Mites of the genus Neharpyrhynchus Fain (Acariformes, Harpirhynchidae) from Neotropical birds

Abstract Three new species of parasitic mites of the genus Neharpyrhynchus Fain (Acariformes, Harpirhynchidae) are described from Neotropical birds: Neharpyrhynchus chlorospingus sp. n. from Chlorospingus pileatus (Passeriformes, Emberizidae) from Costa Rica, Neharpyrhynchus mironovi sp. n. from Dacnys cayana (Passeriformes, Thraupidae) and Neharpyrhynchus tangara sp. n. from Tangara cayana (Thraupidae) both from Brazil. Neharpyrhynchus trochilinus (Fain) is recorded from 3 new host species of the family Trochilidae (Apodiformes), Panterpe insignis and Eugenes fulgens from Costa Rica, and Amazilia lactea from Brazil. Emended diagnosis of the genus and a key to species are provided; all records of Neharpyrhynchus species are summarized.


Introduction
Mites of the genus Neharpyrhynchus Fain (Acariformes, Harpirhynchidae) are permanent and highly specialized parasites of birds, as is the case for all other representatives of the subfamily Harpirhynchinae (Bochkov 2008). Th e subgenus Neharpyrhynchus Fain was established by Fain (1972) in the genus Harpirhynchus Mégnin. Later, Fain (1995) proposed full generic status for Neharpyrhynchus and simultaneously revised this genus, which included fi ve species at that time. Th e life-cycle of these mites was described by Moss et al. (1968) as exemplifi ed by N. novoplumaris Moss et al. Th e last revision of the genus Neharpyrhynchus was recently provided by Martinu et al. (2008). To date this genus includes 11 species belonging to fi ve species groups established in that revision: baile (3 species), hippolae (3 species), pilirostris (1 species), plumaris (3 species), and squamiferus (1 species). In our opinion, however, there are no characters discriminating the pilirostris and hippolae species groups. Moreover, such diff erential characters were not provided even by Martinu et al. (2008) and in their key, N. pilirostris is placed in the same couplet with N. pari, a species from the group hippolae. We, therefore, include all species of the hippolae group in the pilirostris group.
Most species of the genus are known from European passerines and only two species are known from Neotropical birds, N. baile Bochkov et al. from Turdus leucomelas (Passeriformes, Turdidae) (Bochkov et al. 2007) and N. trochilinus (Fain) from hummingbirds (Fain 1972(Fain , 1995. In this paper, we describe three new species from Neotropical birds and provide new records for N. trochilinus. Additionally, an emended diagnosis of the genus and a key to its species are given. Th e diagnostic characters of species groups we recognize in the genus Neharpyrhynchus and all records of these mites are given in Tables 1 and 2, respectively.

Present paper
N. trochilinus (Fain, 1972) Hummingbird Trochilidae (Apodiformes) South America (?) Fain (1972Fain ( , 1995 "  (Fain, 1972) *Temenuchus pagodarum (Gmelin) Sturnidae (Passeriformes) India (?) Fain (1972Fain ( , 1995 with a camera lucida, and measurements were taken using a calibrated ocular micrometer. Drawings were made by A. V. Bochkov. In the species description, names of the leg and idiosomal setae follow Grandjean (1939Grandjean ( , 1944 as adapted by Kethley (1990). Names of the palpal setae follow Grandjean (1946) as adapted by Bochkov (2008). All measurements are given in micrometers (μm) and were made according to the standard method (Bochkov et al. 2007): body length = maximum length of the body up to the anterior extremity of the palpal tibia; body width = maximum width taken at whatever level it occurs; gnathosomal length = length taken ventrally from the gnathosomal base to the anterior extremity of the palpal tibia; gnathosomal width = maximum width taken at whatever level it occurs; length of dorsal shield = maximum length, measured in the median line of the shield; and width of dorsal shield = maximum width taken at whatever level it occurs. Th e scientifi c names of birds follow the checklist of Clements et al. (2010).  Fritsch, 1954: 193, fi gs 11, 12, by original designation.
Males. Gnathosoma as in female. Idiosoma rhomboid in outline. Anterior sclerotized area of propodosoma absent. Dorsal shield well developed, occupying most part of dorsal idiosomal surface. Genital opening situated in middle part of dorsal shield. Genital setae 3 pairs. Penis originating behind genital opening. Situations of dorsal idiosomal setae typical for subfamily. Setae 3a present. Legs I and II well developed, without basal lobes, with 5 articulate segments each. Setation of tibia and tarsi as in females, three other proximal segments with setae. Legs III with two segments, both bearing setae; legs IV with one segment.
Remarks. Th e sclerotized area on the anterior part of the propodonotum was incorrectly named as the propodosomal (=propodonotal) shield by Martinu et al. (2008). In Harpirhynchidae, actually, the true propodonotal shield is fused with the hysteronotal shield or its remnants to form a common large shield, which can be referred to as the dorsal shield (Bochkov 2008). Th e sclerotized area in the anterior part of the propodosoma situated anterior to the dorsal shield is formed de novo and probably helps to fi x the subcapitulum when the female attaches to a feather (Fig. 1). Anterior region of propodonotum covered by short irregularly situated folds, without scales or tubercles (Fig. 6A). Dorsal shield entire, 165 long in midline (160-170), 300 at maximum width (300-330) ( Fig. 2A). Anterior and posterior margins of dorsal shield widely concave. Ventral surface of idiosoma with indistinct transverse striations, without scales or verrucosities (Fig. 2B). Setal lengths: vi, ve, and si -all distinctly barbed, subequal in length, 150-160; se, c2, and 1a -all smooth, 10-12; h1 whip-like, 250 (230-260); 1b smooth, about 40; 3a present, about 20. Base of legs I with distinctly developed and slightly attenuated fl eshy lobe; base of legs II with moderately developed rounded lobe. Leg I with 2 articulated segments (Fig. 3B). Leg II with 2 articulated segments (Fig. 3C). Legs III, IV with one segment, each bearing 4 (more rarely 5) long setae. One ventral seta of leg III and 2 ventral seta of leg IV 100-120 long, about half the length of other setae situated dorsally or dorsoterminally, 200-250 long. Male. Unknown. Etymology. Th e species name is derived from the generic name of the host and is a noun in apposition.

