Three new feather mite genera of the Protolichus generic group (Astigmata, Pterolichidae) from...

Three new feather mite genera of the Protolichus genericgroup (Astigmata, Pterolichidae) from parrots

(Aves, Psittaciformes) of the Old World

Sergey V. Mironov1* and Jacek Dabert21Zoological Institute, Russian Academy of Sciences, Universitetskaya emb. 1, 199034, Saint Petersburg, Russia;

2Department of Animal Morphology, A. Mickiewicz University, Umultowska 89, Poznañ, 61-614, Poland

AbstractThree new genera of feather mites of the Protolichus generic group (Pterolichidae, Pterolichinae), restricted to parrots, are estab-lished based on re-examination of type series and some previously unidentified mite specimens deposited in the collection ofE. Trouessart (Muséum National d�Histoire Naturelle, Paris, France): Loriprotolichus gen. nov. with the type speciesLoriprotolichus charmosynae sp. nov. from Charmosyna papou (Scopoli, 1786) and Loriprotolichus falculiger (Trouessart,1884) comb. nov. from Glossopsitta concinna (Shaw, 1791); Calyptolichus gen. nov. with type species Calyptolichus favettei(Trouessart, 1899) comb. nov.; Nestorilichus gen. nov. with the type species Nestorilichus atyeoi sp. nov. from Nestor meri-dionalis (Gmelin, 1788). Several taxonomic changes were implemented in the course of re-examination of the type series. Theholotype of Pterolichus (Protolichus) falculiger Trouessart, 1884 is designated. Re-examination of the full syntype series ofP. (Protolichus) favettei Trouessart, 1899 representing samples from different hosts, recovered two distinct pterolichid speciesof the Protolichus generic group. The sample clearly corresponding to the original description (Trouessart, 1899) is fixed asthe lectotype and nomenclatural type of P. (Protolichus) favettei; the second species is described as a new species Nestorilichusatyeoi. Protolichus (Mesolichus) favettei cordifora Favette et Trouessart, 1904 is synonymized with P. (Protolichus) favettei.

KeywordsFeather mites, Pterolichidae, systematics, host-parasite associations, Psittaciformes

*Corresponding author: astigmata@zin.ru

Introduction

Feather mites of the suborder Astigmata are the most commonand numerous ectoparasitic mites living permanently onplumage and skin of birds. Approximately 2500 species in 300genera and 38 families are known throughout the world andhave been recorded from almost all avian orders (Gaud andAtyeo 1996, Dabert and Mironov 1999, Mironov 2003,Proctor 2003).

The feather mite family Pterolichidae includes about 330species and is the most species-rich family of the superfami-ly Pterolichoidea. Mites of this family are specialized to in-habit the surface of feathers with large vanes, i.e. flight andcovert feather of the wing and tail, and are associated exclu-sively with non-passerine avian orders. Of the feather mitehigher taxa occurring on parrots (Psittaciformes), the familyPterolichidae is the most numerous with over 120 species in34 genera known from these hosts worldwide. Pterolichids

restricted to parrots comprise three morphologically distinctgeneric groups termed the Protolichus, Rhytidelasma andPsittophagus generic groups (Gaud and Atyeo 1996; Mironovand Pérez 2003; Mironov et al. 2003a, b). Of these, the Proto-lichus group (with about 70 species in 22 genera) is the mostspecies rich and morphologically diverse grouping. Over thepast twenty years, the diversity and taxonomy of this genericgroup has been extensively investigated in South America(Atyeo et al. 1984; Atyeo 1988, 1989a, b, c; Atyeo and Pérez1990; Gaud and Atyeo 1996; Mironov et al. 2005) and inAfrica (Gaud and Mouchet 1959, Gaud 1980). In contrast,mites of Protolichus group in the Australian and Indo-Ma-layan regions have been poorly explored (Atyeo and Pérez1982, Atyeo and Gaud 1987, Mironov et al. 2003b, Dabert etal. 2006). In the two latter regions, the group in question isrepresented by five genera: the monotypic genus SideroferusGaud et Atyeo, 1996, and three species of Titanolichus Gaudet Atyeo, 1996, are known from ground parrots (Psittacinae,

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New genera of feather mites of the Protolichus generic group

Platycercini); two species of Uropsittacolichus Atyeo et Pé-rez, 1982 are known from parrots of the tribe Cyclopsittini(Psittacinae); six species of Protolichus Mégnin et Trouessart,1884 are known from lories and lorikeets (Psittacidae, Lori-inae); and representatives of the genus Apexolichus Gaud etAtyeo, 1996 (8 species) have been recorded from variousgroups of psittaciforms including cockatoos (Cacatuidae) andtwo subfamilies of psittacids, Psittacinae and Loriinae. Such awide and patchwork distribution of one genus of feather mitesamong host taxa raises suspicions that the genus Apexilochusin its current sense is an artificial rather than a monophyletictaxon.

In the course of our study of pterolichid feather mites asso-ciated with parrots (Mironov et al. 2003a, b, 2005; Dabert etal. 2004, 2006) we collected abundant new material from var-ious museums and also re-examined material from the Troues-sart collection, which was created at the end of the 19th cen-tury. In the present paper, based on re-examination of typeseries of three mite species formerly referred to the genusApexolichus by Gaud and Atyeo (1996) and on some addi-tional collections, we re-describe these species and establishthree new pterolichid genera belonging to the Protolichusgeneric group.

Materials and methods

The main material used in the present study was loaned fromTrouessart collections deposited in the Muséum Nationald�Histoire Naturelle (Paris, France). Some additional speci-mens originated from the collections of the authors and fromthe Uebersee Museum (Bremen, Germany). Diagnoses anddescriptions of taxa follow the recent standard schemes usedfor pterolichid mites of the group in question (Gaud and Atyeo1996, Mironov et al. 2003a, Dabert et al. 2006).

