Human lice

The human louse Pediculus humanus L. has a long history, associated with human evolution and the geographic expansion of early humans (Ashfaq et al., Reference Ashfaq, Prosser, Nasir, Masood, Ratnasingham and Hebert2015). In terms of its origin, it perhaps diverged from P. schaeffi, from chimpanzees or their direct ancestors onto early hominids (Herd et al., Reference Herd, Barker and Shao2015). The separation of P. humanus into 2 subspecies, the head louse (P. humanus capitis DeGeer) and the body louse (P. humanus humanus L.), poses an interesting question regarding the timing and reasons for this evolutionary change, with DNA evidence placing it around 72 000–42 000 years ago and associating it with the earliest use of clothing (Kittler et al., Reference Kittler, Kayser and Stoneking2003). Body lice spread infectious diseases, including epidemic typhus (Rickettsia prowazekii), trench fever (Bartonella quintana) and louse-borne relapsing fever (Borrelia recurrentis) (Badiaga and Brouqui, Reference Badiaga and Brouqui2012).

The fossil record of P. humanus is linked with the record of its host, as it lives and breeds on humans for its entire life cycle. The earliest evidence of head lice comes from both South America and Asia, suggesting links with early human migrations (Figure 1). Head louse nits from human hair were found in northeast Brazil in deposits of ca. 10 000 years BP (Araújo et al., Reference Araújo, Ferreira, Guidon, Maures da Serra Freire, Reinhard and Dittmar2000), and from 9000-year-old human hair remains were recovered from Nahal Hemar cave in the southern Levant (Mumcuoglu and Zias, Reference Mumcuoglu and Zias1991).

Figure 1.

The Holocene record of the human louse, Pediculus humanus L. and its subspecies, P. humanus capitis DeGeer and P. humanus humanus L. In some instances (e.g. South American records), the location point represents multiple sites. For more information, see the relevant discussion in the paper.

Figure 1 Long description

The global distribution of Pediculus humanus subspecies and archaeological periods using different markers. Circles represent Pediculus humanus L., triangles for Pediculus humanus capitis DeGeer and squares for Pediculus humanus humanus L. Archaeological periods are indicated by color-coded markers: Paleolithic/Lithic, Neolithic, Bronze/pre-contact, Iron Age/pre-contact, Roman Age/pre-contact, Medieval/pre-contact and Post-medieval/pre-contact. The markers are scattered across various regions including North America, South America, Europe, Asia and parts of Africa. Clusters are visible in Europe and the Middle East, with isolated points in North and South America and Asia. The map provides a visual representation of historical and geographical data related to human lice and archaeological periods.

The understanding of the distribution record of the human louse over time suffers from taphonomic limitations, with lice from mummies providing some of the best-preserved assemblages. It is not coincidental that the first publication on human head lice discussed their presence on mummies in the Cairo Museum collection (Ruffer, Reference Ruffer1914), the majority of which probably were sourced from the Valley of the Kings in the early 20th century. Ewing (Reference Ewing1924) also mentioned human lice from 4th-century CE Egyptian mummies. Head louse nits were recovered from the hair of Egyptian mummies, with the earliest evidence from the Old Kingdom (ca. 2686–2181 BCE) at Abydos (Fletcher, Reference Fletcher1994, Reference Fletcher1998). Additional information comes from later pharaonic mummies (e.g. a XXII Dynasty (ca. 800 BCE) mummy; Huchet, Reference Huchet2016). Delousing combs, used since the pre-Dynastic period (Del Río Álvarez, Reference Del Río Álvarez2018), anti-louse recipes from the Ebers papyrus, written around 3500 years ago (Bryan, Reference Bryan1930), and body shaving practised by the priests against lice (Herodotus Histories 2.37) attest to infestations and measures of control (Herodotus, Reference Herodotus1920; Kislev and Simchoni, Reference Kislev, Simchoni, Aviram, Foerster, Netzer and Stiebel2007). In Asia, additional early head lice records were recovered from a female mummy from Loulan, in Xinjiang, China, ca. 3800 BP (Wen et al., Reference Wen, Zhaoyong, Zhije, Yehua, Jianghua and Zhiya1987). Although generally preservation is largely inimical for the recovery of ectoparasites, an abdomen of P. humanus was found in the Late Minoan IB destruction layer (ca. 1600–1450 BCE), from Aghia Aikaterini in Chania on the Aegean island of Crete (Panagiotakopulu in press).

Southern Levant records include head lice from between the teeth of ivory combs dated ca. 1150–1200 BCE from a pit at Lachish, Ashkelon (Vainstub et al., Reference Vainstub, Mumcuoglu, Hasel, Hesler, Lavi, Rabinovich, Goren and Garfinkel2022), and from Roman combs from the Cave of the Pool, at En Gedi (Mumcuoglu and Hadas, Reference Mumcuoglu and Hadas2011), from Moa in the Arava region and from Ein Hatzeva (Amanzougaghene et al., Reference Amanzougaghene, Mumcuoglu, Fenollar, Alfi, Yeslyurt, Raoult and Mediannikov2016). Body lice, P. humanus humanus, were recovered from clothing in the Jewish fortress at Masada, destroyed by the Roman army ca. 73 CE. Head lice and nits were recovered from combs from Byzantine (ca. 6th–8th century CE) Wadi al-Far’a in the northern Levant (Mumcuoglu and Zias, Reference Mumcuoglu and Zias1989). Also from the Roman period, research at Wadi Halfa in ancient Nubia, ca. 350–550 CE, revealed that 40% of the 218 mummies studied were infested with head lice (Armelagos, Reference Armelagos1969).

