Earthworm

An earthworm is a terrestrial invertebrate

that belongs to the phylum Annelida. They
exhibit a tube-within-a-tube body plan; they
are externally segmented with
corresponding internal segmentation; and
they usually have setae on all segments.[1]
They occur worldwide where soil, water,
and temperature allow.[2]
 Earthworm

An unidentified earthworm species with a well-

developed clitellum

Scientific classification

Kingdom: Animalia

Phylum: Annelida

Class: Clitellata

Order: Opisthopora
Suborder: Lumbricina

Earthworms are commonly found in soil,

eating a wide variety of organic matter.[3]
This organic matter includes plant matter,
living protozoa, rotifers, nematodes,
bacteria, fungi, and other
microorganisms.[4] An earthworm's
digestive system runs the length of its
body.[5]

An earthworm respires (breathes) through

its skin. It has a double transport system
made of coelomic fluid that moves within
the fluid-filled coelom and a simple, closed
circulatory system.
It has a central and peripheral nervous
system. Its central nervous system
consists of two ganglia above the mouth,
one on either side, connected to a nerve
running along its length to motor neurons
and sensory cells in each segment. Large
numbers of chemoreceptors concentrate
near its mouth.

Circumferential and longitudinal muscles

edging each segment let the worm move.
Similar sets of muscles line the gut, and
their actions move digesting food toward
the worm's anus.[6]
Earthworms are hermaphrodites: each
carries male and female reproductive
organs. When mating, two individual
earthworms will exchange sperm and
fertilize each other's eggs. Each individual
has both male and female genital pores.
As invertebrates, they lack a true skeleton,
but they maintain their structure with fluid-
filled coelom chambers that function as a
hydrostatic skeleton.

"Earthworm" is the common name for the

largest members of Oligochaeta (which is
a class or subclass depending on the
author). In classical systems, they were in
the order Opisthopora since the male
pores opened posterior to the female
pores, although the internal male
segments are anterior to the female.
Theoretical cladistic studies have placed
them in the suborder Lumbricina of the
order Haplotaxida, but this may soon
change. Folk names for the earthworm
include "dew-worm", "rainworm",
"nightcrawler", and "angleworm" (from its
use as fishing bait).

Larger terrestrial earthworms are also

called megadriles (translates to "big
worms") as opposed to the microdriles
("small worms") in the semiaquatic
families Tubificidae, Lumbricidae, and
Enchytraeidae. The megadriles are
characterized by a distinct clitellum (more
extensive than that of microdriles) and a
vascular system with true capillaries.[7]

Anatomy

Form and function

Earthworm head
Depending on the species, an adult
earthworm can be from 10 mm (0.39 in)
long and 1 mm (0.039 in) wide to 3 m
(9.8 ft) long and over 25 mm (0.98 in)
wide, but the typical Lumbricus terrestris
grows to about 360 mm (14 in) long.[8]
Probably the longest worm on confirmed
records is Amynthas mekongianus that
extends up to 3 m (10 ft) [9] in the mud
along the banks of the 4,350 km (2,703 mi)
Mekong River in Southeast Asia.

From front to back, the basic shape of the

earthworm is a cylindrical tube-in-a-tube,
divided into a series of segments (called
metameres) that compartmentalize the
body. Furrows are generally[10] externally
visible on the body demarking the
segments; dorsal pores and
nephridiopores exude a fluid that moistens
and protects the worm's surface, allowing
it to breathe. Except for the mouth and
anal segments, each segment carries
bristlelike hairs called lateral setae[11] used
to anchor parts of the body during
movement;[12] species may have four pairs
of setae on each segment or more than
eight sometimes forming a complete
circle of setae per segment.[11] Special
ventral setae are used to anchor mating
earthworms by their penetration into the
bodies of their mates.[13]
Generally, within a species, the number of
segments found is consistent across
specimens, and individuals are born with
the number of segments they will have
throughout their lives. The first body
segment (segment number 1) features
both the earthworm's mouth and,
overhanging the mouth, a fleshy lobe
called the prostomium, which seals the
entrance when the worm is at rest, but is
also used to feel and chemically sense the
worm's surroundings. Some species of
earthworm can even use the prehensile
prostomium to grab and drag items such
as grasses and leaves into their burrow.
An adult earthworm develops a belt-
shaped glandular swelling, called the
clitellum, which covers several segments
toward the front part of the animal. This is
part of the reproductive system and
produces egg capsules. The posterior is
most commonly cylindrical like the rest of
the body, but depending on the species, it
may also be quadrangular, octagonal,
trapezoidal, or flattened. The last segment
is called the periproct; the earthworm's
anus, a short vertical slit, is found on this
segment.[11]
The exterior of an individual segment is a
thin cuticle over the skin, commonly
pigmented red to brown, which has
specialized cells that secrete mucus over
the cuticle to keep the body moist and
ease movement through the soil. Under
the skin is a layer of nerve tissue, and two
layers of muscles—a thin outer layer of
circular muscle, and a much thicker inner
layer of longitudinal muscle.[14] Interior to
the muscle layer is a fluid-filled chamber
called a coelom[15] that by its
pressurization provides structure to the
worm's boneless body. The segments are
separated from each other by septa (the
plural of "septum")[16] which are perforated
transverse walls, allowing the coelomic
fluid to pass between segments.[17] A pair
of structures called nephrostomes are
located at the back of each septum; a
nephric tubule leads from each
nephrostome through the septum and into
the following segment. This tubule then
leads to the main body fluid filtering organ,
the nephridium or metanephridium, which
removes metabolic waste from the
coelomic fluid and expels it through pores
called nephridiopores on the worm's sides;
usually, two nephridia (sometimes more)
are found in most segments.[18] At the
centre of a worm is the digestive tract,
which runs straight through from mouth to
anus without coiling, and is flanked above
and below by blood vessels (the dorsal
blood vessel and the ventral blood vessel
as well as a subneural blood vessel) and
the ventral nerve cord, and is surrounded
in each segment by a pair of pallial blood
vessels that connect the dorsal to the
subneural blood vessels.

Many earthworms can eject coelomic fluid

through pores in the back in response to
stress; the Australian Didymogaster
sylvaticus (known as the "blue squirter
earthworm") can squirt fluid as high as
30 cm (12 in).[19][17]

Nervous system
Nervous system of the anterior end of an earthworm

Central nervous system

The CNS consists of a bilobed brain

(cerebral ganglia, or supra-pharyngeal
ganglion), sub-pharyngeal ganglia, circum-
pharyngeal connectives and a ventral
nerve cord.

Earthworms' brains consist of a pair of

pear-shaped cerebral ganglia. These are
located in the dorsal side of the alimentary
canal in the third segment, in a groove
between the buccal cavity and pharynx.

