The head and body lice of humans are genetically distinct (Insecta: Phthiraptera, Pediculidae): evidence from double infestations

Little is known about the population genetics of the louse infestations of humans. We used microsatellite DNA to study 11 double infestations, that is, hosts infested with head lice and body lice simultaneously. We tested for population structure on a host, and for population structure among seven hosts that shared sleeping quarters. We also sought evidence of migration among louse populations. Our results showed that: (i) the head and body lice on these individual hosts were two genetically distinct populations; (ii) each host had their own populations of head and body lice that were genetically distinct to those on other hosts; and (iii) lice had migrated from head to head, and from body to body, but not between heads and bodies. Our results indicate that head and body lice are separate species.     

Phylogenomics and host-switching patterns of Philopteridae (Psocodea: Phthiraptera) feather lice

Philopteridae feather lice are a group of ectoparasitic insects which have intimate relationships with their avian hosts. Feather lice include an enormous number of described species; however, the relationships of major lineages have been clouded by homoplasious characters due to convergent evolution. In this study, a comprehensive phylogenomic analysis of the group is performed which includes 137 feather louse species. Several other analyses are also completed including dating analysis, cophylogenetic reconstructions, and ancestral character estimation to understand the evolution of complex morphological and ecological traits. Phylogenetic results recover high support for the placement of major feather louse lineages, but with lower support for long-branched enigmatic genera found at the base of the tree. The results of dating analyses suggest modern feather lice began to diversify approximately 49 million years ago following the adaptive radiation of their avian hosts. Cost-based cophylogenetic reconstructions recover a high frequency of host switching, while congruence-based methods indicate a significant level of congruence between host and parasite trees. Ancestral state reconstructions favour a generalist ancestor and water bird host at the root. The analyses completed provide insight into the evolution of a diverse group of ectoparasitic insects which infest a wide variety of avian hosts. The results represent the most comprehensive phylogenetic hypothesis of the group to date and provide a framework for future classification of the family into natural groupings.     

Unexpected distribution patterns of Carduiceps feather lice (Phthiraptera: Ischnocera: Philopteridae) on sandpipers (Aves: Charadriiformes: Scolopacidae)

The louse genus Carduiceps Clay & Meinertzhagen, 1939 is widely distributed on sandpipers and stints (Calidrinae). The current taxonomy includes three species on the Calidrinae (Carduiceps meinertzhageni, Carduiceps scalaris, Carduiceps zonarius) and four species on noncalidrine hosts. We estimated a phylogeny of four of the seven species of Carduiceps (the three mentioned above and Carduiceps fulvofasciatus) from 13 of the 29 hosts based on three mitochondrial loci, and evaluated the relative importance of flyway differentiation (same host species has different lice along different flyways) and flyway homogenization (different host species have the same lice along the same flyway). We found no evidence for either process. Instead, the present, morphology‐based, taxonomy of the genus corresponds exactly to the gene‐based phylogeny, with all four included species monophyletic. Carduiceps zonarius is found both to inhabit a wider range of hosts than wing lice of the genus Lunaceps occurring on the same group of birds, and to occur on Calidris sandpipers of all sizes, both of which are unexpected for a body louse. The previously proposed family Esthiopteridae is found to be monophyletic with good support. The concatenated dataset suggests that the pigeon louse genus Columbicola may be closely related to the auk and diver louse genus Craspedonirmus. These two genera share some morphological characters with Carduiceps, but no support was obtained for grouping these three genera together. Based on mitochondrial data alone, the relationships among genera within this proposed family cannot be properly assessed, but some previously suggested relationships within this proposed family are confirmed.     

Louse infestations of tree shrews (Tupaia glis)

Nine adult tree shrews, Tupaia glis, recently imported from West Malaysia were visually examined for ectoparasites while under general anesthesia. Three shrews were infested by the sucking louse, Sathrax durus , and six shrews had louse ova belonging to this species; two shrews had neither lice nor ova. A total of 20 adult female, 10 adult male, and three third instar nymphal lice was collected. Lice were located on the head, flanks, and dorsal body of shrews while ova were recorded mainly from the anterior flanks but also from some adjacent host sites. The tree shrews appeared to tolerate the lice well although louse vector capacity was not assessed. The last date that lice were recorded from shrews was 22 days after colony set-up, and the last date on which seemingly viable ova were recorded was 64 days after set-up showing that the infestations were ultimately lost.     

