New records and a new species of chewing lice (Phthiraptera, Amblycera, Ischnocera) found on Columbidae (Columbiformes) in Pakistan

The chewing lice (Phthiraptera) of Columbidae (Columbiformes) from Pakistan are studied. Six species of chewing lice with new host records are recorded and one new species of the genus Colpocephalum is described from Columba livia in the Karachi region. All the columbid chewing lice from Pakistan are keyed out and the new species is illustrated and compared with the closest allied species.     

Chewing Lice of Swan Geese (Anser cygnoides): New Host-Parasite Associations

Chewing lice (Phthiraptera) that parasitize the globally threatened swan goose Anser cygnoides have been long recognized since the early 19th century, but those records were probably biased towards sampling of captive or domestic geese due to the small population size and limited distribution of its wild hosts. To better understand the lice species parasitizing swan geese that are endemic to East Asia, we collected chewing lice from 14 wild geese caught at 3 lakes in northeastern Mongolia. The lice were morphologically identified as 16 Trinoton anserinum (Fabricius, 1805), 11 Ornithobius domesticus Arnold, 2005, and 1 Anaticola anseris (Linnaeus, 1758). These species are known from other geese and swans, but all of them were new to the swan goose. This result also indicates no overlap in lice species between older records and our findings from wild birds. Thus, ectoparasites collected from domestic or captive animals may provide biased information on the occurrence, prevalence, host selection, and host-ectoparasite interactions from those on wild hosts.     

Sea louse infection of juvenile sockeye salmon in relation to marine salmon farms on Canada's west coast

Background

Pathogens are growing threats to wildlife. The rapid growth of marine salmon farms over the past two decades has increased host abundance for pathogenic sea lice in coastal waters, and wild juvenile salmon swimming past farms are frequently infected with lice. Here we report the first investigation of the potential role of salmon farms in transmitting sea lice to juvenile sockeye salmon (Oncorhynchus nerka).

Methodology/Principal Findings

We used genetic analyses to determine the origin of sockeye from Canada''s two most important salmon rivers, the Fraser and Skeena; Fraser sockeye migrate through a region with salmon farms, and Skeena sockeye do not. We compared lice levels between Fraser and Skeena juvenile sockeye, and within the salmon farm region we compared lice levels on wild fish either before or after migration past farms. We matched the latter data on wild juveniles with sea lice data concurrently gathered on farms. Fraser River sockeye migrating through a region with salmon farms hosted an order of magnitude more sea lice than Skeena River populations, where there are no farms. Lice abundances on juvenile sockeye in the salmon farm region were substantially higher downstream of farms than upstream of farms for the two common species of lice: Caligus clemensi and Lepeophtheirus salmonis, and changes in their proportions between two years matched changes on the fish farms. Mixed-effects models show that position relative to salmon farms best explained C. clemensi abundance on sockeye, while migration year combined with position relative to salmon farms and temperature was one of two top models to explain L. salmonis abundance.

Conclusions/Significance

This is the first study to demonstrate a potential role of salmon farms in sea lice transmission to juvenile sockeye salmon during their critical early marine migration. Moreover, it demonstrates a major migration corridor past farms for sockeye that originated in the Fraser River, a complex of populations that are the subject of conservation concern.     

Correlated evolution of host and parasite body size: tests of Harrison's rule using birds and lice

Large-bodied species of hosts often harbor large-bodied parasites, a pattern known as Harrison's rule. Harrison's rule has been documented for a variety of animal parasites and herbivorous insects, yet the adaptive basis of the body-size correlation is poorly understood. We used phylogenetically independent methods to test for Harrison's rule across a large assemblage of bird lice (Insecta: Phthiraptera). The analysis revealed a significant relationship between louse and host size, despite considerable variation among taxa. We explored factors underlying this variation by testing Harrison's rule within two groups of feather-specialist lice that share hosts (pigeons and doves). The two groups, wing lice (Columbicola spp.) and body lice (Physconelloidinae spp.), have similar life histories, despite spending much of their time on different feather tracts. Wing lice showed strong support for Harrison's rule, whereas body lice showed no significant correlation with host size. Wing louse size was correlated with wing feather size, which was in turn correlated with overall host size. In contrast, body louse size showed no correlation with body feather size, which also was not correlated with overall host size. The reason why body lice did not fit Harrison's rule may be related to the fact that different species of body lice use different microhabitats within body feathers. More detailed measurements of body feathers may be needed to explore the precise relationship of body louse size to relevant components of feather size. Whatever the reason, Harrison's rule does not hold in body lice, possibly because selection on body size is mediated by community-level interactions between body lice.     