A pair of circum-pharyngeal connectives

from the brain encircle the pharynx and
then connect with a pair of sub-pharyngeal
ganglia located below the pharynx in the
fourth segment. This arrangement means
the brain, sub-pharyngeal ganglia and the
circum-pharyngeal connectives form a
nerve ring around the pharynx.

The ventral nerve cord (formed by nerve

cells and nerve fibers) begins at the sub-
pharyngeal ganglia and extends below the
alimentary canal to the most posterior
body segment. The ventral nerve cord has
a swelling, or ganglion, in each segment,
i.e. a segmental ganglion, which occurs
from the fifth to the last segment of the
body. There are also three giant axons, one
medial giant axon (MGA) and two lateral
giant axons (LGAs) on the mid-dorsal side
of the ventral nerve cord. The MGA is
0.07 mm in diameter and transmits in an
anterior-posterior direction at a rate of
32.2 m/s. The LGAs are slightly narrower
at 0.05 mm in diameter and transmit in a
posterior-anterior direction at 12.6 m/s.
The two LGAs are connected at regular
intervals along the body and are therefore
considered one giant axon.[20][21]

Peripheral nervous system

Eight to ten nerves arise from the

cerebral ganglia to supply the
 prostomium, buccal chamber and
pharynx.

Three pairs of nerves arise from the

subpharyangeal ganglia to supply the
second, third and fourth segment.

Three pairs of nerves extend from each

segmental ganglion to supply various
structures of the segment.

The sympathetic nervous system consists

of nerve plexuses in the epidermis and
alimentary canal. (A plexus is a web of
connected nerve cells.) The nerves that
run along the body wall pass between the
outer circular and inner longitudinal
muscle layers of the wall. They give off
branches that form the intermuscular
plexus and the subepidermal plexus.
These nerves connect with the
cricopharyngeal connective.

Movement

An earthworm crawling over asphalt.

On the surface, crawling speed varies both

within and among individuals. Earthworms
crawl faster primarily by taking longer
"strides" and a greater frequency of
strides. Larger Lumbricus terrestris worms
crawl at a greater absolute speed than
smaller worms. They achieve this by
taking slightly longer strides but with
slightly lower stride frequencies.[22]

Touching an earthworm, which causes a

"pressure" response as well as (often) a
response to the dehydrating quality of the
salt on human skin (toxic to earthworms),
stimulates the subepidermal nerve plexus
which connects to the intermuscular
plexus and causes the longitudinal
muscles to contract. This causes the
writhing movements observed when a
human picks up an earthworm. This
behaviour is a reflex and does not require
the CNS; it occurs even if the nerve cord is
removed. Each segment of the earthworm
has its own nerve plexus. The plexus of
one segment is not connected directly to
that of adjacent segments. The nerve cord
is required to connect the nervous
systems of the segments.[23]

The giant axons carry the fastest signals

along the nerve cord. These are
emergency signals that initiate reflex
escape behaviours. The larger dorsal giant
axon conducts signals the fastest, from
the rear to the front of the animal. If the
rear of the worm is touched, a signal is
rapidly sent forwards causing the
longitudinal muscles in each segment to
contract. This causes the worm to shorten
very quickly as an attempt to escape from
a predator or other potential threat. The
two medial giant axons connect with each
other and send signals from the front to
the rear. Stimulation of these causes the
earthworm to very quickly retreat (perhaps
contracting into its burrow to escape a
bird).

The presence of a nervous system is

essential for an animal to be able to
experience nociception or pain. However,
other physiological capacities are also
required such as opioid sensitivity and
central modulation of responses by
analgesics.[24] Enkephalin and α-
endorphin-like substances have been
found in earthworms. Injections of
naloxone (an opioid antagonist) inhibit the
escape responses of earthworms. This
indicates that opioid substances play a
role in sensory modulation, similar to that
found in many vertebrates.[25]

Sensory reception

Photosensitivity
Earthworms do not have eyes (although
some worms do); however, they do have
specialized photosensitive cells called
"light cells of Hess". These photoreceptor
cells have a central intracellular cavity
(phaosome) filled with microvilli. As well
as the microvilli, there are several sensory
cilia in the phaosome which are
structurally independent of the
microvilli.[26] The photoreceptors are
distributed in most parts of the epidermis
but are more concentrated on the back
and sides of the worm. A relatively small
number occurs on the ventral surface of
the first segment. They are most
numerous in the prostomium and reduce
in density in the first three segments; they
are very few in number past the third
segment.[23]

Epidermal receptor (Sense organ)

These receptors are abundant and

distributed all over the epidermis. Each
receptor shows a slightly elevated cuticle
which covers a group of tall, slender and
columnar receptor cells. These cells bear
small hairlike processes at their outer
ends and their inner ends are connected
with nerve fibres. The epidermal receptors
are tactile in function. They are also
concerned with changes in temperature
and respond to chemical stimuli.
Earthworms are extremely sensitive to
touch and mechanical vibration.

Buccal receptor (Sense organ)

These receptors are located only in the

epithelium of the buccal chamber. These
receptors are gustatory and olfactory
(related to taste and smell). They also
respond to chemical stimuli.
(Chemoreceptor)

Digestive system

The gut of the earthworm is a straight tube

that extends from the worm's mouth to its
anus. It is differentiated into an alimentary
canal and associated glands which are
embedded in the wall of the alimentary
canal itself. The alimentary canal consists
of a mouth, buccal cavity (generally
running through the first one or two
segments of the earthworm), pharynx
(running generally about four segments in
length), esophagus, crop, gizzard (usually),
and intestine. [27]

Food enters at the mouth. The pharynx

acts as a suction pump; its muscular walls
draw in food. In the pharynx, the
pharyngeal glands secrete mucus. Food
moves into the esophagus, where calcium
(from the blood and ingested from
previous meals) is pumped in to maintain
proper blood calcium levels in the blood
and food pH. From there the food passes
into the crop and gizzard. In the gizzard,
strong muscular contractions grind the
food with the help of mineral particles
ingested along with the food. Once
through the gizzard, food continues
through the intestine for digestion. The
intestine secretes pepsin to digest
proteins, amylase to digest
polysaccharides, cellulase to digest
cellulose, and lipase to digest fats.[6]
Earthworms use, in addition to the
digestive proteins, a class of surface
active compounds called drilodefensins,
which help digest plant material.[28]
Instead of being coiled like a mammalian
intestine, in an earthworm's intestine a
large mid-dorsal, tongue-like fold is
present, called typhlosole which increases
surface area to increase nutrient
absorption by having many folds running
along its length. The intestine has its own
pair of muscle layers like the body, but in
reverse order—an inner circular layer within
an outer longitudinal layer.[29]

Circulatory system
Earthworms have a dual circulatory
system in which both the coelomic fluid
and a closed circulatory system carry the
food, waste, and respiratory gases. The
closed circulatory system has five main
blood vessels: the dorsal (top) vessel,
which runs above the digestive tract; the
ventral (bottom) vessel, which runs below
the digestive tract; the subneural vessel,
which runs below the ventral nerve cord;
and two lateroneural vessels on either side
of the nerve cord.[30]

The dorsal vessel is mainly a collecting

structure in the intestinal region. It
receives a pair commissural and dorsal
intestines in each segment. The ventral
vessel branches off to a pair of ventro-
tegumentaries and ventro-intestinals in
each segment. The subneural vessel also
gives out a pair of commissurals running
along the posterior surface of the septum.