There are several records of lice from Europe, with nits of body lice attached to wool fabric recovered from Iron Age Hallstätt (ca. 800–450 BCE) in Salzburg in Austria (Hundt, Reference Hundt1960). An Iron Age human louse was found at Buiston Crannog in Scotland (Kenward et al., Reference Kenward, Hill, Jaques, Kroupa, Large and Crone2000), and lice were also recovered from later phases of the site. Various Roman sites across Britain provided evidence for human lice, including Carlisle (Smith and Tetlow, Reference Smith, Tetlow and Howard-davis2009), Lincoln (Carrott et al., Reference Carrott, Issitt, Kenward, Large, McKenna and Skidmore1995) and York General Accident (Hall and Kenward, Reference Hall and Kenward1990). The assemblages from Roman Castellum Fectio, in the Netherlands, also included human lice (Van den Bos, Reference Van den Bos, Brinkkemper, Bull, Engels, Hakbijl, Schepers, van Dinter, van Reenen and van Geel2014). Head lice from combs were recovered from Roman Ribchester (Large et al., Reference Large, Kenward, Carrott, Nicholson and Kent1994). The most poignant example of head lice in the Roman period is probably the evidence for nits from the hair of a victim of the 79 CE volcanic eruption of Vesuvius, recovered from Herculaneum in southern Italy (Capasso and Di Tota, Reference Capasso and Di Tota1998).

Most records of human lice come from the medieval period, with specimens recovered primarily from sites in Britain and the North Atlantic islands. Human lice were also present in insect faunas from several sites in Hull: Blanket Row (Carrott et al., Reference Carrott, Hall, Jaques, Johnstone, Kenward and Rowland2001), High Street (Jaques et al., Reference Jaques, Hall, Kenward and Carrott2003) and Magistrates Court (Hall et al., Reference Hall, Carrott, Jaques, Johnstone, Kenward, Large and Usai2000). Lincoln (Carrott et al., Reference Carrott, Issitt, Kenward, Large, McKenna and Skidmore1995), Whitby (Hall et al., Reference Hall, Jaques, Kenward and Large1993) and several sites in York (Anglo-Scandinavian Micklegate (Kenward and Hall, Reference Kenward and Hall2000), Anglo-Scandinavian Parliament Street (Hall and Kenward, Reference Hall and Kenward2000), Low Petergate (Hall et al., Reference Hall, Kenward, Girvan and McKenna2007), Walmgate (Jaques et al., Reference Jaques, Hall, Kenward, Rowland and Carrott2001; Hall et al., Reference Hall, Kenward, Jaques, Carrott and Rowland2002)) provided similar evidence. Human lice were present at Viking-period Tuquoy on Westray in the Orkney Islands (Sadler, Reference Sadler1993) and medieval Aberdeen in Scotland (Kenward and Hall, Reference Kenward, Hall, Cameron and Stones2001). Medieval Glenarm Deer Park Farms, in Antrim, Northern Ireland, provided evidence for high frequency of human lice with 585 individuals (Allison et al., Reference Allison, Hall and Kenward1999; Kenward et al., Reference Kenward, Hall, Allison, Carrott, Lynn and McDowell2011), and further records come from Fishamble Street in Dublin, Ireland (Reilly, Reference Reilly2024).

The recovery of human lice from early medieval deposits from 7th- to 8th-century Ribe in Jutland (Allison and Kenward, Reference Allison, Kenward and Sindbæk2022), and from 11th-century Viborg also in Denmark (Kenward, Reference Kenward, Iversen, Robinson, Hjermind and Christensen2005) offers a glimpse of conditions in potential departure points in continental Viking towns and ports.

Evidence of head lice during this period comes from 12th- to 14th-century combs in the Netherlands (Schelvis, Reference Schelvis1994), while the mummy of Ferdinand II of Aragon, King of Naples (1467–1496 CE), was also infested with head lice (Fornaciari et al., Reference Fornaciari, Giuffra, Marinozzi, Picchi and Masetti2009). Mercury may have been used to control the infestation (Fornaciari et al., Reference Fornaciari, Marinozzi, Gazzaniga, Giuffra, Picchi, Giusiani and Masetti2011).

The fossil human lice records in post-medieval Britain are limited to London, at Cutler’s Gardens (Girling, Reference Girling1984). Body lice were recovered from the clothing of the shipwrecked crew of Willem Barentsz, who overwintered on Novaya Zemlya in the 16th century (Hakbijl and De Groot, Reference Hakbijl, De Groot, Ashworth, Buckland and Sadler1997), while head lice were identified in 17th- and 18th-century privy deposits at the Intendant’s Palace in Quebec City, Canada (Bain et al., Reference Bain, Bouchard-perron, Auger and Simoneau2009).

When the Norse colonized the North Atlantic islands, human lice were introduced with their hosts. Human presence at Goðatættur, on Papey off the east coast of Iceland, is verified on the basis of synanthropic taxa, including human lice (Buckland et al., Reference Buckland, Edwards, Blackford, Dugmore, Sadler, Sveinbjarnardóttir, Butlin and Roberts1995). P. humanus is present on medieval and post-medieval sites throughout Iceland (e.g. medieval Nesstofa (Amorosi et al., Reference Amorosi, Buckland, Magnússon, McGovern, Sadler, Luff and Rowley-Conwy1994), medieval and post-medieval Reykholt (Buckland Reference Buckland and Sveinbjarnardóttir2012), Stóraborg (Buckland and Perry, Reference Buckland and Perry1989), Svalbard (Amorosi et al., Reference Amorosi, Buckland, Ólafsson, Sadler, Skidmore, Morris and Rackham1992), Skriðuklaustur (Konráðsdóttir, Reference Konráðsdóttir2012) and early modern Vatnsfjörður (Forbes et al., Reference Forbes, Dussault and Bain2013)).

In Greenland, Norse human louse records range from the Eastern Settlement, including the bishop’s farm at Garðar (Buckland et al., Reference Buckland, Edwards, Panagiotakopulu and Schofield2009) and Tatsip Ataa in Narsaq (Dussault et al., Reference Dussault, Forbes and Bain2014), to the Western Settlement with a range of different sites (e.g. Tugmeralik (Sadler, Reference Sadler1990, Reference Sadler1991), Sandnes (Buckland and Sadler, Reference Buckland and Sadler1989) and Gården Under Sandet (Panagiotakopulu et al., Reference Panagiotakopulu, Skidmore and Buckland2007)) (Figure 2). A body louse was recovered from Sandnes, prompting comparisons with the contemporary account of the funeral of the Archbishop of Canterbury, Thomas Becket (ob. 1170 CE) (Sveinbjarnardóttir and Buckland, Reference Sveinbjarnardóttir and Buckland1982). One medieval chronicler reported that ‘The vermin boiled over like water in a simmering cauldron’, referring to lice crawling from the goat-haired shirt in which Becket’s corpse was dressed at his funeral (Zinsser, Reference Zinsser1935).

