A new genus and new species of sucking louse (Phthiraptera: Anoplura: Enderleinellidae) from the particoloured flying squirrel (Hylopetes alboniger) in Laos, and an updated identification key to enderleinellid genera.

Hylophthirus spinosus, new genus and new species (Phthiraptera: Anoplura: Enderleinellidae), is described from specimens collected from the particoloured flying squirrel, Hylopetes alboniger in Khammouane Province, Laos (Lao Peoples Democratic Republic). Both sexes of the new louse are illustrated with stacked microphotographs, scanning electron micrographs and line drawings. An updated morphological identification key to the six genera (Atopophthirus, Enderleinellus, Hylophthirus, Microphthirus, Phthirunculus and Werneckia) now recognised within the family Enderleinellidae is presented. The new genus is unique within the Enderleinellidae in having massive spines (modified setae) on some abdominal sternites, partially bulbous 5th antennal segments, mesothoracic spiracles borne on protuberances and the morphology of the genitalia of both sexes. Tables showing all genera of sucking lice that include species parasitising sciurids (squirrels, chipmunks, susliks and marmots), and all known species of enderleinellids, with their known host associations and geographical distributions are included and briefly discussed in relation to the new genus and species.

SUCKING LICE (PHTHIRAPTERA: ANOPLURA) PARASITIZING MONGOLIAN RODENTS WITH THE DESCRIPTION OF A NEW SPECIES OF HOPLOPLEURA FROM MOUNTAIN VOLES (ALTICOLA SPP.).

The sucking louse fauna associated with Mongolian mammals is inadequately known. We provide a list of 25 species of sucking lice recorded from Mongolian rodents including previously published records, and new records of specimens collected during an expedition to northwestern Mongolia in 2015. Hoplopleura inagakii Ono and Hasegawa and Polyplax cricetulis Chin are newly recorded from Mongolia and 2 new host associations in Mongolia are recorded for Hoplopleura acanthopus (Burmeister). We describe Hoplopleura altaiensis n. sp., from the Gobi Altai mountain vole, Alticola barakshin Bannikov (type host) with an additional specimen from Alticola strelzowi (Kastchenko) (Strelzow's mountain vole). Both sexes of the new species are illustrated with scanning electron micrographs and line drawings. We note small morphological differences in the shape of the female subgenital plate between specimens prepared for scanning electron microscopy versus those prepared for light microscopy following DNA extraction.

A new species of sucking louse (Psocodea: Phthiraptera: Anoplura: Hoplopleuridae) from the pale field rat, Rattus tunneyi (Rodentia: Muridae), in Australia.

We describe and illustrate a new species of sucking louse, Hoplopleura tunneya new species, from the Australian pale field rat, Rattus tunneyi Thomas (Rodentia: Muridae). Currently, 22 species of the genus Hoplopleura Enderlein, 1904 (Phthiraptera: Anoplura: Hoplopleuridae) are known from Australian endemic rodents. Among the seven new endemic rodent species of the genus Rattus in Australia, R. tunneyi is one of five hosts to Hoplopleura lice. In addition, we give a list of all the species of Hoplopleura known from Australian endemic rodents. Including the introduced species Polyplax spinulosa, the total number of sucking louse species known from Australian endemic rodents is now 24.

Host Associations of Ectoparasites of the Gray Mouse Lemur, Microcebus murinus, in Northwestern Madagascar.

Eight species of ectoparasites were collected during 225 gray mouse lemur, Microcebus murinus (J. F. Miller), captures, in Ankarafantsika National Park, Madagascar, in 2010-2011. The ixodid tick, Haemaphysalis lemuris Hoogstraal, was the most common ectoparasite and was mostly represented by nymphs. Other ectoparasites recorded include the polyplacid sucking louse, Lemurpediculus madagascariensis Durden, Kessler, Radespiel, Zimmermann, Hasiniaina, and Zohdy; the ixodid tick, Haemaphysalis simplex Neumann; an undescribed laelapid mite in the genus Aetholaelaps; another laelapid belonging to the genus Androlaelaps; the chigger mite Schoutedenichia microcebi Stekolnikov; an undescribed species of atopomelid mite in the genus Listrophoroides; and an undescribed species of psoroptid mite in the genus Cheirogalalges. Except for the 2 species of ticks and 1 species of chigger, these ectoparasites may be host-specific to M. murinus. Total tick (H. lemuris and H. simplex) infestation was significantly greater in August than October, whereas louse (L. madagascariensis) infestation was significantly greater in October. There was no significant difference in tick infestations between male and female lemurs, but male lemurs had significantly more lice than female lemurs. Reproductive status was not a significant predictor of tick infestation in males and females.