The pumping action on the dorsal vessel

moves the blood forward, while the other
four longitudinal vessels carry the blood
rearward. In segments seven through
eleven, a pair of aortic arches ring the
coelom and acts as hearts, pumping the
blood to the ventral vessel that acts as the
aorta. The blood consists of ameboid cells
and haemoglobin dissolved in the plasma.
The second circulatory system derives
from the cells of the digestive system that
line the coelom. As the digestive cells
become full, they release non-living cells
of fat into the fluid-filled coelom, where
they float freely but can pass through the
walls separating each segment, moving
food to other parts and assist in wound
healing.[31]

Excretory system

The excretory system contains a pair of

nephridia in every segment, except for the
first three and the last ones.[32] The three
types of nephridia are: integumentary,
septal, and pharyngeal. The integumentary
nephridia lie attached to the inner side of
the body wall in all segments except the
first two. The septal nephridia are attached
to both sides of the septa behind the 15th
segment. The pharyngeal nephridia are
attached to the fourth, fifth and sixth
segments.[32] The waste in the coelom
fluid from a forward segment is drawn in
by the beating of cilia of the nephrostome.
From there it is carried through the septum
(wall) via a tube which forms a series of
loops entwined by blood capillaries that
also transfer waste into the tubule of the
nephrostome. The excretory wastes are
then finally discharged through a pore on
the worm's side.[33]

Respiration

Earthworms have no special respiratory

organs. Gases are exchanged through the
moist skin and capillaries, where the
oxygen is picked up by the haemoglobin
dissolved in the blood plasma and carbon
dioxide is released. Water, as well as salts,
can also be moved through the skin by
active transport.

Life and physiology

At birth, earthworms emerge small but
fully formed, lacking only their sex
structures which develop in about 60 to 90
days. They attain full size in about one
year. Scientists predict that the average
lifespan under field conditions is four to
eight years, while most garden varieties
live only one to two years.

Reproduction

Earthworm copulation
Earthworm cocoons from L. terrestris

An earthworm cocoon from L. rubellus

Several common earthworm species are

mostly parthenogenetic, meaning that
growth and development of embryos
happens without fertilization. Among
lumbricid earthworms, parthenogenesis
arose from sexual relatives many times.[34]
Parthenogenesis in some Aporrectodea
trapezoides lineages arose 6.4 to
1.1 million years ago from sexual
ancestors.[35] A few species exhibit
pseudogamous parthogenesis, meaning
that mating is necessary to stimulate
reproduction, even though no male genetic
material passes to the offspring.[36]

Earthworm mating occurs on the surface,

most often at night. Earthworms are
hermaphrodites; that is, they have both
male and female sexual organs. The
sexual organs are located in segments 9
to 15. Earthworms have one or two pairs
of testes contained within sacs. The two
or four pairs of seminal vesicles produce,
store and release the sperm via the male
pores. Ovaries and oviducts in segment 13
release eggs via female pores on segment
14, while sperm is expelled from segment
15. One or more pairs of spermathecae
are present in segments 9 and 10
(depending on the species) which are
internal sacs that receive and store sperm
from the other worm during copulation. As
a result, segment 15 of one worm exudes
sperm into segments 9 and 10 with its
storage vesicles of its mate. Some species
use external spermatophores for sperm
transfer.

In Hormogaster samnitica and

Hormogaster elisae transcriptome DNA
libraries were sequenced and two sex
pheromones, Attractin and Temptin, were
detected in all tissue samples of both
species.[37] Sex pheromones are probably
important in earthworms because they live
in an environment where chemical
signaling may play a crucial role in
attracting a partner and in facilitating
outcrossing. Outcrossing would provide
the benefit of masking the expression of
deleterious recessive mutations in
progeny[38] (see Complementation).

Copulation and reproduction are separate

processes in earthworms. The mating pair
overlap front ends ventrally and each
exchanges sperm with the other. The
clitellum becomes very reddish to pinkish
in colour. Sometime after copulation, long
after the worms have separated, the
clitellum (behind the spermathecae)
secretes material which forms a ring
around the worm. The worm then backs
out of the ring, and as it does so, it injects
its own eggs and the other worm's sperm
into it. Thus each worm becomes the
genetic father of some of their offspring
(due to its own sperm transferred to other
earthworm) and the genetic mother
(offsprings from its own egg cells) of the
rest. As the worm slips out of the ring, the
ends of the cocoon seal to form a vaguely
onion-shaped incubator (cocoon) in which
the embryonic worms develop. Hence
fertilization is external. The cocoon is then
deposited in the soil. After three weeks, 2
to 20 offspring hatch with an average of 4.
Development is direct i.e. without
formation of any larva.

Locomotion
Close up of an earthworm in garden soil

Earthworms travel underground by means

of waves of muscular contractions which
alternately shorten and lengthen the body
(peristalsis). The shortened part is
anchored to the surrounding soil by tiny
clawlike bristles (setae) set along its
segmented length. In all the body
segments except the first, last and
clitellum, there is a ring of S-shaped setae
embedded in the epidermal pit of each
segment (perichaetine). The whole
burrowing process is aided by the
secretion of lubricating mucus. As a result
of their movement through their lubricated
tunnels, worms can make gurgling noises
underground when disturbed. Earthworms
move through soil by expanding crevices
with force; when forces are measured
according to body weight, hatchlings can
push 500 times their own body weight
whereas large adults can push only 10
times their own body weight.[39]
Regeneration

Earthworms have the ability to regenerate

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) published only 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; thus two worms may grow from
one.[40]

Lumbricus terrestris (Linnaeus, 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.[41]
An unidentified Tasmanian earthworm
shown growing a replacement head has
been reported.[42]

Taxonomy and distribution

Within the world of taxonomy, the stable
'Classical System' of Michaelsen (1900)
and Stephenson (1930) was gradually
eroded by the controversy over how to
classify earthworms, such that Fender and
McKey-Fender (1990) went so far as to
say, "The family-level classification of the
megascolecid earthworms is in chaos."[43]
Over the years, many scientists have
developed their own classification
systems for earthworms, which led to
confusion, and these systems have been
and still continue to be revised and
updated. The classification system used
here which was developed by Blakemore
(2000), is a modern reversion to the
Classical System that is historically proven
and widely accepted.[44]

Categorization of a megadrile earthworm

into one of its taxonomic families under
suborders Lumbricina and Moniligastrida
is based on such features as the makeup
of the clitellum, the location and
disposition of the sex features (pores,
prostatic glands, etc.), number of gizzards,
and body shape.[44] Currently, over 6,000
species of terrestrial earthworms are
named, as provided in a species name
database,[45] but the number of synonyms
is unknown.

