Systematic revision of the feather mite genera Apexolichus Gaud et Atyeo and Titanolichus Gaud et...

Systematic revision of the feather mite genera Apexolichus Gaud et Atyeo and Titanolichus Gaud et Atyeo

(Astigmata, Pterolichidae), parasites of parrots of the Old World (Psittaciformes, Psittacidae)

Jacek Dabert1*, Serge V. Mironov2 and Rainer Ehrnsberger3

1Department of Animal Morphology, A. Mickiewicz University, Umultowska 89, 61-614, Poznañ, Poland; 2Zoological Institute, Russian Academy of Sciences, Universitetskaya quay 1, 199034, Saint Petersburg, Russia;

3Institute of Nature Conservation and Environmental Education, University of Vechta, Driverstrasse 22, D-49377, Vechta, Germany

AbstractWe present a systematic revision and phylogenetic analysis of feather mites of the closely related genera Apexolichus Gaud etAtyeo, 1996 and Titanolichus Gaud et Atyeo, 1996 (Pterolichidae, Pterolichinae). New diagnoses for Apexolichus andTitanolichus and keys to all known species are provided. We redescribe all species of the genus Protolichus Mégnin etTrouessart, 1884, described by E. Trouessart over a century ago, and which have been transferred to these genera. Two newspecies are described: Apexolichus psephoti sp. nov. from Psephotus varius Clark A.H., 1910 (type host) and P. haematonotus(Gould, 1838); and Titanolichus triangulifer sp. nov. from Barnardius zonaris (Shaw, 1805) (type host) and Northiella haemato-gaster (Gould, 1838). New synonymies are proposed: Apexolichus splendens (Favette et Trouessart, 1904) syn. nov. is a jun-ior synonym of A. velifer (Trouessart, 1899); A. distensis (Favette et Trouessart, 1904) syn. nov. is a junior synonym of A. affi-nis (Mégnin et Trouessart, 1884). Apexolichus platycerci (Mironov, Dabert et Proctor, 2003) comb. nov. is a new combinationfor this species, which was formerly placed in the genus Titanolichus. Phylogenetic analysis (MP) confirms the monophyly ofthe genera Apexolichus and Titanolichus. Both of these genera, together with the monotypic genus Sideroferus Gaud et Atyeo,1996 (being a sister group of Titanolichus), form a clade within the Protolichus generic group and are restricted in their hostassociations to parrots of the tribe Platycercini (Psittacidae, Psittacinae), members of which occur in the Australia and Indo-Malayan region. Representatives of each genus are restricted to a particular genus or a few genera of Platycercini. Host asso-ciations of these genera are briefly discussed.

KeywordsFeather mites, Pterolichidae, phylogenetic analysis, host-parasite associations, systematic revision, new species, Psittaciformes

*Corresponding author: [email protected]

Introduction

Feather mites are the most common and numerous ectosym-biotic mites of the suborder Astigmata that live permanentlyon plumage and skin of birds. The approximately 2500 speciesin 450 genera and 34�38 families are known throughout theworld and recorded from almost all avian orders; nevertheless,experts consider that the currently known number of speciesrepresents not more than 15% of the actual extant fauna (Gaudand Atyeo 1996, Proctor 2003).

With about 400 species, the feather mite family Pteroli-chidae is the most species-rich family of the superfamily Pte-rolichoidea. Mites of this family are adapted to inhabit feath-

ers with large vanes, i.e. flight and covert feathers of the wingand tail feathers, and are associated exclusively with non-passerine avian orders. Among feather mites occurring on par-rots (Psittaciformes), the family Pterolichidae is the most nu-merous with over 120 species and 28 genera known fromthese hosts worldwide. Pterolichids restricted to parrots com-prise three morphologically distinct generic groups (Gaud andAtyeo 1996; Mironov and Pérez 2003; Mironov et al. 2003a,b). Among them, the Protolichus group with about 70 speciesin 21 genera is the most numerous and diverse. The presentpaper provides a taxonomic revision and phylogenetic analy-sis of two pterolichid genera, Apexolichus Gaud et Atyeo,1996 and Titanolichus Gaud et Atyeo, 1996 from the Pro-

DOI: 10.2478/s11686-008-0007-x© 2008 W. Stefañski Institute of Parasitology, PASActa Parasitologica, 2008, 53(1), 46–80; ISSN 1230-2821

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Systematic revision of genera Apexolichus and Titanolichus

tolichus generic group. Both genera are associated withpsittacid parrots (Psittaciformes, Psittacidae) in the Australianand Indo-Malayan regions.

The first systematic review focused on pterolichid mitesassociated with parrots was the revision of the genus Proto-lichus Mégnin et Trouessart, 1884 by Favette and Trouessart(1904); in that time this artificial genus incorporated all large-sized pterolichids associated with parrots. In the second partof the 20th century, investigations of diversity of the Proto-lichus group were carried out most extensively in the NewWorld (Atyeo et al. 1984; Atyeo 1988, 1989a-c; Atyeo andPérez 1990). To a lesser extent, these mites were explored inAfrica (Gaud 1980). Much less is known about mites of par-rots distributed in Australia and the Indo-Malayan region(Atyeo and Pérez 1982, Mironov et al. 2003b, Dabert et al.2006, Mironov and Dabert 2007). Up to the present study, 21species in eight genera have been recorded from psittaciformsin the two latter regions: Apexolichus (5 species), Calyptoli-chus Mironov et Dabert, 2007 (1 species), LoriprotolichusMironov et Dabert, 2007 (2 species), Nestorilichus Mironovet Dabert, 2007 (1 species), Protolichus (6 species), Sidero-pherus Gaud et Atyeo, 1996 (1 species), Titanolichus (3 spe-cies), and Uropsittacolichus Atyeo et Pérez, 1982 (2 species).

The genera Apexolichus (9 species) and Titanolichus (1species) were established by Gaud and Atyeo (1996) in thecourse of the taxonomic revision of supraspecific taxa offeather mites of the world. Both genera were based on speciesoriginally described by Trouessart and coauthors (Mégnin andTrouessart 1884, Trouessart 1899, Favette and Trouessart 1904)in the context of the artificial genus Protolichus. Using theconcept of Gaud and Atyeo (1996), reliable discrimination ofthese two genera is quite problematic, because it was basedonly on structure of tarsi II in heteromorph males: in Titano-lichus, tarsus II has dorsobasal apophysis, while in Apexo-lichus, apophyses on this segment are absent. In practice,identification is complicated by rather continuous polymor-phism of males. Mironov et al. (2003b) provisionally sug-gested discrimination of these genera based on the presenceand absence of solenidion σ2 on genua II. However, furtherinvestigation (Dabert et al. 2006) showed that this featurevaries among species of both genera. Finally, re-investigationof three most odd-looking �Apexolichus� species resulted inthem being moved into three separate genera, Calyptolichus,Loriprotolichus and Nestorilichus (Mironov and Dabert 2007).Nevertheless, the problem remained: whether Titanolichusshould be synonymized with Apexolichus or whether thesetaxa are clearly separate.

In the course of our study dealing with feather mites asso-ciated with parrots of the Old World (Mironov et al. 2003a, b,2005; Dabert et al. 2004, 2006) we accumulated a number ofrepresentatives of the Protolichus group from Indo-Malayanand Australian regions and were able to re-examine all re-tained material of the genus �Protolichus� from the Trouessartcollection. The material from this collection contained typespecimens or syntype series of all species referred by Gaudand Atyeo (1996) to Apexolichus and Titanolichus. The pres-

ent paper gives the taxonomic revision of the genera Apexo-lichus and Titanolichus and includes a phylogenetic analysisbased on morphological characters, re-examination of allaccessible type specimens, revision of generic diagnoses, con-structing of keys, redescriptions of old species, and a reviewof host associations of these genera with psittacines of theAustralia and Indo-Malayan regions.

Materials and methods

The most important materials used in the study were loanedfrom the Trouessart collection (specimens mounted on slides)deposited in the Muséum National d�Histoire Naturelle (Paris,France). Other material was collected by the authors from drymuseum skins of parrots in the following museums: ÜberseeMuseum (Bremen, Germany) and State Museum, Departmentof Natural History (Oldenburg, Germany). Mites from skinswere sampled by the �scratching technique� (Gaud and Atyeo1996) and preserved in 70% ethanol. For light microscopymite samples were processed in 10% lactic acid (3�5 days attemperature 40°C), and then mounted on slides in Faure medi-um (Evans 1992). Remounting of old microslides, the medi-um of which was dissolved by distilled water, was carried outby the same processing. Diagnoses and descriptions of taxafollow the recent standard schemes used for pterolichid mitesof the group in question (Gaud and Atyeo 1996; Mironov et al.2003b, 2005). All measurements in species descriptions aregiven in micrometres.

It is necessary to add that materials of Trouessart had beenpreviously examined by Prof. Warren T. Atyeo (University ofGeorgia, Athens, USA) in the 1980s, and lectotypes or para-lectotypes were marked on slide labels for many species of�Protolichus�. Unfortunately, any data concerning designa-tions of lectotypes or simply syntype series for species refer-red to the genera Apexolichus and Titanolichus have neverbeen published and formally they are not valid. Nevertheless,in most cases we followed the concept of Atyeo in designationof lectotypes.

Abbreviations used in accession collection numbers andfor pointing out the depositories of specimens: AMU � A.Mickiewicz University (Poznañ, Poland), UMB � ÜberseeMuseum (Bremen, Germany), QM � Queensland Museum(South Brisbane, Australia), TRT � collections of the MuséumNational d�Histoire Naturelle (Paris, France), ZISP � Zoologi-cal Institute, Russian Academy of Sciences (Saint Peters-burg, Russia). Taxonomic system and Latin names of hostsused in the present study follow Del Hoyo et al. (1997).

Phylogenetic analysis of the genera Apexolichus andTitanolichus included all species formally referred to thesegenera by recent investigators (Gaud and Atyeo 1996, Miro-nov et al. 2003b, Dabert et al. 2006). Six other genera of theProtolichus group, associated with parrots from the Australianand Indo-Malayan regions (Calyptolichus, Loriprotolichus,Nestorilichus, Protolichus, Sideropherus, and Uropsittacoli-chus), were also included as representatives of potentially sis-

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ter groups or closely related outgroups. Among them, the gen-era Loriprotolichus and Protolichus were each represented bytwo species, and the genus Uropsittacolichus was represent-ed by one species; the remaining genera are monotypic. Thus,all genera of the Protolichus group known from parrots inAustralia and Indo-Malaya were used in the analysis. Theprimitive pterolichid genus Mayracarus Atyeo, 1992 (Ptero-lichinae) associated with megapodid hosts (Galliformes, Me-gapodiidae) was used as a distant outgroup and to root the tree.

