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Gustafsson, Daniel R., Bush, Sarah E. (2021): Descriptions of seven new species of Brueelia Kéler 1936 (Phthiraptera: Ischnocera: Philopteridae) from North American sparrows (Aves: Passeriformes: Passerellidae), and review of host use by Brueelia vulgata. Journal of Natural History 54 (33-34): 2071-2112, DOI: 10.1080/00222933.2020.1836280, URL: http://dx.doi.org/10.1080/00222933.2020.1836280

Adaptive radiation is an important mechanism of organismal diversification, and can be triggered by new ecological opportunities. Although poorly studied in this regard, parasites present an ideal system to study adaptive radiations because of their close associations with host species. Both experimental and comparative studies suggest that the ectoparasitic wing lice of pigeons and doves have undergone an adaptive radiation, resulting in differences in both body size and overall coloration. Here we show that long-distance dispersal by dove hosts was important for parasite diversification, providing new ecological opportunities for parasites to speciate by host-switching. We further show that these new opportunities for host-switching decreased over time, and cospeciation became the more dominant mode of parasite speciation. Taken together, our results suggest that host dispersal, followed by host-switching, provided novel ecological opportunities that facilitated adaptive radiation by parasites.

Six new species of chewing lice in the Resartor-group (Brueelia-complex) are described and illustrated. They are: Aratricerca cerata n. sp. ex Zosterops capensis Sundevall, 1850; Aratricerca macki n. sp. ex Melidectes princeps Mayr & Gilliard, 1951 and Ptiloprora perstriata perstriata (de Vis, 1898); Aratricerca madagascariensis n. sp. ex Randia pseudozosterops Delacour & Berlioz, 1931; Turdinirmoides janigai n. sp. ex Prunella collaris nipalensis (Blyth, 1843) and P. collaris fennelli Deignan, 1964; Turdinirmoides rozsai n. sp. ex Carpodacus subhimachala (Hodgson, 1836); and Timalinirmus curvus n. sp. ex Yuhina castaniceps plumbeiceps (Godwin-Austen, 1877). A key to the species of Aratricerca, Turdinirmoides and Timalinirmus is provided.

SARS coronavirus 2 (SARS-CoV-2) is a novel viral pathogen that causes a clinical disease called coronavirus disease 2019 (COVID-19). Although most COVID-19 cases are asymptomatic or involve mild upper respiratory tract symptoms, a significant number of patients develop severe or critical disease. Patients with severe COVID-19 commonly present with viral pneumonia that may progress to life-threatening acute respiratory distress syndrome (ARDS). Patients with COVID-19 are also predisposed to venous and arterial thromboses that are associated with a poorer prognosis. The present study identified the emergence of a low-density inflammatory neutrophil (LDN) population expressing intermediate levels of CD16 (CD16Int) in patients with COVID-19. These cells demonstrated proinflammatory gene signatures, activated platelets, spontaneously formed neutrophil extracellular traps, and enhanced phagocytic capacity and cytokine production. Strikingly, CD16Int neutrophils were also the major immune cells within the bronchoalveolar lavage fluid, exhibiting increased CXCR3 but loss of CD44 and CD38 expression. The percentage of circulating CD16Int LDNs was associated with D-dimer, ferritin, and systemic IL-6 and TNF-α levels and changed over time with altered disease status. Our data suggest that the CD16Int LDN subset contributes to COVID-19–associated coagulopathy, systemic inflammation, and ARDS. The frequency of that LDN subset in the circulation could serve as an adjunct clinical marker to monitor disease status and progression.

A recent paper in this journal concerning parasites of rock pigeons (Columba livia) published by Ali and colleagues exemplifies a growing trend of misidentified parasites in the literature, despite increased online resources that should help facilitate accurate identification. In the Ali et al. paper, a pigeon louse in the genus Columbicola (Phthiraptera: Ischnocera) is misidentified as Menopon gallinae, which is a parasite of chickens (Gallus gallus) and their relatives; moreover, this louse is from an entirely different suborder of lice (Phthiraptera: Amblycera). Another louse is misidentified as Goniodes dissimilis, another parasite of chickens and junglefowl. In addition, photographs of cestodes from pigeons in the same paper are not sufficient to confirm identification. Misidentifications are fueled, in part, by increasing pressure to publish coupled with a decrease in taxonomic expertise. We consider the downstream consequences of misidentification and suggest guidelines for authors, reviewers, and editors that could help to improve the reliability of specimen-based research.

