Louse: Difference between revisions

Phthiraptera; Temporal range: Cenomanian–present PreꞒ Ꞓ O S D C P T J K Pg N
	Light micrograph of Fahrenholzia pinnata
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Psocodea
Suborder:	Troctomorpha
Infraorder:	Nanopsocetae
Parvorder:	Phthiraptera; Haeckel, 1896
Clades
	Amblycera; Anoplura; Rhyncophthirina; Ischnocera; Trichodectera?;

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Louse (pl.: lice) is the common name for any member of the clade Phthiraptera, which contains nearly 5,000 species of wingless parasitic insects. Phthiraptera has variously been recognized as an order, infraorder, or a parvorder, as a result of developments in phylogenetic research.^[1]^[2]^[3]

Lice are obligate parasites, living externally on warm-blooded hosts which include every species of bird and mammal, except for monotremes, pangolins, and bats. Lice are vectors of diseases such as typhus.

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. Genetic evidence indicates that lice are a highly modified lineage of Psocoptera (now called Psocodea), commonly known as booklice, barklice or barkflies. The oldest known fossil lice are from the Cretaceous.^[4]

Humans host two species of louse—the head louse and the body louse are subspecies of Pediculus humanus; and the pubic louse, Pthirus pubis. The body louse has the smallest genome of any known insect; it has been used as a model organism and has been the subject of much research. Lice were ubiquitous in human society until at least the Middle Ages. They appear in folktales, songs such as The Kilkenny Louse House, and novels such as James Joyce's Finnegans Wake. They commonly feature in the psychiatric disorder delusional parasitosis. A louse was one of the early subjects of microscopy, appearing in Robert Hooke's 1667 book, Micrographia.

Morphology and diversity

Lice are divided into two groups: sucking lice, which obtain their nourishment from feeding on the sebaceous secretions and body fluids of their host; and chewing lice, which are scavengers, feeding on skin, 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 use phoresis to hitch a ride on a fly and attempt to find a new host.^[6]

Sucking lice range in length from 0.5 to 5 mm (1⁄64 to 13⁄64 in). They have narrow heads and oval, flattened bodies. They have no ocelli, 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.^[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 from 0.5 to 6 mm (1⁄64 to 15⁄64 in). 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 Ischnocera, but club-shaped in the suborder Amblycera. The legs are short and robust, and terminated by one or two claws. Some species of chewing lice house symbiotic bacteria in bacteriocytes in their bodies. These may assist in digestion because if the insect is deprived of them, it will die. Lice are usually cryptically coloured to 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 are parthenogenetic, with young developing from unfertilized eggs. A louse's egg is commonly called a nit. Many lice attach their eggs to their hosts' hair with specialized saliva; 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.^[5] Lice are exopterygotes, 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.^[5]

Humans host three different kinds of lice: head lice, body lice, and pubic lice. 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 lice 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 stronger T cell immune response or larger uropygial 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 into New Zealand host 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, but speciation has 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 in domestic hens is probably an attempt by the birds to rid themselves of lice.^[7] Lice may transmit microbial diseases and helminth parasites,^[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 the thermoregulation effect 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 of Psocodea (formerly Psocoptera), the order that contains booklice, barklice and barkflies. Within Psocodea, lice are within the suborder Troctomorpha, and most closely related to the family Liposcelididae.^[26] The oldest confirmed fossil louse is Archimenopon myanmarensis, an amblyceran from the Cretaceous amber from Myanmar.^[4] Another early representative of the group is a bird louse, Megamenopon rasnitsyni, from Eckfelder Maar, Germany, which dates to the Eocene, around 44 million years ago.^[27] Saurodectes vrsanskyi from the Early Cretaceous (Aptian) Zaza Formation of Buryatia, Russia, has also been suggested to be a louse, but this is tentative.^[28]

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

Cladogram showing the position of Phthiraptera within Psocodea:^[1]

Classification

Phthiraptera is clearly a monophyletic grouping, united as the members are by a number of derived features including their parasitism on warm-blooded vertebrates and the combination of their metathoracic ganglia with their abdominal ganglia to form a single ventral nerve junction.^[30] The order has traditionally been divided into two suborders, the sucking lice (Anoplura) and the chewing lice (Mallophaga); however, subsequent classifications suggest that the Mallophaga are paraphyletic and four suborders were then recognized:^[31]

Anoplura: sucking lice, occurring on mammals exclusively
Rhynchophthirina: parasites of elephants and warthogs
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 within Psocoptera, Phthiraptera, in 2021 de Moya et al. proposed reducing the rank of Phthiraptera to infraorder, and the four suborders to parvorder.^[1] These changes were accepted by Psocodea Species File and others, with the exception of placing Phthiraptera under the infraorder Nanopsocetae, as a parvorder, with the four subgroups listed above. These classifications are likely to change in the future as a result of ongoing phylogenetic research.^[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 the Antarctic, where penguins carry 15 species of lice (in the genera Austrogonoides and Nesiotinus).^[32] The oldest known record of the group is Megamenopon rasnitsyni from 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]

Ricinus bombycillae, an amblyceran louse from a Bohemian waxwing
Trinoton anserinum, an amblyceran louse from a mute swan
Bovicola limbata, an ischnoceran louse from goats. The species is sexually dimorphic, with the male smaller than the female.

Phylogeny

Lice have been the subject of significant DNA 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, Pediculus and Pthirus: head lice (Pediculus humanus capitis), body lice (Pediculus humanus humanus), and pubic lice (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.^[33] 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.^[33] Because parasites rely on their hosts, host–parasite cospeciation events are likely.

Genetic evidence suggests that human ancestors acquired pubic lice from gorillas approximately 3-4 million years ago.^[33] 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 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 of anthropologists 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.^[37]

In human culture

In social history

Lice have been intimately associated with human society throughout history. In the Middle Ages, they were essentially ubiquitous. At the death of Thomas Becket, 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".^[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 an ointment made from pork grease, incense, lead, and aloe.^[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 early microscope.^[41]

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 Robert Boyle.^[42]^[43]

In 1935 the Harvard medical researcher Hans Zinsser wrote the book Rats, 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]

Soldiers in the trenches of the First World War suffered severely from lice, and the typhus 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. "Delousing" became a euphemism for the extermination of Jews in concentration camps such as Auschwitz under the Nazi regime.^[46]

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".^[47]

In science

The human body louse Pediculus humanus humanus has (2010) the smallest insect genome known.^[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 ideal model organisms to study the molecular mechanisms behind the transmission of pathogens and vector competence.^[49]

In literature and folklore

James Joyce's 1939 book Finnegans Wake has the character Shem the Penman infested with "foxtrotting fleas, the lieabed lice, ... bats in his belfry".^[52]

Clifford E. Trafzer's A Chemehuevi Song: The Resilience of a Southern Paiute Tribe retells 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] 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 Dubliners on Double Dubliners (1972) among others.^[54]^[55]

Robert Burns 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." John Milton in Paradise Lost 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." John Ray recorded a Scottish proverb, "Gie a beggar a bed and he'll repay you with a Louse." In Shakespeare's Troilus and Cressida, Thersites compares Menelaus, brother of Agamemnon, 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 name woodlouse or wood-louse is given to crustaceans of the suborder Oniscidea within the order Isopoda, unrelated to lice. The Oxford English Dictionary's earliest citation of this usage is from 1611.^[57]

