New records of feather mites ( Acariformes : Astigmata ) from non-passerine birds ( Aves ) in Brazil

We present the results of our investigation of feather mites (Astigmata) associated with non-passerine birds in Brazil. The studied birds were obtained from roadkills, airport accidents, and from capitivity. Most ectoparasites were collected from bird specimens by washing. A total of 51 non-passerine species from 20 families and 15 orders were examined. Of them, 24 species were assessed for feather mites for the first time. In addition, 10 host associations are recorded for the first time in Brazil. A total of 101 feather mite species were recorded, with 26 of them identified to the species level and 75 likely representing undescribed species; among the latter samples, five probably represent new genera. These records allowed the first inference about the host and mite association of many species, as well as the first discussion about the geographical distribution of some feather mite taxa along the host distribution.


INTRODUCTION
Feather mites (Astigmata: Analgoidea and Pterolichoidea) are the most diverse and abundant group of ectosymbionts living on bird plumage (Gaud and Atyeo 1996;Proctor 2003).With the exception of the family Pyroglyphidae (Analgoidea), in which some representatives live in nests rather than on a host's body, the majority of feather mites have morphological adaptations for permanent living in specific microhabitats on birds, such as downy, covert or flight feathers, cavities inside quills of the flight feathers, and the skin surface (Dabert and Mironov 1999).Depending on the host species, each type of microhabitat can harbor a variety of feather mite species, often with specific microhabitat partitioning or coexistence without intense competition (Atyeo and Pérez 1988;Pérez 1995;Bochkov and Mironov 2008;Hernandes and Mironov 2015).This considerably enhances the mite diversity which can be found in a single bird species.
While most cases of feather mite transfer occur by physical contact between birds of the same species (e.g., during parental care and copulation), which often links the evolutionary path of the groups and may to some extent mirror their phylogenetic trees (Dabert and Mironov 1999;Dabert 2005), there are exceptional cases of interspecific tranfers that are poorly understood (Mironov and Dabert 1999;Hernandes et al. 2014).Feather mites are often reported as ectocommensals, living harmlessly on the bird's body, feeding on the uropygial oil produced by the birds (Blanco et al. 2001).The few records of parasitism by feather mites occur mainly in groups that live on the skin surface and on the downy feathers, close to or in contact with the skin, such as the analgoid families Analgidae, Psoroptoididae, Dermationidae, and Epidermoptidae, with the latter two families being true parasites by any criteria (Fain 1965;Mironov 2013;Hernandes et al. 2014;Soares et al. 2016).
More than 2,400 feather mite species have been described worldwide, although this may represent only about 20% of the extant total (Gaud and Atyeo 1996;Mironov 2003;Proctor 2003).Many species have been described from birds of the Palearctic and Afrotropic regions (Gaud and Atyeo 1996), while birds from other tropical regions, specifically the Neotropics, still harbor a myriad of undescribed mites (Valim et al. 2011;Barreto et al. 2012).Brazil is currently the country with the richest avifauna of the world, with nearly 1,900 species (CBRO 2014).However, less than 20% of the bird species from this country has been explored for their feather mites (Valim et al. 2011).The lack of knowledge and the huge bird diversity makes Brazil one of the best places for feather mite taxonomic discoveries, specifically for bird orders other than Passeriformes, which are still poorly sampled in feather mite studies mainly due to the general difficulty in collecting these birds.Here, we Below we present the feather mites using the sequence and classification proposed by Gaud and Atyeo (1996), with further modifications (e.g., OConnor 2009), as the superfamily Freyanoidea is included in the superfamily Pterolichoidea.Bird sequence and taxonomy follows Del Hoyo (1992,1994,1996,1997,1999,2001,2002) for suprafamilial ranks and CBRO (2014) for family, genera and species.

RESULTS
In total, we examined 108 specimens of 51 bird species in 20 families and 15 orders, representing 2.7% of the 1,902 species occurring in Brazil and 6.1% of the 837 nonpasserines from the country (CBRO 2014).Twenty-four bird species were assessed for the first time concerning their feather mites, and 10 new host-simbiont associations were discovered for Brazil (Table 1).As for the mites, 101 species from 18 families were recorded (Table 2); from report feather mite associations with non-passerine birds from Brazil.