It is important to note that many specimens in the Trou-essart collection were re-examined by W.T. Atyeo (Universityof Georgia, Athens, USA) in the 1980s, and lectotype or para-lectotype marks were made on slide labels for some species of�Protolichus�. Unfortunately, Atyeo�s study was not complet-ed. Data on designations of lectotypes or syntype series forspecies referred to the genera Apexolichus and Titanolichushave never been published, and therefore formally such des-ignations are not valid. Even many new combinations sur-mised for Protolichus species moved to Apexolichus havenever formally been given in subsequent publications, includ-ing the review of feather mite genera of the world (Gaud andAtyeo 1996). Nevertheless, in most cases we followed theunpublished concepts of W.T. Atyeo in designation of lecto-types.

Abbreviations used in accession collection numbers andfor pointing out the depositories of specimens: AMU � A. Mic-kiewicz University (Poznañ, Poland), TRT � Trouessart col-lections of the Muséum National d�Histoire Naturelle (Paris,France), UBR � Uebersee Museum (Bremen, Germany); ZISP� the Zoological Institute of the Russian Academy of Sci-ences (Saint Petersburg, Russia). Taxonomic system and Latin

names of hosts used in the present study follow del Hoyo et al.(1997).

Systematics

Pterolichidae Trouessart et Mégnin, 1884Pterolichinae Trouessart et Mégnin, 1884

Loriprotolichus gen. nov.

Type species: Loriprotolichus charmosynae sp. nov.Description (both sexes): Prodorsal shield covering almost

all prodorsum, encompassing bases of scapular setae se, si;lateral margins of anterior part of shield with ledge; posteriorangles extending laterally, long and acute; scapular shields assmall narrow bands (Figs 1A; 2A; 3A). Hysterosoma withextensive hysteronotal shield; humeral shields bearing basesof setae c2 and cp. Two vertical setae vi present, their basesclose to each other. Scapular setae si well-developed, spiculi-form or setiform, situated closer to midline than to respectivesetae se. Idiosomal setae c2 spiculiform; setae c3 piliform;setae h1 absent. Cupules ia posterolateral to setae c2; cupulesim posterior to hysteronotal gland openings gl. Bases of epi-merites I, II without sclerotized inflations at bases. Each tar-sus with pair of minute apicodorsal spines; tarsi, tibiae, andgenua of legs I, II without noticeable apophyses on ventralsurface; tarsi I, II slightly longer than corresponding tibiae;solenidion ω1 approximately at midlevel of tarsi I, II; setae baof legs I, II slightly distal to corresponding solenidion ω1;solenidion σ2 on genu I present; ambulacral discs minutelydentate on distal margin (Fig. 4C, D).

Male: Gnathosoma with slightly convex lateral margins;palps without lateral extensions. Opisthosomal lobes well-developed, relatively short (slightly longer than wide at base),rounded apically; anterior margin of terminal cleft semi-round-ed (Fig. 4A, B). Inner, posterior and postero-lateral margins oflobes with narrow entire membrane. Idiosomal setae d2 spic-uliform; setae e2 as thick and long macrosetae; setae f2 ex-tending beyond lobar apices, with or without membranousextension at base; setae ps2 setiform, much shorter than setaef2; setae ps1 setiform, situated on dorsal surface of opistho-somal lobes; setae e1 sickle-shaped or L-shaped, variable inposition between levels of gland openings gl and setae h2(Fig. 4B, H). Epimerites I fused as a Y. Coxal fields I�IV open,coxal fields IV and lateral part of fields III with large sclero-tized areas; other fields without sclerotized areas. Bases oftrochanters I�II flanked by narrow sclerotized bands connect-ing bases of respective epimerites. Paragenital apodemes pres-ent, long and heavily sclerotized, spreading from level oftrochanters III to base of opisthosomal lobes, anterior ends ofapodemes connected by narrow bands to inner tips of epimer-ites IIIa, posterior ends of apodemes fused with opisthoven-tral shields; inner margins of apodemes with poorly sclero-tized areas extending to midline (Figs 1A; 3B; 4A). Anal suck-ers circular, corolla dentate, each sucker surrounded by ovatemembrane, posterior margin of anal field flanked by sclero-tized bands connecting opisthoventral shields. Genital appa-

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Sergey V. Mironov and Jacek Dabert

ratus at level of trochanters III. Genital papillae situated oninner margins of paragenital apodemes at level of genitalapparatus. Cupules ih present. Legs I, II subequal in size; legsIV slightly thicker than legs III. Tarsus IV longer than wide,similar in form and size to tarsus III; setae d, e reduced up topale patches on small spine-like apical extensions (Fig. 4G).

Female: Hysteronotal shield entire (not split into pygidialfragment and main body), covering almost all dorsal surfaceof hysterosoma (Fig. 2A). Lateral sclerotized bands of hys-terosoma absent. Setae d2, e2 short, spiculiform or setiform;setae f2, ps1, ps2 setiform. Epimerites I free, posterior tipsconverging. Oviporus long, extending from level of scapularshields to midlevel between trochanters III and IV. Epigyniumpresent, represented by thick bow-shaped sclerite. Copulatoryopening terminal, on small cone-like extension.

Hosts: Lories of the genera Charmosyna Wagler, 1832 andGlossopsitta Bonaparte, 1854 (Psittacidae, Loriinae).

Differential diagnosis

Within the Protolichus generic group the new genus is mostsimilar in general appearance to Genoprotolichus Gaud etAtyeo, 1996 by having opisthosomal lobes in males relative-ly short and wide and rounded apically. Loriprotolichus dif-fers from that genus by the following characters: in bothsexes, bases of epimerites I, II without sclerotized inflations,prodorsal shield with long and acute posterior angles; inmales, epimerites I fused into a Y, setae ps1 setiform, setae e1sickle-shaped, setae d, e of tarsus IV reduced to weakly scler-otized patches and situated on spine-like extensions; in fe-males, hysteronotal shield entire. The structure of the prodor-

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Fig. 1. Loriprotolichus charmosynae sp. nov., male: A � dorsal view, B � ventral view

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sal shield in both sexes and form of setae e1 in males areunique features of this genus within the Protolichus genericgroup.

Etymology

A combination of the name for the host subfamily and ofProtolichus.

