Belgian Journal ofEntomology 5 (2003) : 37-77

A review of feather of the genus Neopteronyssus (Astigmata Pteronyssidae) associated with woodpeckers (Piciformes Picidae) of the Old World

by SERGE V. MIR.ONOV

Zoological Institute, Russian Academy of Sciences, Universitetskaya emb. I, Saint-Petersburg, 199034, Russia (e-mail: @zin.ro).

Abstract

Eight new species of the feather genus Neopteronyssus MIRONOV, 2002 are described form various woodpecker species (Piciformes, Picidae) from Africa, Asia and Indonesia: Neopteronyssus bilineatus sp. n. from Dendrocopos canicapillus (BL YI'H), N. campetherinus sp. n. from Campethera c. caroli {MALHERBE), N. gecinuli sp. n. from Gecinulus grantia (McCLELLAND), N. javanicus sp. n. from Dryocopus j. javensis {HORSFIELD), N. faini sp. n. from Dinopium ra.fflesi (VIGORS & HORSFIELD), N maritus sp. n. from Dryocopus martius (LINNAEUS), N. muelleripicinus sp. n. from Mulleripicus p. pulverulentus (TEMMINCK), and N. vittatus sp. n. from Picus vittatus VIEILLOT. An improved diagnosis of the genusNeopteronyssus, and a key to all recently known species are provided. Host parasite associations of the genus and distribution of its representatives among woodpeckers of the Old World are briefly discussed. Key words: Acari, , Pteronyssidae, Neopteronyssus, systematics.

Introduction

The family Pteronyssidae OUDEMANS, 1941 includes about 13 0 species currently arranged in 22 genera (F ACCINI & ATYEO, 1981; GAUD, 1991; GAUD & ATYEO, 1996; M.IRONOV, 1989, 2001). Representatives of the family are widely distributed on birds of different families of the orders Passeriformes and Piciformes, and a few species are recorded on the Coraciiformes. The genus Neopteronyssus M.IRONOV, 2002 is one of 10 pteronyssid genera recently known from the piciforms. This genus is specific to the woodpeckers, Picidae, and its representatives are mainly known from woodpeckers distributed in Europe and Africa, and only few records of this genus have been made in North America and East Asia (CERNY & SCHUMILO, 1973; FACCINI & 38 S.V.MIR.ONOV

ATYEO, 1981; MlR.ONOV, 1989; GAUD, 1991). Up to now, the genus Neopteronyssus has included eight species associated with woodpeckers of five genera: Campethera GRAY, Dendrocopos KOCH, Dryocopus BolE, Picoides LACEPEDE, and Picus LINNAEUS. In the course of our taxonomic and biodiversity study of the family Pteronyssidae, carried out during past twelve years {MIR.ONOV, 1990, 1992, 1993, 2001; MIR.ONOV & KOPIJ, 1999, 2000}, a vast number of new pteronyssid taxa, mainly from the passerines of Africa and Southeast Asia, have been recovered and described. In the frame of this continuing study, the main objectives of the present paper are to describe new species of the genus Neopteronyssus from woodpeckers of the Old World, to provide a new diagnosis of the genus taking into consideration characters on newly found representatives, to construct a key to all known species, and to discuss in general lines host associations and a distribution of the taxon in question.

Material and methods

The main part of the material used in the present study was borrowed in a loan from the University of Georgia (Athens, USA). Lesser part of examined material is a feather mite collection deposited in the Zoological Institute, Russian Academy of Sciences (Saint Petersburg, Russia). A new diagnosis of genus and descriptions of new species are given in the standard formats used for pteronyssid taxa of respective rank (FACCINI & ATYEO, 1981; Mironov, 1992, 2001}. The general morphological terms, leg and idiosomal chaetotaxy follow GAUD & ATYEO (1996). All measurements in the descriptions are given in micrometers (J.lm). Since the number of specimens in most type series was restricted, a full set of measurements is given only for the holotype (male} and one paratype (female). The range of idiosomal size (length, width) is displayed for other paratype specimens. Scientific names of birds follow "A Complete Checklist of the Birds of the World" of Howard & Moore (I 991 ). Abbreviations used in collection data and to point out where type materials of new species are deposited: AMNH - American Museum of Natural History, New York, USA; MZUM- Museum of Zoology, University ofMichigan, Ann Arbor, USA; NU - Nebraska University, Lincoln, USA; UGA - University of Georgia, Athens, USA; ZISP - Zoological Institute, Saint Petersburg, Russia. Where a sample is provided with two numbers, the first refers to a collection number of a mite specimen, the second is a collection number of a respective host specimen.

Systematic part Family Pteronyssidae OUDEMANS, 1941 Genus Neopteronyssus MIRONOV, 2002 Pteronyssus: CANESTRINI, 1886: 269 (part.); CANESTRINI & KRAMER, 1899: 78 (section obtusae, part.); VITZTIIUM, 1929: 97 (part.); GAUD&MOUCHET, • Neopteronyssus from woodpeckers of the Old World 39

1959: 513 (part.); GAUD & TILL, 1961: 276 (part.); CERNY & SCHUMILO, 1973: 89 (part.); FACCINI & ATYEO, 1981: 55; MlRONOV, 1989: 106; GAUD, 1991: 116. Neopteronyssus MIRONOV, 2002: 142.

Type species: Dermaleichus picinus KOCH, 1941 from Dryocopus martius (LJNNAEUS), by original designation. Both sexes. Epirnerites I fused Y -shaped, sternum about 112 total length of epimerites. Seta vi present. Prodorsal shield variously shaped: large triangle covering most part of prodorsum (Fig. I) or narrow, longitudinal plate ih median part of propodosoma (Figs 13, 14); this shield not fused with scapular shields, not encompassing setae cl, scapular setae se, si on or off the shield. Setae c2 narrowly lanceolate, needle-like, or setiform, short. Setae c3 lanceolate, short. Setae dp2 of palpae simple, hair-like. Setae ba of tarsi I-ll hair-like. Solenidion allonger than {J)] on legs I. Ventral membrane of tarsus I long, about 112-3/4 length of segment. Entire surface of coxal fields I, IT with striated tegument. Male. Opisthosomal lobes small, bluntly rounded or trapezoidal in shape, terminal cleft triangular or weakly marked. Terminal margin of lobes without membranes (Fig. 3a). Supranal concavity usually ovate, very short, open posteriorly into terminal cleft. Setae c2 on medial margins of humeral shields. Setae psl anterior to setae h3. Hysteronotal shield without internal sclerotized ridges. Coxal fields m open. Transventral sclerite and epiandrium commonly absent (Fig. 2); if present (pycnospilus species group), epiandrium crescent­ shaped and fused with posterior margin of transventral sclerite (Figs 46, 48, 50). Anal discs circular, large. Adanal shields represented by 1-3 small variously shaped sclerites anterior to anal slit, or these sclerites absent. Narrow adanal membranes present. Setae h3 narrowly lanceolate, short, or represented by long thin setae. Tarsus m elongated, with acute apical processus (Fig. 3). Setae r commonly longer than tarsus m. Female. Idiosoma moderately elongated. Set ofhysteronotal shields variable, . represented by three main types: two large shields - the anterior hysteronotal . plate of trapezoid form and opisthosomal plate extending to posterior end of body and commonly bearing several longitudinal ridges (picinus group) (Fig. 5); three large shields - the anterior hysteronotal plate a variously-shaped polygon and pair of opisthosomal plates commonly extending to posterior end of the body (pycnospilus group) (Figs 51-54); small unpaired or paired sclerite situated in the centre of opisthosoma and not extending to poster margin of the body (pici group) (Figs 27, 33, 34). Hysteronotal gland openings gl situated on lateral margins of opisthosomal plates, or on striated tegument (pici group). Epigynium semicircular or almost rectangular (Fig. 6). Terminal copulatory extension present, usually represented by short tube or hemispheric protrusion. The genus recently includes 16 species arranged into three species groups: picinus, pici, and pycnospilus. Hosts. Woodpeckers of the genera: Campethera GRAY, Dendrocopos KOCH, Dinopium RAFINESQUE, Dryocopus Born, Gecinulus Blyth, • 40 S.V.MIRONOV

Muelleripicus BONAPARTE, Picoides LACEPEDE, and Picus LINNAEUS. Taxonomic remark. As the matter of fact, the taxon carrying at present time a valid generic name Neopteronyssus Mm.ONOV, 2002 was recognised and morphologically characterised by FACCINI & ATYEO {1981 ), but it was treated by these authors as the genus Pteronyssus RoBIN, 1877. However, Dermaleichus picinus KocH, 1841 originally designated by ROBIN (in ROBIN, MEGNIN, 1877) as a type species of Pteronyssus, was actually misidentified by this author. This misidentification was not recognised by subsequent authors for more than a hundred of years. In the generic revision of Pteronyssidae, FACCINI & ATYEO {1981) for the first time pointed out"that these two species belonged to reliably distinct genera, but did not formally resolve the problem from the point of view of the Code of Zoological Nomenclature. In regard to the genus Pteronyssus, these authors based on the true species Dermaleichus picinus KOCH, 1841, but not on the species, which ROBIN had really used as a type species. The taxonomic problem caused by the misidentified type species of the genus Pteronyssus was solved only by the taxonomic action of correct fixation {Mm.ONOV, 2002), carried out according to the rules of the recent Code of Zoological Nomenclature. The species really used by ROBIN as a type and recently carrying a valid name Pteronyssus robini (F ACCINI & ATYEO, 1981) was fixed as the type species of the Pteronyssus. Since the generic name for the taxon Pteronyssus sensu FACCINI & ATYEO, 1981 was preoccupied, this taxon was given a new valid name, Neopteronyssus.