The families, with their known

distributions or origins:[44]

Acanthodrilidae

Ailoscolecidae – the Pyrenees and the

southeast USA

Almidae – tropical equatorial (South

America, Africa, Indo-Asia)

Benhamiinae – Ethiopian, Neotropical (a

possible subfamily of Octochaetidae)
Criodrilidae – southwestern Palaearctic:
Europe, Middle East, Russia and Siberia
to Pacific coast; Japan (Biwadrilus);
mainly aquatic

Diplocardiinae/-idae – Gondwanan or
Laurasian? (a subfamily of
Acanthodrilidae)

Enchytraeidae – cosmopolitan but

uncommon in tropics (usually classed
with Microdriles)

Eudrilidae – Tropical Africa south of the

Sahara

Exxidae – Neotropical: Central America

and the Caribbean
Glossoscolecidae – Neotropical: Central
and South America, Caribbean

Haplotaxidae – cosmopolitan
distribution (usually classed with
Microdriles)

Hormogastridae – Mediterranean

Kynotidae – Malagasian: Madagascar

Lumbricidae – Holarctic: North America,

Europe, Middle East, Central Asia to
Japan

Lutodrilidae – Louisiana the southeast

USA

Megascolecidae

Microchaetidae – Terrestrial in Africa

especially South African grasslands
 Moniligastridae – Oriental and Indian
subregion

Ocnerodrilidae – Neotropics, Africa;

India

Octochaetidae – Australasian, Indian,

Oriental, Ethiopian, Neotropical

Octochaetinae – Australasian, Indian,

Oriental (subfamily if Benhamiinae is
accepted)

Sparganophilidae – Nearctic,
Neotropical: North and Central America

Tumakidae – Colombia, South America

As an invasive species
From a total of around 7,000 species, only
about 150 species are widely distributed
around the world. These are the peregrine
or cosmopolitan earthworms.[46]

Ecology

Permanent vertical burrow

Earthworms are classified into three main

ecophysiological categories: (1) leaf litter-
or compost-dwelling worms that are
nonburrowing, live at the soil-litter
interface and eat decomposing organic
matter (epigeic) e.g. Eisenia fetida; (2)
topsoil- or subsoil-dwelling worms that
feed (on soil), burrow and cast within the
soil, creating horizontal burrows in upper
10–30 cm of soil (endogeic); and (3)
worms that construct permanent deep
vertical burrows which they use to visit the
surface to obtain plant material for food,
such as leaves (anecic, meaning "reaching
up"), e.g. Lumbricus terrestris.[47]

Earthworm populations depend on both

physical and chemical properties of the
soil, such as 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 favour neutral to slightly
acidic soils. Lumbricus terrestris is still
present in a pH of 5.4, Dendrobaena
octaedra at a pH of 4.3 and some
Megascolecidae are present in extremely
acidic humic soils. Soil pH may also
influence the numbers of worms that go
into diapause. The more acidic the soil, the
sooner worms go into diapause, and
remain in diapause the longest time at a
pH of 6.4.

Earthworms are preyed upon by many

species of birds (e.g. robins, starlings,
thrushes, gulls, crows), snakes, wood
turtles, mammals (e.g. bears, boars, foxes,
hedgehogs, pigs, moles[48]) and
invertebrates (e.g. ants,[49] flatworms,
ground beetles and other beetles, snails,
spiders, and slugs). Earthworms have
many internal parasites, including
protozoa, platyhelminthes, mites, and
nematodes; they can be found in the
worms' blood, seminal vesicles, coelom, or
intestine, or in their cocoons (e.g. the mite
Histiostoma murchiei is a parasite of
earthworm cocoons[50]).

Nitrogenous fertilizers tend to create

acidic conditions, which are fatal to the
worms, and dead specimens are often
found on the surface following the
application of substances such as DDT,
lime sulphur, and lead arsenate. In
Australia, changes in farming practices
such as the application of
superphosphates on pastures and a
switch from pastoral farming to arable
farming had a devastating effect on
populations of the giant Gippsland
earthworm, leading to their classification
as a protected species. Globally, certain
earthworms populations have been
devastated by deviation from organic
production and the spraying of synthetic
fertilizers and biocides, with at least three
species now listed as extinct but many
more endangered.[51] This earthworm
activity aerates and mixes the soil, and is
conducive to mineralization of nutrients
and their uptake by vegetation. Certain
species of earthworm come to the surface
and graze on the higher concentrations of
organic matter present there, mixing it with
the mineral soil. Because a high level of
organic matter mixing is associated with
soil fertility, an abundance of earthworms
is generally considered beneficial by
farmers and gardeners.[52][53] As long ago
as 1881 Charles Darwin wrote: "It may be
doubted whether there are many other
animals which have played so important a
part in the history of the world, as have
these lowly organized creatures."[54]

Devil's coach horse beetle preying on Lumbricus sp.

Also, while, as the name suggests, the

main habitat of earthworms is in soil, they
are not restricted to this habitat. 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. Aporrectodea limicola,
Sparganophilus spp., and several others
are found in mud in streams. Some
species are arboreal, some aquatic and
some euryhaline (salt-water tolerant) and
littoral (living on the sea-shore, e.g.
Pontodrilus litoralis).[55] Even in the soil
species, special habitats, such as soils
derived from serpentine, have an
earthworm fauna of their own.
Vermicomposting of organic "wastes" and
addition of this organic matter to the soil,
preferably as a surface mulch, will provide
several species of earthworms with their
food and nutrient requirements, and will
create the optimum conditions of
temperature and moisture that will
stimulate their activity.

