31, ScienceDaily: Working On The 'Porsche Of Its Time': New Model For Species Determination Offered] Quote: "...two species of dinosaur that are members of the same genera varied from each other by just 2.2 percent. Translation of the percentage into an actual number results in an average of just three skeletal differences out of the total 338 bones in the body. Amazingly, 58 percent of these differences occurred in the skull alone. "This is a lot less variation than I'd expected", said Novak..."
- [http://www.sciencedaily.com/releases/2003/08/030808081854.htm 2003-08-08, ScienceDaily: Cross-species Mating May Be Evolutionarily Important And Lead To Rapid Change, Say Indiana University Researchers] Quote: "...the sudden mixing of closely related species may occasionally provide the energy to impel rapid evolutionary change..."
- [http://www.sciencedaily.com/releases/2004/01/040109064407.htm 2004-01-09 ScienceDaily: Mayo Researchers Observe Genetic Fusion Of Human, Animal Cells; May Help Explain Origin Of AIDS] Quote: "...The researchers have discovered conditions in which pig cells and human cells can fuse together in the body to yield hybrid cells that contain genetic material from both species... "What we found was completely unexpected", says Jeffrey Platt, M.D."
- [http://www.sciencedaily.com/releases/2000/09/000913211733.htm 2000-09-18, ScienceDaily: Scientists Unravel Ancient Evolutionary History Of Photosynthesis] Quote: "...gene-swapping was common among ancient bacteria early in evolution..."
- [http://plato.stanford.edu/entries/species/ Stanford Encyclopedia of Philosophy entry]
- [http://www.barcodinglife.org/ Barcoding of species] rank22 rank22 ms:Spesies ja:種 (生物) th:สปีชีส์

Acanthocephala

:For the plant genus, see Parodia.
Archiacanthocephala

Palaeacanthocephala

Eoacanthocephala The Acanthocephala (gr. Acanthus - thorn Kephale - head) is a phylum of parasitic worms, characterised by the presence of an evertable proboscis, armed with spines, which it uses to pierce and hold the gut wall of its host. Acanthocephalans typically have complex life cycles, involving a number of hosts, including invertebrates, fishes, amphibians, birds, and mammals. About 850 species have been described.

Morphological Characteristics

There are several morphological characteristics that distinguish acanthocephalans from other phyla of parasitic worms.

Digestion

Acanthocephalans lack a mouth or alimentary canal. This is a feature they share with the cestoda (tapeworms), although the two groups are not related. Adult stages live in the intestines of their host and uptake nutrients which have been digested by the host, directly, through their body surface.

Proboscis

The most notable feature of the acanthocephala is the presence of an anterior, protrudible proboscis that is usually covered with spiny hooks (hence the common name). The proboscis bears rings of recurved hooks arranged in horizontal rows, and it is by means of these hooks that the animal attaches itself to the tissues of its host. The hooks may be of two or three shapes, usually, longer, more slender hooks are arranged along the length of the proboscis, with several rows of more sturdy, shorter basal hooks around the base of the proboscis. The proboscis is used to pierce the gut wall of the final host, and hold the parasite fast while it completes its life cycle. Like the body, the proboscis is hollow, and its cavity is separated from the body cavity by a septum or proboscis sheath. Traversing the cavity of the proboscis are muscle-strands inserted into the tip of the proboscis at one end and into the septum at the other. Their contraction causes the proboscis to be invaginated into its cavity. The whole proboscis apparatus can also be, at least partially, withdrawn into the body cavity, and this is effected by two retractor muscles which run from the posterior aspect of the septum to the body wall. muscle

Phylogenetic Relationships

Acanthocephalans are highly adapted to a parasitic mode of life, and have lost many organs and structures through evolutionary processes. This makes determining relationships with other higher taxa through morphological comparison problematic. Phylogenetic analysis of the 18S ribosomal gene has revealed that the Acanthocephala are most closely related to the rotifers, or may even belong in that phylum. The two are included among the Platyzoa.

Size

The size of the animals varies greatly, from forms a few millimetres in length to Gigantorhynchus gigas, which measures from 100 to 650 mm.

Skin

The body surface of the acanthocephala is peculiar. Externally, the skin has a thin cuticle covering the epidermis, which consists of a syncytium with no cell walls. The syncytium is traversed by a series of branching tubules containing fluid and is controlled by a few wandering, amoeboid nuclei. Inside the syncytium is a irregular layer of circular muscle fibres, and within this again some rather scattered longitudinal fibres; there is no endothelium. In their micro-structure the muscular fibres resemble those of nematodes. Except for the absence of the longitudinal fibres the skin of the proboscis resembles that of the body, but the fluid-containing tubules of the proboscis are shut off from those of the body. The canals of the proboscis open into a circular vessel which runs round its base. From the circular canal two sac-like projections called the lemnisci run into the cavity of the body, alongside the proboscis cavity. Each consists of a prolongation of the syncytial material of the proboscis skin, penetrated by canals and sheathed with a muscular coat. They seem to act as reservoirs into which the fluid which is used to keep the proboscis "erect" can withdraw when it is retracted, and from which the fluid can be driven out when it is wished to expand the proboscis.