The pigeon louse Columbicola columbae is a longstanding and important model for studies of ectoparasitism and host-parasite coevolution. However, a deeper understanding of its evolution and capacity for rapid adaptation is limited by a lack of genomic resources. Here, we present a high-quality draft assembly of the C. columbae genome, produced using a combination of Oxford Nanopore, Illumina, and Hi-C technologies. The final assembly is 208 Mb in length, with 12 chromosome-size scaffolds representing 98.1% of the assembly. For gene model prediction, we used a novel clustering method (wavy_choose) for Oxford Nanopore RNA-seq reads to feed into the MAKER annotation pipeline. High recovery of conserved single-copy orthologs (BUSCOs) suggests that our assembly and annotation are both highly complete and highly accurate. Consistent with the results of the only other assembled louse genome, Pediculus humanus, we find that C. columbae has a relatively low density of repetitive elements, the majority of which are DNA transposons. Also similar to P. humanus, we find a reduced number of genes encoding opsins, G protein-coupled receptors, odorant receptors, insulin signaling pathway components, and detoxification proteins in the C. columbae genome, relative to other insects. We propose that such losses might characterize the genomes of obligate, permanent ectoparasites with predictable habitats, limited foraging complexity, and simple dietary regimes. The sequencing and analysis for this genome were relatively low cost, and took advantage of a new clustering technique for Oxford Nanopore RNAseq reads that will be useful to future genome projects.

The species of the chewing louse genus Guimaraesiella Eichler, 1949 parasitic on drongos (Dicruridae) are reviewed and placed in the new subgenus Dicrurobates, which is described herein together with eight new species, including one species from non-dicrurid hosts. The new species are: Guimaraesiella (Dicrurobates) carbonivora n. sp. from Dicrurus bracteatus carbonarius Bonaparte, 1850; Guimaraesiella (Dicrurobates) latitemporalis n. sp. from Dicrurus hottentottus brevirostris (Cabanis, 1851) and Dicrurus hottentottus ssp. (Linnaeus, 1766); Guimaraesiella (Dicrurobates) lurida n. sp. from Dicrurus leucophaeus Vieillot, 1817; Guimaraesiella (Dicrurobates) luzonica n. sp. from Dicrurus balicassius (Linnaeus, 1766); Guimaraesiella (Dicrurobates) nana n. sp. from Dicrurus hottentottus samarensis Vaurie, 1947; Guimaraesiella (Dicrurobates) regis n. sp. from Dicrurus annectans (Hodgson, 1836), Dicrurus paradiseus paradiseus (Linnaeus, 1766) and Dicrurus paradiseus rangoonensis (Gould, 13836); Guimaraesiella (Dicrurobates) transvaalensis n. sp. from Dicrurus adsimilis apivorus Clancey, 1976; and Guimaraesiella (Dicrurobates) campanula n. sp. from Oriolus larvatus rolleti Salvadori, 1864 and Prionops plumatus poliocephalus (Stanley, 1814). Also, Guimaraesiella (Dicrurobates) sexmaculata (Piaget, 1880) and Guimaraesiella (Dicrurobates) dicruri (Ansari, 1955) are redescribed and illustrated. A key to identify adults of all 10 species included in the subgenus is provided.

The pigeon louse Columbicola columbae is a longstanding and important model for studies of ectoparasitism and host-parasite coevolution. However, a deeper understanding of its evolution and capacity for rapid adaptation is limited by a lack of genomic resources. Here, we present a high-quality draft assembly of the C. columbae genome, produced using a combination of Oxford Nanopore, Illumina, and Hi-C technologies. The final assembly is 208 Mb in length, with 12 chromosome-size scaffolds representing 98.1% of the assembly. For gene model prediction, we used a novel clustering method (wavy_choose) for Oxford Nanopore RNA-seq reads to feed into the MAKER annotation pipeline. High recovery of conserved single-copy orthologs (BUSCOs) suggests that our assembly and annotation are both highly complete and highly accurate. Consistent with the results of the only other assembled louse genome, Pediculus humanus, we find that C. columbae has a relatively low density of repetitive elements, the majority of which are DNA transposons. Also similar to P. humanus, we find a reduced number of genes encoding opsins, G protein-coupled receptors, odorant receptors, insulin signaling pathway components, and detoxification proteins in the C. columbae genome, relative to other insects. We propose that such losses might characterize the genomes of obligate, permanent ectoparasites with predictable habitats, limited foraging complexity, and simple dietary regimes. The sequencing and analysis for this genome were relatively low-cost, and took advantage of a new clustering technique for Oxford Nanopore RNAseq reads that will be useful to future genome projects.

Grooming by birds is thought to serve essential anti-parasite functions. While preening has been well studied, little is known about the function of scratching in birds. We conducted a series of experiments to determine the effectiveness of scratching for controlling feather lice (Columbicola columbae) on Rock Pigeons (Columba livia). First, we used a hobbling technique to impair scratching. After 6 mo, hobbled birds had significantly more lice than controls that could scratch. In addition, lice on hobbled birds were concentrated on the birds’ heads and necks (i.e. the regions that birds scratch). Secondly, we tested the role the claw plays in scratching by declawing nestlings. Once mature, declawed pigeons had significantly more lice than control birds with claws. Moreover, lice on declawed birds were concentrated on the head and neck. Next, we tested whether the flange found on the middle claw of many bird species enhances scratching. We experimentally manipulated the flange; however, the number and location of lice on birds without flanges was not significantly different than that on control birds with intact flanges. Finally, we tested whether scratching removes parasites directly or indirectly by “flushing” them onto body regions where they can be preened. When we impaired scratching (with hobbles) and preening (with “bits”) we found that scratching no longer reduced the number of lice on birds. Our results indicated that scratching and preening work synergistically; scratching reduces parasite load by flushing lice onto regions of the body where they can be eliminated by preening.