References

^ ^a ^b ^c ^d de Moya, Robert S; Yoshizawa, Kazunori; Walden, Kimberly K O; Sweet, Andrew D; Dietrich, Christopher H; Kevin P, Johnson (2021-06-16). Buckley, Thomas (ed.). "Phylogenomics of Parasitic and Nonparasitic Lice (Insecta: Psocodea): Combining Sequence Data and Exploring Compositional Bias Solutions in Next Generation Data Sets". Systematic Biology. 70 (4): 719–738. doi:10.1093/sysbio/syaa075. ISSN 1063-5157. PMID 32979270.
^ ^a ^b Johnson, Kevin P.; Smith, Vincent S. (2021). "Psocodea species file online, Version 5.0". Retrieved 2021-11-03.
^ ^a ^b "Catalogue of Life, Phthiraptera Haeckel, 1896". 2021. Archived from the original on 2021-11-05. Retrieved 2021-11-03.
^ ^a ^b Zhang, Y.; Rasnitsyn, A. P.; Zhang, W.; Song, F.; Shih, C.; Ren, D.; Wang, Y.; Li, H.; Gao, T. (2024). "Stem chewing lice on Cretaceous feathers preserved in amber". Current Biology. 34 (4): 916–922.e1. Bibcode:2024CBio...34E.916Z. doi:10.1016/j.cub.2024.01.027. PMID 38320551.
^ ^a ^b ^c Hoell HV, Doyen JT, Purcell AH (1998). Introduction to Insect Biology and Diversity (2nd ed.). Oxford University Press. pp. 407–409. ISBN 978-0-19-510033-4.
^ Harbison CW (2008). Ecology and Evolution of Transmission in Feather-feeding Lice (Phthiraptera: Ischnocera). Department of Biology (Doctoral thesis). University of Utah. pp. 83–87. ISBN 978-0-549-46429-7.
^ ^a ^b ^c ^d ^e Capinera JL (2008). Encyclopedia of Entomology. Springer Science & Business Media. pp. 838–844. ISBN 978-1-4020-6242-1.
^ Gullan PJ, Cranston PS (2014). The Insects: An Outline of Entomology. Wiley. pp. 41–42. ISBN 978-1-118-84615-5.
^ Smith V. "Phthiraptera: Summary". Phthiraptera.info. Retrieved 25 October 2015.^{[permanent dead link‍]}
^ Mumcuoglu KY (1999). "Prevention and treatment of head lice in children". Paediatric Drugs. 1 (3): 211–8. doi:10.2165/00128072-199901030-00005. PMID 10937452. S2CID 13547569.
^ Rózsa (1997). "Patterns in the abundance of avian lice (Phthiraptera: Amblycera, Ischnocera)" (PDF). Journal of Avian Biology. 28 (3): 249–254. doi:10.2307/3676976. JSTOR 3676976.
^ Rékási J, Rozsa L, Kiss BJ (1997). "Patterns in the distribution of avian lice (Phthiraptera: Amblycera, Ischnocera)" (PDF). Journal of Avian Biology. 28 (2): 150–156. CiteSeerX 10.1.1.506.730. doi:10.2307/3677308. JSTOR 3677308.
^ Felsõ B, Rózsa L (August 2006). "Reduced taxonomic richness of lice (Insecta: Phthiraptera) in diving birds". The Journal of Parasitology. 92 (4): 867–9. doi:10.1645/ge-849.1. PMID 16995408. S2CID 31000581.
^ Felső B, Rózsa L (2007). "Diving behaviour reduces genera richness of lice (Insecta: Phthiraptera) of mammals" (PDF). Acta Parasitologica. 52: 82–85. doi:10.2478/s11686-007-0006-3. S2CID 38683871.
^ Møller AP, Rózsa L (January 2005). "Parasite biodiversity and host defenses: chewing lice and immune response of their avian hosts". Oecologia. 142 (2): 169–76. Bibcode:2005Oecol.142..169M. doi:10.1007/s00442-004-1735-8. PMID 15503162. S2CID 12992951.
^ Møller AP, Erritzøe J, Rózsa L (June 2010). "Ectoparasites, uropygial glands and hatching success in birds". Oecologia. 163 (2): 303–11. Bibcode:2010Oecol.163..303M. doi:10.1007/s00442-009-1548-x. PMID 20043177. S2CID 11433594.
^ ^a ^b Ròzsa L (November 1993). "Speciation patterns of ectoparasites and "straggling" lice". International Journal for Parasitology. 23 (7): 859–64. doi:10.1016/0020-7519(93)90050-9. PMID 8314369.
^ Paterson AM, Palma RL, Gray RD (1999). "How Frequently Do Avian Lice Miss the Boat? Implications for Coevolutionary Studies". Systematic Biology. 48: 214–223. doi:10.1080/106351599260544.
^ MacLeod CJ, Paterson AM, Tompkins DM, Duncan RP (April 2010). "Parasites lost - do invaders miss the boat or drown on arrival?". Ecology Letters. 13 (4): 516–27. Bibcode:2010EcolL..13..516M. doi:10.1111/j.1461-0248.2010.01446.x. PMID 20455925.
^ Rózsa L, Rékási J, Reiczigel J (1996). "Relationship of host coloniality to the population ecology of avian lice (Insecta: Phthiraptera)" (PDF). Journal of Animal Ecology. 65 (2): 242–248. Bibcode:1996JAnEc..65..242R. doi:10.2307/5727. JSTOR 5727.
^ Brown CR, Brown MB, Rannala B (1995). "Ectoparasites reduce long-term survivorship of their avian host" (PDF). Proceedings of the Royal Society of London B. 262 (1365): 313–319. doi:10.1098/rspb.1995.0211. S2CID 13869042.
^ Barlett CM (1993). "Lice (Amblycera and Ischnocera) as vectors of Eulimdana spp. (Nematoda: Filarioidea) in Charadriiform birds and the necessity of short reproductive periods in adult worms". Journal of Parasitology. 75 (1): 85–91. doi:10.2307/3283282. JSTOR 3283282.
^ Booth DT, Clayton DH, Block BA (1993). "Experimental demonstration of the energetic cost of parasitism in free-ranging hosts" (PDF). Proceedings of the Royal Society of London B. 253 (1337): 125–129. Bibcode:1993RSPSB.253..125B. doi:10.1098/rspb.1993.0091. S2CID 85731062. Archived from the original (PDF) on 2010-06-10. Retrieved 2010-08-14.
^ Clayton (1990). "Mate choice in experimentally parasitized rock doves: lousy males lose" (PDF). American Zoologist. 30 (2): 251–262. doi:10.1093/icb/30.2.251. Archived from the original (PDF) on 2010-06-10. Retrieved 2010-08-14.
^ Garamszegi LZ, Heylen D, Møller AP, Eens M, De Lope F (2005). "Age-dependent health status and song characteristics". Behavioral Ecology. 16 (3): 580–591. doi:10.1093/beheco/ari029.
^ Light JE, Smith VS, Allen JM, Durden LA, Reed DL (September 2010). "Evolutionary history of mammalian sucking lice (Phthiraptera: Anoplura)". BMC Evolutionary Biology. 10 (1): 292. Bibcode:2010BMCEE..10..292L. doi:10.1186/1471-2148-10-292. PMC 2949877. PMID 20860811.
^ ^a ^b Wappler T, Smith VS, Dalgleish RC (August 2004). "Scratching an ancient itch: an Eocene bird louse fossil". Proceedings. Biological Sciences. 271 Suppl 5 (suppl_5): S255-8. doi:10.1098/rsbl.2003.0158. PMC 1810061. PMID 15503987.
^ Rasnitsyn AP, Zherikhin VV (1999). "First fossil chewing louse from the lower Cretaceous of Baissa, Transbaikalia (Insecta, Pediculida= Phthiriaptera, Saurodectidae fam. n.)". Russian Entomological Journal. 8 (4): 253–5.
^ Johnson, Kevin P.; Matthee, Conrad; Doña, Jorge (2022). "Phylogenomics reveals the origin of mammal lice out of Afrotheria". Nature Ecology & Evolution. 6 (8): 1205–1210. Bibcode:2022NatEE...6.1205J. doi:10.1038/s41559-022-01803-1. ISSN 2397-334X. PMID 35788706.
^ Ax P (2013). Multicellular Animals: Volume II: The Phylogenetic System of the Metazoa. Springer Science & Business Media. pp. 303–307. ISBN 978-3-662-10396-8.
^ ^a ^b Smith V. "Phthiraptera.info". International Society of Phthirapterists. Retrieved 25 October 2015.
^ Banks JC, Paterson AM (2004). "A penguin-chewing louse (Insecta : Phthiraptera) phylogeny derived from morphology". Invertebrate Systematics. 18 (1): 89–100. doi:10.1071/IS03022. S2CID 53516244.
^ ^a ^b ^c ^d Reed DL, Light JE, Allen JM, Kirchman JJ (March 2007). "Pair of lice lost or parasites regained: the evolutionary history of anthropoid primate lice". BMC Biology. 5 (7): 7. doi:10.1186/1741-7007-5-7. PMC 1828715. PMID 17343749.
^ ^a ^b Parry W (7 November 2013). "Lice Reveal Clues to Human Evolution". LiveScience. Retrieved 25 October 2015.
^ ^a ^b Kittler R, Kayser M, Stoneking M (August 2003). "Molecular evolution of Pediculus humanus and the origin of clothing". Current Biology. 13 (16): 1414–7. Bibcode:2003CBio...13.1414K. doi:10.1016/S0960-9822(03)00507-4. PMID 12932325. S2CID 15277254.
^ Shao R, Zhu XQ, Barker SC, Herd K (2012). "Evolution of extensively fragmented mitochondrial genomes in the lice of humans". Genome Biology and Evolution. 4 (11): 1088–101. doi:10.1093/gbe/evs088. PMC 3514963. PMID 23042553.
^ ^a ^b Light JE, Allen JM, Long LM, Carter TE, Barrow L, Suren G, et al. (December 2008). "Geographic distributions and origins of human head lice (Pediculus humanus capitis) based on mitochondrial data". The Journal of Parasitology. 94 (6): 1275–81. doi:10.1645/GE-1618.1. PMID 18576877. S2CID 9456340.
^ Kowalski TJ, Agger WA (January 2009). "Art supports new plague science". Clinical Infectious Diseases. 48 (1): 137–8. doi:10.1086/595557. PMID 19067623.
^ Harvey, Katherine (2019-04-09). "Medieval people were surprisingly clean (apart from the clergy)". Aeon. Retrieved 2022-10-13.
^ Elliott L (2004). Clothing in the Middle Ages. Crabtree. p. 29. ISBN 978-0-7787-1351-7.
^ Hooke R. "Microscopic view of a louse". The Royal Society. Retrieved 17 October 2015.
^ Sarasohn LT (2010). The Natural Philosophy of Margaret Cavendish: Reason and Fancy During the Scientific Revolution. JHU Press. pp. 165–167. ISBN 978-0-8018-9443-5. The Bear-men were to be her Experimental Philosophers, the Bird-men her Astronomers, the Fly- Worm- and Fish-men her Natural Philosophers, the Ape-men her Chymists, the Satyrs her Galenick Physicians, the Fox-men her Politicians, the Spider- and Lice-men her Mathematicians, the Jackdaw- Magpie- and Parrot-men her Orators and Logicians, the Gyants her Architects, &c.
^ Cavendish M (1668). The Description of a New World, Called the Blazing-World. A. Maxwell.
^ Zinsser H (2007) [1935]. Rats, Lice and History. Transaction Publishers. ISBN 978-1-4128-0672-5.
^ Altschuler DZ (1990). "Zinsser, Lice and History". HeadLice.org. Retrieved 19 October 2015.
^ Evans RJ. "The Great Unwashed". Gresham College. Retrieved 17 October 2015.
^ Weinstein P (26 February 2013). "'The Great Unwashed': Entomophobia/Delusionary Parasitosis; Illusionary Parasitosis". The Great Plagues: Epidemics in History from the Middle Ages to the Present Day. University of Sydney Department of Medical Entomology. Archived from the original on 17 May 2016. Retrieved 17 October 2015.
^ Kirkness EF, Haas BJ, Sun W, Braig HR, Perotti MA, Clark JM, et al. (July 2010). "Genome sequences of the human body louse and its primary endosymbiont provide insights into the permanent parasitic lifestyle". Proceedings of the National Academy of Sciences of the United States of America. 107 (27): 12168–73. Bibcode:2010PNAS..10712168K. doi:10.1073/pnas.1003379107. PMC 2901460. PMID 20566863.
^ Pittendrigh BR, Berenbaum MR, Seufferheld MJ, Margam VM, Strycharz JP, Yoon KS, et al. (March 2011). "Simplify, simplify: Lifestyle and compact genome of the body louse provide a unique functional genomics opportunity". Communicative & Integrative Biology. 4 (2): 188–91. doi:10.4161/cib.4.2.14279. PMC 3104575. PMID 21655436.
^ White W (1859). Notes & Queries. Oxford University Press. pp. 275–276.
^ Pierce H (2004). "Unseemly pictures: political graphic satire in England, c. 1600-c. 1650" (PDF). University of York.
^ Joyce J (1939). Finnegans Wake. Faber. p. 180.
^ Trafzer CE (2015). A Chemehuevi Song: The Resilience of a Southern Paiute Tribe. University of Washington Press. pp. 22–25. ISBN 978-0-295-80582-5.
^ ^a ^b Carroll J. "Songs of Clare: The Kilkenny Louse House". Clare Library. Retrieved 19 October 2015.
^ Scott B (2013). "My Colleen by the Shore" (PDF). Veteran. Retrieved 25 October 2015.
^ Twinn CR (1942). Insect Life in the Poetry and Drama of England: With Special Reference to Poetry (PhD thesis). University of Ottawa. hdl:10393/21088.
^ "Woodlouse". Oxford English Dictionary (Online ed.). Oxford University Press. (Subscription or participating institution membership required.)