MATERIALS AND METHODS
We analyzed mites from 51 bird species.The majority of these birds (43 species) were roadkills and airplane casualties from various localities in Brazil.Ten bird species were represented by detached feathers collected from captive individuals.The dead birds were washed to remove ectosymbionts using water and detergent (Clayton and Walther 1997), then the water was passed through filter paper and the captured material was examined under a dissecting microscope.Retrieved mites were put into 30% lactic acid at 50°C for 24 hours, then mounted in microscopic slides using Hoyer's medium and heated at 50°C for 5 days.The slides were then sealed with varnish and labeled.By means of light microscope with differential interference contrast (DIC) mites were identified to supraspecific taxa using the dichotomous keys in Gaud and Atyeo (1996) and to species, when possible, using relevant literature for each taxon.Voucher specimens Table 1.Feather mite species recorded on non-passerine birds in Brazil in this study.N is the number of analyzed birds.ORIG = origin of the host: Wwild, C -captivity.SF. = feather mite superfamilies: A -Analgoidea, P -Pterolichoidea.Birds: (*) birds with first record of feather mites; (**) birds with first record of feather mites for Brazil.Feather mites: (*) first record for Brazil and the host, (**) first record for Brazil, (***) first record only for the host species.1).Feather mites were recorded for the first time in Brazil on the following bird families: Ardeidae, Falconidae, Tytonidae, and Nyctibiidae.The richest feather mite families recorded were Pterolichidae (27 species), Xolalgidae (12 species), Gabuciniidae and Kramerellidae (nine species each) (Table 2).The fact that only non-passerine birds were analyzed explains the higher number of pterolichoidean mites (62 species) in comparison with analgoideans (39 species).Pterolichidae was also the family with the highest number of probable new genera (four out of five supposed new genera), all belonging to the subfamily Pterolichinae.

Corrections to previously published records of feather mites from Brazil
In our study we found some mistakes and missed records in the previous checklist of feather mites from Brazil (Valim et al. 2011).These mistakes were mostly caused by inaccurate locality information originally reported for the birds.After analyses of each case, eight feather mite species were added to the list and four were removed, and eight bird species were added and six previously indicated as hosts were removed.

Bird species excluded
• Pteroglossus torquatus Gmelin, 1788 (Piciformes: Ramphastidae) is removed from the list.Valim et al. (2011: 299) listed this bird as one of the hosts of Ramphasto bius chiasma.However, this host occurs only in Central America and was originally reported from Costa Rica, Honduras, and Mexico (Atyeo et al. 1987: 156), not from Brazil.• Ramphastos cuvieri (Wagler, 1827) (Piciformes: Ramphastidae) is considered a subspecies of R. tucanus Linnaeus, 1758 by the majority of the bird taxonomists (Vitor Q. Piacentini pers.comm.), and therefore, the record of this bird as a host species of the mite Ramphastobius loricatus (Pteronyssidae) was removed from the list.• Crypturellus boucardi (Sclater, 1859) (Tinamidae) from Mexico was indicated as the type host of Mesosathes meniscurus (Crypturoptidae) and was mistakenly included in the appendix list of Brazilian hosts (Valim et al. 2011: 317).This bird is endemic to Central America and does not occur in Brazil.
Genus Dinalloptes Gaud & Mouchet, 1957 Dinalloptes chelionatus Atyeo & Peterson, 1966 (Figures 3 and 4) Remarks.The genus Megninia currently includes more than 30 species associated with galliforms, cuculiforms, coliiforms, and passeriforms (Mironov and Galloway 2002a).Mironov and Galloway (2002a) noted that Meg ninia badly needs a revision because it includes within it morphologically distinct mites from several orders of birds.The species recorded here represents an undescribed species most similar to M. ginglymura (Méginin, 1877), a mite already reported on domestic chickens in Brazil (Amaral et al. 1975;Gaud et al. 1985).This is the first record of a feather mite on this host.Remarks.The undescribed species recorded here is most similar to M. cubitalis (Mégnin, 1877) and is another species recorded on domestic chickens in Brazil (Amaral et al. 1975;Gaud et al. 1985).This is the first record of a feather mite on this host.Remarks.Megniniella gallinulae has been recorded in the New World from Cuba on G. galeata and also in the Old World on G. chloropus (Linnaeus, 1758), G. angulata Sundevall, 1850, and Amaurornis phoenicurus phoenicurus (Černý 1967;Gaud 1958;1968a;Gaud and Mouchet 1959a;Mironov and Galloway 2002b;Wang and Fan 2010).This is the first record of Megniniella in Brazil and the first record of feather mites on G. galleata from Brazil, since this bird species was split from G. chloropus (see Remsen et al. 2016: Proposal 416, September 2009).
Remarks.The genus Scutalges includes three species described from cuculids from Africa (Gaud 1966).The species recorded here represents an undescribed species most similar to S. carreti (Gaud and Mouchet, 1959).This is the first record of Scutalges outside Africa and also the first record of feather mites from P. cayana in Brazil.
Family Dermationidae Fain, 1965Subfamily Dermationinae Fain, 1965 Genus and species undetermined Remarks.Two dermationine genera have been previously recorded on swifts (Apodidae): Apodicoptes Fain, 1965and Passeroptes Fain, 1965(Fain 1965;Mironov et al. 2005a).This mite recorded here belongs to an undescribed genus due to a unique set of features: the opisthosoma is rounded in males, tibiae III and IV lacking apicoventral expansions, and femora III and IV have well pronunciated retrograde spike-like apophyses in both sexes.This is the first record of a dermationid on C. meridionalis.
Genus Paddacoptes Fain, 1965 Paddacoptes talpacoti (Fain, 1965)  Remarks.The genus Paddacoptes was originally established as a subgenus of Passeroptes (Fain 1965) but was elevated to full generic rank by Gaud and Atyeo (1996).It includes seven species recorded on passeriforms (five species) and columbiforms (two species) (Fain 1965).Paddacoptes talpacoti was originally described from Columbina talpacoti from Brazil.Although there were no illustrations originally provided for that species, this mite perfectly fits the description (Fain 1965).This is the first report of this species since its original description.Remarks.This species is most similar to Passeroptes inermis Fain, 1965, which was described from Calocitta formosa Swainson, 1827 (Passeriformes: Corvidae) (Fain 1965).This is the first record of a feather mite on C. melanochloros and the first record of a dermationid mite on a bird of the family Picidae.