Allopreening occurs when 1 bird preens another bird. The behavior is normally directed at the head and neck of the recipient, i.e., regions that the bird cannot self-preen. Studies of penguins, pigeons, and other groups of birds suggest that allopreening plays a role in the control of ectoparasites, such as ticks and feather lice. However, it is not known whether allopreening increases in response to increases in parasite load, or whether it is a programmed response that occurs independently of parasite load. We conducted a laboratory experiment using wild-caught rock pigeons (

Five new species of chewing lice in the genus

Five new species of

Two large taxonomic revisions of chewing lice belonging to the Brueelia-complex were published independently in 2017: Gustafsson & Bush (August 2017) and Mey (September 2017). However, Mey (2017) was incorrectly dated “Dezember 2016” on the title page. These two publications described many of the same taxonomic units under different names and therefore, the names in Gustafsson and Bush (2017) have priority over the synonyms in Mey (2017). Here we clarify some of the resulting taxonomic confusion. Firstly, we confirm the availability of the genera Guimaraesiella Eichler, 1949 and Acronirmus Eichler, 1953, as well as the status of Nitzschinirmus Mey & Barker, 2014 as a junior synonym of Guimaraesiella. Nine genera were described and simultaneously placed as juniors synonyms by Mey (2017: 182). We agree with his synonymy in seven of them: Australnirmus Mey, 2017 under Saepocephalum Gustafsson & Bush, 2017; Couanirmus Mey, 2017 under Couala Gustafsson & Bush, 2017; Estrildinirmus Mey, 2017 under Mirandofures Gustafsson & Bush, 2017; Harpactiacus Mey, 2017 under Harpactrox Gustafsson & Bush, 2017; Leiothrichinirmus Mey, 2017 under Resartor Gustafsson & Bush, 2017; Manucodiacus Mey, 2017 under Manucodicola Gustafsson & Bush, 2017, and Protonirmus Mey, 2017 under Ceratocista Gustafsson & Bush, 2017. Furthermore, Mey (2017) described and placed Pomatostomiacus as an absolute junior synonym of Sychraella Gustafsson & Bush, 2017; here, we argue that Pomatostomiacus is actually a junior synonym of Anarchonirmus Gustafsson & Bush, 2017. Also, Mey (2017) described and placed Timalinirmus as a probable junior synonym of Turdinirmoides Gustafsson & Bush, 2017; here we argue that Timalinirmus is a valid genus.We place ten more genera from Mey (2017) as junior synonyms, as follows: Callaenirmus Mey, 2017 and Philemoniellus Mey, 2017 under Guimaraesiella Eichler, 1949; Carpodaciella Mey, 2017 under Turdinirmoides Gustafsson & Bush, 2017; Cinclosomatiellum Mey, 2017 under Maculinirmus Złotorzycka, 1964; Koanirmus Mey, 2017 and Tesonirmus Mey, 2017 under Couala Gustafsson & Bush, 2017; Garrulaxeus Mey, 2017 under Priceiella (Camurnirmus) Gustafsson & Bush, 2017; Lycocoranirmus Mey, 2017 under Corvonirmus Eichler, 1944; Neosittiella Mey, 2017 and Plesionirmus Mey, 2017 under Brueelia Kéler, 1936. We accept Melinirmus Mey, 2017 as valid, and Mohoaticus Mey, 2017 as a valid subgenus of Guimaraesiella Eichler, 1949. Also, we provisionally accept Ptilononirmus Mey, 2017 as valid but, until a proper redescription determines its true status, we categorize Ptilononirmus as genus inquirenda. We accept most species described by Mey (2017) as valid, except for two which we place as junior synonyms: Callaenirmus kokakophilus Mey, 2017 under Brueelia callaeincola Valim & Palma, 2015, and Mohoaticus pteroacariphagus Mey, 2017 under Guimaraesiella (Mohoaticus) diaprepes (Kellogg & Chapman, 1902). We agree with Mey’s assessment that four of his new species are junior synonyms of previously described taxa. Furthermore, among the species (subspecies) described by Mey (2017) as new, we establish 31 new generic (subgeneric) combinations, and we regard 16 species as species inquirenda, and three as incertae sedis.

