Two new genera and five new species of the feather mite subfamily Proctophyllodinae (Astigmata: Proctophyllodidae) from suboscine birds in Brazil

Two new genera and five new species of feather mites of the family Proctophyllodidae are described from passerine birds from Brazil: Tyranniphyllodes pitangi gen. n., sp. n. from Pitangus sulphuratus (Tyrannidae); Atrichophyllodes delalandi gen. n., sp. n. from Corythopis delalandi (Tyrannidae); A. mentalis gen. n., sp. n. from Dysithamnus mentalis (Thamnophilidae); Anisophyllodes candango sp. n. from Elaenia chiriquensis (Tyrannidae); and Platyacarus sittasomi sp. n. from Sittasomus griseicapillus (Dendrocolaptidae). The discovery of these taxa might give data for a better understanding of the evolution of the family Proctophyllodidae in general and the dispersion of these mites on passerines in South America in paricular.


Introduction
Proctophyllodidae, a family of feather mites mostly associated with passerines (Passeriformes) and hummingbirds (Apodiformes), is the most species-rich family within the superfamily Analgoidea and comprises three subfamilies: Proctophyllodinae, Pterodectinae, and Rhamphocaulinae (Gaud and Atyeo 1996). The Neotropical region has one of the richest bird faunas in the world, hosting many basal lineages of Passeriformes (Ericson et al. 2002). In this sense, one should expect to find numerous early derivative proctophyllodid mites in this part of the world. Exploring this fauna is thus very important for understanding the evolution and cospeciation processes of this family. However, the diversity of proctophyllodids living on passerine birds on the entire continent of South America, and more specifically in Brazil, is poorly known. Data on this family are scattered among a dozen systematic papers, and most of these deal with species of the subfamily Fiocruz, Rio de Janeiro, Brazil; ZISP, Zoological Institute of the Russian Academy of Sciences, St Petersburg, Russia.

Differential diagnosis
The new genus, Tyranniphyllodes, is most closely similar to the genus Platyacarus in the following features: in males, opisthosomal lobes short and wide, terminal lamellae relatively small (shorter than distance h3:h3), legs III and IV are subequal in size and not hypertrophied, their segments not modified; in both sexes, the prodorsal shield with bluntangular posterior margin. Tyranniphyllodes gen. n. can be readily separated from Platyacarus by the following combination of features: in both sexes, hysteronotal setae d2, e2 and trochanteral setae sRIII are absent; in males, sheath of aedeagus is bifurcated apically, and opisthogastral shield is H-shaped. In males of Platyacarus, the sheath of the aedeagus is ensiform like the aedeagus proper, opisthogastral shield is absent and area between genital apparatus and anal suckers bears two pairs of small sclerites. It is necessary to stress that the absence of setae sRIII is a unique feature of Tyranniphyllodes gen. n. differentiating it from all other genera of Proctophyllodinae; it is also interesting to note that the H-shaped opisthogastral shield in this genus resembles that in the genus Joubertophyllodes Atyeo and Gaud, 1971 and the pinnatus species group of the genus Proctophyllodes.

Description
Both sexes. Moderately elongated proctophyllodids. Prodorsal shield developed in median part of prodorsum, strongly narrowed in anterior end, with short and acute posterior angles, with blunt-angular posterior margin. Scapular setae si and se arranged in transverse line. Humeral shields rudimentary, represented only by small plate lateral to epimerites III and dorsally not developed. Setae c2 situated ventrally on striated tegument. Epimerites I connected by posterior ends into narrow U. Epimerites IIa straight, not bent to median line. Vertical setae ve, lateral hysteronotal setae d2 and e2 absent. Solenidion s1 of genu I slightly longer than v3 on tarsi I, situated in median part of the segment (Figure 3). Setae wa anterior to setae la and ra on legs I and II (Figures 3, 4). Femora II with blunt-angular outer protrusion, other segments of legs I and II without processes and other modifications. Seta sR absent on trochanters III ( Figure 5).
Male. Prodorsal shield completely or partly split into anterior and posterior pieces by transverse band of soft tegument (Figures 1, 9); scapular setae se and si situated on posterior piece. Opisthosomal lobes scarcely expressed; posterior margin of opisthosoma between setae h3 concave and with pair of relatively small tongue-shaped terminal lamellae; dorsal surface of lamellae ornamented. Supranal concavity well developed. Setae h3 setiform, much shorter than macrosetae h2. Setae h1 posterior to level of setae ps2. Coxal fields I-IV open, without extensive sclerotized areas. Genital organ at level of trochanters IV; genital arch large, with widely divergent branches; sheath of aedeagus longer than arch, and bifurcated apically. Opisthogastral shield H-shaped, its transverse branch with median extension directed anteriorly and fused to base of genital sheath, setae g situated on its transverse branch ( Figure 11). Genital acetabula at level of genital arch apex. Anal suckers Figures 1, 2. Tyranniphyllodes pitangi gen. n., sp. n., male: dorsal (1) and ventral (2) views. os, opisthogastral shield.