External links

National Pesticide Information Center – Understanding and Controlling Lice
body and head lice on the University of Florida/Institute of Food and Agricultural Sciences Featured Creatures Web site
crab louse on the University of Florida/Institute of Food and Agricultural Sciences Featured Creatures Web site
Pediculus humanus capitis head louse facts, myths, life cycle at MetaPathogen
Parasitic Insects, Mites and Ticks: Genera of Medical and Veterinary Importance Wikibooks

[deMoya2021-1] Moya, Robert S; Yoshizawa, Kazunori; Walden, Kimberly K O; Sweet, Andrew D; Dietrich, Christopher H; Kevin P, Johnson (2021-06-16). Buckley, Thomas (ed.). "Phylogenomics of Parasitic and Nonparasitic Lice (Insecta: Psocodea): Combining Sequence Data and Exploring Compositional Bias Solutions in Next Generation Data Sets". Systematic Biology. 70 (4): 719–738. doi:10.1093/sysbio/syaa075. ISSN 1063-5157. PMID 32979270.

[Johnson2021-2] Johnson, Kevin P.; Smith, Vincent S. (2021). "Psocodea species file online, Version 5.0". Retrieved 2021-11-03.

[catlife-3] "Catalogue of Life, Phthiraptera Haeckel, 1896". 2021. Archived from the original on 2021-11-05. Retrieved 2021-11-03.

[Archimenopon-4] Zhang, Y.; Rasnitsyn, A. P.; Zhang, W.; Song, F.; Shih, C.; Ren, D.; Wang, Y.; Li, H.; Gao, T. (2024). "Stem chewing lice on Cretaceous feathers preserved in amber". Current Biology. 34 (4): 916–922.e1. Bibcode:2024CBio...34E.916Z. doi:10.1016/j.cub.2024.01.027. PMID 38320551.

[IIBD-5] Hoell HV, Doyen JT, Purcell AH (1998). Introduction to Insect Biology and Diversity (2nd ed.). Oxford University Press. pp. 407–409. ISBN 978-0-19-510033-4.

[6] Harbison CW (2008). Ecology and Evolution of Transmission in Feather-feeding Lice (Phthiraptera: Ischnocera). Department of Biology (Doctoral thesis). University of Utah. pp. 83–87. ISBN 978-0-549-46429-7.

[Capinera-7] Capinera JL (2008). Encyclopedia of Entomology. Springer Science & Business Media. pp. 838–844. ISBN 978-1-4020-6242-1.

[GullanCranston-8] Gullan PJ, Cranston PS (2014). The Insects: An Outline of Entomology. Wiley. pp. 41–42. ISBN 978-1-118-84615-5.

[9] Smith V. "Phthiraptera: Summary". Phthiraptera.info. Retrieved 25 October 2015.^{[permanent dead link‍]}

[10] Mumcuoglu KY (1999). "Prevention and treatment of head lice in children". Paediatric Drugs. 1 (3): 211–8. doi:10.2165/00128072-199901030-00005. PMID 10937452. S2CID 13547569.

[11] Rózsa (1997). "Patterns in the abundance of avian lice (Phthiraptera: Amblycera, Ischnocera)" (PDF). Journal of Avian Biology. 28 (3): 249–254. doi:10.2307/3676976. JSTOR 3676976.

[12] Rékási J, Rozsa L, Kiss BJ (1997). "Patterns in the distribution of avian lice (Phthiraptera: Amblycera, Ischnocera)" (PDF). Journal of Avian Biology. 28 (2): 150–156. CiteSeerX 10.1.1.506.730. doi:10.2307/3677308. JSTOR 3677308.

[13] Felsõ B, Rózsa L (August 2006). "Reduced taxonomic richness of lice (Insecta: Phthiraptera) in diving birds". The Journal of Parasitology. 92 (4): 867–9. doi:10.1645/ge-849.1. PMID 16995408. S2CID 31000581.

[14] Felső B, Rózsa L (2007). "Diving behaviour reduces genera richness of lice (Insecta: Phthiraptera) of mammals" (PDF). Acta Parasitologica. 52: 82–85. doi:10.2478/s11686-007-0006-3. S2CID 38683871.

[15] Møller AP, Rózsa L (January 2005). "Parasite biodiversity and host defenses: chewing lice and immune response of their avian hosts". Oecologia. 142 (2): 169–76. Bibcode:2005Oecol.142..169M. doi:10.1007/s00442-004-1735-8. PMID 15503162. S2CID 12992951.

[16] Møller AP, Erritzøe J, Rózsa L (June 2010). "Ectoparasites, uropygial glands and hatching success in birds". Oecologia. 163 (2): 303–11. Bibcode:2010Oecol.163..303M. doi:10.1007/s00442-009-1548-x. PMID 20043177. S2CID 11433594.

[Rozsa1993-17] Ròzsa L (November 1993). "Speciation patterns of ectoparasites and "straggling" lice". International Journal for Parasitology. 23 (7): 859–64. doi:10.1016/0020-7519(93)90050-9. PMID 8314369.

[18] Paterson AM, Palma RL, Gray RD (1999). "How Frequently Do Avian Lice Miss the Boat? Implications for Coevolutionary Studies". Systematic Biology. 48: 214–223. doi:10.1080/106351599260544.

[19] MacLeod CJ, Paterson AM, Tompkins DM, Duncan RP (April 2010). "Parasites lost - do invaders miss the boat or drown on arrival?". Ecology Letters. 13 (4): 516–27. Bibcode:2010EcolL..13..516M. doi:10.1111/j.1461-0248.2010.01446.x. PMID 20455925.

[20] Rózsa L, Rékási J, Reiczigel J (1996). "Relationship of host coloniality to the population ecology of avian lice (Insecta: Phthiraptera)" (PDF). Journal of Animal Ecology. 65 (2): 242–248. Bibcode:1996JAnEc..65..242R. doi:10.2307/5727. JSTOR 5727.

[21] Brown CR, Brown MB, Rannala B (1995). "Ectoparasites reduce long-term survivorship of their avian host" (PDF). Proceedings of the Royal Society of London B. 262 (1365): 313–319. doi:10.1098/rspb.1995.0211. S2CID 13869042.

[22] Barlett CM (1993). "Lice (Amblycera and Ischnocera) as vectors of Eulimdana spp. (Nematoda: Filarioidea) in Charadriiform birds and the necessity of short reproductive periods in adult worms". Journal of Parasitology. 75 (1): 85–91. doi:10.2307/3283282. JSTOR 3283282.

[23] Booth DT, Clayton DH, Block BA (1993). "Experimental demonstration of the energetic cost of parasitism in free-ranging hosts" (PDF). Proceedings of the Royal Society of London B. 253 (1337): 125–129. Bibcode:1993RSPSB.253..125B. doi:10.1098/rspb.1993.0091. S2CID 85731062. Archived from the original (PDF) on 2010-06-10. Retrieved 2010-08-14.

[24] Clayton (1990). "Mate choice in experimentally parasitized rock doves: lousy males lose" (PDF). American Zoologist. 30 (2): 251–262. doi:10.1093/icb/30.2.251. Archived from the original (PDF) on 2010-06-10. Retrieved 2010-08-14.

[25] Garamszegi LZ, Heylen D, Møller AP, Eens M, De Lope F (2005). "Age-dependent health status and song characteristics". Behavioral Ecology. 16 (3): 580–591. doi:10.1093/beheco/ari029.

[26] Light JE, Smith VS, Allen JM, Durden LA, Reed DL (September 2010). "Evolutionary history of mammalian sucking lice (Phthiraptera: Anoplura)". BMC Evolutionary Biology. 10 (1): 292. Bibcode:2010BMCEE..10..292L. doi:10.1186/1471-2148-10-292. PMC 2949877. PMID 20860811.

[Scratching_an_ancient_itch:_an_Eoce-27] Wappler T, Smith VS, Dalgleish RC (August 2004). "Scratching an ancient itch: an Eocene bird louse fossil". Proceedings. Biological Sciences. 271 Suppl 5 (suppl_5): S255-8. doi:10.1098/rsbl.2003.0158. PMC 1810061. PMID 15503987.

[28] Rasnitsyn AP, Zherikhin VV (1999). "First fossil chewing louse from the lower Cretaceous of Baissa, Transbaikalia (Insecta, Pediculida= Phthiriaptera, Saurodectidae fam. n.)". Russian Entomological Journal. 8 (4): 253–5.

[p273-29] Johnson, Kevin P.; Matthee, Conrad; Doña, Jorge (2022). "Phylogenomics reveals the origin of mammal lice out of Afrotheria". Nature Ecology & Evolution. 6 (8): 1205–1210. Bibcode:2022NatEE...6.1205J. doi:10.1038/s41559-022-01803-1. ISSN 2397-334X. PMID 35788706.

[Ax-30] Ax P (2013). Multicellular Animals: Volume II: The Phylogenetic System of the Metazoa. Springer Science & Business Media. pp. 303–307. ISBN 978-3-662-10396-8.

[info-31] Smith V. "Phthiraptera.info". International Society of Phthirapterists. Retrieved 25 October 2015.

[32] Banks JC, Paterson AM (2004). "A penguin-chewing louse (Insecta : Phthiraptera) phylogeny derived from morphology". Invertebrate Systematics. 18 (1): 89–100. doi:10.1071/IS03022. S2CID 53516244.

[Reed_D.L._2007-33] Reed DL, Light JE, Allen JM, Kirchman JJ (March 2007). "Pair of lice lost or parasites regained: the evolutionary history of anthropoid primate lice". BMC Biology. 5 (7): 7. doi:10.1186/1741-7007-5-7. PMC 1828715. PMID 17343749.

[Parry-34] Parry W (7 November 2013). "Lice Reveal Clues to Human Evolution". LiveScience. Retrieved 25 October 2015.

[Kittler_R._2003-35] Kittler R, Kayser M, Stoneking M (August 2003). "Molecular evolution of Pediculus humanus and the origin of clothing". Current Biology. 13 (16): 1414–7. Bibcode:2003CBio...13.1414K. doi:10.1016/S0960-9822(03)00507-4. PMID 12932325. S2CID 15277254.

[Shao2012-36] Shao R, Zhu XQ, Barker SC, Herd K (2012). "Evolution of extensively fragmented mitochondrial genomes in the lice of humans". Genome Biology and Evolution. 4 (11): 1088–101. doi:10.1093/gbe/evs088. PMC 3514963. PMID 23042553.

[Light-37] Light JE, Allen JM, Long LM, Carter TE, Barrow L, Suren G, et al. (December 2008). "Geographic distributions and origins of human head lice (Pediculus humanus capitis) based on mitochondrial data". The Journal of Parasitology. 94 (6): 1275–81. doi:10.1645/GE-1618.1. PMID 18576877. S2CID 9456340.

[Kowalski-38] Kowalski TJ, Agger WA (January 2009). "Art supports new plague science". Clinical Infectious Diseases. 48 (1): 137–8. doi:10.1086/595557. PMID 19067623.

[39] Harvey, Katherine (2019-04-09). "Medieval people were surprisingly clean (apart from the clergy)". Aeon. Retrieved 2022-10-13.