Key to Loriprotolichus species (males)1. Sickle-shaped setae e1 situated on the hysterosoma proper,approximately equidistant from levels of setae d2 and e2;scapular shields not connected to prodorsal shield (Figs 1A;4H) ................................................... L. charmosynae sp. nov.� Sickle-shaped setae e1 situated on inner margins of opistho-somal lobes, approximately at level of setae h2; scapularshields not connected to prodorsal shield (Figs 3A; 4B) ........................ L. falculiger (Favette et Trouessart, 1904) comb. nov.

Loriprotolichus charmosynae sp. nov. (Figs 1; 2; 4H)

Type material: Holotype male and paratype female (TRT34A14) ex Charmosyna papou (Scopoli, 1786), Psittacidae,New Guinea, no other data.

Description (male, holotype): Idiosoma: length 410, width258. Subcapitulum with pair of oblique crests. Prodorsalshield: anterior part uniformly punctured, with ledge on later-al margins; posterior part with wavy transverse striae, totallength 104, width of anterior part 82, width of posterior part170. Setae si spiculiform, 46�48 long. Distance betweenscapular setae: se:se 60, si:si 22. Scapular shields fused to pos-terior angles of prodorsal shield. Hysterosoma 305 long. Hys-teronotal shield: length from anterior margin to lobar apices308, width at anterior margin 202, anterior two thirds withnumerous striae, posterior part uniformly punctured. Opistho-somal lobes slightly curved medially; terminal cleft semi-

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Fig. 2. Loriprotolichus charmosynae sp. nov., female: A � dorsal view, B � ventral view

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ovate, length 56, greatest width 36; interlobar membrane de-veloped on inner margins of lobes and absent in median partof the cleft. Supranal concavity absent. Subhumeral setae c3about 60 long. Lateral setae: c2 spiculiform, 62 long; d2 spic-uliform 54, e2 thick spiculiform, 148�150 long. Setae f2 seti-form with narrow membranous enlargement in basal part; tipsof these setae extending beyond lobar apices. Setae e1 L-shaped, enlarged in apical half, their bases situated near later-al margins of hysteronotal shield, approximately at mid-dis-tance between levels of setae d2 and e2 (Fig. 1A). Setae ps1setiform situated slightly posterior to level of setae h2. Dorsalmeasurements: c2:d2 126, d2:e2 104, e2:h3 63, e1:e2 42, e2:e2109, e1:e1 134, ps1:ps1 68. Cupules im posterior to level ofopenings gl. Bases of epimerites II without inflations. Genitalapparatus between levels of trochanters III and IV, 32 × 20.

Both pairs of genital papillae at level of apex of genital appa-ratus. Paragenital apodemes: anterior tips extended to level ofsetae g and connected to inner tips of epimerites IIIa; innermargin of apodemes with sclerotized area almost touching atmidline and flanking anal field from anterior margin (Fig.1B). Ventral measurements: 3b:3a 17, 3a:g 5, g:4a 62, 4a:ps392. Cupules ih at level of setae h2. Diameter of anal sucker 23(opisthosoma of holotype is naturally deformed, one suckerhas not been developed). Segments of legs I, II without anyapophyses. Setae d, e of tarsi IV as weakly sclerotized patch-es on pair of apical spines of this segment (Fig. 4G).

Female (paratype): Idiosoma: length 421, width 240. Pro-dorsal shield: form and position of scapular setae as in male,all surface uniformly punctured, length along midline 112,greatest width of anterior part 70, width of posterior part 186.

Fig. 3. Loriprotolichus falculiger comb. nov., male: A � dorsal view, B � ventral view

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New genera of feather mites of the Protolichus generic group 391

Fig. 4. Loriprotolichus, details of male: A-G � L. falculiger; H � L. charmosynae. A � opisthosoma, ventral view; B � opisthosoma, dorsalview; C � leg I; D � leg II; E � leg III; F � leg IV; G � tarsus IV; H � opisthosoma, dorsal view

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Setae si spiculiform, 32�33 long. Distance between scapularsetae: se:se 60, si:si 19. Scapular shields as in male. Hystero-soma 295 long. Hysteronotal shield entire, length 292, widthof anterior part 192, anterior margin straight, surface with net-work ornamentation (Fig. 2A). Setae c3 about 40�42 long.Setae d2, f2 setiform, e2 spiculiform; length of lateral hys-teronotal setae: c2 30, d2 16, e2 25, f2 12. Dorsal measure-

ments: c2:d2 116, d2:e2 123, e2:h3 50, e2:e2 114, h3:h3 41.Cupules im slightly posterior to level of openings gl. Bases ofepimerites II without inflations. Epigynium bow-shaped 23 ×60. Genital papillae situated between levels of setae 3a and g.Cupules ih postero-mesal to setae ps2. Ventral apophyses oftibiae and genua I, II indistinct.

Fig. 5. Calyptolichus favettei comb. nov., male: A � dorsal view, B � ventral view

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Etymology

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

Loriprotolichus falculiger (Trouessart, 1884) comb. nov.(Figs 3; 4A-G)

Pterolichus (Protolichus) falculiger Trouessart (1884): 531;Canestrini and Kramer (1899): 58; Gaud and Atyeo (1996):117; Protolichus (Mesolichus) falculiger: Favette and Trou-essart (1904): 154, fig. 16.

Material examined: Holotype male (TRT 34F15) ex Glos-sopsitta concinna (Shaw, 1791) (Nannodes australis in origi-nal label), Psittacidae, Australia, no other data; holotype des-ignated here, see remarks.

