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Louse (: lice) is the common name for any member of the Phthiraptera, which contains nearly 5,000 species of wingless . Phthiraptera was previously recognized as an order, until a 2021 genetic study determined that they are a highly modified lineage of the order , whose members are commonly known as booklice, barklice or barkflies.

Lice are obligate parasites, living externally on warm-blooded hosts, which include every species of and , except for , , and . Chewing lice live among the hairs or feathers of their host and feed on skin and debris, whereas sucking lice pierce the host's skin and feed on blood and other secretions. They usually spend their whole life on a single host, cementing their eggs, called nits, to hairs or feathers. The eggs hatch into nymphs, which moult three times before becoming fully grown, a process that takes about four weeks.

Humans host two species of louse—the and the are subspecies of Pediculus humanus; and the , pubis. Lice are vectors of diseases such as . Lice were ubiquitous in human society until at least the . They appear in folktales, songs such as The Kilkenny Louse House, and novels such as 's .

The body louse has the smallest of any known insect; it has been used as a and has been the subject of much research. They commonly feature in the psychiatric disorder delusional parasitosis. A louse was one of the early subjects of , appearing in 's 1667 book, .

The oldest known fossil lice are from the .


Classification
Lice are classified as an called Phthiraptera, named by in 1896. Phthiraptera was previously recognized as an order, and was considered the to the order , which contains booklice, barklice and barkflies, all within the greater group called . However, a 2021 genetic study determined that Phthiraptera lice are a highly modified lineage within the Psocoptera group, thus rendering Psocoptera and necessating a classification change. Phthiraptera is now nested within the , belonging to the now-ranked order . Within Psocodea, the Phthiraptera lice are most closely related to family .

A showing the position of Phthiraptera within is shown below:


Internal classification
Phthiraptera is clearly a grouping, united as the members are by a number of derived features, including their parasitism on vertebrates and the combination of their with their ganglia to form a single ventral nerve junction.
(2025). 9783662103968, Springer Science & Business Media. .

The order has traditionally been divided into two suborders, the sucking lice () and the chewing lice (). However, subsequent classifications indicated that the Mallophaga are , and four suborders were then recognized.

Upon finding that Phthiraptera was nested within in 2021, de Moya et al. reduced the rank of Phthiraptera to , and the were reduced down to . Also, based on the nature of one of the parvorders - - they proposed creation of a fifth parvorder to be created and split off from Ischnocera. Here are the five proposed parvorders within Phthiraptera:


Evolutionary history
The oldest confirmed fossil louse is Archimenopon myanmarensis, an from the Cretaceous amber from . Another early representative of the group is a bird louse, Megamenopon rasnitsyni, from , Germany, which dates to the , around 44 million years ago. Saurodectes vrsanskyi from the Early Cretaceous () of , Russia, has also been suggested to be a louse, but this is tentative.

Placental mammal lice had a single common ancestor that lived on with this arising from host-switching from an ancient avian host.


Description
Nearly 5,000 species of louse have been identified, about 4,000 being parasitic on birds and 800 on mammals. Lice are present on every continent in all the habitats that their host animals occupy. They are found even in the , where carry 15 species of lice (in the genera and ).

File:Ricinus bombycillae (Denny, 1842).JPG| Ricinus bombycillae, an amblyceran louse from a File:Trinoton anserinum (Fabricius, 1805).JPG| Trinoton anserinum, an amblyceran louse from a File:Lice image01.jpg| limbata, an ischnoceran louse from . The species is sexually dimorphic, with the male smaller than the female.

Lice are divided into two groups: sucking lice, which obtain their nourishment from feeding on the and body fluids of their host; and chewing lice, which are , feeding on , fragments of feathers or hair, and debris found on the host's body. Many lice are specific to a single species of host and have co-evolved with it. In some cases, they live on only a particular part of the body. Some animals are known to host up to fifteen different species, although one to three is typical for mammals, and two to six for birds. Lice generally cannot survive for long if removed from their host.

(1998). 9780195100334, Oxford University Press.
If their host dies, lice can opportunistically use to hitch a ride on a fly and attempt to find a new host.

Sucking lice range in length from . They have narrow heads and oval, flattened bodies. They have no , and their compound eyes are reduced in size or absent. Their antennae are short with three to five segments, and their mouthparts, which are retractable into their head, are adapted for piercing and sucking. There is a cibarial pump at the start of the gut; it is powered by muscles attached to the inside of the cuticle of the head. The mouthparts consist of a proboscis which is toothed, and a set of stylets arranged in a cylinder inside the proboscis, containing a salivary canal () and a food canal ().