Genus
Genus Psittophagoides Fain, 1964 Psittophagoides brotogeris Fain & Bochkov, 2003 Material examined.1 male and 7 females ex Brotogeris chiriri Remarks.Psittophagoides brotogeris was described from Brotogeris versicolurus (Statius Müller, 1776) from the Brazilian Amazon (Fain and Bochkov 2003) and is here recorded for the first time on B. chiriri.Remarks.Among galliform hosts, Rivoltasia was previously recorded only on hosts of the families Phasianidae and Odontophoridae (Fain 1965;Forrester and Spalding 2003).This is the first record of the Dermationidae on a cracid bird.The only previous record of Rivoltasia in Brazil is that of R. bifurcata (Rivolta, 1876) on domestic chickens (Reis 1939).
Remarks.Five species of Toxerodectes have been recorded on trochilids from Brazil (Park and Atyeo 1973;Park and Atyeo 1974;Kanegae et al. 2008;Hernandes 2013a).This undescribed species represents the second species of Toxerodectes recorded on T. glaucopis; Toxero dectes biscutatus has been recorded in Brazil on this host species from Paraná state (Hernandes 2013a).
Remarks.The genus Chiasmalges includes four species recorded from New World psittacids (Trouessart 1899;Gaud and Atyeo 1967;Pérez and Ramirez 1996;Mironov et al. 2005b;Valim et al. 2011).This is the first record of Chiasmalges on A. ararauna and also in Brazil.Remarks.Two species of Paralgopsis are known: P. paradoxus (Trouessart, 1899) and P. ctenodontus (Gaud, 1968), from psittacids of Colombia and Brazil, respectively (Gaud 1968b).This is the first record of feather mites on C. fasciolata, and the first record of a feather mite of the subfamily Paralgopsinae on Galliformes.Recently, molecular based studies showed that the subfamily Paralgopsinae belongs to the Psoroptoididae, rather than the Pyroglyphidae (Klimov and OConnor 2013;Klimov et al. 2015).Remarks.The genus Paralgopsis was previously recorded on P. leucophthalma from captivity in Brazil (Jardim et al. 2012).
Remarks.From the 13 described species of Fainalges, three were described from Brazilian birds: F. annulifer (Trouessart, 1899) from Deroptyus accipitrinus (Linnaeus, 1758) (Psittacidae); F. intermedius (Trouessart, 1899) from Aratinga solstitialis (Linnaeus, 1766) (Psittacidae); and F. trichocheylus Gaud and Berla, 1964 from Melanerpes flavifrons (Vieillot, 1818) (Picidae).The last host record is probably a case of contamination, because Fainalges species occur exclusively on New World parrots (Gaud and Berla 1964;Pérez 1996;Mironov et al. 2005b).This is the first record of the genus Fainalges on both A. chloroptera and A. ararauna.Remarks.This species is most similar to F. vulgaris Pérez 1995, which was described from Psittacara holochlo ra from Mexico (Pérez 1995).This is the first record of the genus Fainalges on P. leucophthalma.Remarks.The genus Glaucalges comprises three species, two of them recorded from owls (Strigiformes), and one from an African musophagid (Cuculiformes) (Gaud 1980;Gaud and Atyeo 1981;Dabert et al. 2008).This mite is distinctly different from the description of G. tytonis Dabert et al. 2008collected on T. alba (Scopoli, 1769) from Europe (Dabert et al. 2008) and seems closely related to G. attenuatus (Buchholz, 1869) instead, based on the length of the setae e2.This is the first record of feather mites on T. furcata and of Glaucalges in Brazil.Remarks.The genus Glaucalges was recorded on A. cunicularia from Florida (Forrester and Spalding 2003).This is the first record of feather mites on A. cunicularia in Brazil.