Key to species of Neopteronyssus MIRONOV, 2002 Males

1. Transventral sclerite and epiandrium absent (Fig. 2). Tarsus IV with two dorsobasal teeth (Fig. 4) ...... 2 Tranventral sclerite present; epiandrium present, fused with posterior margin of transventral sclerite (Fig. 39). Tarsus IV without dorsobasal teeth (Fig. 41). (pycnospilus group) ...... 12 2. Hysteronotal shield well sclerotized, with ornamentation of small lacunae and/or transverse striation. (picinus group) ...... 3 Hysteronotal shield weakly sclerotized, in some species its margins scarcely distinct, surface without ornamentation of lacunae and striation. (pici group) ...... 9 3. Prodorsal shield large triangular, encompassing bases of scapular setae and covering most part of prodorsum (Figs 1, 7) ...... 4 Prodorsal shield of another form, not encompassing bases of scapular setae, or connected with them only by weakly sclerotized lateral extensions (Figs 8, 13, 19) ...... 5

4. Width of prodorsal shield about 3/4 of hysteronotal shield width. Adanal shield represented by 3 separate sclerites (Figs 1, 2) ...... N. picinus {KOCH, 1841) Neopteronyssus from woodpeckers of the Old World 41

-. Prodorsal shield very large, equal or slightly wider than hysteronotal shield at anterior margin. Adanal shields represented by 5 separate sclerites (Figs 7, 9} ...... N.javanicus sp. n. 5. Prodorsal shield not extending beyond level of scapular setae, posterior margin of the shield without any median extension (Fig. 19} ...... 6 Prodorsal shield with ovate, onion- or finger-like extension stretching beyond the level of scapular setae (Figs 13, 14) ...... 8 6. Prodorsal shied almost rectangular, posterior margin straight or slightly convex. Adanal shield represented by pair of longitudinal ~~lerites (Figs 8, 20} ...... 7 Prodorsal shield enlarged in posterior part by 1/3, posterior margin semicircular. Adanal shield represented by a single median sclerite (Figs 19, 21} ...... N. muelleripicinus sp. n. 7. Supranal concavity nearly closed. All surface of hysteronotal shield with sparsely disposed sinuous transverse dashes (Fig. 8) . . . N. martius sp. n. Supranal concavity widely opened. Hysteronotal shield with tight transverse striation in posterior half (Fig. 20} ...... N. vittatus sp. n. 8. Posterior end ofprodorsal shield with narrow, finger-like median extension (Fig. 13} ...... · ...... N. gecinuli sp. n. Posterior end of prodorsal shield with wide extension terminating by very narrow tip (Fig. 14) ...... N. faini sp. n. 9. Pro dorsal shield well sclerotized, represented by longitudinal plate situated anterior to scapular seta row. Margins of hysteronotal shield distinct, its anterior margin deeply concave (Figs 29, 30) ...... ·...... 10 Prodorsal shields weakly sclerotized, its margins scarcely distinct; this shield occupies almost prodorsum and encompasses the bases of scapular setae. Margins of hysteronotal shield weakly distinct, anterior margin of this shield convex (Figs 25, 35) ...... 11 10. Length of idiosoma less than 470 J.lm, genua m completely extending beyond the level oflobar apices (Figs 29, 31} ...... P. elongatus {BUCHHOLZ, 1869) Length ofidiosoma longer than 470 J.lm, genua m extending beyond lobar apices by distal part only (Figs 30, 32) ...... P. pici (SCOPOLI, 1763) 11. Posterior half of hysteronotal shield with pair of heavy sclerotized longitudinal bars encompassing openings gl (Fig. 25) .. N. bilineatus sp. n. Posterior part of hysteronotal shield without any ornament (Fig. 3 5) ...... N. yungipicinus (MJRONOV, 1987) 12. Supranal concavity short (less 40 J.lm), not extending by the anterior end to the level of setae e2 (Fig. 44) ...... N. pachyprichus (GAUD, 1991) Supranal concavity forming very long median groove (over 60 llm), extending by anterior end to the level of setae el (Figs 42, 43) ...... 13 13. Tips of epiandrium extending to midlevel of genital apparatus (Fig. 46) .. 42 S.V.MIRONOV

...... N. malaconotus (GAUD, 1991) Tips of epiandrium very short, not extending to apex of genital apparatus (Fig. 50) ...... 14 14. Prodorsal shield not separated by transverse band of soft tegument into two parts ·(Fig. 42) ...... N. dicronotus (GAUD, 1991) Prodorsal shield completely separated by transverse band of soft tegument at level of scapular setae into anterior and posterior plates (Fig. 38) . . 15 15. Length of idiosoma less than 380 f.1m. Anterior part of supranal concavity significantly thinner than posterior part (Fig. 45) .."· ...... N. pycnospilus {GAUD & ATYEO, 1959) Length of idiosoma more than 400 f.1m. Anterior part of supranal concavity monotonously attenuates to anterior end (Fig. 3 8) ...... N. campetherinus sp. n.

Females

1. Hysteronotal shield represented by large unpaired anterior piece and large paired or unpaired pygidial piece (Figs 5, 24). Anterior hysteronotal piece always in a form of inverted trapezium, opisthosomal piece(s) commonly extending to posterior margin of opisthosoma (picinus group) ...... 2 Another arrangement of hysteronotal shields ...... 8 2. Prodorsal shield is large triangular, covering almost prodorsum and encompassing bases of scapular setae (Fig. 5) ...... : ...... 3 Prodorsal shield is a relatively small plate situated mainly anterior to the row of scapular setae, not encompassing their bases or connected with them by weakly scletotized lateral extensions (Figs 17, 23) ...... 4 3. Width of prodorsal shield less than width of anterior hysteronotal plate. Opisthosomal plate of hysteronotal shield almost parallel-sided (Fig. 5) ...... N. picinus (KOCH, 1841) Prodorsal shield very large, subequal or slightly wider than anterior hysteronotal plate. Opisthosomal plate of hysteronotal shield enlarged in posterior half, i.e. posteriorly to the level of setae e2 (Fig. 1 ~) ...... N. Javanlcus sp. n. 4. Opisthosomal plate represented by two widely separated longitudinal sclerites of lanceolate form; surface of these sclerites without ornamentation (Fig. 24) ...... N. vittatus sp. n. Opisthosomal plate entire, with several longitudinal sclerotized ridges (Fig 17) ...... 5 5. Posterior end of opisthosomal plate is two times wider than anterior end. Prodorsal shield extending beyond the row of scapular setae by median extension (Figs 18) ...... 6 Posterior end of opisthosomal plate is 1.2-1.5 times wider than anterior end. Prodorsal shield not extending beyond row of scapular setae (Figs 12, Neopteronyssus from woodpeckers of the Old World 43

• 23) ...... 7 6. Posterior median extension of prodorsal shield finger-like, weakly sclerotized. Opisthosomal plate extending to bases of setae h2 (Fig.l7) ...... N. gecinuli sp. n. Posterior median extension of prodorsal shield wide, terminating by narrow tip. Opisthosomal plate not extending to bases of setae h2 (Fig. 18) · · · · · · · · · · · · · · · · · · · · · · · · · · · · ...... N.faini sp. n. 7. Opisthosomal plate with deep angle-like incision on posterior margin. Subhumeral setae c3 subequal in length to trochanters m (Fig. 12) ...... N. martius sp. n. -- Opisthosomal plate without incision on posterior margin. Subhumeral setae c3 is two times longer than trochanter m (Fig. 23) ...... N. muel/eripicinus sp. n. 8. Prodordsal shield is well sclerotized and situated only anterior to scapular seta row (Figs 33, 34), or it covers almost prodorsum and is weakly sclerotized (Figs 27, 37) (pici group) ...... 9 Prodorsal shield is well sclerotized, occupies only median part of prodorsum; this shield always extends beyond scapular setae, but not encompasses their bases (Figs 51-54) (pycnospilus group)...... 12 9. Hysteronotal shields entire, in a form ofX or inverted Y, situated in central area ofhysterosoma (Figs 33, 34) ...... 10 Hysteronotal shield represented by pairs of narrow longitudinal sclerotized bands situated in median part ofhysterosoma (Fig. 27) ...... 11 10. Hysteronotal shield X-shaped, with very long anterior branches (Fig. 33) ...... N. e/ongatus {BUCHHOLZ, 1869) Hysteronotal shield as inverted Y (Fig. 34) ..... N. pici (SCOPOLI, 1763) 11. Longitudinal hysteronotal bands extending by anterior ends to level of dorsolateral setae d2 (Fig. 37) ...... N. ynigipicinus {MIR.ONOV, 1987) Longitudinal hysteronotal bands extending by anterior ends to level of humeral setae cp. (Fig. 27) ...... N. bilineatus sp. n. 12. Hysteronotal shield entire, in a form of large inverted Y (Figs 51, 53) . 13 Hysteronotal shield represented by unpaired anterior plate and paired opisthosomal plates, or only by a pair of opisthosomal plates (Figs 52, 54) ...... 14 13. Posterior ends of hysteronotal shield not extending to bases of setae h2, h3. Prodorsal shield not separated by transverse furrow of soft tegument into two parts (Fig. 51) ...... N. dicronotus (GAUD, 1961) Posterior ends of hysteronotal shield extending to bases of setae h2, h3. Prodorsal shield almost completely separated into two pieces by deep lateral incisions allevel of scapular setae (Fig. 53) ...... N. pachyphrychus (GAUD, 1961) 14. Hysteronotal shield represented only by pair of longitudinal shields G 44 S.V. MIR.ONOV

extending by anterior ends to level of trochanters IV (Fig. 52) ...... N. malaconotus (GAUD, 1961) Hysteronotal shield represented by anterior plate in a form of polygon and pair of opisthosomal plates ...... 15 15. Anterior plate of hysteronotal shield weakly connected with opisthosomal plates, posterior margin of anterior plate with semicircular incision (Fig. 28) ...... N. campetherinus sp. n. Anterior plate of hysteronotal shield clearly separated from opisthosomal plates {Fig. 54) ...... N. pycnospilus (GAUD & MOUCHET, 1959)

Species group picinus Diagnosis Male. Opisthosoma with small trapezium-like lobes; terminal cleft well developed; interlobar membrane narrow, with triangular incision; transventral sclerite and epiandrium absent; tarsus IV with 2 dorsobasal teeth (Figs 1-4). Female. Dorsal hysteronotal shields represented by two large pieces: anterior hysteronotal plate in a form of inverted trapezium, opisthosomal plate commonly entire, provided with longitudinal sclerotized ridges and extending to posterior end of the body ( opisthosomal plate paired and lacking ridges in N. vittatus). Remark. This species group includes 7 species; formerly it was based only on the type species of the genus {MIRONOV, 2002).

