Molecular phylogenetics of the avian feather louse Philopterus-complex (Phthiraptera: Philopteridae).

The avian feather louse Philopterus-complex (Phthiraptera: Ischnocera: Philopteridae) currently contains 12 genera that have been grouped together because of shared morphological characteristics. Although previously lumped into a single genus (Philopterus), more recent morphological treatments have separated the group into several different genera. Here we evaluate the status of these genera using DNA sequence data from 118 ingroup specimens belonging to ten genera in the Philopterus-complex: Australophilopterus Mey, 2004, Cinclosomicola Mey 2004, Clayiella Eichler, 1940, Corcorides Mey, 2004, Mayriphilopterus Mey, 2004, Paraphilopterus Mey 2004, Philopteroides Mey 2004, Philopterus Nitzsch, 1818, Tyranniphilopterus Mey, 2004, and Vinceopterus Gustafsson, Lei, Chu, Zou, and Bush, 2019. Our sampling includes 97 new louse-host association records. Our analyses suggest that the genus Debeauxoecus Conci, 1941, parasitic on pittas (Aves: Pittidae), is outside of the Philopterus-complex, and that there is strong support for the monophyly of a group containing the remaining genera from the complex. Some diverse genera, such as Philopterus (sensu stricto) and Mayriphilopterus are supported as monophyletic, whereas the genera Australophilopterus, Philopteroides, and Tyranniphilopterus are not. The present study is the largest phylogenetic reconstruction of avian lice belonging to the Philopterus-complex to date and suggests that further generic revision is needed in the group to integrate molecular and morphological information.

Integrative taxonomic description of the chewing louse Tricholipeurus lipeuroides infesting Odocoileus virginianus veraecrucis white-tailed deer in Veracruz, Mexico.

Chewing lice (Phthiraptera: Amblycera, Ischnocera) represent a component of the ectoparasite fauna associated with large sized mammals as deers. However, the diversity of chewing louse species infesting deer remains to be fully characterized in the Neotropics. Little is known about the chewing lice infesting the extant fourteen subspecies of white-tailed deer (Odocoileus virginianus) in Mexico. Known to infest white-tailed deer (WTD) in Canada and the United States (U.S.), Tricholipeurus lipeuroides is a chewing louse species that was originally described in the nineteenth century infesting O. v. mexicanus in Mexico. For the first time, infestation of O. v. veraecrucis, a Neotropical WTD subspecies in Mexico, with T. lipeuroides is reported herein. An integrative taxonomic approach was taken by combining morphological and molecular analyses to describe the T. lipeuroides infestion of O. v. veraecrucis. Ecological parameters of the T. lipeuroides infestations were also calculated. The prevalence was 91.7% of the 56 O. v. veraecrucis (29 females and 27 males) inspected while under chemical restraint that were sampled at 3 sites in the central region of Veracruz state in Mexico. The amplification and sequencing of previously reported T. lipeuroides Cytochrome Oxidase Subunit I gene confirmed the identity of all the chewing louse life stages. These results are discussed in the context of comparative analyses on the emergence of novel chewing lice-deer associations.

New host/parasite record for very rare chewing louse Cuculotogaster arabicus (Clay, 1938) (Phthiraptera: Ischnocera) on endemic mountain partridges of Arabian Peninsula.

The endemic avifauna of Arabia is unique and characteristic through the whole region. Little is known about these birds and their ectoparasites. The Arabian partridge Alectoris melanocephala (Rüppell, 1835) and Philby's partridge Alectoris philbyi Lowe, 1934 are two endemic species which are distributed through the Sarawat Mountains in Saudi Arabia and Yemen. Captive breeding population of these birds were examined for chewing lice at the National Wildlife Research Center (NWRC) near Ta'if. Only one rare species of louse, Cuculotogaster arabicus (Clay 1938), was found to infest these birds. The occurrence of such species is considered a first record of the genus Cuculotogaster from Saudi Arabia, and the association of C. arabicus with Philby's partridge is considered a new host/parasite association. The clear diagnostic characters, high definition photos, and drawing of male genitalia are available through this paper.