Protonyssus mironovi
Remarks.This species was also recently described from this material (Hernandes and Pedroso 2016).Remarks.In the revision of the subfamily Ingrassiinae, Gaud and Atyeo (1981) recognized a single species in this genus, P. stellaris (Buchholz, 1869), which was described from Botaurus stellaris (Linnaeus, 1758) and recorded from several other species of herons, egrets, and bitterns (Ardeidae).However, they acknowledged that there might actually be several distinct species.The species we found on N. nycticorax does not correspond to P. stellaris.This is the first record of Pteralloptes in Brazil.
Remarks.This species from S. sibilatrix also does not correspond to P. stellaris and has features that clearly distinguish it from the Pteralloptes sp.A reported on N. nycticorax above.This is the first record of feather mites on S. sibilatrix.

Remarks. Ascouracarus chordeili was described on
Chordeiles rupestris from Brazilian Amazon (Mironov and Fain 2003).This is the first record of feather mites on L. semitorquatus.
Remarks.The genus Ascouracarus comprises four species, three of them from the Caprimulgiformes and one from Psittaciformes (Gaud and Kolebinova 1973;Gaud and Atyeo 1976;Dabert and Ehrnsberger 1992; Pedroso and Hernandes | Feather mites from non-passerine birds in Brazil Mironov and Fain 2003).This quill mite represents an undescribed species, and is the first record of Ascouracaridae on a member of the order Cuculiformes.
Remarks.This species is most similar to A. chordeili Mironov & Fain, 2003 but represents an undescribed species.This is the first record of the Ascouracaridae on birds of the family Nyctibiidae and the first record of feather mites on N. griseus from Brazil.
Remarks.This species was described on Nothura maculosa from Argentina (Alzuet and Brandetti 1990), and is here recorded for the first time in Brazil.

Remarks.
As in some psittacids (Pérez 1995) and cuckoos (Mironov et al. 2015), swifts (Apodidae) are known to host more than one species of the same genus (Peterson et al. 1980).

Remarks. Rhynchocaulus paradoxus was described on
Streptoprocne zonaris from Mato Grosso state in Brazil (Gaud and Berla 1963) and is recorded here for the first time on S. biscutata.

Remarks. Hyperaspidacarus tridentatus has already been recorded in Brazil on
Columbina squammata and on C. passerina (Atyeo and Smith 1983).This is the first record of H. tridentatus on C. talpacoti from Brazil.
Remarks.Falculifer leptotilae was described from L. jamaicensis (Linnaeus, 1766) from Jamaica and Mexico and also recorded on L. verreauxi Bonaparte, 1855 from Trinidad, Guyana, and Colombia (Gaud and Barré 1992).This species has also been previously recorded on L. rufaxilla from Tocantins state in Brazil (Enout et al. 2012).Remarks.The genus Falculifer comprises 12 species restricted to columbiform birds (Gaud 1976;Gaud and Barré 1992).This is the first record of feather mites on P. picazuro.Remarks.This is the first record of feather mites on P. picazuro.Remarks.This is the first record of feather mites on P. speciosa.
Remarks.This species was recently described from this host in Brazil (Hernandes et al. 2016 Remarks.This genus currently includes 12 species from birds of the families Accipitridae, Sagittariidae, and Otitidae (Gaud and Atyeo 1974;Gaud 1983Gaud , 1968a;;Proctor et al. 2006); it is recorded for the first time in Brazil.Remarks.This is the first record of feather mites on M. chimachima.