To describe a new genus and two new species of chewing lice from Southeast Asian trogons (Trogoniformes). These lice belong in the Philopterus-complex. Slide-mounted lice were examined in a light microscope, illustrated by means of a drawing tube, and described using standard procedures. The new genus and species were successfully described. The genus Vinceopterus n. gen. is described from two species of Southeast Asian trogons (Trogoniformes: Harpactes). It presently comprises two species: Vinceopterus erythrocephali n. sp. from three subspecies of the Red-headed Trogon Harpactes erythrocephalus (Gould, 1834), and Vinceopterus mindanensis n. sp. from two subspecies of the Philippine Trogon Harpactes ardens (Temminck, 1826). Vinceopterus belongs to the Philopterus-complex, and thus likely constitutes a genus of head lice. Vinceopterus is the second new genus of chewing lice discovered on Southeast Asian trogons in recent years, the first genus of presumed head lice on trogons worldwide, and the fifth genus of chewing lice known from trogons globally. A translated and revised key to the Philopterus-complex is provided, as well as notes on the various chewing lice genera known from trogons.

The chewing louse subgenus Cicchinella new subgenus is erected and described for species of Guimaraesiella Eichler, 1949, parasitizing Old World babblers (Leiothrichidae, Pellorneidae, Timaliidae). The subgenus is divided into three species groups based on chaetotaxy, head and genitalia of both sexes. Two species are redescribed: Guimaraesiella (Cicchinella) sehri (Ansari, 1955) from Trochalopteron lineatum lineatum (Vigors, 1831) and T. lineatum setafer (Hodgson, 1836), and Guimaraesiella (Cicchinella) avinus (Ansari, 1956) from Trochalopteron subunicolor subunicolor Blyth, 1843. In addition, 12 new species are described and illustrated: Guimaraesiella (Cicchinella) falcifrons n. sp. from Actinodura cyanouroptera sordidior (Sharpe, 1888); Guimaraesiella (Cicchinella) iuga n. sp. from Alcippe peracensis peracensis Sharpe, 1887; Guimaraesiella (Cicchinella) gombakensis n. sp. from Turdinus abbotti abbotti (Blyth, 1845); Guimaraesiella (Cicchinella) mcgrewi n. sp. from Alcippe morrisonia Swinhoe, 1863; Guimaraesiella (Cicchinella) tenella n. sp. from Cyanoderma ruficeps davidi (Oustalet, 1899); Guimaraesiella (Cicchinella) retusa n. sp. from Trochalopteron milnei sinianum Stresemann, 1930; Guimaraesiella (Cicchinella) philiproundi n. sp. from Trochalopteron melanostigma schistaceum (Deignan, 1938) and Trochalopteron peninsulae Sharpe, 1887; Guimaraesiella (Cicchinella) hannesundinae n. sp. from Heterophasia picaoides wrayi (Ogilvie-Grant, 1910); Guimaraesiella (Cicchinella) pallidobrunneis n. sp. from Heterophasia melanoleuca melanoleuca (Blyth, 1859); Guimaraesiella (Cicchinella) hampuslybecki n. sp. from Heterophasia auricularis (Swinhoe, 1864); Guimaraesiella (Cicchinella) scottvillai n. sp. from Liocichla steerii Swinhoe, 1877 and Guimaraesiella (Cicchinella) ambusta n. sp. from Leiothrix argentauris rubrogularis Kinnear, 1925. A key to identify the species of Cicchinella, and both a checklist and a host-louse list of the species of Cicchinella known from the Old World babblers are provided.

Five new species of Guimaraesiella Eichler, 1949 are described and illustrated from hosts in the Eurylaimidae and Calyptomenidae. They are Guimaraesiella corydoni n. sp. from Corydon sumatranus laoensisMeyer de Schauensee, 1929; Guimaraesiella latirostris n. sp. from Eurylaimus ochromalusRaffles, 1822; Guimaraesiella cyanophoba n. sp. from Cymbirhynchus macrorhynchus malaccensisSalvadori, 1874 and C. m. siamensisMeyer de Schauensee and Ripley, 1940; Guimaraesiella altunai n. sp. from Calyptomena viridis caudacutaSwainson, 1838; and Guimaraesiella forcipata n. sp. from Eurylaimus steerii steeriiSharpe, 1876. These represent the first species of Guimaraesiella described from the Calyptomenidae and Eurylaimidae, as well as the first species of this genus described from the Old World suboscines. acceptedVersion

Ecological speciation occurs when local adaptation generates reproductive isolation as a by-product of natural selection1–3. Although ecological speciation is a fundamental source of diversification, the mechanistic link between natural selection and reproductive isolation remains poorly understood, especially in natural populations2–6. Here we show that experimental evolution of parasite body size over four years (ca. 60 generations) leads to reproductive isolation in natural populations of feather lice on birds. When lice are transferred to pigeons of different sizes they rapidly evolve differences in body size that are correlated with host size. These size differences trigger mechanical mating isolation between lice that are locally adapted to the different sized hosts. Size differences among lice also influence the outcome of competition between males for access to females. Thus, body size directly mediates reproductive isolation through its influence on both inter-sexual compatibility and intra-sexual competition. Our results confirm that divergent natural selection acting on a single phenotypic trait can cause reproductive isolation to emerge from a single natural population in real time.