(2025). 9781118846155, Wiley.
The thoracic segments are fused, the abdominal segments are separate, and there is a single large claw at the tip of each of the six legs.
(2025). 9781402062421, Springer Science & Business Media. .

Chewing lice are also flattened and can be slightly larger than sucking lice, ranging in length from . They are similar to sucking lice in form but the head is wider than the thorax and all species have compound eyes. There are no ocelli and the mouthparts are adapted for chewing. The antennae have three to five segments and are slender in the suborder , but club-shaped in the suborder . The legs are short and robust, and terminated by one or two claws. Some species of chewing lice house bacteria in in their bodies. These may assist in digestion because if the insect is deprived of them, it will die. Lice are usually to match the fur or feathers of the host. A louse's colour varies from pale beige to dark grey; however, if feeding on blood, it may become considerably darker.

Female lice are usually more common than males, and some species are , with young developing from unfertilized eggs. A louse's is commonly called a nit. Many lice attach their eggs to their hosts' hair with specialized ; the saliva/hair bond is very difficult to sever without specialized products. Lice inhabiting birds, however, may simply leave their eggs in parts of the body inaccessible to preening, such as the interior of feather shafts. Living louse eggs tend to be pale whitish, whereas dead louse eggs are yellower. Lice are , being born as miniature versions of the adult, known as nymphs. The young moult three times before reaching the final adult form, usually within a month after hatching.

Humans host three different kinds of lice: , , and . Head lice and body lice are subspecies of Pediculus humanus, and pubic lice are a separate species, Pthirus pubis. Lice infestations can be controlled with , and medicated shampoos or washes.


Ecology
The average number of lice per host tends to be higher in large-bodied bird species than in small ones. Lice have an aggregated distribution across bird individuals, i.e. most lice live on a few birds, while most birds are relatively free of lice. This pattern is more pronounced in territorial than in colonial—more social—bird species. Host organisms that dive under water to feed on aquatic prey harbour fewer taxa of lice. Bird taxa that are capable of exerting stronger antiparasitic defence—such as stronger immune response or larger —harbour more taxa of Amblyceran lice than others. Reductions in the size of host populations may cause a long-lasting reduction of louse taxonomic richness, for example, birds introduced into host fewer species of lice there than in Europe. Louse sex ratios are more balanced in more social hosts and more female-biased in less social hosts, presumably due to the stronger isolation among louse subpopulations (living on separate birds) in the latter case. The extinction of a species results in the extinction of its host-specific lice. Host-switching is a random event that would seem very rarely likely to be successful, but has occurred over evolutionary time-scales so it must be successfully accomplished sometimes.

Lice may reduce host life expectancy if the infestation is heavy, but most seem to have little effect on their host. The habit of dust bathing in is probably an attempt by the birds to rid themselves of lice. Lice may transmit microbial diseases and parasites, but most individuals spend their whole life cycle on a single host and are only able to transfer to a new host opportunistically. Ischnoceran lice may reduce the effect of the plumage; thus heavily infested birds lose more heat than others. Lice infestation is a disadvantage in the context of sexual rivalry.


In human culture

In social history
Lice have been intimately associated with human society throughout history. In the , they were essentially ubiquitous. At the death of , Archbishop of Canterbury in 1170, it was recorded that "The vermin boiled over like water in a simmering cauldron, and the onlookers burst into alternate weeping and laughing". The clergy often saw lice and other parasites as a constant reminder of human frailty and weakness. Monks and nuns would purposely ignore grooming themselves and suffer from infestations to express their religious devotion. A mediaeval treatment for lice was an made from pork grease, , , and .
(2025). 9780778713517, Crabtree. .

's 1667 book, , illustrated a human louse, drawn as seen down an early .

Margaret Cavendish's satirical The Description of a New World, Called The Blazing-World (1668) has "Lice-men" as "mathematicians", investigating nature by trying to weigh the air like the real scientist .

(2025). 9780801894435, JHU Press. .

In 1935 the Harvard medical researcher wrote the book Rats, Lice and History, alleging that both body and head lice transmit typhus between humans.

(2025). 9781412806725, Transaction Publishers.
Despite this, the modern view is that only the body louse can transmit the disease.

Soldiers in the trenches of the First World War suffered severely from lice, and the they carried. The Germans boasted that they had lice under effective control, but themselves suffered badly from lice in the Second World War on the Eastern Front, especially in the Battle of Stalingrad. "" became a for the extermination of Jews in concentration camps such as under the Nazi regime.