Aetacarus
Genus Capitolichus Gaud & Atyeo, 1975 Capitolichus sp.(Figures 31 and 32 Remarks.There are four named species in this genus (Gaud and Atyeo 1975;Alzuet et al. 1988), collected from Piciformes (Capitonidae), Passeriformes (Cotingidae, Tyrannidae), and Trogoniformes (Trogonidae).Forrester and Spalding (2003) reported an undetermined species of Capitolichus from Dryocopus pileatus (Linnaeus, 1758) (Picidae).The mite from C. campestris represents an undescribed species.This is the first record of feather mites on C. campestris and also the first record of the genus Capitolichus in Brazil.Remarks.The genus Hieracolichus occurs on birds of the family Accipitridae and includes nine named species (Gaud and Atyeo 1975;Gaud 1983;Mironov et al. 2007).This is the first record of feather mites on H. meridionalis and the first record of the genus Hieracolichus in Brazil.Remarks.This is the first record of feather mites on I. plumbea.
Remarks.Piciformobia guirae was described from G. guira from Argentina, and was also recorded on the same host species in Brazil (Alzuet et al. 1988).
Remarks.This is a second species of the genus Piciformobia from G. guira.
Remarks.This mite represent a second undescribed species of the genus Paragabucinia on H. albicollis.The species Paragabucinia brasiliensis Hernandes, 2014 was previously described on the same host in Brazil (Hernandes 2014).
Genus Tocolichus Gaud & Atyeo, 1975 Tocolichus sp.(Figures 35 and 36 Remarks.Two species are known in this genus: T. allepimerus, described from Selenidera maculirostris (Lichtenstein 1823) from Brazil; and T. ramphastinus (Mégnin and Trouessart, 1884), reported from many species of toucans (Gaud and Atyeo 1975).This mite represents an undescribed species of the genus Tocolichus and is the first record of feather mites on R. toco.Remarks.There are six named species of Dermonoton (Gaud and Mouchet 1959b;Gaud 1980).The mite recorded here represents an undescribed species.This is the first record of feather mites on T. furcata and the first record of the genus Dermonoton in Brazil.
Remarks.The species D. parallelus was recorded on A. cunicularia in Florida, USA (Forrester and Spalding 2003).The mite from A. cunicularia reported here represents an undescribed species.This is the first record of feather mites on A. cunicularia from Brazil.Remarks.Hernandes and OConnor (2015) recently described the first feather mite from an owl in Brazil, an ascouracarid, Cystoidosoma hermaphroditus, described from M. choliba.This species of Dermonoton clearly represents an undescribed species and is the first record of the genus Dermonoton on M. choliba.Remarks.This mite represents an undescribed species of Dermonoton and is the first record of feather mites on P. koeniswaldiana.
Genus Kramerella Trouessart, 1916 Kramerella quadrata Gaud, 1980 (Figure 39  Remarks.Kramerella quadrata was described from T. alba affinis (Blyth, 1862) from Cameroon in Africa (Gaud 1980) and was also recorded on T. alba from Florida, USA (Florrester and Spalding 2003).It is recorded for the first time in Brazil.Remarks.This species seems most similar to Kram erella oti Lönnfors, 1937 that was described on Asio otus (Linnaeus, 1758) from Africa (Gaud 1980).This is the first record of feather mites on Asio stygius.