Phylogeny and classification, origins, and evolution of host associations of lice

Lice are highly successful ectoparasites. Most species of mammals and birds are infested by at least 1 but up to 6 species of lice. Current opinion is that lice evolved from free-living Psocoptera (booklice, barklice and psocids). It is generally agreed that there are 4 main groups of lice: Anoplura, Amblycera, Ischnocera and Rhyncophthirina. In contrast, there is no agreement on the phylogenetic relationships of these groups and their classification. In particular, there is much debate over the validity of the taxon Mallophaga, which is almost certainly paraphyletic. For many years the sister-group of the Boopiidae, which almost exclusively infest Australasian marsupials, was thought to be a group of lice that now infest marsupials in South America. This, however, is almost certainly incorrect; the sister-group of the Boopiidae probably contains bird-infesting lice from the Menoponidae (Amblycera). Thus, menoponid lice transferred from birds to mammals and from these arose the Boopiidae. Transfers of lice between mammals and birds have occurred on other occasions during the evolution of the lice; 2 of the 4 main groups of lice, the Ischnocera and Amblycera, contain families that infest birds and families that infest mammals. Strict cospeciation and coevolution was thought to predominate among the lice; however, detailed studies indicate this to be incorrect. Consequently, the axiom that lice and their hosts invariably coevolve should be abandoned. Ironically, biologists may learn more about the evolutionary biology of hosts when host-switching has occurred. Some evidence exists for competition between species of lice; this interaction may determine whether or not the transfer of a species of louse to an atypical hose (a potential host-switch) is successful. Thus, the extincion of populations of lice (that result in uninfested hosts) may facilitate host-switching and perhaps the evolution of new taxa of lice. In contrast, extinction of hosts unfortunately often leads to the extinction of species of lice.     

Oust the louse: leaping behaviour removes sea lice from wild juvenile sockeye salmon Oncorhynchus nerka

We conducted a manipulative field experiment to determine whether the leaping behaviour of wild juvenile sockeye salmon Oncorhynchus nerka dislodges ectoparasitic sea lice Caligus clemensi and Lepeophtheirus salmonis by comparing sea‐lice abundances between O. nerka juveniles prevented from leaping and juveniles allowed to leap at a natural frequency. Juvenile O. nerka allowed to leap had consistently fewer sea lice after the experiment than fish that were prevented from leaping. Combined with past research, these results imply potential costs due to parasitism and indicate that the leaping behaviour of juvenile O. nerka does, in fact, dislodge sea lice.     

Sea-lice infection models for fishes

As free-living sea-lice larvae are difficult to sample directly, lice abundances on fish have recently been used to study larvae in the water. In the KLV problem, juvenile wild salmon migrate past a salmon farm, and the change of infection with distance along the migration route is used to estimate larvae production from the farm. In the farm problem, time-varying infection of sea-cage fish is used to estimate the time-variation of free-living larvae in waters near the farm. Both inverse problems require good forward models for infection. In the farm problem, hosts are relatively large and lice pathogenesis is seldom mortal, whereas in the KLV problem hosts are small and lice-induced host mortality can affect lice abundance; thus, infection models for the farm problem are special cases of models for the KLV problem. Here I give an infection model for the KLV problem that explicitly includes lice clumping and host mortality, showing that Krkosek et al. (Proc R Soc B 272:689-696, 2005) (KLV) probably underestimated larvae production by the salmon farm, and further, that if lice development rates were known from laboratory data, lice abundance field data could be directly inverted for lice-induced host mortality during migration. If lice-induced host mortality is negligible, or if lice are Poisson distributed, infection models of arbitrary complexity reduce to Erlang models. I give two useful Erlang models with their solutions for non-zero initial conditions.     

Chewing lice from high‐altitude and migrating birds in Yunnan,China, with descriptions of two new species of Guimaraesiella

In total, 366 birds representing 55 species in 24 families and eight orders, were examined for chewing lice (Phthiraptera: Amblycera, Ischnocera) in two high‐altitude localities in Yunnan Province, China. In Ailaoshan, almost all of the birds examined were resident passeriforms, of which 36% were parasitized by chewing lice. In Jinshanyakou, most birds were on migration, and included both passerine and non‐passerine birds. Of the passerine birds caught in Jinshanyakou, only one bird (0.7%) was parasitized by chewing lice. The prevalence of Myrsidea and Brueelia‐complex lice on birds caught in Ailaoshan was higher than in previous reports. Of the chewing lice identifiable to species level, three represent new records for China: Actornithophilus hoplopteri (Mjöberg, 1910), Maculinirmus ljosalfar Gustafsson & Bush, 2017 and Quadraceps sinensis Timmermann, 1954. In total, 17 new host records are included, of which we describe two as new species in the Brueelia‐complex: Guimaraesiella (Cicchinella) ailaoshanensis sp. nov. ex Schoeniparus dubius dubius (Hume, 1874) and G. (C.) montisodalis sp. nov. ex Fulvetta manipurensis tonkinensis Delacour & Jabouille, 1930. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:9FC3D8EE‐2CED‐4DBE‐A1DB‐471B71260D27 .     

