Acarina 20 (2): 67–107 © Acarina 2012

A review of PROSTIGMATA (Acariformes: TROMBIDIFORMES) permanently associated with birds M. Skoracki1, S. A. Zabludovskaya2 and A. V. Bochkov3, 4 *

1Department of Animal Morphology, Faculty of Biology, Adam Mickiewicz University, Umultowska 89, 61–614 Poznan, Poland 2Schmalghausen Institute of Zoology, National Academy of Ukraine, B. Khmelnitzky str. 15, Kiev, Ukraine 3Zoological Institute of the Russian Academy of Sciences, Universitetskaya Emb. 1, 199034 Saint Pe- tersburg, Russia; e-mail: [email protected] 4Museum of Zoology, University of Michigan, 1109 Geddes Ave., Ann Arbor, Michigan 48109, USA *Corresponding author

ABSTRACT: Prostigmatan mites (Acariformes: Prostigmata) permanently parasitizing birds are reviewed at the familial level. Keys to all genera and lists of included species along with verified host records are provided. This ecological group includes about 500 species in 5 families: Cheyletidae (Cheletosomatini, Ornithocheyletiini, and Metacheyletiini), Syringophilidae, Harpirhynchidae (Harpirhynchinae and Harpypalpinae), Cloacaridae (Pneumophaginae), and Ereynetidae (Speleognathinae). The original and reference data on the external morphology, phylogeny, systematics and the host-parasite relationships of these mites are sum- marized.

KEY WORDS: Prostigmata, birds, mites, symbionts, systematics, host-parasite relationships

INTRODUCTION Prostigmatic mites (Acari: Acariformes: Pros- tion of host phylogeny and in deciphering of the tigmata) permanently associated with birds are po- host biogeography. Such investigations were un- tentially very species-rich groups supposedly in- dertaken on various groups of symbionts, includ- cluding about 5000 species (Kethley and Johnston ing acariform mites, and often showed a high level 1975; Moss 1979), but nearly 10 % (about 500 of phylogenetic congruence between hosts and species) of these mites are recognized to date. symbionts (Klassen 1992). Although some investi- These mites occupy a great diversity of microhab- gations conducted on astigmatan mites did not re- itats on the host body: skin, quills, intracutaneous veal phylogenetically congruent pattern, they pro- layers, and respiratory tract, and are very interest- vided valuable results for the host biogeography ing in the global evolutionary sense as well as in (Bochkov and OConnor 2005). At the same time, several parasitological aspects, including host- as a taxonomically poorly studied group, the pros- parasite relationships. tigmatans permanently associated with birds have This ecological group includes mites of five never been involved in this kind of investigations. families belonging to three phylogenetically dis- The data concerning systematics, phylogeny tant superfamilies: Cheyletidae, Harpirhynchidae, and host-parasite relationships of acariform mites and Syringophilidae of the superfamily Cheyle- permanently associated with mammals and birds toidea, Cloacaridae (Cloacaroidea), and Ereyneti- were recently revised (Proctor 2003; Bochkov dae (Tydeoidea) (Mironov and Bochkov 2009). 2009, 2010), but bird-associated prostigmatans The absence of taxonomic revisions and identifi- were not considered in these works. cation keys for most bird-associated prostigmatans In this paper we provide the family level re- seriously impedes their phylogenetic and ecologi- view of bird-associated prostigmatans, keys to all cal investigations. genera, and check-lists of their species along with Most prostigmatic inhabitants of bird are true verified host records. The data on phylogeny, biol- parasites feeding on live host tissues, but the ma- ogy and host-parasite associations of these mites jority of cheyletids living in feather quills are are briefly discussed. predators feeding on other quill-inhabiting mites and thus should be considered as commensals or MATERIAL AND METHODS even mutualists. The materials used in this work were obtained Taking into consideration the high specificity from acarological collections housed in various of permanently parasitizing prostigmatans to avian museums. Data on bird-associated prostigmatan hosts and their non-random distribution on host families including numbers of recognized species taxa (Fain 1994a; Bochkov 2009; Skoracki 2011), and genera, hosts, microhabitats, and distribution data about these mites could be used for the valida- are summarized in Table 1. Idiosomal and leg seta-

