On Identity of the Type Species of The

Acarina 17 (1): 89–100 © Acarina 2009

On identity of the type species of the feather mite genus Anhemialges Gaud, 1958 (Astigmata: Analgidae), with the description of a new genus of the analgid subfamily Megniniinae S. V. Mironov Zoological Institute, Russian Academy of Sciences, Universitetskaya quay 1, Saint Petersburg 199034, Russia; e-mail: [email protected]

ABSTRACT: This paper demonstrates that the genus Anheminalges Gaud, 1958 (Analgidae: Analginae) is based on the misidentified type species “Anhemialges longipes (Trouessart, 1899)”. As indicated originally (Gaud 1958) and subsequently, this species commonly occurs on the barn swallow, Hirundo rustica (Passeriformes), and some other hirundinid species worldwide. Examination of the holotype specimen of Megninia longipes Trouessart, 1899 described from the horned screamer, Anhima cornuta (Anseri- formes), from South America revealed that it does not correspond to the species occurring on hirundinids and belongs to the subfamily Megniniinae rather than Analginae. The species actually involved in the misidentification, Anhemialges gaudi sp. n. (=A. longipes sensu Gaud, 1958, non Trouessart, 1899), is fixed here as the type species of the genus Anhemialges (Article 70.3.2, ICZN). A new genus, Anhimomegninia gen n., is established based on Megninia longipes. The genus Anhemialges is provided with an emended diagnosis and the species Anhimomegninia longipes (Trouessart, 1899), comb. n. is redescribed based on the holotype.

KEY WORDS: feather mites, Analgidae, Anhemialges, Anhimomegninia, new taxa, systematics, taxonomic correction

Introduction per Gaud (1953) reported this species from H. The feather mite genus Anhemialges Gaud, smithi in Oubangui-Chari (recently the Central 1958 (Analgidae) is a morphologically well out- African Republic). It is necessary to stress that lined genus of the subfamily Analginae (Gaud Trouessart (1899) actually described M. longipes 1958; Gaud and Till 1961; Gaud and Atyeo 1996). from the horned screamer Anhima cornuta (Lin- As for most Analgidae, representatives of this ge- naeus) (Anseriformes: Anhimidae) in Guyana. It nus live on down feathers and on the downy parts is impossible to guess why Gaud (1958) identified of body covert feathers (Mironov 1999). This ge- the mite species he found on the barn swallow as nus currently includes only four described species M. longipes. The original description (Trouessart from passerines of the families Hirundinidae and 1899) was very brief and without figures, and Zosteropidae. However, taking in account that Gaud probably relied on the redescription of this Atyeo (in: McClure and Ratanaworabhan 1973) species (Bonnet 1924), which included figures of reported unidentified Anhemialges species from the posterior end of opisthosoma and tarsus III of 23 hosts of eight passerine families and even one male. The finding of M. longipes on a host, which species from a piciform host in South-Eastern is phylogenetically very distant from the type host Asia, it is possible to suggest that this genus is and taken from a different continent, also did not probably as diverse as Analges Nitzsch, 1818, disturb Gaud. It is only possible to suggest that he most the species-richest genus in Analgidae, and considered the record of M. longipes from the also has a very wide host range. horned screamer as the result of accidental con- Investigation of the type material of the type tamination, because Trouessart collected feather species of this genus, Anhemialges longipes mite samples from museum skins. In establishing (Trouessart, 1899), revealed confusion in the iden- the new genus Anhemialges, Gaud (1958) even tity of the type species of the genus Anhemialges did not mention the type host of M. longipes. (see below for material and methods). It appeared Further, Gaud and Mouchet (1959) reported that “A. longipes” in sense of Gaud (1958) and A. longipes from the three more species of swal- subsequent authors does not correspond to the lows, H. nigrita Gray G.R., Cecropis abyssinica type specimen, and the genus Anhemialges was unitatis Sclater et Mackworth-Praed, and Psalido- actually based on a misidentified and undescribed procne pristoptera petiti Sharpe et Bouvier, in mite species. Cameroon. It is interesting that these authors indi- Gaud (1958) established the genus Anhemial- cated Hirundo senegalensis Linnaeus as the type ges in the course of systematic investigations of host instead of Anhima cornuta in their paper. Ap- feather mites in Morocco. Originally Gaud includ- parently they based their statement about the host ed in this genus a sole species, Megninia longipes species on the paper of Bonnet (1924), who re- Trouessart, 1899, which he collected in Morocco vised the genus Megninia Berlese, 1881 and listed from the barn swallow Hirundo rustica Linnaeus for M. longipes the three quite different hosts: (Passeriformes: Hirundinidae). In a bit earlier pa- Anh. cornuta, Streptoprocne zonaris Shaw (Apod-

