Search

We report two host-specific lineages of scab mites of the genus Caparinia, parasitizing European and African hedgehogs. Based on morphology, these mite lineages are closely related sister groups. The morphological differences, however, are subtle and do not provide clear-cut evidence for the existence of separate species. CO1 divergence between these lineages was 7.4-7.8%, well above the CO1 barcoding gaps or thresholds commonly used to separate species, whereas divergence of five nuclear genes was very low, 0.06-0.53%, suggesting that these lineages could belong to a single species with gene flow between them. Thus, there is a conflict between the mitochondrial (CO1) gene and nuclear genes (i.e mito-nuclear discordance). We attribute this conflict to the 'gray zone' where species delimitation is ambiguous due to substantial gene flow. We also report another 'gray zone' species, Psoroptes ovis (a species of veterinary importance), whose within-species CO1 distances reached 6.0%. We provide a detailed morphological description and figures of C. ictonyctis stat. res. from the African hedgehog, using light and SEM microscopy and give morphometric data for this species and its sister species, Caparinia tripilis from Europe. For all known species of Caparinia, we document their host associations and give a key to species of the world based on results of our morphological and molecular analyses and a nearly exhaustive study of museum specimens.

Background: The cox1-barcoding approach is currently extensively used for high-throughput species delimitation and discovery. However, this method has several limitations, particularly when organisms have large effective population sizes. Paradoxically, most common, abundant, and widely distributed species may be misclassified by this technique., Results: We conducted species delimitation analyses for two host-specific lineages of scab mites of the genus Caparinia, having small population sizes. Cox1 divergence between these lineages was high (7.4-7.8%) while that of nuclear genes was low (0.06-0.53%). This system was contrasted with the medically important American house dust mite, Dermatophagoides farinae, a globally distributed species with very large population size. This species has two distinct, sympatric cox1 lineages with 4.2% divergence. We tested several species delimitation algorithms PTP, GMYC, ABGD, BPP, STACEY and PHRAPL, which inferred different species boundaries for these entities. Notably, STACEY recovered the Caparinia lineages as two species and D. farinae as a single species. BPP agreed with these results when the prior on ancestral effective population sizes was set to expected values, although delimitation of Caparinia was still equivocal. No other cox1 species delimitation algorithms inferred D. farinae as a single species, despite the fact that the nuclear CPW2 gene shows some evidence for introgression between the cox1 groups. This indicates that the cox1-barcoding approach may result in excessive species splitting., Conclusions: Our research highlights the importance of using nuclear genes and demographic characteristics to infer species boundaries rather than relying on a single-gene barcoding approach, particularly for putative species having large effective population sizes.

The fur mite genus Dubininetta Fain & Lukoschus, 1978 (Acariformes: Listrophoridae) represented by ectoparasites of moles (Soricomorpha: Talpidae) is revised. An amended diagnosis of this genus is provided. Two new species are described from moles in Vietnam: Dubininetta euroscaptor n. sp. from Euroscaptor parvidens (Miller) and Dubininetta abramovi n. sp. from Euroscaptor subanura Kawada, Son & Can. The insufficiently described D. taiwanensis (Fain & Lukoschus, 1978) from Mogera insularis (Swinhoe) (type-host) is redescribed based on the type-specimens and numerous newly collected in the Russian Far East specimens from Mogera robusta Nehring. The host ranges are considered for all species of Dubininetta. A key to the five species presently known in this genus is provided.

Eleven species of myocoptid mites (Acariformes: Myocoptidae) are recorded from African rodents. Among them, three species are described as new for science: Myocoptes lophuromys sp. nov. from Lophuromys woosnami Thomas, 1906 (Rodentia: Muridae) from Uganda, Trichoecius hylomyscus sp. nov. from Hylomyscus anselli Bishop, 1979 (Rodentia: Muridae) from Tanzania (type host and locality) and Hylomyscus stella (Thomas, 1911) from Gabon, and Trichoecius faini sp. nov. from Hylomyscus arcimontanus Carlton and Stanley, 2005 from Tanzania. Gliricoptes graphiuri Fain, 1970 syn. nov. is synonymized with Gliricoptes vulcanorum Fain, 1970, and the female of Trichoecius angolensis Fain, 1972 is described for the first time. New hosts and localities are provided for four species: Myocoptes grammomys Fain, 1970 from Grammomys dryas (L., 1758) (Rodentia: Muridae) from Burundi and Uganda, Myocoptes spinulatus Fain, 1970 from Dendromus mystacalis (Heuglin, 1863) (Rodentia: Muridae) from Tanzania, Crinicastor congolensis Fain, 1970 from Grammomys macmillani (Wroughton, 1907) (Rodentia: Muridae) from Tanzania, and Trichoecius otomys Fain, 1970 from Otomys typus (Heuglin, 1877) (Rodentia: Muridae) from Uganda. Three species are recorded from new localities: Gliricoptes vulcanorum from Tanzania, Trichoecius angolensis from South Africa, and Trichoecius lootensi Fain, 1970 from Tanzania., (© The Authors 2016. Published by Oxford University Press on behalf of Entomological Society of America. All rights reserved. For Permissions, please email: journals.permissions@oup.com Version of Record, first published online December 16, 2016 with fixed content and layout in compliance with Art. 8.1.3.2 ICZN.)

A new species of parasitic mites, Myocoptes (Comyocoptes) nesokia sp. nov. (Acariformes: Myocoptidae) is described from Nesokia indica (Gray, 1830) (Rodentia: Muridae) from Pakistan. Females of this species differ from the morphologically similar species, Myocoptes (Comyocoptes) striatus Fain, 1970, by the presence of a verrucose pattern in the middle part of the hysteronotum, longer setae cp (120-140 μm vs. 55-60 μm), and by the absence of scales on the opisthosomal cuticle posterior to the level of setal bases ps3.

Mites of the family Myocoptidae (Acariformes: Sarcoptoidea) of the former USSR are revised based on the collection of the Zoological Institute RAS (St. Petersburg, Russia). Seventeen described species are recorded. Four species are described as new for science: Trichoecius dubininae sp. nov. from Lasiopodomys gregalis (Pallas) (Rodentia: Cricetidae) from Russia (Chelyabinsk Prov., type locality), Kazakhstan and Kirghizia, T. lemmus sp. nov. from Lemmus sibiricus (Kerr) (Rodentia: Cricetidae) from Russia (Taymyr peninsula), T. meriones sp. nov. from Meriones meridianus (Pallas) (Rodentia: Muridae) from Turkmenia, and Myocoptes meriones sp. nov. from Meriones tamariscinus (type host) from Kirghizia (type locality) and Meriones libycus (Lichtenstein) from Tajikistan. In addition, the presence of 7 more species in the fauna of the former USSR is considered highly probable. Myocoptids of the former USSR represent all 6 genera currently recognized in the family. Keys to species, their hosts and localities are provided and all data are summarized in tabular format. A new name Apocalyptoides nom. nov. is proposed for the preoccupied generic name Apocalypsis Bochkov, 2010 not Butler (1876).

Host associations of permanent ectoparasitic mite Marsupialges misonnei Fain, 1963 (Acariformes: Psoroptidae: Marsupialginae) are analyzed. This species was first recorded from an ethanol-preserved museum specimen of Caluromys philander (Linnaeus, 1758) (Didelphimorphia: Didelphidae) originating from French Guiana. We discovered specimens of M. misonnei from both species known in the carnivore genus Nasua (Carnivora: Procyonidae): N. narica (Linnaeus, 1766) from Panama (collected in the field) and N. nasua (Linnaeus, 1766) from Brazil (collected from dry museum specimen). Two alternative hypotheses about an initial host of this mite (bare-tailed woody opossum or coatis) are discussed. We argue that M. misonnei was originally parasitic on Nasua spp. and occasionally contaminated C. philander from these hosts in the collecting process.

A new species Crocidurobia faini n. sp. (Acariformes: Myobiidae) recorded on Crocidura denti Dollman (type-host) (Soricomorpha: Soricidae), Crocidura cf. niobe Thomas, Crocidura cf. littoralis Heller, and Crocidura sp. from DR Congo is described. This new species differs in both sexes from all other species of the subgenus Crocidurobia Jameson, 1970 by short setae c2, more than three times shorter than setae se. Females of the new species differ from those of the close species Crocidurobia toroensis Dusbabek, 1983 by setal bases e2 situated slightly anterior to the level of setal bases e1; males of the new species differ by the genital shield bearing five pairs of setae situated anterior to the level of setal bases d1.