Two New Species of Sucking Lice (Psocodea: Phthiraptera: Hoplopleuridae) from Chestnut Mice, Pseudomys gracilicaudatus and Pseudomys nanus (Rodentia: Muridae), in Australia.

We describe two new species of sucking lice in the genus Hoplopleura Enderlein, 1904 (Psocodea: Phthiraptera: Hoplopleuridae) from Australia: Hoplopleura gracilicaudatusa n. sp. from the eastern chestnut mouse Pseudomys gracilicaudatus (Gould) (Rodentia: Muridae), and Hoplopleura nanusa n. sp. from the western chestnut mouse Pseudomys nanus (Gould) (Rodentia: Muridae). Pseudomys Gray is the most speciose genus of rodents endemic to Australia with 24 species; however, only two Pseudomys species have been reported previously to be hosts of sucking lice. The description of the new species in the present study doubles the number of sucking louse species known to parasitize Pseudomys mice and increases the total number of sucking louse species known from endemic Australian rodents from 21 to 23. Pseudomys gracilicaudatus and P. nanus are closely related murines that diverged ~1 MYA with distinct and widely separated extant geographic distributions. The two new Hoplopleura species described in the present study share some morphological characters and likely co-evolved and co-speciated with their chestnut mouse hosts.

Eight New Species of Sucking Lice (Psocodea: Phthiraptera) From Endemic Murine Rodents in Australia and an Updated Identification Key.

Based on a comprehensive study of museum specimens, eight new species of sucking lice of the genus Hoplopleura Enderlein, 1904 (Psocodea: Phthiraptera: Hoplopleuridae), are described from six genera of Australian Old Endemic rodents: Conilurus Ogilby, 1838 (Rodentia: Muridae), Leggadina Thomas, 1910 (Rodentia: Muridae), Leporillus Thomas, 1906 (Rodentia: Muridae), Mesembriomys Palmer, 1906 (Rodentia: Muridae), Pogonomys Milne-Edwards, 1877 (Rodentia: Muridae), and Xeromys Thomas, 1889 (Rodentia: Muridae). The description of these new species increases the number of sucking louse species from endemic Australian rodents from 13 to 21 and extends the records of sucking lice to all of the 14 genera of endemic rodents in Australia. Our results show that sucking lice are much more diverse among rodents in Australia than previously known. Furthermore, the Australian Hoplopleura species are host specific-each Hoplopleura species, including the eight new species described in the present study, parasitizes only a single host species, except Hoplopleura irritans Kuhn and Ludwig, 1967 (Psocodea: Phthiraptera: Hoplopleuridae) and Hoplopleura melomydis Weaver, 2017 (Psocodea: Phthiraptera: Hoplopleuridae), each of which is found on two host species. An updated dichotomous key for identifying Australian Hoplopleura species is included.

Two New Species of Sucking Lice (Phthiraptera: Anoplura: Hoplopleuridae and Polyplacidae) from Grant's Rock Mouse, Micaelamys granti, in South Africa.

Two new species of sucking lice (Phthiraptera: Anoplura), Hoplopleura granti n. sp. (Hoplopleuridae) and Polyplax megacephalus n. sp. (Polyplacidae), are described from Grant's rock mouse, Micaelamys granti (Wroughton), from Northern Cape Province, South Africa. Adults of both sexes are illustrated via line drawings and differential interference contrast microscopy images, and are compared with previously described related species that parasitize related hosts: Hoplopleura patersoni Johnson from Aethomys chrysophilus (de Winton) (red veld rat), Aethomys kaiseri (Noack) (Kaiser's aethomys), and Micaelamys namaquensis (A. Smith) (Namaqua rock mouse); Hoplopleura aethomydis Kleynhans from M. namaquensis; Polyplax praomydis Bedford from A. chrysophilus and M. namaquensis; and Polyplax solivaga Johnson from A. chrysophilus. It is not known if these new species of lice are vectors of any pathogens.