Reconstruction of phylogenetic relationships between taxawas based on morphological characters and carried out usingthe maximum parsimony method as implemented in PAUP 4software (Swofford 1998). The maximum parsimony analysiswas performed with branch-and-bound algorithm, which guar-antees finding the most parsimonious tree. All characters wereequally weighted and treated as unordered. Support for theclades was estimated by decay/support indices (Bremer 1988)using AutoDecay software (Eriksson 1998).

Results

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

Apexolichus Gaud et Atyeo, 1996

Type species: Pterolichus (Protolichus) affinis Mégnin etTrouessart, 1884

Description: Both sexes. Prodorsal shield occupying mesalpart of prodorsum and extending to or beyond scapular setae;anterior part strongly sclerotized; posterior part poorly scle-rotized or not expressed (Figs 1B; 3B). Scapular shields rep-resented by poorly sclerotized bands or small plates on later-al margins of propodosoma. Hysterosoma with extensive hys-teronotal shield. Humeral shields well-developed. Two verti-cal setae vi present, their bases close to each other. Scapularsetae moved to midline, distance between setae se approxi-mately equal to or less than width of gnathosoma. Setae si andc2 spiculiform; setae c3 setiform; setae h1 absent. Cupules iapostero-lateral to bases of setae c2, situated on humeral shield;cupules im variously positioned in relation to hysteronotalgland openings gl, but always close to that level. Each tarsuswith two minute apicodorsal spines, one of them may be sit-uated on proximal part of ambulacral stalk and have a biden-tate apex; tarsi I with blunt angular ventral margin, tarsi I, IIapproximately 1.5�2 times longer than corresponding tibiae;solenidion ω1 closer to base of tarsi I, II than to apex; setaeba slightly distal to corresponding solenidion ω1; condy-lophores heavily developed; ambulacral discs minutely den-tate on distal margin. Solenidion σ2 of genu I present or absent.

Male: Epimerites I free. Bases of epimerites II with orwithout ball-shaped and heavily sclerotized inflations (Figs1A; 4A). Gnathosoma trapezoidal in shape, lateral marginsblunt-angular without lateral spines; basal podomeres of palps

with or without lateral expansions. Opisthosomal lobes well-developed, longer than wide, with blunt posterior ends, wide-ly separated from each other by semi-ovate terminal cleft;anterior margin of terminal cleft concave or blunt-angular;interlobar membrane present, occupies most part of terminalcleft; terminal membrane separated from interlobar mem-brane, lateral membranes commonly present (absent in A. affi-nis). Setae d2, e2 long, spiculiform; setae f2 setiform or withnarrow membranous extension, commonly curved; setae ps2setiform, always longer than setae f2; setae ps1 wide foliform,membraneous; setae e1 setiform, situated posterior to level ofsetae e2. Paragenital apodemes present, long, spread com-monly from midlevel of coxal fields III to opisthosomal lobeswhere they fuse with opisthoventral shields; middle parts ofapodemes distant from each other, inner margins of these partswith sclerotized areas or connected to each other by widetransverse bridge. Anal suckers circular, corolla dentate, eachdisc surrounded by membrane with radial striation. Genital pa-pillae situated between anterior parts of paragenital apodemesor on their inner margins. Coxal fields I�IV open, withoutlarge sclerotized areas. Bases of trochanters I, II flanked bynarrow sclerotized bands connecting bases of respectiveepimerites. Cupules ih present. Legs II slightly (homeomorphmales) or noticeably longer than legs I (heteromorph males);legs III and IV similar in size, not hypertrophied. Tibiae andgenua I with spine-shaped apicoventral apophyses; tibiae andgenua II with more weakly expressed apophyses than on legsI or almost lacking them (Figs 2A, B; 6A, B; 10A, B). Setaera, la of tarsus I proximal or at the same level as seta wa. TarsusIV short, not more than 1.5 times longer than tibia IV, similarin size to tarsus III; modified setae d, e of tarsus IV as smallbarrel-shaped structure with apical disc. Polymorphism ofmales continuous, expressed mainly in size of legs I, II andtheir apophyses.

Female: Epimerites I free. Bases of epimerites II with ball-shaped inflations (Fig. 3A). Gnathosoma trapezoidal as inmales. Hysteronotal shield split into pygidial fragment andmain body; lateral sclerotized bands not split from the shield.Oviporus at level of coxal fields III. Epigynum shaped as thickbow-like sclerite or absent. Setae d2, e2, f2 spiculiform or seti-form, ps1, ps2 simple setiform. Copulatory opening terminal,on small cone-like extension. Apicoventral apophyses of tib-iae and genua I, II developed much more poorly than in males, commonly as blunt extensions, or may be practically indis-tinct.

Hosts: parrots of the genera Lathamus Lesson, 1830, Pla-tycercus Vigors, 1825, Prosopeia Bonaparte, 1854, and Pse-photus Gould, 1845 (Psittacinae, Platycercini).

Key to species

Males

1. Basal segment of palps with lateral extension directedanteriorly and bearing setae dp2 (Fig. 5C). Tibia I withdorsobasal blunt-angular extension (Fig. 6A). Setae d2

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short (45�55), approximately equal to half-distance be-tween setae se ........................ A. velifer (Trouessart, 1899)

− Basal segment of palps without lateral extension or withlateral crest extending backward. Tibia I without dor-sobasal blunt-angular extension. Setae d2 long (65�120),equal or longer than distance between setae se ................ 2

2. Basal segment of palps with lateral crest extending back-ward (Fig. 16C) ............................................................... 3

− Basal segment of palps without lateral extension ............ 53. Tarsus II with large rounded dorsobasal extension bearing

solenidion ω1 and seta ba. Femur I with spine-like ventralprocess. Tibia and genu II without apicoventral spine-like

process (Fig. 16A, B) ................................... A. platycerci(Mironov et al., 2003b)

− Tarsus II without any dorsobasal extension (Figs 10B;19B). Femur I without spine-like ventral process. Tibiaand genu II with apicoventral spine-like process ............. 4

4. Tarsus I with small spine-like retrograde dorsobasalprocess (Fig. 10A). Setae d2 as long as half the distancebetween levels of setae d2 and e2 .................... A. lathamiMironov et al., 2003b

− Tarsus I without dorsobasal spine (Fig. 19A). Length ofsetae d2 almost equal to distance between setae d2 and e2............................................................. A. psephoti sp. nov.

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Fig. 1. Apexolichus affinis (Mégnin et Trouessart, 1884), male: A � ventral view, B � dorsal view. Setal designations after Gaud and Atyeo(1996)


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5. Interlobar membrane occupies entire terminal cleft, withshallow incision on posterior margin; dorsal surface of thismembrane smooth (Figs 11B; 12A). Genital apodemesconnected to each other by large sclerotized bridge poste-rior to genital apparatus. Solenidion σ2 of genu I absent.Setae d2 shorter than c2 ................................ A. neglectus(Favette et Trouessart, 1904)

− Interlobar membrane with deep triangular incision, dorsalsurface of this membrane with a few striae running paral-lel to the free margin of membrane (Fig. 1B). Genitalapodemes not connected to each other posterior to genitalapparatus. Solenidion σ2 of genu I present. Setae d2 equalor longer than setae c2 ......................................... A. affinis(Mégnin et Trouessart, 1884)

Females

1. Epigynum present (Fig. 13D) .......................................... 2− Epigynum absent (Fig. 13B) ............................................ 4

2. Solenidion σ2 of genu I absent ....................... A. platycerci(Mironov et al., 2003b)

− Solenidion σ2 of genu I present, albeit very minute in size......................................................................................... 3

3. Length of idiosoma 440�465, length of setae si 64�70(Fig. 19D) ........................................... A. psephoti sp. nov.

− Length of idiosoma 470�490, length of setae si 80�90(Fig. 3B) ............... A. affinis (Mégnin et Trouessart, 1884)

4. Setae si longer than distance between setae se. Tips ofsetae d2 extending to level of cupules im (Fig. 13A).Solenidion σ2 of genu I absent ....................... A. neglectus(Favette et Trouessart, 1904)

− Setae si equal or shorter than distance between setae se.Tips of setae d2 not extending to level of cupules im.Solenidion σ2 of genu I present, minute ........................ 5

5. Setae c2 equal or longer than distance between setae se(Fig. 7C). Setae e2 1.4�1.5 times longer than setae d2.Idiosoma 400�455 long .... A. lathami Mironov et al., 2003b

− Setae c2 shorter than distance between setae se (Fig. 7A).

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Fig. 2. Apexolichus affinis (Mégnin et Trouessart, 1884), legs of male: A � leg I, dorsal view; B � leg II, dorsal view; C � leg III, dorsal view;D � leg IV, dorsal view; E � tarsus IV, ventral view. Setal designations after Gaud and Atyeo (1996)


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Setae d2 and e2 subequal in length. Idiosoma 540�600long ....................................... A. velifer (Trouessart, 1899)

Apexolichus affinis (Mégnin et Trouessart, 1884) (Figs 1�3)

Pterolichus (Protolichus) affinis Mégnin et Trouessart, 1884:262.Apexolichus affinis: Gaud & Atyeo, 1996: 117, fig. 339.Protolichus chiragricus: Favette & Trouessart, 1904: 142 (inpart).Protolichus velifer: Favette & Trouessart, 1904: 144 (in part).

Protolichus distensis Favette et Trouessart, 1904: 147, fig. 18,syn. nov.Apexolichus distensis: Gaud & Atyeo, 1996: 117.

Material examined: Male lectotype, 10 female paralectotypes(TRT 34G2), ex Platycercus elegans (Gmelin, 1788) (�Platy-cercus pennantii� in original label), Australia, no other data;lectotype designated here; 4 males and 4 females (UBRA.1166) ex Pl. elegans, Australia, Victoria, 8 November 1905,coll. unknown; 1 male, 1 female (QM, no number) ex Pl. cale-donicus (Gmelin, 1788), Australia, Tasmania, Richmond, 12

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Fig. 3. Apexolichus affinis (Mégnin et Trouessart, 1884), female: A � ventral view; B � dorsal view


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July 1978, O. Seeman; 1 male, 4 females (UMB 1/1047) exPl. eximius (Kuhl, 1820), Australia, S. Australia, AdelaideZoo, May 1956, dead in transport; 1 male, 3 females (TRT34G12) ex Pl. flaveolus (Gould, 1837), Australia, no otherdata; 2 males, 4 females (UMB 23365) ex Pl. icterotes (Kuhl,1820), Australia, S. Australia, Gawler Range, August 1955,H.O. Wagner.

Diagnosis: Solenidion σ2 of genua I present. Male: basalsegment of palps without lateral extensions; hysteronotalshield monotonously punctured, terminal cleft with roundedanterior margin; interlobar membrane with deep triangularincision, dorsal surface of this membrane with a few striaegoing parallel to free margin of membrane; sclerotized areason inner margins of paragenital apodemes not forming entiretransverse bridge; setae d2 longer than c2; bases of epimeritesII without inflations; tibiae and genua I, II with distinct spine-shaped ventral processes; setae d, e of tarsi IV approximatelyat midlevel of segment. Female: epigynum present, setae d2,e2 thin spiculiform, cupules im anterior to openings gl, lengthof idiosoma 470�505, length of setae: si 80�90, c2 65�75 (inspecimens from P. icterotes: si 65�75, c2 60�65).

