INFLUENCE OF GROOMING ON PERMANENT ARTHROPOD ASSOCIATES OF BIRDS: CATTLE EGRETS, LICE, AND MITES.

Birds have a diverse community of "permanent" arthropods that complete their entire life cycle on the body of the host. Because some of these arthropods are parasites that reduce host fitness, birds control them by grooming, which consists of preening with the beak and scratching with the feet. Although preening is the primary component of grooming, scratching is essential for controlling arthropods on the head and neck, which cannot be preened. Several unrelated groups of birds have evolved comb-like pectinate claws on the middle toenail of each foot. We tested the role of these claws in the control of arthropods by experimentally removing teeth from the claws of captive western cattle egrets (Bubulcus ibis) infested with chewing lice (Insecta: Phthiraptera), feather mites (Acari: Sarcoptiformes), and nasal mites (Acari: Mesostigmata). After a period of 4 mo, we compared the abundance of arthropods on experimental birds to that of control birds with intact teeth. We used video to quantify the grooming rates of the captive birds, which groomed twice as much as wild birds. Experimental and control birds did not differ significantly in grooming time. Both groups virtually eradicated the chewing lice, but not feather mites or nasal mites. We found no support for the hypothesis that pectinate claws increase the efficiency of arthropod control by grooming. Experiments with wild birds are needed to test the hypothesis further under conditions in which birds devote less time to grooming.

The fitness landscape of a community of Darwin's finches.

Divergent natural selection should lead to adaptive radiation-that is, the rapid evolution of phenotypic and ecological diversity originating from a single clade. The drivers of adaptive radiation have often been conceptualized through the concept of "adaptive landscapes," yet formal empirical estimates of adaptive landscapes for natural adaptive radiations have proven elusive. Here, we use a 17-year dataset of Darwin's ground finches (Geospiza spp.) at an intensively studied site on Santa Cruz (Galápagos) to estimate individual apparent lifespan in relation to beak traits. We use these estimates to model a multi-species fitness landscape, which we also convert to a formal adaptive landscape. We then assess the correspondence between estimated fitness peaks and observed phenotypes for each of five phenotypic modes (G. fuliginosa, G. fortis [small and large morphotypes], G. magnirostris, and G. scandens). The fitness and adaptive landscapes show 5 and 4 peaks, respectively, and, as expected, the adaptive landscape was smoother than the fitness landscape. Each of the five phenotypic modes appeared reasonably close to the corresponding fitness peak, yet interesting deviations were also documented and examined. By estimating adaptive landscapes in an ongoing adaptive radiation, our study demonstrates their utility as a quantitative tool for exploring and predicting adaptive radiation.

Co-parasitism in the face of predation: Effects of natural enemies on a neotropical mockingbird.

Co-parasitism is ubiquitous and has important consequences for the ecology and evolution of wild host populations. Studies of parasite co-infections remain limited in scope, with few experimental tests of the fitness consequences of multiple parasites, especially in natural populations. We measured the separate and combined effects of Philornis seguyi nest flies and shiny cowbirds Molothrus bonariensis on the fitness of a shared host, the chalk-browed mockingbird (Mimus saturninus) in Argentina. Using a two-factor experimental approach, we manipulated the presence of nest flies and cowbirds in mockingbird nests and assessed their effects on mockingbird haemoglobin levels, begging and provisioning rates, body size, and fledging success. We also monitored rates of nest predation in relation to parasitism by flies and cowbirds. Nest flies reduced the haemoglobin concentration, body size, and fledging success of mockingbirds, likely because mockingbirds did not compensate for parasitism by begging more or feeding their nestlings more. Cowbirds also reduced the fledging success of mockingbirds, even though they had no detectable effect on haemoglobin or body size. Nests with cowbirds, which beg more than mockingbirds, attracted more nest predators. There was no significant interaction between the effects of flies and cowbirds on any component of mockingbird fitness. The combined effects of nest flies and cowbirds were strictly additive. In summary, we show that nest flies and cowbirds both reduce host fitness, but do not have interactive effects in co-parasitized nests. Our results further suggest that predators exacerbate the effects of nest flies and cowbirds on their hosts. Our study shows that the fitness consequences of co-parasitism are complex, especially in the context of community-level interactions.