67 M. Skoracki, S.A. Zabludovskaya, A.V. Bochkov

A B Fig. 1. Scheme of eleutherengone gnathosoma: A — in dorsal view; B — in ventral view (from Bochkov 2009, with minor modifications). tion of these families is represented in Tables 2–5. Prostigmata includes parasitic forms. Most of the examined materials are deposited Parasites permanently associated with birds in the following collections: Adam Mickiewicz belong to two phylogenetically distant parvorders, University (Poznan, Poland); Royal Belgian Insti- Eupodina (Ereynetidae) and Eleutherengona tute of Natural Sciences (Brussels, Belgium); (Cheyletidae, Cloacaridae, Harpirhynchidae, and Royal Museum for Central Africa (Tervuren, Bel- Syringophilidae). gium); Zoological Institute, Russian Academy of The only possible paleontological record of Sciences (Saint-Petersburg, Russia). these bird-associated mites is the finding of eggs In the taxonomic part, the gnathosomal seta- on a feather from the Lower Cretaceous period tion follows Grandjean (1946), the leg and idio- (northeast Brazil) (Martill and Davis 1998). Au- somal setation follows Grandjean (1939, 1944). thors believed that these eggs were laid by feather The universal setal nomenclature of idiosoma pro- mites (Astigmata: Psoroptidia). These spherical posed by Grandjean (1939) was adopted for Pros- eggs are more similar to those of Cheyletoidea, tigmata by Kethley (1990). Schemes of external than to the strongly elongated eggs of feather mites mite morphology are provided in Figs. 1 and 2. (Bochkov 2008). Proctor (2003), however, sup- The bird systematics follows Clements et al. posed that this feather dropped to water and these (2011). Data about all described species of bird- eggs were laid by Ostracoda. associated prostigmatans are summarized in the Appendix (deposited at Acarina web-site: http:// Key to families of Prostigmata — permanent parasites of birds insects.ummz.lsa.umich.edu/acarina/). 1. Subcapitulum of gnathosoma distinctly devel- SYSTEMATICS AND PHYLOGENY oped (Fig. 1). Palps present. At least some of idio- Suborder Trombidiformes Reuter, 1909 somal and leg setae filiform ...... 2 Infraorder Prostigmata Kramer, 1887 — Subcapitulum of gnathosoma strongly reduced According to the system of acariform mites or absent. Palps absent. Idiosomal and leg setae proposed by Mironov and Bochkov (2008), the represented by alveoli or fleshy-like projections ... suborder Trombidiformes is separated into four Cloacaridae Camin, Moss, Oliver et Singer, 1967 infraorders, Sphaerolichida, Bimichaelida, Oeh- (Pneumophagus Fain et Smiley, 1989) (Fig. 10) serichestida, and Prostigmata. Only the infraorder 2. Propodonotal setae si represented by unmodi-

68 A review of Prostigmata permanently associated with birds

A B

C

D

Fig. 2. Scheme of eleutherengone mite: A — idiosoma in dorsal view; B — same in ventral view; C — setation of legs I–IV; D — tarsus I (from Bochkov 2009, with minor modifications).