Based on multilocus phylogenetic analyses (18S, 28S, EF1-α, SRP54, HSP70, CO1, 10 860 nt aligned), we show that the house dust mite subfamily Guatemalichinae is nested within non-onychalgine pyroglyphid mites and forms the sister group to the genus Sturnophagoides (bootstrap support 100, posterior probability 1.0). Because high bootstrap support values may be misleading in the presence of incongruence, we evaluate robustness of the Guatemalichinae+Sturnophagoides clade using: (1) internode certainty indices to estimate the frequency of conflicting bipartitions in maximum-likelihood bootstrap trees, (ii) consensus networks to investigate conflict among different loci; and (iii) statistical hypothesis testing based on information theory, both multi-scale and regular bootstrap. Results suggest that this grouping is very well supported given the data. The molecular analyses were integrated with detailed morphological study using scanning electron and light microscopy. We suggest that the subfamilial status of Guatemalichinae should be reconsidered, and this lineage should be placed within the subfamily Dermatophagoidinae. The latter subfamily is currently accepted in the literature as a monophyletic group but was here inferred as paraphyletic and was not supported by any morphological synapomorphy. The paraphyly involved the most species-rich and medically important genus, Dermatophagoides. Our findings suggest the need for a comprehensive revision of the higher-level relationships of pyroglyphid house dust mites using both DNA sequences and morphology coupled with a broad taxonomic sampling., (© The Willi Hennig Society 2015.)

The fur-mite genus Soricilichus Fain, 1970 (3 species) (Acariformes: Chirodiscidae) represented by permanent symbionts of the African shrews of the subfamily Crocidurinae (Soricomorpha: Soricidae) is revised. The external morphology of these species was investigated with light and scanning electron microscopy. Based on the type specimens and newly obtained samples, 2 recognized species, S. scutisorex Fain, 1970 from Scutisorex somereni Thomas and S. kivuensis Fain, 1981 from Crocidura sp.-both are from DR of Congo-are redescribed. A new species S. sylvisorex sp. nov. found on shrews of the genera Sylvisorex (S. granti Thomas (type host), S. lunaris Thomas, S. vulcanorum Hutterer and Verheyen) and Crocidura (C. denti Dollman, C. cf. niobe, Crocidura sp.), collected in the DR Congo is also described. An amended generic diagnosis, including description of female immature stages, and a key to species are provided.

Sixteen species of the superfamily Sarcoptoidea (Acariformes: Psoroptidia) belonging to 10 genera of the families Atopomelidae, Listrophoridae, Chirodiscidae, and Listropsoralgidae are recorded in Brazil. Among them, three species, Prolistrophorus hylaeamys sp. nov. from Hylaeamys laticeps (Lund, 1840) (Cricetidae: Sigmodontinae) from Minas Gerais, Lynxacarus serrafreirei sp. nov. from Galictis cuja (Molina, 1782) (Carnivora: Mustelidae) from Rio de Janeiro (Listrophoridae), and Didelphoecius micoureus sp. nov. (Atopomelidae) from Micoureus paraguayanus (Tate, 1931) (Didelphimorphia: Didelphidae) from Minas Gerais are described as new for science. Three species of the family Listrophoridae, Prolistrophorus bidentatus Fain et Lukoschus, 1984 from Akodon cursor (Winge, 1887) (Rodentia: Cricetidae) (new host), Prolistrophorus ctenomys Fain, 1970 from Ctenomys torquatus Lichtenstein, 1830 (Rodentia: Ctenomyidae) (new host), and Leporacarus sylvilagi Fain, Whitaker et Lukoschus, 1981 from Sylvilagus brasiliensis (Linnaeus, 1758) (Lagomorpha: Leporidae) (new host) -from Minas Gerais and Rio Grande do Sul, and one species of the family Chirodiscidae, Parakosa tadarida McDaniel and Lawrence, 1962 from Molossus molossus (Pallas, 1766) (Chiroptera: Molossidae) are recorded for the first time in Brazil. The previously unknown female of Didelphoecius validus Fain, Zanatta-Coutinho et Fonseca, 1996 (Atopomelidae) from Metachirus nudicaudatus (Geoffroy, 1803) (Didelphimorphia: Didelphidae) from Minas Gerais is described. All data on host-parasite associations of sarcoptoids in Brazil are summarized. Totally, 61 sarcoptoid species of 8 families are recorded in Brazil.

Mites of the subfamily Harpirhynchinae (Acariformes: Cheyletoidea: Harpirhynchidae) associated with neognathous birds (Aves: Neognathae) in the New World are revised. In all, 68 species in 8 genera are recorded. Among them, 27 new species and 1 new genus are described as new for science: Harpyrhynchoides gallowayi Bochkov, OConnor and Klompen sp. nov. from Columba livia (Columbiformes: Columbidae) from Canada (Manitoba), H. zenaida Bochkov, OConnor and Klompen sp. nov. from Zenaida macroura (Columbiformes: Columbidae) from USA (Michigan), H. calidris Bochkov, OConnor and Klompen sp. nov. from Calidris minutilla (Charadriiformes: Scolopacidae) from USA (Kansas), H. actitis Bochkov, OConnor and Klompen sp. nov. from Actitis macularius (Charadriiformes: Scolopacidae) from Canada (British Columbia), H. charadrius Bochkov, OConnor and Klompen sp. nov. from Charadrius vociferus (Charadriiformes: Charadriidae) from USA (Texas), H. pluvialis Bochkov, OConnor and Klompen sp. nov. from Pluvialis dominica (Charadriiformes: Charadriidae) from USA (Ohio), H. bubulcus Bochkov, OConnor and Klompen sp. nov. from Bubulcus ibis (Pelecaniformes: Ardeidae) from USA (Florida), H. ixobrychus Bochkov, OConnor and Klompen sp. nov. from Ixobrychus exilis (Pelecaniformes: Ardeidae) from USA (Michigan), H. puffinus Mertins sp. nov. from Puffinus gravis (Procellariformes: Procellariidae) from USA (Florida), H. megascops Bochkov, OConnor and Klompen sp. nov. from Megascops asio (Strigiformes: Strigidae) from USA (Michigan), H. athene Bochkov, OConnor and Klompen sp. nov. from Athene canicularia (Strigiformes: Strigidae) from USA (Texas), H. coccyzus Bochkov, OConnor and Klompen sp. nov. from Coccyzus americanus (Cuculiformes: Cuculidae) from USA (Michigan), H. crotophaga Bochkov, OConnor and Klompen sp. nov. from Crotophaga ani (Cuculiformes: Cuculidae) from Suriname; Crassacarus Bochkov, OConnor and Klompen, gen. nov.: Crassacarus alexfaini Bochkov, OConnor and Klompen sp. nov. (type of genus) from Cardinalis cardinalis (type host) from USA (Michigan), Passerina ciris (unknown locality in North America) (Passeriformes: Cardinalidae), and Setophaga petechia (Passeriformes: Parulidae) from USA (Michigan), C. tinae Bochkov, OConnor and Klompen sp. nov. from Carduelis tristis (Passeriformes: Fringillidae) from USA (Wyoming), C. fritschi Bochkov, OConnor and Klompen sp. nov. from Bombycilla cedrorum (Passeriformes: Bombycillidae) from USA (Michigan), C. sialia Bochkov, OConnor and Klompen sp. nov. from Sialia currucoides (Passeriformes: Turdidae) from USA (Wyoming), C. melanerpes Bochkov, OConnor and Klompen sp. nov. from Melanerpes formicivorus (Piciformes: Picidae) from USA (Kansas); Neharpyrhynchus turdus Bochkov, OConnor and Klompen sp. nov. from Turdus migratorius (Passeriformes: Turdidae) from USA (Michigan), N. campylorhynchus Bochkov, OConnor and Klompen sp. nov. from Campylorhynchus brunneicapillus (Passeriformes: Troglodytidae) from USA (unknown locality), N. spizella Bochkov, OConnor and Klompen sp. nov. from Spizella passerina (Passeriformes: Emberizidae) from USA (various localities), N. quiscalus Bochkov, OConnor and Klompen sp. nov. from Quiscalus quiscula (Passeriformes: Icteridae) from USA (Michigan), N. agelaius Bochkov, OConnor and Klompen sp. nov. from Agelaius phoeniceus (Passeriformes: Icteridae) from USA (Michigan), N. bombycilla Bochkov, OConnor and Klompen sp. nov. Bombycilla cedrorum (Passeriformes: Bombycillidae) from USA (Michigan), N. vireo Bochkov, OConnor and Klompen sp. nov. from Vireo olivaceus (Passeriformes: Vireonidae) from USA (Florida), N. picidarum Bochkov, OConnor and Klompen sp. nov. from Colaptes auratus (type host), Melanerpes formicivorus, Melanerpes uropygidialis, and Picoides pubescens (Piciformes: Picidae) from USA (various localities); Perharpyrhynchus charadrius Bochkov, OConnor and Klompen sp. nov. from Charadrius vociferus (Charadriiformes: Charadriidae) from USA (Michigan). Harpyrhynchoides oenae lamorali (Fain, 1972) syn. nov. is synonymized with Harpyrhynchoides oenae (Fain, 1972). Harpirhynchoides agapornis (Fain, 1972) comb. nov. and Crassacarus cylindripalpus (Fritsch, 1954) comb. nov. are transferred from the subgenus Pseudoharpirhynchus Fain, Bochkov and Mironov, 1999 (type species Harpirhynchus agapornis Fain, 1972) of the genus Harpirhynchus Megnin 1877. The subgenus Pseudoharpirhynchus syn. nov. is synonymized with the genus Harpyrhynchoides. Diagnoses for the subfamily and all genera recorded in the New Word (Anharpyrhynchus Fain, 1972, Crassacarus gen. nov., Harpirhynchus, Harpyrhynchoides Fain, 1972, Fainharpirhynchus Bochkov and Galloway, 2013, Neharpyrhynchus Fain, 1972, Perharpyrhynchus Fain, 1972, Trichorhynchiella Fain, 1995) are provided. Keys to all harpirhynchine genera and all their species occurring in the New World are also given. A list of all harpirhynchine species and their hosts is compiled.