Adaptive radiation occurs when the members of a single lineage evolve different adaptive forms in response to selection imposed by competitors or predators. Iconic examples include Darwin's finches, Caribbean anoles, and Hawaiian silverswords, all of which live on islands. Although adaptive radiation is thought to be an important generator of biodiversity, most studies concern groups that have already diversified. Here, we take the opposite approach. We experimentally triggered diversification in the descendants of a single population of host-specific parasites confined to different host “islands.” We show rapid adaptive divergence of experimentally evolving feather lice in response to preening, which is a bird's main defense against ectoparasites. We demonstrate that host defense exerts strong phenotypic selection for crypsis in lice transferred to different colored rock pigeons (Columba livia). During four years of experimental evolution (∼60 generations), the lice evolved heritable differences in color. Strikingly, the observed color differences spanned the range of phenotypes found among congeneric lice adapted to other species of birds. To our knowledge, this is the first real-time demonstration that microevolution is fast enough to simulate millions of years of macroevolutionary change. Our results further indicate that host-mediated selection triggers rapid divergence in the adaptive radiation of parasites, which are among the most diverse organisms on Earth.

Twelve new species of chewing lice in the genus Priceiella Gustafsson & Bush, 2017, are described from Old World “babblers” (Leiothrichidae, Paradoxornithiidae, Pellorneidae and Timaliidae). Eight species belong to the subgenus Thescelovora Gustafsson & Bush, 2017: Priceiella ( Thescelovora ) calcicola n. sp. from Turdinus crispifrons crispifrons ; Priceiella ( Thescelovora ) fuscicaena n. sp. from Malacopteron magnum magnum and Malacopteron cinereum cinereum ; Priceiella ( Thescelovora ) orichalca n. sp. from Turdinus brevicaudatus leucostictus and four other host species; Priceiella ( Thescelovora ) chanthaburiana n. sp. from Megapomatorhinus hypoleucos tickelli and Pomatorhinus schisticeps klossi ; Priceiella ( Thescelovora ) macrocephala n. sp. from Megapomatorhinus hypoleucos wrayi ; Priceiella ( Thescelovora ) coleyae n. sp. ex Stachyris striolata tonkinensis ; Priceiella ( Thescelovora ) austini n. sp. ex Pomatorhinus ruficollis intermedius ; and Priceiella ( Thescelovora ) ornata n. sp. ex Psittiparus gularis fokiensis . The remaining four species belong to the subgenus Camurnirmus Gustafsson & Bush, 2017: Priceiella ( Camurnirmus ) najeri n. sp. from Garrulax monileger stuarti and three other hosts; Priceiella ( Camurnirmus ) bohsae n. sp. from Garrulax strepitans ; Priceiella ( Camurnirmus ) lindquistae n. sp. ex Ianthocincla chinensis chinensis ; Priceiella ( Camurnirmus ) sonorae n. sp. ex Garrulax maesi maesi . We redescribe and illustrate Priceiella ( Camurnirmus ) rhinocichlae (Eichler, 1957). Keys to species of two subgenera, a comprehensive checklist of all the species of Priceiella , and a host-louse list are provided.

Ten new species of chewing lice in the genus Brueelia Kéler, 1936, are described from hosts in the families Paridae, Regulidae, and Sittidae. They are: Brueelia johnsoni n. sp. from Poecile sclateri eidos (Peters, 1927); Brueelia juniperi n. sp. from Baeolophus ridgwayi ridgwayi (Richmond, 1902); Brueelia kabulica n. sp. from Sitta tephronota tephronota Sharpe, 1872; Brueelia mpumalangensis n. sp. from Melaniparus niger niger (Vieillot, 1818); Brueelia nazae n. sp. from Parus cinereus caschmirensis Hartert, 1905; Brueelia oxyrhyncha n. sp. from Sitta nagaensis nagaensis Godwin-Austen, 1874; Brueelia picea. sp. from Parus major excelsus Buvry, 1857; Brueelia ragusica n. sp. from Sitta neumayer neumayer Michahelles, 1830; Brueelia regulicida n. sp. from Regulus calendula grinnelli Palmer, 1897; Brueelia sittacola n. sp. from Sitta carolinensis carolinensis Latham, 1790. Brueelia regulicida is the first Brueelia-complex louse to be described from the host family Regulidae. Collectively, the Brueelia of parid, regulid, and sittid hosts show two peculiar patterns. Firstly, lice on closely related hosts appear to be distantly related. Secondly, lice on most hosts in these families appear to be more closely related to lice on other host families than to each other. This contradicts the traditional view that Brueelia-complex lice on closely related hosts are themselves closely related. Potentially, the tendency of the hosts to participate in mixed-species feeding flocks may explain some of these patterns.