89 S.V. Mironov iformes: Apodidae) and H. senegalensis. No hemialge longipes sensu Gaud, 1958, non Troues- doubts that the first host was taken from the paper sart, 1999) and fix it here as the type species of the of Trouessart (1899), while two others were added genus Anhemialges. I also provide in the present by Bonnet (1924) based on his own material. The paper an emended diagnosis of the genus Anhemi- two latter records apparently represented mites alges, establish a new genus Anhimomegninia gen. from the genera Cypselalges Gaud et Atyeo, 1991 n. with the type species Megninia longipes Troues- (a genus specific to swifts) andAnhemialges sensu sart, 1899 and redescribe the latter species as Anhi- Gaud, 1958, respectively. Thus, the “redescrip- momegninia longipes (Trouessart, 1899) comb. n. tion” of M. longipes given by Bonnet (1924) was MATERIAL AND METHODS apparently based on thee different species. In their review of feather mites living in sub- The type specimen of Megninia longipes was loaned from the Muséum national d’Histoire na- Saharan Africa, Gaud and Till (1961: Fig. 108A, turelle (Paris, France); other specimens used in the B) for the first time illustrated the mite from hirun- present study belong to the collection of the Zoo- dinids that they considered to be A. longipes. In logical Institute of the Russian Academy of Sci- that work they also mentioned that the genus An- ences (Saint Petersburg, Russia). hemialges included two species. Unfortunately The diagnoses and descriptions are given in these authors did not name the second species, and the modern formats used for analgid mites (Gaud it is only possible to guess that they probably and Atyeo 1991; Mironov and Galloway 2002a, meant Megninia aestivalis subintegra Berlese 2002b). General morphological terms, leg and idi- (1883), which was described from the martins osomal chaetotaxy follow Gaud and Atyeo (1996). Delichon urbicum (Linnaeus) and Riparia riparia All measurements are in micrometres (µm). Meas- (Linnaeus) (Hirundinidae) in Italy and was illus- urement standards for particular structures are as trated quite well (Berlese 1883: fasc. 26, No 1). follows: (i) idiosoma is measured from its anterior Since the 1960s, a number of subsequent au- margin to lobar apices in males and to posterior thors reported A. longipes (sensu Gaud, 1958) margin of opisthosoma in females, (ii) prodorsal from various hirundinids, mainly from common shield length is the greatest length from anterior European species (Černy 1967, Arutunjan and end to level of posterior angles and width is the Mironov 1983; Mironov 1983, 1996; Kolarova greatest width of its posterior part; (iii) hystero- and Mitov 2008). In the review of suprageneric soma is measured from the level of the sejugal fur- feather mite taxa of the World, Gaud and Atyeo row to lobar apices in males and to the posterior (1996) once more gave the drawings of the male margin of opisthosoma in females; (iv) hysterono- and female of A. longipes based on specimens tal shield length is the greatest length from the an- from some hirundinids. terior margin to lobar apices; width is measured at Investigation of the holotype specimen of the anterior margin; (v) distance between setae of Megninia longipes deposited in the Trouessart col- different pairs is the shortest distance between the lection (Muséum national d’Histoire naturelle, transverse levels formed by setae of correspond- Paris, France, slide 28i9) has shown that this spe- ing pairs. cies does not correspond to the species that com- Depositories of examined material: TRT — monly occurs on Hirundo rustica. Moreover, the the Muséum national d’Histoire naturelle, Paris, specimen of Megninia longipes described by France; ZISP — the Zoological Institute of the Trouessart (1899) even does not belong to the sub- Russian Academy of Sciences, Saint Petersburg, family Analginae. It represents a quite distinctive Russia. Systematics and scientific names of birds species of the subfamily Megniniinae and deserves follow Dickinson (2003). to be treated as a separate genus related to Megnin- iella Gaud, 1958. Thus, Gaud (1958) actually mis- SYSTEMATICS identified a mite species, which he used as the type Family Analgidae Trouessart et Mégnin, 1884 species for the genus Anhemialges. This species Subfamily Analginae Trouessart et Mégnin, has never been formally described, only its figures 1884 were reproduced in three papers (Gaud and Till Genus Anhemialges Gaud, 1958 1961; Arutunjan and Mironov 1983; Gaud and Gaud 1958: 36; Gaud and Mouchet 1959: Atyeo 1996). Applying Article 70.3.2 of the Inter- 159; Gaud and Till, 1961: 192, Arutunjan and Mi- national Code of Zoological Nomenclature (ICZN ronov 1996: 232; Gaud and Atyeo 1996 (Pt. I): 53, 1999), I describe Anhemialges gaudi sp. n. (=An- (Pt. II): 26.