Host sympatry and body size influence parasite straggling rate in a highly connected multihost,multiparasite system

Parasite lineages commonly diverge when host lineages diverge. However, when large clades of hosts and parasites are analyzed, some cases suggest host switching as another major diversification mechanism. The first step in host switching is the appearance of a parasite on an atypical host, or “straggling.” We analyze the conditions associated with straggling events. We use five species of colonially nesting seabirds from the Galapagos Archipelago and two genera of highly specific ectoparasitic lice to examine host switching. We use both genetic and morphological identification of lice, together with measurements of spatial distribution of hosts in mixed breeding colonies, to test: (1) effects of local host community composition on straggling parasite identity; (2) effects of relative host density within a mixed colony on straggling frequency and parasite species identity; and (3) how straggling rates are influenced by the specifics of louse attachment. Finally, we determine whether there is evidence of breeding in cases where straggling adult lice were found, which may indicate a shift from straggling to the initial stages of host switching. We analyzed more than 5,000 parasite individuals and found that only ~1% of lice could be considered stragglers, with ~5% of 436 host individuals having straggling parasites. We found that the presence of the typical host and recipient host in the same locality influenced straggling. Additionally, parasites most likely to be found on alternate hosts are those that are smaller than the typical parasite of that host, implying that the ability of lice to attach to the host might limit host switching. Given that lice generally follow Harrison's rule, with larger parasites on larger hosts, parasites infecting the larger host species are less likely to successfully colonize smaller host species. Moreover, our study supports the general perception that successful colonization of a novel host is extremely rare, as we found only one nymph of a straggling species, which may indicate successful reproduction.     

云南省吸虱昆虫区系及物种多样性

吸虱是寄生于真兽类哺乳动物体表的专性吸血寄生虫,广布于世界各地。云南省已知吸虱昆虫9科13属44种,分别占中国已知吸虱科、属、种的81.82%,59.09%,45.83%。文章参考大量相关文献,从分类阶元、特有物种、动物地理区划和宿主动物4个方面分析云南省吸虱的物种多样性。云南省吸虱特有种有13种,占云南省已知吸虱种类的29.55%,27种为东洋种,15种为古北和东洋两界兼有种,广布种9种。吸虱在5个地理小区的分布,以横断山中部和横断山南部2个地理小区的吸虱物种多样性较高,其它3个区的物种多样性较低。相对于全国而言,云南省吸虱物种多样性较高,吸虱的宿主动物种类丰富。但蚤、恙螨和革螨等其它体表寄生虫相比较,兽类宿主动物体表吸虱的物种多样性明显低于其它体表寄生虫,1科(属)阶元的吸虱其宿主多为相对一致的1个科(属)动物阶元,反映了吸虱宿主特异性较高的事实,吸虱昆虫与其对应的宿主动物已经形成了比较稳定的"一对一"的寄生关系,这是吸虱昆虫与其宿主动物协同进化的生态学表现。     

Molecular phylogeny and novel host associations of avian chewing lice (Insecta: Phthiraptera) from South Africa

Compared with Europe and the Americas, the ectoparasites of African birds are poorly understood, despite the avian fauna being relatively well known. Notably, previous studies documenting the host associations and genetic diversity of parasitic chewing lice of southern African birds have been limited in geographic and taxonomic scope. Recent field expeditions exploring the avian diversity in South Africa facilitated an opportunity to obtain louse specimens from a taxonomically diverse host assemblage. This study is the first to investigate avian louse host associations and diversity across a large portion of South Africa encompassing several distinct habitat types, while incorporating molecular genetic data (from portions of the mitochondrial COI and nuclear EF‐1α genes) for ectoparasite phylogenetic analyses. From 1105 South African bird individuals and 170 species examined for lice, a total of 105 new louse–host associations were observed. Morphological and genetic examination of lice with these new host associations reveals a maximum of 66 louse species new to science. Results of this study support the observation that examining museum specimens is a useful way to investigate louse diversity and host associations.     