Earthworms are environmental indicators

of soil health. Earthworms feed on the
decaying matter in the soil and analyzing
the contents of their digestive tracts gives
insight into the overall condition of the
soil. The earthworm gut accumulates
chemicals, including heavy metals such as
cadmium, mercury, zinc, and copper. The
population size of the earthworm indicates
the quality of the soil, as healthy soil would
contain a larger number of earthworms.[56]

Environmental impacts

The major benefits of earthworm activities

to soil fertility for agriculture can be
summarized as:

Biological: In many soils, earthworms

play a major role in the conversion of
large pieces of organic matter into rich
humus, thus improving soil fertility. This
is achieved by the worm's actions of
 pulling below the surface deposited
organic matter such as leaf fall or
manure, either for food or to plug its
burrow. Once in the burrow, the worm
will shred the leaf, partially digest it and
mingle it with the earth. Worm casts
(see bottom right) can contain 40
percent more humus than the top 9
inches (230 mm) of soil in which the
worm is living.[57]

Faeces in the form of casts

Chemical: In addition to dead organic
matter, the earthworm also ingests any
other soil particles that are small
enough—including sand grains up to
1
⁄20 inch (1.3 mm)—into its gizzard,
wherein those minute fragments of grit
grind everything into a fine paste which
is then digested in the intestine. When
the worm excretes this in the form of
casts, deposited on the surface or
deeper in the soil, minerals and plant
nutrients are changed to an accessible
form for plants to use. Investigations in
the United States show that fresh
earthworm casts are five times richer in
available nitrogen, seven times richer in
available phosphates, and 11 times
richer in available potassium than the
surrounding upper 6 inches (150 mm) of
soil. In conditions where humus is
plentiful, the weight of casts produced
may be greater than 4.5 kilograms
(9.9 lb) per worm per year.[57]

Physical: The earthworm's burrowing

creates a multitude of channels through
the soil and is of great value in
maintaining the soil structure, enabling
processes of aeration and drainage.[58]
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)".[59] Thus, the
earthworm not only creates passages
for air and water to traverse the soil, but
also modifies the vital organic
component that makes a soil healthy
(see Bioturbation). Earthworms promote
the formation of nutrient-rich casts
(globules of soil, stable in soil mucus)
that have high soil aggregation and soil
fertility and quality.[57] In podzol soils,
earthworms can obliterate the
characteristic banded appearance of the
soil profile by mixing the organic (LFH),
eluvial (E) and upper illuvial (B) horizons
to create a single dark Ap horizon.[60][61]
Earthworms accelerate nutrient cycling in
the soil-plant system through
fragmentation & mixing of plant debris –
physical grinding & chemical digestion.[57]
The earthworm's existence cannot be
taken for granted. Dr. W. E. Shewell-Cooper
observed "tremendous numerical
differences between adjacent gardens",
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.[62]

Darwin estimated that arable land

contains up to 53,000 per acre
(130,000/ha) (of worms, but more recent
research has produced figures suggesting
that even poor soil may support 250,000
per acre (620,000/ha), whilst rich fertile
farmland may have up to 1,750,000 per
acre (4,300,000/ha), meaning that the
weight of earthworms beneath a farmer's
soil could be greater than that of the
livestock upon its surface. Richly organic
topsoil populations of earthworms are
much higher – averaging 500 per square
metre (46/sq ft) and up to 400 g2 – such
that, for the 7 billion of us, each person
alive today has support of 7 million
earthworms.[63]
The ability to break down organic
materials and excrete concentrated
nutrients makes the earthworm a
functional contributor in restoration
projects. In response to ecosystem
disturbances, some sites have utilized
earthworms to prepare soil for the return
of native flora. Research from the Station
d'écologie Tropicale de Lamto asserts that
the earthworms positively influence the
rate of macroaggregate formation, an
important feature for soil structure.[64] The
stability of aggregates in response to
water was also found to be improved
when constructed by earthworms.[64]
Though not fully quantified yet,
greenhouse gas emissions of earthworms
likely contribute to global warming,
especially since top-dwelling earthworms
increase the speed of carbon cycles and
have been spread by humans into many
new geographies.[65]

Economic impact

Earthworms being raised at La Chonita Hacienda in Mexico

Various species of worms are used in
vermiculture, the practice of feeding
organic waste to earthworms to
decompose food waste. These are usually
Eisenia fetida (or its close relative Eisenia
andrei) or the brandling worm, commonly
known as the tiger worm or red wiggler.
They are distinct from soil-dwelling
earthworms. In the tropics, the African
nightcrawler Eudrilus eugeniae[66] and the
Indian blue Perionyx excavatus are used.

Earthworms are sold all over the world; the

market is sizable. According to Doug
Collicutt, "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."[67]

Earthworms provide an excellent source of

protein for fish, fowl and pigs but were
also used traditionally for human
consumption. Noke is a culinary term used
by the Māori of New Zealand, and refers to
earthworms which are considered
delicacies for their chiefs.

See also
Drilosphere, the part of the soil
influenced by earthworm secretions and
castings
 The Formation of Vegetable Mould
through the Action of Worms, an 1881
book by Charles Darwin

Soil life

Vermicompost

Vermifilter

Vermifilter toilet

Worm charming

References
1. Edwards, Clive A.; Lofty, J. R. (1977).
Biology of Earthworms. London: Chapman
& Hall. pp. preface. ISBN 0-412-14950-8.
2. Coleman, David C.; Crossley, D.A.; Hendrix,
Paul F. (2004). Fundamentals of Soil
Ecology. Amsterdam; Boston: Elsevier
Academic Press. p. 170. ISBN 0-12-
179726-0.

3. Bonkowski, Michael; Griffiths, Bryan S.;

Ritz, Karl (November 2000). "Food
preferences of earthworms for soil fungi".
Pedobiologia. 44–6 (6): 667.
doi:10.1078/S0031-4056(04)70080-3 (http
s://doi.org/10.1078%2FS0031-4056%280
4%2970080-3) .

4. Lofty, Clive A.; Lofty, J. R. (1977). Biology

of Earthworms. London: Chapman & Hall.
p. 80. ISBN 0-412-14950-8.
5. Edwards, Clive A.; Lofty, J.R. (1977).
Biology of Earthworms. London: Chapman
& Hall. p. 19. ISBN 0-412-14950-8.

6. Cleveland P. Hickman Jr.; Larry S. Roberts;

Frances M Hickman (1984). Integrated
Principles of Zoology (https://archive.org/
details/integratedprinci7ed0hick/page/3
44) (7th ed.). Times Mirror/Mosby
College Publishing. p. 344 (https://archive.
org/details/integratedprinci7ed0hick/pag
e/344) . ISBN 978-0-8016-2173-4.
7. Omodeo, Pietro (2000). "Evolution and
biogeography of megadriles (Annelida,
Clitellata)". Italian Journal of Zoology. 67–
2 (2): 179–201.
doi:10.1080/11250000009356313 (http
s://doi.org/10.1080%2F11250000009356
313) . S2CID 86293273 (https://api.sema
nticscholar.org/CorpusID:86293273) .

8. Blakemore 2012, p. xl.

9. Blakemore, R. J.; et al. (2007).

"Megascolex (Promegascolex)
mekongianus Cognetti, 1922 – its extent,
ecology and allocation to Amynthas
(Clitellata/Oligochaeta: Megascolecidae)"
(http://opuscula.elte.hu/PDF/Tomus36/3_
Blakemore.pdf) (PDF). Opuscula
Zoologica.
10. Edwards & Bohlen 1996, p. 11.