[40] Elliott L (2004). Clothing in the Middle Ages. Crabtree. p. 29. ISBN 978-0-7787-1351-7.

[41] Hooke R. "Microscopic view of a louse". The Royal Society. Retrieved 17 October 2015.

[Sarasohn2010-42] Sarasohn LT (2010). The Natural Philosophy of Margaret Cavendish: Reason and Fancy During the Scientific Revolution. JHU Press. pp. 165–167. ISBN 978-0-8018-9443-5. The Bear-men were to be her Experimental Philosophers, the Bird-men her Astronomers, the Fly- Worm- and Fish-men her Natural Philosophers, the Ape-men her Chymists, the Satyrs her Galenick Physicians, the Fox-men her Politicians, the Spider- and Lice-men her Mathematicians, the Jackdaw- Magpie- and Parrot-men her Orators and Logicians, the Gyants her Architects, &c.

[43] Cavendish M (1668). The Description of a New World, Called the Blazing-World. A. Maxwell.

[44] Zinsser H (2007) [1935]. Rats, Lice and History. Transaction Publishers. ISBN 978-1-4128-0672-5.

[45] Altschuler DZ (1990). "Zinsser, Lice and History". HeadLice.org. Retrieved 19 October 2015.

[46] Evans RJ. "The Great Unwashed". Gresham College. Retrieved 17 October 2015.

[47] Weinstein P (26 February 2013). "'The Great Unwashed': Entomophobia/Delusionary Parasitosis; Illusionary Parasitosis". The Great Plagues: Epidemics in History from the Middle Ages to the Present Day. University of Sydney Department of Medical Entomology. Archived from the original on 17 May 2016. Retrieved 17 October 2015.

[48] Kirkness EF, Haas BJ, Sun W, Braig HR, Perotti MA, Clark JM, et al. (July 2010). "Genome sequences of the human body louse and its primary endosymbiont provide insights into the permanent parasitic lifestyle". Proceedings of the National Academy of Sciences of the United States of America. 107 (27): 12168–73. Bibcode:2010PNAS..10712168K. doi:10.1073/pnas.1003379107. PMC 2901460. PMID 20566863.

[49] Pittendrigh BR, Berenbaum MR, Seufferheld MJ, Margam VM, Strycharz JP, Yoon KS, et al. (March 2011). "Simplify, simplify: Lifestyle and compact genome of the body louse provide a unique functional genomics opportunity". Communicative & Integrative Biology. 4 (2): 188–91. doi:10.4161/cib.4.2.14279. PMC 3104575. PMID 21655436.

[White1859-50] White W (1859). Notes & Queries. Oxford University Press. pp. 275–276.

[51] Pierce H (2004). "Unseemly pictures: political graphic satire in England, c. 1600-c. 1650" (PDF). University of York.

[52] Joyce J (1939). Finnegans Wake. Faber. p. 180.

[Trafzer2015-53] Trafzer CE (2015). A Chemehuevi Song: The Resilience of a Southern Paiute Tribe. University of Washington Press. pp. 22–25. ISBN 978-0-295-80582-5.

[Clare-54] Carroll J. "Songs of Clare: The Kilkenny Louse House". Clare Library. Retrieved 19 October 2015.

[55] Scott B (2013). "My Colleen by the Shore" (PDF). Veteran. Retrieved 25 October 2015.

[56] Twinn CR (1942). Insect Life in the Poetry and Drama of England: With Special Reference to Poetry (PhD thesis). University of Ottawa. hdl:10393/21088.

[57] "Woodlouse". Oxford English Dictionary (Online ed.). Oxford University Press. (Subscription or participating institution membership required.)

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[15]

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[21]