Two new mite species of the subfamily Harpirhynchinae Dubinin, 1957 (Acariformes: Harpirhynchidae) are described from passerine birds (Aves: Passeriformes): Harpirhynchoides artamus n. sp. from Artamus fuscus Vieillot (Artamidae) from an unknown locality in South Asia and Neharpyrhynchus domrowi n. sp. from three host species of the family Meliphagidae, Acanthorhynchus tenuirostris (Latham) (type-host) from Australia (New South Walles), Ptiloprora perstriata (De Vis) and Myzomela rosenbergii Schlegel from Papua New Guinea.

Tachornithoglyphus gen. nov. (Acariformes: Pyroglyphidae) is established for Tachornithoglyphus tachornis (Cruz, Cuervo and Dusbabek, 1984), comb. nov. (transferred from Guatemalichus), collected from nests of the Antillean palm swift, Tachornis phoenicobia (Apodiformes: Apodidae) in Cuba. The new genus differs from the other four genera of the subfamily Guatemalichinae, Guatemalichus Fain and Wharton, 1970, Pottocola Fain, 1971, Fainoglyphus Atyeo and Gaud, 1977, and Capitonocoptes Fain and Gaud, 1984, mainly by having the length of solenidion σ1 of genu I less than one-third that of the segment (vs. solenidion σ1I at least half as long as genu I), by the absence of famulus ε on tarsus I (vs. present), and by coxal apodemes Ia separated from each other and contiguous to the lateral parts of the epigynal arch (vs. posterior tips of apodemes Ia fused to each other and with the median part of the epigynal arch, or separated and contiguous to or fused with the median part of the epigynal arch). A detailed redescription of adults and tritonymphs of T. tachornis is provided.

Carcinopodacarus polymorphus gen. n. et sp. n. (Acariformes: Dermationidae: Dermationinae) is described from the guira cuckoo Guira guira (Gmelin) (Cuculiformes: Cuculidae) in Brazil. The new genus differs from the closest genus, Psittophagoides Fain, 1964, by the following features: in both sexes, the anterior spines of trochanters I and II are absent (vs present in Psittophagoides), setae d2 are distinctly developed (vs only alveoli), and genual setae mGI are absent (vs present); in males, the hysteronotal shield is split transversally at the level of trochanters III (vs hysteronotal shield entire); in females, the platelets situated posterior to the propodonotal shield are absent (vs present), the metapodosomal sclerites are present (vs absent), and the adanal shields are fused anteriorly to each other (vs separated from each other). In this species, andropolymorphism is detected for the first time for the family. It involves various characters but the most impressive feature is the structure of legs III. In hetero- and mesomorphic males, these legs are strongly hypertrophied and have a distinct ventral spur on femora III; in homeomorphic males, legs III are not modified and subequal to legs IV.

A new species of the genus Lemuralges Fain, 1963 (Acariformes: Psoroptidae: Makialginae) is described from the Malagasy lemur Propithecus diadema (Bennett) (Primates: Indriidae) based on all postembryonic instars. This new species differs from the only known species in this genus, Lemuralges intermedius Fain, 1963, by the following features: both sexes of L. propithecus sp. n. show a pair of medioventral projections of the subcapitulum (vs without projections in L. intermedius) and the propodonotal shield is slightly ornamented (vs unornamented); in males the hysteronotal shield is completely covered by longitudinal striae (vs median part without striae), setae c2 are 120-140 µm long (vs 200-210 µm long), and femur III has a short transverse furrow dorsally (vs a longitudinal furrow); in females, setae h2 are, at least, 2 times shorter than h3 (vs slightly longer, or subequal to, h3), tibia IV has a ventro-apical projection (vs without projection). Larvae and protonymphs of the new species show some unique developmental delays. Female and male tritonymphs differ by their external morphology.

Two new species of parasitic acariform mites (Acariformes) are described from the Delacour's marmoset rat Hapalomysdelacouri Thomas (Rodentia: Muridae) in Vietnam: Afrolistrophorus (Afrolistrophorus) hapalomyssp. n. (Listrophoridae) and Radfordia (Radfordia) mirabilissp. n. (Myobiidae). Based on morphological evidences, we show that species of both mite genera associated with Hapalomys Blyth do not demonstrate clear phylogenetic links with respective congeners from rodents of the closest genus Chiropodomys Peters (Rodentia: Muridae).

Two new species of Paracoroptes Lavoipierre, 1955 (Acariformes: Psoroptidae: Paracoroptinae) are described: Paracoroptes miopithecus sp. n. from Miopithecus talapoin (Schreber) and Paracoroptes piliocolobus sp. n. from Piliocolobus badius (Kerr) (Primates: Cercopithecidae)--both are from the Democratic Republic of the Congo. A key to all 6 species of the genus is provided and host-parasite relationships of its representatives are discussed.

The zumpti species group of the genus Harpyrhynchoides (Harpirhynchidae), parasites of passerines, is revised. A key to the species of this group is provided and data on host associations and geographic distribution of its constituent species are summarized. This group includes six previously recognized species: Harpyrhynchoides alaudinus Bochkov, 2000, H. brevis (Ewing, 1911) comb. nov., H. heatherae Bochkov and Galloway, 2013, H. rubeculinus (Cherny and Sixl, 1971), H. vulgaris Bochkov and Galloway, 2004, and H. zumpti (Fain, 1972). Three species from North American passerines are described as new: H. setophaga sp. nov. from Setophaga ruticilla (Parulidae), H. xanthocephalus sp. nov. from Xanthocephalus xanthocephalus (Icteridae), and H. spizella sp. nov. from Spizella passerina (Emberizidae). Additionally, H. brevis is redescribed based on samples from Coccothraustes vespertinus (type host) and Loxia curvirostra (Passeriformes: Fringillidae) from North America. Harpyrhynchoides kirgizorum Fain et al. 1999 syn. nov. is synonymized with H. zumpti.