Etymology
Contraction of the host family, Tyrannidae, and Proctophyllodes, the type genus of the mite family Proctophyllodidae.

Etymology
The specific epithet derives from the generic name of the host and is a noun in the genitive case.

Differential diagnosis
The new genus, Atrichophyllodes, is similar to Nycteridocaulus Atyeo (1966) in the following characters in males: terminal lamellae are wide and short, almost semicircular, genital arch is much wider than long, legs III and IV subequal in size and not hypertrophied; epimerites I free. Atrichophyllodes gen. n. is readily distinguished from the latter genus by the absence of idiosomal setae d2 and e2, and by the branches of genital apparatus forming a low arch (rather than a bat-shaped structure, as in Nycteridocaulus). This genus also resembles Tyranniphyllodes gen. n. (see above) in the absence of hysterosomal setae d2 and e2; however, it differs from Tyranniphyllodes gen. n. by having a sword-shaped aedeagus sheath, a trochanteral seta sRIII, a uniquely shaped prodorsal shield (in both sexes), and by males lacking an opisthogastral shield.

Description
Both sexes. Moderately elongated proctophyllodids. Prodorsal shield entire, covering nearly the entire prodorsum, with acute antero-lateral extensions, posterior angles rounded. Scapular setae si and se arranged in transverse line. Humeral shields well-developed dorsally, encompassing bases of setae c2 and cp. Epimerites I free, their posterior parts close to each other. Epimerites IIa L-shaped, bent to median line. Hysteronotal setae d2 and e2 absent. Solenidion s1 of genu I slightly longer than v3 on tarsi I, situated in median part of the segment ( Figure 17). Setae wa anterior to setae la and ra on legs I and II (Figures 17,18). Segments of legs I and II without processes or other modifications.
Male. Scapular setae se and si situated on prodorsal shield. Opisthosomal lobes short and wide; posterior margin of lobes with semi-circular terminal lamellae. Supranal concavity well expressed. Setae h3 setiform, much shorter than macrosetae h2. Setae h1 anterior to the level of setae ps2. Coxal fields I-IV open, without extensive sclerotized areas. Genital organ at level of trochanters IV; genital arch large and low, sheath of aedeagus approximately twice longer than arch. Adanal shields present, situated between anal suckers and setae ps3, weakly connected to each other anterior to anal opening. Genital acetabula at level of genital arch apex, acetabular plates surrounding these acetabula present or absent. Anal suckers cylindrical, corolla dentate. Pregenital and paragenital plates absent. Opisthoventral shields not developed. Legs III and IV subequal, not hypertrophied. Tarsus IV without apical claw-like process, modified setae d and e buttonlike.
Female. Lobar region of opisthosoma clearly separated from hysterosoma, opisthosomal lobes well developed, with long terminal appendages. Anterior hysteronotal and lobar shields separated by narrow band of striated tegument. Supranal concavity absent. Macrosetae h2 setiform, long. Epigynium horseshoe-shaped, large. Translobar sclerites present. Legs III and IV subequal in size; segments without modifications; solenidia w of tibiae III and IV subequal in length.

Etymology
Contraction of a (without, G.), tricho (hair, G.), and Proctophyllodes to point out the absence of dorsal setae d2 and e2.

Etymology
The specific epithet was directly taken from the specific name of the type host and is a noun in the genitive case.