The Chewing Lice (Insecta, Phthiraptera) Fauna of the Swainson's Warbler, Limnothlypis swainsonii (Aves, Parulidae).

We examined Swainson's warblers (Limnothlypis swainsonii (Audubon, 1834), Aves: Parulidae) for lice fauna during 2 yr at three study sites in Arkansas, USA. A total of 66 individuals were examined; eight birds (10.6%) were parasitized with 16 lice of two new species belonging to two genera Myrsidea Waterson, 1915 (Amblycera: Menoponidae) and Brueelia Kéler, 1936 (Ischnocera: Philopteridae). Parasitological parameter data are given on the prevalence of lice on Swainson's warblers. Species descriptions and illustrations are provided for Myrsidea bensoni sp. nov. and Brueelia limnothlypiae sp. nov.; including a key for females of the genus Myrsidea that parasitize Parulidae (Passeriformes).

Chewing lice (Phthiraptera: Amblycera, Ischnocera) from Red Sea gulls with new host-parasite records.

Knowledge about chewing lice from marine birds of the Red Sea is minimal. Five species of gulls were examined for chewing lice in three different localities of the Saudi Arabian Red Sea coast. Two gull species were examined for lice for the first time (Larus armenicus Buturlin, 1934 and Larus michahellis Naumann, 1840) and their lice represent new host-louse associations. Four species and two subspecies of lice were identified from 159 specimens collected. Actornithophilus piceus lari (Packard, 1870) and Austromenopon transversum (Denny, 1842) (suborder: Amblycera), and Quadraceps punctatus (Burmeister, 1838) and Saemundssonia lari (O. Fabricius, 1780) (suborder: Ischnocera) were recorded for the first time from Saudi Arabia and Red Sea birds. Taxonomic and ecological notes, type hosts, data on specimens examined, collecting localities, an identification key, and photographs of each species and subspecies are given. 

Three new species of chewing lice of the genus Emersoniella Tendeiro, 1965 (Insecta: Phthiraptera: Ischnocera: Philopteridae) from Papua New Guinean kingfishers and kookaburras (Aves: Coraciiformes: Alcedinidae).

Three new species of the ischnoceran louse genus Emersoniella (Phthiraptera) are described from four species of New Guinean kingfishers and kookaburras (Coraciiformes: Alcedinidae: Halcyoninae). They are: Emersoniella crassicarina n. sp. ex Dacelo gaudichaud Quoy & Gaimard (rufous-bellied kookaburra) and Dacelo leachii intermedia Salvadori (blue-winged kookaburra); E. reninoda n. sp. ex Melidora macrorrhina macrorhina Lesson (hook-billed kingfisher); and E. persei n. sp. ex Tanysiptera danae Sharpe (brown-headed paradise-kingfisher). In addition, we illustrate Emersoniella regis Emerson & Price, Emersoniella halcyonis Tendeiro, and the male genitalia of Emersoniella galateae Emerson & Price, as well as provide a complete host-louse checklist, and an updated key to all seven species of this genus. 

Chewing lice (Phthiraptera: Amblycera and Ischnocera) from wild birds in southern Vietnam, with descriptions of two new species.