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@@ Line 7: / Line 7: @@
 {{Automatic taxobox
 | name = Phthiraptera
-| fossil_range = {{fossilrange|Eocene|present|earliest=Cretaceous}}
+| fossil_range = {{fossilrange|Cenomanian|present|earliest=Cretaceous}}
 | image = Fahrenholzia pinnata.JPG
-| image_caption = [[micrograph|Light micrograph]] of ''[[Fahrenholzia pinnata]]''
+| image_caption = [[Light micrograph]] of ''[[Fahrenholzia pinnata]]''
 | taxon = Phthiraptera
-| authority = [[Ernst Haeckel|Haeckel]], 1896
+| authority = [[Haeckel]], 1896
 | display_parents = 2
 | subdivision_ranks = Clades
@@ Line 17: / Line 17: @@
 *[[Amblycera]]
 *[[Anoplura]]
-*[[Haematomyzus|Rhyncophthirina]]
+*[[Rhyncophthirina]]
 *[[Ischnocera]]
 *[[Trichodectera]]?
 | subdivision_ref = <ref name="deMoya2021"/>
 }}
-'''Louse''' ({{plural form}}: '''lice''') is the common name for any member of the [[clade]] '''Phthiraptera''', which contains nearly 5,000 species of wingless [[Parasitism|parasitic]] [[insect]]s. Phthiraptera has variously been recognized as an [[order (biology)|order]], [[infraorder]], or a [[parvorder]], as a result of developments in phylogenetic research.<ref name="deMoya2021"/><ref name=Johnson2021/><ref name=catlife/>
+'''Louse''' ({{plural form}}: '''lice''') is the common name for any member of the [[clade]] '''Phthiraptera''', which contains nearly 5,000 species of wingless [[parasitic]] [[insect]]s. Phthiraptera has variously been recognized as an [[order (biology)|order]], [[infraorder]], or a [[parvorder]], as a result of developments in phylogenetic research.<ref name="deMoya2021"/><ref name=Johnson2021/><ref name=catlife/>
 Lice are [[obligate parasite]]s, living externally on warm-blooded [[Host (biology)|hosts]] which include every species of [[bird]] and [[mammal]], except for [[monotreme]]s, [[pangolin]]s, and [[bat]]s. Lice are [[Vector (epidemiology)|vectors]] of diseases such as [[typhus]].
-Chewing lice live among the hairs or feathers of their host and feed on skin and debris, while 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 '''[[Head louse#Eggs/Nits|nits]]''', to hairs or feathers. The eggs hatch into [[Nymph (biology)|nymphs]], which moult three times before becoming fully grown, a process that takes about four weeks.
+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 '''[[Head louse#Eggs/Nits|nits]]''', to hairs or feathers. The eggs hatch into [[Nymph (biology)|nymphs]], 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 of [[Psocoptera]] (now called [[Psocodea]]), commonly known as booklice, barklice or barkflies. The oldest known fossil lice are from the [[Paleogene]], though [[molecular clock]] estimates suggest that they originated earlier, during the [[Cretaceous]].
+Genetic evidence indicates that lice are a highly modified lineage of [[Psocoptera]] (now called [[Psocodea]]), commonly known as booklice, barklice or barkflies. The oldest known fossil lice are from the [[Cretaceous]].<ref name=Archimenopon>{{Cite journal|last1=Zhang |first1=Y. |last2=Rasnitsyn |first2=A. P. |last3=Zhang |first3=W. |last4=Song |first4=F. |last5=Shih |first5=C. |last6=Ren |first6=D. |last7=Wang |first7=Y. |last8=Li |first8=H. |last9=Gao |first9=T. |year=2024 |title=Stem chewing lice on Cretaceous feathers preserved in amber |journal=Current Biology |volume=34 |issue=4 |pages=916–922.e1 |doi=10.1016/j.cub.2024.01.027 |pmid=38320551 |bibcode=2024CBio...34E.916Z }}</ref>
-Humans host two species of louse—the [[head louse]] and the [[body louse]] are subspecies of ''[[Pediculus humanus]]''; and the [[Crab louse|pubic louse]], ''[[Pthirus]] pubis''. The body louse has the smallest [[genome]] of any known insect; it has been used as a [[model organism]] and has been the subject of much research.
+Humans host two species of louse—the [[head louse]] and the [[body louse]] are subspecies of ''[[Pediculus humanus]]''; and the [[pubic louse]], ''[[Pthirus]] pubis''. The body louse has the smallest [[genome]] of any known insect; it has been used as a [[model organism]] and has been the subject of much research.
 Lice were ubiquitous in human society until at least the [[Middle Ages]]. They appear in folktales, songs such as ''The Kilkenny Louse House'', and novels such as [[James Joyce]]'s ''[[Finnegans Wake]]''. They commonly feature in the psychiatric disorder [[delusional parasitosis]]. A louse was one of the early subjects of [[Microscope|microscopy]], appearing in [[Robert Hooke]]'s 1667 book, ''[[Micrographia]]''.
 ==Morphology and diversity==
-Lice are divided into two groups: sucking lice, which obtain their nourishment from feeding on the [[Sebaceous gland|sebaceous secretions]] and body fluids of their host; and chewing lice, which are [[scavenger]]s, feeding on [[skin]], 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.<ref name=IIBD>{{cite book | vauthors = Hoell HV, Doyen JT, Purcell AH |year=1998 |title=Introduction to Insect Biology and Diversity |edition=2nd |publisher= [[Oxford University Press]] |pages= 407–409|isbn= 978-0-19-510033-4}}</ref> If their host dies, lice can opportunistically use [[Phoresis (biology)|phoresis]] to hitch a ride on a fly and attempt to find a new host.<ref>{{cite thesis | vauthors = Harbison CW | date = 2008 |title=Ecology and Evolution of Transmission in Feather-feeding Lice (Phthiraptera: Ischnocera) | degree = Doctoral  | work = Department of Biology | publisher = University of Utah |url=https://books.google.com/books?id=RUfFjqPoQTEC&pg=PA83 |year=2008 |isbn=978-0-549-46429-7 |pages=83–87}}</ref>
+Lice are divided into two groups: sucking lice, which obtain their nourishment from feeding on the [[Sebaceous gland|sebaceous secretions]] and body fluids of their host; and chewing lice, which are [[scavenger]]s, feeding on [[skin]], 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.<ref name=IIBD>{{cite book | vauthors = Hoell HV, Doyen JT, Purcell AH |year=1998 |title=Introduction to Insect Biology and Diversity |edition=2nd |publisher= [[Oxford University Press]] |pages= 407–409|isbn= 978-0-19-510033-4}}</ref> If their host dies, lice can opportunistically use [[phoresis]] to hitch a ride on a fly and attempt to find a new host.<ref>{{cite thesis | vauthors = Harbison CW | date = 2008 |title=Ecology and Evolution of Transmission in Feather-feeding Lice (Phthiraptera: Ischnocera) | degree = Doctoral  | work = Department of Biology | publisher = University of Utah |url=https://books.google.com/books?id=RUfFjqPoQTEC&pg=PA83 |isbn=978-0-549-46429-7 |pages=83–87}}</ref>
-Sucking lice range in length from {{convert|0.5|to|5|mm|in|frac=64|abbr=on}}. They have narrow heads and oval, flattened bodies. They have no [[Simple eye in invertebrates|ocelli]], and their compound eyes are reduced in size or absent. Their [[Antenna (biology)|antennae]] are short with three to five segments, and their mouthparts, which are retractable into their head, are adapted for piercing and sucking.<ref name=Capinera/> 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 ([[ventral]]ly) and a food canal ([[dorsally]]).<ref name=GullanCranston>{{cite book| vauthors = Gullan PJ, Cranston PS |title=The Insects: An Outline of Entomology |date=2014 |publisher=Wiley |isbn=978-1-118-84615-5 |pages=41–42}}</ref> 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.<ref name=Capinera>{{cite book| vauthors = Capinera JL |title=Encyclopedia of Entomology |url=https://books.google.com/books?id=i9ITMiiohVQC&pg=PA838 |year=2008 |publisher=Springer Science & Business Media |isbn=978-1-4020-6242-1 |pages=838–844}}</ref>
+Sucking lice range in length from {{convert|0.5|to|5|mm|in|frac=64|abbr=on}}. They have narrow heads and oval, flattened bodies. They have no [[ocelli]], and their compound eyes are reduced in size or absent. Their [[Antenna (biology)|antennae]] are short with three to five segments, and their mouthparts, which are retractable into their head, are adapted for piercing and sucking.<ref name=Capinera/> 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 ([[ventral]]ly) and a food canal ([[dorsally]]).<ref name=GullanCranston>{{cite book| vauthors = Gullan PJ, Cranston PS |title=The Insects: An Outline of Entomology |date=2014 |publisher=Wiley |isbn=978-1-118-84615-5 |pages=41–42}}</ref> 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.<ref name=Capinera>{{cite book| vauthors = Capinera JL |title=Encyclopedia of Entomology |url=https://books.google.com/books?id=i9ITMiiohVQC&pg=PA838 |year=2008 |publisher=Springer Science & Business Media |isbn=978-1-4020-6242-1 |pages=838–844}}</ref>
-Chewing lice are also flattened and can be slightly larger than sucking lice, ranging in length from {{convert|0.5|to|6|mm|in|frac=64|abbr=on}}. 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 [[Ischnocera]], but club-shaped in the suborder [[Amblycera]]. The legs are short and robust, and terminated by one or two claws. Some species of chewing lice house [[Symbiosis|symbiotic]] bacteria in [[bacteriocyte]]s in their bodies. These may assist in digestion because if the insect is deprived of them, it will die.
+Chewing lice are also flattened and can be slightly larger than sucking lice, ranging in length from {{convert|0.5|to|6|mm|in|frac=64|abbr=on}}. 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 [[Ischnocera]], but club-shaped in the suborder [[Amblycera]]. The legs are short and robust, and terminated by one or two claws. Some species of chewing lice house [[symbiotic]] bacteria in [[bacteriocyte]]s in their bodies. These may assist in digestion because if the insect is deprived of them, it will die.
-Lice are usually [[Crypsis|cryptically coloured]] to match the fur or feathers of the host.<ref name=Capinera/><ref>{{cite web |url=http://phthiraptera.info/classification/23 |title=Phthiraptera: Summary | vauthors = Smith V |publisher=Phthiraptera.info |access-date=25 October 2015}}</ref> A louse's color varies from pale beige to dark gray; however, if feeding on blood, it may become considerably darker.
+Lice are usually [[Crypsis|cryptically coloured]] to match the fur or feathers of the host.<ref name=Capinera/><ref>{{cite web |url=http://phthiraptera.info/classification/23 |title=Phthiraptera: Summary |vauthors=Smith V |publisher=Phthiraptera.info |access-date=25 October 2015 }}{{Dead link|date=August 2023 |bot=InternetArchiveBot |fix-attempted=yes }}</ref> 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 [[Parthenogenesis|parthenogenetic]], with young developing from unfertilized eggs. A louse's [[egg]] is commonly called a nit. Many lice attach their eggs to their hosts' hair with specialized [[saliva]]; 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 [[Personal grooming|preening]], such as the interior of feather shafts. Living louse eggs tend to be pale whitish, whereas dead louse eggs are yellower.<ref name=IIBD/> Lice are [[Exopterygota|exopterygote]]s, being born as miniature versions of the adult, known as [[nymph (biology)|nymphs]]. The young moult three times before reaching the final adult form, usually within a month after hatching.<ref name=IIBD/>
+Female lice are usually more common than males, and some species are [[parthenogenetic]], with young developing from unfertilized eggs. A louse's [[egg]] is commonly called a nit. Many lice attach their eggs to their hosts' hair with specialized [[saliva]]; 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 [[Personal grooming|preening]], such as the interior of feather shafts. Living louse eggs tend to be pale whitish, whereas dead louse eggs are yellower.<ref name=IIBD/> Lice are [[exopterygote]]s, being born as miniature versions of the adult, known as [[nymph (biology)|nymphs]]. The young moult three times before reaching the final adult form, usually within a month after hatching.<ref name=IIBD/>
