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Leech
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For other uses, see Leech (disambiguation).

Leech

Temporal range: Silurian–recent

PreꞒ

Pg

N
 Hirudo medicinalis sucking blood

Helobdella sp.

Scientific classification

Kingdom: Animalia

Phylum: Annelida

Class: Clitellata

Subclass: Hirudinea
Lamarck, 1818

Leeches are segmented parasitic or predatory worms that belong to

the phylum Annelida and comprise the subclass Hirudinea. They are closely
related to the oligochaetes, which include the earthworm, and like them have
soft, muscular, segmented bodies that can lengthen and contract. Both
groups are hermaphrodites and have a clitellum, but leeches typically differ
from the oligochaetes in having suckers at both ends and in having ring
markings that do not correspond with their internal segmentation. The body is
muscular and relatively solid, and the coelom, the spacious body cavity found
in other annelids, is reduced to small channels.
The majority of leeches live in freshwater habitats, while some species can be
found in terrestrial or marine environments. The best-known species, such as
the medicinal leech, Hirudo medicinalis, are hematophagous, attaching
themselves to a host with a sucker and feeding on blood, having first secreted
the peptide hirudin to prevent the blood from clotting. The jaws used to pierce
the skin are replaced in other species by a proboscis which is pushed into the
skin. A minority of leech species are predatory, mostly preying on small
invertebrates.
The eggs are enclosed in a cocoon, which in aquatic species is usually
attached to an underwater surface; members of one family, Glossiphoniidae,
exhibit parental care, the eggs being brooded by the parent. In terrestrial
species, the cocoon is often concealed under a log, in a crevice or buried in
damp soil. Almost seven hundred species of leech are currently recognised,
of which some hundred are marine, ninety terrestrial and the remainder
freshwater.
Leeches have been used in medicine from ancient times until the 19th century
to draw blood from patients. In modern times, leeches find medical use in
treatment of joint diseases such as epicondylitis and osteoarthritis, extremity
vein diseases, and in microsurgery, while hirudin is used as
an anticoagulant drug to treat blood-clotting disorders.

Contents

 1Diversity and phylogeny

o 1.1Evolution
 2Anatomy and physiology
o 2.1Reproduction and development
o 2.2Feeding and digestion
o 2.3Nervous system
o 2.4Gas exchange
o 2.5Movement
 3Interactions with humans
o 3.1Bites
o 3.2In human culture
o 3.3Water pollution
 4References
 5General bibliography
 6External links

Diversity and phylogeny

Haemadipsa zeylanica, a terrestrial leech

Placobdelloides siamensis, a parasite of turtles in Thailand. The ventral face (right) shows many
young leeches.[1]

Some 680 species of leech have been described, of which around 100 are
marine, 480 freshwater and the remainder terrestrial.[2][3] Among Euhirudinea,
the true leeches, the smallest is about 1 cm (1⁄2 in) long, and the largest is the
giant Amazonian leech, Haementeria ghilianii, which can reach 30 cm (12 in).
Except for Antarctica,[2] leeches are found throughout the world but are at their
most abundant in temperate lakes and ponds in the northern hemisphere. The
majority of freshwater leeches are found in the shallow, vegetated areas on
the edges of ponds, lakes and slow-moving streams; very few species tolerate
fast-flowing water. In their preferred habitats, they may occur in very high
densities; in a favourable environment with water high in organic pollutants,
over 10,000 individuals were recorded per square metre (over 930 per square
foot) under rocks in Illinois. Some species aestivate during droughts, burying
themselves in the sediment, and can lose up to 90% of their bodyweight and
still survive.[4] Among the freshwater leeches are the Glossiphoniidae, dorso-
ventrally flattened animals mostly parasitic on vertebrates such as turtles, and
unique among annelids in both brooding their eggs and carrying their young
on the underside of their bodies.[5]
The terrestrial Haemadipsidae are mostly native to the tropics and
subtropics,[6] while the aquatic Hirudinidae have a wider global range; both of
these feed largely on mammals, including humans.[4] A distinctive family is
the Piscicolidae, marine or freshwater ectoparasites chiefly of fish, with
cylindrical bodies and usually well-marked, bell-shaped, anterior
suckers.[7] Not all leeches feed on blood; the Erpobdelliformes, freshwater or
amphibious, are carnivorous and equipped with a relatively large, toothless
mouth to ingest insect larvae, molluscs, and other annelid worms, which are
swallowed whole.[8] In turn, leeches are prey to fish, birds, and invertebrates.[9]
The name for the subclass, Hirudinea, comes from the
Latin hirudo (genitive hirudinis), a leech; the element -bdella found in many
leech group names is from the Greek βδέλλα bdella, also meaning
leech.[10] The name Les hirudinées was given by Jean-Baptiste Lamarck in
1818.[11] Leeches were traditionally divided into two infraclasses,
the Acanthobdellidea and the Euhirudinea (true leeches).[12] The Euhirudinea
are divided into the proboscis-bearing Rhynchobdellida and the rest, including
some jawed species, the "Arhynchobdellida", without a proboscis.[13]
The phylogenetic tree of the leeches and their annelid relatives is based
on molecular analysis (2019) of DNA sequences. Both the former classes
"Polychaeta" (bristly marine worms) and "Oligochaeta" (including the
earthworms) are paraphyletic: in each case the complete groups (clades)
would include all the other groups shown below them in the tree.
The Branchiobdellida are sister to the leech clade Hirudinida, which
approximately corresponds to the traditional subclass Hirudinea. The main
subdivision of leeches is into the Rhynchobdellida and the Arhynchobdellida,
though the Acanthobdella are sister to the clade that contains these two
groups.[13]
Anne  
lida  
  "Polychaeta" (exc. "Oligochaeta")
Clite  
llata  
 