Description (male, holotype): Idiosoma: length 582, width365. Subcapitulum without spines or crests. Prodorsal shield:anterior piece uniformly punctured, lateral margins with smallledge; posterior part with wavy transverse striation; totallength 117, width of anterior part 90, width of posterior part224. Setae si 72�75 long. Distance between scapular setae:se:se 63, si:si 21. Scapular shields as narrow bands with acuteinner ends, separated from prodorsal shield. Hysterosoma 415long. Hysteronotal shield: length from anterior margin tolobar apices 372, width at anterior margin 212, anterior twothirds with numerous striae, posterior part uniformly punc-tured. Opisthosomal lobes slightly curved medially; terminalcleft semi-ovate, length 72, greatest width 37; interlobar mem-brane relatively large on inner margins of lobes and almostreduced on anterior margin of the cleft. Supranal concavity

Fig. 6. Calyptolichus favettei comb. nov., details of male: A � opisthosoma, ventral view; B � opisthosoma, dorsal view; C � leg I; D � legII; E � leg III; F � leg IV

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absent. Subhumeral setae c3 setiform, 58�60 long. Lateralidiosomal setae: c2 spiculiform, 95�97 long; d2 setiform 75,e2 thick spiculiform, 175�177 long; f2 setiform, extendingbeyond lobar apices. Setae e1 L-shaped, slightly enlarged indistal half, their bases situated on inner margins of opisthoso-mal lobes at level of setae h2 (Figs 3A; 4B). Setae ps1 seti-form. Dorsal measurements: c2:d2 168, d2:e2 124, e2:h3 81,e2:e1 57, e2:e2 129, e1:e1 56, ps1:ps1 85. Cupules im poste-rior to level of openings gl. Bases of epimerites II withoutinflations. Genital apparatus between levels of trochanters IIIand IV, 38 × 16. Both pairs of genital papillae at level of apexof genital apparatus. Paragenital apodemes: anterior tips ex-tended to level of setae 3a, encompassing their bases and con-nected to inner tips of epimerites IIIa, inner margin of apo-demes with sclerotized area almost touching at midline andflanking anal field from anterior margin (Figs 3B; 4A).Ventral measurements: 3b:3a 39, g:4a 77, 4a:ps3 112, setae 3aand g at same transverse level. Cupules ih at level of setae h2.Diameter of anal suckers 23. Segments of legs I, II withoutany apophyses. Setae d, e of tarsi IV as weakly sclerotizedpatches on apical spines (Fig. 4G).

Female unknown.

Remarks

The sole male used for the description of Protolichus fal-culiger (Trouessart 1884, Favette and Trouessart 1904) is theholotype by monotypy (under the Article 73.1.2 of ICZN),rather than the �lectotype� as it was hand-marked by W.T.Atyeo on the label.

Although Gaud and Atyeo (1996) placed P. falculiger intothe genus Apexolichus, the combination of the specific epithet�falculiger� with that generic name was not formally given inthe text.

Calyptolichus gen. nov.

Type species: Pterolichus (Protolichus) favettei Trouessart,1899.

Description (both sexes): Prodorsal shield occupying me-sal part of prodorsum and extending beyond scapular setae se,si; anterior part of shield strongly sclerotized, posterior part(beyond scapular setae) poorly sclerotized (Figs 5A; 7A). Scap-ular shields as narrow oblique bands. Hysterosoma with ex-tensive hysteronotal shield; humeral shields with poorly dis-tinct margins, bearing bases of setae c2 and cp. Two verticalsetae vi present, their bases close to each other. Scapular setaesi well-developed, spiculiform, their position relative to se dif-ferent in males and females; setae c2, spiculiform; setae c3piliform; setae h1 absent. Cupules ia postero-lateral to basesof setae c2; cupules im posterior to hysteronotal gland open-ings gl. Bases of epimerites II with heavily sclerotized ball-like inflations. Each tarsus with two minute apicodorsal spines(in males one of these spines with bidentate apex); tarsi I, IIwith blunt angular ventral margin, approximately 1.5 timeslonger than corresponding tibiae; solenidion ω1 closer to base

of tarsi I, II than to apex; setae ba slightly distal to correspond-ing solenidion ω1; solenidion σ2 of genu I present; condy-lophores heavily developed; ambulacral discs minutely den-tate on distal margin (Fig. 6C, D).

Male: Gnathosoma trapezoidal in general shape, lateralmargins without any processes; palps without lateral exten-sions. Bases of scapular setae si adjacent to bases of respectivesetae se; both pairs of scapular setae very close to midline.Opisthosomal lobes well-developed, narrow and acute api-cally, widely separated from each other by almost rectangu-lar terminal cleft; anterior margin of terminal cleft with pair oflarge blunt-angular extensions; interlobar, terminal, and later-al membranes absent (Figs 5A; 6B). Setae d2, e2 long, spic-uliform; setae f2 with narrow membranous extensions, theirtips not extending beyond lobar apices; setae ps2 setiform,longer than setae f2; setae ps1 enlarged, membranous, approx-imately triangular in form; setae e1 setiform, situated poste-rior to e2. Epimerites I free, posterior tips close to each other.Coxal fields I�IV open, without large sclerotized areas. Basesof trochanters I, II flanked by narrow sclerotized bands con-necting bases of respective epimerites. Paragenital apodemespresent, long and heavily sclerotized, spreading from level oftrochanters III to base of opisthosomal lobes, anterior ends ofapodemes free, noticeably convergent to midline; posteriorends of apodemes fused with opisthoventral shields (Fig. 6A).Anal suckers circular, corolla dentate, each sucker surround-ed by ovate membrane with radial striation, posterior marginof anal field flanked by sclerotized bands connecting opistho-ventral shields. Genital apparatus at level of trochanters IV.Genital papillae situated medial to inner margins of paragen-ital apodemes. Cupules ih present. Legs II slightly longer thanlegs I; legs III and IV similar in size, not hypertrophied. TibiaeI, II and genua I with well-developed spine-like ventralprocess, genua II with weakly developed ventral process.Tarsus IV elongated, similar in form and size to tarsus III;modified setae d, e as membranous buttons (Fig. 6F).

Female: Gnathosoma trapezoidal as in male. Both pairs ofscapular setae noticeably moved to midline, distance of setaesi from midline same as to respective setae se. Hysteronotalshield entire, covering almost entire dorsal surface of hys-terosoma (Fig. 7A). Lateral sclerotized bands of hysterosomaabsent. Setae d2 setiform; e2 spiculiform; setae f2 and ps2 spic-uliform, very short; setae ps1 setiform. Epimerites I free, pos-terior tips close to each other. Oviporus situated at level ofhumeral shields and trochanters III. Epigynium present, rep-resented by thick bow-shaped sclerite. Copulatory openingterminal, on little cone-like extension.