A total of 239 individuals of 50 bird species were examined for chewing lice (Insecta: Phthiraptera) in southern Vietnam. Fifty-six birds of 20 species were parasitised by 15 species of lice belonging to 10 genera from two suborders, Amblycera: Menacanthus, Meromenopon, Myrsidea, and Ischnocera: Alcedoecus, Brueelia, Cuculicola, Meropoecus, Penenirmus, Philopteroides and Philopterus. Thirteen louse samples from Passeriformes were identified to genus only because they contain inadequate material. A total of 29 host-louse associations were found, of which nine are new, including: (1) two new species of the genus Brueelia, which are described and named in this paper: Brueelia binhchauensis from Megalaima lineata (Vieillot, 1816) (Piciformes: Megalaimidae), and Brueelia malacocincla from Malacocincla abbotti Blyth, 1845 (Passeriformes: Pellorneidae); (2) first records of lice from Cyornis hainanus (Ogilvie-Grant, 1900); and (3) the first record of Myrsidea claytoni Hellenthal & Price, 2003 from Cymbirhynchus macrorhynchos (Gmelin, 1788) (Passeriformes: Eurylaimidae), here regarded as a case of natural host-switching. A portion of the mitochondrial cytochrome oxidase I (COI) gene for some species of chewing lice was sequenced in order to assess their genetic divergences.

Co-phylogeography and morphological evolution of sika deer lice (Damalinia sika) with their hosts (Cervus nippon).

Lice are obligate parasites of mammals and birds and have become an important model for studies of host-parasite co-evolution and co-phylogenetics. Population genetic and phylogeographic studies represent an important bridge between microevolution and co-phylogenetic patterns. We examine co-phylogeographic patterns in sika deer and their parasitic lice. Co-phylogeographic patterns in deer and lice were evaluated using homologous regions of mitochondrial COI sequences. The phylogeographic breaks recovered for deer populations matched those of previous studies. Comparisons of the phylogeographic tree topology for deer lice with that of their hosts revealed a significant level of congruence. However, comparisons of genetic distances between deer and lice suggested that one of the estimated co-divergence events is more likely a recent host switch. Taking into account genetic divergence, there is not strong evidence for complete phylogeographic co-divergence between deer and their parasitic lice. However, mitochondrial phylogenies only track genetic structure of female lineages, and the incongruence between deer and louse phylogeography may be explained by louse migration mediated by male deer. Morphological analysis of head shape variation based on an elliptic Fourier descriptor showed that overall morphological variation contained phylogenetic signal, suggesting that in general morphology of these lice evolves congruent to population history.

Two new species of Cotingacola Carriker, Phthiraptera: Ischnocera: Philopteridae) from Amazonian Brazil, with comments on host-specificity.

This paper describes two new species of Cotingacola Carriker, 1956, C. lutzae n. sp. and C. engeli n. sp., from the cinereous mourner Laniocera hypopyrra (Vieillot) and black-necked red cotinga Phoenicircus nigricollis Swainson (Aves: Passeriformes: Cotingidae), respectively. These species are fully illustrated and compared with their morphologically closest relatives. With the addition of these two new species, this Neotropical chewing louse genus now includes ten species. We also present records from Brazil for two previously described species, C. stotzi Clayton & Price, 1998 and C. parmipapillae Carriker, 1956. The re-examination of a specimen identified as 'Cotingacola sp.' in a recent phylogenetic study confirmed that it is C. parmipapillae and that Querula purpurata (Statius Müller) regularly hosts two Cotingacola species, which means that at least three cotingid hosts carry more than one species of Cotingacola. We comment on the variability in the host-specificity of Cotingacola species.

PHYLOGENETIC RELATIONSHIPS OF GUIMARAESIELLA AND PRICEIELLA (PHTHIRAPTERA: ISCHNOCERA) FROM BABBLERS AND NON-BABBLERS (PASSERIFORMES).

Babblers (Passeriformes: Leiothrichidae, Pellorneidae, Timaliidae) are parasitized by more genera of lice of the Brueelia complex than any other group of songbirds. However, the relationships of these louse groups are poorly known. We here try to resolve the relationships between Guimaraesiella (Guimaraesiella), Guimaraesiella (Cicchinella), and their putative sister group Priceiella by using mitochondrial cytochrome c subunit 1 (COI), 12S, and 16S sequences. Our data indicate that G. (Cicchinella) forms a monophyletic group of lice from babblers, but the relationship between G. (Guimaraesiella), G. (Cicchinella), and Priceiella could not be resolved. Moreover, the position of the third lineage of babbler-specific lice, containing only the aberrant species Guimaraesiella montisodalis, is unresolved. Morphologically, this species is different from all other Guimaraesiella in several characters and may represent a distinct lineage. We present some data indicating that (1) the Nanling Mountain range may be a biogeographical barrier to chewing lice and (2) host participation in mixed-species feeding flocks may influence host associations in Brueelia-complex chewing lice.