1. Neopteronyssus picinus (KoCH, 1841) (Figs 1-6)

DermaleichuspicinusKocH, 1841: fasc. 33, N 16, 17. Pteronyssus picinus: 0UDEMANS, 1937: 2202 (part.); FACCINI & ATYEO, 1981: 57,fig.45-48;MIRONOV, 1989: 107,fig. 1,4,2,4,5,3. Neopteronyssuspicinus: MIR.ONOV, 2002: 143. Pteronyssus gaudi CERN¥, 1963: 651. Material examined. 1 male, 3 females (UGA 3366, AMNH 552 708) from Dryocopus martius (LINNAEUS), Gennany, Renthendorf, 8.V.1832, coli. unknown; 16 males, 20 females (ZISP 4233), same host, Russia, Leningrad Prov., Gumbaritzy, 22. VIIT.l981, S. V. MmoNov; 4 males, 4 females (ZISP 4240), same host, Russia, Volzhsko-Kamskij kraj (recently Tatarstan), Lubyany, 15.XI.1958, coli. unknown; 2 females (ZISP 1487), same host, Russia, Moscow Prov., Zvenigorod, IQ.VIIT.l948, A LANGE; 10 males, 10 females (ZISP 4242), same host, Moldavia, Chemovitzy Prov., Vizhnitzkij dist, 27. VII.l964, M LUNKASHU; 1 male (ZISP1823), same host, Russia, Primorye, Sudzukhin reserve, 17 .IV.l944, M VOLKOV A. This species is reliably known only from the type host, the black woodpecker Dryocopus martius, in Europe and the Russian Far East {FACCINI & ATYEO, 1981; MIR.ONOV, 1989, 2002). Records of this species from other species of European woodpeckers given by some earlier authors (Canestrini & Kramer, 1899; Oudemans, 1929, 1937; Vitzthum 1929} are obviously incorrect, because this species for a long time was erroneously considered as a synonym of other pteronyssid species associated with these birds (Mm.ONOV, 2002). Neopteronyssus from woodpeckers of the Old World 45

so..-m

Figs 1-4. Neopteronyssus picinus, male. 1 : idiosoma, dorsal view, 2: idiosoma, ventral view, 3: tarsus ID, dorsal view, 4: tarsus IV, dorsal view. Taxonomic remark. This species was originally designated as a type species of the genus Pteronyssus ROBIN, 1877, but it actually was misidentified by ROBIN (in ROBIN & MEGNIN, 1877). FACCINI & ATYEO (1981) treated this species in the sense of KocH (1841) as the type species of Pteronyssus. After the correct fixation of the type species of the genus Pteronyssus {MIR.ONOV, 2002), taxon Pteronussus sensu FACCOO & ATYEO, 1981 was given a new name, Neopteronyssus, and Dermaleichus picinus KocH had retained as a type species. • 46 S.V. MIRONOV

Figs 5-6. Neopteronyssus picinus, female. 5: dorsal view, 6: ventral view.

2. Neopteronyssusjavanicus MIRONOV sp. n. (Figs 7, 9, 11)

Type material. Male holotype, 2 male and 2 female paratypes UGA 9453, AMNH 447677) from Dryocopus j. javensis (HORSFIELD), SW Borneo, Riam, 1o40'S, Ill o 54' E, 26.XI.1935, J.J. MENDEN. Holotype, 1 male and 1 female paratypes - MZUM, 1 male and 1 female paratypes- ZISP. Additional material. 1 male, 2 females (UGA 9454, AMNH 552592) from the same host, Malaysia, Trang prov., Lamra, 9.1.1910, H. C. ROBINSON. Description Male (holotype). Length of idiosoma 557, width of idiosoma 263 (idiosomal size in 2 paratypes 550-562 x 250-265). Prodorsal shield large, triangular, covering most part of prodorsum, encompassing bases of scapular setae, Neopteronyssus from woodpeckers ofthe Old World 47

Figs 7-8. Males of the genus Neopteronyssus, dorsal view of idiosoma. 7 : Neopteronyssus javanicus, 8: N. martius. posterior part with transverse striation, posterior margin almost straight, length along median line 164, width at posterior margin 210, setae se separated by 106. Length of hysterosoma 380. Setae c2 needle-like, 60 in length. Subhumeral setae c3 lanceolate, with acute apex, 62 in length. Hysteronotal shield: anterior margin straight, greatest length 380, width at anterior margin 208, surface with numerous pit-like lacunae in median part of the shield, and sinuous transverse striation in anterior and lateral parts (Fig 7). Dorsal setae el slightly anterior to level of dorsal hysteronotal openings gl. Opisthosoma with short trapezoid lobes carrying setae h2 and h3 on apices. Terminal cleft about 22 in length, 40 in width (distance between setae h3}, interlobar membrane narrow, incision in interlobar membrane 15. Distance between setae and openings: c2-d2 133, d2-e2 135, d2-gl 10, el-gl 4-6, h2-h2 55. Setae h3 lanceolately enlarged in basal part. Transventral sclerite absent, anterior ends of epimerites illa with sclerotized areas of irregular form, epiandrium absent. Setae 3a anterior to 3b. Genital arch 50 in length, 28 in width, tip of aedeagus enlarged, genital setae g indistinct, setae 4a slightly posterior at base of the arch. Adanal membranes narrow. Adanal shield represented by 5 sclerites: pair of triangular sclerites carrying setae ps3, pair of longitudinal sclerites anterior 48 S.V.MIRONOV

•

200J.1m

9

Figs 9-10. Males ofthe genusNeopteronyssus, ventral view ofhysterosoma. 9: Neopteronyssus javanicus. 10 : N. martius.

to these setae, and angle-like sclerite at anterior end of anal slit (Fig. 9). Tarsus m 78 in length; seta r 1.5-2 times longer than this segment, seta s narrowly lanceolate in apical part. Tarsus IV with 2 acute dorsobasal teeth. Female (paratype). Length of idiosoma 550, width of idiosoma 276 (idiosomal size in 2 other paratypes 542-554 x 265-275). Prodorsal shield as in the male, 164 x 209, setae se separated by 110. Length of hysterosoma 3 55. Setae c2 needle-like, 68 in length. Subhumeral setae c3 narrowly lanceolate, with acute apex, 82 in length. Arrangement of hysteronotal shields: anterior hysteronotal plate and opisthosomal plate separated by crescent-shaped transverse furrow of soft tegument (Fig. 11 ). Anterior hysteronotal plate a large inverted trapezium, its anterior margin straight, posterior margin concave, surface with transverse striation, greatest length 175, length along median line 140, width at anterior margin 225. Opisthosomal plate extending to bases of setae h2, h3, surface with 5 longitudinal heavy sclerotized ridges, posterior margin with little incision between setae psi, setae e2 and openings gl on lateral margins of the plate, greatest length of opisthosomal plate 21 0, greatest width 178. Terminal copulatory extension represented by small obliterated cone between setae h3, about 22 in length, 31 in width at base. Distance between setae and seta rows: c2-d2 116, d2-e2 15, e2-.f2 110, el-e2 86, h2-h2 86. Epigynium semicircular, 58 in length, 98 in width.

Differential diagnosis. The new species is closely related to the type species of the genus, Neopteronyssus picinus. Both sexes of N. javanicus are Neopteronyssus from woodpeckers of the Old World 4Sf

200~o~m

Figs 11-12. Females of the genus Neopteronyssus, dorsal view ofidiosoma. 11 : Neopteronyssus javanicus, 12 : N. martius. distinguished by having the larger prodorsal shield, which is equal or slightly wider than the anterior margin of hysteronotal shield, the males differ by the adanal shield consisting of 5 separate sclerites (Figs 7, 9), and the females differ by the opisthosomal plate enlarged in a posterior half, beyond the level of setae e2 (Fig. 11 ). In N. picinus, the width of prodorsal shield is less than the width of hysteronotal shield; in the males of this species, the adanal shield consists of 3 sclerites (Figs 1, 2); in the females, the opisthosomal plate is almost parallel-sided (Fig. 5).

Etymology. The specific epithet derives from the species name of the host.