"Oligochaeta" (exc. Lumbriculidae)
   
   

Lumbriculidae (blackworms)
  Branchiobdellida
  symbionts

Hirudinida  
parasitic   Acanthobdella
Euhirudinea Arhynchobdellida  
     

Erpobdellidae
 
 

Hirudiniformes

Rhynchobdellida  
   

Glossiphoniidae
Oceanobdelliformes  
  ectoparasitic

Piscicolidae
  
 

Ozobranchid

Evolution

Fossil of a possible leech from the Silurian of Wisconsin

The most ancient annelid group consists of the free-living polychaetes that
evolved in the Cambrian period, being plentiful in the Burgess Shale about
500 million years ago. Oligochaetes evolved from polychaetes and the
leeches branched off from the oligochaetes. Both the oligochaetes and the
leeches, having no hard parts, do not fossilise well.[14] The oldest leech fossils
are from the Jurassic period around 150 million years ago, but a fossil with
external ring markings found in Wisconsin in the 1980s, with what appears to
be a large sucker, seems to extend the group's evolutionary history back to
the Silurian, some 437 million years ago.[15][16]

Anatomy and physiology

Leeches show a remarkable similarity to each other in morphology, very
different from typical annelids which are cylindrical with a fluid-filled space,
the coelom (body cavity). In leeches, the coelom is reduced to four slender
longitudinal channels, and the interior of the body is filled with a
solid dermis in between the various organs. Typically, the body is dorso-
ventrally flattened and tapers at both ends. Longitudinal and circular muscles
in the body wall are supplemented by diagonal muscles, giving the leech the
ability to adopt a large range of body shapes and show great flexibility. Most
leeches have a sucker at both the anterior (front) and posterior (back) ends,
but some primitive leeches have a single sucker at the back.[17][18]
Leech anatomy in cross-section: the body is solid, the coelom (body cavity) reduced to channels,
with circular, longitudinal, and transverse muscles making the animal strong and flexible. [19]