11. Sims & Gerard 1985, pp. 3–6.

12. Edwards & Bohlen 1996, p. 3.

13. Feldkamp, J. (1924). "Feldkamp, J.

"Untersuchungen über die
Geschlechtsmerkmale und die Begattung
der Regenwurmer Zoologische
Jahrbücher". Anatomie. 46: 609–632.

14. Edwards & Bohlen 1996, p. 8-9.

15. Edwards & Bohlen 1996, p. 1.

16. Sims & Gerard 1985, p. 8.

17. Edwards & Bohlen 1996, p. 12.

18. Edwards & Bohlen 1996, p. 6.

19. Myrmecofourmis (2018). "Meet the
squirting earthworm" (https://ghostarchiv
e.org/varchive/youtube/20211030/PuQEA
uM9De8) . Youtube. Archived from the
original (https://www.youtube.com/watc
h?v=PuQEAuM9De8) (video) on 2021-10-
30.

20. "Experiment: Comparing speeds of two

nerve fiber sizes" (https://backyardbrains.
com/experiments/comparingNerveSpeed
#prettyPhoto) . BackyardBrains. Retrieved
April 4, 2015.
21. Drewes, C. D.; Landa, K. B.; McFall, J. L.
(1978). "Giant nerve fibre activity in intact,
freely moving earthworms" (https://doi.or
g/10.1242%2Fjeb.72.1.217) . The Journal
of Experimental Biology. 72: 217–227.
doi:10.1242/jeb.72.1.217 (https://doi.org/
10.1242%2Fjeb.72.1.217) . PMID 624897
(https://pubmed.ncbi.nlm.nih.gov/62489
7) .
22. Quillin, K.J. (1999). "Kinematic scaling of
locomotion by hydrostatic animals:
ontogeny of peristaltic crawling by the
earthworm lumbricus terrestris". Journal
of Experimental Biology. 202 (6): 661–
674. doi:10.1242/jeb.202.6.661 (https://d
oi.org/10.1242%2Fjeb.202.6.661) .
PMID 10021320 (https://pubmed.ncbi.nl
m.nih.gov/10021320) .

23. "Earthworm-nervous system" (http://crono

don.com/BioTech/Earthworm_NS.html) .
Cronodon. Retrieved April 3, 2015.
24. Elwood, R.W. (2011). "Pain and suffering in
invertebrates?" (https://doi.org/10.1093%
2Filar.52.2.175) . ILAR Journal. 52 (2):
175–84. doi:10.1093/ilar.52.2.175 (http
s://doi.org/10.1093%2Filar.52.2.175) .
PMID 21709310 (https://pubmed.ncbi.nl
m.nih.gov/21709310) .

25. Smith, J.A. (1991). "A question of pain in

invertebrates" (https://doi.org/10.1093%2
Filar.33.1-2.25) . ILAR Journal. 33 (1–2):
25–31. doi:10.1093/ilar.33.1-2.25 (https://
doi.org/10.1093%2Filar.33.1-2.25) .
26. Röhlich, P.; Aros, B.; Virágh, Sz. (1970).
"Fine structure of photoreceptor cells in
the earthworm, Lumbricus Terrestris".
Zeitschrift für Zellforschung und
Mikroskopische Anatomie. 104 (3): 345–
357. doi:10.1007/BF00335687 (https://do
i.org/10.1007%2FBF00335687) .
PMID 4193489 (https://pubmed.ncbi.nlm.
nih.gov/4193489) . S2CID 22771585 (http
s://api.semanticscholar.org/CorpusID:227
71585) .

27. Edwards & Bohlen 1996, p. 13.

28. Liebeke, Manuel; Strittmatter, Nicole;
Fearn, Sarah; Morgan, A. John; Kille, Peter;
Fuchser, Jens; Wallis, David; Palchykov,
Vitalii; Robertson, Jeremy (2015-08-04).
"Unique metabolites protect earthworms
against plant polyphenols" (https://www.n
cbi.nlm.nih.gov/pmc/articles/PMC45328
35) . Nature Communications. 6: 7869.
Bibcode:2015NatCo...6.7869L (https://ui.a
dsabs.harvard.edu/abs/2015NatCo...6.7
869L) . doi:10.1038/ncomms8869 (http
s://doi.org/10.1038%2Fncomms8869) .
PMC 4532835 (https://www.ncbi.nlm.nih.
gov/pmc/articles/PMC4532835) .
PMID 26241769 (https://pubmed.ncbi.nl
m.nih.gov/26241769) .

29. Edwards & Bohlen 1996, pp. 13–15.

30. Sims & Gerard 1985, p. 10.

31. Cleveland P. Hickman Jr.; Larry S. Roberts;

Frances M Hickman (1984). Integrated
Principles of Zoology (https://archive.org/
details/integratedprinci7ed0hick/page/3
44) (7th ed.). Times Mirror/Mosby
College Publishing. pp. 344–345 (https://a
rchive.org/details/integratedprinci7ed0hic
k/page/344) . ISBN 978-0-8016-2173-4.