Etymology
The specific epithet was directly taken from the specific name of the type host and is a noun in apposition.
Genus Anisophyllodes Atyeo, 1967 The genus Anisophyllodes was originally based on a single species, Anisophyllodes pipromorphae Atyeo, 1967 collected from Mionectes (5 Pipromorpha) oleagineus (Lichtenstein, 1823) (Tyrannidae) from Trinidad and Tobago (Atyeo 1967). Atyeo (1969) later transferred Alloptes intermedius described by Trouessart and Neumann (1888), on Elaenia martinica (Linnaeus, 1766), to Anisophyllodes. The latter species was also reported by Cerny and Lukoschus (1975) from E. flavogaster (Thunberg, 1822). The main diagnostic features separating this genus from other proctophyllodine genera are as follows: in both sexes, epimerites I free with diverging posterior tips; in males, opisthosoma attenuate to posterior end and with truncated opisthosomal lobes usually separated by slitlike terminal cleft; genital arch and aedeagus short, situated at the level of trochanters IV; setae g and ps3 arranged in a longitudinal rectangle; paired or unpaired pregenital plates are present; adanal shields present. Dimorphism of males was also noted for this genus: in heteromorph males, the opisthosomal lobes bear short and wide terminal lamellae and legs IV are thicker than legs III; in homeomorph males, the opisthosomal lobes lack terminal lamellae and legs III, IV are subequal in size. It is necessary to note that the original diagnosis (Atyeo 1967) stated that dorsal setae e1 are absent in Anisophyllodes; however, this character actually varies among species. In A. intermedius and in a new species described below these setae are present.

Etymology
The specific epithet derives from candango (African origin), a common name meaning those who live, but were not actually born in Brasilia (the capital of Brazil), the type locality of this mite species; the name is a noun in apposition.
Genus Platyacarus Kudon, 1982a Twelve known species of the genus Platyacarus were described in a series of four papers by Kudon (1982aKudon ( , 1982bKudon ( , 1982cKudon ( , 1982d. Representatives of this genus occur only on birds of the family Dendrocolaptidae. They were recorded on these hosts across Central and South America (Kudon 1982e): Brazil (17 host species), Ecuador (three), Guyana (one), Colombia (one), Venezuela (three), Panama (two), French Guyana (one), Surinam (one), and Costa Rica (one). Mites of Platyacarus may be distinguished from other proctophyllodine genera by the following combination of characters: setae ve absent; epimerites I normally fused into a U; setae f2 absent; in males, opisthosomal lobes short, provided with terminal lamellae of triangle or tongue-shaped form; aedeagus is ensiform; genital arch is small, with strongly divergent branches; adanal shields represented by one or two pairs of small sclerites; setae g and ps3 arranged into longitudinal rectangle; legs III-IV subequal.

Differential diagnosis
Platyacarus sittasomi sp. n. may be referred to the oligolaccius species group (Kudon 1982a) by most features, except for the structure of the prodorsal shield in both sexes (Figures 41,49) and primary spermaduct in females ( Figure 51). It differs from the two species previously included in this group, P. oligolaccius Kudon, 1982a andP. dontocoronius Kudon, 1982a, by the following characters: in both sexes, the lateral margins of the prodorsal shield without incision around scapular setae se; in males, the aedeagus is short (25) extending beyond genital setae g by the distal third only, and anterior margin of hysteronotal shield straight; in females, the terminal cleft is narrow (60-63622-27), approximately three times longer than wide, and the primary spermaduct is not expanded in the proximal part. In both sexes of two formerly known species, the prodorsal shield has deep lateral incisions around setae se; in the males, the aedeagus is much longer (50 in P. oligolaccius and 41 in P. dontocoronius), almost extending to the level of anal setae ps3, and anterior margin of hysteronotal shield is concave; in the females, the terminal cleft is approximately as long as wide (47 long in P. oligolaccius and 38 in P. dontocoronius), and the primary spermaduct is abruptly expanded near the head of spermatheca.

Etymology
Specific epithet derives from the generic name of the type host and is a noun in the genitive case. Figures 47,48. Platyacarus sittasomi sp. n., male: lobar region, dorsal (47) and ventral (48) views. ias, inner anal shield; oas, outer anal shield.