Like other annelids, the leech is a segmented animal, but unlike other
annelids, the segmentation is masked by external ring markings
(annulations).[20] The number of annulations varies, both between different
regions of the body and between species.[17] In one species, the body surface
is divided into 102 annuli,[21] but the body consists of 33 segments, a number
constant across all leech species. Of these segments, the first five are
designated as the head and include the anterior brain,
several ocelli (eyespots) dorsally and the sucker ventrally. The following 21
mid-body segments each contain a nerve ganglion, and between them
contain two reproductive organs, a single female gonopore and nine pairs
of testes. The last seven segments contain the posterior brain and are fused
to form the animal's tail sucker.[17]
The body wall consists of a cuticle, an epidermis and a thick layer of
fibrous connective tissue in which are embedded the circular muscles, the
diagonal muscles and the powerful longitudinal muscles. There are also
dorso-ventral muscles. The coelomic channels run the full length of the body,
the two main ones being on either side; these have taken over the function of
the hemal system (blood vessels) in other annelids. Part of the
lining epithelium consists of chloragogen cells which are used for the storage
of nutrients and in excretion. There are 10 to 17 pairs
of metanephridia (excretory organs) in the mid-region of the leech. From
these, ducts typically lead to a urinary bladder, which empties to the outside at
a nephridiopore.[19]
Reproduction and development
Further information: Leech embryogenesis
Leeches are protandric hermaphrodites, with the male reproductive organs,
the testes, maturing first and the ovaries later. In hirudinids, a pair will line up
with the clitellar regions in contact, with the anterior end of one leech pointing
towards the posterior end of the other; this results in the male gonopore of
one leech being in contact with the female gonopore of the other. The penis
passes a spermatophore into the female gonopore and sperm is transferred
to, and probably stored in, the vagina.[22]
Some jawless leeches (Rhynchobdellida) and proboscisless
leeches (Arhynchobdellida) lack a penis, and in these, sperm is passed from
one individual to another by hypodermic injection. The leeches intertwine and
grasp each other with their suckers. A spermatophore is pushed by one
through the integument of the other, usually into the clitellar region. The
sperm is liberated and passes to the ovisacs, either through the coelomic
channels or interstitially through specialist "target tissue" pathways. [22]
Some time after copulation, the small, relatively yolkless eggs are laid. In
most species, an albumin-filled cocoon is secreted by the clitellum and
receives one or more eggs as it passes over the female gonopore.[22] In the
case of the North American Erpobdella punctata, the clutch size is about five
eggs, and some ten cocoons are produced.[23] Each cocoon is fixed to a
submerged object, or in the case of terrestrial leeches, deposited under a
stone or buried in damp soil. The cocoon of Hemibdella soleae is attached to
a suitable fish host.[22][24] The glossiphoniids brood their eggs, either by
attaching the cocoon to the substrate and covering it with their ventral surface,
or by securing the cocoon to their ventral surface, and even carrying the
newly hatched young to their first meal.[25]
When breeding, most marine leeches leave their hosts and become free-living
in estuaries. Here they produce their cocoons, after which the adults of most
species die. When the eggs hatch, the juveniles seek out potential hosts when
these approach the shore.[25] Leeches mostly have an annual or biannual life
cycle.[22]
Feeding and digestion
About three quarters of leech species are parasites that feed on the blood of a
host, while the remainder are predators. Leeches either have a pharynx that
they can protrude, commonly called a proboscis, or a pharynx that they
cannot protrude, which in some groups is armed with jaws.[26]
In the proboscisless leeches, the jaws (if any) of Arhynchobdellids are at the
front of the mouth, and have three blades set at an angle to each other. In
feeding, these slice their way through the skin of the host, leaving a Y-shaped
incision. Behind the blades is the mouth, located ventrally at the anterior end
of the body. It leads successively into the pharynx, a short oesophagus,
a crop (in some species), a stomach and a hindgut, which ends at an anus
located just above the posterior sucker. The stomach may be a simple tube,
but the crop, when present, is an enlarged part of the midgut with a number of
pairs of ceca that store ingested blood. The leech secretes an
anticoagulant, hirudin, in its saliva which prevents the blood from clotting
before ingestion.[26] A mature medicinal leech may feed only twice a year,
taking months to digest a blood meal.[18]
Leech bites on a cow's udder

The bodies of predatory leeches are similar, though instead of a jaw many
have a protrusible proboscis, which for most of the time they keep retracted
into the mouth. Such leeches are often ambush predators that lie in wait until
they can strike prey with the proboscises in a spear-like fashion.[27] Predatory
leeches feed on small invertebrates such as snails, earthworms and insect
larvae. The prey is usually sucked in and swallowed whole. Some
Rhynchobdellida however suck the soft tissues from their prey, making them
intermediate between predators and blood-suckers.[26]