Hosts: Cockatoos of the genus Calyptorhynchus Desma-rest, 1826 (Cacatuidae).

Within the Protolichus group, the genus Calyptolichus is mostsimilar to the genus Apexolichus based on the strongly scler-otized inflations of epimerites II in females, and well-devel-oped ventral spine-like processes on tibiae and genua I, II in

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males. Calyptolichus differs from Apexolichus in the follow-ing combination of features. In males, the opisthosomal lobesare narrow and very acute apically, devoid of any terminalmembranes, and separated by wide rectangular terminal cleft,anterior margin of terminal cleft with a pair of short tooth-likeprocess; posterior margin of anal field flanked by sclerotizedband; modified setae d, e of tarsus IV are button-like. Infemales, hysteronotal shield is not split into main body andpygidial fragment.

Remarks

The genus is monotypic, but it will probably be found to in-clude more species. Gaud and Atyeo (1996: 117) noted thatthey found on various Calyptorhynchus species Apexolichuscordifora (Favette et Trouessart, 1904) and two unnamedspecies of Apexolichus, males of which had attenuate opistho-

somal lobes without interlobar membranes. Unfortunately,these unnamed species have never been described and theplace of their depositories is unknown to us.

Etymology

Combination of abbreviations of the generic name of the hostsand Protolichus.

Calyptolichus favettei (Trouessart, 1899) comb. nov. (Figs5�7)

Pterolichus (Protolichus) favettei Trouessart (1899): 6; Canes-trini and Kramer (1899): 57. Protolichus (Mesolichus) favettei:Favette and Trouessart (1904): 152. Protolichus (Mesolichus)favettei cordifora Favette and Trouessart (1904): 153, fig. 23,syn. nov. Apexolichus cordifora: Gaud and Atyeo (1996): 117.

Fig. 7. Calyptolichus favettei comb. nov., female: A � dorsal view, B � ventral view

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Type material examined: Lectotype male and paralecto-type female (TRT 34G6) ex Probosciger aterrimus (Gmelin,1788) (Microglossus aterrimus in original label), Cacatuidae,New Guinea, no other data; lectotype designated here. Para-lectotype male (TRT 34G8) ex Nestor notabilis Gould, 1856,Psittacidae, New Zealand, no other data. Holotype male (bymonotypy) of Protolichus favettei cordifora (TRT 34G10) exNorthiella haematogaster haematorrhous (Gould, 1838) (Pse-photus xanthorrhous in original label), Psittacidae, Australia,no other data.

Additional material: 4 males, 4 females (ZISP 4355-4362)ex Calyptorhynchus funereus (Shaw, 1794), Cacatuidae, East-ern Australia, no other data; 4 males, 4 females (AMU 01746-01749), same data.

Description (male, lectotype, measurements for 2 para-lectotypes in brackets): Idiosoma: length from anterior marginto tips of opisthosomal lobes 697 (630�680), width 335 (310�322). Subcapitulum without spines or crests. Prodorsal shield:anterior piece elongated, slightly enlarged anteriorly and withrounded posterior border; posterior part with poorly distinctmargin; total length of prodorsal shield 190 (174�177), great-est width of anterior piece 90 (70�80) (Fig. 5A). Setae si 100(102�110) long. Distance between scapular setae: se:se 50(54�56), si:si 24 (23�25). Inner ends of scapular shields con-nected to sclerotized inflations of epimerites II. Hysterosoma477 (458�465) long. Hysteronotal shield: length from anteri-or margin to lobar apices 458 (440�458), width at anteriormargin 215 (215�218), surface uniformly punctured. Opistho-somal lobes narrow with acute apices, widely separated byalmost rectangular terminal cleft, anterior margin of terminalcleft with pair of short angle-shaped extensions (Fig. 6A, B);length of cleft 94 (86�90), width 110 (82�90). Supranal con-cavity absent. Subhumeral setae c3 40 (38�46) long. Lateralsetae c2 spiculiform, 57 (54�71) long; setae d2, e2 broken inall type series (in additional material these setae 62�70 and125�135, respectively); setae f2 setiform with membranousextension. Setae ps1 triangular, with very acute angle directedposteriorly. Dorsal measurements: c2:d2 236 (242�248), d2:e290 (80�91), e2:h3 100 (94�102), e2:e1 23 (18�22), e2:e2 139(134�138), e1:e1 107 (92�102), ps1:ps1 110 (95�106). Cu-pules im posterior to level of openings gl. Epimerites I al-most touching at their posterior tips. Bases of epimerites IIwith heavily sclerotized ball-shaped inflations. Genital appa-

ratus at level of anterior part of trochanters IV, 30 × 17 (30�34× 15�17). Posterior pair of genital papillae at level of apex ofgenital apparatus. Paragenital apodemes: anterior ends strong-ly converging, extending to midlevel between setae 3a and g(Figs 5B; 6A). Ventral measurements: 3b:3a 17 (20�27), 3a:g40 (34�44), g:4a 77 (82�85), 4a:ps3 75. Cupules ih at level ofsetae ps2. Diameter of anal suckers 30 (21�25). Legs I, IIsubequal in size. Tibiae I, II and genua I with distinct ventralspine-like processes, genu II with poorly expressed ventralprocess. Setae d, e button-shaped with little nipple, seta e sit-uated at midlevel of segment, seta d closer to its base (Fig. 6C-F).