Of lice and math: using models to understand and control populations of head lice

In this paper we use detailed data about the biology of the head louse (pediculus humanus capitis) to build a model of the evolution of head lice colonies. Using theory and computer simulations, we show that the model can be used to assess the impact of the various strategies usually applied to eradicate head lice, both conscious (treatments) and unconscious (grooming). In the case of treatments, we study the difference in performance that arises when they are applied in systematic and non-systematic ways. Using some reasonable simplifying assumptions (as random mixing of human groups and the same mobility for all life stages of head lice other than eggs) we model the contagion of pediculosis using only one additional parameter. It is shown that this parameter can be tuned to obtain collective infestations whose characteristics are compatible with what is given in the literature on real infestations. We analyze two scenarios: One where group members begin treatment when a similar number of lice are present in each head, and another where there is one individual who starts treatment with a much larger threshold ("superspreader"). For both cases we assess the impact of several collective strategies of treatment.     

Potential natural vegetation and pre‐anthropic pollen records on the Azores Islands in a Macaronesian context

This paper discusses the concept of potential natural vegetation (PNV) in the light of the pollen records available to date for the Macaronesian biogeographical region, with emphasis on the Azores Islands. The classical debate on the convenience or not of the PNV concept has been recently revived in the Canary Islands, where pollen records of pre‐anthropic vegetation seemed to strongly disagree with the existing PNV reconstructions. Contrastingly, more recent PNV model outputs from the Azores Islands show outstanding parallelisms with pre‐anthropic pollen records, at least in qualitative terms. We suggest the development of more detailed quantitative studies to compare these methodologies as an opportunity for improving the performance of both. PNV modelling may benefit by incorporating empirical data on past vegetation useful for calibration and validation purposes, whereas palynology may improve past reconstructions by minimizing interpretative biases linked to differential pollen production, dispersal and preservation.     

Individual fish tank arrays in studies of Lepeophtheirus salmonis and lice loss variability

In studies of the salmon louse Lepeophtheirus salmonis (Kr?yer, 1837), experimental design is complicated by a highly variable and unpredictable lice loss among common experimental tanks and a substantial rate of host transfer within tanks. When fish hosting L. salmonis are maintained in individual tanks, unspecific effects such as host transfer, louse predation by cohabitant hosts and agonistic host interactions are excluded. This study suggests that it is possible to maintain Atlantic salmon Salmo salar infected with L. salmonis in an array of small, single fish tanks and, by doing so, provide an experimental system in which the loss of motile pre-adult and adult stages of L. salmonis is predictable. Here, lice can be collected shortly after detachment for detailed studies or to provide mortality curves of lice from individual fish. This represents an experimental approach improving precision in studies of L. salmonis, such as drug and vaccine efficacy assays, RNA interference (RNAi) studies and host-parasite interactions. The natural loss of pre-adult/adult L. salmonis from the system was higher for males than females. The loss of females appeared to be a process somewhat selective against large individuals. Inherent qualities of the host appeared to be of little significance in explaining the variability in loss of preadult/adult lice.     

Ancient Human Genomes and Environmental DNA from the Cement Attaching 2,000-Year-Old Head Lice Nits

Over the past few decades, there has been a growing demand for genome analysis of ancient human remains. Destructive sampling is increasingly difficult to obtain for ethical reasons, and standard methods of breaking the skull to access the petrous bone or sampling remaining teeth are often forbidden for curatorial reasons. However, most ancient humans carried head lice and their eggs abound in historical hair specimens. Here we show that host DNA is protected by the cement that glues head lice nits to the hair of ancient Argentinian mummies, 1,500–2,000 years old. The genetic affinities deciphered from genome-wide analyses of this DNA inform that this population migrated from north-west Amazonia to the Andes of central-west Argentina; a result confirmed using the mitochondria of the host lice. The cement preserves ancient environmental DNA of the skin, including the earliest recorded case of Merkel cell polyomavirus. We found that the percentage of human DNA obtained from nit cement equals human DNA obtained from the tooth, yield 2-fold compared with a petrous bone, and 4-fold to a bloodmeal of adult lice a millennium younger. In metric studies of sheaths, the length of the cement negatively correlates with the age of the specimens, whereas hair linear distance between nit and scalp informs about the environmental conditions at the time before death. Ectoparasitic lice sheaths can offer an alternative, nondestructive source of high-quality ancient DNA from a variety of host taxa where bones and teeth are not available and reveal complementary details of their history.     