69 M. Skoracki, S.A. Zabludovskaya, A.V. Bochkov fied simple setae. Genital papillae absent. Tibia of is located on the ventral side of the trochanter-fe- legs I with or without solenidion φ ...... 3 mur-genu, the absence of the eupathidia on the — Propodonotal setae si represented by bothridia palpal tarsus, the absence of the separate hyster- (Fig. 11A). Genital papillae present. Tibia of legs I onotal shield, the completely fused genital and with complicate ereynetal organ (sensory complex, anal openings in females, the absence of the Fig. 12A) ...... Ereynetidae (Speleognathinae) aggenital setae in both sexes, the absence of setae 3. Distal part of hypostome fused with subcapitu- ft, vsI, solenidion φI, σI, coxal setae 1b (reversed lum. Peritremes situated on subcapitulum, not lin- in Ophioptinae), 3b, and 4c (reversed in Harpypal- ear. Palpal trochanter, femur and genu not fused. pinae). Setae scx very small, covered by lateral extensions Family Cheyletidae Leach, 1815 of propodonotum. Tibia and genu I with 1 solen- idion each ...... 4 Figs. 3–5 — Distal part of hypostome not fused with subca- Type genus: Cheyletus Latreille, 1796 pitulum. Peritremes situated at base of subcapitu- Diagnosis. Gnathosoma moderately or dis- lum, linear. Palpal trochanter, femur and genu tinctly developed (1/5–1/3 of idisosomal length). fused to each other. Setae scx distinctly developed, Subcapitulum bearing setae elc. p, n, ao1 and ao2. situated dorsally. Tibia and genu I without solen- Peritremes situated on rostral part of stylophore, idion ...... Harpirhynchidae distinctly segmented, generally M-shaped. Distal Dubinin, 1957 (Figs. 8, 9) part of hypostome fused with stylophore. Hypos- 4. Subcapitulum not deeply submerged in idioso- tomal lips present. Palps linear, consisting of 5 ma, subcapitular apodeme absent. Claw of palpal segments, with various projections in some para- tibia present. Palpal tibia and tarsus separated. Se- sitic forms. Palpal tibia with distinct claw. Palpal tae 4a present. Setae a’I absent ...... tarsus strongly reduced. Palpal setation: femur ...... Cheyletidae Leach, 1815 (Figs. 3–5) with setae d, v’, v”, genu — d, l”, tibia — d, lT, l’, — Subcapitulum deeply submerged in idiosoma, and tarsus with eupathidia acm, sul, ul’, ul”, and subcapitular apodeme distinctly developed. Claw solenidion ω. Eupathidia acm and sul” comb-like of palpal tibia absent. Palpal tibia and tarsus fused. in most predaceous mites and smooth in most par- Setae 4a absent. Setae a’I present ...... asites. Idiosoma generally rhomb-like or ovate in .... Syringophilidae Lavoipierre, 1953 (Figs. 6, 7) outlline. Bases of trochanters inserted on ventral side of idiosoma. In Cheyletiellini, idiosoma bear- Parvorder Eleutherengona Oudemans, 1909 ing 1 pair of distinctly developed lateral scapular Superfamily Cheyletoidea Leach, 1815 lobes. Eyes absent; or 1 pair of lens-like eyes pres- The supefamily Cheyletoidea includes cur- ent on propodonotum in some free-living preda- rently five families, Cheyletidae, Syringophilidae, tors. Propodonotal shield present, distinctly devel- Harpirhynchidae, Psorergatidae, and Demodici- oped. Hysteronotal and pygidial shields present or dae; mites of the three first families are associated absent. Ventral side of idiosoma generally devoid with birds (Bochkov 2009). The phylogeny of of shields but in males of some predatory forms these mites was recently reconstructed by Bochk- weakly sclerotized pseudosternal shield present. ov (2002, 2008). There are two main phylogenetic Opisthosoma moderately developed. In females, lineages in this superfamily. Lineage I includes the anal and genital-ovipore openings situated ven- families Cheyletidae and Syringophilidae, where- trally, close to posterior end of body or terminally, as lineage II is represented by three families, adnate and covered by pair of folds. In males, ae- Harpirhynchidae (Psorergatidae–Demodicidae). deagus tube-like or comma-like and pointed api- The monophyly of lineage I is supported by the cally, genital and anal openings fused (or anal following synapomorphies: the peritremes are sit- opening absent). In males of free-living forms, uated on the rostral part of the stylophore, distinct- genital opening situated terminally or ventro ter- ly segmented, and arch-like or M-shaped; the dis- minally. In parasitic males, genital opening dor- tal part of the hypostomal apex is fused with the sally displaced, and in some bird parasites, it lo- stylophore, and the hypostomal lips are present. cated in anterior part of idiosoma. Distance Lineage II could be characterized by the fol- between coxal fields II and III distinctly shorter lowing synapomorphies: the presence of the than idiosomal width (excluding tribe Bakini). In strongly sclerotized pharyngeal bulb, the absence most taxa, 3 pairs of cupules present: im, ip, and of adoral setae ao1 and ao2, the palpal tibia-tarsus ih. In some parasites, these cupules absent. Idio-

70 A review of Prostigmata permanently associated with birds

Fig. 4. Ornithocheyletia chirovi (Cheyletidae: Ornithochey- letiini), female in dorsal view.