Female (paralectotype): Idiosoma, length × width, 595 ×322. Prodorsal shield: anterior part as almost rectangular plateslightly enlarged anteriorly; posterior part as transverse ovalwith poorly distinct margin; total length along midline 137,greatest width of anterior part 72 (Fig. 7A). Setae si spiculi-form, 84�85 long. Distance between scapular setae: se:se 74,si:si 38. Scapular shields as in male. Hysterosoma 425 long.Hysteronotal shield: pygidial fragment demarked but not sep-arated from the main body of the shield, total length 403,width of anterior part 233, anterior margin straight, surfacewith network ornamentation. Setae c3 35 long. Setae d2 seti-form, e2, f2 thin spiculiform; length of setae: c2 46, d2 19�20,e2 25, f2 22. Dorsal measurements: c2:d2 174, d2:e2 158,e2:h3 60, e2:e2 130, h3:h3 52. Cupules im posterior to level ofopenings gl. Bases of epimerites II with sclerotized inflations.Epigynium bow-shaped 36 × 75. Genital papillae approxi-mately at level of setae g. Cupules ih postero-mesal to setaeps2. Ventral apophyses of tibiae and genua I, II weakly dis-tinct.

Remarks

According to the original description by Trouessart (1899: 6),the clearest feature distinguishing Protolichus favettei was thepair of small lobes of triangular form situated on the anteriormargin of terminal cleft (Figs 5; 6A, B). Originally, Trouessartlisted four parrot hosts for this species: Nestor notabilis andN. meridionalis (Gmelin, 1788) from New Zealand, Psepho-tus xanthorrhous (recent valid name Northiella haematogas-ter haematorrhous) from Australia (Psittacidae), and Micro-glossum aterrimus (recent name Probosciger aterrimus) (Ca-catuidae) from New Guinea (Table I). Further, Favette and

Table I. Data and contents of slides designated by Trouessart as Protolichus favettei

No. Host (original spelling) Identification and remarks by Trouessart Designation in recent study

34G6* Microglossum aterrimus Protolichus favettei (type) Calyptolichus favettei (lectotype male, paralectotype female)

34G7* Nestor meridionalis Protolichus c.f. favettei sp. nov. Nestorilichus atyeoi sp. nov. (holotype male, paratypes: 1 male and 1 female)

34G8* Nestor notabilis Protolichus favettei sp. nov. Calyptolichus favettei (paralectotype male)34G9 Nestor montanus Protolichus favettei sp. nov. Nestorilichus atyeoi sp. nov. (paratype male)34G10* Psephotus xanthorrhous Protolichus favettei var. cordifora Calyptolichus favettei (paralectotype male)

*Specimens of the original type series of P. favettei.

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Trouessart (1904) added one more host, Nestor montanusBuller (a synonym of N. meridionalis); as for the sample pre-viously collected from Psephotus xanthorrhous, these authorsused the sole male for the description of a separate subspecies,Protolichus favettei cordifora. Regarding the hosts Psephotusxanthorrhous and Microglossum aterrimus, Favette and Trou-essart (1904) considered these associations to be accidentaland suggested that the true hosts of these mites are parrots ofthe genus Nestor. A curious thing regarding the four host asso-ciations given in the original papers (Trouessart 1899) is thatnone of them represents true hosts of the mite species charac-terized by the pair of angular extensions in terminal cleft. Aswas discovered by Gaud and Atyeo (1996), who treated thisspecies under the name �Apexolichus cordifora�, and was alsoconfirmed by our collecting, the true host of the mite species

with the mentioned structure of opisthosoma is the cockatooCalyptorhynchus funereus (Cacatuidae). As it is easy possibleto guess, Trouessart (1899) made such extreme errors in hostassociations, because he always collected materials from drybird skins, and the mites on non-Calyptorhynchus hosts weresimply museum-based contaminations.

Specimens of �Protolichus favettei� from each host men-tioned in the two primary papers (Trouessart 1899, Favetteand Trouessart 1904) have fortunately been retained in theTrouessart collection (Table I). We have found that this mate-rial actually contains two species that should be treated as dif-ferent pterolichine genera, although the general shapes of opi-sthosomal lobes in males of these species look superficiallysimilar. In one species, the males truly possess a pair of smalltriangular extension in the terminal cleft (Figs 5; 6A, B) that

Fig. 8. Nestorilichus atyeoi sp. nov., male: A � dorsal view, B � ventral view

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correspond to the original description; while in other speciesthey are absent (Figs 8; 9A, B). The first species was detect-ed on Probosciger aterrimus (TRT 34G6) and Nestor nota-bilis (TRT 34G8), and the second species was detected onlyon Nestor species (TRT 34G7, 34G9). Formally, an exemplarfrom any one of these species may be chosen for fixation ofthe lectotype and nomenclature type (the type specimen bear-ing the name) of P. favettei. Following the Article 74 of ICZNand its recommendations, we consider the former specimens,corresponding to the original description and subsequentredescription (Trouessart 1899, Favette and Trouessart 1904),as the true Protolichus favettei. Taking into consideration thatthe slide from Probosciger aterrimus also bears the Troues-sart�s remark �typ� made by the hand of Trouessart, we des-ignate the male specimen in the slide (TRT 34G6) as the lec-totype of P. favettei, the female specimen from this slide andthe male specimen from Nestor notabilis (TRT 34G8) as theparalectotypes. The second pterolichid species is described inthe present paper under the name Nestorilichus atyeoi gen.nov., sp. nov. It is worthy to note that Favette and Trouessart(1904: 153) did recognize a variation of P. favettei fromN. montanus (TRT 34G9), which differed by �some detailsof terminal cleft� from P. favettei of their latest concept, butthis variation was neither named nor characterized by theauthors.

The subspecies Protolichus favettei cordifora Favette etTrouessart, 1904 was based on a sole male from Psephotusxanthorrous, which initially (Trouessart 1899) was includedin the type series of P. favettei (Table I ). Under the Article73.1.2 of ICZN, this specimen is the holotype of P. f. cordifo-ra by monotypy. Favette and Trouessart (1904: 153, fig. 23)stressed in the original description of P. f. cordifora that thisspecies differs from the nominal subspecies by only the dark-coloured heart-like sclerite in area of epimerites I; this �u-nique feature� caused the specific epithet for this subspecies.Examination of that specimen (TRT 34G10), which was re-mounted and cleaned by W.T. Atyeo, does not show any darksclerite in the mite; apparently, this �heart-like sclerite� wasjust a grain of detritus in the original slide. By all other char-acters, this specimen completely corresponds to the speci-mens of P. favettei of the concept fixed in the present paper;therefore we designate P. f. cordifora as a junior synonym ofP. favettei.