In the psychiatric disorder delusional parasitosis, patients express a persistent irrational fear of animals such as lice and mites, imagining that they are continually infested and complaining of itching, with "an unshakable false belief that live organisms are present in the skin".


In literature and folklore
's 1939 book has the character Shem the Penman infested with " fleas, the lieabed lice, ... bats in his belfry".

Clifford E. Trafzer's A Song: The Resilience of a Tribe retells the story of Sinawavi ()'s love for Poowavi (Louse). Her eggs are sealed in a basket woven by her mother, who gives it to Coyote, instructing him not to open it before he reaches home. Hearing voices coming from it, however, Coyote opens the basket and the people, the world's first human beings, pour out of it in all directions.

(2025). 9780295805825, University of Washington Press. .

The Irish songwriter John Lyons (b. 1934) wrote the popular song The Kilkenny Louse House. The song contains the lines "Well we went up the stairs and we put out the light, Sure in less than five minutes, I had to show fight. For the fleas and the bugs they collected to march, And over me stomach they formed a great arch". It has been recorded by Christie Purcell (1952), Mary Delaney on From Puck to Appleby (2003), and the on (1972) among others.

dedicated a poem to the louse, inspired by witnessing one on a lady's bonnet in church: "Ye ugly, creepin, blastid wonner, Detested, shunn'd, by saint and sinner, How dare ye set your fit upon her, sae fine lady! Gae somewhere else, and seek your dinner on some poor body." in mentioned the biblical plague of lice visited upon pharaoh: "Frogs, lice, and flies must all his palace fill with loathed intrusion, and filled all the land." recorded a Scottish proverb, "Gie a beggar a bed and he'll repay you with a Louse." In 's Troilus and Cressida, compares , brother of , to a louse: "Ask me not what I would be, if I were not Thersites; for I care not to be the louse of a lazar, so I were not Menelaus."


In science
The human Pediculus humanus humanus has the smallest insect known, and this louse can transmit certain diseases that the closely related human ( P. h. capitis) cannot. With their simple life history and small genomes, the pair make ideal to study the molecular mechanisms behind the transmission of and vector competence.

Lice have been the subject of significant research in the 2000s that led to discoveries on human evolution. The three species of sucking lice that parasitize human beings belong to two genera, and : ( Pediculus humanus capitis), ( Pediculus humanus humanus), and ( Pthirus pubis). Human head and body lice (genus Pediculus) share a common ancestor with chimpanzee lice, while pubic lice (genus Pthirus) share a common ancestor with gorilla lice. Using phylogenetic and cophylogenetic analysis, Reed et al. hypothesized that Pediculus and Pthirus are sister taxa and monophyletic. In other words, the two genera descended from the same common ancestor. The age of divergence between Pediculus and its common ancestor is estimated to be 6-7 million years ago, which matches the age predicted by chimpanzee-hominid divergence. Because parasites rely on their hosts, hostparasite cospeciation events are likely.

Genetic evidence suggests that human ancestors acquired pubic lice from approximately 3-4 million years ago. Unlike the genus Pediculus, the divergence in Pthirus does not match the age of host divergence that likely occurred 7 million years ago. Reed et al. propose a Pthirus species host-switch around 3-4 million years ago. While it is difficult to determine if a parasitehost switch occurred in evolutionary history, this explanation is the most parsimonious (containing the fewest evolutionary changes).

Additionally, the DNA differences between head lice and body lice provide corroborating evidence that humans used clothing between 80,000 and 170,000 years ago, before leaving Africa. Human head and body lice occupy distinct ecological zones: head lice live and feed on the scalp, while body lice live on clothing and feed on the body. Because body lice require clothing to survive, the divergence of head and body lice from their common ancestor provides an estimate of the date of introduction of clothing in human evolutionary history.

The mitochondrial genome of the human species of the body louse ( Pediculus humanus humanus), the head louse ( Pediculus humanus capitis) and the pubic louse ( ) fragmented into a number of at least seven million years ago. Analysis of mitochondrial DNA in human body and hair lice reveals that greater genetic diversity existed in African than in non-African lice. Human lice can also shed light on human migratory patterns in prehistory. The dominating theory of regarding human migration is the Out of Africa Hypothesis. Genetic diversity accumulates over time, and mutations occur at a relatively constant rate. Because there is more genetic diversity in African lice, the lice and their human hosts must have existed in Africa before anywhere else.


Woodlouse
The name or wood-louse is given to of the Oniscidea within the order , unrelated to lice. The Oxford English Dictionarys earliest citation of this usage is from 1611.


See also
  • Pest (organism)
  • Use of DNA in forensic entomology


External links

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