Leech attacking a slug

Blood-sucking leeches use their anterior suckers to connect to hosts for

feeding. Once attached, they use a combination of mucus and suction to stay
in place while they inject hirudin into the hosts' blood. In general, blood-
feeding leeches are non host-specific, and do little harm to their host,
dropping off after consuming a blood meal. Some marine species however
remain attached until it is time to reproduce. If present in great numbers on a
host, these can be debilitating, and in extreme cases, cause death.[25]
Leeches are unusual in that they do not
produce amylases, lipases or endopeptidases.[26] This lack of endopeptidases
means the mechanism of protein digestion cannot follow the same sequence
as it would in all other animals in which endopeptidases first split proteins
into peptides, and the exopeptidases then degrade the peptides.[28] Leeches
produce intestinal exopeptidases which remove amino acids from the long
protein molecules one by one, possibly aided
by proteases from endosymbiotic bacteria in the hindgut.[29] This evolutionary
choice of exopeptic digestion in Hirudinea distinguishes these carnivorous
clitellates from oligochaetes, and may explain why digestion in leeches is so
slow.[26]
A deficiency of digestive enzymes and of B complex vitamins is compensated
for by enzymes and vitamins produced by endosymbiotic microflora. In Hirudo
medicinalis, these supplementary factors are produced by an
obligatory mutualistic relationship with two bacterial species, Aeromonas
veronii and a still-uncharacterised Rikenella species. Non-bloodsucking
leeches, such as Erpobdella punctata, are host to three bacterial
symbionts, Pseudomonas, Aeromonas, and Klebsiella spp. (a slime
producer). The bacteria are passed from parent to offspring in the cocoon as it
is formed.[28]
Nervous system
A leech's nervous system is formed of a few large nerve cells; their large size
makes leeches convenient as model organisms for the study of invertebrate
nervous systems. The main nerve centre consists of the cerebral ganglion
above the gut and another ganglion beneath it, with connecting nerves
forming a ring around the pharynx a little way behind the mouth. A nerve cord
runs backwards from this in the ventral coelomic channel, with 21 pairs
of ganglia in segments six to 26. In segments 27 to 33, other paired ganglia
fuse to form the caudal ganglion.[30] Several sensory nerves connect directly to
the cerebral ganglion; there are sensory and motor nerve cells connected to
the ventral nerve cord ganglia in each segment.[18]
Leeches have between two and ten pigment spot ocelli, arranged in pairs
towards the front of the body. There are also sensory papillae arranged in a
lateral row in one annulation of each segment. Each papilla contains many
sensory cells. Some rhynchobdellids have the ability to change colour
dramatically by moving pigment in chromatophore cells; this process is under
the control of the nervous system but its function is unclear as the change in
hue seems unrelated to the colour of the surroundings.[30]
Leeches can detect touch, vibration, movement of nearby objects, and
chemicals secreted by their hosts; freshwater leeches crawl or swim towards
a potential host standing in their pond within a few seconds. Species that feed
on warm-blooded hosts move towards warmer objects. Many leeches avoid
light, though some blood feeders move towards light when they are ready to
feed, presumably increasing the chances of finding a host.[18]
Gas exchange
Leeches live in damp surroundings and in general respire through their body
wall. The exception to this is in the Piscicolidae, where branching or leaf-like
lateral outgrowths from the body wall form gills. Some rhynchobdellid leeches
have an extracellular haemoglobin pigment, but this only provides for about
half of the leech's oxygen transportation needs, the rest occurring by
diffusion.[19]
Movement
Leeches move using their longitudinal and circular muscles in a modification
of the locomotion by peristalsis, self-propulsion by alternately contracting and
lengthening parts of the body, seen in other annelids such as earthworms.
They use their posterior and anterior suckers (one on each end of the body) to
enable them to progress by looping or inching along, in the manner
of geometer moth caterpillars. The posterior end is attached to the substrate,
and the anterior end is projected forward peristaltically by the circular muscles
until it touches down, as far as it can reach, and the anterior end is attached.
Then the posterior end is released, pulled forward by the longitudinal muscles,
and reattached; then the anterior end is released, and the cycle
repeats.[31][18] Leeches explore their environment with head movements and
body waving.[32] The Hirudinidae and Erpobdellidae can swim rapidly with up-
and-down or sideways undulations of the body; the Glossiphoniidae in
contrast are poor swimmers and curl up and fall to the sediment below when
disturbed.[33]
 

Leeches move by looping using their front and back

[31]
suckers.

Video of looping movement

Interactions with humans

Leeches can be removed by hand, since they do not burrow into the skin or leave the head in the
wound.[34][35]