(tc), (p), (u), vs, ω1 (only in males); IV, (tc), (p), (u), vs, ω1 (only in males); tibiae I–IV — I, d, (l), Fig. 3. Metacheletoides numidae (Cheyletidae: Cheletosoma- tini), male in dorsal view (from Bochkov and Skoracki 2012 (v), φ; II, d, l”, (v), φ; III, d, l”, (v) + φ (only in with minor modifications). males); IV, d, l”, (v) + φ (only in males); genua I–IV — I, d, l’, σ; II–IV, d, l’; femora I–IV — d, v; somal setation (maximum set): scx, vi, ve, si, se, trochanters I–IV — I, II, IV, v; III, l, v; coxae I–IV c1, c2, d1, d2, e1, e2, f1, f2, h1, h2, h3, ps1, ps2, — I, 1a, 1b, 1c; II, 2c; III, 3a, 3b, 3c; IV, 4a, 4b, ps3, ag1, ag2, ag3, g1, g2. Setae scx covered dor- 4c. Solenidion ω1 II situated ventrally. Larva, pro- sally by extending lateral margins of propodono- to- and tritonymphs present; deutonymph absent. tum. In males, 2 pairs of aggenital setae, 2 pairs of Male moulting from protonymph. genital setae, and 3 pairs of pseudoanal setae pres- Detailed historical reviews of the cheyletid ent. In some taxa idiosomal chaetome neotrichous systematics were provided by Volgin (1969) and and many neotrichous dorsal setae strongly modi- Bochkov (2009). Therefore we discuss it here very fied. In most taxa, legs slender, consisting of 5 ar- briefly. The excellent taxonomic revision of the ticulated segments, and their tarsi having weakly family Cheyletidae prepared by Volgin (1969) developed pretarsi bearing 2 smooth lateral claws could be considered as a base of the modern chey- and empodium with tenent hairs. Pretarsi separat- letid systematics. According to his system, the ed from respective tarsi by dorsal knob. Leg length family consisted of ten tribes and the two subfam- usually 60–70% of idiosomal length. In Metachey- ilies Cheyletinae and Cheyletiellinae. The revision letia, legs IV lost or primordial. In free-living of the family Cheyletidae prepared by Summers forms, leg coxae distinctly bordered, and in some and Price (1970) appeared almost simultaneously cases well sclerotized. In many parasitic mites, leg with the Volgin’s monograph but was not so com- coxae represented exclusively by coxal apodemes. prehensive. The latter work did not introduce any Projections of leg segments present only in some principal improvements to the cheyletid system parasitic species. Maximum set of leg setae (paired and the tribes and subfamilies established by Vol- setae are in parentheses): tarsi I–IV — I, ft, (tc), gin were unceremoniously ignored by these au- a”, (p), (u), vs, ω1; II, (tc), (p), (u), vs, ω1; III, thors.

71 M. Skoracki, S.A. Zabludovskaya, A.V. Bochkov

A B

Fig. 5. Metacheyletia ngaii (Cheyletidae: Metacheyletiini), female: A — in dorsal view, B — in ventral view (from Bochkov and Skoracki 2011 with minor modifications).

