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Phthiraptera
Temporal range:Cenomanian–present
Light micrographofFahrenholzia pinnata
Scientific classificationEdit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Psocodea
Suborder: Troctomorpha
Infraorder: Nanopsocetae
Parvorder: Phthiraptera
Haeckel,1896
Clades[1]

Louse(pl.:lice) is the common name for any member of thecladePhthiraptera,which contains nearly 5,000 species of winglessparasiticinsects.Phthiraptera has variously been recognized as anorder,infraorder,or aparvorder,as a result of developments in phylogenetic research.[1][2][3]

Lice areobligate parasites,living externally on warm-bloodedhostswhich include every species ofbirdandmammal,except formonotremes,pangolins,andbats.Lice arevectorsof diseases such astyphus.

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, callednits,to hairs or feathers. The eggs hatch intonymphs,which moult three times before becoming fully grown, a process that takes about four weeks. Genetic evidence indicates that lice are a highly modified lineage ofPsocoptera(now calledPsocodea), commonly known as booklice, barklice or barkflies. The oldest known fossil lice are from theCretaceous.[4]

Humans host two species of louse—thehead louseand thebody louseare subspecies ofPediculus humanus;and thepubic louse,Pthiruspubis.The body louse has the smallestgenomeof any known insect; it has been used as amodel organismand has been the subject of much research. Lice were ubiquitous in human society until at least theMiddle Ages.They appear in folktales, songs such asThe Kilkenny Louse House,and novels such asJames Joyce'sFinnegans Wake.They commonly feature in the psychiatric disorderdelusional parasitosis.A louse was one of the early subjects ofmicroscopy,appearing inRobert Hooke's 1667 book,Micrographia.

Morphology and diversity

Lice are divided into two groups: sucking lice, which obtain their nourishment from feeding on thesebaceous secretionsand body fluids of their host; and chewing lice, which arescavengers,feeding onskin,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.[5]If their host dies, lice can opportunistically usephoresisto hitch a ride on a fly and attempt to find a new host.[6]

Sucking lice range in length from0.5 to 5 mm (164to1364in). They have narrow heads and oval, flattened bodies. They have noocelli,and their compound eyes are reduced in size or absent. Theirantennaeare short with three to five segments, and their mouthparts, which are retractable into their head, are adapted for piercing and sucking.[7]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 (ventrally) and a food canal (dorsally).[8]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.[7]

Chewing lice are also flattened and can be slightly larger than sucking lice, ranging in length from0.5 to 6 mm (164to1564in). 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 suborderIschnocera,but club-shaped in the suborderAmblycera.The legs are short and robust, and terminated by one or two claws. Some species of chewing lice housesymbioticbacteria inbacteriocytesin their bodies. These may assist in digestion because if the insect is deprived of them, it will die. Lice are usuallycryptically colouredto match the fur or feathers of the host.[7][9]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 areparthenogenetic,with young developing from unfertilized eggs. A louse'seggis commonly called a nit. Many lice attach their eggs to their hosts' hair with specializedsaliva;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 topreening,such as the interior of feather shafts. Living louse eggs tend to be pale whitish, whereas dead louse eggs are yellower.[5]Lice areexopterygotes,being born as miniature versions of the adult, known asnymphs.The young moult three times before reaching the final adult form, usually within a month after hatching.[5]

Humans host three different kinds of lice:head lice,body lice,andpubic lice.Head lice and body lice are subspecies ofPediculus humanus,and pubic lice are a separate species,Pthirus pubis.Lice infestations can be controlled withlice combs,and medicated shampoos or washes.[10]

Ecology

The average number of lice per host tends to be higher in large-bodied bird species than in small ones.[11]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.[12] Host organisms that dive under water to feed on aquatic prey harbour fewer taxa of lice.[13][14] Bird taxa that are capable of exerting stronger antiparasitic defence—such as strongerT cellimmune response or largeruropygial glands—harbour more taxa of Amblyceran lice than others.[15][16] Reductions in the size of host populations may cause a long-lasting reduction of louse taxonomic richness,[17]for example, birds introduced intoNew Zealandhost fewer species of lice there than in Europe.[18][19]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.[20]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, butspeciationhas occurred over evolutionary time-scales so it must be successfully accomplished sometimes.[17]