Figure 2.

Ectoparasites including Pediculus humanus L., the human louse (indicated by the red arrow) and Bovicola ovis Schrank, the sheep louse (indicated by the blue arrows) from Gården Under Sandet in the Western Settlement, Greenland.

However, the earliest records of human lice in Greenland come from the Saqqaq site of Qeqertasussuk ca. 2400–1400 BCE (Skidmore, pers. comm.). Thule Inuit records of head lice extend to the 15th-century Qilaqitsoq mummies (Bresciani et al., Reference Bresciani, Haarløv, Nansen, Moller, Hart Hansen and Gulløv1989). Head and body lice were recovered from 12th- to 14th-century Cape Grinnell and Qaqaitsut Thule houses (Dussault et al., Reference Dussault, Bain and LeMoine2014bb). In Alaska, human lice were recovered from the pre-contact Yup’ik site at Nunalleq ca. 14th–17th centuries (Forbes et al., Reference Forbes, Dussault and Bain2014) and from Aleutian mummies (Horne, Reference Horne1979).

When Ewing (Reference Ewing1924) studied 20 prehistoric American Indian mummies from the southwestern USA and Peru, he found evidence of lice and nits. He also highlighted morphological differences between lice from different regions and cited Kellogg’s (Reference Kellogg1913) hypothesis that an American derivative of the Old World species evolved using Ateles sp., Spider monkeys, as an intermediate host. Early research from the Americas involved North American Indian mummies in Arizona State Museum collections with lice and nits recovered from 2 Anasazi mummies from Painted Cave, ca. 1150–1250 CE, and 6 Hohokam mummies from Ventana Cave, ca. 1000–1400 CE (Birkby in El-Najjar et al., Reference El-Najjar, Mulinski, Reinhard, Cockburn, Cockburn and Reyman1998).

More recent work on mummies includes research on head lice from the Chiribaya culture (ca. 1000–1360 CE) in southern Peru and coastal northern Chile. Hair and skulls from mummies at several sites in southern Peru, Chiribaya Alta, Algodonal and El Yaral, yielded evidence of nits and adults. Male mummies from Chiribaya Alta and El Yaral had the highest density of infestations (Reinhard and Buikstra, Reference Reinhard and Buikstra2003). Additional records of head lice came from a Maitas Chiribaya mummy from the Azapa Valley, northern Chile, ca. 1100–1200 BP (Arriaza et al., Reference Arriaza, Standen, Reinhard, Araújo, Heukelbach and Dittmar2013) and a Chiribaya Baja mummy in the Moquegua Valley, southern Peru, ca. 800–1050 BP (Reinhard and Buikstra, Reference Reinhard and Buikstra2003). Head lice nits, sub-adults and adults, were recovered from most mummies along the coast of Arica-Camarones in northern Chile (Arriaza et al., Reference Arriaza, Standen, Reinhard, Araújo, Heukelbach and Dittmar2013). Similar infestations were recorded in 6 of 7 Chinchorro mummies from Camarones Site 15D (ca. 2000 BCE) (Rivera et al., Reference Rivera, Mumcuoglu, Matheny and Matheny2008). Severe infestations of head lice and nits were present on 22 Inca-period mummies from Camarones, ca. 1370–1450 CE (Arriaza et al., Reference Arriaza, Aravena, Núñez and Standen2022). Adult head lice, unhatched and hatched eggs, were present in 23 out of 41 combs from archaeological sites from northern Chile, ca. 1250–1650 CE (Arriaza et al., Reference Arriaza, Standen, Heukelbach, Cassman and Olivares2014).

Evidence for delousing and grooming comes from the recovery of head lice from coprolites at Lovelock Cave in Nevada dated to around 3000 years ago (Napton, Reference Napton1969; Reinhard and Bryant, Reference Reinhard and Bryant1992) and at Danger Cave in Utah ca. 130–300 CE (Fry, Reference Fry, Jennings and Sweeney1977). Head lice were also recovered from coprolites at La Cueva de los Muertos Chiquitos in Rio Zape ca. 600–800 CE, indicating similar practices (Reinhard et al., Reference Reinhard, Brooks, Brooks and Largent1989).

Pubic lice

Pthirus pubis (L.), the pubic or crab louse, is a cosmopolitan species, and its infestation is currently considered a sexually transmitted disease. It is found attached to pubic hairs and, in some cases, in other areas of the body, armpits, beard, eyelashes, etc. (Clay, Reference Clay and Smith1973). Pthirus probably moved from archaic gorillas to hominins around 3–4 million years ago (Reed et al., Reference Reed, Light, Allen and Kirchman2007). Its fossil record includes specimens in both hemispheres.

Two records of pubic lice were recovered from Britain. The earliest British record comes from Roman Carlisle, ca. 100–200 CE (Kenward, Reference Kenward1999), and another specimen was found in 18th-century London (Girling, Reference Girling1984). King Ferdinand of Aragon, in addition to head lice, had acquired a pubic lice infestation, and nits and adults were recovered from his mummy (Fornaciari et al., Reference Fornaciari, Giuffra, Marinozzi, Picchi and Masetti2009). The Norse probably introduced it to Iceland, and there are post-medieval records from 17th-century Reykholt (Buckland et al., Reference Buckland, Sadler, Sveinbjarnardóttir, Morris and Rackham1992). Additional records from Greenland were recovered from Thule houses at Cape Grinnell (Dussault et al., Reference Dussault, Bain and LeMoine2014b). From South America, there are 2 different recorded infestations, both from pre-Columbian prehistoric indigenous populations. Pubic lice nits were recovered from 2000-year-old mummies from San Pedro de Atacama in Chile, while P. pubis adults were found on a 1000-year-old Chiribaya Baja mummy from the Moquegua Valley, which was also infested with head lice (Rick et al., Reference Rick, Rocha, Dittmar, Coimbra, Reinhard, Bouchet, Ferreira and Araújo2002).