Fain et al. (1997) and Gerson et al. (1999)���������� re- Key to bird-associated genera of Cheyletidae vised the family at the generic level and provided a list of all described species. 1. Legs IV present, normally developed. Idiosoma The most important synapomorphies of Chey- rhomb-like in outline ...... 2 letidae are the ventral situation of solenidion on — Legs IV absent or primordial. Idiosoma ovate tarsi II, male moulting from protonymph, and the in outline ...... (Metacheyletiini Fain, 1980) absence of setae a’ of tarsus I (Bochkov 2008). Metacheyletia Fain, 1972 The attempt of the cladistic analysis of the 2. Eupathidium sul” comb-like or serrate. Palpal family Cheyletidae was undertaken by Bochkov claw widely opened, slightly curved laterally, with and Fain (2001). According to the result of this basal angle(s). At least some idiosomal setae of analysis with some corrections (Bochkov 2004), females ultralong (excluding normally always the family Cheyletidae includes 14 tribes most of long c2 and h2). Male genital opening situated ter- which were established by Volgin (1969): Ac- minally. Tarsi of legs I much longer than wide aropsellini, Bakini, Cheletogenini, Cheletosoma- (Cheletosomatini Volgin, 1969) ...... 3 tini, Chelonotini, Cheyletiini, Cheyletiellini, — Eupathidium sul” smooth. Palpal claw strongly “Cheyletini”, Cheletomorphini, Metacheyletiini, curved ventrally and without basal angles. Idiosom- Niheliini, Ornithocheyletiini, Teinocheylini, and a al setae of females moderately long (excluding nor- unnamed tribe represented by the genus Cau- mally always long c2 and h2). Male genital opening dacheles. The relationships between these tribes situated dorsally. Tarsi of legs I slightly longer than remain unresolved. The bird parasites and nidi- wide (Ornithocheyletiini Volgin, 1969) ...... 8 colous mites belong to the tribes Cheletosomatini, 3. Eupathidium sul” smooth or slightly serrate. Metacheyletiini, and Ornithocheyletiini. Setae c3 present ...... 5