Host behaviour drives parasite genetics at multiple geographic scales: population genetics of the chewing louse,Thomomydoecus minor

Pocket gophers and their symbiotic chewing lice form a host–parasite assemblage known for a high degree of cophylogeny, thought to be driven by life history parameters of both host and parasite that make host switching difficult. However, little work to date has focused on determining whether these life histories actually impact louse populations at the very fine scale of louse infrapopulations (individuals on a single host) at the same or at nearby host localities. We used microsatellite and mtDNA sequence data to make comparisons of chewing‐louse (Thomomydoecus minor) population subdivision over time and over geographic space where there are different potential amounts of host interaction surrounding a zone of contact between two hybridizing pocket‐gopher subspecies. We found that chewing lice had high levels of population isolation consistent with a paucity of horizontal transmission even at the very fine geographic scale of a single alfalfa field. We also found marked genetic discontinuity in louse populations corresponding with host subspecies and little, if any, admixture in the louse genetic groups even though the lice are closely related. The correlation of louse infrapopulation differentiation with host interaction at multiple scales, including across a discontinuity in pocket‐gopher habitat, suggests that host behaviour is the primary driver of parasite genetics. This observation makes sense in light of the life histories of both chewing lice and pocket gophers and provides a powerful explanation for the well‐documented pattern of parallel cladogenesis in pocket gophers and chewing lice.     

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.     

Revision of Dennyus (Collodennyus) lice (Phthiraptera: Menoponidae) from swiftlets, with descriptions of new taxa and a comparison of host–parasite relationships

Lice of the subgenus Dennyus ( Collodennyus ) are host specific, permanent parasites of swiftlets (Aves: Apodidae). As a prelude to a test of the hypothesis that these lice have cospeciated with their hosts, we revise the taxonomy of the subgenus, redescribing the seven previously recognized species, and adding thirteen new species and three new subspecies. All twenty-three of these louse taxa are found on swiftlets (Apodiformes: Apodidae), with four from hosts of the genus Collocalia , eighteen from Aerodramus , and one from Hydrochous . Successful identification is associated in most cases with females; males are only tenuously separable. A complete host–parasite list for the subgenus Collodennyus is provided, as well as a key for the identification of these taxa. Limited morphological variation within the subgenus has prevented ready extraction of discrete characters for cladistic analysis. In the absence of such characters, a cluster analysis of female and male lice is presented. Comparison of a dendrogram for Dennyus ( Collodennyus ) with a molecular phylogeny for the swiftlet hosts suggests that the history of the swiftlet–louse association has been complex, including episodes of host switching and independent speciation by the lice.     

Dispersal ecology versus host specialization as determinants of ectoparasite distribution in brood parasitic indigobirds and their estrildid finch hosts

Brood parasitic birds offer a unique opportunity to examine the ecological and evolutionary determinants of host associations in avian feather lice (Phthiraptera). Brood parasitic behaviour effectively eliminates vertical transfer of lice between parasitic parents and offspring at the nest, while at the same time providing an opportunity for lice associated with the hosts of brood parasites to colonize the brood parasites as well. Thus, the biology of brood parasitism allows a test of the relative roles of host specialization and dispersal ecology in determining the host-parasite associations of birds and lice. If the opportunity for dispersal is the primary determinant of louse distributions, then brood parasites and their hosts should have similar louse faunas. In contrast, if host-specific adaptations limit colonization ability, lice associated with the hosts of brood parasites may be unable to persist on the brood parasites despite having an opportunity for colonization. We surveyed lice on four brood parasitic finch species (genus Vidua), their estrildid finch host species, and a few ploceid finches. While Brueelia lice were found on both parasitic and estrildid finches, a molecular phylogeny showed that lice infesting the two avian groups belong to two distinct clades within Brueelia. Likewise, distinct louse lineages within the amblyceran genus Myrsidea were found on estrildid finches and the parasitic pin-tailed whydah (Vidua macroura), respectively. Although common on estrildid finches, Myrsidea lice were entirely absent from the brood parasitic indigobirds. The distribution and relationships of louse species on brood parasitic finches and their hosts suggest that host-specific adaptations constrain the ability of lice to colonize new hosts, at least those that are distantly related.     

Sucking louse (Hoplopleura erratica: Insecta, Anoplura) exchange between individuals of a wild population of Eastern chipmunks, Tamias striatus, in central Tennessee, U.S.A.