NEW SPECIES AND NEW RECORDS OF PRICEIELLA (PHTHIRAPTERA: ISCHNOCERA: BRUEELIA-COMPLEX) FROM SOUTH CHINA.

Seven new species of chewing lice in the genus PriceiellaGustafsson and Bush, 2017, are described and illustrated based on specimens collected in south China. They are Priceiella (Camurnirmus) tanydrepanus n. sp. from Garrulax castanotis castanotis (Ogilvie-Grant, 1899); Priceiella (Camurnirmus) nanlingensis n. sp. from Garrulax maesi maesi (Oustalet, 1890); Priceiella (Thescelovora) brutifrons n. sp. from Turdinus brevicaudatus stevensi (Kinnear, 1925); Priceiella (Thescelovora) chuae n. sp. from Pellorneum albiventre cinnamomeus (Rippon, 1900); Priceiella (Thescelovora) catanachei n. sp. from Stachyris strialata swinhoei Rothschild, 1903; Priceiella (Thescelovora) dehongensis n. sp. from Stachyris nigriceps yunnanensis La Touche, 1921; and Priceiella (Thescelovora) rotundiceps n. sp. from Pomatorhinus ruficollis styani Seebohm, 1884. Several of these new species constitute the second species of Priceiella known from the host, suggesting that local endemism may be high in this louse genus. We also provide new host records for Priceiella (Thescelovora) coleyaeGustafsson et al., 2018, and Priceiella (Thescelovora) austiniGustafsson et al., 2018, and correct the type host subspecies for the latter. Finally, we amend the subgenus descriptions of CamurnirmusGustafsson and Bush, 2017, and ThescelovoraGustafsson and Bush, 2017. As a result of these changes, Priceiella najeriGustafsson et al., 2018, is moved to subgenus Thescelovora. An updated checklist of the genus Priceiella is provided.

Evolution of cryptic coloration in ectoparasites.

Cryptic coloration is a classic example of evolution by natural selection. However, it has been studied almost exclusively in predator-prey systems, despite the fact that it may evolve in other groups, such as ectoparasites. The principle defense of hosts against ectoparasites is grooming behavior, which has a visual component. Host-imposed selection should lead to the evolution of background matching if it helps ectoparasites escape from grooming. Here we use sister taxa comparisons to show that avian feather lice (Phthiraptera: Ischnocera) have evolved coloration that matches the host's plumage, except in the case of head lice, which are protected from grooming. We also show covariation of parasite and host color within a single species of louse. Thus, cryptic coloration has evolved both within and between species of feather lice. Other examples of the evolution of crypsis presumably exist among the 70,000 known species of ectoparasites that collectively represent five animal phyla.

Five New Species of Guimaraesiella (Phthiraptera: Ischnocera) from Broadbills (Aves: Passeriformes: Calyptomenidae: Eurylaimidae).

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 laoensis Meyer de Schauensee, 1929 ; Guimaraesiella latirostris n. sp. from Eurylaimus ochromalus Raffles, 1822 ; Guimaraesiella cyanophoba n. sp. from Cymbirhynchus macrorhynchus malaccensis Salvadori, 1874 and C. m. siamensis Meyer de Schauensee and Ripley, 1940 ; Guimaraesiella altunai n. sp. from Calyptomena viridis caudacuta Swainson, 1838 ; and Guimaraesiella forcipata n. sp. from Eurylaimus steerii steerii Sharpe, 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.

New Genus and Two New Species of Chewing Lice (Phthiraptera: Ischnocera) Parasitizing New Guinean Peltops (Passeriformes: Artamidae).

