Earthworm is the usual name for the largest members of 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 posterior to the female pores, even though the internal male segments are anterior to the female. Theoretical cladistic studies have placed them instead in the suborder Lumbricina of the order Haplotaxida, but this may again soon change. Folk names for the earthworm include "dew-worm", "rainworm", "night crawler" and "angleworm" (due to its use as fishing bait).

Earthworms are also called megadriles (or big worms), as opposed to the microdriles (or small worms) in the families Tubificidae, Lumbriculidae, and Enchytraeidae, among others. 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.

Anatomy

The basic morphology of an earthworm is a tube, the digestive system, within a tube, the muscular outer body. The body is annular, formed of segments that are most specialized in the anterior. Earthworms have a simple 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" (aortic arches) 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. Earthworms eat in a unique way: their mouth cavity connects directly into the digestive tract without any intermediate processes. Most earthworms are decomposers feeding on undecayed leaf and other plant matter, others are more geophagous.

Reproduction

Earthworm reproduction

Regeneration

Earthworms have the facility to replace or replicate lost segments, but this ability varies between species and depends on the extent of the damage. Stephenson (1930) devoted a chapter of his monograph to this topic, while G.E. Gates spent 20 years studying regeneration in a variety of species, but ?because little interest was shown?, Gates (1972) only published a few of his findings that, nevertheless, show it is theoretically possible to grow two whole worms from a bisected specimen in certain species. Gates?s reports included:

• Eisenia fetida (Savigny, 1826) with head regeneration, in an anterior direction, possible at each intersegmental level back to and including 23/24, while tails were regenerated at any levels behind 20/21 http://www.biolbull.org/cgi/reprint/96/2/129.pdf.
• Lumbricus terrestris Linneus, 1758 replacing anterior segments from as far back as 13/14 and 16/17 but tail regeneration was never found.
• Perionyx excavatus Perrier, 1872 readily regenerated lost parts of the body, in an anterior direction from as far back as 17/18, and in a posterior direction as far forward as 20/21.
• Lampito mauritii Kinberg, 1867 with regeneration in anterior direction at all levels back to 25/26 and tail regeneration from 30/31; head regeneration was sometimes believed to be caused by internal amputation resulting from Sarcophaga sp. larval infestation.
• Criodrilus lacuum Hoffmeister, 1845 also has prodigious regenerative capacity with ?head? regeneration from as far back as 40/41.http://links.jstor.org/sici?sici=0003-0031(195310)50%3A2%3C414%3AORCOEO%3E2.0.CO%3B2-Y

An unidentified Tasmanian earthworm shown growing a second head is reported here: http://www.qvmag.tas.gov.au/zoology/invertebrata/printarchive/printtext/inv20aitems.html#20blakemore.

Behavior

Rainstorms

Earthworms are seen on the surface after large rain storms flood the soil because, despite needing a moist environment to allow the diffusion of gases across their skin membrane, where the soil becomes saturated they begin to drown. To protect themselves they escape to the surface but if the ground is un-naturally hard they may become stranded and die from exposure. This is why they are seen in places like driveways after a storm. However, this theory is not applicable to certain earthworm species that can survive immersion for several days in oxygenated water.

An earthworm being eaten by an American Robin.

Another theory is that 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. Since the relative humidity is higher during and after rain, they do not become dehydrated. This is a dangerous activity in the daytime, since earthworms die quickly when exposed to direct sunlight with its strong UV content, and are more vulnerable to predators such as birds.

A further theory is that, as there are many other organisms in the ground as well, and their respiration increases carbon dioxide, this gas may dissolve into the rainwater to form carbonic acid. As the soil becomes too acidic for the worms, they seek a more neutral environment on the surface.

Locomotion and importance to soil

Close up of an earthworm in garden soil

Benefits

The major benefits of earthworm activities to soil fertility can be summarized as:

• Biological. In many soils, earthworms play a major role in converting large pieces of organic matter (e.g. dead leaves) into rich humus, and thus improving soil fertility. This is achieved by the worm's actions of pulling down below any organic matter deposited on the dried dirt, such as leaf fall or manure, 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.

Faeces in form of casts

• 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 (1.25mm) across?into its gizzard 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, minerals and plant nutrients are 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)" ^[2]. 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.

See Bioturbation.

The earthworm's existence cannot be taken 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), 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 invasive species

North America

A total of approximately 182 earthworm taxa in 12 families are reported from America north of Mexico, i.e., USA & Canada, of which 60 (ca. 33%) are exotic/introduced. Only two genera of Lumbricid earthworms are indigenous to North America while introduced genera have spread to areas where earthworms did not formerly exist, especially in the north where forest development relies on a large amount of undecayed leaf matter. When worms decompose that leaf layer, the ecology may shift making the habitat unsurvivable for certain species of trees, ferns and wildflowers. Currently there is no economically feasible method for controlling invasive earthworms in forests. Earthworms normally spread slowly, but can be quickly introduced by human activities such as construction earthmoving, or by fishermen releasing bait, or by plantings from other areas. ^[3]

Australia

Australia has 650 known species of native earthworm that survive in both rich and in nutrient-poor conditions where they may be sensitive to changes in the environment. Introduced species are commonly found in agricultural environments along with persistent natives. Most of the 75 or so exotics have been accidentally introduced into Australia. The total species numbers are predicted to exceed 2,000. ^[4]

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 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 acidic humus. Aporrectodea limicola and Sparganophilus and several others are found in mud in streams. Some species are arborial. Even in the soil species, there are special habitats, such as soils derived from serpentine which have an earthworm fauna of their own.