90 The type species of the feather mites and a new genus of the analgid mites

Type species: Megninia longipes non Troues- shield), flanked laterally by wide adanal mem- sart, 1899 by monotypy (ex Hirundo rustica, Mo- branes stretching from adanal shield to bases of rocco) (=Anhemialges gaudi sp. n. ex H. rustica). setae ps2. Area between anal field and posterior Diagnosis. Both sexes. Medium sized analg- ends of paragenital apodemes covered with large ines. Prodorsal shield occupying median part of gastral shield (paired or unpaired). Anal suckers prodorsum, shaped as narrow trapezium, with pair cylindrical, low, corolla not dentate. of longitudinal median ridges, with pair of acute Legs III hypertrophied, slightly thicker and suprategumental extensions on posterior margin about 1.5 times longer than legs IV. Femur, genu of this shield (Figs. 1A, 2A). Laterocoxal setae and tibia III cylindrical, without apophyses. Tarsus present, setiform. Scapular shields with suprateg- III elongated subequal in length to corresponding umental extension on inner margin. Hysteronotal tibia and slightly curved, setae s and w of tarsus III setae absent: c1, h1. Vertical setae ve present, ru- lanceolate, spiculiform, or blade-like, other setae dimentary. Epimerites I fused into a Y with long of this segment setiform. Tarsus IV subequal in sternum, anterior and central parts of these length to corresponding tibia, conical and curved, epimerites strongly thickened (Figs. 1B, 2B). Fe- with bidentate apical process; seta d button-like, mur I with large hook-like lateral apophysis seta e stick-like, minute, situated on apical process. rounded apically, trochanter I with tooth-like lat- Pretarsi III, IV developed as on legs I, II. eral apophysis opposing to femoral apophysis Males are strongly intraspecifically variable (Figs. 3A, C); femur II with small hook-like lat- in body size and particularly in size of opistho- eral apophysis or without it. Tarsi I, II with cuff- somal lobes, but polymorphism is continuous, like (machete-like) ventral apophyses, tibiae I, II without the contrasting qualitative features ob- with spine-like ventral apophyses. Tarsus I with 8 served in Analges or Hemialges Trouessart et setae (ba, d, e, f, la, ra, wa, s), tarsus II with 7 se- Neumann, 1888 that allows designation of hetero- tae, ventral seta wa absent (Figs. 3A, B, D). morph and homeomorph forms. Male. Idiosoma moderately elongated; lateral Female. Hysteronotal shield absent. Setae e1, margins of opisthosoma slightly attenuate posteri- d1 present or absent (variation occurs among indi- orly, without deep incisions. Opisthosoma bilo- viduals of the same species). Epigynum present, bate; opisthosomal lobes of moderate size, roughly bow-shaped, free from epimerites, distant from rectangular or angular, separated by rectangular or posterior tips of epimerites I, II. Lateral flaps of trapezoidal terminal cleft, which is 2–3 times wid- oviporus not sclerotized. Epimerites IIIa short, not er than opisthosomal lobe width; terminal cleft oc- extending to posterior end of oviporus flaps. cupied by interlobar membrane with deep median Other included species: Anhemialges subinte- incision and short extensions extending beyond ger (Berlese, 1883) from Delichon urbicum (Lin- lobar apices. Lateral membranes absent. Hyster- naeus) (Hirundinidae); A. albidus (Tyrrell, 1882)1 onotal shield with acute or rectangular anterior comb. n. from Tachycineta bicolor (Vieillot) (Hi- angles, anterior margin straight or moderately rundinidae), A. gracillimus (Bonnet, 1924) from concave. Setae c2 situated on humeral shields. Se- Zosterops maderaspatanus (Linnaeus) (Z. mada- tae d2, e2 represented by macrosetae, these setal gascariensis in the original description) (Zoster- pairs far removed from each other and both distant opidae). Only type hosts are listed. from anterior angles of hysteronotal shield: setae Anhemialges gaudi sp. n. d2 situated at level of trochanters III, setae e2 posterior to level of trochanters IV. Setae h3 on lobar Figs. 1–3 apices, setae h2 on lateral margins of lobes. Setae Megninia longipes (non Trouessart, 1899): ps1 on lateral margins of terminal cleft, far ante- Bonnet 1924: 174, fig. 33, 34 (part.); Gaud 1953: rior to level of setae h3. 211; Radford 1953: 207 (part.) [misidentifica- Coxal field III closed, rarely may be open in tion]. mesal angles. Genital apparatus at level of tro- Anhemialges longipes (non Trouessart, 1899): chanters III. Epiandrium present or absent, thin Gaud 1958: 36; Gaud and Mouchet 1959: 159; bow-shaped if present, paragenital apodemes represented by narrow longitudinal bands flanking 1 This species was originally described by Tyrrell (1882) in genital apparatus laterally. Genital shield absent. the genus Megninia Berlese, 1881. Formally, Černý (1967) for the first time used the combination Anhemialges albidus, Anal field (ventral area of opisthosoma bearing but it was given in the unpublished museum catalog of the anal opening, anal suckers, setae ps3, and adanal Cuban fauna.

91 S.V. Mironov

A B

Fig. 1. Anhemialges gaudi, sp.n., male holotype. A — dorsal view, B — ventral view.