El co-parasitismo es ubicuo y tiene consecuencias importantes para la ecología y la evolución de las poblaciones de hospedadores silvestres. Los estudios de coinfecciones por parásitos poseen un alcance limitado, con pocos trabajos experimentales que analicen las consecuencias del parasitismo múltiple sobre la eficacia biológica, especialmente en poblaciones naturales. Medimos los efectos separados y combinados de las moscas de los nidos Philornis seguyi y el tordo renegrido (Molothrus bonariensis) sobre la eficacia biológica de un hospedador compartido, la calandria grande (Mimus saturninus) en Argentina. Utilizando un enfoque experimental de dos factores, manipulamos la presencia de moscas de nido y tordos en los nidos de calandrias y evaluamos sus efectos sobre los niveles de hemoglobina, las tasas de solicitud de alimento y aprovisionamiento, el tamaño corporal y el éxito de supervivencia de los pichones de calandria. También monitoreamos las tasas de depredación de nidos en relación con el parasitismo de moscas y tordos. Las moscas de los nidos redujeron la concentración de hemoglobina, el tamaño corporal y el éxito de supervivencia de los pichones de calandrias, probablemente porque las calandrias no compensaron el parasitismo solicitando más alimento o alimentando más a sus pichones. Los tordos también redujeron el éxito de supervivencia de las calandrias, aunque no tuvieron un efecto detectable sobre la hemoglobina o el tamaño corporal. Los nidos con tordos, quienes solicitaron más alimento más que las calandrias, atrajeron a más depredadores de nidos. El aumento de la depredación de nidos asociado a la alta intensidad de solicitud de alimento puede explicar por qué las calandrias en Argentina no solicitan más alimento en respuesta al parasitismo de las moscas. No hubo una interacción significativa entre los efectos de las moscas y los tordos en ningún componente de la eficacia biológica de las calandrias. Los efectos combinados de las moscas de los nidos y los tordos fueron estrictamente aditivos. En resumen, mostramos que las moscas de los nidos y los tordos reducen la eficacia biológica del hospedador, pero no tienen efectos interactivos en los nidos co-parasitados. Nuestros resultados sugieren además que los depredadores exacerban los efectos de las moscas de los nidos y los tordos sobre sus hospedadores. Nuestro estudio muestra que las consecuencias del co-parasitismo sobre la eficacia biológica son complejas, especialmente en el contexto de las interacciones a nivel comunitario.

DOES PREENING BEHAVIOR REDUCE THE PREVALENCE OF AVIAN FEATHER LICE (PHTHIRAPTERA: ISCHNOCERA)?

Animals defend themselves against parasites in many ways. Defenses, such as physiological immune responses, are capable of clearing some infections. External parasites that do not feed on blood, however, are not controlled by the physiological immune system. Instead, ectoparasites like feather-feeding lice (Phthiraptera: Ischnocera) are primarily controlled by behavioral defenses such as preening. Here we test the hypothesis that birds able to preen are capable of clearing infestations of feather lice. We experimentally manipulated preening ability in a captive population of rock pigeons (Columba livia) that were infested with identical numbers of feather lice (Columbicola columbae or Campanulotes compar or both). We then monitored the feather louse infestations for 42 wk. Birds with impaired preening remained infested throughout the experiment; in contrast, the prevalence of lice on birds that could preen normally decreased by 50%. These data indicate that it is indeed possible for birds to clear themselves of feather lice, and perhaps other ectoparasites, by preening. We note, however, that captive birds spend more time preening than wild birds, and that they are less likely to be reinfested than wild birds. Thus, additional studies are necessary to determine under what circumstances wild birds can clear themselves of ectoparasites by preening.

Grooming Time Predicts Survival: American Kestrels, Falco sparverius, on a Subtropical Island.