Four native species of parasitic mites belonging to the genus Schizocarpus Trouessart, 1896 (Acariformes: Chirodiscidae) are recorded on the North American beaver Castor canadensis Kuhl, 1820 (Rodentia: Castoridae) from Russia. Totally ten beavers from all three main geographically isolated populations of in Russia (Leningrad Province, Voronezh Biosphere Reserve (beaver farm) and Khabarovsk Territory) were examined. Additionally, in captivity (Voronezh beaver farm) eight species were recorded switched from the Eurasian beaver Castor fiber Linnaeus, 1758 on C. canadensis.

A new species of the genus Neocheyletiella Baker, 1949 (Acariformes: Cheyletidae) is described from the zebra finch, Taeniopygia guttata (Viellot, 1817) (Passeriformes: Estrildidae), from a laboratory colony at the Massachusetts Institute of Technology, Cambridge, MA. The setal additions in the ontogeny of the new species, Neocheyletiella parvisetosa Mertins & Bochkov, and the main differential characters of all 17 known species of the genus Neocheyletiella are provided in tabular format. Keys to females and males of Neocheyletiella spp. also are given., (© 2014 Entomological Society of America.)

A new subgenus Neotomobia n. subg. (Acariformes: Myobiidae: Radfordia) is established for species parasitising rodents of the subfamily Neotominae (Rodentia: Cricetidae): Radfordia subuliger Ewing, 1938 (type-species), Radfordia eremici Fain & Bochkov, 2002, Radfordia neotomae Jameson & Whitaker, 1975 and Radfordia hamiltoni Jameson & Whitaker, 1975. Three new species are described: Radfordia peromyscus n. sp. from Peromyscus megalops Merriam from Mexico, Radfordia onychomys n. sp. from Onychomys leucogaster (Wied-Neuwied) from the USA and Radfordia megadontomys n. sp. from Megadontomys thomasi (Merriam) from Mexico.

Six fur-mite species of the genus Prolistrophorus Fain, 1970 (Acariformes: Listrophoridae) were recorded from Central and South American rodents of the subfamily Sigmodontinae (Rodentia: Cricetidae). Among them, Prolistrophorus (Aprolistrophorus) parabidentatus sp. nov. from Akodon azarae from Argentina and Prolistrophorus (Aprolistrophorus) tylomys sp. nov. from Tylomys nudicaudus from Guatemala are described as new for science. New hosts are recorded for the following species: Prolistrophorus (Prolistrophorus) grassii (Radford, 1954) from Zygodontomys brevicauda from Colombia, P. (P.) frontalis (Hirst, 1921) from Oligoryzomys sp. from Argentina, P. (P.) argentinus (Hirst, 1921) from Melanomys caliginosus, Akodon affinis from Colombia and Scapteromys aquaticus from Argentina, Prolistrophorus (Beprolistrophorus) hirstianus Fain, 1973 from Scapteromys aquaticus from Argentina.

Seven new species of intracutaneous parasites of the subfamily Harpirhynchinae Dubinin (Acariformes: Cheyletoidea: Harpirhynchidae) are described: Anharpyrhynchus elizae sp. nov. from Cyanocitta cristata (type host), Cyanocitta stelleri (Passeriformes: Corvidae), and Colaptes auratus (Piciformes: Picidae) from the USA; Anharpyrhynchus apodus sp. nov. from Lichmera indistincta (type host) and Phylidonyris novaehollandiae (Passeriformes: Meliphagidae) from Australia; Anharpyrhynchus lukoschusi sp. nov. from Manorina flavigula (type host) and Certhionyx pectoralis (Passeriformes: Meliphagidae) from Australia; Fainharpirhynchus legatus sp. nov. from Legatus leucophaius (Passeriformes: Tyrannidae) from Trinidad and Tobago; Fainharpirhynchus mossi sp. nov. from Myiarchus crinitus (Passeriformes: Tyrannidae) from an unknown locality in North America; Perharpyrhynchus caprimulgus sp. nov. from Caprimulgus fossii (Caprimulgidae: Caprimulgiformes) from Mozambique; Perharpyrhynchus elseyornis sp. nov. from Elseyornis melanops (Charadriiformes: Charadriidae) from Australia. Keys to females of the genera Anharpyrhynchus, Fainharpirhynchus, and Perharpyrhynchus are provided and data on hosts and distributions of all known intracutaneous harpirhynchines are summarized. In addition, a new non-subcutaneous harpirhynchine is described: Trichorhynchiella myiarchus sp. nov. from feather bases of Myiarchus crinitus (Passeriformes: Tyrannidae) from an unknown locality in North America.

The subfamily Harpypalpinae Fain, 1972 (Acariformes: Cheyletoidea: Harpirhynchidae) is revised. Diagnoses of the subfamily and its two constituent genera, Harpypalpus Dubinin, 1957 and Harpypalpoides Lombert and Moss, 1979, and keys to females of all known species are provided. Data on harpypalpine hosts and distribution are summarized, and nine new species are described: Harpypalpus lonchura sp. nov. from Lonchura castaneothorax (Gould) (Estrildidae) in Australia, Harpypalpus pyrrhula sp. nov. from Pyrrhula pyrrhula (Linnaeus) (Fringillidae) in the Netherlands, Harpypalpus sturnus sp. nov. from Sturnus vulgaris Linnaeus (Sturnidae) in the Netherlands, Harpypalpus taeniopygia sp. nov. from Taeniopygia guttata (Vieillot) (Estrildidae) in Australia, Harpypalpoides hirundinoides sp. nov. from Hirundo rustica Linnaeus (Hirundinidae) in the Netherlands, Harpypalpoides johnstoni sp. nov. from Fringilla coelebs Linnaeus (Fringillidae) (type host) in the Netherlands and Fringilla montifringilla Linnaeus in Kazakhstan, Harpypalpoides regulus sp. nov. from Regulus regulus (Linnaeus) (Regulidae) in the Netherlands, Harpypalpoides sitta sp. nov. from Sitta pygmaea Vigors (Sittidae) in the USA (Arizona, California), and Harpypalpoides sylvia sp. nov. from Sylvia atricapilla (Linnaeus) (Sylviidae) in the Netherlands.

The external morphology of adult and immature stages of mange mites of the genus Chorioptes was investigated with the aid of light and scanning electron microscopy. A molecular phylogeny of this genus was inferred based on six genes (18S, 28S rDNA, EF1-α, SRP54, HSP70, and CO1). The validity of four species (Ch. bovis, Ch. panda, Ch. texanus, and Ch. sweatmani sp. nov. described from the moose from Sweden, Finland, and Russia) was confirmed based on morphology and a Bayesian species delimitation analysis incorporating both gene tree uncertainties and incomplete lineage sorting via the coalescent process model in BPP. Sequence data for Ch. crewei and Ch. mydaus was not available but their morphology strongly suggests their validity. The six valid Chorioptes species are diagnosed using type and non-type specimens, and a key to species is provided. Ch. sweatmani differs from closely related Ch. texanus by the following features: in males, the body length, including the gnathosoma, is 380-405 μm (vs. 220-295 in Ch. texanus), the idiosoma is 3-4 times longer than setae cp (vs. 1.3-1.6 times longer), legs III are approximately three times longer than setae sRIII (vs. 1.8-2 times longer), the apical spur of tarsus III is curved (vs. straight), a spur near seta fIII base is not developed (vs. small but distinct); in females, setae h2 are 1.4-1.5 times shorter than legs IV (vs. about two times longer). Hosts and distribution records of Chorioptes species are summarized.

The subfamily Onychalginae Fain, 1988 (Acariformes: Pyroglyphidae) is a monophyletic lineage comprising the closest parasitic relatives of the free-living pyroglyphid house dust mites. Onychalgine mites parasitize passerine birds of the families Estrildidae, Passeridae, and Ploceidae in Africa and South America (a single record of Onychalges spinitarsis from a piciform bird requires confirmation). We revise this subfamily based on external morphology of adults and immature stages using light and scanning electron microscopy and give a key to species. Onychalginae includes 2 genera: Onychalges Gaud and Mouchet, 1959 (6 species) and Paramealia Gaud, 1968 (1 species). The genus Kivuicola Fain, 1971 syn. nov. is synonymized with Onychalges, and its single species K. kivuana Fain, 1971 syn. nov. is considered as a putative synonym of O. odonturus Gaud, 1968. Onychalges spinitarsis (Fain and Gaud, 1984) is considered as a species inquirenda.