Behavior is usually the first line of defense against parasites. Antiparasite behaviors, such as grooming, or outright avoidance, have been shown to reduce the risk of parasitism in a wide variety of host–parasite systems. However, despite the central importance of antiparasite behavior, little is known about the extent to which prior exposure to parasites improves effectiveness. Here, we report the results of a 2-year study designed to test whether exposure to parasites can "prime" behavior, loosely analogous to priming of the immune system. We tested whether preening improves with experience by infesting captive-bred rock pigeons (Columba livia) with 2 common species of rock pigeon feather lice. We infested "primed" birds in Years 1 and 2 of the study and "nonprimed" birds only in Year 2. Birds with lice preened about a third more, on average, than birds without lice. Birds subsequently cleared of lice resumed preening at the same rate as birds that never had lice. Thus, our results confirm that preening is an inducible, reversible defense that is partly triggered by the presence of lice. Surprisingly, primed birds did not differ significantly from nonprimed birds in the overall rate or the efficacy of preening. Primed and nonprimed birds preened at similar rates and had similar numbers of lice at the end of the study. Our results therefore provide little evidence that antiparasite behavior improves with experience, at least in the case of preening as a defense against feather lice.

Songbirds host one of the largest, and most poorly understood, groups of lice: the Brueelia-complex. The Brueelia-complex contains nearly one-tenth of all known louse species (Phthiraptera), and the genus Brueelia has over 300 species. To date, revisions have been confounded by extreme morphological variation, convergent evolution, and periodic movement of lice between unrelated hosts. Here we use Bayesian inference based on mitochondrial (COI) and nuclear (EF-1α) gene fragments to analyze the phylogenetic relationships among 333 individuals within the Brueelia-complex. We show that the genus Brueelia, as it is currently recognized, is paraphyletic. Many well-supported and morphologically unified clades within our phylogenetic reconstruction of Brueelia were previously described as genera. These genera should be recognized, and the erection of several new genera should be explored. We show that four distinct ecomorphs have evolved repeatedly within the Brueelia-complex, mirroring the evolutionary history of feather-lice across the entire order. We show that lice in the Brueelia-complex, with some notable exceptions, are extremely host specific and that the host family associations and geographic distributions of these lice are significantly correlated with our understanding of their phylogenetic history. Several ecological phenomena, including phoresis, may be responsible for the macroevolutionary patterns in this diverse group.

Data is presented in support of a phylogenetic reconstruction of one of the largest, and most poorly understood, groups of lice: the Brueelia-complex (Bush et al., 2015 [1]). Presented data include the voucher information and molecular data (GenBank accession numbers) of 333 ingroup taxa within the Brueelia-complex and 30 outgroup taxa selected from across the order Phthiraptera. Also included are phylogenetic reconstructions based on Bayesian inference analyses of combined COI and EF-1α sequences for Brueelia-complex species and outgroup taxa.

The chewing louse fauna of pigeons and doves in Japan is reviewed based on published records and new collections. An updated checklist of the chewing lice of Japanese pigeons and doves is provided, and 3 new species are described: Columbicola asukae n. sp. and Coloceras nakamurai n. sp., both from Columba janthina Temminck, 1830 (Japanese wood pigeon), and Columbicola lemoinei n. sp. from Treron formosae permagnus Stejneger, 1887, and Treron formosae medioximus (Bangs, 1901) (whistling green-pigeons). This checklist includes data on the first records of Coloceras chinense (Kellogg and Chapman, 1902), Coloceras piriformis (Tendeiro, 1969), and Columbicola guimaraesi Tendeiro, 1965, in Japan. New host records of Hohorstiella sp. from Columba janthina and Treron formosae permagnus, and Coloceras sp. from Treron sieboldii sieboldii (Temminck, 1835) (white-bellied green-pigeon) are provided.

Birds have many kinds of internal and external parasites, including viruses, bacteria and fungi, as well as protozoa, helminths and arthropods. Because parasites have negative effects on host fitness, selection favours the evolution of anti-parasite defences, many of which involve behaviour. We provide a brief review of anti-parasite behaviours in birds, divided into five major categories: (i) body maintenance, (ii) nest maintenance, (iii) avoidance of parasitized prey, (iv) migration and (v) tolerance. We evaluate the adaptive significance of the different behaviours and note cases in which additional research is particularly needed. We briefly consider the interaction of different behaviours, such as sunning and preening, and how behavioural defences may interact with other forms of defence, such as immune responses. We conclude by suggesting some general questions that need to be addressed concerning the nature of anti-parasite behaviour in birds.This article is part of the Theo Murphy meeting issue ‘Evolution of pathogen and parasite avoidance behaviours’.

Our helminthological examination of murid rodents on Luzon Island, Philippines, revealed a remarkable diversity of Hymenolepis Weinland, 1858. Here we describe two new species based on specimens from murid rodents Rattus everetti (Günther) and Apomys datae (Meyer) collected from Luzon Island. Hymenolepis alterna n. sp. differs from all known species of Hymenolepis in having irregularly alternating genital pores. This feature has not been reported from any previously known member of Hymenolepis. Additionally, Hymenolepis alterna n. sp. also differs from other Hymenolepis spp. in the relative position of both poral and antiporal dorsal osmoregulatory canals which are shifted towards the middle of the proglottis in relation to the ventral canals on both sides of the proglottides, and in having curved or twisted external seminal vesicle, covered externally by a dense layer of intensely stained cells. Hymenolepis bilaterala n. sp. differs from all known species of Hymenolepis in the relative position of both poral and antiporal dorsal osmoregulatory canals, which are shifted bilaterally towards the margins of proglottides in relation to the ventral canals, and in possession of testes situated in a triangle and eggs with very thin outer coat. A total of seven species of Hymenolepis are known from the Philippine archipelago. This total includes the cosmopolitan species Hymenolepis diminuta (Rudolphi, 1819), which was likely introduced to the island with invasive rats. Strikingly, all seven known species occur on the island of Luzon alone. By comparison, only six Hymenolepis spp. are known from the whole Palaearctic and seven from the Nearctic despite a much better level of knowledge of rodent helminths in these zoogeographical regions, as well as vast territories, diverse landscapes and very rich rodent fauna. This suggests that Hymenolepis spp. may have undergone an unusually active radiation in the Philippines. Possible explanations of this phenomenon are discussed.