Flying squirrels and their ectoparasites: disseminators of epidemic typhus

Information gathered during the past decade indicates that the eastern flying squirrel, Glaucomys volans, is a zoonotic reservoir of Rickettsia prowazekii - causative agent of louse-borne (epidemic) typhus. The sporadic cases o f typhus that have occurred in the USA in association with flying squirrels provide evidence that flying squirrels can transmit R. prowazekii infection to humons. Strains of R. prowazekii, isolated from flying squirrels multiply readily in human body lice, but flying squirrel lice, although readily infected, are very host specific and tend not to bite humans. It may be that the infection is spread to humans in infective ectoporasite faeces aerosolized when the flying squirrels groom themselves. As Joseph McDade emphasizes in this article, current concepts of typhus epidemiology and control must be re-evaluated to take into account this zoonotic aspect.     

Chewing Lice (Insecta: Phthiraptera) from Migratory Birds of the Curonian Spit

The fauna of chewing lice (Insecta: Phthiraptera) from migratory birds was studied in 2008–2009 and 2017 on the Curonian Spit, at Rybachy Biological Station of the Zoological Institute, Russian Academy of Sciences. Altogether, 35 species of chewing lice were collected off 2010 birds of 65 species. An annotated list of species is presented, including 11 species new to the Russian Federation, 29 species new to the Northwest of Russia, and 2 new host records: Menacanthus eurysternus from Carduellis carduellis and Ricinus frenatusfromCarduellis spinus.     

Allee effects may slow the spread of parasites in a coastal marine ecosystem

Allee effects are thought to mediate the dynamics of population colonization, particularly for invasive species. However, Allee effects acting on parasites have rarely been considered in the analogous process of infectious disease establishment and spread. We studied the colonization of uninfected wild juvenile Pacific salmon populations by ectoparasitic salmon lice (Lepeophtheirus salmonis) over a 4-year period. In a data set of 68,376 fish, we observed 85 occurrences of precopular pair formation among 1,259 preadult female and 613 adult male lice. The probability of pair formation was dependent on the local abundance of lice, but this mate limitation is likely offset somewhat by mate-searching dispersal of males among host fish. A mathematical model of macroparasite population dynamics that incorporates the empirical results suggests a high likelihood of a demographic Allee effect, which can cause the colonizing parasite populations to die out. These results may provide the first empirical evidence for Allee effects in a macroparasite. Furthermore, the data give a rare detailed view of Allee effects in colonization dynamics and suggest that Allee effects may dampen the spread of parasites in a coastal marine ecosystem.     

External parasites of raptors (Falconiformes and Strigiformes): identification in an ex situ population from Mexico

Raptorial birds harbor a variety of ectoparasites and the mayority of them are host specific. The aim of this study was to identify the ectoparasites of captive birds of prey from Mexico, as well as to verify their impact in the health of infested birds. Raptorial birds were confiscated and kept in captivity at the Centro de Investigación y Conservación de Vida Silvestre (CIVS) in Los Reyes La Paz, Mexico State. Seventy-four birds of prey (66 Falconiformes and eigth Strigiformes) of 15 species were examined for the presence of ectoparasites. We examined both juvenile and adult birds from both sexes. The overall prevalence was 16.2%; 66.7% of raptors were infested with a single type of external parasite. Lice were the most prevalent ectoparasites (91.7%), followed by feather mites and fleas (8.3%). Degeeriella fulva (72.7%), Craspedorrhynchus sp. (45.4%) and Strigiphilus aitkeni (9.1%) (Ischnocera, Philopteridae) were recovered from wings, head and neck regions of red-tailed hawk (Buteo jamaicensis), Swainson's hawk (B. swainsoni), Harris's hawk (Parabuteo unicinctus) and Barn owl (Tyto alba). Low lice infestation level was observed. Nymphs and females of feather mites Kramerella sp. (Pterolichoidea, Kramerellidae) were recovered solely from Barn owl (T. alba); while one Caracara (Caracara cheriway) was infested by the sticktight flea Echidnophaga gallinacea (Siphonaptera, Pulicidae). No clinical signs were observed in any infested bird. Probably the periodic use of organophosphorates was responsible of the low prevalence and lice infestation levels. The diversity of external parasites illustrates the importance of detailed revision of incoming and long-term captive raptors as part of responsible captive management. Five new hosts and geographic records are presented.     