32. Farabee, H.J. "Excretory System" (https://

web.archive.org/web/20120730161917/ht
tp://www.emc.maricopa.edu/faculty/farab
ee/BIOBK/BioBookEXCRET.html) .
Archived from the original (http://www.em
c.maricopa.edu/faculty/farabee/biobk/bio
bookexcret.html) on 30 July 2012.
Retrieved 29 July 2012.
33. Cleveland P. Hickman Jr.; Larry S. Roberts;
Frances M Hickman (1984). Integrated
Principles of Zoology (https://archive.org/
details/integratedprinci7ed0hick/page/3
45) (7th ed.). Times Mirror/Mosby
College Publishing. pp. 345–346 (https://a
rchive.org/details/integratedprinci7ed0hic
k/page/345) . ISBN 978-0-8016-2173-4.
34. Domínguez J, Aira M, Breinholt JW,
Stojanovic M, James SW, Pérez-Losada M
(2015). "Underground evolution: New
roots for the old tree of lumbricid
earthworms" (https://www.ncbi.nlm.nih.go
v/pmc/articles/PMC4766815) . Mol.
Phylogenet. Evol. 83: 7–19.
doi:10.1016/j.ympev.2014.10.024 (https://
doi.org/10.1016%2Fj.ympev.2014.10.02
4) . PMC 4766815 (https://www.ncbi.nlm.
nih.gov/pmc/articles/PMC4766815) .
PMID 25463017 (https://pubmed.ncbi.nl
m.nih.gov/25463017) .
35. Fernández R, Almodóvar A, Novo M,
Simancas B, Díaz Cosín DJ (2012).
"Adding complexity to the complex: new
insights into the phylogeny, diversification
and origin of parthenogenesis in the
Aporrectodea caliginosa species complex
(Oligochaeta, Lumbricidae)". Mol.
Phylogenet. Evol. 64 (2): 368–79.
doi:10.1016/j.ympev.2012.04.011 (https://
doi.org/10.1016%2Fj.ympev.2012.04.01
1) . PMID 22542691 (https://pubmed.ncb
i.nlm.nih.gov/22542691) .
36. Cosín D.J.D., Novo M., Fernández R.
(2011) Reproduction of Earthworms:
Sexual Selection and Parthenogenesis. In:
Karaca A. (eds) Biology of Earthworms.
Soil Biology, vol 24. Springer, Berlin,
Heidelberg, pp. 76ff. [1] (https://www.ucm.
es/data/cont/media/www/pag-26131/Re
production%20of%20earthworms%20(201
1).%20Cap.%20Libro.pdf)
37. Novo M, Riesgo A, Fernández-Guerra A,
Giribet G (2013). "Pheromone evolution,
reproductive genes, and comparative
transcriptomics in mediterranean
earthworms (annelida, oligochaeta,
hormogastridae)" (https://doi.org/10.109
3%2Fmolbev%2Fmst074) . Mol. Biol. Evol.
30 (7): 1614–29.
doi:10.1093/molbev/mst074 (https://doi.o
rg/10.1093%2Fmolbev%2Fmst074) .
PMID 23596327 (https://pubmed.ncbi.nl
m.nih.gov/23596327) .
38. Bernstein H, Hopf FA, Michod RE (1987).
"The molecular basis of the evolution of
sex". Molecular Genetics of Development.
Adv. Genet. Advances in Genetics. Vol. 24.
pp. 323–70. doi:10.1016/S0065-
2660(08)60012-7 (https://doi.org/10.101
6%2FS0065-2660%2808%2960012-7) .
ISBN 978-0-12-017624-3. PMID 3324702
(https://pubmed.ncbi.nlm.nih.gov/33247
02) .
39. Quillan, K.J. (2000). "Ontogenetic scaling
of burrowing forces in the earthworm
Lumbricus terrestris" (http://jeb.biologists.
org/content/203/18/2757.full.pdf+html) .
Journal of Experimental Biology. 203 (Pt
18): 2757–2770.
doi:10.1242/jeb.203.18.2757 (https://doi.
org/10.1242%2Fjeb.203.18.2757) .
PMID 10952876 (https://pubmed.ncbi.nl
m.nih.gov/10952876) . Retrieved April 4,
2015.
40. Gates, G. E. (April 1949). "Regeneration in
an Earthworm, Eisenia Foetida (Savigny)
1826. I. Anterior Regeneration" (https://we
b.archive.org/web/20070401005634/htt
p://www.biolbull.org/cgi/reprint/96/2/12
9.pdf) (PDF). The Biological Bulletin. 96
(2): 129–139. doi:10.2307/1538195 (http
s://doi.org/10.2307%2F1538195) .
JSTOR 1538195 (https://www.jstor.org/st
able/1538195) . PMID 18120625 (https://
pubmed.ncbi.nlm.nih.gov/18120625) .
Archived from the original (http://www.biol
bull.org/cgi/reprint/96/2/129.pdf) (PDF)
on 2007-04-01.
41. Gates, G. E. (1 January 1953). "On
Regenerative Capacity of Earthworms of
the Family Lumbricidae". The American
Midland Naturalist. 50 (2): 414–419.
doi:10.2307/2422100 (https://doi.org/10.
2307%2F2422100) . JSTOR 2422100 (http
s://www.jstor.org/stable/2422100) .

42. "Invertebrata 20a items" (https://web.archi

ve.org/web/20060622131046/http://www.
qvmag.tas.gov.au/zoology/invertebrata/pr
intarchive/printtext/inv20aitems.html#20b
lakemore) . Archived from the original (htt
p://www.qvmag.tas.gov.au/zoology/invert
ebrata/printarchive/printtext/inv20aitems.
html#20blakemore) on 2006-06-22.
Retrieved 2006-11-20.
43. Fender & McKey-Fender (1990). Soil
Biology Guide. Wiley-Interscience.
ISBN 978-0-471-04551-9.

44. Blakemore, R.J. (2006) (March 2006).

"Revised Key to Worldwide Earthworm
Families from Blakemore (2000) plus
Reviews of Criodrilidae (including
Biwadrilidae) and Octochaetidae" (http://w
ww.annelida.net/earthworm/Introductor
y%20Key%20to%20the%20Revised%20Fa
milies%20of%20Earthworms.pdf) (PDF).
A Series of Searchable Texts on
Earthworm Biodiversity, Ecology and
Systematics from Various Regions of the
World. annelida.net. Retrieved May 15,
2012.
45. "Earthworms" (http://earthworms.elte.h
u/) . Earthworms.elte.hu. Retrieved
19 March 2022.

46. [ref Blakemore (2016) Cosmopolitan

Earthworms]

47. Earthworms: Renewers of

Agroecosystems (SA Fall, 1990 (v3n1)) (ht
tp://www.sarep.ucdavis.edu/NEWSLTR/v3
n1/sa-9.htm) Archived (https://web.archi
ve.org/web/20070713122050/http://www.
sarep.ucdavis.edu/NEWSLTR/v3n1/sa-9.
htm) 2007-07-13 at the Wayback
Machine
48. Gould, Edwin; McShea, William; Grand,
Theodore (1993). "Function of the Star in
the Star-Nosed Mole, Condylura cristata".
Journal of Mammalogy. 74 (1): 108–116.
doi:10.2307/1381909 (https://doi.org/10.
2307%2F1381909) . ISSN 0022-2372 (http
s://www.worldcat.org/issn/0022-2372) .
JSTOR 1381909 (https://www.jstor.org/st
able/1381909) .

49. Dejean, A.; Schatx, B. (1999). "Prey

Capture Behavior of Psalidomyrmex
procerus (Formicidae; Ponerinae), a
Specialist Predator of Earthworms
(Annelida)". Sociobiology: 545–554.
ISSN 0361-6525 (https://www.worldcat.or
g/issn/0361-6525) .
50. Oliver, James H. (1962). "A Mite Parasitic
in the Cocoons of Earthworms" (https://w
ww.jstor.org/stable/3275424) . The
Journal of Parasitology. 48 (1): 120–123.
doi:10.2307/3275424 (https://doi.org/10.
2307%2F3275424) . ISSN 0022-3395 (http
s://www.worldcat.org/issn/0022-3395) .
JSTOR 3275424 (https://www.jstor.org/st
able/3275424) . PMID 14481811 (https://
pubmed.ncbi.nlm.nih.gov/14481811) .
51. Blakemore, R.J. (2018) (2018). "Critical
Decline of Earthworms from Organic
Origins under Intensive, Humic SOM-
Depleting Agriculture" (https://doi.org/10.
3390%2Fsoilsystems2020033) . Soil
Systems. Soil Systems 2(2): 33. 2 (2): 33.
doi:10.3390/soilsystems2020033 (https://
doi.org/10.3390%2Fsoilsystems202003
3) .