3. Neopteronyssus martius MIRONOV sp. n. (Figs 8, 10, 12) Type material. Male holotype, 1 male and 3 female paratypes (ZISP 1483) from Dryocopus martius (LINNAEUS), Kazakhstan Republic, North Kazakhstan Prov., 13.IX.l951, BEZUKLADNIKOV. Holotype and paratypes­ ZISP. 50 S. V. MIRONOV

Description Male (holotype).Length ofidiosoma 517, width ofidiosoma 218 (idiosomal size in one paratype 495 x 180). Prodorsal shield as almost rectangular plate situated anterior to row of scapular setae and not encompassing their bases, with weakly convex posterior margin (Fig. 8), length of the shield along median line 92, greatest width 55. Setae se separated by 71. Length of hysterosoma 348. Setae c2 needle-like, thin, 46 in length. Subhumeral setae c3 narrowly lanceolate, with acute apex, 44 in length. Hysteronotal shield: anterior margin deeply concave, greatest length 340, width at anterior margin 156, surface with sparsely disposed sinuous transver~e dashes (Fig 8). Dorsal setae el slightly anterior to level of dorsal hysteronotal openings gl. Opisthosoma with short trapezoidal lobes canying setae h2 and h3 on apices. Terminal cleft triangular, about 28 in length, 24 in width, interlobar membrane narrow, incision in interlobar membrane 26. Distance between setae and openings: c2-d2 123, d2-e2 133, d2-gl 82, el-gl 4-6, h2-h2 42. Setae h3 lanceolately enlarged in basal part. Transventral sclerite absent, anterior ends of epimerites m with sclerotized areas of irregular form, epiandrium absent. Setae 3a anterior to 3b . .Genital arch 45 in length, 30 in width, tip of aedeagus enlarged, genital setae g indistinct. Adanal membranes narrow. Adanal shield represented by pair of longitudinal sclerites anterior to setae ps3 (Fig. 10). Tarsus m 66 in length; seta r 2-2.5 times longer than this segment, seta s narrowly lanceolate in apical part. Tarsus IV with 2 acute and short dorsobasal teeth. Female (paratype). Length of idiosoma 570, width of idiosoma 250 (idiosomal size in 2 other paratypes 550-555 x 220-230). Prodorsal shield as in the male, 106 x 71, setae se separated by 84. Length of hysterosoma 380. Setae c2 needle-like, thin, 40 in length. Subhumeral setae c3 narrowly lanceolate, with acute apex, 45 in length. Arrangement of hysteronotal shields: anterior hysteronotal plate and opisthosomal plate separated by crescent­ shaped transverse furrow of soft tegument (Fig. 12). Anterior hysteronotal plate a large inverted trapezium, its anterior margin deeply concave, posterior margin almost straight, surface with sparsely disposed transverse sinuous lines, greatest length of the plate158, length along median line 120, width at anterior margin 175. Opisthosomal plate slightly enlarged in posterior part, extending to bases of setae h2, h3, surface with 5 longitudinal heavy sclerotized ridges, posterior margin with angular incision between setae psl, setae e2 and openings gl on lateral margins of the plate, greatest length 200, greatest width 126. Terminal copulatory extension represented by large hemisphere between setae h3, about 20 in length, 46 in width at base. Distance between setae and seta rows: c2-d2 128, d2-e2 115, e2-fl 88, el-e2 54-10, h2-h2 93. Epigynium semicircular, 58 in length, 98 in width.

Differential diagnosis. The new species is similar to Neopteronyssus picinus and N. javanicus, mentioned above. Both sexes of N. martius easily differ from these species by having a small, almost rectangular prodorsal shield, which is not encompassing the bases of scapular setae, and deeply concave Neopteronyssus from woodpeckers of the Old World 51

200~m

14 Figs 13-14. Males of the genus Neopteronyssus, dorsal view of idiosoma. 13 : Neopteronyssus gecinuli, 14 : N. faini. anterior margin of hysteronotal shield (Figs 8, 10, 12). In N. picinus and N. javanicus, the prodorsal shield is large triangular, covering almost prodorsum and encompassing bases of scapular setae, the anterior margin of hysteronotal shield in males and this margin of anterior hysteronotal plate in females are straight (Figs 1, 4, 7, 11).

Etymology. The specific epithet is directly accepted from the species name ofthe host.

4. Neopteronyssus gecinuli MIRONOV sp. n. (Figs 13, 15, 17)

Type material. Male holotype, 3 male and 4 female paratypes (UGA 9500, AMNH- no number) from Gecinulus grantia (MCCLELLAND), SE China, Kwangtung prov., Tokien, Autumn 1910, P. D. BERGEN. Holotype, 1 male and 2 female paratypes - MZUM, 2 male and 2 female paratypes - ZISP. 52 S.V. MIRONOV

Figs 15-16. Malesofthe genusNeopteronyssus, ventral view ofhysterosoma. 15: Neopteronyssus gecinuli, 16 : N. faini. Description Male {holotype). Length ofidiosoma 535, width ofidiosoma 276 (idiosomal size in 3 paratypes 525-540 x 270-280). Prodorsal shield as a re(!tangular plate situated anterior to row of scapular setae, with weakly sclerotized finger-like extension on posterior margin, length of main part of the shield 102, length including posterior extension 133, greatest width 70 (Fig. 13). Setae se separated by 90. Length of hysterosoma. 348. Setae c2 needle-like, 45 in length. Subhumeral setae c3 narrowly lanceolate, with acute apex, 51 in length. Hysteronotal shield: anterior margin concave, almost surface with transverse striation, greatest length 340, width at anterior margin 177. Dorsal setae el and openings. gl approximately at the same transverse level. Opisthosoma with short trapezoid lobes carrying setae h2 and h3 on apices. Supranal concavity widely opened into terminal cleft, length of the cleft including concavity 64, width of cleft at level of setae h3 54. Interlobar membrane narrow, length of incision in the membrane 20. Distance between setae and openings: c2-d2 102, d2-e2 144, d2-gl 80, el-gl 0-6, h2-h2 72. Setae h3 hair-like. Transventral sclerite absent, anterior ends of epimerites ma with T -shaped sclerotized area, epiandrium absent. Setae 3a anterior to 3b. Genital apparatus 26 in length, 10 in width, aedeagus with coiled, very thin apical part. Genital setae g indistinct. Setae 4a slightly posterior at base of the arch. Adanal membranes narrow. Adanal shield represented by single median sclerite situated anterior to of anal slit (Fig. 15). Tarsus m 75 in length; seta r slightly shorter than this segment, seta s slightly enlarged. Tarsus IV with 2 acute dorsobasal teeth. Female (P.aratype). Length of idiosoma 522, width of idiosoma 267 (idiosomal size in other paratypes 515-526 x 265-270). Prodorsal shield almost Neopteronyssus from woodpeckers of the Old World 53

•

Figs 17-18. Females the genus Neopteronyssus, dorsal view of idiosoma. 17 : Neopteronyssus gecinuli, 18 : N. faini. as in the male, but posterior margin slightly convex, length of the main along median line 110, length including extension 133, greatest width 100. Setae se separated by 95. Length of hysterosoma 348. Setae c2 needle-like, 45 in length. Subhumeral setae c3 narrowly lanceolate, with acute apex, 57 in length. Arrangement of hysteronotal shields: anterior hysteronotal plate and opisthosomal plate separated by transverse band of soft tegument (Fig. 17). Anterior hysteronotal plate a large inverted trapezium, its anterior margin concave, posterior margin straight, surface with sparse transverse striation, greatest length 120, length along median line 80, width at anterior margin 177. Opisthosomal plate extending to bases of setae h2, h3, attenuate in anterior part, surface with ?longitudinal ridges (median ridge weakly sclerotized and in some specimens scarcely visible), posterior margin with shallow incision, greatest length 200, greatest width 110. Setae e2 on lateral margins of opisthosomal plate, openings gl on striated tegument. Terminal copulatory extension as short tube situated asymmetrically between setae h3, about 20 in length, 11 in width at apex. Distance between setae and seta rows: c2-d2 102, d2-e2 115, e2-j2 93, el-e2 68, h2-h2 98. Epigynium semicircular, 57 in length, 106 in width. 54 S.V. MlR.ONOV

.Differential diagnosis. Among the picinus species group, N. gecinuli is most similar to Neopteronyssus faini (see below) by having a tight transversal striation on hysteronotal shield in males and narrowed anterior end of opisthosomal plate in females. Both sexes of N. gecinuli differ from that species. by having a finger like extension on the posterior margin of prodorsal shield, the females are distinguished by the opisthosomal plate extending to bases ofmacrochaetae h2, h3 (Figs 13, 11).1nN.faini, the posterior extension of prodorsal shield is wide (Fig. 14), in females of this species, the opisthosomal plate does not extend to bases of marochaetae situated on the posterior margin of the body.

Etymology. The specific epithet derives from the generic name of the type host.