Dispersal is a fundamental component of the life history of most species. Dispersal influences fitness, population dynamics, gene flow, genetic drift and population genetic structure. Even small differences in dispersal can alter ecological interactions and trigger an evolutionary cascade. Linking such ecological processes with evolutionary patterns is difficult, but can be carried out in the proper comparative context. Here, we investigate how differences in phoretic dispersal influence the population genetic structure of two different parasites of the same host species. We focus on two species of host-specific feather lice (Phthiraptera: Ischnocera) that co-occur on feral rock pigeons (Columba livia). Although these lice are ecologically very similar, "wing lice" (Columbicola columbae) disperse phoretically by "hitchhiking" on pigeon flies (Diptera: Hippoboscidae), while "body lice" (Campanulotes compar) do not. Differences in the phoretic dispersal of these species are thought to underlie observed differences in host specificity, as well as the degree of host-parasite cospeciation. These ecological and macroevolutionary patterns suggest that body lice should exhibit more genetic differentiation than wing lice. We tested this prediction among lice on individual birds and among lice on birds from three pigeon flocks. We found higher levels of genetic differentiation in body lice compared to wing lice at two spatial scales. Our results indicate that differences in phoretic dispersal can explain microevolutionary differences in population genetic structure and are consistent with macroevolutionary differences in the degree of host-parasite cospeciation.

Comparisons of host and parasite phylogenies often show varying degrees of phylogenetic congruence. However, few studies have rigorously explored the factors driving this variation. Multiple factors such as host or parasite morphology may govern the degree of phylogenetic congruence. An ideal analysis for understanding the factors correlated with congruence would focus on a diverse host–parasite system for increased variation and statistical power. In this study, we focused on the Brueelia-complex, a diverse and widespread group of feather lice that primarily parasitise songbirds. We generated a molecular phylogeny of the lice and compared this tree with a phylogeny of their avian hosts. We also tested for the contribution of each host–parasite association to the overall congruence. The two trees overall were significantly congruent, but the contribution of individual associations to this congruence varied. To understand this variation, we developed a novel approach to test whether host, parasite or biogeographic factors were statistically associated with patterns of congruence. Both host plumage dimorphism and parasite ecomorphology were associated with patterns of congruence, whereas host body size, other plumage traits and biogeography were not. Our results lay the framework for future studies to further elucidate how these factors influence the process of host–parasite coevolution.

Five new species of chewing lice in the genus BrueeliaKeler, 1936, are described from North American jays and allies. They are Brueelia mexicana n. sp. from Aphelocoma woodhouseii cyanotisRidgway, 1887; Brueelia bonnevillensis n. sp. from Aphelocoma woodhouseii nevadaePitelka, 1945; Brueelia diblasiae n. sp. from Cyanocitta stelleri frontalis (Ridgway, 1873); Brueelia tempestwilliamsae n. sp. from Gymnorhinus cyanocephalaWied-Neuwied, 1841; Brueelia mcnewae n. sp. from Nucifraga columbiana (Wilson, 1811). An identification key to the Brueelia on corvid hosts is provided.

Nine new species of chewing lice in the genus BrueeliaKeler, 1936, are described from North American hosts. They are Brueelia thorini n. sp. from Haemorhous mexicanus frontalis (Say, 1822) and Haemorhous mexicanus potosinus Griscom, 1928; Brueelia straseviciusi n. sp. from Haemorhous purpureus (Gmelin, 1789); Brueelia mattsonae n. sp. from Coccothraustes vespertinus brooksi (Grinnell, 1917); Brueelia novemstriata n. sp. from Icterus wagleri wagleri Sclater, 1857, and I. parisorum Bonaparte, 1838; Brueelia benkmani n. sp. from Pheucticus melanocephalus (Swainson, 1827); Brueelia arizonae n. sp. from Passerina caerulea (Linnaeus, 1758); Brueelia hellstromi n. sp. from Piranga ludoviciana (Wilson, 1811); Brueelia dolorosa n. sp. from Spinus pinus pinus (Wilson, 1810); and Brueelia melancholica n. sp. from Spinus tristis (Linnaeus, 1758). Brueelia limbata (Burmeister, 1838) is redescribed and illustrated from North American material. A key for these North American species is provided.