AbstractAnimals have evolved a variety of adaptations to care for their body surfaces, such as grooming behavior, which keeps the integument clean, parasite-free, and properly arranged. Despite extensive research on the grooming of mammals, birds, and arthropods, the survival value of grooming has never been directly measured in natural populations. We monitored grooming and survival in a population of marked American kestrels (Falco sparverius) on San Salvador Island, Bahamas. We found a strong association between time spent grooming and survival over a 2-year period. The quadratic relationship we show is consistent with stabilizing natural selection on grooming time. To our knowledge, this is the first evidence for a correlation between grooming time and survival in a natural population. Grooming time may predict the survival of many animal taxa, but additional studies are needed to determine the shape and strength of the relationship among birds, mammals, and arthropods.

The terroir of the finch: How spatial and temporal variation shapes phenotypic traits in DARWIN'S finches.

The term terroir is used in viticulture to emphasize how the biotic and abiotic characteristics of a local site influence grape physiology and thus the properties of wine. In ecology and evolution, such terroir (i.e., the effect of space or "site") is expected to play an important role in shaping phenotypic traits. Just how important is the pure spatial effect of terroir (e.g., differences between sites that persist across years) in comparison to temporal variation (e.g., differences between years that persist across sites), and the interaction between space and time (e.g., differences between sites change across years)? We answer this question by analyzing beak and body traits of 4388 medium ground finches (Geospiza fortis) collected across 10 years at three locations in Galápagos. Analyses of variance indicated that phenotypic variation was mostly explained by site for beak size (η 2 = 0.42) and body size (η 2 = 0.43), with a smaller contribution for beak shape (η 2 = 0.05) and body shape (η 2 = 0.12), but still higher compared to year and site-by-year effects. As such, the effect of terroir seems to be very strong in Darwin's finches, notwithstanding the oft-emphasized interannual variation. However, these results changed dramatically when we excluded data from Daphne Major, indicating that the strong effect of terroir was mostly driven by that particular population. These phenotypic results were largely paralleled in analyses of environmental variables (rainfall and vegetation indices) expected to shape terroir in this system. These findings affirm the evolutionary importance of terroir, while also revealing its dependence on other factors, such as geographical isolation.

Long-distance dispersal of pigeons and doves generated new ecological opportunities for host-switching and adaptive radiation by their parasites.

Adaptive radiation is an important mechanism of organismal diversification and can be triggered by new ecological opportunities. Although poorly studied in this regard, parasites are an ideal group in which 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 adaptively radiated, leading to differences in body size and overall coloration. Here, we show that long-distance dispersal by dove hosts was central to parasite diversification because it provided new ecological opportunities for parasites to speciate after host-switching. We further show that among extant parasite lineages host-switching decreased over time, with cospeciation becoming 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.

Epigenetic effects of parasites and pesticides on captive and wild nestling birds.

Anthropogenic changes to the environment challenge animal populations to adapt to new conditions and unique threats. While the study of adaptation has focused on genetic variation, epigenetic mechanisms may also be important. DNA methylation is sensitive to environmental stressors, such as parasites and pesticides, which may affect gene expression and phenotype. We studied the effects of an invasive ectoparasite, Philornis downsi, on DNA methylation of Galápagos mockingbirds (Mimus parvulus). We used the insecticide permethrin to manipulate P. downsi presence in nests of free-living mockingbirds and tested for effects of parasitism on nestling mockingbirds using epiGBS, a reduced-representation bisulfite sequencing (RRBS) approach. To distinguish the confounding effects of insecticide exposure, we conducted a matching experiment exposing captive nestling zebra finches (Taeniopygia guttata) to permethrin. We used zebra finches because they were the closest model organism to mockingbirds that we could breed in controlled conditions. We identified a limited number of differentially methylated cytosines (DMCs) in parasitized versus nonparasitized mockingbirds, but the number was not more than expected by chance. In contrast, we saw clear effects of permethrin on methylation in captive zebra finches. DMCs in zebra finches paralleled documented effects of permethrin exposure on vertebrate cellular signaling and endocrine function. Our results from captive birds indicate a role for epigenetic processes in mediating sublethal nontarget effects of pyrethroid exposure in vertebrates. Environmental conditions in the field were more variable than the laboratory, which may have made effects of both parasitism and permethrin harder to detect in mockingbirds. RRBS approaches such as epiGBS may be a cost-effective way to characterize genome-wide methylation profiles. However, our results indicate that ecological epigenetic studies in natural populations should consider the number of cytosines interrogated and the depth of sequencing in order to have adequate power to detect small and variable effects.