Remarks. Three species are recognized in the genus
Petitota (Gaud and Mouchet 1959a;Atyeo and Philips 1984;Mironov 1997).This genus is considered rare in mite collections (Atyeo and Philips 1984).The specimens reported here represent an undescribed species and the first records of Petitota on M. choliba and in Brazil.
Remarks.The genus Pseudogabucinia includes five species reported from birds of the families Ciconiidae, Falconidae, Accipitridae, Gruidae, and Otididae (Gaud 1968a;Gaud 1982;Gaud and Mouchet 1961;Dubinin 1956).From these species, only P. intermedia (Mégnin & Trouessart, 1884) has been recorded on Accipitridae and Falconidae (Gaud 1988).This is the first feather mite recorded on E. leucurus and represents an undescribed species.Remarks.The genus Ardeacarus is typical from ardeids and includes a single named species, A. ardeae (Canestrini, 1878) originally described from Ixobrychus minutus (Linnaeus, 1766) (Ardeidae) from Europe (Dubinin 1951;Gaud 1981a) and later reported from many ardeids around the world.Our specimens represent an undescribed species of Ardeacarus.This is the first record of feather mites on B. pinnatus.Remarks.This is the first record of a feather mite on B. striata from Brazil.The single species of Ardeacarus recorded from this host was A. ardeae from Africa and Russia (Dubinin 1956;Černý 1967;Gaud 1981a).This mite represents another undescribed species in the genus Ardeacarus.Remarks.This is the first record of a feather mite on this host from Brazil.Currently, the single species of Ardeacarus recorded from this host is A. ardeae from Africa (Gaud 1981a).This mite represents another undescribed species in the genus Ardeacarus.Remarks.This is the first record of feather mites on Syrigma sibilatrix.This mite represents another undescribed species in the genus Ardeacarus.
Remarks.The single species of Ardeacarus recorded from this host was A. ardeae from Africa, Russia, Cuba, and USA (Dubinin 1956;Černý 1967;Gaud 1981a).This mite represents another undescribed species in the genus Ardeacarus.Remarks.Ardeacaris ardeae has been recorded on N. nycticorax from Africa, Russia, and Cuba (Dubinin 1956;Černý 1967;Gaud 1981a).This is the first record of feather mites on N. nycticorax from Brazil and also represents another undescribed species in the genus Ardeacarus.Remarks.There are two named Ardeialges species: A. dermogaster Gaud & Mouchet, 1959, described from Trigiornis leucolophus (Jardine, 1846) (Ardeidae) from Africa (Gaud and Mouchet 1959b); and A. herodias (Dubinin, 1951), described from Ardea cinerea Linnaeus, 1758 from Russia but also recorded on other ardeids from Africa (Gaud 1981a).This is the first record of the genus Ardeialges on N. nycticorax and the first record of this genus in Brazil.Remarks.This feather mite represents an undescribed pterolichine genus, which is recognized by a set of unique characteristics: absence of setae f2, coxal fields I and II sclerotized and by the absence of setae e1 on females.This is the first record of feather mites from P. obscura.
Remarks.This species seems most similar to the genus Tetraolichus by the absence of the setae f2 (Mironov et al. 2010).This feather mite could not be placed in any known pterolichine genus.This is the first record of a feather mite from A. cujubi.Remarks.This mite has the appearance of Pterolichus and also seems similar to Tetraolichus because of the absence of the hysterosomal setae f2; however, this mite does not fit any known pterolichine genus.This is the first record of feather mites on both P. tomentosa and C. blumenbachii.Remarks.This species also lacks setae f2 and is similar to Tetraolichus and represents a new species that probably belongs to the same genus as the undescribed species recorded above.This is the first record of feather mites from C. fasciolata.Remarks.This mite also represents an undescribed species.This is the first record of feather mites on F. xanthopterygius.Remarks.This mite species represents an undescribed species from the mesomexicana species group (Atyeo and Pérez 1988;Mironov and Pérez 2003).This is the first record of the genus Neorhytidelasma on A. ararauna.
Remarks.This mite species also represents an undescribed species from the mesoxicana species group (Atyeo and Pérez 1988;Mironov and Pérez 2003).This mite is most similar to Neorhytidelasma sp.A reported above.This is the first record of the genus Neorhytidelasma on A. hyacinthinus.Remarks.This species also belongs to the mesomexi cana species group (Atyeo et al. 1988;Mironov and Pérez 2003) and is very close to N. mesomexicana Atyeo et al. 1988, which was described from P. holochlorus (Sclater, 1859) (Atyeo and Pérez 1988;Atyeo et al. 1988).This is the first record of the genus Neorhytidelasma on P. leucophthalmus.Remarks.This species is virtually undistinguishable from P. obtusus and might indeed be that species because the feathers were collected from captive birds, often kept with Old World Galliformes.Therefore, our record might not represent a natural association.Remarks.The genus Pterolichus is until now reported exclusively from galliform birds (Atyeo and Gaud 1992;Mironov et al. 2010).Coragyps atratus is the commonest necrophagous bird found in Brazil and could possibly have acquired these mites by foraging on dead phasianid birds, common in rural and urban areas.Therefore, although this is the first record of Pterolichidae on a bird of the family Cathartidae, it may not represent a natural association.