When Gaud and Atyeo (1996) established a new genusApexolichus with eight species, which united representativesof the �Protolichus lunula� group (sensu Favette and Troues-sart), characterized by unmodified legs I and II in males, theytreated Protolichus favettei and P. cordifora as the two validspecies, although the only one new combination, Apexolichuscordifora, was formally given in the text. Gaud and Atyeo alsopointed that the true host of A. cordifora (sensu these authors)is Calyptorhynchus funereus (Cacatuidae). Under the nameP. favettei they apparently treated the species associated withNestor species. As far as is possible to understand the conceptof these authors, they ignored the clear characteristics of P. fa-vettei mentioned in the original and subsequent re-descrip-

tions (Trouessart 1899, Favette and Trouessart 1904) (Figs 5;6A, B), and applied the name P. cordifora to the mite speci-mens with triangular extensions in the terminal cleft, probablybecause they wanted to retain both �favettei� and �cordifora�as valid specific names. We have found that Atyeo had madethe hand mark �lectotype� of P. favettei on the specimen fromN. montanus (TRT 34G9). This is inappropriate according tothe ICZN. This specimen does not belong to the syntype seriesconsisting of samples from the four primary mentioned hosts(Trouessart 1899), because this sample was added later (Fa-vette and Trouessart 1904). In that case it could be only a�neotype� and not a �lectotype�. It is a moot point, however,because these hand-made marks in the collection were neverpublished and thus have no official authority.

Nestorilichus gen. nov.

Type species: Nestorilichus atyeoi sp. nov. Description (both sexes): Prodorsal shield occupying me-

sal part of prodorsum and extending beyond scapular setae se,si; anterior part strongly sclerotized; posterior part (beyondscapular setae) poorly sclerotized (Figs 8A; 10A). Scapularshields as narrow oblique bands. Hysterosoma with extensivehysteronotal shield; humeral shields with poorly distinguishedmargins, bearing bases of setae c2 and cp. Two vertical setaevi present, their bases close to each other. Both pairs of scapu-lar setae se, si very close to midline. Setae si, c2, d2, e2 spic-uliform; setae c3 piliform; setae h1 absent. Cupules ia pos-tero-lateral to bases of setae c2; cupules im posterior to hys-teronotal gland openings gl. Each tarsus with two minute apicodorsal spines (in males one spine with bidentate apex);tarsi I, II with blunt-angular ventral margin, approximately 1.5times longer than corresponding tibiae; solenidion ω1 closerto base of tarsi I, II than to apex; setae ba slightly distal to cor-responding solenidion ω1; solenidion σ2 of genu I present;ambulacral discs minutely dentate on distal margin.

Male: Gnathosoma trapezoidal in general shape, lateralmargins of subcapitulum with small rounded processes; palpswithout lateral extensions. Scapular setae se, si considerablymoved to midline. Opisthosomal lobes well-developed, nar-row, bluntly rounded apically, widely separated from eachother by terminal cleft, anterior margin of terminal cleft con-cave; interlobar membrane developed only in anterior part ofterminal cleft; terminal and lateral membranes absent (Fig.9B). Setae f2 setiform with narrow membranous extension;setae ps2 setiform; setae ps1 crescent-shaped, membranous;setae e1 setiform, situated posterior to level of setae e2.Epimerites I fused as a Y. Bases of epimerites II with heavilysclerotized ball-like inflations. Coxal fields I�IV open, with-out large sclerotized areas. Bases of trochanters I, II flankedby narrow sclerotized bands connecting bases of respectiveepimerites. Paragenital apodemes present, long and well scler-otized, spreading from level of inner tips of epimerites III toopisthosomal lobes; anterior ends convergent towards mid-line, posterior ends of apodemes fused with opisthoventralshields; middle parts of apodemes connected by wide trans-

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verse bridge separating genital and anal fields (Fig. 9A). Analsuckers circular, corolla dentate, each sucker surrounded bymembrane with radial striation. Genital apparatus at level oftrochanters IV. Genital papillae situated anterior to genitalapparatus, off paragenital apodemes. Cupules ih present. LegsII slightly longer than legs I; legs III and IV similar in size, nothypertrophied; segments of legs I, II without any processes.Tarsus IV elongated, similar in form and size to tarsus III;modified setae d, e of tarsus IV as barrel-shaped suckers.

Female: Gnathosoma trapezoidal, subcapitulum withoutlateral spines. Both pairs of scapular setae noticeably moved to

midline, setae si approximately equidistant from midline andrespective setae se. Hysteronotal shield split into pygidial frag-ment and main body (Fig. 10A). Lateral sclerotized bands ofhysterosoma absent. Setae f2, spiculiform, short; setae ps1,ps2, setiform. Epimerites I free. Bases of epimerites II withoutball-like inflations. Lateral sclerotized bands absent. Ovipor-us situated at level of humeral shields and trochanters III.Epigynium present, represented by thick bow-shaped sclerite.Copulatory opening terminal, on small cone-like extension.

Hosts: Parrots of the genus Nestor Lesson, 1830 (Psit-tacidae, Psittacinae, Nestorini).

Fig. 9. Nestorilichus atyeoi sp. nov., details of male: A � opisthosoma, ventral view; B � opisthosoma, dorsal view; C � leg I; D � leg II; E �leg III; F � leg IV

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Within the Protolichus generic group, Nestorilichus is mostsimilar to Apexolichus based on the structure of prodorsal andhysteronotal shields and lateral setae c2, d2, e2 in both sexes,and barrel-like form of setae d, e of tarsus IV in males (Figs8A; 9F; 10A). The new genus differs from Apexolichus by thefollowing combination of characters: in males, ventral spine-like processes of tibiae and genua I, II are absent, opisthoso-mal lobes narrow, without terminal membranes, interlobarmembrane is developed only on anterior margin of the termi-nal cleft; in females, bases of epimerites II without sclerotizedinflations.

Etymology

The generic epithet is a combination of contractions of thegeneric name of the hosts and Protolichus.

Nestorilichus atyeoi sp. nov.