A New Species of Sucking Louse from the Mandrill from Gabon with a Review of Host Associations and Geographical Distributions, and Identification Keys to Members of the Genus pedicinus (Phthiraptera: Anoplura: Pedicinidae).

Members of the sucking louse genus Pedicinus are ectoparasites of cercopithecid primates in Africa, Asia, and Gibraltar. Pedicinus gabonensis n. sp. is described on the basis of adult male and female specimens collected from the mandrill (Mandrillus sphinx) in Gabon. The new species is compared morphologically with other members of the genus Pedicinus, and a nuclear elongation factor 1 alpha gene sequence is provided. Host associations and geographical distributions of the 18 previously recognized species of the genus and of P. gabonensis n. sp. are reviewed. Updated identification keys are provided for males and females of all known valid species of Pedicinus.

Rapid host expansion of an introduced parasite, the spiny rat louse Polyplax spinulosa (Psocodea: Phthiraptera: Polyplacidae), among endemic rodents in Australia.

BACKGROUND: Historical European exploration and colonization resulted in the introduction of four species of rodents to the Australian continent from Eurasia: the brown rat, Rattus norvegicus, the black rat, R. rattus, the Pacific rat, R. exulans, and the house mouse, Mus musculus. The spread of these rodents created opportunities for their co-introduced sucking lice to parasitize and adapt to endemic rodents in Australia. METHODS: We collected sucking lice from rodent specimens in seven museums across Australia. We identified the spiny rat louse, Polyplax spinulosa, based on morphology. We sequenced the mitochondrial cox1 and rrnL genes of P. spinulosa specimens and constructed a phylogenetic tree with rrnL sequences. RESULTS: We examined 989 rodent specimens of 54 species and collected 2111 adult sucking lice and 1064 nymphal sucking lice. We found that P. spinulosa had nearly doubled its host range by parasitizing at least six endemic rodent species in Australia. The other two introduced lice, P. serrata and Hoplopleura pacifica, however, have apparently failed to expand to any endemic rodents in Australia. Our analysis of mitochondrial rrnL gene sequences divided P. spinulosa into two genotypes (European vs Southeast Asian), which differ by 7.5%; both genotypes were introduced into Australia and then expanded their host ranges to include endemic rodents. CONCLUSIONS: The earliest record of a European ship landing in Australia was in 1606, followed by British settlement in 1788. The expansion of P. spinulosa to at least six endemic rodent species in Australia has therefore occurred in the time frame of 200 to 400 years, which is extremely rapid relative to its host expansion to eight native rat species in Eurasia in ~ 16 million years since it diverged from P. serrata. The host expansion of P. spinulosa is remarkable for a blood-sucking louse and is in stark contrast to the absence of host expansion by P. serrata and H. pacifica. Comparison among these three introduced sucking lice indicated that both louse-specific factors and host-specific factors can contribute to the success or failure of host expansion.

A New Species of Sucking Louse from the Long-tailed Ground Squirrel, Urocitellus undulatus, from Mongolia, with a Key to Species, and a Review of Host Associations and Geographical Distributions of Members of the Genus Linognathoides (Psocodea: Anoplura: Polyplacidae).

Members of the genus Linognathoides are ectoparasites of ground squirrels and marmots (Rodentia: Sciuridae) in the Nearctic, Palearctic, and Afrotropical regions. Linognathoides urocitelli n. sp. is described based on adult male and female and third-instar nymphal specimens collected from the long-tailed ground squirrel (Urocitellus undulatus) in Mongolia. The new species is compared morphologically to other members of the genus Linognathoides. Additionally, DNA sequences of a 610-base pair (bp) fragment of the nuclear 18S rDNA gene, a 452-bp fragment of the mitochondrial 12S rrnS gene, and a 363-bp fragment of the mitochondrial 16S rrnL gene are provided. Host associations and geographical distributions of the 11 previously recognized species of the genus, and of Linognathoides urocitelli n. sp., are reviewed. A dichotomous identification key to adults of all known species in the genus Linognathoides is provided.

A new species of sucking louse Hoplopleura villosissima n. sp. (Psocodea: Phthiraptera: Hoplopleuridae) and a new host record of the spiny rat louse Polyplax spinulosa Burmeister, 1839 (Psocodea: Phthiraptera: Polyplacidae) from the long-haired rat Rattus villosissimus Waite (Rodentia: Muridae) in Australia.