Genotyping of Human Lice Suggests Multiple Emergences of Body Lice from Local Head Louse Populations

Background

Genetic analyses of human lice have shown that the current taxonomic classification of head lice (Pediculus humanus capitis) and body lice (Pediculus humanus humanus) does not reflect their phylogenetic organization. Three phylotypes of head lice A, B and C exist but body lice have been observed only in phylotype A. Head and body lice have different behaviours and only the latter have been involved in outbreaks of infectious diseases including epidemic typhus, trench fever and louse borne recurrent fever. Recent studies suggest that body lice arose several times from head louse populations.

Methods and Findings

By introducing a new genotyping technique, sequencing variable intergenic spacers which were selected from louse genomic sequence, we were able to evaluate the genotypic distribution of 207 human lice. Sequence variation of two intergenic spacers, S2 and S5, discriminated the 207 lice into 148 genotypes and sequence variation of another two intergenic spacers, PM1 and PM2, discriminated 174 lice into 77 genotypes. Concatenation of the four intergenic spacers discriminated a panel of 97 lice into 96 genotypes. These intergenic spacer sequence types were relatively specific geographically, and enabled us to identify two clusters in France, one cluster in Central Africa (where a large body louse outbreak has been observed) and one cluster in Russia. Interestingly, head and body lice were not genetically differentiated.

Conclusions

We propose a hypothesis for the emergence of body lice, and suggest that humans with both low hygiene and head louse infestations provide an opportunity for head louse variants, able to ingest a larger blood meal (a required characteristic of body lice), to colonize clothing. If this hypothesis is ultimately supported, it would help to explain why poor human hygiene often coincides with outbreaks of body lice. Additionally, if head lice act as a reservoir for body lice, and that any social degradation in human populations may allow the formation of new populations of body lice, then head louse populations are potentially a greater threat to humans than previously assumed.     

Immunization of rabbits with a midgut extract of the human body louse Pediculus humanus humanus: the effect of induced resistance on the louse population

Abstract. Resistance to human body lice, Pediculus humanus humanus L, induced by feeding on rabbits immunized with an extract of louse gut was studied. The mortality of lice fed on immunized rabbits was 73%, significantly higher than that of lice fed on control rabbits (52%) ( P < 0.01). The proportion of dead nymphs and female lice with ruptured guts was significantly higher in lice fed on immunized rabbits (P<0.01). The size of the bloodmeal was 35% greater in female lice fed on control rabbits than on immunized rabbits. Lice fed on immunized rabbits laid 40% less eggs than those fed on the controls, they also demonstrated a significant decrease in the number of eggs per female over time ( P < 0.01). 86% of the eggs laid by lice fed on immunized animals hatched, compared with 92% hatching of eggs laid by the lice fed on control animals (P< 0.01). With the exception of the first bloodmeal the percentage of hatched eggs which were laid between any two bloodmeals was significantly smaller (p<0.01) in the lice fed on immunized rabbits than in the control group. The first nymphal stage of lice fed on immunized rabbits took an average of 5.2 days to moult to the second stage, compared with 4 days for those fed on control rabbits.     

Changes in physiological parameters and feeding behaviour of Atlantic salmon Salmo salar infected with sea lice Lepeophtheirus salmonis

Atlantic salmon Salmo salar L. artificially infected with salmon lice Lepeophtheirus salmonis (Kr?yer 1837) recovered from detrimental physiological changes and skin damage induced by preadult lice as the parasites matured. Growth rates of Atlantic salmon remained unaffected by lice infection, but food consumption decreased with increasing feeding and movement of the lice prior to and post-mating, correlating with the appearance of head erosions and detrimental changes in physiological integrity. Food consumption of the fish increased as the lice moulted to the adult stage and gravid female lice settled in a posterior location on the fish, subsequently reducing the impact of infection and allowing recovery of the skin damage. However, the impact of preadults was limited, as the decrease in food consumption of fish at 21 d post-infection had no effect on either the specific growth rate or condition factor of the fish. Furthermore, the intensity of lice infections at each of the sample days was not correlated with food consumption, specific growth rate or any of the haematological or physiological parameters measured, either before or after infection, indicating that lice intensity was independent of social dominance/subordinance. This work has provided the first evidence that infected fish can recover from the detrimental changes caused by lice infection, even when they are still infected with lice. If fish can survive the preadult stage of lice, then the mortal impact of lice infections is greatly reduced.