Gaud and Till 1961: 193, fig. 108A, B; Černý Additional material. 1 male, 1 female (ZISP 1967: 17; Arutunjan and Mironov 1983: 232, fig. 16 653) ex H. rustica, FRANCE, La Roche sur 9, 1–4; Mironov 1996: 255; Gaud and Atyeo 1996 Yon, August, 1962, J. Gaud; 1 female (ZISP (Pt. II): 26, fig. 37A–E; Kolarova and Mitov 2008: 16 654), same host, FRANCE, Dinan, September 94. [misidentification]. 1967, J. Gaud; 1 male (ZISP 16 655), same host, Type material. Male holotype, (ZISP 4481), FRANCE, Angouléme, August 1963, J. Gaud. All 7 males and 6 female paratypes (ZISP 4482 – specimens are identified by J. Gaud as Anhemial- 4493) ex Hirundo rustica Linnaeus (Passerifor- ges longipes. mes: Hirundinidae), RUSSIA, Kaliningrad Pro- Description. Male (Holotype, range of mea- vince, Rybachy, 55°05’ N, 20°44’ E, 27 May surements for 8 paratypes in parentheses). Idioso- 1982, S.V. Mironov. ma, length × width, 370 × 183 (365–410 ×

92 The type species of the feather mites and a new genus of the analgid mites

A B

Fig. 2. Anhemialges gaudi sp. n., female paratype. A — dorsal view, B — ventral view.

180–215), length of hysterosoma 268 (265–305). and forms short extensions on lobar apices, inci- Gnathosoma without spines on lateral margins of sion in interlobar membrane angular, 26 (20–27) subcapitulum. Prodosal shield: narrowly trapezoi- long. Setae d1, e1 present. Setae e1 posterior to dal, length 75 (70–85), width 78 (75–80), posterior hysteronotal gland openings gl. Setae ps1 at level margin concave, with pair of acute suprategumen- of setae h2 or slightly anterior. Setae h3 flattened tal extensions about 6 (6–8) long, setae se sepa- and enlarged in medial part, greatest wide 8 (8–10). rated by 62 (60–88). Scapular shield with angular Distance between hysteronotal setae: c2:d2 75 suprategumental extensions in postero-mesal an- (70–76), d2:e2 100 (90–105), e2:h2 71 (69–85), gle. Hysteronotal shield: length 260 (255–290), h2:h3 24 (20–24), ps1:h3 24 (22–28), ps2:ps2 82 width at anterior margin 100 (96–115), lateral (80–102), h2:h2 75 (72–90), h3: h3 68 (65–75), margins, anterior margin slightly concave, surface d1:d2 27 (25–33), e1:e2 31 (22–35). evenly punctate, without other ornamentation. Sternum about 2/3 of total length of epimerites Terminal cleft: parallel-sided, anterior margin I, basal part of these epimerites strongly thickened. concave, with unclear border, length 48 (42–53), Coxal fields II and III with large triangular sclero- width 47 (45–55). Supranal concavity large semi- tized areas in lateral parts. Epimerites IVa wide ovate, with poorly distinct lateral borders. Interlo- triangular, with setae 4a on anterior end. Genital bar membrane occupies most space between lobes arch 22 × 26 (17–22 × 24–26), aedeagus minute,

93 S.V. Mironov

G B A

I F

Fig. 3. Anhemialges gaudi sp. n., details. A–D — male, G–I — female. A — leg I, dorsal view, B — tarsus I, ventral view, C — trochanter and femur I, ventral view, D — leg II, dorsal view, E — genu, tibia and tarsus III, dorsal view, F — tibia and tarsus IV, dorsal view, G — tarsus III, dorsal view, H — tarsus IV, dorsal view, I — primary specrmaduct and head of spermatheca. co — copulatory opening, hs — dead of spermatheca, pd — primary spermaduct. much shorter than arch. Epiandrum thin, bow- covering anterior half of anal opening, setae ps3 shaped, 18 (16–20) long, 37 (35–40) wide. Para- setiform, situated near anterior margin of adanal genital apodemes flank area with epiandrum and shield. Adanal membranes wide, stretching from genital apparatus from level of setae 3a to midlev- lateral ends of adanal shield to bases of setae ps2, el of trochanters IV. Gastral shield trapezoidal, 65 base of membranes with narrow and poorly scle- (50–66) long (in small-sized individuals may be rotized adanal apodemes. Anal suckers low cylin- split longitudinally). Adanal shield triangular, drical, 14 (13–17) in diameter. Cupules ih approx-