Adult Hoplopleura erratica sucking lice were labelled using a setal clipping technique. Host exchange for lice marked in this manner was monitored through a small population of dye-marked Eastern chipmunks, Tamias striatus, the normal host of this ectoparasite, in a deciduous woodland in central Tennessee, U.S.A. This was accomplished by frequent host live-trapping, anaesthetization and pelage examinations. Louse transfer was most prevalent during the host summer mating period, presumably because of increased chipmunk contacts. At this time, 66·7% of marked adult lice within the host population transferred. Exchanges between opposite sex hosts were more common and adult males donated most lice; no juvenile chipmunks were recorded as louse donors. A greater proportion of male than female lice transferred. Using the exclusive boundary strip method, all exchanging chipmunks had overlapping ranges or shared at least one boundary line.     

GENETIC DIFFERENTIATION AMONG CHEWING LOUSE POPULATIONS (MALLOPHAGA: TRICHODECTIDAE) IN A POCKET GOPHER CONTACT ZONE (RODENTIA: GEOMYIDAE)

Genetic variation among populations of chewing lice (Geomydoecus actuosi) was examined in relation to chromosomal and electrophoretic variation among populations of their hosts (Thomomys bottae) at a contact zone. Louse demes were characterized by low levels of genetic heterozygosity (H? = 0.039) that may result from founder effects during primary infestation of hosts, compounded by seasonal reductions in louse population size. Louse populations sampled from different hosts showed high levels of genetic structuring both within and among host localities. Microgeographic differentiation of louse populations is high (mean F_ST = 0.092) suggesting that properties of this host–parasite system promote differentiation of louse populations living on different individual hosts. Among-population differentiation in lice (F_ST = 0.240) was similar to that measured among host populations (F_ST = 0.236), suggesting a close association between gene flow in pocket gophers and gene flow in their lice.     

Relative importance of demographics, locale, and seasonality underlying louse and flea parasitism of raccoons (Procyon lotor)

A variety of demographic, seasonal, and site-specific variables may influence parasitism, but the relative importance of these variables is generally unclear. We measured the relative ability of host characteristics, season, and site to explain louse (Trichodectes octomaculatus) and flea (Orchopeas howardi) infestation across 10 populations of raccoons (Procyon lotor). Lice are highly dependent on specific hosts and are predicted to display a relatively strong relationship with factors intrinsic to the host, when compared to fleas, which can infest multiple species and survive off-host for weeks without feeding. We developed a priori models that represented explicit hypotheses and contrasted their ability to predict infestation patterns. While the abundance of lice was seasonal, models that included solely host age and sex best predicted prevalence and abundance, in part because males were infested with 3 times the number of lice than were females. Conversely, flea prevalence and abundance, which peaks sharply in the spring, was best predicted by season; factors intrinsic to the host were relatively unimportant for predicting abundance. These, and other, recent findings emphasize the need to simultaneously assess the relative importance of multiple ecological variables between parasite species when attempting to describe general trends and constraints of host-parasite associations.     

Experimental evidence for costs due to chewing lice in the European bee-eater (Merops apiaster)

Animals frequently host organisms on their surface which can be beneficial, have no effect or a negative effect on their host. Ectoparasites, by definition, are those which incur costs to their host, but these costs may vary. Examples of avian ectoparasites are chewing lice which feed exclusively on dead feather or skin material; therefore, costs to their bird hosts are generally considered small. Theoretically, many possible proximate effects exist, like loss of tissue or food, infected bites, transmission of microparasitic diseases or reduced body insulation due to loss of feathers, which may ultimately also have fitness consequences. Here, we experimentally examined a possible negative impact of 2 feather-eating louse species (Meropoecus meropis and Brueelia apiastri) on male and female European bee-eaters (Merops apiaster) by removing or increasing louse loads and comparing their impact to a control group (lice removed and immediately returned) after 1 month. A negative effect of chewing lice was found on body mass and sedimentation rate and to a lesser extent on haematocrit levels. Males and females lost more weight when bearing heavy louse loads, and were more susceptible to infestations as indicated by the higher sedimentation rate. Our results further suggest differences in sex-specific susceptibility.     

Population biology of swift (Apus apus) ectoparasites in relation to host reproductive success

Abstract.