BACKGROUND: The sucking louse fauna of endemic Australian rodents has been under-studied for decades. Sixty-five species of native rodents have been recorded in Australia. However, only 11 species of lice have been reported from 11 species of endemic Australian rodents. RESULTS: We describe a new species of sucking louse, Hoplopleura villosissima Wang (Psocodea: Phthiraptera: Hoplopleuridae), and report a new host record of the spiny rat louse, Polyplax spinulosa Burmeister, 1839 (Psocodea: Phthiraptera: Polyplacidae), from the long-haired rat, Rattus villosissimus Waite (Rodentia: Muridae), which is endemic to Australia. CONCLUSIONS: This study is the first record of sucking louse from R. villosissimus and the first record of a species of Polyplax Enderlein, 1904 from an endemic Australian rodent. This study brings the total number of sucking louse species in endemic Australian rodents from 11 to 13. Previously, only the introduced brown rat, Rattus norvegicus Berkenhout and the black rat, Rattus rattus Linnaeus were recorded as the hosts of P. spinulosa in Australia. Because R. villosissimus overlaps with R. rattus in distribution but not with R. norvegicus, we propose that P. spinulosa transferred to R. villosissimus from R. rattus.

A New Species of Sucking Louse (Phthiraptera: Anoplura: Polyplacidae) From the Gray Mouse Lemur, Microcebus murinus (Primates: Cheirogaleidae), in Madagascar.

Lemurpediculus madagascariensis sp. nov. (Phthiraptera: Anoplura: Polyplacidae) is described from the Gray Mouse lemur, Microcebus murinus (J. F. Miller) (Primates: Cheirogaleidae), from Ankarafantsika National Park, Madagascar. Lemurs were trapped using Sherman Live Traps and visually inspected for lice, which were preserved in 90% ethanol. Adults of both sexes and the third-instar nymph of the new species are illustrated and distinguished from the four previously known species of Lemurpediculus: L. verruculosus (Ward); L. petterorum Paulian; L. claytoni Durden, Blanco, and Seabolt; and L. robbinsi Durden, Blanco, and Seabolt. It is not known if the new species of louse is a vector of any pathogens or parasites.

Two New Species of Sucking Lice (Phthiraptera: Anoplura: Polyplacidae) From Endangered, Hibernating Lemurs (Primates: Cheirogaleidae).

Lemurpediculus robbinsi sp. nov. is described from Crossley's dwarf lemur, Cheirogaleus crossleyi A. Grandidier, and Lemurpediculus claytoni sp. nov. is described from Sibree's dwarf lemur, Cheirogaleus sibreei Forsyth Major, from Madagascar. Both sexes of each new louse species are illustrated and distinguished from the two previously known species of Lemurpediculus: L. verruculosus (Ward) and L. petterorum Paulian. With the addition of two new species to the genus, an amended description of Lemurpediculus is provided. The two hosts of the new louse species are morphologically similar, endangered, obligately hibernating lemurs. These two species of lemurs are sometimes sympatric in rainforests in eastern Madagascar. Despite the morphological similarity of the two host species, their lice are morphologically distinct and are easiest to identify based on the shape of the subgenital plate of the female and the shape of the genitalia in the male. Both new species of lice should be considered to be endangered because their hosts are endangered. It is not known if either of the new species of lice are vectors of pathogens or parasites to their hosts.

Host-Parasite Associations in Small Mammal Communities in Semiarid Savanna Ecosystems of East Africa.

Despite the established importance of rodents as reservoirs of vector-borne zoonoses in East Africa, there is relatively limited information regarding the infestation parameters and host associations of ectoparasites that vector many such pathogens among small mammals in this region. Between 2009 and 2013, small mammals were live-trapped in the semiarid savanna of Kenya. A subset of these individual hosts, including 20 distinct host taxa, was examined for ectoparasites, which were identified to species. Species of fleas, ticks, mites, and sucking lice were recorded. Based on these data, we calculated host-specific infestation parameters, documented host preferences among ectoparasites, conducted a rarefaction analysis and extrapolation to determine if ectoparasites were adequately sampled, and assessed nestedness for fleas to understand how pathogens might spread in this system. We found that the flea community structure was significantly nested. Understanding the ectoparasite network structure may have significant human relevance, as at least seven of the ectoparasite species collected are known vectors of pathogens of medical importance in the region, including Yersinia pestis, Rickettsia spp., and Theileria parva, the causative agents of plague, spotted fevers and other rickettsial illnesses in humans, and theileriosis, respectively.