Description: Male (lectotype). Idiosoma, length × width,440 × 211 (in additional material, 4 males from P. elegans,idiosomal size 445�470 × 215�240). Gnathosoma: basal seg-ment of palps without lateral expansions; ventral side of sub-capitulum without spines. Prodorsal shield: longitudinal plateenlarged in anterior part, not extending to row of scapularsetae; length of shield 155, greatest width 43. Striated surfacearound scapular setae se, si slightly punctured. Setae si spic-uliform, 68 long. Distance between scapular setae: se:se 58,si:si 26. Scapular shields very small, developed only on lateralmargins of propodosoma. Hysterosoma 328 long. Hysteronotalshield: greatest length 314, greatest width 236, anterior marginstraight, surface uniformly punctured. Opisthosomal lobesroughly triangular, with narrowed and rounded posterior ends,with small terminal membrane mesal to bases of setae h3, with-out lateral membranes. Supranal concavity not expressed.Terminal cleft large, with rounded anterior end, length of cleft68, width at level of setae ps1 95. Interlobar membrane occu-pies anterior part of terminal cleft, with angular incision, dor-sal surface with a few transverse striae near free margin; lengthof incision 49. Setae c3 setiform, about 75 long. Lateral setae c2 spiculiform, 66 long, d2, e2 broken in lectotype (in addi-tional material, 4 males from Pl. elegans, length of lateral setae:c2 58�68, d2 65�72, e2 120�132). Setae f2 narrowly foliform*.Setae ps1 ovate (Fig. 1A, B). Dorsal measurements (distancebetween transverse setal rows or bases of setae): c2:d2 122,d2:e2 117, e2:h3 80, e2:e1 25, e2:e2 142, e1:e1 91, ps1:ps1 105.Cupules im and openings gl at same transverse level.

Epimerites I free, posterior tips close to each other. Basesof epimerites II without inflations. Genital apparatus at levelof anterior margin of trochanters IV, 25 × 22. Paragenitalapodemes: anterior tips extended to level of setae 3a, middle

parts with inner sclerotized areas but not fused into entiretransverse bridge. Ventral measurements: 3b:3a 11, 3a:g 34,g:4a 50, 4a:ps3 75. Cupules ih at level of setae ps2. Diameterof anal suckers 18.

Legs I, II subequal in size. Genua and tibiae I, II with dis-tinct ventral spine-like apophyses (Fig. 2A, B). Solenidion σ2on genu I present. Tarsus IV: setae d, e barrel-shaped, situat-ed at midlevel of segment (Fig. 2E).

Female (measurements for 10 paralectotypes): Idiosoma,length × width, 470�505 × 265�290. Prodorsal shield: almostrectangular plate, not extended to scapular setae si, se; lengthalong midline 88�97, greatest width 62�66. Setae si spiculi-form, 84�91 long. Distance between scapular setae: se:se72�79, si:si 29�35. Scapular shields developed only on later-al margins of propodosoma. Hysterosoma 322�341 long.Hysteronotal shield: length 280�292, width 200�215, anteri-or margin straight, surface with network pattern. Pygidialshield unpaired, clearly separated from hysteronotal shield.Setae c3 36�44 long. Setae d2, e2 thin spiculiform, f2 seti-form; length of lateral hysteronotal setae: c2 67�77, d2 30�32,e2 23�31, f2 18�24. Dorsal measurements: c2:d2 124�134,d2:e2 139�150, e2:h3 46�50, e2:e2 113�122, h3:h3 42�50.Cupules im anterior to level of openings gl. Bases of epimer-ites II with heavily sclerotized inflations. Epigynum bow-shaped 20�30 × 50�63 (Fig. 3A). Cupules ih postero-mesal tosetae ps2. Solenidion σ2 on genu I present. Ventral apophy-ses of tibia I, II weakly expressed, apophyses on genua I, IIindistinct.

Remarks: Protolichus affinis, the type species of the genusApexolichus, was originally described by Mégnin and Trou-essart (1884) from two parrot species, Platycercus elegans (inoriginal description Pl. pennanti) and Pl. flaveolus, both fromAustralia. In subsequent studies, Trouessart re-examined thematerial from these hosts and identified the specimens he hadpreviously referred to Protolichus affinis as different species,resulting in confusion about host associations and even absurdtaxonomical situation. Thus, a few years later, Trouessart(1899) described a new species, Protolichus velifer, and syn-onymized the older species, P. affinis, to this new name. Thiswas an illogical and unlawful action from the point of view oftaxonomic rules. Trouessart listed three species as the hostsof P. velifer: Eunymphicus cornutus from New Caledonia,Pl. flaveolus from Australia, and Prosopeia personata fromFiji. From this information it is possible to understand thatTrouessart referred to P. velifer the specimens of P. affinisfrom Pl. flaveolus. It is necessary to stress here that Pl. ele-gans was not mentioned as the host of P. velifer.

Further, in a subsequent revision of the genus Protolichus,the situation with P. affinis became even more confused. Inthis revision, Favette and Trouessart (1904) established onemore new species, Protolichus chiragricus, and declared P. af-finis also as the synonym of that species. These authors listedfor P. chiragricus three hosts from Australia: Pezoporus wal-

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*In microslides these setae are often visible edge-on, therefore they may look like setiform ones with membranous basal extension.


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licus, Platycercus elegans and Pl. flaveolus. This probablymeans that all specimens of P. affinis from Pl. elegans and atleast some specimens from Pl. flaveolus were referred toP. chiragricus. Thus, P. affinis was shut down as a validspecies and declared as a synonym of two different juniorspecies.

Gaud and Atyeo (1996) did not comment this confused sit-uation but restored Protolichus affinis as a valid species, andhad chosen it as a type species for the genus Apexolichus.Specimens from Platycercus elegans (TRT 34G2) were hand-marked by Atyeo as a syntype series. A male lectotype ofP. affinis is designated in the present study.

Protolichus distensis was described by Favette and Trou-essart (1904) from Platycercus eximius from South Australiaand Tasmania. This material is absent from the collection ofTrouessart and is apparently lost. Gaud and Atyeo (1996)treated it as a valid species under the name Apexolichus dis-tensis. Our specimens collected from the type host, Pl. exim-ius from Australia, are conspecific to Apexolichus affinis. Inaddition, the general image of a male (a photo from micro-scope) of P. distensis given by Favette and Trouessart (1904:fig. 18) also allows us to refer that specimen to Apexolichusaffinis, based on such clearly recognized features as follows:the wide terminal cleft with rounded anterior end, the deep

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Fig. 4. Apexolichus velifer (Trouessart, 1899), male: A � ventral view; B � dorsal view


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incision in interlobar membrane, the absence of sclerotizedbridge between paragenital apodemes, and the absence of thebasal inflations on epimerites II. As it is said in the previousremark, Protolichus affinis was �shut down� by its author(Trouessart 1899); therefore it is not surprising that mites fromPlatycercus eximius were described in the subsequent revisionof the genus Protolichus as a new species.

Apexolichus velifer (Trouessart, 1899) (Figs 4�6; 7A, B)

Pterolichus (Protolichus) velifer Trouessart, 1899: 5 (in part);Canestrini & Kramer, 1899: 57 (in part).Protolichus velifer: Favette & Trouessart, 1904: 144, figs 1,13, 15.Apexolichus velifer Gaud & Atyeo, 1996: 117.Protolichus splendens Favette & Trouessart, 1904: 17, fig. 17;Gaud & Atyeo, 1996: 117, syn. nov.

Material examined: Lectotype male and paralectotype female(TRT 34G13) ex Prosopeia personata (Gray G.R, 1848), IslesFiji, no other data; 4 males and 2 females (UMB 1538) exP. tabuensis (Gmelin, 1788), Isles Fiji, 1871, coll. unknown.

Diagnosis: Solenidion σ2 of genua I present. Male: basalsegment of palps with lateral expansions directed anteriorlyand bearing setae dp2; hysteronotal shield with network pat-tern in anterior part; anterior margin of terminal cleft blunt-angular; interlobar membrane with deep right-angular inci-sion, dorsal surface of this membrane with a few striae nearfree margin of membrane; setae d2 about half the length ofsetae c2; paragenital apodemes posterior to genital apparatusconnected to each other by wide and heavily sclerotized trans-verse bridge; bases of epimerites II with inflations; tibiae andgenua I, II with distinct spine-shaped ventral apophyses, tibiaI with short and blunt dorsobasal apophysis; setae d, e of tarsiIV situated approximately at midlevel of segment. Epigynum

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Fig. 5. Apexolichus velifer (Trouessart, 1899), details of male: A � genito-anal region; B � dorsal opithosoma; C � gnathosoma, ventral; D �variability of shape of setae ps1, individuals from Prosopeia personata (1, 2) and from P. tabuensis (3�6)


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absent, setae d2, e2 short spiculiform, cupules im anterior toopenings gl, length of idiosoma 540�600, length of setae: si100�105, c2 70�75.

Description: Male (lectotype). Idiosoma, length × width,682 × 365. Gnathosoma: basal segment of palps with lateralexpansions directed anteriorly and bearing setae dp2; ventralside of subcapitulum with pair of flat spines at base of palpsand pair of narrow crests along lateral margins (Fig. 5C).Prodorsal shield: split into two pieces, anterior piece shaped aslongitudinal plate enlarged in anterior part; posterior piece inform of transverse plate bearing setae se, si on anterior mar-gin; total length of prodorsal shield 154, greatest width ofanterior piece 62. Setae si spiculiform, 110 long. Distancebetween scapular setae: se:se 90, si:si 42. Scapular shields asnarrow poorly sclerotized bands. Hysterosoma 465 long.Hysteronotal shield: greatest length 465, greatest width 236,anterior margin straight, anterior part with faint network pat-tern. Opisthosomal lobes almost straight, with rounded ends,with narrow terminal membrane on posterior margin of lobesand with small triangular lateral membrane at level of setaeh2. Supranal concavity open posteriorly to terminal cleft.Terminal cleft with blunt-angular anterior end and with slight-ly divergent lateral margins, length of cleft (excluding con-cavity) 85, width at level of setae ps1 120. Interlobar mem-brane occupies most of terminal cleft, with blunt angular inci-sion, dorsal surface with a few transverse striae in median

part; length of incision 37. Setae c3 setiform, about 70 long.Lateral setae c2, d2, e2 thick spiculiform, 90, 48, and 152�155in length, respectively. Setae f2 narrowly lanceolate. Setae ps1of roughly triangular form (in some specimens inner and pos-terior angle may be rounded (Fig. 5B, D). Dorsal measure-ments: c2:d2 150, d2:e2 196, e2:h3 110, e2:e1 26, e2:e2 117,e1:e1 51, ps1:ps1111. Cupules im slightly anterior to level ofopenings gl.