5. Neopteronyssus faini MIRONOV sp. n. (Figs 14, 16, 18)

Type material. Male holotype, 4 male and 4 female paratypes (NU 8391) from Dinopium ra.fflesi (VIGORS & HORSFIEID), Malaya, Mallaca, no other collection data. Holotype, 2 male,and 2 female paratypes - MZUM, 2 male and 2 female paratypes - ZISP. Description. Male (holotype). Length ofidiosoma 468, width ofidiosoma 240 (idiosomal size in 4 paratypes 452-470 x 240- 248). Prodorsal shield as rectangular plate situated anterior to row of scapular setae, with wide extension on posterior margin having narrow tip (Fig. 14), length of the shield including extension along median line 122, greatest width 69. Setae se separated by 84. Length of hysterosoma 307. Setae c2 bristle-like, 40 in length. Subhumeral setae c3 narrowly lanceolate, with acute apex, 44 in length. Hysteronotal shield: anterior margin concave, almost surface with transverse striation, greatest length 302, width at anterior margin 150. Dorsal setae el and openings gl approximately at the same transverse level. Opisthosoma with short trapezoidal lobes carrying setae h2 and h3 on apices. Terminal cleft about 33 in length, 44 in width, supranal concavity widely open into terminal cleft. Interlobar membrane narrow, length of triangular incision in the membrane 22. Distance between setae and openings: c2-d2 104, d2-e2 124, d2-gl68, el-gl 0-3, h2-h2 69. Setae h3 hair-like. Transventral sclerite absent, anterior ends of epimerites Ilia with T -shaped sclerotized areas, epiandrium absent. Setae 3a anterior to 3b. Genital apparatus 11 in length, 10 in width, aedeagus with coiled, very thin apical part. Genital setae g indistinct. Adanal membranes narrow. Adanal shield represented by single median sclerite situated anterior to anal slit (Fig. 16). Tarsus m 75 in length; seta r slightly shorter than this segment, seta s narrowly lanceolate in apical part. Tarsus IV with 2 acute dorsobasal teeth. Female (paratype). Length of idiosoma 464, width· of idiosoma 225 (idiosomal size in other 3 aratypes 457-470 x 220-230). Prodorsal shield as in the male, length along median line 126, greatest width 69. Setae se separated Neopteronyssus from woodpeckers of the Old World 55 by 82. Length of hysterosoma 308. Setae c2 bristle-like, 37 in length. Subhiimeral setae c3 narrowly lanceolate, with acute apex, 47 in length. Arrangement of hysteronotal shields: anterior hysteronotal plate and opisthosomal plate separated by transverse band of soft tegument. Anterior hysteronotal plate a large inverted trapezium, its anterior margin deeply concave, posterior margin almost straight, surface with sparse transverse striation (Fig.· 18), greatest length 124, length along median line 100, width at anterior margin 157. Opisthosomal plate attenuate in anterior part, not extending to bases of setae h2, h3, surface with 5 longitudinal ridges (median one weakly sclerotized), posterior margin with triangular incision, setae e2 on lateral margins of opisthosomal plate, greatest length 144, greatest width 93. Openings gl on striated tegument. Terminal copulatory extension as small tube situated asymmetrically between setae h3, about 10 in length, 12 in width. Distance between setae and seta rows: c2-d2 95, d2-e2 84, e2-j2 90, el-e2 44- 50, h2-h2 85. Epigynium crescent-shaped, 53 in length, 80 in width.

Differential diagnosis. The new species is most closely related to Neopteronyssus gecinuli by the structure of the prodorsal and hysteronotal shields. Both sexes of N. faini are distinguished from N. gecinuli by having a wide extension on posterior margin of prodorsal shield (Fig. 14), and the females differs by shorter opisthosomal plate, which is not extending to posterior end of the body (Fig. 18). In. N. gecinuli, the extension of the prodorsal shield is narrow, finger-like (Fig. 13) and in the females, the opisthosomal plate extends to bases of macrochaetae h2, h3 (Fig. 15).

Etymology. The species in named in an honour of the great Belgian acarologist, Prof., Dr. A. FAIN (Institute royal des Sciences naturelles de Belgique, Bruxelles, Belgique).

6. Neopteronyssus muelleripicinus MIRONOV sp. n. (Figs 19, 21, 23)

Type material. Male holotype and female paratype (UGA 9515, AMNH 483020) from Mulleripicus p. pulverulentus (TEMMINCK), Palavan Island, Mt. Talakahigan, , 15.V.1962, J. RAMos & P. GoNZALES. Holotype, paratype- MZUM. Additional material. 2 males, 2 females (UGA 9511, AMNH 552534) from M. f. ju/vus {QUOY & GAIMARD), Celebes, Lembeh Island, IV.1895, coil. unknown. Description Male (holotype). Length ofidiosoma 495, width ofidiosoma 227. Prodorsal shield a small plate not extending beyond row of scapular setae, posterior end of the shield rounded, length along median line 106, grea~est width 66. Setae se separated by 82. Length of hysterosoma 325. Setae c2 needle-like, 49 in length. Subhumeral setae c3 narrowly lanceolate, with acute apex, 64 in length. Hysteronotal shield: anterior margin slightly concave, all surface with · 56 S.V. Mm.ONOV

200~m

19

Figs 19-20. Males of the genus Neopteronyssus, dorsal view of idiosoma. 19 : Neopteronyssus muelleripicinus, 20 : N. vittatus. transverse striation (Fig. 19), greatest length 308, width at anterior margin 158. Dorsal setae el slightly posterior to openings gl. Opisthosoma with short trapezoidallobes carrying setae h2 and h3 on apices. Terminal cleft about 28 in length, 40 in width. Interlobar membrane narrow, incision in the membrane 22 in length. Distance between setae and openings: c2-d2 126, d2-e2 118, d2-gl 68, gl-el 4-8, h2-h2 51. Setae h3 hair-like. Transventral sclerite absent, anterior ends of epirnerites ma with T -shaped sclerotized areas, epiandrium absent. Setae 3a slightly anterior to 3b. Genital apparatus 37 in length, 10 in width, tip of aedeagus bifurcate. Genital setae g indistinct. Adanal membranes narrow. Adanal shield represented by single median sclerite situated anterior to anal slit (Fig. 21 ). Tarsus m 78 in length; seta r 2. 5-3 times longer than this segment, seta s slightly enlarged. Tarsus IV with 2 acute dorsobasal teeth. Female (paratype). Length of idiosoma 526, width of idiosoma 265. Prodorsal shield as in the male, encompassing bases of scapular setae by Neopteronyssus from woodpeckers of the Old World 57

Figs 21-22. Males the genus Neopteronyssus. ventral view ofhysterosoma. 21 : Neopteronyssus muelleripicinus, 22 : N. vittatus. weakly sclerotized lateral extensions, length of the shield along median line 110, greatest width 66. Setae se separated by 86. Length of hysterosoma 348. Setae c2 needle-like, 40 in length.. Subhumeral setae c3 narrowly lanceolate, with acute apex, 77 in length. Arrangement of hysteronotal shields: anterior hysteronotal plate, opisthosomal plate separated by narrow transverse band of soft tegument, and additional pair of small sclerites situated lateral to each opisthosomal plate (Fig. 23). Anterior hysteronotal plate a large inverted trapezium, it.s anterior and posterior margins slightly concave, surface with transverse striation, greatest length 168, length along median line 129, width at anterior margin 180. Opisthosomal plate slightly enlarged posterior, extending to bases of setae h2, h3, surface with 5 longitudinal ridges (median ridge weakly sclerotized than lateral ones), posterior margin almost straight, setae e2 and openings g/ on lateral margins of opisthosomal plate, greatest length 157, greatest width 95. Terminal copulatory extension as small hemisphere situated asymmetrically between setae h3, about 6 in length. Distance between setae and seta rows: c2-d2 126, d2-e2 94, e2-fl18, el-e2 71, h2-h2 82. Epigynium crescent-shaped, 64 in length, 88 in width.

Differential diagnosis. Nepteronyssus muelleripicinus is most closely related to two previous species, N. gecinuli and N. faini, by having tightly striated hysteronotal shield in males (Figs 13, 14, 19). Both sexes of N. muelleripicinus differs from these species by the absence of an extension on the posterior margin of prodorsal shield, the males are distinguished by having bifurcated apex of aedeagus and almost closed supranal concavity (Figs 19, 58 s. V. MIR.ONOV

Figs 23-24. Females the genus Neopteronyssus, dorsal view of idiosoma. 23 : Neopteronyssus muellerlpicinus, 24 : N. vittatus. 21), the females differ by longer subhumeral setae c3 (about 75) and opisthosomal plate not enlarged posteriorly (Fig. 23). In N. gecinuli and N. faini, the prodorsal shield has a median extension extending beyond the row of scapular setae; in the males, the apex of aedeagus is attenuated and supranal concavity widely open into terminal cleft (Figs 15-18); in the females, the setae c3 (45-60) in length, and opisthosomal plate two times large in posterior part than in anterior one (Figs 17, 18).

Etymology. The specific epithet derives from the generic name of the type host.

7. Neopteronyssus vittatus MIRONOV sp. n. (Figs 20, 22, 24) Type material. Male holotype, 1 male and 2 female paratypes (UGA 9484, AMNH 549329) from Picus vittatus VmiLLOT, Indonesia, Java Sea, Kangean Is., circa 1890, E. PRILLWITZ. Holotype, 1 female paratype - MZUM, 1 male and 1 female paratypes- ZISP. Additional material. 2 males 1 females (UGA 759), same host, Malaya,. Rantau Panjang, USA MedResUnit, 23.ill.1967, coli. unknown; 2 males, 2 females (NU 6309), same host, Malaya, Selangot, Rantau Panjang, 11.V.1961, coli. unknown. Neopteronyssus from woodpeckers of the Old World 59