-Humans host three different kinds of lice: [[Head louse|head lice]], [[Body louse|body lice]], and [[Crab louse|pubic lice]]. 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 [[Nit comb|lice combs]], and medicated shampoos or washes.<ref>{{cite journal | vauthors = Mumcuoglu KY | title = Prevention and treatment of head lice in children | journal = Paediatric Drugs | volume = 1 | issue = 3 | pages = 211–8 | year = 1999 | pmid = 10937452 | doi = 10.2165/00128072-199901030-00005 | s2cid = 13547569 }}</ref>
+Humans host three different kinds of lice: [[head lice]], [[body lice]], and [[Crab louse|pubic lice]]. 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 [[Nit comb|lice combs]], and medicated shampoos or washes.<ref>{{cite journal | vauthors = Mumcuoglu KY | title = Prevention and treatment of head lice in children | journal = Paediatric Drugs | volume = 1 | issue = 3 | pages = 211–8 | year = 1999 | pmid = 10937452 | doi = 10.2165/00128072-199901030-00005 | s2cid = 13547569 }}</ref>
 ==Ecology==
 The average number of lice per host tends to be higher in large-bodied bird species than in small ones.<ref>{{cite journal | vauthors = Rózsa  | year = 1997| title = Patterns in the abundance of avian lice (Phthiraptera: Amblycera, Ischnocera) | url = http://www.zoologia.hu/list/patterns%20in%20the%20abundance.pdf | journal = Journal of Avian Biology | volume = 28 | issue = 3| pages = 249–254 | doi=10.2307/3676976| jstor = 3676976}}</ref> 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.<ref>{{cite journal | vauthors = Rékási J, Rozsa L, Kiss BJ | year = 1997 | title = Patterns in the distribution of avian lice (Phthiraptera: Amblycera, Ischnocera) | url = http://www.zoologia.hu/list/patterns%20in%20the%20distribution.pdf | journal = Journal of Avian Biology | volume = 28 | issue = 2| pages = 150–156 | doi=10.2307/3677308 | jstor = 3677308 | citeseerx = 10.1.1.506.730 }}</ref>
-Host organisms that dive under water to feed on aquatic prey harbor fewer taxa of lice.<ref>{{cite journal | vauthors = Felsõ B, Rózsa L | title = Reduced taxonomic richness of lice (Insecta: Phthiraptera) in diving birds | journal = The Journal of Parasitology | volume = 92 | issue = 4 | pages = 867–9 | date = August 2006 | pmid = 16995408 | doi = 10.1645/ge-849.1 | s2cid = 31000581 }}</ref><ref>{{cite journal | vauthors = Felső B, Rózsa L | year = 2007 | title = Diving behaviour reduces genera richness of lice (Insecta: Phthiraptera) of mammals | url = http://www.zoologia.hu/list/felso_rozsa2.pdf | journal = Acta Parasitologica | volume = 52 | pages = 82–85 | doi=10.2478/s11686-007-0006-3| s2cid = 38683871 }}</ref>
+Host organisms that dive under water to feed on aquatic prey harbour fewer taxa of lice.<ref>{{cite journal | vauthors = Felsõ B, Rózsa L | title = Reduced taxonomic richness of lice (Insecta: Phthiraptera) in diving birds | journal = The Journal of Parasitology | volume = 92 | issue = 4 | pages = 867–9 | date = August 2006 | pmid = 16995408 | doi = 10.1645/ge-849.1 | s2cid = 31000581 }}</ref><ref>{{cite journal | vauthors = Felső B, Rózsa L | year = 2007 | title = Diving behaviour reduces genera richness of lice (Insecta: Phthiraptera) of mammals | url = http://www.zoologia.hu/list/felso_rozsa2.pdf | journal = Acta Parasitologica | volume = 52 | pages = 82–85 | doi=10.2478/s11686-007-0006-3| s2cid = 38683871 }}</ref>
-Bird taxa that are capable of exerting stronger antiparasitic defense—such as stronger [[T cell]] immune response or larger [[uropygial gland]]s—harbor more taxa of Amblyceran lice than others.<ref>{{cite journal | vauthors = Møller AP, Rózsa L | title = Parasite biodiversity and host defenses: chewing lice and immune response of their avian hosts | journal = Oecologia | volume = 142 | issue = 2 | pages = 169–76 | date = January 2005 | pmid = 15503162 | doi = 10.1007/s00442-004-1735-8 | s2cid = 12992951 | bibcode = 2005Oecol.142..169M }}</ref><ref>{{cite journal | vauthors = Møller AP, Erritzøe J, Rózsa L | title = Ectoparasites, uropygial glands and hatching success in birds | journal = Oecologia | volume = 163 | issue = 2 | pages = 303–11 | date = June 2010 | pmid = 20043177 | doi = 10.1007/s00442-009-1548-x | s2cid = 11433594 | bibcode = 2010Oecol.163..303M }}</ref>
+Bird taxa that are capable of exerting stronger antiparasitic defence—such as stronger [[T cell]] immune response or larger [[uropygial gland]]s—harbour more taxa of Amblyceran lice than others.<ref>{{cite journal | vauthors = Møller AP, Rózsa L | title = Parasite biodiversity and host defenses: chewing lice and immune response of their avian hosts | journal = Oecologia | volume = 142 | issue = 2 | pages = 169–76 | date = January 2005 | pmid = 15503162 | doi = 10.1007/s00442-004-1735-8 | s2cid = 12992951 | bibcode = 2005Oecol.142..169M }}</ref><ref>{{cite journal | vauthors = Møller AP, Erritzøe J, Rózsa L | title = Ectoparasites, uropygial glands and hatching success in birds | journal = Oecologia | volume = 163 | issue = 2 | pages = 303–11 | date = June 2010 | pmid = 20043177 | doi = 10.1007/s00442-009-1548-x | s2cid = 11433594 | bibcode = 2010Oecol.163..303M }}</ref>
-Reductions in the size of host populations may cause a long-lasting reduction of louse taxonomic richness,<ref name=Rozsa1993>{{cite journal | vauthors = Ròzsa L | title = Speciation patterns of ectoparasites and "straggling" lice | journal = International Journal for Parasitology | volume = 23 | issue = 7 | pages = 859–64 | date = November 1993 | pmid = 8314369 | doi = 10.1016/0020-7519(93)90050-9 }}</ref> for example, birds introduced into [[New Zealand]] host fewer species of lice there than in Europe.<ref>{{cite journal | vauthors = Paterson AM, Palma RL, Gray RD | year = 1999 | title = How Frequently Do Avian Lice Miss the Boat? Implications for Coevolutionary Studies | journal = Systematic Biology | volume = 48 | pages = 214–223 | doi=10.1080/106351599260544 | doi-access = free }}</ref><ref>{{cite journal | vauthors = MacLeod CJ, Paterson AM, Tompkins DM, Duncan RP | title = Parasites lost - do invaders miss the boat or drown on arrival? | journal = Ecology Letters | volume = 13 | issue = 4 | pages = 516–27 | date = April 2010 | pmid = 20455925 | doi = 10.1111/j.1461-0248.2010.01446.x }}</ref> 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.<ref>{{cite journal | vauthors = Rózsa L, Rékási J, Reiczigel J | year = 1996 | title = Relationship of host coloniality to the population ecology of avian lice (Insecta: Phthiraptera) | url = http://www.zoologia.hu/list/relationship%20of%20host%20coloniality.pdf | journal = Journal of Animal Ecology | volume = 65 | issue = 2| pages = 242–248 | doi=10.2307/5727 | jstor = 5727 }}</ref> 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 [[speciation]] has occurred over evolutionary time-scales so it must be successfully accomplished sometimes.<ref name=Rozsa1993/>
+Reductions in the size of host populations may cause a long-lasting reduction of louse taxonomic richness,<ref name=Rozsa1993>{{cite journal | vauthors = Ròzsa L | title = Speciation patterns of ectoparasites and "straggling" lice | journal = International Journal for Parasitology | volume = 23 | issue = 7 | pages = 859–64 | date = November 1993 | pmid = 8314369 | doi = 10.1016/0020-7519(93)90050-9 }}</ref> for example, birds introduced into [[New Zealand]] host fewer species of lice there than in Europe.<ref>{{cite journal | vauthors = Paterson AM, Palma RL, Gray RD | year = 1999 | title = How Frequently Do Avian Lice Miss the Boat? Implications for Coevolutionary Studies | journal = Systematic Biology | volume = 48 | pages = 214–223 | doi=10.1080/106351599260544 | doi-access = free }}</ref><ref>{{cite journal | vauthors = MacLeod CJ, Paterson AM, Tompkins DM, Duncan RP | title = Parasites lost - do invaders miss the boat or drown on arrival? | journal = Ecology Letters | volume = 13 | issue = 4 | pages = 516–27 | date = April 2010 | pmid = 20455925 | doi = 10.1111/j.1461-0248.2010.01446.x | bibcode = 2010EcolL..13..516M }}</ref> 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.<ref>{{cite journal | vauthors = Rózsa L, Rékási J, Reiczigel J | year = 1996 | title = Relationship of host coloniality to the population ecology of avian lice (Insecta: Phthiraptera) | url = http://www.zoologia.hu/list/relationship%20of%20host%20coloniality.pdf | journal = Journal of Animal Ecology | volume = 65 | issue = 2| pages = 242–248 | doi=10.2307/5727 | jstor = 5727 | bibcode = 1996JAnEc..65..242R }}</ref> 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 [[speciation]] has occurred over evolutionary time-scales so it must be successfully accomplished sometimes.<ref name=Rozsa1993/>
 Lice may reduce host life expectancy if the infestation is heavy,<ref>{{cite journal | vauthors = Brown CR, Brown MB, Rannala B | year = 1995 | title = Ectoparasites reduce long-term survivorship of their avian host | url = http://www.rannala.org/reprints/1995/Brown1995a.pdf | journal = Proceedings of the Royal Society of London B | volume = 262 | issue = 1365| pages = 313–319 | doi=10.1098/rspb.1995.0211| s2cid = 13869042 }}</ref> but most seem to have little effect on their host. The habit of dust bathing in [[Chicken|domestic hens]] is probably an attempt by the birds to rid themselves of lice.<ref name=Capinera/> Lice may transmit microbial diseases and [[helminth]] parasites,<ref>{{cite journal | vauthors = Barlett CM |year=1993 |title=Lice (''Amblycera'' and ''Ischnocera'') as vectors of ''Eulimdana'' spp. (Nematoda: Filarioidea) in Charadriiform birds and the necessity of short reproductive periods in adult worms |journal=Journal of Parasitology |volume=75 |issue=1 |pages=85–91  |jstor=3283282 |doi=10.2307/3283282}}</ref> but most individuals spend their whole life cycle on a single host and are only able to transfer to a new host opportunistically.<ref name=Capinera/> Ischnoceran lice may reduce the [[thermoregulation]] effect of the plumage; thus heavily infested birds lose more heat than others.<ref>{{cite journal | vauthors = Booth DT, Clayton DH, Block BA | year = 1993 | title = Experimental demonstration of the energetic cost of parasitism in free-ranging hosts | url = http://darwin.biology.utah.edu/PubsHTML/PDF-Files/18.pdf | journal = Proceedings of the Royal Society of London B | volume = 253 | issue = 1337 | pages = 125–129 | doi = 10.1098/rspb.1993.0091 | bibcode = 1993RSPSB.253..125B | s2cid = 85731062 | access-date = 2010-08-14 | archive-url = https://web.archive.org/web/20100610162227/http://darwin.biology.utah.edu/PubsHTML/PDF-Files/18.pdf | archive-date = 2010-06-10 | url-status = dead }}</ref>
@@ Line 56: / Line 56: @@
 ==Evolution==
-Phthiraptera lice are members of [[Psocodea]] (formerly [[Psocoptera]]), the [[order (biology)|order]] that contains booklice, barklice and barkflies. Within Psocodea, lice are within the [[suborder]] [[Troctomorpha]], and most closely related to the family [[Liposcelididae]].<ref>{{cite journal | vauthors = Light JE, Smith VS, Allen JM, Durden LA, Reed DL | title = Evolutionary history of mammalian sucking lice (Phthiraptera: Anoplura) | journal = BMC Evolutionary Biology | volume = 10 | issue = 1 | pages = 292 | date = September 2010 | pmid = 20860811 | pmc = 2949877 | doi = 10.1186/1471-2148-10-292 }}</ref> The oldest confirmed fossil louse is a bird louse, ''[[Megamenopon|Megamenopon rasnitsyni]]'', from [[Eckfelder Maar]], Germany, which dates to the [[Eocene]], around 44 million years ago.<ref name="Scratching an ancient itch: an Eoce">{{cite journal | vauthors = Wappler T, Smith VS, Dalgleish RC | title = Scratching an ancient itch: an Eocene bird louse fossil | journal = Proceedings. Biological Sciences | volume = 271 Suppl 5 | issue = suppl_5 | pages = S255-8 | date = August 2004 | pmid = 15503987 | pmc = 1810061 | doi = 10.1098/rsbl.2003.0158 }}</ref> ''[[Saurodectes|Saurodectes vrsanskyi]]'' from the Early Cretaceous ([[Aptian]]) [[Zaza Formation]] of [[Buryatia]], Russia, has also been suggested to be a louse, but this is tentative.<ref>{{cite journal | vauthors = Rasnitsyn AP, Zherikhin VV | title = First fossil chewing louse from the lower Cretaceous of Baissa, Transbaikalia (Insecta, Pediculida= Phthiriaptera, Saurodectidae fam. n.). | journal = Russian Entomological Journal | date = 1999 | volume = 8 | issue = 4 | pages = 253–5 }}</ref>