Bases of epimerites II with heavily sclerotized inflations.Genital apparatus at level of trochanters IV, 34 × 16. Paragen-ital apodemes: anterior tips extended to level of humeral setaecp, middle parts between genital apparatus and anal fieldfused to each other by large and heavily sclerotized transversebridge (Fig. 5A). Ventral measurements: 3b:3a 11, 3a:g 54,g:4a 94, 4a:ps3 144. Cupules ih anterior to level of setae h2.Diameter of anal suckers 22. Legs I, II subequal in size. Tibiaeand genua I, II with distinct spine-like ventral apophyses; tibiaI with blunt dorsobasal apophysis. Solenidion σ2 on genu Ipresent. Setae d, e of tarsus IV approximately at midlevel ofsegment.

Female (paralectotype): Idiosoma, length × width, 570 ×365. Prodorsal shield: almost rectangular plate, not extendedto level of scapular setae si, se; length along midline 118,greatest width 88. Setae si missing in paralectotype (spiculi-form, 100 long in 2 specimens from additional material). Dis-tance between scapular setae: se:se 105, si:si 51. Scapular

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Fig. 6. Apexolichus velifer (Trouessart, 1899), legs of male, dorsal view: A � leg I; B � leg II; C � leg IV


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Fig. 7. Females of Apexolichus velifer (Trouessart, 1899) (left) and A. lathami Mironov, Dabert et Proctor, 2003 (right): A and C � dorsal view;B and D � ventral view of oviporal region. Figures C and D after Mironov et al. (2003b), modified


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shields narrow, poorly sclerotized. Hysterosoma 378 long.Hysteronotal shield: length 298, width 250, anterior marginstraight, anterior part transversally striated, median part withnetwork pattern. Pygidial shields unpaired, clearly separatedfrom hysteronotal shield. Setae c3 56 long. Setae d2, e2, f2thin spiculiform; length of lateral setae: c2 75, d2 35, e2 36,f2 14. Dorsal measurements: c2:d2 133, d2:e2 155, e2:h3 58,e2:e2 159, h3:h3 51. Cupules im anterior to level of openingsgl. Bases of epimerites II with heavily sclerotized inflations.Epigynum absent. Cupules ih postero-mesal to setae ps2.Solenidion σ2 on genu I present. Ventral apophyses of tibiaeand genua I, II poorly developed.

Remarks: In the original description of Protolichus veli-fer and in the revision of the genus Protolichus (Touessart1899, Favette and Trouessart 1904), three host species werelisted for this mite: Eunymphicus cornutus from New Cal-

edonia, Platycercus flaveolus from Australia, and Prosopeiapersonata from Fiji. Mentioning of Pl. flaveolus as the hostwas apparently the result of re-identification of the samplesfrom that host (TRT 34G12), which were originally identifiedby Trouessart (1884) as Protolichus affinis Trouessart, 1884(see remark for Apexolichus affinis). As for the material fromE. cornutus referred by Trouessart (1899) to P. velifer, it eitherwas not retained in the collection or this author actually meantthe samples (TRT 34G11) that were subsequently described asthe subspecies Protolichus velifer neglectus Favette et Trou-essart, 1904. Gaud and Atyeo (1996) declared Prosopeia per-sonata as the type host and marked the only retained male andfemale as lectotype and paralectotype of A. velifer.

Protolichus splendens was described by Favette and Trou-essart (1904) from Prosopeia tabuensis from the Fijian Is-lands based on a single male. Gaud and Atyeo (1996) treated

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Fig. 8. Apexolichus lathami Mironov, Dabert et Proctor, 2003, male: A � ventral view; B � dorsal view. After Mironov et al. (2003b), modi-fied


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this mite as a valid species of Apexolichus. Type material ofthis species was not retained in the collection of Trouessart.Surprisingly, in the differential diagnosis of P. splendens,Favette and Trouessart compared this species to P. distensis,rather to P. velifer, although the similarity of paragenital apo-demes in males of P. velifer and P. splendens is quite remark-able and obvious (see Favette and Trouessart 1904: figs 13,17). Comparison of type material of P. velifer from Prosopeiapersonata and mites collected from Pr. tabuensis shows nodifference between samples from these two Prosopeia spe-cies; therefore we consider Apexolichus splendens as a juniorsynonym of A. velifer.

Apexolichus lathami Mironov, Dabert et Proctor, 2003(Figs 7C, D; 8�10)

Apexolichus lathami Mironov, Dabert et Proctor, 2003:187,figs 1�3, 7, 8.

Material examined: Male holotype (QM S48358), 1 male and4 female paratypes (QM S38359) ex Lathamus discolor(Shaw, 1790), Australia, Queensland, Spreyton, 17 October1998, B. Gartell; 3 males, 2 females (AMU OL-59), samehost, Australia, no other data; 1 male, 2 females (AMU-OL-66), Australia, no other data.

Diagnosis: Solenidion σ2 of genua I present. Male: basalsegment of palps with lateral crest extending backward; hys-teronotal shield monotonously punctured; anterior margin ofterminal cleft rounded; interlobar membrane with short andwide incision, dorsal surface of this membrane with a few

transverse striae in anterior part; setae d2 equal to or slightlylonger than setae c2; paragenital apodemes posterior to geni-tal apparatus connected to each other by wide sclerotizedtransverse bridge; bases of epimerites II without inflations;tibiae I with short and acute dorsobasal apophysis and spine-like ventral apophysis, tibia II with spine-like ventral apophy-ses, genua I, II with poorly expressed ventral apophyses; setaed, e of tarsi IV approximately at midlevel of segment. Female:epigynum absent, setae d2, e2 thin spiculiform, cupules im atlevel of openings gl, length of idiosoma 400�455, length ofsetae: si 62�70, c2 60�70.

Remark: This species is known only from the type host,Lathamus discolor, in Australia (Mironov et al. 2003b).

Apexolichus neglectus (Favette et Trouessart, 1904) (Figs11�13A, B)

Protolichus velifer neglectus Favette et Trouessart, 1904: 145,fig. 14.Apexolichus neglectus: Gaud & Atyeo, 1996: 117.

Material examined: Male lectotype, 3 male and 5 female para-lectotypes (TRT 34G11) ex Eunymphicus cornutus (Gmelin,1788), New Caledonia, no other data.

Diagnosis: Solenidion σ2 of genua I absent. Male: basalsegment of palps without lateral extension; hysteronotalshield monotonously punctured; anterior margin of terminalcleft rounded; interlobar membrane with shallow incision,dorsal surface of this membrane smooth; setae d2 equal orslightly shorter than setae c2; paragenital apodemes posterior

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Fig. 9. Apexolichus lathami Mironov, Dabert et Proctor, 2003, details of male: A � dorsal opisthosoma; B � genito-anal region


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to genital apparatus connected to each other by wide sclero-tized transverse bridge; bases of epimerites II with heavilysclerotized inflations; tibiae and genua I, II with spine-shapedventral apophyses; setae d, e of tarsi IV closer to base of seg-ment. Female: epigynum absent, setae d2, e2 thin spiculiform,cupules im slightly anterior to openings gl, length of idioso-ma 475�515, length of setae: si 98�106, c2 90�94.

Description: Male (lectotype, measurements for 3 para-lectotypes in brackets). Idiosoma, length × width, 550 × 295(533�558 × 297�310). Gnathosoma: basal segment of palpswithout lateral expansions. Prodorsal shield: split into twopieces; anterior piece shaped as longitudinal plate enlarged inanterior part; posterior piece shaped as transverse and slight-ly curved plate bearing setae se, si on anterior margin; totallength of shield 152 (150�180), greatest width of anteriorpiece 82 (77�85). Setae si spiculiform, 95 (90�98) long.Distance between scapular setae: se:se 73 (71�76), si:si 34(31�35). Scapular shields as narrow poorly sclerotized bands.Hysterosoma 390 (388�396) long. Hysteronotal shield: great-est length 384 (380�390), greatest width 186 (185�202), ante-rior margin straight, surface uniformly punctured. Opistho-somal lobes slightly curved medially, with rounded ends, shortterminal membrane on posterior margin of lobes and smallangular lateral membrane at level of setae h2. Supranal con-cavity open posteriorly to terminal cleft. Terminal cleft muchshorter than wide, with concave anterior margin, length of

cleft 62 (60�70), greatest width 100 (88�102). Interlobarmembrane occupies entire terminal cleft, with shallow inci-sion on posterior margin; length of incision 12 (12�14). Setaec3 setiform, about 55 long. Lateral setae c2, d2, e2 thick spiculiform, 91 (90�100), 62 (60�66), and 110 (100�110) inlength, respectively. Setae f2 setiform. Setae ps1 roughly cres-cent-shaped (Fig. 12A). Dorsal measurements: c2:d2 143(129�140), d2:e2 154 (150�167), e2:h3 77 (74�80), e2:e1 22(15�24), e2:e2 123 (120�130), e1:e1 60 (58�62), ps1:ps1 96(94�103). Cupules im slightly anterior to level of openings gl.Epimerites I free, their posterior parts closely adjacent. Basesof epimerites II with heavily sclerotized inflation. Genitalapparatus at level of anterior margin of trochanters IV, 29 × 14(29�35 ×14�17). Paragenital apodemes: anterior tips extend-ed to tips of epimerites IIIa, middle parts of apodemes con-nected to each other by large sclerotized bridge. Ventral meas-urements: 3b:3a 7 (7�13), 3a:g 34 (32�36), g:4a 65 (62�69),4a:ps3 95 (90�98). Cupules ih posterior to level of setae ps2.Diameter of anal suckers 26 (25�27). Legs I, II similar in size.Tibiae and genua I, II with spine-like ventral apophyses (Fig.12B, C). Solenidion σ2 on genu I absent. Tarsus IV: setae d,e situated near base of segment (Fig. 12D).

Female (5 paralectotypes): Idiosoma, length × width, 477�515 × 310�328. Prodorsal shield: longitudinal rectangular plateslightly enlarged in anterior part, not extended to scapular setaesi, se; length along midline 100�104, greatest width 77�83; sur-

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Fig. 10. Apexolichus lathami Mironov, Dabert et Proctor, 2003, legs of male, dorsal view: A � leg I; B � leg II; C � tibia and tarsus III; D �tibia and tarsus IV


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face posterior to bases of scapular setae may be slightly punc-tured. Setae si spiculiform, 98�106 long. Distance betweenscapular setae: se:se 83�87, si:si 36�39. Scapular shields nar-row, poorly sclerotized. Hysterosoma 328�340 long. Hystero-notal shield: length 252�280, width 214�220, anterior marginstraight, entire surface with network pattern (Fig. 13A). Pygid-ial shields unpaired, clearly separated from hysteronotalshield. Setae c3 50�54 long. Setae d2, e2, f2 thin spiculiform;length of lateral setae: c2 90�94, d2 42�44, e2 40�42, f2 15�17.Dorsal measurements: c2:d2 130�132, d2:e2 143�148, e2:h346�58, e2:e2 128�139, h3:h3 48�50. Cupules im slightly ante-rior to level of openings gl. Bases of epimerites II with heavilysclerotized inflations. Epigynum absent (Fig. 13B). Cupules ih

postero-mesal to setae ps2. Solenidion σ2 on genu I absent.Ventral apophyses of tibiae and genua I, II distinct, short.