The small genus Trichorhynchiella Fain, 1995, including two species of permanent bird ectoparasites belonging to the family Harpirhynchidae (Acariformes: Cheyletoidea), is revised. The generic diagnosis is emended. The type species, Trichorhynchiella paddae Fain, 1995 from Lonchura oryzivora (L, 1758) (Passeriformes: Estrildidae) from Southeast Asia and Trichorhynchiella phleocryptes sp. n. from Phleocryptes melanops (Viellot, 1817) (Passeriformes: Furnariidae) from Argentina are redescribed and described, respectively. The phenomenon of ovoviviparity inherent to females of this genus is briefly discussed. In almost immobile physogastric females of Trichorhynchiella, ovoviviparity eliminates necessity to attach eggs to the host body.

Five new species and one new genus of the subfamily Harpirhynchinae (Acariformes: Harpirhynchidae) are described from birds in Canada: Harpyrhynchoides heatherae sp. nov. from Junco hyemalis (Passeriformes: Emberizidae), H. botaurus sp. nov. from Botaurus lentiginosus (Pelecaniformes: Ardeidae), H. phalaropus sp. nov. from Phalaropus lobatus (Charadriiformes: Scolopacidae), Neharpyrhynchus loxia sp. nov. from Loxia curvirostra (Passeriformes: Fringillidae), and Fainharpirhynchus contopus gen. nov., sp. nov. from Contopus cooperi (Passeriformes: Tyrannidae). Additionally, 3 species were recorded in Canada (Manitoba) for the first time: Harpyrhynchoides tracheatus (Fritsch, 1954) from Buteo jamaicensis (Accipitriformes: Accipitridae) (new host), H. modestus (Fain, 1976) from Columba livia (Columbiformes: Columbidae) (new host), and Neharpyrhynchus pilirostris (Berlese et Trouessart, 1889) from Passer domesticus (Passeriformes: Passeridae).

Various data considered as possible cases where metazoans return form the obligate parasitism to free mode of life (a violation of ecological interpretation of Dollo's law). It is shown that among temporary and phase parasites their return to the free mode of life is a rare but not exceptional phenomenon. It is possibly to conclude that these cases are displayed only by those ectoparasitic arthropods, ancestors of which were predators. The only, although incompletely proven cases of the violation of Dollo's law among permanent parasites is suggested to be acariform mites of the family Pyroglyphidae (Acariformes: Psoroptoidea).

Three new harpirhynchid species of the subfamily Harpirhynchinae (Acariformes: Harpirhynchidae) are described from North American birds: Harpyrhynchoides aegolius sp. n. from Aegolius acadicus (Strigiformes: Strigidae), Harpyrhynchoides accipiter sp. n. from Accipiter striatus (Accipitriformes: Accipitridae), and Neharpyrhynchus icterus sp. n. from Icterus galbula (Passeriformes: Icteridae). Three species are recorded from new hosts from North America for the first time: Harpyrhynchoides tracheatus (Fritsch, 1954) from Buteo lineatus (Accipitriformes: Accipitridae), Harpyrhynchoides pectinifer (Lawrence, 1959) from Colaptes auratus (Piciformes: Picidae), and Harpyrhynchoides rubeculinus (Cerný & Sixl, 1971) from Catharus ustulatus (Passeriformes: Turdidae). Neharpyrhynchoides novoplumaris (Moss et al., 1968) previously recorded from Cardinalis cardinalis (Passeriformes: Cardinalidae) from the United States is recollected from this host.

The family Listropsoralgidae Fain, 1965 (Acariformes: Sarcoptoidea) is represented by the permanent skin ectoparasites associated with the South American and Australian marsupials (12 species) and the South American rodents of the family Echimyidae (1 species). The phylogenetic relationships of these mites (12 ingroup and 2 outgroup species) are reconstructed on the basis of the maximum parsimony (MP) and Bayesian analyses (BA) of 76 morphological characters. MP analysis confirmed monophyly of the listropsoralgid genera, the strict consensus of 18 trees generated by MP has the following pattern: Petauralges (Listropsoralgoides, Didelphialges, Listropsoralges) with poor resolution among species of the genus Listropsoralges. The same tree was generated by BA. Both successive and implied weighting strategies resulted in 7 MP trees: Petauralges (Listropsoralgoides (Didelphialges (Listropsoralges))). The relationships between species of the genus Listropsoralges received the poorest resolution: L. caenolestes (L. monodelphis, L. vossi, L. faini, L. brevisetosa (L. thylamys (L. marmosa-L. caluromys))). The host-parasite relationships of listropsoralgids are briefly discussed. The family Listropsoralgidae is taxonomically revised and to date includes 13 species in 4 genera. Six species and one genus are described as new: Listropsoralges brevisetosus sp. n. from Marmosa murina (Didelphimorphia: Didelphidae) from Peru, Listropsoralges similis sp. n. from Caluromys derbianus (Didelphidae) from Panama, Listropsoralges thylamys sp. n. from Thylamys venustus (Didelphidae) from Bolivia, Listropsoralges vossi sp. n. from Monodelphis domestica (Didelphidae) from Brazil, Listropsoralges caenolestes sp. n. from Caenolestes fuliginosus (Paucituberculata: Caenolestidae) from Ecuador, and Didelphialges metachirus gen. n., sp. n. from Metachirus nudicaudatus (Didelphidae) from Peru. The female of Listropsoralges faini Bochkov and Wauthy, 2009 is described for the first time.

Six species of 3 genera belonging to the fur mite family Listrophoridae were recorded on skins of South American rodents of the cricetid subfamily Sigmodontinae housed in the Bavarian State Collection of Zoology (Munich, Germany). Among them, Amlistrophorus geoxus sp. nov. from Geoxus valdivianus from Chile is described as a new for science, and males of Prolistrophorus amazonicus amazonicus Fain, 1971 are recorded for the first time. The full generic status for the subgenus Amlistrophorus of the genus Prolistrophorus proposed by Fain et al. (1996) is not supported, and Prolistrophorus musculinus Fain, 1973 stat. nov. (formerly a subspecies of P. amazonicus) from Mus musculus (Rodentia: Muridae) from Suriname is raised to species status. New hosts are recorded for the following species: Prolistrophorus argentinus (Hirst, 1921) from Holochilus brasiliensis and H. chacarius from Argentina, P. amazonicus from Calomys callosus from Argentina and Bolivia, C. laucha and C. musculinus from Argentina, P. akodon Fain and Lukoschus, 1982 from Akodon montensis from Argentina, P. nectomys Fain, 1971 from Nectomys palmipes from Peru and Melanomys caliginosus from Panama, and Sclerolistrophorus oxymycteris Fain, 1976 from Oryzomys laticeps from Brazil.

A new mange mite species, Macropodicoptes mironovi n. g., n. sp. (Acariformes: Sarcoptidae), is described from Wallabia bicolor (Desmarest) (Diprotodontia: Macropodidae) from the Taronga Western Plains Zoo at Dubbo, NSW, Australia. This is the fourth species and third genus allocated to the sarcoptid subfamily Diabolicoptinae, whose representatives exclusively parasitise marsupials. The new genus differs from Diabolicoptes Fain & Domrow, 1974, in both sexes, by the proportions of the gnathosoma, the shape of many hysterosomal setae and the absence of setae e1, pRII-II, sRIII and solenidion ω3I; and, in males, by the lack of fusion of coxal apodemes I.

Host-parasite relationships of mammals and astigmatan mites (Acariformes: Astigmata) belonging to the parvorder Psoroptidia are analyzed. The absolute majority of mammal-associated psoroptidians belongs to the paraphyletic superfamily Sarcoptoidea. Mites of the family complex Psoroptidae (Lobalgidae, Psoroptidae, and Paracoroptinae) shifted from birds to placental mammals independently from each other. Mites of the family complex Sarcoptidae, including all other sarcoptoid families, derived from the common stalk of Psoroptidia independently from the Psoroptid complex. Mites of the sarcoptid complex shifted from nidicoly in mammalian nests to the permanent parasitism on these hosts. They are widely represented on both marsupial and placental mammals and are absent on Monotremata.