+Phthiraptera lice are members of [[Psocodea]] (formerly [[Psocoptera]]), the [[order (biology)|order]] that contains booklice, barklice and barkflies. Within Psocodea, lice are within the [[suborder]] [[Troctomorpha]], and most closely related to the family [[Liposcelididae]].<ref>{{cite journal | vauthors = Light JE, Smith VS, Allen JM, Durden LA, Reed DL | title = Evolutionary history of mammalian sucking lice (Phthiraptera: Anoplura) | journal = BMC Evolutionary Biology | volume = 10 | issue = 1 | pages = 292 | date = September 2010 | pmid = 20860811 | pmc = 2949877 | doi = 10.1186/1471-2148-10-292 | doi-access = free | bibcode = 2010BMCEE..10..292L }}</ref> The oldest confirmed fossil louse is ''[[Archimenopon|Archimenopon myanmarensis]]'', an [[amblycera]]n from the Cretaceous amber from [[Myanmar]].<ref name=Archimenopon /> Another early representative of the group is a bird louse, ''[[Megamenopon|Megamenopon rasnitsyni]]'', from [[Eckfelder Maar]], Germany, which dates to the [[Eocene]], around 44 million years ago.<ref name="Scratching an ancient itch: an Eoce">{{cite journal | vauthors = Wappler T, Smith VS, Dalgleish RC | title = Scratching an ancient itch: an Eocene bird louse fossil | journal = Proceedings. Biological Sciences | volume = 271 Suppl 5 | issue = suppl_5 | pages = S255-8 | date = August 2004 | pmid = 15503987 | pmc = 1810061 | doi = 10.1098/rsbl.2003.0158 }}</ref> ''[[Saurodectes|Saurodectes vrsanskyi]]'' from the Early Cretaceous ([[Aptian]]) [[Zaza Formation]] of [[Buryatia]], Russia, has also been suggested to be a louse, but this is tentative.<ref>{{cite journal | vauthors = Rasnitsyn AP, Zherikhin VV | title = First fossil chewing louse from the lower Cretaceous of Baissa, Transbaikalia (Insecta, Pediculida= Phthiriaptera, Saurodectidae fam. n.). | journal = Russian Entomological Journal | date = 1999 | volume = 8 | issue = 4 | pages = 253–5 }}</ref>
+Placental mammal lice had a single common ancestor that lived on [[Afrotheria]] with this arising from host-switching from an ancient avian host.<ref name="p273">{{cite journal |last1=Johnson |first1=Kevin P. |last2=Matthee |first2=Conrad |last3=Doña |first3=Jorge |date=2022 |title=Phylogenomics reveals the origin of mammal lice out of Afrotheria |journal=Nature Ecology & Evolution |volume=6 |issue=8 |pages=1205–1210 |doi=10.1038/s41559-022-01803-1 |pmid=35788706 |bibcode=2022NatEE...6.1205J |issn=2397-334X}}</ref>
 [[Cladogram]] showing the position of Phthiraptera within [[Psocodea]]:<ref name="deMoya2021">{{Cite journal|last1=de Moya|first1=Robert S|last2=Yoshizawa|first2=Kazunori|last3=Walden|first3=Kimberly K O|last4=Sweet|first4=Andrew D|last5=Dietrich|first5=Christopher H|last6=Kevin P|first6=Johnson|date=2021-06-16|editor-last=Buckley|editor-first=Thomas|title=Phylogenomics of Parasitic and Nonparasitic Lice (Insecta: Psocodea): Combining Sequence Data and Exploring Compositional Bias Solutions in Next Generation Data Sets|url=https://academic.oup.com/sysbio/article/70/4/719/5912026|journal=Systematic Biology|language=en|volume=70|issue=4|pages=719–738|doi=10.1093/sysbio/syaa075|pmid=32979270|issn=1063-5157|doi-access=free}}</ref>
@@ Line 102: / Line 104: @@
 ===Classification===
-Phthiraptera is clearly a [[Monophyly|monophyletic]] grouping, united as the members are by a number of derived features including their parasitism on [[warm-blooded]] vertebrates and the combination of their [[Metathoracic ganglion|metathoracic]] [[Ganglion|ganglia]] with their [[abdominal ganglia]] to form a single ventral nerve junction.<ref name=Ax>{{cite book| vauthors = Ax P |title=Multicellular Animals: Volume II: The Phylogenetic System of the Metazoa |url=https://books.google.com/books?id=x7vtCAAAQBAJ&pg=PA303 |year=2013|publisher=Springer Science & Business Media |isbn=978-3-662-10396-8 |pages=303–307 }}</ref> The order has traditionally been divided into two suborders, the sucking lice (Anoplura) and the [[Mastication|chewing]] lice (Mallophaga); however, subsequent classifications suggest that the Mallophaga are [[paraphyletic]] and four suborders were then recognized:<ref name=info>{{cite web |url=http://phthiraptera.info/ |title=Phthiraptera.info | vauthors = Smith V |publisher=International Society of Phthirapterists |access-date=25 October 2015}}</ref>
+Phthiraptera is clearly a [[monophyletic]] grouping, united as the members are by a number of derived features including their parasitism on [[warm-blooded]] vertebrates and the combination of their [[Metathoracic ganglion|metathoracic]] [[ganglia]] with their [[abdominal ganglia]] to form a single ventral nerve junction.<ref name=Ax>{{cite book| vauthors = Ax P |title=Multicellular Animals: Volume II: The Phylogenetic System of the Metazoa |url=https://books.google.com/books?id=x7vtCAAAQBAJ&pg=PA303 |year=2013|publisher=Springer Science & Business Media |isbn=978-3-662-10396-8 |pages=303–307 }}</ref> The order has traditionally been divided into two suborders, the sucking lice (Anoplura) and the [[chewing]] lice (Mallophaga); however, subsequent classifications suggest that the Mallophaga are [[paraphyletic]] and four suborders were then recognized:<ref name=info>{{cite web |url=http://phthiraptera.info/ |title=Phthiraptera.info | vauthors = Smith V |publisher=International Society of Phthirapterists |access-date=25 October 2015}}</ref>
 * [[Anoplura]]: [[sucking]] lice, occurring on mammals exclusively
-* [[Haematomyzus|Rhynchophthirina]]: parasites of elephants and [[warthog]]s
+* [[Rhynchophthirina]]: parasites of elephants and [[warthog]]s
 * [[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
@@ Line 111: / Line 113: @@
 Upon finding that Phthiraptera was nested within [[Psocoptera]], Phthiraptera, in 2021 de Moya et al. proposed reducing the rank of Phthiraptera to [[infraorder]], and the four [[suborder]]s to [[parvorder]].<ref name="deMoya2021"/> These changes were accepted by Psocodea Species File and others, with the exception of placing Phthiraptera under the infraorder [[Nanopsocetae]], as a parvorder, with the four subgroups listed above. These classifications are likely to change in the future as a result of ongoing [[phylogenetic]] research.<ref name=Johnson2021/><ref name=catlife/>
-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.<ref name=info/> They are found even in the [[Antarctic]], where [[penguin]]s carry 15 species of lice (in the genera ''[[Austrogonoides]]'' and ''[[Nesiotinus]]'').<ref>{{cite journal | vauthors = Banks JC, Paterson AM |title=A penguin-chewing louse (Insecta : Phthiraptera) phylogeny derived from morphology |journal=Invertebrate Systematics |date=2004 |volume=18 |issue=1 |pages=89–100 |doi=10.1071/IS03022 |s2cid=53516244 |url=https://semanticscholar.org/paper/16d5d420c889243028da7dde813779ff82b1b458 }}</ref> The oldest known record of the group is ''[[Megamenopon|Megamenopon rasnitsyni]]'' from 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.<ref name="Scratching an ancient itch: an Eoce"/>
+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.<ref name=info/> They are found even in the [[Antarctic]], where [[penguin]]s carry 15 species of lice (in the genera ''[[Austrogonoides]]'' and ''[[Nesiotinus]]'').<ref>{{cite journal | vauthors = Banks JC, Paterson AM |title=A penguin-chewing louse (Insecta : Phthiraptera) phylogeny derived from morphology |journal=Invertebrate Systematics |date=2004 |volume=18 |issue=1 |pages=89–100 |doi=10.1071/IS03022 |s2cid=53516244 }}</ref> The oldest known record of the group is ''[[Megamenopon|Megamenopon rasnitsyni]]'' from 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.<ref name="Scratching an ancient itch: an Eoce"/>
 <gallery mode="nolines">
 File:Ricinus bombycillae (Denny, 1842).JPG|''Ricinus bombycillae'', an amblyceran louse from a [[Bohemian waxwing]]
 File:Trinoton anserinum (Fabricius, 1805).JPG|''Trinoton anserinum'', an amblyceran louse from a [[mute swan]]
-File:Lice image01.jpg|''[[Bovicola]] limbata'', an ischnoceran louse from [[goat]]s. The species is [[sexual dimorphism|sexually dimorphic]], with the male smaller than the female.
+File:Lice image01.jpg|''[[Bovicola]] limbata'', an ischnoceran louse from [[goat]]s. The species is [[sexually dimorphic]], with the male smaller than the female.
 </gallery>
@@ Line 123: / Line 125: @@
 Lice have been the subject of significant [[DNA]] 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, ''Pediculus'' and ''Pthirus'': head lice (''Pediculus humanus capitis''), body lice (''Pediculus humanus humanus''), and pubic lice (''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.<ref name="Reed D.L. 2007"/> 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–human last common ancestor|chimpanzee-hominid divergence]].<ref name="Reed D.L. 2007"/> Because parasites rely on their hosts, host{{endash}}parasite cospeciation events are likely.
-Genetic evidence suggests that human ancestors acquired pubic lice from [[gorilla]]s approximately 3-4 million years ago.<ref name="Reed D.L. 2007">{{cite journal | vauthors = Reed DL, Light JE, Allen JM, Kirchman JJ | title = Pair of lice lost or parasites regained: the evolutionary history of anthropoid primate lice | journal = BMC Biology | volume = 5 | issue = 7 | pages = 7 | date = March 2007 | pmid = 17343749 | pmc = 1828715 | doi = 10.1186/1741-7007-5-7 }}</ref> 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 parasite{{endash}}host switch occurred in evolutionary history, this explanation is the most parsimonious (containing the fewest evolutionary changes).<ref name="Reed D.L. 2007"/>
+Genetic evidence suggests that human ancestors acquired pubic lice from [[gorilla]]s approximately 3-4 million years ago.<ref name="Reed D.L. 2007">{{cite journal | vauthors = Reed DL, Light JE, Allen JM, Kirchman JJ | title = Pair of lice lost or parasites regained: the evolutionary history of anthropoid primate lice | journal = BMC Biology | volume = 5 | issue = 7 | pages = 7 | date = March 2007 | pmid = 17343749 | pmc = 1828715 | doi = 10.1186/1741-7007-5-7 | doi-access = free }}</ref> 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 parasite{{endash}}host switch occurred in evolutionary history, this explanation is the most parsimonious (containing the fewest evolutionary changes).<ref name="Reed D.L. 2007"/>
-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.<ref name=Parry>{{cite web | vauthors = Parry W |title=Lice Reveal Clues to Human Evolution |url=http://www.livescience.com/41028-lice-reveal-clues-to-human-evolution.html |publisher=LiveScience |access-date=25 October 2015 |date=7 November 2013}}</ref> 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.<ref name=Parry/><ref name="Kittler R. 2003">{{cite journal | vauthors = Kittler R, Kayser M, Stoneking M | title = Molecular evolution of Pediculus humanus and the origin of clothing | journal = Current Biology | volume = 13 | issue = 16 | pages = 1414–7 | date = August 2003 | pmid = 12932325 | doi = 10.1016/S0960-9822(03)00507-4 | s2cid = 15277254 | doi-access = free }}</ref>