Remark: This species is known only from Eunymphicuscornutus in New Caledonia (Favette and Trouessart 1904); orig-inally it was described as a subspecies of Protolichus velifer.

Apexolichus platycerci (Mironov, Dabert et Proctor, 2003)comb. nov. (Figs 13C, D; 14�17)

Titanolichus platycerci Mironov, Dabert et Proctor, 2003: 189,figs 4�6, 9, 10.

Material examined: Male holotype (QM S48360), 3 males and7 female paratypes (QM S48361) ex Platycercus adscitus

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Fig. 11. Apexolichus neglectus (Favette et Trouessart, 1904), male: A � ventral view; B � dorsal view


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(Latham, 1790), Australia, Queensland, St. Lucia, 16.04.1999,H. Proctor; 4 males, 7 females, same host (AMU BR-100),Australia, no other data.

Diagnosis: Solenidion σ2 of genua I absent. Male: basalsegment of palps with lateral crest extending backward; hys-teronotal shield with network pattern in anterior part; anteri-or margin of terminal cleft blunt-angular in shape; interlobarmembrane with short and wide incision, dorsal surface of thismembrane smooth; setae d2 2�2.5 times longer than setae c2;paragenital apodemes posterior to genital apparatus connect-ed to each other by wide transverse bridge; bases of epimeritesII without inflations; legs II much longer than legs I; tarsi IIwith large dorsobasal apophysis bearing solenidion ω1 andseta ba on rounded apex; tibiae, genu and femur I with spine-like ventral apophyses, tibiae and genua II without ventralapophyses; setae d, e of tarsi IV inserted approximately atmidlevel of segment. Female: epigynum present; setae d2, e2

thin spiculiform, cupules im anterior to level of openings gl,length of idiosoma 470�490, length of setae: si 65�70, c250�55.

Remark: This species is known only from Platycercusadscitus in Australia. Originally it was described in the con-text of the genus Titanolichus based on modification of tarsiII, which resembled that in heteromorphic males of the lattergenus (Mironov et al. 2003b). The present comparative studyshowed that this similarity to Titanolichus is superficial, andthat this species should be moved to the genus Apexolichusbased on structure of modified setae of tarsi IV in males andepimerites II in females.

Apexolichus psephoti sp. nov. (Figs 18; 19)

Type material: Male holotype (UMB 1/1019), 5 male and 8female paratypes ex Psephotus varius Clark A.H., 1910, S.

Fig. 12. Apexolichus neglectus (Favette et Trouessart, 1904), details of male: A � dorsal opisthosoma; B � leg I, dorsal view; C � leg II, dor-sal view; D � leg IV, dorsal view

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Australia, Gawler Ranges, 08.1955, H.O.Wagner. Holotype, 4male and 7 female paratypes � AMU; 1 male and 1 femaleparatypes � UMB.

Additional material: 12 males and 8 females (UMB 23347)ex Psephotus haematonotus (Gould, 1838), S. Australia, Gaw-ler Ranges, 08.1955, H.O. Wagner.

Diagnosis: Solenidion σ2 of genua I present, minute. Male: basal segment of palps with lateral crest extending back-ward; hysteronotal shield with network pattern; anterior mar-gin of terminal cleft roughly concave; interlobar membranewith shallowly concave posterior margin, dorsal surface with

transverse striae near margin; setae d2 about 1.5 times longerthan setae c2; paragenital apodemes posterior to genital appa-ratus connected to each other by wide sclerotized transversebridge; bases of epimerites II without inflations; legs I and IIsubequal; tibiae and genua I, II with spine-like ventral apoph-yses; setae d, e of tarsi IV closer to base of segment. Fe-male: epigynum present, setae d2, e2 thin spiculiform, cupulesim anterior to level of openings gl, length of idiosoma440�465, length of setae: si 64�70, c2 50�65.

Description: Male (holotype, measurements for 5 para-types in brackets). Idiosoma, length × width, 500 × 245

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Fig. 13. Females of Apexolichus neglectus (Favette et Trouessart, 1904) (left) and A. platycerci (Mironov, Dabert et Proctor, 2003) (right):A and C � dorsal view; B and D � ventral view of oviporal region. Figures C and D after Mironov et al. (2003b), modified


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(470�505 × 220�265). Gnathosoma: basal segment of palpswith lateral crest extending backward; subcapitulum withoutventral spines (Fig. 18A). Prodorsal shield: longitudinal platenot extended to bases of scapular setae se, si, posterior end ofthis plate narrowed, not wider than distance between setae si;length 85 (80�90), greatest width 45 (45�50). Setae si spic-uliform, 70 (68�75) long. Distance between scapular setae:se:se 50 (48�50), si:si 22 (18�22). Scapular shields poorlyexpressed. Hysterosoma 350 (340�365) long. Hysteronotalshield: greatest length 335 (320�345), greatest width 178(165�178), anterior margin slightly convex, central part withfaint network pattern (Fig. 18B). Opisthosomal lobes slight-ly divergent, with rounded ends, with narrow terminal mem-brane on posterior margin of lobes forming short angularextension at level of setae h2. Supranal concavity open pos-teriorly to terminal cleft. Terminal cleft wider than long, withslightly convex and rough anterior margin, length of cleftexcluding supranal concavity 42 (42�45), greatest width(anterior to setae ps1) 94 (72�85). Interlobar membrane

occupies most part of terminal cleft, with shallowly concaveposterior margin, dorsal surface with a few transverse striaenear margin; length of incision 8 (5�10). Setae c3 setiform,about 58 (48�60) long. Lateral setae c2, d2, e2 thick spiculi-form, 64 (62�68), 110 (105�125), and 130 (122�138) long,respectively. Setae f2 setiform. Setae ps1 foliform, ovate (Fig.18B). Dorsal measurements: c2:d2 140 (135�142), d2:e2 115(110�125), e2:h3 70 (65�70), e2:e1 26 (24�26), e2:e2 90(85�91), e1:e1 55 (53�55), ps1:ps1 85 (82�85). Cupules imand openings gl at same transverse level. Bases of epimeritesII without sclerotized inflation. Genital apparatus at level of trochanters IV, 30 × 18 (28�30 × 15�18). Paragenital apo-demes: anterior tips extended to level of humeral setae cp,middle parts connected to each other by large transverse scler-otized bridge. Ventral measurements: 3b:3a 15 (10�15), 3a:g40 (38�40), g:4a 50 (50�52), 4a:ps3 88 (84�88). Cupules ih atlevel of setae h2. Diameter of anal suckers 18 (17�18). Legs IIslightly longer that legs I. Tibiae I, II and genu I with long andacute spine-like ventral apophyses; genu II with blunt spine-

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Fig. 14. Apexolichus platycerci (Mironov, Dabert et Proctor, 2003), male: A � ventral view; B � dorsal view. After Mironov et al. (2003b),modified


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like apophysis (Fig. 19A, B). Solenidion σ2 on genu I present,minute. Setae d, e of tarsus IV closer to base of segment (Fig.19C).

Female (8 paratypes): Idiosoma, length × width, 440�465× 235�270. Prodorsal shield: almost rectangular plate, notextended to scapular setae si, se; length along midline 75�80,greatest width 50�55. Setae si spiculiform, 64�70 long. Dis-tance between scapular setae: se:se 65�67, si:si 22�28. Scap-ular shields narrow, poorly sclerotized. Hysterosoma 295�

325 long. Hysteronotal shield: length 260�270, width 180�190, anterior margin slightly concave, surface with networkpattern (Fig. 19D). Paired pygidial shields poorly outlined,covered with transverse striae. Setae c3 35�42 long, setae c2thick spiculiform, d2, e2 thin spiculiform, f2 setiform; lengthof lateral setae: c2 50�65, d2 20�23, e2 21�25, f2 10�18.Dorsal measurements: c2:d2 115�120, d2:e2 124�132, e2:h348�52, e2:e2 100�110, h3:h3 45�50. Cupules im anterior tolevel of openings gl. Bases of epimerites II with heavily scler-

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Fig. 15. Apexolichus platycerci (Mironov, Dabert et Proctor, 2003), details of male: A � genito-anal region; B � dorsal opisthosoma

Fig. 16. Apexolichus platycerci (Mironov, Dabert et Proctor, 2003), details of male: A � leg II, dorsal view; B � tibia and tarsus II, ventral view;C � gnathosoma, ventral view


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otized inflations. Epigynum present, bow-shaped, 15�18 ×45�50 (Fig. 19E). Cupules ih postero-mesal to setae ps2. Sole-nidion σ2 on genu I present, minute (recognizable under highmagnification).

Etymology: The specific epithet is derived from the gener-ic name of the host.

Titanolichus Gaud et Atyeo, 1996

Type species: Pterolichus (Protolichus) chiragricus Mégninet Trouessart, 1884

Description: Both sexes. Prodorsal shield occupying medianpart of prodorsum and extending to or beyond scapular setae;anterior part strongly sclerotized; posterior part poorly scler-otized or not expressed (Figs 20B; 24B). Scapular shields rep-resented by transverse bands. Hysterosoma with extensivehysteronotal shield. Humeral shields well-developed. Twovertical setae vi present, their bases close to each other. Scap-ular setae moved to midline, distance between setae se ap-proximately equal to width of gnathosoma. Setae si and c2

spiculiform; setae c3 setiform; setae h1 absent. Cupules iapostero-lateral to bases of setae c2, situated on humeral shield;cupules im positioned at level of hysteronotal gland openingsgl or posterior. Each tarsus with two minute apicodorsalspines, one of them may be situated on basal part of ambu-lacral stalk; tarsi I with blunt angular ventral margin. Tarsi I, II1.5�2 times longer than corresponding tibiae; solenidion ω1closer to base of tarsi I, II than to apex; setae ba slightly dis-tal to corresponding solenidion ω1; condylophores heavilydeveloped; ambulacral discs minutely dentate on distal mar-gin. Solenidion σ2 of genu I present or absent.