De,cription Male (holotype). Length ofidiosoma 472, width ofidiosoma 236 (idiosomal size in I paratype 476 x 240). Prodorsal shield a small rectangular plate, not extending beyond row of scapular setae, posterior :margin slightly convex (Fig. 20), length along median line 90, greatest width 66. Setae se separated by 79. .. Length of hysterosoma 320. Setae c2 needle-like, 46 in length. Subhumeral setae c3 narrowly lanceolate, with acute apex, 3 5 in length. Hysteronotal shield: anterior margin concave, almost surface with transverse striation, greatest length 306, width at anterior margin 128. Dorsal setae el slightly anterior to openings gl. Opisthosoma with short trapezoidal lobes carrying setae h2 and h3 on apices. Terminal cleft about 37 in length, 53 in width, interlobar membrane narrow, incision in interlobar the membrane 31 in length. Distance between setae and openings: c2-d2 100, d2-e2 136, d2-g/88, el-gl 0- 6, h2-h2 69. Setae h3 hair-like. Transventral sclerite absent, anterior ends of epimerites ma with L-shaped sclerotized areas, epiandrium absent (Fig. 22). Setae 3a anterior to 3b. Genital apparatus 35 in length, 17 in width, tip of aedeagus slightly enlarged, genital setae g at midlevel of genital apparatus. Adanal membranes narrow. Adanal shield represented by pair of longitudinal sclerites situated anterior to anal slit. Tarsus m 78 in length; seta r slightly longer than this segment, seta s narrowly lanceolate. Tarsus IV with 2 acute dorsobasal teeth. Female (paratype). Length of idiosoma 490, width of idiosoma 236 (idiosomal size in 1 paratype 487 x 238). Prodorsal shield as in the male, length of the shield along median line 95, greatest width 66. Setae se separated by 75. Length of hysterosoma 321. Setae c2 needle-like, 26 in length. Subhumeral setae c3 narrowly lanceolate, with acute apex, 3 5 in length. Arrangement of hysteronotal shields: anterior hysteronotal plate and pair of opisthosomal plates (Fig 24). Anterior hysteronotal plate a large inverted trapezium, its anterior · concave, posterior margins with median incision, surface with sparsely disposed transverse striation, greatest length 168, length along median line 129, width at anterior margin 180. Opisthosomal plates lanceolate, widely separated from each other, 100-105 in length, 25-30 in width, not extending to bases of setae h2, h3, setae e2 on lateral margins of opisthosomal plates, and openings gl on striated tegument near opisthosomal plates. Terminal copulatory extension as small cone situated asymmetrically between setae h3, about 16 in length. Distance between setae and seta rows: c2-d2 98, d2-e2 89, e2-f2 50, el-e2 44, h2-h2 88. Epigynium semicircular, 51 in length, 95 in width.

Differential diagnosis. The males of Neopteronyssus vittatus are most similar to N. muelleripicinus by having the prodorsal shield without median extension and widely opened supranal concavity, while the females clearly differs from all known species of the picinus group by the structure of the opisthosomal plate, which is separated into two longitudinal sclerites (Fig. 24). The males of N. vittatus are distinguished from N. mue/leripicinus by having the adanal shield consisting of a pair of longitudinal sclerites and short subhumeral setae c3, which are shorter than trochanters m (Fig. 22). In the • 60 S.V.MIR.ONOV

• males of N. mue/leripicinus, the adanal shield is represented by single median sclerite, and the setae c3 equal or slightly longer than trochanters m (Fig. 21 ). Etymology. The· specific epithet is directly accepted from the species name of the host.

Species group pici

Male. Opisthosoma with very small rounded lobes, terminal cleft and interlobar membrane weakly expressed, transventral sclerite and epiandrium absent (Figs 25, 29-32), tarsus IV with 2 little dorsobasai teeth. Female. Dorsal hysteronotal shields are represented by a relatively small unpaired or paired sclerite(s) situated in the median part of opisthosoma and not extending to poster end of the body (Figs 27, 33, 34). Remark This species group includes 4 species.

8. Neopteronyssus bilineatus MIRONOV sp. n. (Figs 25-27)

Type material. Male holotype, female paratype (NU 16524) from Dendrocopos canicapillus {BLY1H), Thailand, Chanthaburi prov., Knao Soi Dao Tai, 4.X.1966, coil. unknown; 1 male, 1 female paratypes (NU 16525), same collection data. Holotype and 1 female paratype - MZUM, 1 male and 1 female paratypes - ZISP. Description Male (holotype). Length ofidiosoma 392, width ofidiosoma 192 (idiosomal size in male paratype 398 x 204). Prodorsal shield large triangular, covering almost prodorsum, encompassing bases of scapular setae, weakly sclerotized, posterior margin straight (Fig. 25), length along median line 122, width at posterior margin about 135. Setae se separated by 66. Length of hysterosoma 245. Setae c2 hair-like, 28 in length. Subhumeral setae c3 lanceolate, 26 in length. Hysteronotal shield: anterior margin convex, most area of the shield weakly sclerotized, opisthosomal part with pair of longitudinal heavy sclerotized bands (Fig. 25), greatest length of the shield 250, width at anterior margin 135. Dorsal setae el slightly anterior to openings gl. Opisthosoma with very short rounded lobes carrying setae h2 and h3 on terminal margin. Terminal cleft about 10 in length, 3 3 in width, supranal concavity almost closed. Interlobar membrane very narrow, interrupted in anterior end of the cleft, incision in the membrane 30 in length. Distance between setae and openings: c2-d2 100, d2-e2 93, d2-gl62, el-g/8, h2-h2 42. Setae h3 hair-like. Transventral sclerite absent, anterior ends of epimerites ma as oblique T, epiandrium absent. Setae 3b slightly anterior to 3a. Genital apparatus 22 in length, 10 in width, tip of aedeagus thin and curved. Genital setae g at midlevel of genital apparatus. Adanal membranes narrow. Adanal shield represented by single sclerite of irregular form, situated at anterior end of anal slit (Fig. 26). Tarsus m 51 in. length; seta r 3-3.5 times longer than this segment, all ventral setae of this tarsus hair-like. Tarsus IV with 2 acute dorsobasal teeth. Neopteronyssus from woodpeckers of the Old World 61

Figs 25-26. Neopteronyssus bilineatus, male. 25: idiosoma, dorsal view, 26: idiosoma, ventral view.

Female (paratype). Length of idiosoma 446, width of idiosoma 206 (idiosomal size in other paratype 442 x 205). Prodorsal shield as in the male, length along median line 133, width at posterior margin 135. Setae se separated by 68. Length of hysterosoma 321. Setae c2 hair~like, 24 in length. Subhumeral setae c3 lanceolate, 22 in length. Dorsal surface of hysterosoma with two pairs of heavy sclerotized longitudinal sclerites: anterior hysteronotal pair of these sclerites very long, 95~ 100 in length, situated in median part of hysterosoma and extending by anterior ends to level of humeral setae; pigydial pair very short, situated near bases of setae h2, h3 (Fig. 27). Setae e2 lateral to margins of anterior sclerites, openings gl and other dorsal hysteronotal setae on 62 S.V.MIRONOV

•

200~m 200~m

28

Figs 27-28. Femaes ofthe genusNeopteronyssus, dorsal view ofidiosoma. 27: Neopteronyssus bi/ineatus, 28 : N. campetherinus. soft tegument. Terminal copulatory extension small finger-like, situated asymmetrically between setae h3, about 12 x 5 in size. Distance between setae and seta rows: c2-d2 98, d2-e2 104, e2-f2 55, el-e2 22, h2-h2 65. Epigynium semicircular, 55 in length, 85 in width.

Differential diagnosis. The new species is closely related to Neopteronyssus yungipicinus {MIR.ONOV, 1987) described from Dendrocopos kizuki (TEMMINCK) by similar structure of dorsal idiosomal shields. The males of N. bilineatus differ from that species by having a pair of longitudinal sclerotized bands on the opisthosoma (Fig. 25), and the females are distinguished by having the hysteronotal sclerotized band-like sclerites extending to level of humeral setae (Fig. 27). In the males of N. yungipicinus, the hysteronotal shield has no heavy sclerotized areas (Fig. 35), and in the females, the hysteronotal sclerotized bands are shorter, extending by anterior ends to the level of setae el (Fig. 37).

Etymology. The species epithet refers to two pair of dark sclerotized bands on the dorsal surface of hysterosoma in both sexes. Neopteronyssus from woodpeckers of the Old World 63

Figs 29-30. Males of the genus Neopteronyssus, dorsal view of idiosoma. 29 : Neopteronyssus elongatus, 30 : N. pici.

9. Neopteronyssus elongatus (BUCID:IOLZ, 1869) (Fig. 29, 31, 33)

Dermaleichus elongatus BUCHHOLZ, 1869: 28, tab. 2, figs 15, 16. Pteronyssus elongatus: FACCINI & ATYEO, 1981: 57; MIRONOV, 1989: 107, figs 4, 1-2, 5, 1. Neopteronyssus elongatus: MIRONOV, 2002: 143. Pteronyssus gracilis (non NITZSCH, 1818): CANES1RINI & KRAMER, 1899: 79. Pteronyssus longiscutatus CERNY & SCHUMILO, 1973: 92, figs I C, 2C.

Material examined. 4 males, 8 females (ZISP 4262) from Dendrocopus minor (Linnaeus), Russia, Kaliningrad Prov., Rybachiy, 15. VII.1980, S. V. MIRoNov; 8 males, 10 females (ZISP 4260), same host, Russia, Leningrad Prov., Gumbaritzy, 3.VTII.l981, S.V. MIRoNov. This species is known only from the type host, the lesser spotted woodpecker Dendrocopos minor, in Europe {CERNY & ScHUMILo, 1973; FACCINI & ATYEO, 1981; MIRONOV, 1989). • 64 S.V. MIR.ONOV

•

Figs 31-32. Males ofthe genus Neopteronyssus, ventral view ofhysterosoma. 31 : Neopteronyssus elongatus, 32 : N. pici. 10. Neopteronyssus pici (SCOPOLI, 1763) (Figs 30, 32, 34)

Acarus pici SCOPOLI, 1763: 388. Alloptespici: 0UDEMANS, 1929: 695. Pteronyssus pici: FACCINI & ATYEO, 1981: 57; MIRONOV, 1989: 107, figs 2.5, 4.3, 5.2. Neopteronyssuspici: MIR.ONOV, 2002: 143. Acarus gracilisNITZSCH, 1818: 252. Analges gracilis: GIEBEL, 1871: 493. Pteronyssus gracilis: CANES1RINI, 1886: 269; OUDEMANS, 1937: 2202. Pteronyssus speciosus TYRRELL, 1882: 47, tab. 1, fig. 5. Pteronyssus monoplax CERNY, 1969: 290, figs 1, 2; CERNY & SCHUMILO, 1973: 94. Pteronyssus picinus (non KocH, 1841): BERLESE, 1886: fasc. 24, N 5.