Bedbugs

Cimex lectularius L., the common bed bug, is currently a cosmopolitan ectoparasite, which feeds on the blood of humans during the night. It may also feed on the blood of other hosts, including bats, chickens and rabbits (Noble and Noble, Reference Noble and Noble1976). Its earliest record comes from the Workmen’s Village at pharaonic Amarna, a New Kingdom site with a span of around 2 decades in the 14th century BCE, from ca. 1347–1332 BCE (Panagiotakopulu and Buckland, Reference Panagiotakopulu and Buckland1999) (Figure 3A, B). In the British Isles, C. lectularius occurs in the Roman period at Lincoln Brayford Pool (Carrott et al., Reference Carrott, Issitt, Kenward, Large, McKenna and Skidmore1995) and in Carlisle Annetwell Street (Large et al., Reference Large and Kenward1998). Various other Roman specimens identified as Cimex sp., including Alcester in Warwickshire (Osborne, Reference Osborne1971), are likely to be C. lectularius. In addition to the British finds, there is a Roman record from Fréjus in the south of France (Ponel, Reference Ponel and Excoffon2011). Evidence from 10th-century deposits from Norwich (Kenward and Allison, Reference Kenward, Allison and Ayers1994) and from 11th-century deposits in York (Kenward and Hall, Reference Kenward and Hall1995) indicates that bedbugs were re-introduced or expanded their distribution in the medieval and post-medieval period. During the 18th century, London (Girling, Reference Girling1984) and Sheffield (Buckland, pers. comm.) provide further information on bedbugs. C. lectularius records from other parts of Europe include bedbugs infesting the bishop’s residence in Oslo during the 13th century (Marini et al., Reference Marini, Batchelor and Young2019). During the post-medieval period, bedbugs occur in the coffin of a 17th-century bishop buried at Lund Minster, in Sweden (Fägerström et al., Reference Fägerström, Buckland, Lemdahl, Karsten, Lagerås and Manhag2020), from 18th- to 19th-century Dublin in Ireland (Hall et al., Reference Hall, Jaques, Kenward, Carrott and Johnson2004) and in the privy at the Intendant’s Palace in Quebec (Bain et al., Reference Bain, Bouchard-perron, Auger and Simoneau2009), the latter attesting to the progressive cosmopolitanization of bedbugs.

Figure 3.

Ectoparasites from the Workmen’s Village at Amarna, Egypt: (A) head and thorax of Cimex lectularius L.; (B) Cimex lectularius L.; (C) specimens of Pulex irritans L., the human flea; (D) Ctenocephalides felis (Bouché), the cat flea.

In North America, Cimex antennatus Usinger & Ueshima, C. latipennis Usinger & Ueshima and C. pilosellus (Horváth) were identified in sediments from the Paisley Caves of south-central Oregon. C. antennatus was recovered from deposits dating to ca. 5100 cal BP, while the other 2 bedbug taxa were found in deposits dating to ca. 11 000–9400 cal BP (Adams and Jenkins, Reference Adams and Jenkins2017). These Nearctic bedbugs feed on bats, and although C. pilosellus may occasionally feed on humans, they are not human ectoparasites. Nevertheless, they provide insight into the environments of New World hunter-gatherers.

The human flea

The so-called human flea, Pulex irritans L., a species which is shared between man and other animals, including pigs, has been a companion of humans since at least 5700 years ago, and its origins probably lie in the Andean region of South America (Buckland and Sadler, Reference Buckland and Sadler1989). The early attempts to domesticate Cavia porcellus, the guinea pig (Larson and Fuller, Reference Larson and Fuller2014), may have led to the human flea moving to a new host, humans, a new source of blood meals. Finds of P. irritans/simulans from pre-Columbian Chiribaya guinea pig and dog mummies in the Moquegua Valley in Peru, ca. 900 CE (Dittmar et al., Reference Dittmar, Mamat, Whiting, Goldmann, Reinhard and Guillen2003), provide a glimpse of how the human flea may have moved onto humans, as does modern taxonomic research on P. irritans and its congener P. simulans Baker (Lareschi et al., Reference Lareschi, Venzal, Nava, Mangold, Portillo, Palomar-urbina and Oteo-revuelta2018).

The human flea probably started spreading via step-by-step gift exchange of furs, northwards in the Americas, before crossing the Beringian Strait to Eurasia, perhaps feeding on other new hosts. As it needs a relatively permanent nest in order to maintain breeding populations, its initial spread appears to be associated with the settlements of the Neolithic. Its earliest European record comes from Saint-Maximin-la-Sainte-Baume in France ca. 3656 BCE (Remicourt et al., Reference Remicourt, Andrieu-Ponel, Audibert, Baradat, Battentier, Belkacem, Blaise, Bonnardin, Dousse, Caverne, Fernandes, Gerbe, Girard, Lachenal, Lepère, Locatelli, Martin, Paravel, Parisot, Ponel, Pousset, Rué, Schmitt, Sénépart, Tallet and Thirault2012) (Figure 4). Additional Neolithic records were found at Schipluiden in the Netherlands, ca. 3500 cal BCE (Hakbijl, Reference Hakbijl, Louwe Kooijmans and Jongste2006). Human fleas were recorded on the South Tyrolean ice mummy called Ötzi of ca. 3350–3105 BCE (Schedl, Reference Schedl, Bortenschlager and Oeggl2000). A single sample from Chalain in the Jura, in France, ca. 3200 BCE, included 55 human fleas (Yvinec et al., Reference Yvinec, Ponel and Beaucournu2000). The northernmost point in its expansion appears to have been Skara Brae on Orkney ca. 3100–2500 BCE (Buckland and Sadler, Reference Buckland and Sadler1989). After the Neolithic, there is evidence for human fleas from pharaonic Amarna; the records are from both the elite part of the main city, the house of Ranefer, Akhenaten’s chief charioteer and from the Workmen’s Village (Panagiotakopulu, Reference Panagiotakopulu2001; Panagiotakopulu et al., Reference Panagiotakopulu, Buckland and Kemp2010) (Figure 3C). A single house floor sample from the Workmen’s Village yielded 35 human fleas (Panagiotakopulu, Reference Panagiotakopulu2001). These finds, the discussion of pestilence in the Amarna letters, and the fact that the main vector of Plague, Xenopsylla cheopis (Rothschild), is endemic in the Nile Valley led to a hypothesis about the origins and spread of bubonic plague, with human fleas playing a role in the spread of the epidemic (Panagiotakopulu, Reference Panagiotakopulu2004).