When confronted with a parasite or pathogen, hosts can defend themselves by resisting or tolerating the attack. While resistance can be diminished when resources are limited, it is unclear how robust tolerance is to changes in environmental conditions. Here, we investigate the sensitivity of tolerance in a single host population living in a highly variable environment. We manipulated the abundance of an invasive parasitic fly,Philornis downsi, in nests of Galápagos mockingbirds (Mimus parvulus) over four field seasons and measured host fitness in response to parasitism. Mockingbird tolerance toP. downsivaried significantly among years and decreased when rainfall was limited. Video observations indicate that parental provisioning of nestlings appears key to tolerance: in drought years, mockingbirds likely do not have sufficient resources to compensate for the effects ofP. downsi. These results indicate that host tolerance is a labile trait and suggest that environmental variation plays a major role in mediating the consequences of host–parasite interactions.

Two previously unrecognized species of the genus Hymenolepis are described based on specimens obtained from murid rodent species Bullimus luzonicus, Apomys microdon, and Rattus everetti collected on Luzon Island, Philippines. Hymenolepis bicauda n. sp. differs from all known Hymenolepis spp. in relative position of the poral dorsal and ventral osmoregulatory canals, gravid uterus occupying less than half the length of proglottid, relatively few eggs, and the highly characteristic longitudinal split of proglottids at the end of the gravid strobila. Hymenolepis haukisalmii n. sp. differs from all known Hymenolepis spp. in the relative position of both poral and aporal dorsal and ventral osmoregulatory canals and uterus lacking dorsal and ventral diverticula. The shift in the relative position of the dorsal and ventral osmoregulatory canals was not known in Hymenolepis from rodents in other regions of the world and is reminiscent of the situation observed in Hymenolepis erinacei, parasitic in hedgehogs, and members of the genus Talpolepis, parasitic in moles. The cosmopolitan species Hymenolepis diminuta was the only member of the genus previously reported from the Philippines. The genus Hymenolepis Weinland, 1858, includes hymenolep- idid cestodes with an unarmed scolex and rudimentary rostellar apparatus, parasitic primarily in rodents, with a few species known from bats and 1 from hedgehogs. Members of the genus have been reported from the Palearctic, Nearctic, Ethiopian, and Oriental regions (Lopez-Neyra, 1942a, 1942b; Skrjabin and Matevosyan, 1948; Spassky, 1954; Yamaguti, 1959; Hunkeler, 1972; Ryzhikov et al., 1978; Genov, 1984; Gardner, 1985; Schmidt, 1986; Gardner and Schmidt, 1988; Czaplinski and Vaucher, 1994; Mas-Coma and Tenora, 1997; Sawada, 1997; Gulyaev and Melnikova, 2005; Makarikova et al., 2010; Makarikov and Tkach, 2013). To the best of our knowledge, Hymenolepis diminuta (Rudolphi, 1819) is the only species of Hymenolepis previously reported from the Philippines (Tubangui, 1931; Fedorko, 1999). As part of a biodiversity survey of terrestrial vertebrates and their parasites in the Philippines, we found hymenolepidid cestodes belonging to Hymenolepis in three species of murid rodents, namely, the large Luzon forest rat Bullimus luzonicus (Thomas, 1895), the small Luzon forest mouse Apomys microdon Hollister, 1913, and the Philippine forest rat Rattus everetti (G ¨ unther, 1879), collected in Aurora Province, Luzon Island. These 2 species of cestodes, described herein, are morphologically distinct from previously known Hymenolepis species.

Two new species of Eimeria Schneider, 1875, from emerald tree skinks, Lamprolepis smaragdina (Lesson) are described from specimens collected in Papua New Guinea (PNG) and the Philippines. Oocysts of Eimerianuiailan n. sp. from the only L. smaragdina from PNG are ovoidal, with a smooth, colourless, bi-layered wall, measure 23.7 × 19.1 μm, and have a length/width (L/W) ratio of 1.3; both micropyle and oocyst residuum are absent, but a fragmented polar granule is present. Sporocysts are ovoidal to ellipsoidal, 11.9 × 7.0 μm, L/W 1.7, and the wall is composed of two valves joined by a longitudinal suture; neither Stieda nor sub-Stieda bodies are present; a sporocyst residuum is present as a compact mass of granules. Sporozoites are elongate, 14.6 × 2.6 μm, and contain anterior and posterior refractile bodies with a nucleus between them. Oocysts of Eimeriaauffenbergi n. sp. from L. smaragdina collected in the Philippines are ovoidal, with a smooth, colourless, bi-layered wall, measure 19.9 × 15.8 μm, L/W 1.3; both micropyle and oocyst residuum are absent, but one to four polar granules are present. Sporocysts are ovoidal to ellipsoidal, 10.3 × 5.8 μm, L/W 1.8, and the wall is composed of two valves joined by a longitudinal suture; neither Stieda nor sub-Stieda bodies are present; a sporocyst residuum is composed of dispersed granules.