94 The type species of the feather mites and a new genus of the analgid mites imately at midlevel of adanal membranes. Length tory opening on a small conical extension. In of anal field from anterior margin of adanal shield males of A. subinteger, the median part of hyster- to lobar apices 130 (110–135). Distance between onotal shield from its anterior margin to the level ventral setae: 3a:4a 30 (30–35), 3a:g 42 (40–44), of trochanters IV is covered with transverse wavy g:ps3 88 (80–92). striae, incision in interlobar membrane shlallow, Femur II with short lateral apophysis curved and seta w of tarsi III extends only to bases of cor- ventrally (in small-sized individuals may be indis- responding seta s; in females, the copulatory open- tinct). Genual seta cGI long setiform, seta cGII ing is situated on small blunt extension with a little with foliate enlargement at base and with long fili- pit on apex. form apical part. Tarsus III 86 (82–95) long, with Remarks. In establishing the genus Anhemi- narrow crest along lateral margin, blade like setae alges, Gaud (1958) considered “Anhemialges lon- w and s 39 (33–40) and 29 (22–30) long, respec- gipes” collected only from Hirundo rustica in tively; seta r slightly longer than this segment. Morocco and did not mention other hosts. There- Tarsus IV conical, slightly curved, with bidentate fore it is more reasonable to choose this bird spe- apex, 33 (32–38) long. Legs IV extending by am- cies as the type host of the new species; although bulacral discs to level of setae h2. in the earlier publication Gaud (1953) reported Female (range of measurements for 6 para- “Megninia longipes” from H. smithi and H. sene- types). Idiosoma, length × width, 380–400 × galensis. The record for the latter host was cited as 160–180, length of hysterosoma from anterior Trouessart (1899), however this is apparently an margin of humeral shields to posterior end of error and he meant Bonnet (1924), who first indi- opisthosoma 255–270. cated that M. longipes had three different hosts Prodorsal shield shaped as in male, length (including. H. senegalensis), though Bonnet was 78–85, width 72–80, acute suprategumental ex- actually dealing with three different mite species tension 7–9 long, setae se separated by 60–70. (see Introduction). Scapular shields as in male. Setae c2 at level of So far, five hirundinid species of three genera humeral shields, setae d2 at level of trochanters have been indicated as hosts of A. gaudi (Gaud IV, length of setae c2, d2 about ¾ of idiosoma’s and 1958; Gaud Mouchet 1959; Gaud and Till greatest width. Setae e2 not extending to posterior 1961). Taking into consideration that hirundinid margin of opisthosoma, 2–2.5 times shorter than species from the genera Delichon Horsefield et setae c2, d2. Setae e1 absent, setae d1 present or Moore and Tachycineta Cabanis bear other An- absent. Distance between setae: c2:d2 55–62, hemialges species (Tyrrell 1882, Berlese 1883), it d2:e2 100–115, e2:h2 84–90, h5:h3 60–66. Copu- cannot be excluded that Gaud and Till (1961) ac- latory opening on small conical extension near tually dealt with a complex of closely related mites posterior margin of opisthosoma, secondary sper- and that swallows from genera other than Hirundo maducts indistinct, primary spermaduct and head could bear separate mite species. of spermatheca as in Fig. 3I. It is unknown at present, whether the “An- Epimerites I as in male. Epigynum semicircu- hemialges longipes” specimens from H. hirundo lar, with slightly thickened posterior, 24–28 long, collected by Gaud in Morocco still exist, but it is 39–45 wide. Setae 3a on tips of epigynum. Genital potentially possible that they are retained in some papillae mesal to bases of setae g. collection, where he deposited his materials. Tarsus III 45–50 long, tarsus IV 52–57, tibial Etymology. The species is named after Prof. solenidion jIII slightly longer than corresponding Jean Gaud, the greatest French expert in feather tarsus, solenidion jIV slightly shorter than corre- mites. sponding tarsus. Legs IV maximally extending by ambulacral disc to level of setae f2. Subfamily Megniniinae Gaud et Atyeo, 1981 Differential diagnosis. Males of Anhemialg- Genus Anhimomegninia gen. n. es gaudi sp. n. differ from a closely related spe- Type species: Megninia longipes Trouessart, cies, A. subinteger from Delichon urbicum, by 1899 from the horned screamer Anhima cornuta, having a uniformly punctate hysteronotal shield, Guyana. incision in interlobar membrane extending at least Diagnosis. Male. Small-sized megniniines. to midlevel of terminal cleft, and longer seta w of Idiosoma strongly enlarged in area of humeral and tarsi III extending to midlength of corresponding scapular shields. Prodorsal shield occupying me- seta s; females differ by the position of the copula- dian part of prodorsum, shaped as long and narrow