Male: Epimerites I free or fused as a Y (in heteromorphmales of some species). Bases of epimerites II without ball-shaped inflation (Figs 20A; 22A). Gnathosoma trapezoidal inshape, lateral margins blunt-angular without lateral spines;basal podomeres of palps without lateral expansions. Opis-thosoma noticeably narrowed to level of setae e2. Opistho-somal lobes well-developed, widely separated from eachother by semi-ovate or ovate terminal cleft, and distinctlydivergent; posterior end of lobe enlarged, posterior marginblunt or rounded; in heteromorph males, lobes strongly en-

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Fig. 17. Apexolichus platycerci (Mironov, Dabert et Proctor, 2003), legs of male: A � leg I, dorsal view; B � leg I, ventral view; C � tibiaand tarsus III, dorsal view; D � tibia and tarsus IV, dorsal view


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larged apically and whole opisthosoma may have a shape ofwhale tail. Anterior margin of terminal cleft concave or blunt-angular; interlobar membrane present, occupies most part ofterminal cleft; terminal membrane clearly separated frominterlobar membrane, lateral membranes absent. Setae d2, e2long, spiculiform; setae f2 setiform or with narrow membra-nous extension, commonly curved; setae ps2 setiform, equalor longer than setae f2; setae ps1 wide foliform, widenedhook-like or setiform; setae e1 setiform, situated posterior tolevel of setae e2. Paragenital apodemes present, long, spread-ing from level of coxal fields III to opisthosomal lobes wherethey fuse with opisthoventral shields; middle parts of apo-demes between genital apparatus and anal field close to eachother, free or connected to each other by wide transversebridge (Figs 20A; 22A; 24A, D). Anal suckers circular, corol-la dentate, each disc surrounded by membrane with radial stri-ation. Genital papillae situated lateral to genital apparatus onsoft cuticle. Coxal fields I�IV open, without sclerotized areas.Bases of trochanters I, II flanked by narrow sclerotized bands

connecting bases of respective epimerites. Cupules ih present.Legs II slightly (homeomorph males) or much longer thanlegs I (heteromorph males); legs III and IV similar in size, not hypertrophied. Tibiae and genua I, II with spine-shapedapicoventral apophyses; tarsus II in heteromorph males withstrong dorsobasal apophysis, directed backward (Fig. 21A-C).Setae ra, la of tarsus I proximal or at the same level as seta wa.Tarsus IV 1.5�1.7 times longer than tibia IV, similar in size totarsus III; setae d, e of tarsus IV as short spines (Fig. 25D).Polymorphism of males continuous, expressed in structure oftarsi and tibiae II and opisthosomal lobes.

Female: Epimerites I free. Bases of epimerites II withoutstrong inflations. Gnathosoma trapezoidal as in the male. Hys-teronotal shield split into pygidial fragment and main body;pygydial fragment paired (Figs 23; 26B). Lateral sclerotizedbands absent. Oviporus at level of epimerites III. Epigynumpresent, semicircular or bow-shaped. Setae d2, e2 spiculiformor setiform, setae f2, ps1, ps2 setiform. Copulatory openingterminal, situated on small cone-like extension. Apicoventral

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Fig. 18. Apexolichus psephoti sp. nov., male: A � ventral view; B � dorsal view


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apophyses of tibiae and genua I, II developed much weakerthan in males, commonly as blunt extension, or may be prac-tically indistinct.

Hosts: parrots of the genera Pezoporus, Neophema andBarnardius (Psittacinae, Platycercini).

Key to species (males and females)

1. In males, opisthosoma with pair of large lateral angularextensions at level of setae e2. In heteromorph males, tib-iae II with dorsal longitudinal crest, epimerites I free (Figs

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Fig. 19. Apexolichus psephoti sp. nov.: A � leg I of male, ventral view; B � leg II of male, ventral view; C � tarsus IV of male, dorsal view;D � female, dorsal view; E � oviporal region of female


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Fig. 20. Titanolichus chiragricus (Trouessart et Mégnin, 1884), male: A � ventral view of heteromorph male; B � dorsal view of hetero-morph male; C � genito-anal region of homeomorph male; D � dorsal propodosoma of homeomorph male. After Dabert et al. (2006), mod-ified


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24A; 25B). In females, prodorsal shield does not extendto bases of scapular setae (Fig. 26B) .............. Titanolichustriangulifer sp. nov.

− In males, opisthosoma without lateral angular extensionsat level of setae e2. In heteromorph males, tibiae II with-out dorsal crest, epimerites I commonly fused into Y (Fig.20A). In females, prodorsal shield extending beyond basesof scapular setae .............................................................. 2

2. In both sexes, solenidion σ2 on genua I present, minute. Inheteromorph males (with strong dorsobasal extension ontarsi II and with opisthosoma shaped as whale tail) dis-tance between scapular setae and hysteronotal shield(100�140) noticeably exceeds length of setae si (Fig.20A). In homeomorph males, distance between scapularsetae and hysteronotal shield 80�90, incision in interlobarmembrane roughly triangular in shape, 45�50 in length(Fig. 20C). In females, setae f2 about 10 long, cupules imand openings gl approximately at same transverse level(Fig. 23A) ................................... Titanolichus chiragricus(Mégnin et Trouessart, 1884)

− In both sexes, solenidion σ2 on genua I absent. In hetero-morph males, distance between scapular setae (65�85)equal to or less than length of setae si (Fig. 22B). In homeo-morph males, distance between scapular setae and hys-

teronotal shield 60�75, incision in interlobal membraneshallow, 30�35 in length (Fig. 22B). In females, setae f2about 15�25 long, cupules im commonly posterior to levelof openings gl (Fig. 23B) ................. Titanolichus seemaniDabert et al., 2006

Titanolichus chiragricus (Trouessart et Mégnin, 1884)(Figs 20; 21; 23A)Pterolichus (Protolichus) chiragricus Trouessart et Mégnin,1884: 261, Canestrini & Kramer, 1899: 56.Protolichus chiragricus: Favette & Trouessart, 1904: 142, figs9, 10 (in part). Titanolichus chiragricus: Gaud & Atyeo, 1996: 138, fig. 420.

Material examined: Homeomorph male lectotype (TRT 34H6)ex Pezoporus wallicus (Kerr, 1792), New Zealand (error, thehost is restricted to Australia), no other data. Heteromorphmale, 4 homeomorph males, 2 females (QM S64150-S64156)ex P. wallicus, Australia, Tasmania, S.E. Cape Track, 5 July2003, O. Seeman.

Diagnosis: Solenidion σ2 of genua I present. Male: later-al margins of opisthosoma without extensions; terminal cleftexcluding supranal concavity wider than long; interlobar mem-brane with short incision; setae f2 blade-shaped with very nar-

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Fig. 21. Titanolichus chiragricus (Trouessart et Mégnin, 1884), legs of male, dorsal view: A � leg I of heteromorph male; B � leg II of het-eromorph male; C � leg II of homeomorph male; D � tibia and tarsus IV of heteromorph male. After Dabert et al. (2006), modified


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Fig. 22. Titanolichus seemani Dabert, Mironov et Proctor, 2006, male: A � ventral view of heteromorph male; B � dorsal view of heteromorphmale; C � leg II of heteromorph male, dorsal view. After Dabert et al. (2006), modified


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row and acute distal half; middle parts of paragenital apo-demes close to each other but not connected by sclerotizedbridge. Heteromorph males: epimerites I fused as a Y, opistho-soma with whale tail shape, tarsus II with apically acute dor-sobasal apophyses, tibia II without dorsal crest, distancebetween scapular setae and hysteronotal shield definitelyexceeds length of setae si. Homeomorph male: incision in ter-minal cleft roughly V-shaped, 45�50 long. Female: posteriorpart of prodorsal shield encompassing bases of scapular setae;setae f2 about 10 long, cupules im at level of openings gl;length of idiosoma 505�550.

Remark: Protolichus chiragricus, the type species of thegenus Titanolichus, was originally described by Mégnin andTrouessart (1884) from Pezoporus wallicus. Further, Favetteand Trouessart (1904) reported this species from Platycercuselegans and Pl. flaveolus in Australia. As it is possible to sug-gest, Favette and Trouessart by unclear reason re-assigned the

specimens from these two species of Platycercus, which orig-inally (Mégnin and Trouessart 1884) were described as Pro-tolichus affinis, and referred them to P. chiragricus. We re-examined all retained materials from Pl. elegans and Pl. flave-olus in the Trouessart collection and recovered only represen-tatives of Apexolichus affinis (see above remarks for thisspecies). In a recent publication (Dabert et al. 2006) dealingwith new Titanolichus species, Titanolichus chiragricus wasredescribed based on the sole type specimen and additionalmaterial from the type host. This mite species is apparently spe-cific to Pezoporus wallicus.

Titanolichus seemani Dabert, Mironov et Proctor, 2006(Figs 22; 23B)

Material examined: Heteromorph male holotype, 2 homeo-morph male and 3 female paratypes (QM, S64230, S64157-

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Fig. 23. Dorsal view of females of Titanolichus chiragricus (Trouessart et Mégnin, 1884) (A) and Titanolichus seemani Dabert, Mironov etProctor, 2006 (B). After Dabert et al. (2006), modified


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Fig. 24. Titanolichus triangulifer sp. nov., male: A � ventral view of heteromorph male; B � dorsal view of heteromorph male; C � dorsalopisthosoma of homeomorph male; D � genito-anal region of homeomorph male


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64161) ex Neophema chrysogaster (Latham, 1790); 3 hetero-morph males, 1 homeomorph male, and 3 females (TRT34H5) (�Protolichus chiragricus� in original label) ex N. pul-chella (Shaw, 1792), Australia, no other data; 2 heteromorphmales, 5 homeomorph males, 6 females (UMB 22962) exN. splendida (Gould, 1841), Australia, no date, Schulze.

Diagnosis: Solenidion σ2 of genua I absent. Male: lateralmargins of opisthosoma without angle-shaped extensions; ter-minal cleft, excluding supranal concavity, wider than long;interlobar membrane with short incision; setae f2 flattened,blade-shaped with very narrow and acute distal half; middleparts of paragenital apodemes close to each other but not con-nected by wide sclerotized bridge. Heteromorph male: epi-

merites I fused into a Y (rarely remain free as in Fig. 22A),opisthosoma shaped as whale tail, tarsus II with apically acutedorsobasal apophysis, tibia II without dorsal crest, distancebetween scapular setae and hysteronotal shield 65�85, equalor less than length of setae si. Homeomorph male: incision interminal cleft with rounded anterior end, 30�55 long. Female:posterior part of prodorsal shield encompassing bases ofscapular setae; setae f2 about 15�25 long, cupules im posteri-or to level of openings gl (at same level in specimens fromN. splendida); length of idiosoma 460�520.

Remark: Titanolichus seemani was originally described byDabert et al. (2006) from Neophema chrysogaster in Tasma-nia; N. pulchella and N. splendida are new host records for

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Fig. 25. Titanolichus triangulifer sp. nov., legs of male: A � leg I of heteromorph male, ventral view; B � leg II of heteromorph male, ven-tral view; C � leg II of homeomorph male, ventral view; D � tarsus IV of heteromorph male, dorsal view


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this species. It is possible to conclude that T. seemani is prob-ably restricted to hosts of the genus Neophema, small-sizedAustralian parrots.