• 1 We censused ectoparasite populations of adult and nestling swifts over the course of the host's breeding season. Nearly all of the birds were infested with chewing lice and two-thirds of the nests were infested with louse flies. Feather mites were observed but not quantified.
• 2 Lice and louse flies both showed aggregated distributions among hosts. Louse eggs, hatched lice and adult louse flies had negative binomial distributions, whereas the aggregated distribution of louse fly pupae was not adequately described by negative binomial or Poisson models.
• 3 Transmission of lice from parents to offspring was documented. A comparison of the age structure of lice on parents and offspring indicated that most transmission was by nymphal lice.
• 4 Host reproductive success and survival appeared to be independent of the number of lice or louse flies. Neither parasite correlated with the number, body mass, or date of fledging of young birds, nor with the overwinter survival of adults. We caution, however, that experimental manipulations of parasite load are required for a definitive test of the impact of ectoparasites on evolutionary fitness components.

     

Rodent louse diversity,phylogeny, and cospeciation in the Manu Biosphere Reserve,Peru

We investigated the diversity, cophylogenetic relationships, and biogeography of hoplopleurid sucking lice (Phthiraptera: Anoplura) parasitizing rodents (Muridae: Sigmodontinae) in the Manu National Park and Biosphere Reserve. Our morphological and molecular studies reveal that 15 distinct louse species parasitize 19 rodent species. Three of these louse species are new to science, and all but two of the host associations were previously unknown. We find that hoplopleurid lice in South America parasitize multiple host species across a large geographic area, and that Peru represents a new geographic locality for almost all the louse species collected in the present study. Phylogenetic analyses of mitochondrial and nuclear data reveal that the louse family Hoplopleuridae and the genera Hoplopleura and Pterophthirus are not monophyletic, and lice do not appear to group by host tribe, collecting locality, or collection elevation. The lack of monophyly for these apparently natural groups (taxonomic, locality, and elevation) indicates that host switching with or without parasite speciation may be prevalent among hoplopleurid lice. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 598–610.     

HOST SPECIALIZATION DIFFERENTIATES CRYPTIC SPECIES OF FEATHER-FEEDING LICE

Parasite species with differentiated host-specific populations provide a natural opportunity to explore factors involved in parasite diversification. Columbicola macrourae is a species of ectoparasitic feather louse currently recognized from 15 species of New World pigeons and doves. Mitochondrial sequences reveal five divergent haplotype clusters within C. macrourae , suggesting cryptic species. Each cluster is relatively host specific, with only one or a few hosts. We conducted a reciprocal transfer experiment with two of these lineages to test whether host use has an adaptive component. Our results demonstrate that the fitness of each lineage is considerably higher on its native host than on the novel host suggesting that one or more selective agents favor host specialization by the different lineages. In addition, we were able to morphologically separate individual lice from the two experimental lineages using discriminant function analysis. Furthermore, differences in the size of these louse lineages match differences in the size of their respective hosts, paralleling the strong correlation between parasite and host body size across the genus Columbicola . Together, these results suggest that selection in this cryptic species complex reflects selection across the whole genus, and that this selection, in part, contributes to the maintenance of host specialization.     

Cophylogeny on a fine scale: Geomydoecus chewing lice and their pocket gopher hosts, Pappogeomys bulleri

Many species of pocket gophers and their ectoparasitic chewing lice have broadly congruent phylogenies, indicating a history of frequent codivergence. For a variety of reasons, phylogenies of codiverging hosts and parasites are expected to be less congruent for more recently diverged taxa. This study is the first of its scale in the pocket gopher and chewing louse system, with its focus entirely on comparisons among populations within a single species of host and 3 chewing louse species in the Geomydoecus bulleri species complex. We examined mitochondrial DNA from a total of 46 specimens of Geomydoecus lice collected from 11 populations of the pocket gopher host, Pappogeomys bulleri. We also examined nuclear DNA from a subset of these chewing lice. Louse phylogenies were compared with a published pocket gopher phylogeny. Contrary to expectations, we observed a statistically significant degree of parallel cladogenesis in these closely related hosts and their parasites. We also observed a higher rate of evolution in chewing louse lineages than in their corresponding pocket gopher hosts. In addition, we found that 1 louse species (Geomydoecus burti) may not be a valid species, that subspecies within G. bulleri are not reciprocally monophyletic, and that morphological and genetic evidence support recognition of a new species of louse, Geomydoecus pricei.