Figure 4.

The Holocene record of the human flea, Pulex irritans L.

During the Bronze Age, human fleas occurred in the Netherlands, at the site of Eigenblok 1500–1300 cal BCE (Van Dijk and Schelvis, Reference Van Dijk, Schelvis, Jongste and van Wijngaarden2002) and from a bog body from Emmer-Erfscheidenveen, in the Bourtangermoor, dating to between the 12th and 14th centuries BCE (Hakbijl, Reference Hakbijl and van der Sanden1990).

Evidence for human fleas has been recovered from a few Iron Age sites, St Patrick’s Isle on the Isle of Man (Tomlinson et al., Reference Tomlinson, Allison, Innes, Kenward and Freke2002), Dalhousie (Kenward et al., Reference Kenward, Large, Issitt, Crone and O’Sullivan1997) and Black Loch of Myrton (Allison et al., Reference Allison, Davies, Whitehouse, Crone and Cavers2026), both in Scotland, and Assendelver Polder in the Netherlands (Hakbijl, Reference Hakbijl1989).

The Roman Empire’s military campaigns and supply shipments for the army have led to additional records of P. irritans. Evidence from France comes from Gallo-Roman Amiens, Toufreville and Compans (Yvinec et al., Reference Yvinec, Ponel and Beaucournu2000). In Britain, human fleas have been recorded at several sites, e.g. London (Smith, Reference Smith2012) and York (Hall and Kenward, Reference Hall and Kenward1990) and from the Roman forts at Carlisle (Kenward et al., Reference Kenward, Allison, Morgan, Jones, Hutchinson and McCarthy1991) and Ribchester (Carrott et al., Reference Carrott, Fell, Large, Kenward, Kent, Buxton and Howard-davies2000) in northern England and from Bearsden in Scotland (Locke, Reference Locke and Breeze2016).

The larger number of sites and samples with human fleas comes from the medieval period. Although research from the early medieval period is limited, there are records of human fleas from various sites and contexts in Anglo-Scandinavian York (Buckland et al., Reference Buckland, Greig and Kenward1974; Kenward and Hall, Reference Kenward and Hall1995, Reference Kenward and Hall2000; Carrott et al., Reference Carrott, Dobney, Hall, Issitt, Jaques, Kenward, Large and McKenna1997). One of the most unusual finds involves adult human fleas recovered from a feather pillow placed in a boat burial at Øksnes in Vesterålen, northern Norway, ca. 10th century CE (Panagiotakopulu et al., Reference Panagiotakopulu, Buckland and Wickler2018). From London (Smith, Reference Smith2012) to Drum Castle in Scotland (Kenward et al., Reference Kenward, Large and Skidmore1995) and Glenarm Deer Park Farms in northern Ireland (Allison et al., Reference Allison, Hall and Kenward1999; Kenward et al., Reference Kenward, Hall, Allison, Carrott, Lynn and McDowell2011), and from Novgorod in Russia (Reilly, Reference Reilly, Brisbane, Makarov and Nosov2012) to farms in the Western Settlement of Greenland (Sadler, Reference Sadler1990; Panagiotakopulu et al., Reference Panagiotakopulu, Skidmore and Buckland2007), flea infestations were recovered from numerous sites with adequate preservation. Whether this expansion had to do with a warmer climate at the start of the medieval period, an expansion of trade networks, or both, remains unclear (Panagiotakopulu and Buckland, Reference Panagiotakopulu and Buckland2017). However, the spread of human fleas indicates a plethora of appropriate habitats both in urban and rural areas. It also provided a potential secondary vector for the spread of Plague when it reached epidemic proportions in Europe. In the post-medieval period, human fleas were present, and although the number of sites with fleas reduced (Panagiotakopulu and Buckland, Reference Panagiotakopulu and Buckland2017), they were recovered from sites as different as Yup’ik Nunalleq in Alaska (Forbes et al., Reference Forbes, Huchet, McManus-Fry, Coutouly, Masson-McLean, Masson-McLean, Ledger, Britton, Hillerdal and Knecht2020) and the Novaya Zemlya shipwreck (Hakbijl and De Groot, Reference Hakbijl, De Groot, Ashworth, Buckland and Sadler1997).

Fleas

A variety of animal ectoparasites have been recovered during archaeoentomological studies. What has become increasingly evident is that the domestication of various animals brought animal ectoparasites within areas of human habitation.

Cats were probably domesticated at the time of the beginnings of agriculture in the Fertile Crescent (Driscoll et al., Reference Driscoll, Menotti-raymond, Roca, Hupe, Johnson, Geffen, Trut, O’brien and Macdonald2007). As Egypt is one of the regions with a long history of taming cats (Linseele et al., Reference Linseele, Van Neer and Hendrickx2007), it comes as no surprise that the earliest record of cat ectoparasites, Ctenocephalides felis (Bouché), the cat flea, an efficient carrier of various diseases (Moore et al., Reference Moore, André, Šlapeta and Breitschwerdt2024), was recovered from the Workmen’s Village at Amarna (Panagiotakopulu, Reference Panagiotakopulu2004) (Figure 3D). Cats would have been frequenting the areas where the workers lived, although cat fleas also parasitize dogs and other animals (Moore et al., Reference Moore, André, Šlapeta and Breitschwerdt2024). Other records of cat fleas come from France during the medieval period, from Beauvais and Rennes (Yvinec et al., Reference Yvinec, Ponel and Beaucournu2000). A cat flea was found together with human lice and a human flea from a medieval comb in the Netherlands (Schelvis, Reference Schelvis1994), perhaps indicating that combs were also used to clean domestic animals and skins. Additional records were recovered from medieval and post-medieval sites in England, from various contexts in medieval Bristol ca. 12th–15th centuries CE (Smith, Reference Smith, Ford, Brady and Teague2017), and from post-medieval London, at Cutler’s Gardens (Girling, Reference Girling1984). Although the actual data are limited, likely due to taphonomic constraints, cat fleas were probably present with domestic cats in earlier settlements in Europe, from the Roman period onwards (de Martino et al., Reference de Martino, De Cupere, Rovelli, Serventi, Mouraud, Baldoni, Di Corcia, Geiger, Alhaique, Alves and Buitenhuis2025).