52. NSW Department of Primary Industries,

How earthworms can help your soil (htt
p://www.dpi.nsw.gov.au/agriculture/soils/
biology/earthworms)
53. Galveston County Master Gardener
Association, Beneficials in the garden: #38
Earthworms (http://aggie-horticulture.tam
u.edu/galveston/beneficials/beneficial-38
_earthworms.htm)

54. Darwin, Charles (1881). The Formation of

Vegetable Mould through the Action of
Worms, with Observations on their Habits.
John Murray. Found at Project Gutenberg
Etext Formation of Vegetable Mould, by
Darwin (http://www.gutenberg.org/ebook
s/2355)
55. Blakemore, R.J. (2007). "Origin and means
of dispersal of cosmopolitan Pontodrilus
litoralis (Oligochaeta: Megascolecidae)" (h
ttps://www.researchgate.net/publication/
240435104) . European Journal of Soil
Biology.

56. Fründ, Heinz-Christian; Graefe, Ulfert;

Tischer, Sabine (2011), Karaca, Ayten
(ed.), "Earthworms as Bioindicators of Soil
Quality" (http://link.springer.com/10.1007/
978-3-642-14636-7_16) , Biology of
Earthworms, Berlin, Heidelberg: Springer
Berlin Heidelberg, vol. 24, pp. 261–278,
doi:10.1007/978-3-642-14636-7_16 (http
s://doi.org/10.1007%2F978-3-642-14636-
7_16) , ISBN 978-3-642-14635-0, retrieved
2021-02-18
57. Nyle C. Brady; Ray R. Weil (2009).
Elements of the Nature and Properties of
Soils (3rd ed.). Prentice Hall. ISBN 978-0-
13-501433-2.

58. Lines-Kelly, Rebecca (2021). "How

earthworms can help your soil" (https://w
ww.dpi.nsw.gov.au/agriculture/soils/guide
s/soil-biology/earthworms) .
www.dpi.nsw.gov.au. Archived (https://we
b.archive.org/web/20211008175255/http
s://www.dpi.nsw.gov.au/agriculture/soils/
guides/soil-biology/earthworms) from
the original on 2021-10-08. Retrieved
2021-11-30.

59. Mollison, Bill, Permaculture- A Designer's

Manual, Tagari Press, 1988
60. R. T. Fisher (January 1928). "Soil Changes
and Silviculture on the Harvard Forest" (htt
ps://harvardforest.fas.harvard.edu/sites/h
arvardforest.fas.harvard.edu/files/publicat
ions/pdfs/Fisher_Ecology_1928.pdf)
(PDF). Ecology. 9 (1): 6–11.
doi:10.2307/1929537 (https://doi.org/10.
2307%2F1929537) . JSTOR 1929537 (http
s://www.jstor.org/stable/1929537) .
Retrieved 2022-03-19.
61. Langmaid, K. K. (1 February 1964). "Some
Effects of Earthworm Invasion in Virgin
Podzols" (https://www.nrcresearchpress.c
om/doi/pdf/10.4141/cjss64-005) .
Canadian Journal of Soil Science. 44 (1):
34–37. doi:10.4141/cjss64-005 (https://d
oi.org/10.4141%2Fcjss64-005) . Retrieved
19 March 2022.

62. Cooper, Shewell; Soil, Humus And Health

ISBN 978-0-583-12796-7

63. Blakemore, R.J. (2017) (2017-02-12).

"Nature article to commemorate Charles
Darwin's birthday on 12th February" (http
s://vermecology.wordpress.com/2017/0
2/12/nature-article-to-commemorate-charl
es-darwins-birthday-on-12th-feb/) .
VermEcology.
64. Blanchart, Eric (1992-12-01). "Restoration
by earthworms (megascolecidae) of the
macroaggregate structure of a
destructured savanna soil under field
conditions". Soil Biology and
Biochemistry. 24 (12): 1587–1594.
doi:10.1016/0038-0717(92)90155-Q (http
s://doi.org/10.1016%2F0038-0717%289
2%2990155-Q) .

65. Burke, David (December 26, 2019). "The

power of earthworm poop and how it
could influence climate change" (https://w
ww.cbc.ca/news/canada/nova-scotia/eart
hworms-climate-change-carbon-research-
1.5370724) . CBC.
 66. Blakemore, R.J. (2015). "Eco-taxonomic
profile of the iconic vermicomposter - the
'African Nightcrawler', Eudrilus eugeniae
(Kinberg, 1867)" (https://web.archive.org/
web/20161022191344/http://africaninvert
ebrates.org/ojs/index.php/AI/article/view
File/395/421) . African Invertebrates 56:
527-548. Archived from the original (htt
p://africaninvertebrates.org/ojs/index.ph
p/AI/article/viewFile/395/421) on 2016-
10-22.

67. Collicutt, Doug. "Biology of the Night

Crawler (Lumbricus terrestris)" (http://ww
w.naturenorth.com/fall/ncrawler/Night_Cr
awlers_03.html) . NatureNorth. Retrieved
5 June 2022.

Works cited
Blakemore, Robert J. (2012).
Cosmopolitan Earthworms – an Eco-
Taxonomic Guide to the Peregrine
Species of the World. (5th Ed).
Yokohama, Japan: VermEcology.

Edwards, Clive A.; Bohlen, P. J. (1996).

Biology and Ecology of Earthworms (htt
ps://books.google.com/books?id=ad4rD
wD_GhsC) . Springer Science &
Business Media. ISBN 978-0-412-56160-
3.

Sims, Reginald William; Gerard, B

(1985). Earthworms: Keys and Notes for
the Identification and Study of the
Species. London: Published for The
 Linnean Society of London and the
Estuarine and Brackish-Water Sciences
Association by E. J. Brill/Dr. W.
Backhuys.

Further reading
Edwards, Clive A. (ed.) Earthworm
Ecology. Boca Raton: CRC Press, 2004.
Second revised edition. ISBN 0-8493-
1819-X

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

External links
Media related to Earthworms at
Wikimedia Commons

Data related to Lumbricina at

Wikispecies

Chisholm, Hugh, ed. (1911).

"Earthworm" (https://en.wikisource.org/
wiki/1911_Encyclop%C3%A6dia_Britanni
ca/Earthworm) . Encyclopædia
 Britannica (11th ed.). Cambridge
University Press.

Retrieved from
"https://en.wikipedia.org/w/index.php?
title=Earthworm&oldid=1140422576"

This page was last edited on 20 February 2023, at

00:59 (UTC). •
Content is available under CC BY-SA 3.0 unless
otherwise noted.