Nervous System

The central ganglion of the nervous system lies behind the proboscis sheath or septum. It innervates the proboscis and projects two stout trunks posteriorly which supply the body. Each of these trunks is surrounded by muscles, and this nerve-muscle complex is called a retinaculum. In the male at least there is also a genital ganglion. Some scattered papillae may possibly be sense-organs.

Sex

The Acanthocephala are dioecious. There is a structure called the genital ligament which runs from the posterior end of the proboscis sheath to the posterior end of the body. In the male, two testes lie on either side of this. Each opens in a vas deferens which bears three diverticula or vesiculae seminales. The male also possesses three pairs of cement glands, found behind the testes, which pour their secretions through a duct into the vasa deferentia. These unite and end in a penis which opens posteriorly. In the female, the ovaries are found, like the testes, as rounded bodies along the ligament. From these masses of ova dehisce into the body cavity and float in its fluid. Here the eggs are fertilized and segment so that the young embryos are formed within their mother's body. The embryos escape into the uterus through the uterine bell, a funnel like opening continuous with the uterus. At the junction of the bell and the uterus there is a second small opening situated dorsally. The bell "swallows" the matured embryos and passes them on into the uterus, and from there, out of the body via the oviduct. Should the bell swallow any of the ova, or even one of the younger embryos, these are passed back into the body cavity through the second, dorsal, opening. The embryo passes from the body of the female into the alimentary canal of the host and leaves this with the feces.

Other Features

A curious feature shared by both larva and adult is the large size of many of the cells, e.g. the nerve cells and cells forming the uterine bell. Polyploidy is common, with up to 343n having been recorded in some species. The acanthocephalans lack an excretory system, although some species have been shown to possess flame cells (protonephridia).

Life Cycles

General Patterns

Acanthocephalans have complex life cycles, involving a number of hosts, for both developmental and resting stages. Complete life cycles have been worked out for only 25 species. Having been expelled by the female, the acanthocephalan embryo is released along with the feces of the host. For development to occur, the embryo needs to be ingested by an invertebrate, almost always a crustacean (there is one known life cycle which uses a mollusc as a first intermediate host). Inside the intermediate host, the acanthocephalan penetrates the gut wall, moves into the body cavity, encysts, and begins transformation into the infective cystacanth stage. This form has all the organs of the adult save the reproductive ones. The parasite is released when the first intermediate host is ingested. This can be by a suitable final host, in which case the cystacanth develops into a mature adult, or by a paratenic host, which is not suitable for further development, in which the parasite again forms a cyst. When consumed by a suitable final host, the cycstacant excysts, everts its proboscis and pierces the gut wall. It then feeds, grows and develops its sexual organs. Adult worms then mate. The male uses the excretions of its cement glands to plug the vagina of the female, preventing subsequent matings from occurring. Embryos develop inside the female, and the life cycle repeats.

An example - Polymorphus spp.

vagina Polymorphus spp. are parasites of seabirds, particularly the Eider Duck (Somateria mollissima). Heavy infections of up to 750 parasites per bird are common, causing ulceration to the gut, disease and seasonal mortality. Recent research has suggested that there is no evidence of pathogenicity of Polymorphus spp. to intermediate crab hosts. The cystacanth stage is long lived and probably remains infective throughout the life of the crab. The life cycle of Polymorphus spp. normally occurs between sea ducks (e.g. eiders and scoters) and small crabs. Infections found in commercial-sized lobsters in Canada were probably acquired from crabs that form an important dietary item of lobsters. Cystacanths occurring in lobsters can cause economic loss to fishermen. There are no known methods of prevention or control.

External links

- [http://www.fishdisease.net Fishdisease.net] Category:Animals Category:Parasitology category:Parasites

Chaetognatha

- Archisagittoidea
- Sagittoidea Chaetognatha is a phylum of predatory marine worms that are a major component of plankton worldwide. They show some preference for warmer waters. Chaetognaths are transparent and are torpedo shaped sometimes with arrowhead like opaque structures in their heads. They range in size from 3mm to 12cm. The common term for the phylum is Arrow Worms. There are about 100 modern species assigned to 15 genera. Despite the limited diversity, the number of individuals is staggering. Chatognaths are transparent or translucent and are covered by a cuticle. They have fins and a pair of hooked, chitinous, grasping spines on each side of their heads that are used in hunting. The spines are covered with a hood when swimming. They have a distinct head, trunk and tail. All species are hermaphroditic, carrying both eggs and sperm. A few species are known to use neurotoxins to subdue prey. Chaetognaths are traditionally classed as deuterostomes by embryologists. Molecular phylogenists, however, consider them to be protostomes. They have some developmental similarities to nematodes. Although they have a mouth with one or two rows of tiny teeth, compound eyes, and a nervous system, they have no respiratory, circulatory, or excremental systems. Materials are moved about the body cavity by cilia. Waste materials are simply excreted through the