Dog fleas, Ctenocephalides canis (Curtis), were found from Gallo-Roman Amiens in France (Yvinec et al., Reference Yvinec, Ponel and Beaucournu2000). In England, Roman insect assemblages from 1st to 4th-century York yielded dog fleas (e.g. York General Accident (Hall and Kenward, Reference Hall and Kenward1990); York Palmer Lane (Carrott et al., Reference Carrott, Dobney, Hall, Kenward, Milles and Nicholson1992)). Medieval Beauvais and Rennes in France (Yvinec et al., Reference Yvinec, Ponel and Beaucournu2000) and sites in England (e.g. London (Smith, Reference Smith2012), Hull (Jaques et al., Reference Jaques, Hall, Kenward and Carrott2003), Beverley (Kenward and Carrott, Reference Kenward and Carrott2003)) and the late medieval Newport shipwreck on the banks of the River Usk in South Wales (Smith, Reference Smith2014) produced dog fleas. These records clearly indicate the presence of dogs, domesticated at least since the Upper Palaeolithic, within settlements at least since the Roman period, if not earlier.

Additional flea taxa include the northern rat flea, Nosopsyllus fasciatus (Bosc d’Antic) from Iron Age London (Smith, Reference Smith, Grant, Norcott and Stevens2012b), Anglo-Scandinavian Coppergate at York (Kenward and Hall, Reference Kenward and Hall1995) and Glenarm (Allison et al., Reference Allison, Hall and Kenward1999). Ctenophthalmus nobilis (Rothschild) was recovered alongside N. fasciatus in York. Both taxa are found on rodents and have a capacity to spread both Bartonella and Rickettsia bacteria; N. fasciatus is an efficient vector of Plague.

The only archaeological record of the European chicken flea, Ceratophyllus gallinae (Schrank), the most widespread bird ectoparasite in Europe, which became cosmopolitan with the introduction of poultry, was found in the Anglo-Saxon Holywell Row cemetery in Suffolk (Lethbridge Reference Lethbridge1931). The earliest specimens of bird ectoparasites are 2 species of swallow fleas, C. petrochelidoni Wagner and C. celsus Jordan, recovered from Lovelock Cave in Nevada ca. 2600–2400 BCE (Nelson, Reference Nelson1972). Another bird ectoparasite, the bird nest flea, C. vagabundus (Boheman), was found at Qeqertasussuk in Greenland (Böcher and Fredskild, Reference Böcher and Fredskild1993), while the duck flea C. garei has been recovered from early modern assemblages in Iceland (Forbes et al., Reference Forbes, Dussault and Bain2013).

Animal lice

Bovicola bovis (L.) is one of the most widespread livestock lice, often found on cattle, on the shoulders, back and the base of the tail (Noble and Noble, Reference Noble and Noble1976). The cattle louse may be the vector for a range of diseases, and studies have confirmed that it successfully spreads Staphylococcus aureus, Escherichia coli and Listeria (Nahorna and Proskurina, Reference Nahorna and Proskurina2020). Its earliest record comes from Thayngen-Weier in Neolithic Switzerland ca. 2900–2800 BCE (Nielsen et al., Reference Nielsen, Mahler and Rasmussen2000). Further records come from the Iron Age sites of Goldcliff in South Wales (Smith et al., Reference Smith, Osborne, Barrett, Bell, Caseldine and Neumann2000) and Black Loch of Myrton in Scotland (Allison et al., Reference Allison, Davies, Whitehouse, Crone and Cavers2026).

Insect assemblages from the Iron Age Middendelfland, Zuid-Holland, in the Netherlands included 3 ectoparasites on cattle, B. bovis, Solenopotes capillatus (Enderlein) and Linognathus vituli (L.) (Schelvis and Koot, Reference Schelvis and Koot1995), all of which are now cosmopolitan. L. vituli was first described from cattle in Sweden (Kim et al., Reference Kim, Pratt and Stojanovich1986). Additional evidence for B. bovis comes from a single Roman site in North Yorkshire, from Viking Ribe (Allison and Kenward, Reference Allison, Kenward and Sindbæk2022), Anglo-Scandinavian York (Kenward and Hall, Reference Kenward and Hall1995), 14th-century Skegness (Buckland unpubl.) and from Glenarm Deer Park Farms in northern Ireland (Allison et al., Reference Allison, Hall and Kenward1999). Another cattle ectoparasite, Haematopinus eurysternus (Denny), which thrives during the winter, when animals are stalled in barns (Kim et al., Reference Kim, Pratt and Stojanovich1986), was found from the Norse farm Gården Under Sandet, in the Western Settlement, Greenland (Panagiotakopulu et al., Reference Panagiotakopulu, Skidmore and Buckland2007).

Sheep biting lice, Bovicola ovis Schrank (= Damalinia ovis (L.)), have probably been associated with sheep since at least the time they were first domesticated. However, their earliest records are much later, from the Iron Age, at the Black Loch of Myrton (Allison et al., Reference Allison, Davies, Whitehouse, Crone and Cavers2026), Middendelfland (Schelvis and Koot, Reference Schelvis and Koot1995) and from Roman Healam Bridge (Jaques et al., Reference Jaques, Foster, Walker, Martin, Hall, Allison, Carrott, Ambrey, Fell, Fraser, Ross, Speed and Wood2017) (Figure 5). The lack of early records is probably a combined result of preservation and the nature of the contexts studied. Sheep lice became widespread in the medieval period, and they are present at sites across northern Europe. Records include Selby and various sites in York, England (Buckland unpubl.; Kenward and Hall, Reference Kenward and Hall1995), Glenarm in northern Ireland (Allison et al., Reference Allison, Hall and Kenward1999), Ypres in Belgium (Schelvis, Reference Schelvis, Dewilde, Ervynck and Wielemans1999) and Ribe in Denmark (Allison and Kenward, Reference Allison, Kenward and Sindbæk2022). In the North Atlantic region, specimens come from Stóraborg and post-medieval Skálholt in Iceland (Buckland and Perry, Reference Buckland and Perry1989; Konráðsdóttir et al., Reference Konráðsdóttir, Panagiotakopulu and Lucas2021) and from the Western Settlement in Greenland, notably Sandnes and Gården Under Sandet (Sadler, Reference Sadler1990; Panagiotakopulu et al., Reference Panagiotakopulu, Skidmore and Buckland2007) (Figure 2).