Titanolichus triangulifer sp. nov. (Figs 24�26)

Type material: Holotype heteromorph male, paratypes: 2 het-eromorph males, 1 homeomorph male, 5 females ex Barnard-ius zonaris (Shaw, 1805), Australia, S. Australia, GawlerRanges, August 1955, H.O. Wagner (UMB 23360). Holotype,2 male and 4 female paratypes � AMU; other paratypes �UMB.

Additional material: 2 heteromorph males, 4 females(UMB 1/1030) ex Northiella haematogaster (Gould, 1838),Australia, S. Australia, Adelaide Zoo, May 1956, dead intransport, coll. unknown.

Diagnosis: Solenidion σ2 of genua I absent. Male: lateralmargins of opisthosoma with pair of large triangle-shapedextensions; terminal cleft ovate; interlobar membrane withdeep angle-shaped incision; setae f2 setiform; middle parts ofparagenital apodemes connected to each other by wide scler-otized bridge. Heteromorph male: epimerites I free; opistho-somal lobes enlarged apically, with strongly convex lateralmargins; tarsus II with bluntly rounded dorsobasal apophyses,tibia II with dorsal crest; distance between scapular setae andhysteronotal shield (45�55) shorter than length of setae si.Homeomorph male: length of incision in interlobar membrane30�32. Female: posterior part of prodorsal shield not extend-ing to bases of scapular setae; setae f2 about 10 long, cupulesim at level of openings gl, length of idiosoma 520�550.

Description: Heteromorph male (holotype, measurementsfor 2 paratypes in brackets). Idiosoma, length × width, 510 ×

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Fig. 26. Titanolichus triangulifer sp. nov., female: A � oviporal region; B � dorsal view


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240 (505�515 × 235�255). Gnathosoma: basal segment ofpalps without lateral extension; subcapitulum without ventralspines. Prodorsal shield: longitudinal plate not extended tobases of scapular setae se, si, posterior end of this plate nar-rowed, not wider than distance between setae si; length 90(85�90), greatest width 45 (45�50). Setae si spiculiform, 72(70�77) long. Distance between scapular setae: se:se 60(50�60), si:si 27 (20�27). Scapular shields narrow, band-like,poorly sclerotized. Hysterosoma 375 (370�375) long. Hyster-onotal shield: greatest length 355 (350�355), greatest width190 (180�190), anterior margin straight, central part withsparse striated pattern. Lateral margins of opisthosoma withpair of large triangular extensions at level of setae e2 (Fig.24A, B).

Opisthosomal lobes slightly divergent posteriorly, withrounded and enlarged posterior ends and strongly convex lat-eral margins at level of setae h2. Supranal concavity small,open posteriorly to terminal cleft. Terminal cleft wider thanlong, with rounded anterior margin, and with strongly concavelateral margins, length of cleft excluding supranal concavity50 (50�55), greatest width 80 (80�85). Interlobar membraneoccupies most part of terminal cleft; incision in membranedeep angle-shaped, 38 (35�40) long. Setae c3 setiform, 50(40�55) long. Lateral setae c2, d2, e2 thick spiculiform, 78(70�78), 75 (72�75), and 95 (95�110) long, respectively.Setae f2 short setiform, 15 (12�15) long. Setae ps1 thick seti-form, curved. Dorsal measurements: c2:d2 135 (120�140),d2:e2 150 (150�156), e2:h3 64 (60�65), e2:e1 15 (15�19),e2:e2 90 (90�92), e1:e1 40 (40�42), ps1:ps1 26 (24�28).Cupules im and openings gl approximately at same transverselevel. Epimerites I free. Bases of epimerites II without scler-otized inflations. Genital apparatus at level of anterior mar-gin of trochanters IV, 30 × 18 (20�32 × 16�18). Paragenitalapodemes: anterior tips extended to level of setae 3a (in someparatypes to level of humeral setae), middle parts connected toeach other by large heavily sclerotized transverse bridge.Ventral measurements: 3b:3a 15 (15�18), 3a:g 40 (35�40),g:4a 53 (50�58), 4a:ps3 98 (85�105). Cupules ih slightly ante-rior to level of setae h2. Diameter of anal suckers 18 (17�18).Tibiae and genua I, II with large spine-like ventral apophyses;tarsus II with semi-rounded dorsobasal extension bearingsolenidion ω1 and with longitudinal dorsal crest; genu II withdorsal longitudinal crest (Fig. 25A, B). Solenidion σ2 on genuI absent. Setae d, e of tarsus IV short spine-like, situatedapproximately at midlevel of segment (Fig. 25D).

Homeomorph male (1 paratype): Similar to heteromorphmale except for the following features: tarsus II without semi-rounded dorsobasal extension, tibia II without dorsal crest,setae ps1 filiform (Fig. 24C). Measurements: idiosoma, length× width, 490 × 240; prodorsal shield, length × width, 85 × 40;setae si 65 long; distance between scapular setae se:se 48�60,si:si 22; hysterosoma 345 long; hysteronotal shield, length ×width, 320 × 160; terminal cleft wider excluding supranalconcavity 45 long, 68 wide; incision in interlobar membrane32 long; setae: c3 45, c2 50, d2 70, e2 110, f2 12; distancebetween setae: c2:d2 120, d2:e2 132, e2:h3 60, e2:e1 12, e2:e2

88, e1:e1 38, ps1:ps1 24; genital apparatus 25 × 15; distancebetween ventral setae: 3b:3a 15, 3a:g 35, g:4a 50, 4a:ps3 85;diameter of anal suckers 15�18.

Female (5 paratypes): Idiosoma, length × width, 520�550× 260�290. Prodorsal shield: longitudinal plate narrowed pos-teriorly, not extended to scapular setae si, se; length alongmidline 90�95, greatest width 50�55. Setae si spiculiform,55�70 long. Distance between scapular setae: se:se 77�85,si:si 37�40. Scapular shields shaped as narrow transversebands, poorly sclerotized. Hysterosoma 380�390 long. Hyster-onotal shield: length 295�310, width 200�210, anterior mar-gin straight, posterior part with network pattern (Fig. 26B).Pygidial shield paired, weakly outlined. Setae c3 58�62 long.Setae d2 setiform, e2 spiculiform, f2 setiform; length of later-al setae: c2 42�50, d2 25�32, e2 42�50, f2 15�20. Dorsalmeasurements: c2:d2 140�145, d2:e2 160�168, e2:h3 45�50,e2:e2 110�115, h3:h3 45�54. Cupules im and openings glapproximately at same transverse level. Bases of epimerites IIwithout sclerotized inflations. Epigynum present, semi-cir-cular, 42�48 × 85�98 (Fig. 26A). Genital papillae at levels ofsetae g. Cupules ih postero-mesal to setae ps2. Solenidion σ2on genu I absent.

Remark: Samples of Titanolichus triangulifer collectedfrom dry skin specimens of two different Australian parrots,Barnardius zonaris and Northiella haematogaster, awakedoubts that both host associations are natural, because the gen-era Barnardius and Northiella are not very closely related.Taking into consideration that the avian sample of B. zonariswas captured in natural conditions, while that of N. haemato-gaster derived from the Adelaide Zoo where contaminationwas quite probable, we suggest that the first host associationis natural and the second one is probably contamination.

Etymology: The specific epithet points out a pair of trian-gular extensions on lateral margins of opisthosoma in males.

Discussion

Phylogenetic analysis

The reconstruction of phylogenetic relationships between taxawas conducted based on the data matrix included 18 OTUs(species) and 54 morphological characters (Appendices 1, 2).The branch-and-bound procedure provided a single most par-simonious tree (Fig. 27) with the following parameters (ex-cluding uninformative characters): length 105, CI = 0.607, HI= 0.333, RI = 0.746, RC = 0.496. The consistency index wasbetter than the critical value for this number of taxa, indicat-ing a good fit of the data to the tree topology (Sanderson andDonoghue 1989). The tree topology confirms the monophy-ly of genera Apexolichus and Titanolichus. All branches aresupported by Bremer index, although mostly of low value.

The analysis revealed a close relationship of the generaApexolichus and Titanolichus, but proved that they are clear-ly distinct genera. Moreover, it showed that the monotypicgenus Sideroferus and Titanolichus are closer to each otherthan the latter genus and Apexolichus. In the discussion below

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(and in Fig. 27) we consider only unambiguous character-statechanges. Some other character states also characterize partic-ular clades but the interpretation of direction of their evolu-tionary changes is ambiguous (plesiomorphies or apomor-phies). For these reasons these characters are omitted fromboth cladogram and discussion. The clade bearing these threegenera, which we name here the Apexolichus complex, ischaracterized by two synapomorphies (independently derivedin Protolichus): presence of polymorphic males (#3) and longanterior tips of male genital apodemes (#22). The cladeTitanolichus-Sideroferus is supported by three unique synapo-morphies: distal enlargement of opisthosomal lobes in het-eromorph males (#5) and spine-like setae d and e on male tarsiIV (#53, #54). This clade is also supported by another synapo-morphy shared homoplastically with Apexolichus platycerciand A. neglectus: the absence of solenidion σ2 (#52). Accord-ing to our analysis, this solenidion reappeared secondarily inTitanolichus chiragricus. The Apexolichus clade is defined bya single but unique synapomorphy: reduction of scapularshields in females (#16).

Sideroferus lunula (Robin, 1877) is a quite derived spe-cies, characterized by numerous autoapomorphic features,although none of these character states are unique for this

species: setae si in males and females setiform, not longer thandistance se-se (#30, #31), setae d2 in males and e2 in femalessetiform (#34, #37), loss of ventral apophyses on male genuaI and tibiae I, II (#42, #43, #44), presence of dorsal apophy-ses on tarsi I in males (#47). Titanolichus, the sister group ofSideroferus, is characterized by two autoapomorphies: adja-cent posterior parts of male genital apodemes (#24), which isunique for this genus, and large dorsal apophyses on tarsi IIin males (#49).

Within the Apexolichus clade the species Apexolichusplatycerci is probably most deviating species, because itsmales have lost ventral apophyses on tibiae II (#44) and poss-es dorsobasal process on tarsi I (#49), which is similar to thosein the genus Titanolichus. Three species, A. velifer, A. latha-mi, and A. neglectus apparently comprise the most derivedspecies grouping within Apexolichus, because their femaleshave lost epigynum (#20) that is very rare case in Pterolicho-idea as the whole.

The analysis once more confirmed our former hypothesisthat the genus Apexolichus sensu Gaud and Atyeo (1996) wasan artificial paraphyletic grouping. The genera Calyptolichus,Loriprotolichus, and Nestorilichus, based on three species for-merly removed from the genus Apexolichus (Mironov and Da-

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Fig. 27. Reconstructions of phylogenetic relationships between species of Apexolichus-Protolichus complex derived from maximum parsi-mony analysis. The support for particular clades (Bremer indices) is shown in italic below branches. Only unambiguous synapomorphiesare shown: black circles � unique apomorphies, white circles � convergent apomorphies


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bert 2007), represent in the cladogram separate, well-definedlineages. Thus, in our phylogenetic tree, the genus �Apexo-lichus� sensu Gaud and Atyeo (1996) is represented by fourseparate clades of generic rank.