DISCUSSION
All new pterolichine genera were recorded on birds of the family Cracidae, from a bird order (Galliformes) that is known to harbor an exuberant variety of pterolichid genera around the world (Atyeo and Pérez 1991;Atyeo 1992).The only two records of feather mites on cracids in Brazil are also from the family Pterolichidae: Tetraol ichus forficula and Xoloptes minor, both described from the cracid Ortalis squamata (Trouessart and Neumann 1888), although these mite identifications have been reported as questionable (Mironov et al. 2010).
In a survey of feather mites of birds from Canada, Galloway et al. (2014) found a higher number of analgoidean species in comparison with pterolichoideans (94 vs. 39 species), based on exploration of a similar number of passeriform and non-passeriform birds (47 vs. 58).Concerning only non-passerines, those authors recorded a slightly higher number of analgoidean mites than pterolichoideans (52 vs. 42 mite species).But the composition of non-passerines in that paper was quite different with a large number of aquatic bird orders (i.e., Suliformes, Anseriformes, and Charadriiformes) from our study.Many aquatic birds, hummingbirds, and passerines host a higher diversity of analgoidean feather mites living on the flight feathers, such as the families Proctophyllodidae, Alloptidae, and Avenzoaridae (Gaud and Atyeo 1996).On the other hand, in terrestrial nonpasserine birds, which made up the majority of the birds in our study, the superfamily Pterolichoidea is usually more diverse in this microhabitat (e.g., families Crypturoptidae, Falculiferidae, Gabuciniidae, and Pterolichidae).
Other records presented here are also noteworthy, such as the records of Paddacoptes talpacoti (Dermationidae), Hyperaspidacarus tridentatus (Falculiferidae), Pedroso and Hernandes | Feather mites from non-passerine birds in Brazil and Micralges steganonotus (Analgidae) on Columbina talpacoti (Columbidae).A previous study (Moraes et al. 2011) assessed the mite fauna from 51 individuals of this host from São Paulo state, but none of the aforementioned mites were observed.This may indicate a scattered pattern of distribution of these mites on different populations of C. talpacoti in Brazil.
The following records also deserve mention: the first record of the genus Pseudogabucinia (Kramerellidae) on a bird of the family Ardeidae (Syrigma sibilatrix), and the first record of the genus Paralgopsis (Pyroglyphidae) on Crax fasciolata (Galliformes).Paralgopsis had previously been recorded only from inside the quills of psittaciform birds (Gaud 1968b;Jardim et al. 2012), and our record of this genus on a galliform could be the result of contamination, because it was collected from a captive bird.However, information about feather mites from this bird species is scarce in Brazil and requires further investigation.
The records and specimens reported here will provide the material not only for future taxonomic and descriptive research, but also for comparative studies of interpopulation, geographic, and host variation of feather mites.The large number of undescribed genera and species clearly reflects the high proportion of unknown feather mites to be discovered in Neotropical birds and reinforces the need for more taxonomic studies with these mites, especially in Brazil with its rich avifauna.

ACKNOWLEDGEMENTS
We are grateful to Dr. Angelo Pires do Prado (in memorian) and to his student and main collector of the material used in this work, the M.Sc.David Vilas Boas-Filho, as well as his team of students from the University of Campinas (Unicamp), Brazil.We also wish to acknowledge all other collectors: Yoshika Oniki, Edwin O. Willis (in memorian), Carlos O.A. Gussoni, Leopoldo O.F. Bernardi, Fabio A.F. Jacomassa, Gustavo O. Almeida, and Matheus H. Gabriel.Also, thanks to the biologist Rachel B. Santos from the Airport of São Paulo (Congonhas) for donating several dead specimens of birds for study.We are also thankful to Vitor Q. Piacentini (Academy of Natural Sciences of Drexel University, Ornithology Department, Philadelphia, USA) for confirming the occurrence of several bird species.This study was funded by the FAPESP, Sao Paulo Research Foundation (2011/50145-0) and (2016/11671-1).LGAP acknowleges a scholarship from the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES).
ContinuedPedroso and Hernandes | Feather mites from non-passerine birds in Brazil
those, only 26 could be identified to species.Seventy eight mite species represent both first-time associations with corresponding bird host and new records for Brazil; nine species (six of which are already named species) represent first-time records for Brazil but not for the host bird; and four mite records constitute new host associations but not new occurrences of that mite species in Brazil (Table