Figure 5.

The Holocene record of Bovicola ovis Schrank, the sheep louse.

With the exception of the evidence from the Iron Age crannog site of Cults Loch (Allison, Reference Allison, Cavers and Crone2018), the presence of the obligate ectoparasite on goats, Bovicola caprae (Gurlt), is exclusive to the medieval period from rural and urban sites which had adequate preservation and produced other ectoparasites, e.g. Glenarm (Allison et al., Reference Allison, Hall and Kenward1999), London (Smith, Reference Smith2012) and York (Kenward and Hall, Reference Kenward and Hall1995). During the same period, Gården Under Sandet in Greenland (Panagiotakopulu et al., Reference Panagiotakopulu, Skidmore and Buckland2007) yielded records of B. caprae and an additional goat ectoparasite, Linognathus stenopsis (Burmeister).

The earliest record of the horse louse, Bovicola equi (Denny), is recovered from Roman York, General Accident, from the 2nd century CE (Hall and Kenward, Reference Hall and Kenward1990). During the medieval period, records of horse lice come from Ribe (Allison and Kenward, Reference Allison, Kenward and Sindbæk2022), Glenarm (Allison et al., Reference Allison, Hall and Kenward1999) and York (Kenward and Hall, Reference Kenward and Hall1995). In Anglo-Scandinavian Micklegate at York, the insect assemblages included Haematopinus apri Goureau, the boar louse (Kenward and Hall, Reference Kenward and Hall2000), which lives on wild boar and old or hybrid breeds of swine. Glenarm had the highest numbers of specimens of H. apri with 43 individuals from a single context and 94 overall, indicating the presence of pigs in the structure where the ectoparasites were recovered (Allison et al., Reference Allison, Hall and Kenward1999). During the Anglo-Scandinavian period, Coppergate in York also provides a record of a cat biting louse, Felicola subrostratus (Burmeister) (Kenward and Hall, Reference Kenward and Hall1995).

The dog louse Trichodectes canis (DeGeer) was recovered from Nunalleq in Alaska (Forbes et al., Reference Forbes, Huchet, McManus-Fry, Coutouly, Masson-McLean, Masson-McLean, Ledger, Britton, Hillerdal and Knecht2020), while another dog louse, Linognathus setosus Von Olfers, came from Cape Grinnell and Qaqaitsut in Greenland (Dussault et al., Reference Dussault, Bain and LeMoine2014b) together with bird ectoparasites, Nirmus sp. From the midden of the 14th–17th-century Thule site at Kangeq in southwest Greenland, there is evidence for bird ectoparasites, Arctornithophilus sp. and Ricinus sp. (Panagiotakopulu et al., Reference Panagiotakopulu, Schofield, Vickers, Edwards and Buckland2020).

A single record of a rodent louse was recovered from a comb found in the Judean Desert, dated to ca. 1660–1950 CE. Acomys hair recovered from the same comb led to the hypothesis that it was used to groom a spiny mouse (Mumcuoglu et al., Reference Mumcuoglu, Sukenik and Bar-oz2020). A record of a flea that lives on moles, Hystrichopsylla talpae (Curtis), was found at medieval Glenarm (Allison et al., Reference Allison, Hall and Kenward1999).

Animal lice from South America include another record of F. subrostratus (F. felis), recovered from felid coprolites dated ca. 1610–1860 years BP from Furna do Estrago in Brazil (Guerra et al. Reference Guerra, Duarte, Oliviera, Mello and Serra-Freire2001), presumably from the fur of one of the native small cats of South America. Specimens of the South American camelid sucking lice Microthoracius praelongiceps (Neumann) and M. minor were recovered from late Chiribaya mummified llamas (Astos et al., Reference Astos, Leguía and Wheeler1999). In addition to P. irritans/simulans from pre-Columbian guinea pig mummies, the chewing louse Gliricolla porcelli (Schrank), an ectoparasite of guinea pigs, was recovered together with P. simulans fleas and haematophagous mites of the genus Ornithonyssus, from C. porcellus from El Yaral in Peru (Dittmar, Reference Dittmar2000).

Louse flies (keds)

One of the earliest records of louse flies (Hippoboscidae), Lipoptena cervi (L.), the deer ked, was found in skins associated with the mummy of Ötzi in the Alps (Gothe and Schöl, Reference Gothe and Schöl1994). L. cervi parasitizes a variety of cervids and is associated with zoonotic diseases (Välimäki et al., Reference Välimäki, Kaitala, Madslien, Härkönen, Várkonyi, Heikkilä, Jaakola, Ylönen, Kortet and Ytrehus2011). A later record comes from medieval York (Hall et al., Reference Hall, Kenward, Rowland, Jaques and Carrott2001). Another species of the same family, the louse fly Hippobosca longipennis F., which infests dogs and various wild animals, was recovered from a Roman dog mummy from el-Deir, 1st–4th century CE, in Kharga Oasis, Egypt (Huchet et al., Reference Huchet, Callou, Lichtenberg and Dunand2013). Although outside the scope of this paper, it is worth noting the presence of an extinct endoparasitic botfly, Oestridae, Cobboldia russanovi Grunin, associated with a frozen late Pleistocene mammoth in Siberia (Grunin, Reference Grunin1973).