Host-mite associations

As our main goal in phylogenetic analysis was testing themonophyly of genera comprising the Apexolichus complex,we do not carry out formal cophylogenetic analysis of hostassociations of the whole Protolichus group; this is left for afuture study assuming more complete sampling of hosts andmites. Nevertheless, overview of host associations of all rep-resentatives of the Protolichus group, except for the genusProtolichus (Table I), allow to recover clear host specificityat generic and specific level of mites to certain parrot taxa(species, genera or subfamilies of hosts), that suggests animportant participation of cospeciation events in the origin ofobserved host-mite associations. The genus Protolichus is gen-erally restricted to the lories (Loriinae); however, it needs aseparate taxonomic revision and checking of known host as-sociations. The Apexolichus complex is clearly restricted tothe parrots of the tribe Platycercini, distributed mainly in

Australia and Indo-Malaya. Each species of this complex iseither monoxenous or associated with several species of onegenus. Titanolichus triangulifer is associated with hosts fromthe genera Barnardius and Northiella; this latter host is prob-ably a case of an accidental contamination.

The morphologically most derived species of the genusApexolichus (A. velifer, A. neglectus) occur on host from out-side of Australia, on Fijian Islands and New Caledonia, re-spectively. All other mite genera involved in the analysis alsoshow a restriction to certain suprageneric taxon of hosts. Thegenera Nestorilichus and Uropsittacolichus are restricted tothe tribes Nestorini and Cyclopsittacini (Psittacinae), respec-tively. Mites of the genus Loriprotolichus occur on lories(Loriinae); according to recently accumulated host-associa-tion data they do not overlap in distribution among host gen-era with representatives of the genus Protolichus, which isalso restricted to this tribe. The genus Calyptolichus is knownonly from large sized cockatoos of the genus CalyptorhynchusDesmarest, 1826 (Cacatuidae, Cacatuinae, Calyptorhynchini).

Acknowledgements. We wish to thank Dr. Peter-René Becker, thedirector of the Natural History Department in the Übersee Museum

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Table I. Host-parasite associations of feather mite species of the Protolichus generic group (excluding Protolichus) with parrots from Australiaand Indo-Malaya

Mite Host Host tribe/ Host family Location Sourcesubfamily

Apexolichus affinis Platycercus elegans Platycercini Psittacidae Australia Mégnin & Trouessart(1884), present study

" Pl. eximius Platycercini Psittacidae Australia: Tasmania Favette & Trouessart (1904),present study

" Pl. caledonicus Platycercini Psittacidae Australia: Tasmania present study" Pl. flaveolus Platycercini Psittacidae Australia Mégnin & Trouessart (1884),

present study" Pl. icterotis Platycercini Psittacidae Australia present study

A. neglectus Eunymphicus cornutus Platycercini Psittacidae New Caledonia Favette & Trouessart (1904)A. lathami Lathamus discolor Platycercini Psittacidae Australia Mironov et al. (2003a)A. platycerci Platycercus adscitus Platycercini Psittacidae Australia Mironov et al. (2003a)A. velifer Prosopeia personata Platycercini Psittacidae Fijian Islands Trouessart (1898)

" Pr. tabuensis Platycercini Psittacidae Fijian Islands Trouessart (1898),present study

A. psephoti Psephotus varius Platycercini Psittacidae Australia present study" Ps. haematonotus Platycercini Psittacidae Australia present study

Titanolichus chiragricus Pesoporus wallicus Platycercini Psittacidae Australia Mégnin & Trouessart (1884)T. seemani Neophema chrysogaster Platycercini Psittacidae Australia: Tasmania Dabert et al. (2006)

" N. splendida Platycercini Psittacidae Australia present study" N. pulchella Platycercini Psittacidae Australia present study

T. triangulifer Barnardius zonaris Platycercini Psittacidae Australia present study" Northiella haematogaster* Platycercini Psittacidae Australia present study

Sideroferus lunula Melopsittacus undulatus Platycercini Psittacidae Australia Robin & Mégnin (1877),Favette & Trouessart (1904)

Uropsittacolichus elegans Cyclopsitta d. Cyclopsittacini Psittacidae Indonesia: Irian Jaya Atyeo & Pérez (1982)diophthalmica

" C. d. coccineifrons Cyclopsittacini Psittacidae Papua New Guinea Atyeo & Pérez (1982)

Uropsittacolichus diamondi C. gulielmitertii Cyclopsittacini Psittacidae Papua New Guinea Atyeo & Pérez (1982)" Psittaculirostris desmarestii* Cyclopsittacini Psittacidae Papua New Guinea Atyeo & Pérez (1982)

Calyptolichus favettei Calyptorhynchus funereus Calyptorhynchini Cacatuidae Australia Gaud & Atyeo (1996), Mironov & Dabert (2007)

Nestorilichus atyeoi Nestor meridionalis Nestorini Psittacidae New Zealand Mironov & Dabert (2007)Loriprotolichus falculiger Glossopsitta concinna Loriinae Psittacidae Papua New Guinea Trouessart (1884)L. charmosynae Charmosyna papou Loriinae Psittacidae Australia Mironov & Dabert (2007)

*Probable contamination.


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(Bremen, Germany), for making the ornithological collection avail-able for our study. We are also grateful to Dr. Owen Seeman (Queens-land Museum, Brisbane, Australia) for provision with Apexolichusspecimens from some previously unexplored Australian parrots. Thestudy was supported by the Committee of Science Promotion of theUniversity of Vechta (KFN 2004 and 2005), Germany, and in part(for SM) by the Russian Foundation for Basic Research (grant No07-04-00426a).

References

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Atyeo W.T. 1989a. Aralichus hastifolia (Mégnin and Trouessart), aspecies of feather mites (Acarina, Pterolichidae) restricted tospecies of the parrot genus Enicognathus Gray (Aves, Psit-tacidae). Journal of the Kansas Entomological Society, 62,329�334.

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Systematic revision of genera Apexolichus and Titanolichus 79

Appendix 1. Character argumentation for morphological characters used in the phylogenetic analysis of the Apexolichus-Protolichuscomplex

No. Characters

1. Lateral extension of palps: (0) absent, (1) directed anteriorly, (2) directed posteriorly2. Palpal seta dp1: (0) simple, (1) scale-like3. Male polymorphism: (0) absent, (1) present4. Length of opisthosomal lobes in males: (0) not longer than wide, (1) equal or much longer than wide5. Distal enlargement of lobes in heteromorph males: (0) absent, (1) present6. Lobar apices in males: (0) bluntly rounded, (1) acute, (2) obliquely cut, (3) narrowed distally7. Spine in terminal cleft in males: (0) absent, (1) present8. Interlobar membrane in males: (0) present, (1) absent9. Form of interlobar membrane in males: (0) narrow, (1) occupies whole terminal cleft

10. Terminal lamellae in males: (0) present, (1) absent11. Angular lateral membrane at level of h2 in males: (0) absent, (1) present12. Pronotal shield in males: (0) covers almost whole prodorsum, (1) covers median part13. Pronotal shield in females: (0) covers almost whole prodorsum, (1) covers median part14. Splitting of pygidial shield from hysteronotal shield in females: (0) not split, (1) split15. Scapular shields in females: (0) absent, (1) present16. Form of scapular shields in females: (0) large, simple, (1) reduced, (2) large, fused with ball-like structures17. Epimerites I in males: (0) free, (1) fused in heteromorph, free in homeomorph, (2) fused in all forms, (3) touching by posterior halves18. Ball-like inflation of bases of epimeres II in males: (0) absent, (1) present19. Ball-like inflation of bases of epimeres II in females: (0) absent, (1) present20. Epigynum: (0) present, (1) absent21. Genital apodemes in males: (0) absent, (1) present22. Anterior tips of male genital apodemes: (0) short, (1) long23. Male genital apodemes and epimerites IV: (0) not connected, (1) connected24. Posterior parts of male genital apodemes: (0) distant, (1) adjacent25. Male genital apodemes posterior to genital organ: (0) without sclerotized shield(s), (1) a pair of sclerotized shields or single sclero-

tized bridge present26. Corolla of anal suckers in males: (0) smooth, (1) dentate27. Distance between setae se:se in both sexes: (0) longer than width of gnathosoma base, (1) equal to or less than width of gnathosoma

base28. Setae c3 in both sexes: (0) lanceolate, (1) setiform29. Setae h1 in males: (0) present, (1) absent30. Setae si in males: (0) setiform, very short, (1) setiform, not longer than distance se:se, (2) spiculiform, longer than distance se:se31. Setae si in females: (0) setiform, very short, (1) setiform or spiculiform, subequal to se:se, (2) spiculiform, longer than se:se32. Setae c2 in males: (0) setiform, (1) spiculiform33. Setae c2 in females: (0) setiform, (1) spiculiform34. Setae d2 in males: (0) setiform, (1) spiculiform35. Setae d2 in females: (0) setiform, (1) spiculiform, wide, (2) spiculiform, thin36. Setae e2 in males: (0) setiform, short, (1) very large and long spicules37. Setae e2 in females: (0) setiform, (1) spiculiform or lanceolate38. Setae e1 in males: (0) setiform, (1) sickle-like, L-shaped, (2) lanceolate39. Setae f2 in males: (0) setiform, (1) short, curved, with membranous enlargement, (2) long, curved, extended beyond lobar apices, (3)

lanceolate, straight40. Setae ps1 in males: (0) setiform, (1) foliform, (2) triangular41. Cupules ih in males: (0) indistinct, (1) well-developed42. Ventral apophyses on genua I in males: (0) absent, (1) present43. Ventral apophyses on tibiae I in males: (0) absent, (1) present44. Ventral apophyses of tibiae II in males: (0) absent, (1) present45. Dorsal apophyses of tibiae I in males: (0) absent, (1) present46. Dorsal apophyses of tibiae II in males: (0) absent, (1) present47. Dorsal apophyses of tarsi I in males: (0) absent, (1) present48. Shape of dorsal apophyses of tarsi I in males: (0) short dorsobasal, (1) directed anterior, bearing solenidion ω149. Dorsal apophyses of tarsi II in males: (0) absent, (1) present50. Shape of dorsal apophyses of tarsi II in males: (0) hook-like, (1) rounded with solenidion ω151. Ventral extension of tarsi II in males: (0) absent, (1) present52. Solenidion σ2 in both sexes: (0) present, (1) absent53. Setae d on tarsi IV in males: (0) button or button with nipple, (1) barrel-like with cap, (2) spine, (3) pale patch54. Seta e on tarsi IV in males: (0) button or button with nipple, (1) barrel-like with cap, (2) spine, (3) pale patch


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Systematic revision of the feather mite genera Apexolichus Gaud et Atyeo and Titanolichus Gaud et...

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