Four new syringophilid species of Syringophiloidus Kethley, 1970 are described from North American passerines: S. zonotrichia n. sp. from Zonotrichia albicolis (Gmelin) (Emberizidae) on Texas; S. jackowiaki n. sp. from Poecile carolinensis (Auduborn) (Paridae) in Texas; and S. xanthocephalus n. sp. from Xanthocephalus xanthocephalus (Bonaparte) (Icteridae) and S. agelaius n. sp. from Agelaius phoeniceus Linnaeus (Icteridae), both from Arizona. Spizella breweri (Cassin) (Emberizidae) from California is a new host for Syringophiloidus sialius Skoracki, Flannery & Spicer, 2009; and Melospiza lincolnii (Auduborn) (Emberizidae) from Texas and Vermivora ruficapilla (Wilson) (Parulidae) from California are new hosts for S. seiuri (Ckark, 1964). S. daberti Bochkov, Fain & Skoracki, 2004 from Passerina ciris Linnaeus (Cardinalidae) is recorded in the USA for the first time. A table with the host associations and distribution of all of the North American species of Syringophiloidus is given.

Arthropod symbiotes of the Laotian rock-rat, Laonastes aenigmamus (Rodentia:Diatomyidae), from Laos are examined. This host is a member of Diatomyidae previously thought to have gone extinct >10 million yr ago. Permanent symbiotes are represented by 2 species, a new species of sucking louse, Polyplax sp., near rhizomydis (Phthiraptera:Polyplacidae), and a new species of fur mite, Afrolistrophorus sp., near maculatus (Acariformes:Listrophoridae). The temporary parasites are represented by 18 species, i.e., 1 mesostigmatan species, i.e., a new species of Androlaelaps near casalis (Parasitiformes:Laelapidae); immature stages of 2 tick species, Ixodes granulatus and Haemaphysalis sp. (Parasitiformes:Ixodidae); and a rich fauna of chiggers (Acariformes:Trombiculidae) comprising 8 genera and 15 species. It is hypothesized that this host completely lost its initial fauna of ectosymbiotes and that ancestors of the recorded symbiotes switched to this host from rodents of the superfamily Muroidea.

A new species of syringophilid mites, Mironovia lagopus sp. nov. (Acari: Syringophilidae) is described from feather quills of the rock ptarigman Lagopus muta (Montin) (Phasianidae) from Iceland. Females of this new species differ from the closely related Mironovia rouloul Skoracki and Sikora by setae d2, which are subequal or 1.1-1.2 times shorter than setae e2 (vs. d2 1.4-1.5 times longer than e2 in Mironovia rouloul) and by setae ag1, which are 1.3-1.5 times shorter than ag2 (vs. ag1 1.7-1.8 times shorter than ag2). The postembryonic stages of this new species are figured and described in details. In ontogeny of Mironovia spp., the observed pattern of setal appearance is not different from the pattern in other galliform-associated genera such as Syringophilus and Colinophilus, with exception for tarsal setae p'I, II which are absent in protonymphs. The key to all four species of the genus Mironovia is also provided.

Cheyletoid mites are represented by two lineages being presumably monophyletic. The ancestor of Cheyletidae-Syringophilidae lineage probably was a predator preying on other arthropods. It is hypothesized that syringophilids originated from the common ancestor with Cheyletidae, which switched to preying in bird nests. In Cheyletidae, parasitism on birds and mammals originated independently in several phylogenetic lineages (tribes). All cheyletids are permanents ectoparasites, excluding mites of the tribe Chelonotini. In this tribe, immature instars and males are probably predators in squirrel nests. Cheyletoid lineage II is represented by exclusively permanent parasites of vertebrates belonging to three families Harpirhynchidae (Demodicidae-Psorergatidae). It is presumed from the wide distribution of these mites on birds (Harpirhynchidae) and mammals (Psorergatidae and Demodicidae), that the common ancestor of this branch could have occurred on the common ancestor of birds and mammals; however, switching during an early phase of host evolution can not be excluded. A possible reason for the absence of cheyletoids on recent reptiles (excluding snakes) involves peculiarities of their molting. The high probability of loss of mites during reptile molting seems to have prevented original establishment of cheyletoid parasites on these hosts. These mites are probably absent also on crocodilians because of their aquatic mode of life. In birds, the skin has undergone significant evolutionary changes comparable to what is seen in the integument of mammals. This probably allowed to some cheyletoid mites of the family Harpirhynchidae to transfer to intradermal parasitism in capsules similar to those induced by species of Psorergatidae. The indirect argument of the long-time parasitic relationships between vertebrates and cheyletoids serves a find of mite eggs on the dinosaur's feathers from Lower Cretaceous period (northeast Brazil) (Martill, Davis, 1998). Authors believed that these eggs were laid by feather mites (Astigmata: Psoroptidia). These rounded shape eggs, however, are more similar with those of Cheyletoidea, than with the boomerang-shape eggs of feather mites. The position of the subfamily Ophioptinae associated with snakes of the superfamily Colubroidea in the core of the family Harpirhynchidae (bird parasites) is explained by the switching of its ancestor from passerine birds. Certain snakes feed on nestlings and adult birds, and most of these preys are small passerine birds.

Five new quill mite species of the genus Torotrogla Kethley, 1970 (Acari: Syringophilidae) are described from North American passerines: T. aphelocoma sp.n. from Aphelocoma californica (Corvidae) and T. cardinalis sp.n. from Cardinalis cardinalis (Cardinalidae) in Texas; and T. coccothraustes sp.n. from Coccothraustes vespertinus (Fringillidae), T. cyanocitta sp.n. from Cyanocitta stelleri (Corvidae), and T. piranga sp.n. from Piranga ludoviciana (Thraupidae) in California. A key to females of all known Torotrogla spp. and a table with their host associations and distributions are given.