Material examined. 8 males, 12 females (ZISP 4312) from Dendrocopos major {LINNAEUS), Russia, Kaliningrad Prov., Rybachiy, 24.IX..1979, S.V.MmoNov; 10 males, 10 females (ZISP 4330), same host, Russia, Leningrad Prov., Gumbaritzy, 3l.Vll.l981, S. V.MmoNov; 5 males, 7 females (ZISP 4267), same host, Modavia, Lozovoye, 3.X.1966, R. ScHUMILO. 15 males, 20 females (ZISP 4270) from D. leucotos (BECHSTEIN), Moldavia, 1. VI.1967, coli. unknown; 10 males, 8 females (ZISP 4322), same host, Russia, Kaliningrad Prov., Rybachiy, 28.VII.l981, S.V.MIRONOV. 2 male, 2 female (UGA 8261, AMNH387498) Neopteronyssus from woodpeckers of the Old World 65

200J,~m

33

Figs 33-34. Females of the genus Neopteronyssus, dorsal view ofidiosoma. 33 : Neopteronyssus / elongatus, 34 : N. pici.

from D. v. villosus (LINNAEUS), Canada, Ontario, Moose River, Moose Factocy, 15.VI.l926, coli. unknown. 1 male, I female (UGA 8266, AMNH 755535) from Picoides arcticus (SwAJNSON), Canada, New Brunswick, Grand Manan Island, 9.Xl923, A. L. Moss. 10 males, 14 females (ZISP 4307) from Picoides trldacty/is (LINNAEUS), Moldavia, 13.IX.l967, coil. unknown; 8 males, 8 females (ZISP 4292), same host, Russia, Leningrad Prov ., Gumbaritzy, 20.Xl981, S.V.MIRONOV. This species was originally described from the middle spotted woodpecker Dendrocopos medius {LINNAEUS) in Europe (ScoPOLI, 1763). Up to present, it has been recorded from various species of spotted and green woodpeckers of the genera Dendrocopos KOCH, Picoides LACEPEDE, and Picus LINNAEUS in Europe and North America (Table) (CERNY & SCHUMILO, 1973; FACCINI & ATYEO, 1981; MIR.ONOV, 1989). Wide distribution of N. pici and associations with woodpecker species of different genera may be probably explained by the result of natural contaminations between its hosts in the past. 66 S.V. MIRONOV

Figs. 35-37. Neopteronyssus yungipicinus. 35: male, dorsal view ofidiosoma, 36: male, ventral view ofhysterosoma, 37 : female, dorsal view of idiosoma. 11. Neopteronyssus yungipicinus (MIRONOV, 1987) (Figs 35-37)

PteronyssusyungipicinusMm..ONOV, 1987: 531, fig. 2, 1989: 108, figs 2, 6, 4, 5, 5, 4. Neopteronyssus yungipicinus: MIRONOV, 2002: 143.

Material examined. Male holotype, I male and 2 female paratypes (ZISP 934) from Dendrocopos kizuki (TEMMINCK), Russia, Primorye, Sudzukhin reserve, 24. V.l944, M VOLKOVA. This species is known only from the type host, Dendrocopos kizuki, in the type locality {MIRONOV, 1987). Neopteronyssus from woodpeckers of the Old World 67

Figs 38-41. Neopteronyssus campetherinus. male. 38: dorsal view ofidiosoma, 39: ventral view ofidiosoma, 40: tarsus m. dorsal view. 41 :tarsus IV. dorsal view.

Species group pycnospilus

Male. Opisthosoma with wide and truncated lobes, terminal cleft completely occupied by interlobar membrane (Figs 38, 42-45), transventral sclerite present, epiandrium fused with its posterior end (Figs 39, 46, 48), tarsus IV without dorsobasal teeth (Fig. 41 ). Female. Dorsal hysteronotal shields variously shaped, most commonly represented by three large pieces: the anterior hysteronotal plate a variously­ shaped polygon, and pair of opisthosomal plates commonly extending to • 68 S.V.MIRONOV posterior end of the body (Figs 28, 54). Dorsal plates of hysterosoma may be free or connected (Figs 28, 53); anterior hysteronotal plate absent in N. malaconotus (Fig. 52). Remark. This group includes 5 species.

12. Neopteronyssus campetherinus MIRONOV sp. n. (Figs 28, 38-41)

Pteronyssus pycnospilus (variete): GAUD, 199I: 117, 123, fig. 2.

Type material. Male holotype, I male and 3 female paratypes (UGA 937I, AMNH 549634) from Campethera c. caroli ~E), Angola, Canhocia, 9° IS' S, 14 o 35' E, 4.XII.1903, W. J. ANSORGE. Holotype, I female paratype-MZUM, I male and 2 female paratypes- ZISP. Description Male (holotype). Length ofidiosoma 4I5, width ofidiosoma 245 (idiosomal size in I paratype 406 x 235). Prodorsal shield triangular in general shape, covering median part of prodorsum, separated by transverse row of soft tegument at level of scapular setae into anterior and posterior plates (Fig. 3 8); the posterior plate with acute posterior angles, total length of the shield along median line II2, greatest width of posterior fragment II3, anterior plate 77 in length, 55 in width. Setae se separated by 68. Length of hysterosoma 240. Setae c2 hair-like, short, about 25 in length. Subhumeral setae c3 lanceolate, 29 in length. Hysteronotal shield: anterior margin straight, surface without ornamentation, greatest length 204, width at anterior margin 100. Dorsal setae el slightly posterior to openings gl. Opisthosoma rectangular in general shape, with truncated lobes carrying setae h2 and h3 on terminal margin. Terminal cleft triangular, 35 in length 42 in width; fused with long supranal concavity extending by anterior end to level of cupules im; total length of the cleft and supranal concavity I 08. Interlobar membrane without incision, occupying all space of terminal cleft . Distance between setae and openings: c2-d2 95, d2-e2 93, d2-gl 48-5I, el-gl 7-2-IO, h2-h2 78. Setae h3 lanceolate in basal part. Transventral sclerite present, anterior ends of epimerites ma with long and acute sclerotized extensions (Fig. 39). Epiandrium fused with posterior margin of transventral sclerite, tips of epiandrium short, not extending to genital apparatus. Setae 3b slightly anterior to 3a. Genital apparatus 26 in length, 13 in width, tip of aedeagus acute, curved backward. Genital setae g at midlevel of genital apparatus. Adanal membranes wide. Adanal shield represented by longitudinal sclerite situated at anterior end of anal slit. Tarsus m 62 in length; seta r 2.5-3 times longer than this segment, all ventral setae of this tarsus hair­ like (Fig. 40). Tarsus IV without dorsobasal teeth (Fig. 41 ). Female (paratype). Length of idiosoma 495, width of idiosoma 245 (idiosomal size in 2 other paratype 485-490 x 232-240). Prodorsal shield as in the male, but posterior plate without extending posterior angles and with widely rounded posterior margin (Fig. 28); total length of the shield along median line 130, width of posterior plate, anterior plate 71 in length, 58 in Neopteronyssus from woodpeckers of the Old World 69

"'

200 t.~m

45

Figs. 42-45. Males of the genus Neopteronyssus, dorsal view of idiosoma. 42 : Neopteronyssus dicronotus, 43 : N. malaconotus, 44 : N. pachyphrichus, 45 : N. pycnospilus. (Fig. 42 after: GAUD, 1991, modified). width. Setae se separated by 72. Length of hysterosoma 325. Setae c2 hair­ like, 24 in length. Subhumeral setae c3 lanceolate, 3 5 in length. Arrangement of hysteronotal shields: anterior hysteronotal plate as a polygon with deep incision on posterior margin and pair of opisthosomal plates of irregular form (Fig. 28); anterior plate is eonnected by narrow bulks with anterior ends of opisthosomal plates; lateral margins of all plates crossed by oblique striations; total length of hysteronotal shield complex 263, greatest width of anterior part 70 S.V.MlRONOV

501Jm

Figs 46-50. Males of the genus Neopteronyssus. 46: Neopteronyssus malaconotus, ventral view of hysterosoma, 47: same, tarsus Ill, dorsal view, 48 :N. pachyphrichus, ventral view of hysterosoma, 49 : same, tarsus ID, SO: N. pycnospilus, ventral view ofhysterosoma 128. Setae e2 and openings gl on opisthosomal plates. Terminal copulatory extension as small hemisphere situated asymmetrically between setae h3, about 7 in length. Distance between setae and seta rows: c2-d2 128, d2-e2 113, e2-fl 46, el-e2 28, h2-h2 118. Epigynium rectangular, 50 in length, 93 in width.

Differential diagnosis. The males of Neopteronyssus campetherinus are very similar to N. pycnospilus (Figs 38, 45), while females of these species are Neopteronyssus from woodpeckers ofthe Old World 71 easily tlistinguished by the structure of hysteronotal shields. In the females N. campetherinus, the anterior hysteronotal plate is connected with the opistho­ somal plates (Fig. 28), and in. N. pycnospilus these plates are clearly separated (Fig. 54). The males of N. campetherinus, may be reliably distinguished from N. pycnospilus by larger length of idiosoma, (405-415 J.Lm in the former versus 369-380 Jlm in the latter).

Etymology. The species name derives from the generic name of the type host.

Remark. This species was formerly figured by GAUD (1991), but treated as a large-sized variety ofPteronyssus pycnospilus GAUD et MOUCHET, 1959.