Lice may reduce host life expectancy if the infestation is heavy,[21]but most seem to have little effect on their host. The habit of dust bathing indomestic hensis probably an attempt by the birds to rid themselves of lice.[7]Lice may transmit microbial diseases andhelminthparasites,[22]but most individuals spend their whole life cycle on a single host and are only able to transfer to a new host opportunistically.[7]Ischnoceran lice may reduce thethermoregulationeffect of the plumage; thus heavily infested birds lose more heat than others.[23] Lice infestation is a disadvantage in the context of sexual rivalry.[24][25]

Evolution

Phthiraptera lice are members ofPsocodea(formerlyPsocoptera), theorderthat contains booklice, barklice and barkflies. Within Psocodea, lice are within thesuborderTroctomorpha,and most closely related to the familyLiposcelididae.[26]The oldest confirmed fossil louse isArchimenopon myanmarensis,anamblyceranfrom the Cretaceous amber fromMyanmar.[4]Another early representative of the group is a bird louse,Megamenopon rasnitsyni,fromEckfelder Maar,Germany, which dates to theEocene,around 44 million years ago.[27]Saurodectes vrsanskyifrom the Early Cretaceous (Aptian)Zaza FormationofBuryatia,Russia, has also been suggested to be a louse, but this is tentative.[28]

Placental mammal lice had a single common ancestor that lived onAfrotheriawith this arising from host-switching from an ancient avian host.[29]

Cladogramshowing the position of Phthiraptera withinPsocodea:[1]

Psocodea

Classification

Phthiraptera is clearly amonophyleticgrouping, united as the members are by a number of derived features including their parasitism onwarm-bloodedvertebrates and the combination of theirmetathoracicgangliawith theirabdominal gangliato form a single ventral nerve junction.[30]The order has traditionally been divided into two suborders, the sucking lice (Anoplura) and thechewinglice (Mallophaga); however, subsequent classifications suggest that the Mallophaga areparaphyleticand four suborders were then recognized:[31]

  • Anoplura:suckinglice, occurring on mammals exclusively
  • Rhynchophthirina:parasites of elephants andwarthogs
  • Ischnocera:mostly avian chewing lice, with one family parasitizing mammals
  • Amblycera:a primitive suborder of chewing lice, widespread on birds, and also occurring on South American and Australian mammals

Upon finding that Phthiraptera was nested withinPsocoptera,Phthiraptera, in 2021 de Moya et al. proposed reducing the rank of Phthiraptera toinfraorder,and the foursuborderstoparvorder.[1]These changes were accepted by Psocodea Species File and others, with the exception of placing Phthiraptera under the infraorderNanopsocetae,as a parvorder, with the four subgroups listed above. These classifications are likely to change in the future as a result of ongoingphylogeneticresearch.[2][3]

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.[31]They are found even in theAntarctic,wherepenguinscarry 15 species of lice (in the generaAustrogonoidesandNesiotinus).[32]The oldest known record of the group isMegamenopon rasnitsynifrom the Eocene of Germany, but it is essentially a modern form, belonging to Amblycera, so the group as a whole likely has an origin in the Mesozoic.[27]

Phylogeny

Lice have been the subject of significantDNAresearch in the 2000s that led to discoveries on human evolution. The three species of sucking lice that parasitize human beings belong to two genera,PediculusandPthirus:head lice (Pediculus humanus capitis), body lice (Pediculus humanus humanus), and pubic lice (Pthirus pubis). Human head and body lice (genusPediculus) share a common ancestor with chimpanzee lice, while pubic lice (genusPthirus) share a common ancestor with gorilla lice. Using phylogenetic and cophylogenetic analysis, Reed et al. hypothesized thatPediculusandPthirusare sister taxa and monophyletic.[33]In other words, the two genera descended from the same common ancestor. The age of divergence betweenPediculusand its common ancestor is estimated to be 6-7 million years ago, which matches the age predicted bychimpanzee-hominid divergence.[33]Because parasites rely on their hosts, host–parasite cospeciation events are likely.