95 S.V. Mironov band expanded at level of scapular setae, with me- shield narrow, not larger than the distance between dian longitudinal ridges in anterior part, without scapular setae se, epimerites I are fused into a Y, suprategumental extensions on margins. Latero- tarsus III is strongly elongated and bears two coxal setae lcx absent. Vertical setae vi present. blade-like setae (w, s), and ambulacral discs of Idiosomal setae absent: ve, c1, d1, e1, h1. Lateral pretarsi II–IV are strongly reduced compared to margins of opisthosoma slightly attenuate posteri- those on legs I. Males of Anhimomegninia differ orly. Opisthosoma bilobate; lobes angular, moder- from those of Megniniella by having semicircular ately elongated, separated by ovate terminal cleft. ventral extensions on tarsi I, II (a feature which Terminal cleft occupied by interlobar membrane should occur in females), the scapular and humer- with median incision. Lateral membranes absent. al shields fused, and strongly elongated ambu- Hysteronotal shield with roughly rectangular ante- lacral stalks of tarsi III, IV. In Megniniella, the rior part. Scapular and humeral shields fused. Se- ventral extensions of tarsi I, II are acute (as in most tae c2 situated on scapular shields. Setae h3 on genera of Megniniinae), the scapular and humeral lobar apices, setae h2 on lateral margins of opist- shields are free, ambulacral stalks of tarsi I, II are hosomal lobes, setae ps1 on lateral margins of ter- only 1.5–2 times longer than those on tarsi I, II. minal cleft, far anterior to level of setae h3. Semicircular form of the ventral extension on Epimerites I fused into a Y with long sternum. tarsi I, II and central sclerites of ambulacra repre- Coxal field III not closed. Genital apparatus at sented by a pair of longitudinal pieces in all legs level of trochanters III. Epiandrum absent. Para- are unique characters within Megniniinae. These genital apodemes present, flanking genital appara- features should eventually allow to recognize fe- tus laterally. Genital shield absent. Adanal mem- males of this genus, which are unknown at pres- brane entire, horseshoe-shaped, encircling anterior ent. half of anal field. Adanal and gastral shields pres- Etymology. Contraction of Anhima (the ge- ent, situated in anterior part of anal field. Anal neric name of the host of the type species) and the suckers disc-like, corolla not indented. feather mite genus Megninia, gender feminine. Ambulacral discs of legs II–IV narrowly ovate, much smaller than on legs I and provided Anhimomegninia longipes (Trouessart, 1899) comb. n. with thin apical extension (Figs. 6A–D). Central sclerite of ambulacral discs of all legs represented Figs. 4–6 by pair of longitudinal rods. Distal half of ambu- Megninia longipes Trouessart 1899: 26; lacral stalk of tarsi I, II with noticeably convex Canestrini and Kramer 1899: 101; Bonnet 1924: dorsal side. Femora I, II without lateral apophy- 174, figs. 33, 34 (part.); Radford 1953: 207 (part.) ses. Tarsi I, II with semicircular ventral expan- Material examined. Male holotype (by sions (Figs. 6A, B). Tibiae I, II with spine-like monotypy) (TRT 28i9) ex Anhima cornuta (Lin- ventral apophyses. Tarsi I, II with 8 setae. Solen- naeus) (Anseriformes: Anhimidae), Guyana. idion w1 and seta ba of tarsus II closer to apex Description. Male (holotype). Idiosoma, than to base of this segment, solenidion w1 of tar- length × width, 335 × 218, length of hysterosoma sus II 2.5–3 times longer than on tarsus I. Legs III 268–305. Subcapitulum with slightly convex lat- hypertrophied, much thicker and longer than legs eral margins, without lateral spines. Prodosal IV, femur and genu III without apophyses; distal shield a narrow longitudinal plate slightly enlarged end of tibia III with two apophyses; tarsus III between bases of scapular setae and with pair of much thinner that other segments of this leg, short angle-shaped lateral extensions anterior to strongly elongated, with acute apex, setae s and w scapular setae, posterior end slightly extending lanceolate or spiniform, other setae setiform. Tar- beyond level of scapular setae, posterior margin sus IV subequal in length to corresponding tibia, straight, length 80, width at level of lateral exten- with dorsobasal process; setae d, e barrel-shaped, sions 47. Both pair of scapular setae off prodorsal with discoid cap. Ambulacral stalk of tarsi III, IV shield, setae se separated by 53. Setae c2 on inner strongly elongated, 2–3 times longer than on tarsi margins of scapular shields. Hysteronotal shield: I, II. length 222, width at anterior margin 78, anterior Female. Unknown. angles rounded, anterior margin straight, lateral Differential diagnosis. Anhimomegninia gen. margins parallel-sided in anterior part, slightly di- n. is close to Megniniella Gaud, 1958 by having vergent posteriorly, surface punctured, posterior the following characters in male: the prodorsal half with fine longitudinal striae and dashes. Ter-

96 The type species of the feather mites and a new genus of the analgid mites

Fig. 4. Anhimomegninia longipes (Trouessart, 1899), male holotype: dorsal view. minal cleft: ovate, lateral margins well sclerotized, IIIa. Inner tips of epimerites IIIa bearing setae 3a length 47, greatest width 31. Supranal concavity and with narrow extension directed laterally. Ada- longitudinal, poorly outlined, open posteriorly nal membrane large horseshoe-shaped, encircling into terminal cleft. Interlobar membrane occupies anal field (ventral area of opisthosoma bearing most space between lobes, incision in interlobar anal opening, anal suckers, adanal and gastral membrane angular, 29 long. Setae ps1 slightly an- shields) from anterior and lateral sides and extend- terior to midlevel of terminal cleft. Distance be- ing to bases of setae ps2. Adanal shields roughly tween hysteronotal setae c2:d2 80, d2:e2 66, e2:h2 rectangular 15 × 27, with truncate extension on 84, h2:h3 7, ps1:h3 27, ps2:ps2 75, h2:h2 55, h3: posterior margin, covering anterior half of anal h3 42, d1:d2 35. opening. Gastral shield 21 in length along midline, Sternum slightly longer than half of total with two poorly sclerotized branches (adanal apo- length of epimerites I. Coxal fields I–III without demes) stretching posteriorly, along basal margin large sclerotized areas in lateral parts. Epimerites of adanal membrane. Anal suckers disc-like, 116 IVa wide triangular, setae 4a mesal to their ante- in diameter. Cupules ih close to anal suckers. rior ends. Genital arch: length 20, width at base Length of anal field (from anterior margin of gas- 20; aedeagus directed anteriorly, curved, 16 long. tral shield to lobar apices) 108. Distance between Paragenital apodemes flanking genital apparatus ventral setae: 3a:4a 22, 3a:g 33, g:ps3 55. from anterior and lateral sides, connected by thin Femur I with small rounded ledge on lateral longitudinal bands with inner tips of epimerites margin, femur with rounded margin. Genual seta