A new species of sucking louse (Phthiraptera: Anoplura: Linognathidae) from Günther's Dikdik (Madoqua guentheri) in Kenya.

Linognathus samburi n. sp. is described from adult male and female specimens collected from a juvenile female Günther's dikdik (Madoqua guentheri) live-trapped near Olturot Village, Samburu district (Rift Valley Province) in northern Kenya. The new species is distinguished from other species of Linognathus including Linognathus geigyi and Linognathus damarensis, both of which parasitize Kirk's dikdik (Madoqua kirkii). A dichotomous key to the species of Linognathus that are known to parasitize dikdiks is included.

Mapping the social network: tracking lice in a wild primate (Microcebus rufus) population to infer social contacts and vector potential.

BACKGROUND: Studies of host-parasite interactions have the potential to provide insights into the ecology of both organisms involved. We monitored the movement of sucking lice (Lemurpediculus verruculosus), parasites that require direct host-host contact to be transferred, in their host population of wild mouse lemurs (Microcebus rufus). These lemurs live in the rainforests of Madagascar, are small (40 g), arboreal, nocturnal, solitary foraging primates for which data on population-wide interactions are difficult to obtain. We developed a simple, cost effective method exploiting the intimate relationship between louse and lemur, whereby individual lice were marked, without removal from their host, with an individualized code, and tracked throughout the lemur population. We then tested the hypotheses that 1) the frequency of louse transfers, and thus interactions, would decrease with increasing distance between paired individual lemurs; 2) due to host polygynandry, social interactions and hence louse transfers would increase during the onset of the breeding season; and 3) individual mouse lemurs would vary in their contributions to the spread of lice. RESULTS: We show that louse transfers involved 43.75% of the studied lemur population, exclusively males. Louse transfers peaked during the breeding season, perhaps due to increased social interactions between lemurs. Although trap-based individual lemur ranging patterns are restricted, louse transfer rate does not correlate with the distance between lemur trapping locales, indicating wider host ranging behavior and a greater risk of rapid population-wide pathogen transmission than predicted by standard trapping data alone. Furthermore, relatively few lemur individuals contributed disproportionately to the rapid spread of lice throughout the population. CONCLUSIONS: Using a simple method, we were able to visualize exchanges of lice in a population of cryptic wild primates. This method not only provided insight into the previously unseen parasite movement between lemurs, but also allowed us to infer social interactions between them. As lice are known pathogen vectors, our method also allowed us to identify the lemurs most likely to facilitate louse-mediated epidemics. Our approach demonstrates the potential to uncover otherwise inaccessible parasite-host, and host social interaction data in any trappable species parasitized by sucking lice.

Lice and ticks of the eastern rufous mouse lemur, Microcebus rufus, with descriptions of the male and third instar nymph of Lemurpediculus verruculosus (Phthiraptera: Anoplura).

Sucking lice and ticks were collected from live-trapped eastern rufous mouse lemurs, Microcebus rufus Geoffroy, in and around the periphery of Ranomafana National Park, southeastern Madagascar, from 2007 to 2009. Samples of 53 sucking lice (Insecta: Phthiraptera: Anoplura) and 28 hard ticks (Acari: Ixodidae) were collected from 36 lemur captures representing 26 different host individuals. All of the lice were Lemurpediculus verruculosus (Ward) (6 males, 46 females, 1 third instar nymph). Only the holotype female was known previously for this louse and the host was stated to be a "mouse lemur." Therefore, we describe the male and third instar nymph of L. verruculosus and confirm M. rufus as a host (possibly the only host) of this louse. All of the ticks were nymphs and consisted of 16 Haemaphysalis lemuris Hoogstraal, 11 Haemaphysalis sp., and 1 Ixodes sp. The last 2 ticks listed did not morphologically match any of the Madagascar Haemaphysalis or Ixodes ticks for which nymphal stages have been described.

Evolutionary history of mammalian sucking lice (Phthiraptera: Anoplura).