The evolution of animal parasitism in mites of the infraorder Eleutherengona (Prostigmata) is discussed. Parasitism has arisen independently in numerous phyletic lineages or superfamilies of this infraorder. Mites of the family Pterygosomatidae are parasites of terrestrial arthropods and lizards, and species of Myobiidae are exclusively associated with placental and marsupial mammals. Most families of the superfamily Cheyletoidea comprise permanent parasites of vertebrates, and mites of the sister families Cloacaridae and Epimyodicidae, whose phylogenetic relationships with other eleutherengones are unknown, are endoparasites of turtles and small mammals, respectively. Moreover, some families of the diverse cohort Heterostigmata include insect parasitoids or true parasites of insects. Some phylogenetic lineages, such as Heterostigmata and Cheyletoidea, present a series of life forms transitioning between free-living predators, nidicolous predators, fungivorous mites, and, finally, highly-specialized ecto- and endoparasites. The representatives of some phylogenetically distant eleutherengone lineages developed similar adaptations to predation and parasitism. However, in spite of some similarities in these adaptations, the evolutionary trends and pathways for switching to a parasitic mode of life are quite different in particular eleutherengone lineages. With a few exceptions, each eleutherengone lineage is associated with a particular host group. Temporary parasitism by the larval stage only, a life-history pattern characteristic of the Parasitengona, does not occur in these mites, and all active stages are parasitic and live on host, except for the cheyletid tribe Chelonotini where only adult females are parasites. Species in most eleutherengone lineages that parasitize hosts of particular groups are mono- or oligoxenous parasites, and, therefore, good potential models for co-phylogenetic studies. Mites of the family Pterygosomatidae are permanent parasites of lizards and various arthropods that typically live in concealed habitats. This family represents the only example of mites belonging to one exclusively parasitic eleutherengone group occurring on both invertebrate and vertebrate hosts. The most probable explanation of the occurrence of this family on such varied host groups is host switching. It is hypothesized that Pimeliaphilus-like mites parasitized different arthropods living in concealed spaces and switched to parasitism on lizards that sought refuge in similar spaces. This host switch would have been ancient to account for the diversity of lizard parasites seen today. Mites of the family Myobiidae are ectoparasites of mall marsupial and placental mammals. The development of effective structures for attaching to mammalian hair, primarily involving the first pair of legs, has increased in complexity during myobiid evolution. The subfamily Xenomyobiinae includes a single species Xenomyobia hirsuta parasitizing the Peruan marsupial Lestoros inca. Legs I of this species are only slightly modified. Mites of the subfamily Archemyobiinae, tribe Archemyobiini parasitize other South American marsupials. In these mites, the genua of the female legs I bear paired ventral clasping organs. Mites of the tribe Australomyobiinae parasitize Australian marsupials. The last tribe and all other myobiids belonging to the subfamilies Protomyobiinae and Myobiinae have an unpaired attachment organ on genu I. The subfamily Protomyobiinae is separated onto 3 tribes. In more derivative members of the tribes Acanthophthiriini (parasites of bats), Protomyobiini (parasites of "insectivores"), and in all Elephantulobiini (parasites of elephant shrews), the tibia and tarsus of legs I are fused apically. Mites of the subfamily Myobiinae parasitize rodents and are the most morphologically specialized, with 3 apical segments of legs I fused. Basing on the wide distribution of myobiid mites on mammalian taxa, it could be concluded that the origin of parasitism occurred in these mites not later than in the Lower Cretaceous, the assumed time of divergence of marsupial and placental mammals (Carroll, 1993). The host associations of the superfamily Cloacaroidea is still unresolved problem. Mites of this superfamily show a high level of host specificity; however, cloacaroid families and subfamilies are associated with phylogenetically distant lineages of vertebrates, turtles (Cloacarinae), birds (Pneumophaginae), and mammals (Epimyodicidae). The common ancestor of cloacarines may have become associated with turtles before the late Triassic. We believe that parasitism of Pneumophagus bubonis, the single species of Pneumophaginae, in the lungs of Bubo virginianus (Aves: Strigiformes) is a result of an ancient host shift following predation by some birds on live or. dead turtles. All 4 currently known species of the Epimyodicidae are endoparasites in the subcutaneous tissues of voles (Rodentida: Cricetidae), moles, and shrews (Soricomorpha: Talpidae and Soricidae). The sister relationship between Cloacaridae and Epimyodicidae could reflect parasitism by their common ancestor on archaic amniotes before the divergence of the Synapsida (ancestors of mammals) and Diapsida (ancestors of turtles and birds). On the other hand, the ancestors of epimyodicids, retaining some features of their free-living ancestors, could have switched from diapsids onto synapsids considerably later.

The external morphology of 2 closely related mite families, Cloacaridae and Epimyodicidae (Acari: Prostigmata), comprising highly specialized endoparasites of vertebrates, is analyzed. These mites exhibit strong regression of many structures ancestrally present in other Prostigmata as a consequence of their endoparasitic mode of life. The relationships of these 2 families with other taxa in the infraorder Eleutherengona are still not clear. Our reinterpretation of the chelicerae as unfused precludes inclusion of this lineage in the Cheyletoidea as proposed previously. A new superfamily, Cloacaroidea superfam, nov., incertae sedis, within the infraorder Eleutherengona is established for these 2 families, and their host-parasite relationships are briefly discussed. A new cloacarid species, Caminacarus dawsoni n. sp., from Graptemys pseudogeographica (Testudines: Emydidae) from the United States is also described.

Two species of Chirorhynchobiidae, Chirorhynchobia glossophaga sp. n. and Chirorhynchobia matsoni Yunker, 1970 are described and redescribed, respectively. The male of C. glossophaga and the tritonymph of C. matsoni are the first to be described in the family. The mites are ectoparasites living on the wing membrane of phyllostomid bats Glossophaga soricina (Pallas) from Peru and Anoura geoffroyi Gray from several localities. Females of the new species differ from those of C. matsoni by the shorter idiosoma (length 439-450 versus 497-562 in C. matsoni), by the dorsal position of setae 4a (terminal in C. matsoni), and by the narrower posterior projections of the propodonotal shield (width 6 versus 17). Although the morphology of Chirorhynchobiidae is strongly regressive, homologies of the leg and idiosomal setae in these mites with those of other Astigmata are established. The combination of absence of famulus epsilon and solenidia sigmaI-II and presence of solenidia omega1I-II and omega3I in these mites is shared with the Rhyncoptidae-Sarcoptidae clade, suggesting a possible relationship with those groups.

The term synhospitality means the association of two or more closely related parasite species with one host species (Eichler, 1966). The cases of two or three synhospitalic species are known from the same host species, and especially ones where parasites were recorded from different parts of the host range, are quite common. The most ordinary reason causing synhospitality in permanent parasites is the host switching. Nevertheless, there are a number of synhospitality cases, where the parasite complex is monophyletic because evolved on a single host species. The special term--"phylogenetic synhospitality" (FS) is proposed for these cases of synhospitality. Most known cases of FS in acariform mites, permanent parasites of vertebrates, are analysed. It is found out that both astigmatan and prostigmatan parasite mites demonstrate a numbers of FS. The majority of these examples represent parasitism of two or three synhospitalic parasite species. Impressive examples of FS involving a number of synhospitalic species is shown by only astigmatan mites inhabiting the fur of mammals or plumage of birds. Most known examples involving four or more mite species are discussed: 51 mite species of the genus Schizocarpus (Chirodiscidae) parasitizing Castor fiber and C. canadensis (Castoridae); 6 species of Listrophorus spp. (Listrophoridae) from Ondatra zibethicus (Cricetidae); 23 species of Listrophoroides s. 1. (Atopomelidae) from Maxomys surifer (Muridae); 21 species of Cytostethum (Atomelidae) from Potorous tridactylus (Potoridae); 4 species of Listrophoroides (Afrolistrophoroides) from Malacomys longipes (Muridae); 7 species of Fainalges (Xolalgidae) from Aratinga holochlora (Psittacidae); 4 species of Zygepigynia (Pteronyssidae) from Chrysocolaptes lucidus (Picidae). The main reason of FS is that, in spite of the Fahrenholz's rule, the speciation of many parasites proceeds much more intensively than in their hosts because of the more rapid replacement of the parasitic generations. The first factor causing FS is the mite speciation it temporary segregated populations of the host (allopatric speciation). In this case, the "multispecies complexes" appeared after the subsequent reintegration of the host populations formerly isolated. The second factor is the speciation due to the specialization of mites to local microhabitats in the fur or plumage of host (sympatric or synxenic speciation). The second way of speciation is most characteristic for mites with highly specialized attaching structures. The phenomenon of FS more resides in ectoparasites of mammals rather than in feather mites in spite of much more structural complicacy of plumage rather than the fur. The high mobility of birds and wide dispersion of their new generations probably embarrass the process of sympatric speciation in their parasites. As a rule, only really significant geographical barriers play role for population isolation in birds. Thus, it could be concluded that two independent factors or their combination lead to FS. (i) The complex and/or disjunctive host range giving a possibility for allopatric speciation in parasites. (ii) The deep mite specialization to local microhabitats on the host body causing sympatric (synxenic) speciation. Fur of mammals and plumage of birds are very complicated in structure and microconditions and provide a considerable number of different microhabitats for mites inhabiting them. The prevalence of one of these two factors depends on the biological peculiarities of both parasites and their hosts. In mites with lesser specialized attaching organs, for example in atopomelids, allopatric speciation dominates. In mites with strongly specialized attaching organs, for example in listrophorids or chirodiscids, both pathways of speciation may take place. In feather mites, sympatric speciation should be more probable due to quite complicate and various structure of feathers in avian hosts. In fur mites, sympatric speciation is more likely in mites parasitizing hosts with peculiar ecology, for example in semiaquatic rodents possessing quite different fur structure in different parts of the body.