Neharpyrhynchus chlorospingus
Diff erential diagnosis. Th is species belongs to the "plumaris" species group including three species, N. plumaris (Fritsch), N. novoplumaris (Moss et al.), and N. spinus Martinu et al. (Martinu et al. 2008). In females of this group, legs I and II consist of the two articulated segments, palpal setae vF are smooth, the anterior region of the propodonotum is covered by short irregular striations, and setae 3a are present. Within this group, it is close to N. novoplumaris described from Certhia familiaris Linnaeus (Passeriformes, Certhiidae) [type host] and Cardinalis cardinalis (Linnaeus) (Passeriformes, Cardinalidae) from USA (Moss et al. 1968). In females of both of these species setae dG are about half the lenth of l"G. In the other two species of the genus, setae dG and l"G are subequal. Females of N. chlorospingus diff er from N. novoplumaris by the following characters. In N. chlorospingus, palpal setae dF are slightly shorter than l"G, setae se and c2 are about four times shorter than 1b, the posterior margin of the dorsal shield is widely concave. In N. novoplumaris, palpal setae dF are slightly longer than l"G, setae se and c2 are subequal or only slightly shorter than 1b, the posterior margin of the dorsal shield is widely convex.  Type deposition. Holotype and 10 paratypes deposited in the MZUSP, 6 paratypes in the ZISP, 2 paratypes in the UMMZ, and 2 paratypes in the IPCR. Alcohol preserved paratypes deposited in the MZUSP and ZISP.
Male. Unknown. Etymology. Th e species is named in honour of the prominent Russian acarologist Dr. Sergey V. Mironov (ZISP).
Diff erential diagnosis. It is close to species of the group "pilirostris". In all these species, setae vF are smooth, only two articulated segments on legs I and II are present, and setae 3a are absent. Among species of this group, N. mironovi is close to N. pari by the presence of four setae on leg III and by irregular ornamentation of the anterior part of the propodosoma. Th e new species diff ers from N. pari by the following characters. In females of N. mironovi, the palps are distinctly infl ated dorsally, the ornamentation of the anterior part of the propodonotum is scale-like and present only in the posterior half of this region, setae c2 are 50-60 long. In N. pari, the palps are moderately infl ated dorsally, the anterior part of the propodonotum is fully ornamented by verrucosities and setae c2 are 5-6 long. Type deposition. Holotype and 10 paratypes deposited in the MZUSP, six paratypes in the ZISP, 2 paratypes in the UMMZ, and 2 paratypes in the IPCR. Alcohol preserved paratypes deposited in the MZUSP and ZISP.
Male. Unknown. Etymology. Th e species name derives from the generic name of the host and is a noun in apposition.
Diff erential diagnosis. Th is new species is closest to N. mironovi and diff ers by the following characters. In females of N. tanagra, setae c2 are 8-12 long, the posterior margin of the dorsal shield is almost straight. In N. mironovi, setae c2 are 50-60 long, the posterior margin of the dorsal shield is widely concave. Both species are collected from the hosts belonging to the family Th raupidae. (Fain, 1972) Fig. 6D Harpyrhynchus (Neharpyrhynchus) trochilinus Fain, 1972: 55. Neharpyrhynchus trochilinus Fain 1995Bochkov et al. 2007: 38;Martinu et al. 2008: 207, fi g. 1 [types in IRSNB]. Hosts and distribution. This species was briefly diagnosed from both sexes collected from an unidentified species of hummingbird (Trochilidae) that originated from South America (exact locality unknown) and died in the Zoo of Antwerp (Belgium) during its quarantine. Later on, Fain (1995) provided the full description of this species based on the type specimens and newly obtained specimens from Chrysolampis mosquitus (Linnaeus) (Trochilidae) that also originated in South America (without exact locality) and died in the Zoo quarantine. The trochilids, Panterpe insignis, Eugenes fulgens (Costa Rica), and Amazilia lactea (Brazil) are new hosts for this mite species. It is probable, that this species is associated exclusively with hummingbirds and is widely distributed on representatives of this host family.

Neharpyrhynchus trochilinus
Remarks. Th e longitudinally subdivided dorsal shield of this species is an artifact sometimes induced by the mite mounting. In this species, actually, the dorsal shield is entire. It diff ers from the closely related N. baile Bochkov et al. by the following characters. In females of N. trochilinus, setae dF, dG, and l"G are subequal, legs III and IV with 5-6 setae each, setae si and se 25-35 long. In N. baile, setae dF is about 1.5 times longer than dG and l"G, legs III and IV as a rule with 4 setae each, setae si and se are 6-12 long.

Keys to species of the genus Neharpyrhynchus Fain (females)
(based on Martinu et al. 2008) 1 Anterior margin of propodonotum without ornamentation or just with few striations.