97 S.V. Mironov

Fig. 5. Anhimomegninia longipes (Trouessart, 1899), male holotype: ventral view. cGI short, with foliate lateral enlargement, seta the only specimen of M. longipes from Anhima cGII with basal foliate enlargement and filiform cornuta existing in the Trouessart collection, it is apical part (Figs. 6A, B). Tarsus III 129 long, ap- highly probable that Trouessart had in the hands a proximately 1.5 times longer and 2 time thinner sole male, and therefore this specimen is the holo- than corresponding tibia, with straight and acute type by monotypy (ICZN 1999, Article 73.1.2) apical extension; setae w spiculiform, 62 long, Bonnet (1924: 174, Figs. 33, 34) illustrated the seta s lanceolate, 33 long; seta r slightly shorter opisthosoma and tarsus III of “Megninia longipes” than a half of this segment length (Fig. 6C). Tar- male and listed the three host species: Anhima cor- sus IV 24 long, slightly shorter than correspond- nuta, Streptoprocne zonaris and Hirundo senega- ing tibia, with rectangular dorsobasal extension, lensis. The figured structures correspond much setae d, e barrel-like, with discoid apical caps more closely to those in the genus Cypselalges (Fig. 6 D). Gaud et Atyeo, 1991, a genus specific to apodids in Remarks. Trouessart (1899) described Meg- the analgid subfamily Protalginae: the extensions of ninia longipes only from the male and mentioned interlobar membrane protruding beyond lobar api- one host species from one location; unfortunately ces are acute (Fig. 33), tarsus III is cone-like, there is no direct indication in the text that he had straight and relatively thick at base (Fig. 34). In An- a sole male. Since the description is given for a himomegninia, the extensions of the membrane are male and the examined slide still retaining the absent, in Anhemialges they are rounded (if pres- original labels written by the Trouessart’s hand is ent), and in both genera, tarsus III of male is much

98 The type species of the feather mites and a new genus of the analgid mites

A D

B E

Fig. 6. Anhimomegninia longipes (Trouessart, 1899), details of male. A — leg I, dorsal view, B — leg II, dorsal view, C — tibia and tarsus III, dorsal view, D — tibia and tarsus IV, dorsal view, E — genital apparatus and paragenital apodemes. Seta- tion of legs I, II is combined from legs of different sides, because many setae are broken in the holotype specimen. thinner than was illustrated by Bonnet and defi- a natural parasite of screamers, or is this record nitely curved. Therefore it is quite probable that a caused by the accidental contamination. This ques- Cypselalges species from Streptoprocne zonaris tion may be solved by additional investigation was actually used for making these drawings. screamers in relation of their feather mite fauna. Feather mites from the families Alloptidae ACKNOWLEDGEMENTS and Freyanidae are known from both genera of screamers, Anhima Brisson and Chauna Illiger The author thanks Dr. Heather C. Proctor (Dubinin 1950; Peterson 1971), but the finding of (University of Alberta, Edmonton, Canada) for Anhimomegninia longipes is the only record of an critically reviewing the manuscript and Dr. Pavel analgid species on birds from the family Anhingi- B. Klimov (University of Michigan, Ann Arbor, dae. Therefore the question remains: is A. longipes USA) for taxonomic consulting. The investigation