The most frequent louse fly in archaeological deposits is the sheep ked, Melophagus ovinus L., which currently has a cosmopolitan distribution. It largely infests sheep, and occasionally also goats and cattle and may result in poor animal health (Small, Reference Small2005). The larvae of M. ovinus are cemented to the fleece, and high infestations of sheep keds are associated with malnutrition (Cheyne et al., Reference Cheyne, Foster, Spence, Jewell, Milner and Boyd1974). Both puparia and adults of sheep keds are preserved in the archaeological record. The earliest evidence for the species comes from Late Bronze Age northern Italy, ca. 1650 BCE, where puparia of sheep keds were recovered from the site Oppeano 4D (Nicosia et al., Reference Nicosia, Dal Corso, D’aquino, Baldan, Bortolini, Battistel and Polisca2025). In Britain, their earliest records are recovered from Iron Age Buiston in Scotland (Kenward et al., Reference Kenward, Hill, Jaques, Kroupa, Large and Crone2000). There are various records from the Roman period in England (e.g. from Lincoln (Carrott et al., Reference Carrott, Issitt, Kenward, Large, McKenna and Skidmore1995) and Carlisle (Kenward et al., Reference Kenward, Allison, Morgan, Jones, Hutchinson and McCarthy1991)). Sheep keds were widespread in the medieval period. There are various sites in Britain with relevant evidence. Sheep keds were recovered from various sites in London (Smith, Reference Smith2012), Hull Blanket Row (Carrott et al., Reference Carrott, Hall, Jaques, Johnstone, Kenward and Rowland2001), Magistrates Court (Hall et al., Reference Hall, Carrott, Jaques, Johnstone, Kenward, Large and Usai2000) and various sites in York (Anglo-Scandinavian deposits in Micklegate (Kenward and Hall, Reference Kenward and Hall2000), Anglo-Scandinavian Parliament Street (Hall and Kenward, Reference Hall and Kenward2000), Low Petergate (Hall et al., Reference Hall, Kenward, Girvan and McKenna2007), Walmgate (Jaques et al., Reference Jaques, Hall, Kenward, Rowland and Carrott2001; Hall et al., Reference Hall, Kenward, Jaques, Carrott and Rowland2002), etc.). The evidence extends to Fishamble Street in Dublin, Ireland and Ypres in Belgium. M. ovinus is a frequent ectoparasite on Norse sites in the North Atlantic region, where it would have been introduced with domestic animals. Nine sites from medieval Iceland, including Bessastaðir (Amorosi et al., Reference Amorosi, Buckland, Ólafsson, Sadler, Skidmore, Morris and Rackham1992), Goðatættur (Buckland et al., Reference Buckland, Edwards, Blackford, Dugmore, Sadler, Sveinbjarnardóttir, Butlin and Roberts1995) and Nesstofa (Amorosi et al., Reference Amorosi, Buckland, Magnússon, McGovern, Sadler, Luff and Rowley-Conwy1994), yielded evidence for M. ovinus. From the post-medieval period in Iceland, large numbers of M. ovinus were recorded from Stóraborg (Buckland and Perry, Reference Buckland and Perry1989) and Skálholt (Konráðsdóttir et al., Reference Konráðsdóttir, Panagiotakopulu and Lucas2021); washing of wool was assumed to have taken place with the use of stale urine collected in barrels, a practice recorded in the ethnographic record to get rid of sheep keds (Jónasson, Reference Jónasson1961).

Ticks

One of the earliest records of ticks from a synanthropic context comes from the Black Loch of Myrton, where 1 adult and several nymphs of the so-called castor bean tick, Ixodes ricinus L., were recovered, perhaps brought onto the site with vegetation (Allison et al., Reference Allison, Davies, Whitehouse, Crone and Cavers2026). I. ricinus has a wide distribution which ranges from Scandinavia to North Africa and from the Iberian Peninsula to parts of Russia. It prefers humid, deciduous and mixed woodland habitats. It can spread a variety of pathogens to animals, and in Europe, it is the principal vector of Lyme borreliosis and tick-borne encephalitis to humans (Kahl and Gray, Reference Kahl and Gray2023). Additional records of adults and nymphs of this species were present at Anglo-Scandinavian York (Kenward and Hall, Reference Kenward and Hall1995) and Glenarm in northern Ireland (Allison et al., Reference Allison, Hall and Kenward1999). Another tick species, Rhipicephalus sanguineus Latreille, the brown dog tick, was found on a Roman dog mummy from el-Deir in Egypt (Huchet et al., Reference Huchet, Callou, Lichtenberg and Dunand2013). It parasitizes dogs and other mammals and occasionally also bites humans. A single specimen of R. turanicus Pomerantzev, an ectoparasite of different animals which may occasionally bite humans, was recovered from a 17th- to early 20th-century comb from the Judean desert (Mumcuoglu et al., Reference Mumcuoglu, Sukenik and Bar-oz2020).

The best evidence for ticks associated with humans comes from human coprolites from the Antelope Cave site in northern Arizona used by Virgin Anasazi people (Ancestral Puebloans), ca. 680–960 CE. The coprolites contained specimens of crushed ticks, Dermacentor andersoni Stiles (Johnson et al., Reference Johnson, Reinhard, Sianto, Araújo, Gamder and Janovy2008), which were probably killed and swallowed by their hosts. D. andersoni, the Rocky Mountain wood tick, lives on a variety of mammals including humans and is the principal vector of Rocky Mountain spotted fever. Other records from North America include the soft-bodied tick Otobius megnini (Dugès), which parasitizes a variety of animals and could bite humans. It was recovered from the Paisley Caves, in deposits which range from the start of the Holocene (ca. 12672–10390 BP) to the mid Holocene (5520 BP and 4990 BP) (Adams and Jenkins, Reference Adams and Jenkins2020).

In South America, hard ticks, Amblyomma sp., were recovered from felid coprolites from Furna do Estrago in Brazil, together with soft unidentified ticks (Guerra et al. Reference Guerra, Gazêta, Amorim, Duarte and Serra-Freire2003). Additional taxa of ticks, Amblyomma parvitarsum Neumann and Ornithodoros sp., were found from a rock shelter with evidence for hunter-gatherers at Peñas de la Cruz, ca. 8000 BP, in northwestern Argentina, while Argas cf. neghmei Kohls & Hoogstraal was found from Cueva de los Corrales 1, which was used by agropastoralists ca. 770–490 BP in the same region (Nava et al., Reference Nava, Martínez, Arreguez and Guglielmone2013).