BACKGROUND: Sucking lice (Phthiraptera: Anoplura) are obligate, permanent ectoparasites of eutherian mammals, parasitizing members of 12 of the 29 recognized mammalian orders and approximately 20% of all mammalian species. These host specific, blood-sucking insects are morphologically adapted for life on mammals: they are wingless, dorso-ventrally flattened, possess tibio-tarsal claws for clinging to host hair, and have piercing mouthparts for feeding. Although there are more than 540 described species of Anoplura and despite the potential economical and medical implications of sucking louse infestations, this study represents the first attempt to examine higher-level anopluran relationships using molecular data. In this study, we use molecular data to reconstruct the evolutionary history of 65 sucking louse taxa with phylogenetic analyses and compare the results to findings based on morphological data. We also estimate divergence times among anopluran taxa and compare our results to host (mammal) relationships. RESULTS: This study represents the first phylogenetic hypothesis of sucking louse relationships using molecular data and we find significant conflict between phylogenies constructed using molecular and morphological data. We also find that multiple families and genera of sucking lice are not monophyletic and that extensive taxonomic revision will be necessary for this group. Based on our divergence dating analyses, sucking lice diversified in the late Cretaceous, approximately 77 Ma, and soon after the Cretaceous-Paleogene boundary (ca. 65 Ma) these lice proliferated rapidly to parasitize multiple mammalian orders and families. CONCLUSIONS: The diversification time of sucking lice approximately 77 Ma is in agreement with mammalian evolutionary history: all modern mammal orders are hypothesized to have diverged by 75 Ma thus providing suitable habitat for the colonization and radiation of sucking lice. Despite the concordant timing of diversification events early in the association between anoplurans and mammals, there is substantial conflict between the host and parasite phylogenies. This conflict is likely the result of a complex history of host switching and extinction events that occurred throughout the evolutionary association between sucking lice and their mammalian hosts. It is unlikely that there are any ectoparasite groups (including lice) that tracked the early and rapid radiation of eutherian mammals.

Haemodipsus brachylagi n. sp. (Phthiraptera: Anoplura: Polyplacidae), a new sucking louse from the pygmy rabbit in Nevada.

The male and female of Haemodipsis brachylagi n. sp. (Phthiraptera: Anoplura) are described from specimens collected from a pygmy rabbit, Brachylagus idahoensis (Merriam) (Lagomorpha: Leporidae), from Nevada. Morphological features that differentiate the new species from other known species of Haemodipsus are elucidated, and an identification key to both sexes of the 3 species now known from this genus in North America is included. Geographical distributions of the other 4 species of Haemodipsus known from other parts of the world are highlighted.

Epifaunistic arthropod parasites of the four-striped mouse, Rhabdomys pumilio, in the Western Cape Province, South Africa.

Flea, lice, mite, and tick species associated with 510 Rhabdomys pumilio were collected at 9 localities in the Western Cape Province, South Africa. The aims of the study were first to quantify the species richness, prevalence, and relative mean intensity of infestation of epifaunistic arthropod species associated with R. pumilio, and second to determine temporal variations in the mean abundance of the parasitic arthropods. Each mouse was examined under a stereoscopic microscope and its parasites were removed, identified, and quantified. The epifaunal population was made up of more than 25,000 individuals and included 8 flea, 1 sucking louse, 11 mite, and 13 ixodid tick species. Female-biased sex ratios were noted for 9 (30%) of the ectoparasite species. Three undescribed mite and 1 undescribed tick species were recovered, and new locality records for 2 flea, the louse, and 2 mite species were documented. A phoretic host association between a nonparasitic mite species, Psylloglyphus uilenbergi kivuensis, and 3 flea species, Chiastopsylla rossi, Hypsophthalmus temporis, and Listropsylla agrippinae, was recorded. The mean abundance of the parasitic mite and insect species were higher during the cold wet season, whereas ticks were more numerous during the warm dry months. The large number of ectoparasite species on R. pumilio, a locally abundant and regionally widespread species, is of medical and veterinary importance particularly in relation to the transmission of pathogens such as Anaplasma marginale, Babesia caballi, and Babesia canis to domestic animals; Rickettsia conori; Yersinia pestis; and the viral disease Crimean-Congo hemorrhagic fever to humans.