Bites
Leech bites are generally alarming rather than dangerous, though a small
percentage of people have severe allergic or anaphylactic reactions and
require urgent medical care. Symptoms of these reactions include red
blotches or an itchy rash over the body, swelling around the lips or eyes, a
feeling of faintness or dizziness, and difficulty in breathing.[36] An externally
attached leech will detach and fall off on its own accord when it is satiated on
blood, which may take from twenty minutes to a few hours; bleeding from the
wound may continue for some time.[36] Internal attachments, such as inside the
nose, are more likely to require medical intervention.[37]
Bacteria, viruses, and protozoan parasites from previous blood sources can
survive within a leech for months, so leeches could potentially act as vectors
of pathogens. Nevertheless, only a few cases of leeches transmitting
pathogens to humans have been reported.[38][39]
Leech saliva is commonly believed to contain anaesthetic compounds to
numb the bite area, but this has never been proven.[40] Although morphine-like
substances have been found in leeches, they have been found in the neural
tissues, not the salivary tissues. They are used by the leeches in modulating
their own immunocytes and not for anaesthetising bite areas on their
hosts.[41][40] Depending on the species and size, leech bites can be barely
noticeable or they can be fairly painful.[42][43]
In human culture
Further information: Hirudotherapy
The leech appears in Proverbs 30:15 as an archetype of
insatiable greed.[44] More generally, a leech is a persistent social
parasite or sycophant.[45]
The medicinal leech Hirudo medicinalis, and some other species, have been
used for clinical bloodletting for at least 2,500 years: Ayurvedic texts describe
their use for bloodletting in ancient India. In ancient Greece, bloodletting was
practised according to the theory of humours found in the Hippocratic
Corpus of the fifth century BC, which maintained that health depended on a
balance of the four humours: blood, phlegm, black bile and yellow bile.
Bloodletting using leeches enabled physicians to restore balance if they
considered blood was present in excess.[46][47]
Pliny the Elder reported in his Natural History that the horse leech could drive
elephants mad by climbing up inside their trunks to drink blood.[48] Pliny also
noted the medicinal use of leeches in ancient Rome, stating that they were
often used for gout, and that patients became addicted to the
treatment.[49] In Old English, lǣce was the name for a physician as well as for
the animal, though the words had different origins, and lǣcecraft, leechcraft,
was the art of healing.[50]

"Leech finders" from The Costume of Yorkshire by George

Walker, 1814, engraved by Robert Havell
 

Pharmacy leech jar with airholes in the lid. England, 1830–

1870.

 Three leech doctors decide on bloodletting for

their grasshopper patient. Lithograph by F.-J.-V.
Broussais from a cartoon by J. J. Grandville, c. 1832
William Wordsworth's 1802 poem "Resolution and Independence" describes
one of the last of the leech-gatherers, people who travelled Britain catching
leeches from the wild, and causing a sharp decline in their abundance, though
they remain numerous in Romney Marsh. By 1863, British hospitals had
switched to imported leeches, some seven million being imported to hospitals
in London that year.[48]
In the nineteenth century, demand for leeches was sufficient for hirudiculture,
the farming of leeches, to become commercially viable.[51] Leech usage
declined with the demise of humoral theory,[52] but made a small-scale
comeback in the 1980s after years of decline, with the advent of microsurgery,
where venous congestion can arise due to inefficient venous drainage.
Leeches can reduce swelling in the tissues and promote healing, helping in
particular to restore circulation after microsurgery to reattach body
parts.[53][54] Other clinical applications include varicose veins, muscle
cramps, thrombophlebitis, and joint diseases such as epicondylitis
and osteoarthritis.[55][56][57][58]
Leech secretions contain several bioactive substances with anti-
inflammatory, anticoagulant and antimicrobial effects.[57] One active
component of leech saliva is a small protein, hirudin.[59] It is widely used as an
anticoagulant drug to treat blood-clotting disorders, and manufactured
by recombinant DNA technology.[60][61]
In 2012 and 2018, Ida Schnell and colleagues trialled the use
of Haemadipsa leeches to gather data on the biodiversity of
their mammalian hosts in the tropical rainforest of Vietnam, where it is hard to
obtain reliable data on rare and cryptic mammals. They showed that
mammal mitochondrial DNA, amplified by the polymerase chain reaction, can
be identified from a leech's blood meal for at least four months after feeding.
They detected Annamite striped rabbit, small-toothed ferret-badger, Truong
Son munjtac, and serow in this way.[62][63]
Water pollution
Exposure to synthetic estrogen as used in contraceptive medicines, which
may enter freshwater ecosystems from municipal wastewater, can affect
leeches' reproductive systems. Although not as sensitive to these compounds
as fish, leeches showed physiological changes after exposure, including
longer sperm sacs and vaginal bulbs, and decreased epididymis weight.[64]

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General bibliography
 Ruppert, Edward E.; Fox, Richard S.; Barnes,
Robert D. (2004). Invertebrate Zoology, 7th
Edition. Cengage Learning. ISBN 978-81-315-
0104-7.

External links
 Media related to Hirudinea at Wikimedia
Commons
 Data related to Hirudinea at Wikispecies
 The dictionary definition of leech at Wiktionary
-8e8e-71521e70b145

Categories:
 Leeches
 Clitellata
 Extant Silurian first appearances
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