99 S.V. Mironov was supported by the Russian Foundation for Ba- Gaud, J. and Mouchet, J. 1959. Acariens plumicoles sic Research (project No 07-04-00426a). Most of (Analgesoidea) parasites des oiseaux du Came- the material used in the study belongs to the col- roun. II. Analgesidae. Annales de parasitologie lection UFC ZIN No 2-2.20 deposited in the Zoo- humaine et comparée, 34: 149–208. logical Institute RAS (Saint Petersburg, Russia). Gaud, J. and Till, W.M. 1961. Suborder Sarcoptiformes, pp. 180–352. In: Zumpt, F. (Ed.). The arthropod References parasites of vertebrates in Africa south of the Sa- Arutunjan, E.S. and Mironov, S.V. 1983. [New and hara (Ethiopian Region). Volume I (Chelicerata). little known species of feather mites (Analgoidea) Publications of the South African Institute of in the fauna of the USSR]. Zoologicheskiy Sbornik, Medical Research. No L. (Vol. IX). Johannesburg, Institut Zoologii AN Armyanskoy SSR, 19: South Africa. 457 pp. 319–336. [In Russian] ICZN. 1999. International Code of Zoological Nomen- Atyeo, W.T. 1973. Feather mites. In: McClure H.E. and clature, 4th edition. The International Trust for Zoological Nomenclature, London. 305 p. Ratanaworabhan N. (Eds.). Some ectoparasites of Kolarova, N.T. and Mitov, P.G. 2008. Feather Mites of the birds of Asia. Jintana Printing Ldt., Bangkok, the Superfamily Analgoidea (Acari: Astigmata) p. 54–78. from Passerines (Aves: Passeriformes) in South Berlese, A. 1882–1903. Acari, Myriopoda et Scorpio- Dobrudzha, Bulgaria. Acta Zoologica Bulgarica, nes hucusque in Italia reperta. Padova and Portici. Supply 2: 91–102. 101 fascicles. Mironov, S.V. 1983. [Feather mites (Sarcoptiformes, Bonnet, A. 1924. Révision des genres Mégnina, Mesal- Analgoidea) of Passeriformes of the Nizhnyes- ges et genres voisins de la sous-famille des Sar- virsk Preserve], pp. 82–94. In: Bibikova L.A. coptides plumicoles (Première partie). Bulletin de (Ed.). Parazitologicheskie issledovaniya v la Société zoologique de France, 49: 146–188. zapovednikakh. Sbornik nauchnykh trudov TsNIL Canestrini, G. and Kramer, P. 1899. Demodicidae und glavokhoty RSFSR, Moscow. [In Russian] Sarcoptidae. Das Tierreich, 7: 1–193. Mironov, S.V. 1996. [Feather mites of the passerines in Černý, V. 1967. Catálogo de la fauna Cubana. XX. Li- the North-West of Russia]. Parazitologiya, 30: sta de los ácaros parásitos de aves reportadas de 521–539. [In Russian] Cuba. Museo „Felipe Poey“ de la Academia de Mironov, S.V. 1999. Feather mites: general morpho- Ciencias de Cuba, Trabajos de Divulgación, 45: logical adaptations, phylogeny and coevolutionary 1–23. (Unpublished) relationships with birds. Ekologija, 2: 57–66. Dickinson, E.C. 2003 The Howard and Moore Com- Mironov, S.V. and Galloway, T. 2002a. New feather plete Checklist of the Birds of the World, 3rd Edi- mite taxa (Acari: Analgoidea) and mites collected tion. Princeton University Press, Princeton, N.J., from native and introduced birds of New Zealand. 1056 pp. Acarologia, 42: 185–201. Dubinin, V.B. 1950. [Systematic analysis of species of Mironov, S.V. and Galloway, T. D. 2002b. Four new feather mites (Sarcoptiformes, Analgesoidea), feather mite species (Acari: Analgoidea) from parasites of anatids]. Parazitologicheskiy Sbornik, some birds in Canada. The Canadian Entomolo- ZIN AN SSSR, 12: 17–72. [In Russian] gist, 134: 605–618. Gaud, J. 1953. Sarcoptides plumicoles des oiseaux Peterson, P.C. 1971. A revision of the feather mite ge- d’Afrique occidentale et centrale. Annales de pa- nus Brephosceles (Proctophyllodidae: Alloptinae). rasitologie humaine et comparée, 28: 193–226. Bulletin of the University of Nebraska State Mu- Gaud, J. 1958. Acariens plumicoles (Analgesoidea) pa- seum, 9: 89–172. rasites des oiseaux du Maroc. II. Analgidae. Bulle- Radford, C.D. 1953. The mites (Acarina: Analgesidae) tin de la Société de Sciences naturelles et physi- living on or in the feathers of birds. Parasitology, ques du Maroc, 38: 27–49. 42: 199–230. Gaud, J. and Atyeo, W.T. 1991. Huit genres nouveaux Trouessart, E.L. (1898) 1899. Diagnoses préliminaires de la famille Analgidae (Acarina, Analgoidea). d’espèces nouvelles d’Acariens plumicoles. Addi- Acarologia, 32: 163–182. tions et corrections à la sous-famille des Analgési- Gaud, J. and Atyeo, W.T. 1996. Feather mites of the nés. Bulletin de la Société d’études scientifiques World (Acarina, Astigmata): the supraspecific d’Angers, 28: 1–62. taxa. Musée Royal de l’Afrique Centrale, Annales, Tyrrell, J.B. 1882. On some Canadian ectoparasitic Sciences Zoologiques, 277: 1–193 (Part I, text), Sarcoptidae. Transactions of the Ottawa Field- 1–436 (Part II, Figures). naturalists’ Club, 3: 43–48.

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