Ecology

Earthworms are classified into three main ecophysiological categories: (1) leaf litter/compost dwelling worms (epigeic) e.g. Eisenia fetida; (2) topsoil or subsoil dwelling worms (endogeics); and (3) worms that construct permanent deep burrows through which they visit the surface to obtain plant material for food, such as leaves (anecic), e.g. Lumbricus terrestris. ^[5]

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 acidic soil. However, Lumbricus terrestris are still present in a pH of 5.4 and Dendrobaena octaedra at a 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 go into diapause, and remain in diapause the longest time at a 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 both European Robins and American Robins. Some snakes feed on them and mammals such as bears, foxes, hedgehogs and moles eat many earthworms as well. Earthworms are also eaten by many invertebrates such as ground beetles and other beetles, snails, slugs. Earthworms have many internal parasites including Protozoa, Platyhelminthes, Nematodes. They are found in many parts of earthworms' bodies such as blood, seminal vesicles, coelom, intestine, or in the cocoons.

The application of chemical fertilizers, sprays and dusts can have a disastrous effect on earthworm populations. Nitrogenous fertilizers 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 completely wiped out the giant Gippsland earthworm.

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. Adding organic matter, preferably as a surface mulch, on a regular basis will provide them with their food and nutrient requirements, and 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 the UK has no natural predator itself. At present sightings of the New Zealand flatworm 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, which is 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 fetida (or its close relative Eisenia andrei) or the Brandling worm, also known as the Tiger worm or Red Wiggler, and are distinct from soil-dwelling earthworms.

Earthworms are sold all over the world. The earthworm market is sizable. 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."

Earthworms are also sometimes sold as food for human consumption. Noke is a culinary term used by the M?ori of New Zealand to refer to earthworms which are considered delicacies.

A report on biodiversity published by the Irish Government in May 2008 estimated the activities of the earthworm to be worth a minimum of ?723 millon per annum to Irish agriculture.

Taxonomy and main geographic origins of earthworms

Main families :

• Lumbricidae : Temperate Northern Hemisphere from Vancouver Island, Canada to Japan, mostly Eurasia.
• Hormogastridae : Europe.
• Sparganophilidae : North America.
• Almidae : Africa, South America.
• Ocnerodrilidae : Central and South America, Africa.
• Acanthodrilidae : Africa, midland and southeastern North America, Central and South America, Australia and Oceania.
• Octochaetidae : Central/South America, western Africa, India, New Zealand, Australia.
• Exxidae : Central America/Caribbean.
• Megascolecidae : South East Asia, Australasia and Oceania, northwestern North America.
• Glossoscolecidae : Central and northern South America.
• Eudrilidae : Tropical Africa.

References

1. ↑ Charles Darwin, The formation of vegetable mould through the action of worms, with observations on their habits. Found at Project Gutenberg Etext Formation of Vegetable Mould, by Darwin
2. ↑ Mollison, Bill, Permaculture- A Designer's Manual, Tagari Press, 1988
3. ↑ Cosmopolitan Earthworms http://bio-eco.eis.ynu.ac.jp/eng/database/earthworm/COSMO2.doc
4. ↑ Blakemore, Rob, ''Diversity of exotic earthworms in Australia - a status report.'' Transactions of the Royal Zoological Society of New South Wales, 1999
5. ↑ Earthworms: Renewers of Agroecosystems (SA Fall, 1990 (v3n1))

Further reading

Edwards, Clive A. (Ed.) Earthworm Ecology. Boca Raton: CRC Press, 2004. Second revised edition. ISBN 084931819X

Lee, Keneth E. Earthworms: Their Ecology and Relationships with Soils and Land Use. Academic Press. Sydney, 1985. ISBN 0-12-440860-5

Stewart, Amy. The Earth Moved: On the Remarkable Achievements of Earthworms. Chapel Hill, N.C.: Algonquin Books, 2004. ISBN 1-56512-337-9

See also

• Drilosphere
• Lumbricidae
• Megascolecidae
• Sparganophilidae
• Soil life
• Gilbert White
• Noke
• Terra preta and its reference to peregrine earthworm Pontoscolex corethrurus

External links

• Earthworm Digest A great informational website and magazine
• Infography about Earthworms
• How to Make a Worm Farm Good for Composting and Fishing
• Kids Discovery
• Earthworm Information (UC Davis)
• earthworm on Encyclopedia.com
• Biology of the Night Crawler (Lumbricus terrestris)
• WormWatch - Field guide to earthworms
• New Zealand flatworm page (UK Govt.)
• New Zealand flatworm page (HDRA)
• Worm Watch Canadian worm awareness and appreciation site, with detailed worm anatomy.
• Minnesota Invasive Earthworms Minnesota DNR information on the negative impacts of earthworms
• Lumbricidae keys and Dichogastrid checklist
• web-book: Earthworms of Hungary
• Opuscula Zoologica Budapest, online papers on earthworm taxonomy
• A Series of Searchable Texts on Earthworm Biodiversity, Ecology and Systematics from Various Regions of the World
• ''Earthworms as pests and otherwise'' hosted by the UNT Government Documents Department
• Third International Oligochaete Taxonomy Meeting, Platres, Cyprus, April 2nd to 6th, 2007
• Short comments on the Levant earthworm fauna

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