Pterolichus (Protolichus) favettei Trouessart (1899): 6 (inpart); Gaud and Atyeo (1996): 117. Protolichus (Mesolichus)favettei: Favette and Trouessart (1904): 152 (in part).

Type material: Holotype male, paratypes: 1 male and 1female (TRT 34G7) ex Nestor meridionalis (Gmelin, 1788),Psittacidae, New Zealand, no other data; 1 male paratype(TRT 34G9) ex N. meridionalis (N. montanus Buller in orig-inal label), New Zealand, no other data.

Additional material: 1 male, 1 female (ZISP 4359, 4360)ex N. meridionalis New Zealand, no other data; 2 males(UMB-15089), same host species, New Zealand, Stephen Is-land, 1896, coll. unknown.

Description (male, holotype, measurements for 2 para-types in brackets): Idiosoma: length from margin to tips ofopisthosomal lobes 585 (576�582), width 310 (286�310).Gnathosoma: lateral margin of subcapitulum with short spine-like extension, ventral surface of subcapitulum with trans-verse line. Prodorsal shield: anterior part almost rectangular;posterior part as transverse sclerite with poorly outlined bor-ders, its posterior margin slightly concave; total length ofprodorsal shield 161 (149�155), greatest width of anterior part90 (82�86). Setae si 70 (75�77) long.

Distance between scapular setae: se:se 68 (58�69), si:si 30(25�31). Scapular shields not connected to inflations of epi-merites II. Hysterosoma 420 (396�409) long. Hysteronotalshield: length from anterior margin to lobar apices 422 (372�415), width at anterior margin 228 (202�223), surface withnetwork pattern in central part. Opisthosomal lobes attenuateto apex, lobar apex with short and strongly sclerotized termi-nal membrane; lateral margins of terminal cleft almost paral-lel, anterior end of the cleft approximately as blunt-angle,interlobar membrane narrow, developed only in anterior partof the cleft (Figs 8A; 9B); total length of cleft 80 (83�86),width 71 (68�81). Supranal concavity absent. Subhumeralsetae c3 40 (40�45) long. Lateral setae c2 spiculiform, 62(60�65) long, d2 90 (90�92), e2 83 (broken in all paratypes).

Setae f2 setiform with membranous extension along all itslength. Setae ps1 crescent-shaped, membranous (Fig. 9B).Dorsal measurements: c2:d2 136 (130�132), d2:e2 162 (152�162), e2:h3 80 (81�84), e2:e1 17 (15�16), e2:e2 123 (111�114), e1:e1 86 (78�86), ps1:ps1 71 (78�82). Cupules imslightly posterior to level of openings gl. Sternum as half oftotal length of epimerites I. Bases of epimerites II with heavi-ly sclerotized inflations. Genital apparatus at level of tro-chanters IV, 34 × 20 (30�34 ×16�20). Genital papillae slight-ly anterior to genital apparatus. Paragenital apodemes: ante-rior tips almost extending to level of setae 3a, middle partsposterior to genital apparatus connected to each other by widetransverse bridge; this bridge forms the anterior margin ofanal field (Fig. 9A). Ventral measurements: 3b:3a 16 (14�16),3a:g 43 (46�48), g:4a 70 (60�65), 4a:ps3 85 (82�93). Cupulesih slightly anterior to level of setae h2. Diameter of anal suck-ers 23 (23�26). Legs II slightly longer than legs I. Tarsus Iwith ventral ledge in basal third, ventral setae la, ra, wa clos-er to base of this segment (Fig. 9C). Tibiae and genua I, IIwithout ventral processes. Setae d, e of tarsus IV barrel-shaped with distinct apical cap, both situated approximately atmidlevel of segment (Fig. 9F).

Female (paratype): Idiosoma: length 601, width 316. Pro-dorsal shield: anterior piece as almost rectangular longitudinalplate, posterior piece as weakly sclerotized transverse platebearing scapular setae si, se, total length of shield along mid-line 137, greatest width of anterior part 65 (Fig. 10A). Setaesi spiculiform, 84�85 long. Distance between scapular setae:se:se 71, si:si 37. Scapular shields connected by inner endswith sclerotized areas anterior to trochanter II. Hysterosoma403 long. Hysteronotal shield: length of main body 280, widthof anterior part 230, total length 415, anterior margin straight,surface with network ornamentation; pygidial fragment withconvex anterior margin. Setae c3 37�40 long. Setae d2, e2thick spiculiform, f2 thin spiculiform. Length of lateral hys-teronotal setae: c2 43, e2 26, d2 and f2 broken in paratype, inadditional material 57�60 and 23�25. Dorsal measurements:c2:d2 120, d2:e2 144, e2:h3 74, e2:e2 128, h3:h3 57. Cupulesim posterior to level of openings gl. Epigynium bow-shaped,40 × 70. Genital papillae between levels of setae 3a and g.Cupules ih indistinct.

Etymology

The species is named in honor of Prof. W.T. Atyeo (TheUniversity of Georgia, Athens, USA), the preeminent worldexpert on feather mites.

Remarks

The new species described above, Nestorilichus atyeoi, isapparently restricted to parrots of the genus Nestor (Psitta-cinae, Nestorini), as suggested both by the materials of Trou-essart and by additional collections. Nestorilichus atyeoi is thesecond pterolichid mite species that we recognized in the orig-inal syntypes series of Pterolichus (Protolichus) favettei Trou-essart, 1899 (see above remarks on Calyptolichus favettei).

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Aknowledgements. The authors wish to thank Dr. Mark Judson(Muséum National d�Histoire Naturelle, Paris, France) for loaningmaterials from the Trouessart collection, Dr. Peter-René Becker(Uebersee Museum, Bremen, Germany) for making available com-parative materials from Nestor meridionalis, and Dr. Heather C.Proctor (University of Alberta, Canada) for critically reviewing themanuscript. The study was supported in part by the RussianFederation for Basic Research, grant 07-04-00426a for SVM.

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Fig. 10. Nestorilichus atyeoi sp. nov., female: A � dorsal view, B � ventral view

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