The external morphological adaptations to parasitism in acariform mites (Acari: Acariformes), permanently parasiting mammals, are briefly summated and analyzed. According to several external morphological criteria (structures of gnathosoma, idiosoma, setation, legs and life cycle), the following six morphoecotypes were established: skin mites (i)-- Cheyletidae, Chirorhynchobiidae, Lobalgidae, Myobiidae, Myocoptidae (the most part), Rhyncoptidae, Psoroptidae; fur mites (ii)--Atopomelidae, Clirodiscidae, Listrophoridae, Myocoptidae (Trichoecius only); skin burrowing mites (iii)--Sarcoptidae; intradermal mites (iv) - sorergatidae and Demodicidae; interstitial mites (v) - pimyodicidae; respiratory mites (vi) - reynetidae, Gastronyssidae, Lemurnyssidae, Pneumocoptidae. In the case of prostigmatic mites, the detailed reconstruction of the origin and evolution of "parasitic" morphoecotypes is possible due to the tentative phylogenetic hypotheses, which were proposed for the infraorder Eleutherengon, a, including the most part of the permanent mammalian parasites among prostigmatic mites (Kethley in Norton, 1993; Bochkov, 2002). The parasitism of Speleognathinae (Ereynetidae) in the mammalian respiratory tract arose independently of the other prostigmats. It is quite possible that these mites switched on mammals from birds, because they are more widely represented on these hosts than on mammals. The prostigmatic parasitism on mammalian skin seems to be originated independently in myobiids, in the five cheyletid tribes, Cheyletiellini, Niheliini, and Teinocheylini, Chelonotini, Cheyletini, and, probably, in a cheyletoid ansector of the sister families Psorergatidae-Demodicidae (Bochkov, Fain, 2001; Bochkov, 2002). Demodicids and psorergatids developed adaptations to parasitism in the skin gland ducts and directly in the epithelial level, respectively in the process of the subsequent specialization. Mites of the family Epimyodicidae belong to the phylogenetic line independent of other cheyletoids. These mites possess the separate chelicerae and, therefore, can not be included to the superfamily Cheyletoidea. It is not quite clear whether they were skin parasites initially or they directly switched to parasitism from the predation. The phylogeny of sarcoptoid mites (Psoroptidia: Sarcoptoidea) is not developed, however, some hypotheses about origin and the following evolution of their morphoecotypes can be proposed. We belive that astigmatic mites inhabiting the mammalian respiratory tract transferred to parasitism independently of other sarcoptoids. The idiosoma of these mites is not so much flattened dorso-ventrally and has proportions which are similar to hose of free-living astigmatids. Moreover, in the most archaic species, the legs are not shortened or thickened as in the most parasites. The disappearance of many morphological structures in these mites, probably, happened parallely with some other sarcoptoids due to their parasitic mode of life. The skin inhabiting sarcoptoids belong to the "basic" morphoecotype, and all other sarcoptoid morphoecotypes, excluding respiratory mites, are derived from it. Some mites of this morphoecotype live on the concave surfaces of the widened spine-like hairs of the rodents belonging to the family Echimyidae (mites of the subfamily Echimytricalginae), in the mammalian ears (some Psoroptidae) or partially sink into the hair follicles (Rhynocoptidae). Finally, mites of the family Chirorhynchobiidae live on the bat wing edges attaching to them by their "ixodid-like" gnathosoma. The fur-sarcoptoids, probably, originated from the skin mites. This morphoecotype is divided onto two subtypes: mites with the dorso-ventrally flattened idiosoma (subtype I) and mites with the teretial idiosoma (subtype II). Each "fur-mite" family includes mites of the both subtypes. All mites of the first subtype belong to the early derivative lineages in their families. Among listrophorids such early derivative lineage is represented by the subfamily Aplodontochirinae (Bochkov, OConnor, 2006), and among Chirodiscidae--by mites of the subfamilies Chirodiscinae, Schizocoptinae, and Lemuroeciinae. Among the "fur" astigmatid families, the family Atopomelidae. probably, is the most archaic, and the most part of atopomelids belongs to the first subtype. However there are several more specialized atopomelid genera belonging to the second subtype, Atopomelus, Dasyurochirus, Lemuroptes, Murichirus, Metachiroecius etc. We believe that mites of the first subtype are represented by the "intermediate" forms between skin mites and mites of the second subtype. Some skin sarcoptoids transferred from skin parasitism to burrowing of the host skin (Sarcoptidae). The established morphoecotypes are partially corresponding to some morphoecotypes established by Mironov (1987) for feather mites. Our morphoecotypes of skin and skin burrowing mites perfectly correspond to Mironov's epidermoptoid and knemidocoptoid morphoecotypes, respectively. The proctophylloid morphoecotype (mites living on the wing feathers), which is the most widely represented within feather mites, has an analogy among mammalian mites - the subfamily Echimytricalginae. The analgoid (mites living in the down feathers) and dermoglyphoid (mites living in the feather quills) morphoecotypes have no analogues among mammalian mites for the obvious reasons. It is interesting why some microhabitats on the host body are not still occupied by prostigmatic or astigmatic mites. We believe that the nutrition is the main limitative factor here. The parasitic prostigmates evolved from predators and, therefore, feed on content of the living cells. They need the direct contact with the live tissues of the host and they belong, therefore, to the morphoecotypes represented by the respiratory, skin, gland duct, intradermal, and interstitial mites. Whereas, the most part of the skin inhabiting astigmats feed on the dead epithelial scales. For this reason these mites, so easily colonized fur of their hosts and feed on the hair grease there. On the other hand, some sarcoptoids transferred to the true parasitism and feed on the cambial cells of the skin epithelium. More over we do not know exactly about nutrition of rhyncoptids yet.

The paper reports the results of eight-year investigations on the ectoparasites of rodents and insectivores carried out in southern taiga of the Ilmen-Volkhov lowland (Novgorod Region) and Kurgolovsky reserve (Leningrad Region). Twelve species of small mammals were captured including three dominate species--bank vole Clethrionomys glareolus (2722 specimens), common shrew Sorex araneus (1658 specimens), and wood mouse Apodemus uralensis (367 specimens). Parasite community of the bank vole comprises 34 species of mites, ticks, and insects, the community of common shrew comprises 25 species, and the community of A. uralensis includes 28 species. Taxonomic diversity of the ectoparasite communities was shown to be based on the diversity of types of parasitism and ecological nishes of the host body. Permanent ectoparasites are found to be represented by 2 species of lie and 14 species of acariform mites. The group of temporary parasites includes 13 species of fleas, 10 species of gamasid mites. 3 ixodid species and 1 Trombiculidae. There is a common pool of temporary parasites of small mammals in the ecological system of taiga. Significance of different shrew and rodent species as hosts were found to be dependent on the population density in possible hosts and many other factors. Species diversity in the parasite communities of different small mammal species is dependent on the number of possible ecological nishes in the host body. Actual infill of these nishes by ectoparasites is usually lesser than potential one. Species composition of temporary parasites, their occurrence and abundance changes according to season. Interspecific competition in the temporary parasite species can decrease because of the seasonal disjunction of their population peaks. Diversification of the ecological niches of ectoparasites allow simultaneous feeding of more parasite individuals on one host, than in the case of parasitising of single species or several species with similar ecological nishes. The distribution of parasites on their hosts was also studied. The aggregative distribution has been found in ixodid larvae only, and the distribution of fleas was close to the Poisson distribution. Deviations from the aggregative distribution can be an effect of several independent factors, including limited ability of small mammals for providing numerous parasites with food. On the most part of hosts simultaneous parasitizing of no more than 1-3 individuals of each tick, mite, and flea species was registered. Excessive infestation by ectoparasites may probably be limited by effective reactions of self-purification in the mammal hosts.

The mesostigmatid genus Andreacarus Radford, 1953 (Acari: Laelapidae), species of which are obligatory parasites of small mammals, is revised. Andreacarus includes 11 species, four species previously recognized and seven new species described from Madagascar hosts: A. brachyuromys sp. n. from Brachyuromys betsileoensis Bartlett, A. eliurus sp. n. from Eliurus species, A. gymnuromys sp. n. from Gymnuromys roberti Major, A. voalavo sp. n. from Voalavo gymnocaudus Carleton & Goodman, A. nesomys sp. n. from Nesomys rufus Peters-all from nesomyid rodents; A. tenrec sp. n. from Tenrec ecaudatus Schreber (Afrosoricida: Tenrecidae); and A. galidia from Galidia elegans I. Geoffroy Saint-Hilaire (Carnivora: Viverridae). An amended generic diagnosis and key to females are also given. Six species described in Andreacarus from Australian and New Guinean hosts are removed and transferred to the new genus, Juxtalaelaps.