+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.<ref name=Parry>{{cite web | vauthors = Parry W |title=Lice Reveal Clues to Human Evolution |url=http://www.livescience.com/41028-lice-reveal-clues-to-human-evolution.html |publisher=LiveScience |access-date=25 October 2015 |date=7 November 2013}}</ref> 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.<ref name=Parry/><ref name="Kittler R. 2003">{{cite journal | vauthors = Kittler R, Kayser M, Stoneking M | title = Molecular evolution of Pediculus humanus and the origin of clothing | journal = Current Biology | volume = 13 | issue = 16 | pages = 1414–7 | date = August 2003 | pmid = 12932325 | doi = 10.1016/S0960-9822(03)00507-4 | s2cid = 15277254 | doi-access = free | bibcode = 2003CBio...13.1414K }}</ref>
-The mitochondrial genome of the human species of the body louse (''[[Pediculus humanus|Pediculus humanus humanus]]''), the head louse (''[[Pediculus humanus|Pediculus humanus capitis]]'') and the pubic louse (''[[Pthirus pubis]]'') fragmented into a number of minichromosomes, at least seven million years ago.<ref name=Shao2012>{{cite journal | vauthors = Shao R, Zhu XQ, Barker SC, Herd K | title = Evolution of extensively fragmented mitochondrial genomes in the lice of humans | journal = Genome Biology and Evolution | volume = 4 | issue = 11 | pages = 1088–101 | year = 2012 | pmid = 23042553 | pmc = 3514963 | doi = 10.1093/gbe/evs088 }}</ref> Analysis of mitochondrial DNA in human body and hair lice reveals that greater genetic diversity existed in African than in non-African lice.<ref name="Kittler R. 2003"/><ref name=Light>{{cite journal | vauthors = Light JE, Allen JM, Long LM, Carter TE, Barrow L, Suren G, Raoult D, Reed DL | display-authors = 6 | title = Geographic distributions and origins of human head lice (Pediculus humanus capitis) based on mitochondrial data | journal = The Journal of Parasitology | volume = 94 | issue = 6 | pages = 1275–81 | date = December 2008 | pmid = 18576877 | doi = 10.1645/GE-1618.1 | s2cid = 9456340 }}</ref>  Human lice can also shed light on human migratory patterns in prehistory. The dominating theory of [[Anthropology|anthropologists]] regarding human migration is the [[Recent African origin of modern humans|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.<ref name=Light/>
+The mitochondrial genome of the human species of the body louse (''[[Pediculus humanus|Pediculus humanus humanus]]''), the head louse (''[[Pediculus humanus|Pediculus humanus capitis]]'') and the pubic louse (''[[Pthirus pubis]]'') fragmented into a number of minichromosomes, at least seven million years ago.<ref name=Shao2012>{{cite journal | vauthors = Shao R, Zhu XQ, Barker SC, Herd K | title = Evolution of extensively fragmented mitochondrial genomes in the lice of humans | journal = Genome Biology and Evolution | volume = 4 | issue = 11 | pages = 1088–101 | year = 2012 | pmid = 23042553 | pmc = 3514963 | doi = 10.1093/gbe/evs088 }}</ref> Analysis of mitochondrial DNA in human body and hair lice reveals that greater genetic diversity existed in African than in non-African lice.<ref name="Kittler R. 2003"/><ref name=Light>{{cite journal | vauthors = Light JE, Allen JM, Long LM, Carter TE, Barrow L, Suren G, Raoult D, Reed DL | display-authors = 6 | title = Geographic distributions and origins of human head lice (Pediculus humanus capitis) based on mitochondrial data | journal = The Journal of Parasitology | volume = 94 | issue = 6 | pages = 1275–81 | date = December 2008 | pmid = 18576877 | doi = 10.1645/GE-1618.1 | s2cid = 9456340 }}</ref>  Human lice can also shed light on human migratory patterns in prehistory. The dominating theory of [[Anthropology|anthropologists]] 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.<ref name=Light/>
 ==In human culture==
@@ Line 135: / Line 137: @@
 [[File:Louse diagram, Micrographia, Robert Hooke, 1667.jpg|thumb|upright=0.7|left|Drawing of a louse clinging to a human hair. [[Robert Hooke]], ''[[Micrographia]]'', 1667]]
-Lice have been intimately associated with human society throughout history. In the [[Middle Ages]], they were essentially ubiquitous. At the death of [[Thomas Becket]], [[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".<ref name=Kowalski>{{cite journal | vauthors = Kowalski TJ, Agger WA | title = Art supports new plague science | journal = Clinical Infectious Diseases | volume = 48 | issue = 1 | pages = 137–8 | date = January 2009 | pmid = 19067623 | doi = 10.1086/595557 | doi-access = free }}</ref> 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.<ref>{{Cite web |last=Harvey |first=Katherine |date=2019-04-09 |title=Medieval people were surprisingly clean (apart from the clergy) |url=https://aeon.co/essays/medieval-people-were-surprisingly-clean-apart-from-the-clergy |url-status=live |access-date=2022-10-13 |website=[[Aeon magazine|Aeon]] |language=en}}</ref> A mediaeval treatment for lice was an [[ointment]] made from pork grease, [[incense]], [[lead]], and [[aloe]].<ref>{{cite book | vauthors = Elliott L |title=Clothing in the Middle Ages |url=https://books.google.com/books?id=qtq_WSpdo0gC&pg=PT29 |year=2004 |publisher=Crabtree |isbn=978-0-7787-1351-7 |page=29}}</ref>
+Lice have been intimately associated with human society throughout history. In the [[Middle Ages]], they were essentially ubiquitous. At the death of [[Thomas Becket]], [[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".<ref name=Kowalski>{{cite journal | vauthors = Kowalski TJ, Agger WA | title = Art supports new plague science | journal = Clinical Infectious Diseases | volume = 48 | issue = 1 | pages = 137–8 | date = January 2009 | pmid = 19067623 | doi = 10.1086/595557 | doi-access = free }}</ref> 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.<ref>{{Cite web |last=Harvey |first=Katherine |date=2019-04-09 |title=Medieval people were surprisingly clean (apart from the clergy) |url=https://aeon.co/essays/medieval-people-were-surprisingly-clean-apart-from-the-clergy |access-date=2022-10-13 |website=[[Aeon magazine|Aeon]] |language=en}}</ref> A mediaeval treatment for lice was an [[ointment]] made from pork grease, [[incense]], [[lead]], and [[aloe]].<ref>{{cite book | vauthors = Elliott L |title=Clothing in the Middle Ages |url=https://books.google.com/books?id=qtq_WSpdo0gC&pg=PT29 |year=2004 |publisher=Crabtree |isbn=978-0-7787-1351-7 |page=29}}</ref>
 [[Robert Hooke]]'s 1667 book, ''[[Micrographia|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 early [[microscope]].<ref>{{cite web | vauthors = Hooke R |title=Microscopic view of a louse |url=http://prints.royalsociety.org/art/580831/microscopic-view-of-a-louse |publisher=The Royal Society |access-date=17 October 2015}}</ref>
@@ Line 147: / Line 149: @@
 Soldiers in the trenches of the [[First World War]] suffered severely from lice, and the [[typhus]] 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]]. "[[Delousing]]" became a [[euphemism]] for the extermination of Jews in [[concentration camp]]s such as [[Auschwitz]] under the Nazi regime.<ref>{{cite web | vauthors = Evans RJ |author-link1=Richard J. Evans |title=The Great Unwashed |url=http://www.gresham.ac.uk/lectures-and-events/the-great-unwashed |publisher=Gresham College |access-date=17 October 2015}}</ref>
-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".<ref>{{cite web | vauthors = Weinstein P | date = 26 February 2013 |title= 'The Great Unwashed': Entomophobia/Delusionary Parasitosis; Illusionary Parasitosis |url=http://medent.usyd.edu.au/fact/delpara.htm |publisher=University of Sydney Department of Medical Entomology |access-date=17 October 2015 | series = The Great Plagues: Epidemics in History from the Middle Ages to the Present Day }}</ref>
+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".<ref>{{cite web | vauthors = Weinstein P | date = 26 February 2013 | title = 'The Great Unwashed': Entomophobia/Delusionary Parasitosis; Illusionary Parasitosis | url = http://medent.usyd.edu.au/fact/delpara.htm | publisher = University of Sydney Department of Medical Entomology | access-date = 17 October 2015 | series = The Great Plagues: Epidemics in History from the Middle Ages to the Present Day | archive-date = 17 May 2016 | archive-url = https://web.archive.org/web/20160517005233/http://medent.usyd.edu.au/fact/delpara.htm | url-status = dead }}</ref>
 ===In science===
@@ Line 154: / Line 156: @@
 ===In literature and folklore===
-[[File:Mother Louse, Alewife Wellcome L0000658.jpg|thumb|left|upright|Mother Louse, a notorious [[Alewife (trade)|alewife]] in Oxford during the mid-18th century, shown with three lice as a [[coat of arms]]. Image by [[David Loggan]].<ref name="White1859">{{cite book|url=https://books.google.com/books?id=l68ZhT2agbEC&pg=PA275|title=Notes & Queries|publisher=Oxford University Press|year=1859|pages=275–276| vauthors = White W }}</ref><ref>{{Cite document|publisher=University of York| vauthors = Pierce H |title=Unseemly pictures: political graphic satire in England, c. 1600-c. 1650|date=2004|url=http://etheses.whiterose.ac.uk/9864/1/423697_vol1.pdf}}</ref>]]
+[[File:Mother Louse, Alewife Wellcome L0000658.jpg|thumb|left|upright|Mother Louse, a notorious [[Alewife (trade)|alewife]] in Oxford during the mid-18th century, shown with three lice as a [[coat of arms]]. Image by [[David Loggan]].<ref name="White1859">{{cite book|url=https://books.google.com/books?id=l68ZhT2agbEC&pg=PA275|title=Notes & Queries|publisher=Oxford University Press|year=1859|pages=275–276| vauthors = White W }}</ref><ref>{{Cite web|publisher=University of York| vauthors = Pierce H |title=Unseemly pictures: political graphic satire in England, c. 1600-c. 1650|date=2004|url=http://etheses.whiterose.ac.uk/9864/1/423697_vol1.pdf}}</ref>]]
 [[James Joyce]]'s 1939 book ''[[Finnegans Wake]]'' has the character Shem the Penman infested with "[[foxtrot]]ting fleas, the lieabed lice,&nbsp;... bats in his belfry".<ref>{{cite book | vauthors = Joyce J | author-link1 = James Joyce |title=Finnegans Wake |date=1939 |publisher=Faber |page=180}}</ref>
@@ Line 163: / Line 165: @@
 [[Robert Burns]] 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." [[John Milton]] in ''[[Paradise Lost]]'' 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." [[John Ray]] recorded a Scottish proverb, "Gie a beggar a bed and he'll repay you with a Louse."
-In [[William Shakespeare|Shakespeare]]'s ''[[Troilus and Cressida]]'', [[Thersites]] compares [[Menelaus]], brother of [[Agamemnon]], 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."<ref>{{Cite thesis |title=Insect Life in the Poetry and Drama of England: With Special Reference to Poetry | vauthors = Twinn CR |publisher=University of Ottawa | degree= PhD |year=1942 |hdl=10393/21088  }}</ref>
+In [[Shakespeare]]'s ''[[Troilus and Cressida]]'', [[Thersites]] compares [[Menelaus]], brother of [[Agamemnon]], 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."<ref>{{Cite thesis |title=Insect Life in the Poetry and Drama of England: With Special Reference to Poetry | vauthors = Twinn CR |publisher=University of Ottawa | degree= PhD |year=1942 |hdl=10393/21088  }}</ref>
 === Woodlouse ===
@@ Line 184: / Line 186: @@
 }}</ref>
-<ref name=catlife>
+<ref name=catlife>{{Cite web
-{{Cite web| title = Catalogue of Life, Phthiraptera Haeckel, 1896
+| title = Catalogue of Life, Phthiraptera Haeckel, 1896
 | url = https://www.catalogueoflife.org/data/taxon/8MP8P
 | access-date = 2021-11-03
 | date = 2021
+| archive-date = 2021-11-05
+| archive-url = https://web.archive.org/web/20211105084348/https://www.catalogueoflife.org/data/taxon/8MP8P
+| url-status = dead
 }}</ref>
 }}

v t e Human lice and pediculosis
Species	Head louse Crab louse Body louse
Infestation	Pediculosis Pediculosis corporis Phthiriasis
Treatment	Nitpicking Pediculicide Lindane Permethrin Phenothrin Delphinium
Other terms of interest	Cooties Sucking louse Louse