Genetic evidence suggests that human ancestors acquired pubic lice fromgorillasapproximately 3-4 million years ago.[33]Unlike the genusPediculus,the divergence inPthirusdoes not match the age of host divergence that likely occurred 7 million years ago. Reed et al. propose aPthirusspecies host-switch around 3-4 million years ago. While it is difficult to determine if a parasite–host switch occurred in evolutionary history, this explanation is the most parsimonious (containing the fewest evolutionary changes).[33]

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.[34]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.[34][35]

The mitochondrial genome of the human species of the body louse (Pediculus humanus humanus), the head louse (Pediculus humanus capitis) and the pubic louse (Pthirus pubis) fragmented into a number of minichromosomes, at least seven million years ago.[36]Analysis of mitochondrial DNA in human body and hair lice reveals that greater genetic diversity existed in African than in non-African lice.[35][37]Human lice can also shed light on human migratory patterns in prehistory. The dominating theory ofanthropologistsregarding human migration is theOut 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.[37]

In human culture

In social history

Drawing of a louse clinging to a human hair.Robert Hooke,Micrographia,1667

Lice have been intimately associated with human society throughout history. In theMiddle Ages,they were essentially ubiquitous. At the death ofThomas Becket,Archbishop of Canterburyin 1170, it was recorded that "The vermin boiled over like water in a simmering cauldron, and the onlookers burst into alternate weeping and laughing".[38]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.[39]A mediaeval treatment for lice was anointmentmade from pork grease,incense,lead,andaloe.[40]

Robert Hooke's 1667 book,Micrographia: or some physiological descriptions of minute bodies made by magnifying glasses with observations and Inquiries thereupon,illustrated a human louse, drawn as seen down an earlymicroscope.[41]

Margaret Cavendish's satiricalThe 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 scientistRobert Boyle.[42][43]

In 1935 the Harvard medical researcherHans Zinsserwrote the bookRats, Lice and History,alleging that both body and head lice transmit typhus between humans.[44]Despite this, the modern view is that only the body louse can transmit the disease.[45]

Detail showing delousing fromJan Siberechts' paintingCour de ferme( "Farmyard" ), 1662

Soldiers in the trenches of theFirst World Warsuffered severely from lice, and thetyphusthey carried. The Germans boasted that they had lice under effective control, but themselves suffered badly from lice in theSecond World Waron the Eastern Front, especially in theBattle of Stalingrad."Delousing"became aeuphemismfor the extermination of Jews inconcentration campssuch asAuschwitzunder the Nazi regime.[46]

In the psychiatric disorderdelusional 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".[47]

In science

The human body lousePediculus humanus humanushas (2010) the smallest insectgenomeknown.[48]This louse can transmit certain diseases while the human head louse (P. h. capitis), to which it is closely related, cannot. With their simple life history and small genomes, the pair make idealmodel organismsto study themolecular mechanismsbehind the transmission ofpathogensandvectorcompetence.[49]

In literature and folklore

Mother Louse, a notoriousalewifein Oxford during the mid-18th century, shown with three lice as acoat of arms.Image byDavid Loggan.[50][51]

James Joyce's 1939 bookFinnegans Wakehas the character Shem the Penman infested with "foxtrottingfleas, the lieabed lice,... bats in his belfry ".[52]

Clifford E. Trafzer'sAChemehueviSong: The Resilience of aSouthern PaiuteTriberetells the story of Sinawavi (Coyote)'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.[53]

The Irish songwriter John Lyons (b. 1934) wrote the popular[54]songThe 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 onFrom Puck to Appleby(2003), and theDublinersonDouble Dubliners(1972) among others.[54][55]

Robert Burnsdedicated 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."John MiltoninParadise Lostmentioned 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."John Rayrecorded a Scottish proverb, "Gie a beggar a bed and he'll repay you with a Louse." InShakespeare'sTroilus and Cressida,ThersitescomparesMenelaus,brother ofAgamemnon,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."[56]

Woodlouse

The namewoodlouseor wood-louse is given tocrustaceansof thesuborderOniscidea within theorderIsopoda,unrelated to lice. TheOxford English Dictionary's earliest citation of this usage is from 1611.[57]

See also

References

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