13. Neopteronyssus dicronotus (GAUD, 1991) (Figs 42, 51)

Pteronyssus dicronotus GAUD, 1991: 116, figs 3a, 4a, Sa. Neopteronyssus dicronotus: MlR.ONOV, 2002: 143

This species is known only from two males and one female collected from Campethera tullbergi (SJOSTEDT) (=C. taeniolaema) in Zair (GAUD, 1991). The material was not available in the present study.

14. Neopteronyssus malaconotus (GAUD, 1991) (Figs 43, 46, 47, 52)

Pteronyssus malaconotus GAUD, 1991: 118, figs 3b, 4b, Sb. Neopteronyssus malaconotus : M!R.ONOV, 2002: 143

Material examined. 2 males, 4 females (NU 3702) from Campethera notata (LICI-ITENSTEIN), Mozambique, Lake Fuhwa, 23 . vn.I953, coli. unknown. This species is associated with different species of the genus Campethera GRAY in Africa. It was recorded on Campethera caroli {MALHERBE) (type host) in Zaire, C. abingoni (SMITII) in Zaire and Mozambique, and C. notata in Mozambique (GAUD, 1991).

15. Neopteronyssus pachyphrichus (Gaud, 1991) (Figs 44, 48, 49, 53)

Pteronyssus pachyphrichus GAUD, 1991: 120, fig. 3c, 4c, Se. Neopteronyssus pachyphrichus: M!R.ONOV, 2002: 143.

Material examined. I male, 1 female (UGA 9369, AMNH 549608) from Campethera c. caroli, Abanga River and Ogooue River, Gabon, 22.X.I907, W.J. A.NsoRGE. This species is known from C. caroli (type host) in Gabon and Campethera nivosa (SWAINSON) in Zaire (GAUD, 1991). 72 S.V.MlRONOV

200 ~o~m

51 52

Figs 51-54. Females of the genus Neopteronyssus, dorsal view of idiosoma 51 : Neopteronyssus dicronotus, 52 : N. malaconotus, 53 : N. pachyphrichus, 54 : N. pycnospilus. (Fig. 51 after: GAUD, 1991, modified).

16. Neopteronyssus pycnospilus (GAUD & MOUCHET, 1959) (Figs 45, 50, 54) Pteronyssus pycnospilus GAUD & MOUCHET, 1959: 525, figs 10F, 11F; GAUD & TILL, 1961: 278; FACCINI & ATYEO, 1981: 57; GAUD, 1991: 121, figs 3d, 0 Neopteronyssus from woodpeckers of the Old World 73

4d. Neopteronyssuspycnospilus: MlR.ONOV, 2002: 143. Material examined. 1 male, 2 females (UGA 9365, AMNH 549494) from Campethera cailliautii permista (REICHENOW), Gabon. Ogooue River, Lawbarene, 22.X 1907. W.J. ANSORGE. This species is known only from the type host, Campethera cailliautii permista, in Cameroon, Gabon, and Zaire (GAUD, 1991).

Discussion In spite of the perfonned study, the fauna of pteronyssid mite~· associated with the woodpeckers Picidae in the Old World is still explored far incompletely, and in the New World it is practically unexplored. Feather mite species of the genus Neopteronyssus are known now from 25 species of 8 genera of woodpeckers (Table) that is less than one third of picid species occurring in the Old World {HOWARD & MOORE, 1991). Nevertheless, accumulated data on the genus Neopteronyssus allow drawing out some preliminary conclusions in regard to host associations of its species, specificity range, and distribution. In general lines it is possible to state now, that the genus Neopteronyssus is the most numerous and widely distributed pteronyssid genus occurring on the woodpeckers ofthe·Old World and in North America. Other species-rich genera known from woodpeckers, such as Anephippius GAUD, 1981, Conomerus GAUD, 1981, and Zygepigynia GAUD & CORPUZ­ RARos, 1985, are associated with woodpecker genera having more restricted geographical range. Thus, two fanner genera are distributed only on African · genera of woodpeckers (GAUD, 1990a, b; 1991), and the latter genus is specific to the genus Chrysoco/aptes BL Y1H in the South-east Asia (GAUD & CORPUZ-RAROS, 1985). Within the genus Neopteronyssus, the pycnospilus species group is most clearly separated from other representatives of the genus both in its morphological features (see diagnosis above) and host associations. Representatives of this group are restricted to the genus Campethera distributed exclusively in Africa. Based on the host association data (Table), it is possible to note, that certain species of Campethera, for example C. caro/i, are the hosts of several mite species of the pycnospi/us group. It is reasonable to suggest that these mite species could coexist on the same bird individual. Similar coexisting of closely related mite species was observed in other pteronyssid genera. Thus, some European species of green woodpeckers, Picus, may carry simultaneously two species of the genus Pteronyssus, and some species of the genus Chrysocolaptes BL YTH in South-east Asia may carry even four species of Zygepigynia GAUD & CoRPuz-RARos, 1985. At the same time it is necessary to note, that some species of the pycnospilus group are distributed on several species of the genus Campethera. It is possible to suggest, either that most species of this group were fanned on the ancestor of the bird genus Campethera, or that the process of natural contamination and exchanging by pteronyssid species between different Campetera ·species took place in some historical.past. 74 S.V. MlRONOV

• Table. Host associations and distribution ofmites of the genus Neopteronyssus.

Mite spedes and spedes Hostspedes Cowttryor References group geographical region picinus goap N.fainisp. n. Dinopium raf/lesi Malaya present study N. gecinuU sp. n. Geclnulus grantia SE China present study N.javanicus sp. n. Dryocopus j.javensis Borneo Island, present study Malaysia N. martius sp. n. Dryocopus martius Kazakhstan present study N. muelleripicinus sp. n. MueUeripicus p. Palavan Island pt;esent study pulverulentus (( M.f.folvus Celebes Island present study N. piclnus (KOCH. 1841) Dryocopus martius Europe, Russia: CEJoor & SCHUMILO, 1973, Primcnye FACCINI & ATYEO, 1981, MIRONOV, 1989 N. vittatus sp. n. Picus vittatus Indonesia, Malaya present study pidgroup N. biUneatus sp. n. Dendrocopos can/capillus Thailand present study N. elongatus {BucHHOLZ) Dendrocopos minor Europe CBRNY & SCHUMILO, 1973; FACCINI&ATYE0,1981; M!RONOV,1989 N. pie/ (SCOPOLI, 1763) D.leucotos Europe CBRNY&SCHUMIL0,1973; M!RONOV,l989 (( D. major Europe CEJoor & SCHUMILO, 1973; FACCINI&ATYE0,1981; MIRONOV, 1989 (( D. medius Europe CEJoor, 1969; MIRONOV, 1989 (( D. syrlacus Europe CBRNY & SCHUMILO, 1973 (( D. villoms Canada: Ontario TYRRELL, 1882, present study

(( Picoides arcticus Canada: New present study Brunswick

(( P. tridactyUs Europe CBRNY & SCHUMJLO, 1973

(( Picuscanus Europe CBRNY & SCHUMILO, 1973; MIRONOV, 1989

(( P. viridis Europe, Azerbajdzhan MIRONOV,1989 N. yunglplcinus {MIRONOV, D. kizuki Russia: Primorye MIRONOV,1987 1987) pycnospilus group N. campetherae sp. n. Campethera c. caroU Angola present study N. dicronotus (GAUD, 1991) Campethera tullbergi Zaire OAUD, 1991 N. malaconotus (GAUD, C. carol/ Zaire OAUD, 1991 1991) (( C. abingoni Zaire, Mozambique, OAUD, 1991 Transvaal

(( C. notata Mozambique OAUD, 1991 N. pachyphrichus (GAUD, C. carol/ Gabon 0AUD, 1991 1991) (( C. nivosa Zaire OAUD, 1991 N. pycnospllus (OAUD et C. cailllautii permlsta Cameroon, Gabon, GAUD & MouCHET, 1959, MouCHET, 1959) Zaire OAUD,.. 1991 Neopteronyssus from woodpeckers of the Old World 75

The &epresentatives of the picinus group are mainly monoxenous or narrowly oligoxenous (Table). It is most clear visible with an example of species associated with black woodpeckers of the genus Dryocopus. Each examined species of the genus has one specific mite species. It is worth to note, that in different parts of the range of the black woodpecker, Dryocopus martius, this bird has different species of Neopteronyssus. Thus, in Europe and in the Russian Far East (Primorye) it normally bears N. picinus (CERNY & SCHUMILO, 1973; MIR.ONOV, 1989), while in Kazakhstan, this woodpecker carries another species, N. martius. It may mean that the population of D. martius in the area of Kazakhstan was formerly isolated from the re~t part of the host's population, and that had lead to formation of a separate mite species within this local population. Within the pici group, three species, N. hilineatus, N. elongates, and N. yungipicinus, are monoxenous. In a contrast to them, the fourth species, N. pici, is widely distributed on almost medium-sized European, North Asian and North American woodpeckers of the genera Dendrocopos, Picoides, and Picus. This is obviously the result of numerous cases of contamination between these hosts, which probably took place many times in the past. Apparently, the most suitable places for such contaminations are tree holes used by woodpeckers for nesting and hiding, and where accidentally lost mites can probably survive for certain time without a host. At present, it is difficult to guess, what genus of woodpeckers was a primary host of N. pici, green woodpeckes of the genus Picus, or spotted woodpeckers of the genera Dendrocopos and Picoides. This question could be probably solved by subsequent . investigations of the pteronyssid mite fauna form the Picidae distributed in Central and South-east Asia.

Acknowledgements

The author wishes to thank Prof. W.T. ATYEo (The University of Georgia, Athens, USA) for loaning material used in the present study. The study was made in the frame of projects supported by the Russian Foundation for Basic Researches (Grants No 00-04-49323, 00-04-48885, 02-04-63000).

References

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