A MISIDENTIFICATION CRISIS PLAGUES SPECIMEN-BASED RESEARCH: A CASE FOR GUIDELINES WITH A RECENT EXAMPLE (ALI ET AL., 2020).

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 assembled and annotated genome of the pigeon louse Columbicola columbae, a model ectoparasite.

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.

Is Allopreening a Stimulus-Driven Defense Against Ectoparasites?

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 (Columba livia) to test the relationship between ectoparasite load and allopreening rate. We added feather lice (Columbicola columbae) to captive pigeons and tested for changes in allopreening rates compared to control birds with no lice added. Allopreening rates did not change in response to the addition of lice. Interestingly, however, our data revealed a negative correlation between allopreening and self-preening rates.

Parasitism by an invasive nest fly reduces future reproduction in Galápagos mockingbirds.

Organisms allocate limited resources to competing activities such as reproduction, growth, and defense against parasites and predators. The introduction of a novel parasite may create new life history trade-offs. As hosts increase their investment in self-maintenance or defense, the cost of parasitism may carry over to other aspects of host biology. Here, in an experimental field study, we document delayed effects of an introduced nest parasite, Philornis downsi, on reproduction of Galápagos mockingbirds (Mimus parvulus). Parasitism of first nests reduced both the number and size of chicks that parents hatched when they re-nested several weeks later. The delayed effect of P. downsi on future reproduction may have been mediated by behavioral shifts by the parents to avoid or resist parasitism. Our results demonstrate that effects of parasitism can persist even after immediate exposure ends. We draw attention to the potential implications that introduced parasites have for host reproductive strategies.

Temporally varying disruptive selection in the medium ground finch (Geospiza fortis).

Disruptive natural selection within populations exploiting different resources is considered to be a major driver of adaptive radiation and the production of biodiversity. Fitness functions, which describe the relationships between trait variation and fitness, can help to illuminate how this disruptive selection leads to population differentiation. However, a single fitness function represents only a particular selection regime over a single specified time period (often a single season or a year), and therefore might not capture longer-term dynamics. Here, we build a series of annual fitness functions that quantify the relationships between phenotype and apparent survival. These functions are based on a 9-year mark-recapture dataset of over 600 medium ground finches (Geospiza fortis) within a population bimodal for beak size. We then relate changes in the shape of these functions to climate variables. We find that disruptive selection between small and large beak morphotypes, as reported previously for 2 years, is present throughout the study period, but that the intensity of this selection varies in association with the harshness of environment. In particular, we find that disruptive selection was strongest when precipitation was high during the dry season of the previous year. Our results shed light on climatic factors associated with disruptive selection in Darwin's finches, and highlight the role of temporally varying fitness functions in modulating the extent of population differentiation.

Rapid experimental evolution of reproductive isolation from a single natural population.

Ecological speciation occurs when local adaptation generates reproductive isolation as a by-product of natural selection. Although ecological speciation is a fundamental source of diversification, the mechanistic link between natural selection and reproductive isolation remains poorly understood, especially in natural populations. Here, we show that experimental evolution of parasite body size over 4 y (approximately 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 differences in size 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 intersexual compatibility and intrasexual 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.

Host defense triggers rapid adaptive radiation in experimentally evolving parasites.

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.

Annual environmental variation influences host tolerance to parasites.

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 to P. downsi varied 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 of P. 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.

Twelve new species of Guimaraesiella (Phthiraptera: Ischnocera: Philopteridae) from "babblers" (Passeriformes: Leiothrichidae, Pellorneidae, Timaliidae) with a description of a new subgenus and a key to its species.

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.

Anti-parasite behaviour of birds.

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'.

Phoretic dispersal influences parasite population genetic structure.

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.

Twelve new species of Priceiella (Phthiraptera: Ischnocera: Philopteridae) from Old World babblers, with keys to species of two subgenera and checklists of species for the genus.

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.