The genus Sarahcultrix n. gen. (Phthiraptera: Ischnocera) is described and illustrated based on 2 new species of chewing lice from New Guinean birds in the genus Peltops Wagler, 1829 (Passeriformes: Artamidae). These species are: Sarahcultrix ypsilophora n. sp. ex Peltops montanus Stresemann, 1921, and Sarahcultrix sphenura n. sp. ex Peltops blainvillii (Garnot, 1827).

The Genus Brueelia (Phthiraptera: Ischnocera: Philopteridae) of North American Jays and Allies (Aves: Passeriformes: Corvidae), with Descriptions of Five New Species.

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

Brueelia (Phthiraptera: Ischnocera: Philopteridae) of North American Nine-Primaried Oscines (Aves: Passeriformes: Passerida) with Descriptions of Nine New Species.

Nine new species of chewing lice in the genus Brueelia Kéler, 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.

Ten new species of Brueelia Kéler, 1936 (Phthiraptera: Ischnocera: Philopteridae) from nuthatches (Aves: Passeriformes: Sittidae), tits and chickadees (Paridae), and goldcrests (Regulidae).

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.

The chewing lice (Phthiraptera: Ischnocera, Amblycera) of the great cormorant (Phalacrocorax carbo).

The Great Cormorant is a widespread bird species with almost worldwide distribution. Accordingly, its general biology has been investigated thoroughly. Less well known, however, are the chewing lice that live inside the plumage of this diving bird. We examined the two known species of Great Cormorant chewing lice, Eidmanniella pellucida (Rudow, 1869) (Amblycera: Menoponidae) and Pectinopygus gyricornis (Denny, 1842) (Ischnocera: Philopteridae). Taking advantage of the autofluorescence of the cuticle, confocal laser scanning microscopy (CLSM) was used to explore the external morphology of all developmental stages of P. gyricornis. Morphometric analyses revealed a standard increase in body size from first larval instar to the adult. In addition, all instars exhibited increasing body segment differentiation, especially in the abdomen and the head. A total of 277 individuals of Pectinopygus gyricornis and 2 individuals of Eidmanniella pellucida were collected from eleven Great Cormorants from Mecklenburg-Western Pomerania, Germany, in 2015.

Two new species of Philopteroides (Phthiraptera: Ischnocera: Philopteridae) of the beckeri species-group, from New Guinean painted berrypeckers (Aves: Passeriformes: Paramythiidae).

Two new species of the louse genus Philopteroides Mey, 2004 are described and illustrated from two host species of painted berrypeckers, endemic to New Guinea: Philopteroides sinancorellus n. sp. ex Oreocharis arfaki (Meyer, 1875) (tit berrypecker), and Philopteroides gigas n. sp. ex Paramythia montium De Vis, 1892 (crested berrypecker). Both louse species belong to the beckeri species-group based on their short, broad preantennal areas, and shallow median indentations of the hyaline margin. The description of these two new species brings the total number of Philopteroides species to 15. An amended key to the beckeri species-group is included.

Four new species of Brueelia Kéler, 1936 (Phthiraptera: Ischnocera: Philopteridae) from African songbirds (Passeriformes: Sturnidae and Laniidae).

Four new species in the louse genus Brueelia Kéler, 1936 are described from African hosts of the families Sturnidae and Laniidae. They are: Brueelia rigbyi n. sp. ex Corvinella melanoleuca (Jardine, 1831), B. clara n. sp. ex Lamprotornis australis (A. Smith, 1836), B. tkachi n. sp. ex Spreo albicapillus albicapillus Blyth, 1856, and B. coryliventer n. sp. ex Creatophora cinerea (Meuschen, 1787). These four species are characterized by a unique abdominal chaetotaxy and a sinuous thickening of the distal margin of the male mesosome. These four species are very similar, and we consider them to form one species group. A key to the species of the group is provided.