72 A review of Prostigmata permanently associated with birds

— Eupathidium sul” comb-like. Setae c3 absent . Diagnosis. Gnathosoma moderately develdevel-- ...... 4 oped. Subcapitulum deeply inserted into idiosoma, 4. Coxal fields III and IV contiguous. Setae c1 bearing setae elcp, n, ao1 and ao2; subcapitular situated on propodonotal shield ...... apodeme distinctly developed. Peritremes situated ...... Cheletopsis Oudemans, 1904 on rostral part of stylophore, distinctly segmented, — Coxal fields III and IV distinctly separated. Se- generally M-shaped. Hypostomal lobes fused with tae c1 situated off propodonotal shield ...... stylophore. Hypostomal lips present. In many taxa, ...... Eucheletopsis Volgin, 1969 female hypostome bearing 1 pair of hyaline protu- 5. Setae c4 absent. Female hysteronotal shield ab- berances. Palps linear, consisting of 4 segments sent or situated on posterior part of opisthosoma. (tibia and tarsus fused). Palpal tibiotarsus without Setae e1 and f1 absent ...... 6 claw. Palpal setation: femur with setae d, v’,v”, — Setae c4 present. Female hysteronotal shield genu — d, l”, tibiotarsus — d, lT, l’, eupathidia present, situated between levels of setal bases d2 acm, sul, ul’, ul”, and solenidion ω. Idiosoma and e2. Setae e1 and f1 present ...... strongly elongated. Bases of trochanters inserted on ...... Picocheyletus Bochkov et OConnor, 2003 ventral side of idiosoma. Eyes absent. Propodono- 6. Eupathidium sul” serrate. Palpal claw with 1 tal shield present. Hysteronotal and pygidial shields basal angle. Tarsal claws on all legs equal or sub- present or absent. Ventral side of idiosoma gener- equal in size ...... 7 ally devoid of shields. Opisthosoma distinctly de- — Eupathidium sul” smooth. Palpal claw with veloped. Female anal and genital-ovipore openings several basal angles. Tarsal claws on legs I much situated ventrally close to posterior end of body or smaller than claws of tarsi III–IV ...... terminally, adnate, and covered by pair of folds. In ...... Metacheletoides Fain, 1972 males, aedeagus tube-like and pointed apically, 7. In female, small hysteronotal shield present and genital and anal openings fused (or anal opening setae h1 short, much shorter than c2 ...... absent) and situated dorsally. Distance between ...... Cheletosoma Oudemans, 1905 coxal fields II and III subequal or about twice as — In female, hysteronotal shield absent and setae long as idiosomal width. Idiosomal cupules absent. h1 long, subequal to c2 ...... Idiosomal setation (maximum set): scx, vi, ve, si, se, ...... Cheletoides Oudemans, 1904 c1, c2, d1, d2, e2, f1, f2, h1, h2, ps1, ps2, ag1, ag2, 8. Genua III and IV with 1 and without setae, re- ag3, g1, g2. Setae scx covered dorsally by extend- spectively ...... 10 ing lateral margins of propodonotum. In males, 2–3 — Genua III and IV with 2 and with 1–2 setae, pairs of aggenital setae, 2 pairs of genital setae, and respectively ...... 9 2 pairs of pseudoanal setae present. In some taxa, 9. Hysteronotal shield present. Trochanters III and aggenital setae neotrichous. Legs slender, consist- IV without setae. Genu IV with 1 seta ...... ing of 5 articulated segments; their tarsi having ...... Ornithocheyla Volgin, 1964 weakly developed pretarsi bearing 2 smooth lateral — Hysteronotal shield absent. Trochanters III and claws and empodium with tenet hairs. Maximum IV with 2 and 1 setae, respectively. Genu IV with set of leg setae (paired setae are in parentheses): 2 setae ...... Bakericheyla Volgin, 1966 tarsi I–IV — I, ft, (tc), (a), (p), (u), vs, ω1; II, (tc), 10. Palpal femora without ventral processes. (p), (u), vs, ω1; III, (tc), (p), (u), IV, (tc), (p), (u); Propodonotal shield very small. Setae c3 present. In tibiae I–IV — I, d, (l), v, φ; II, d, (l), v, φ; III, d, (l); females, 2 pairs of aggenital setae present. Solen- IV, d, (l); genua I–IV — I, d, l’, σ; II, d, l’; III and IV idion φI present. Trochanter IV, femora III and IV l’, femora I–IV — I and II, d, v, III and IV d; tro- with 1 seta each ...... Neocheyletiella Baker, 1949 chanters I–IV, v; III; coxae I–IV — I, 1a, 1b, 1c; II, — Palpal femora with ventral processes. 2c; III, 3a, 3b, 3c; IV, 4b, 4c. Solenidion ω1II situ- Propodonotal shield distinctly developed, cover- ated dorsally. Larva, proto- and tritonymphs pres- ing part of hysteronotum. Setae c3 absent. In fe- ent; deutonymph absent. Male moulting from males, aggenital setae absent. Solenidion φI ab- tritonymph. sent. Trochanter IV, femora III and IV without The detailed discussion of syringophilid sys- setae ...... Apodicheles Fain, 1979 tematics is provided in the monograph by Skoracki Family Syringophilidae Lavoipierre, 1953 (2011). Therefore, only the principal works are considered below. The most important review for Figs. 6–7 syringophilid systematics was prepared by Keth- Type genus: Syringophilus Heller, 1880 ley (1970). In the work following this revision,

73 M. Skoracki, S.A. Zabludovskaya, A.V. Bochkov

A B

C

Fig. 6. Syringophilus bipectinatus (Syringophilidae: Syringophilinae), female: A — in dorsal view; B — in ventral view; C — Torotrogla sp. (Syringophilinae), hypostomal protuberance.

74 A review of Prostigmata permanently associated with birds

A B

Fig. 7. Picobia heeri (Syringophilidae: Picobiinae), female: A — in dorsal view; B — in ventral view.

Johnston and Kethley (1973) proposed the origi- for the monotypic genus Cuculiphilus. This genus nal variant of syringophilid system based on the was subsequently included in the subfamily Pico- result of their phenetic analysis. This analysis con- biinae (Fain et al. 2000). From the beginning of firmed all syringophilid genera established by XXI century, the biodiversity of syringophilids Kethley (1970) and allowed the division of this has been studied mainly by A.V. Bochkov and M. family into two unequal subfamilies, Syringophili- Skoracki (with collaborators). To date, this family nae and Picobiinae. The third syringophilid sub- includes 278 species of 53 genera versus 24 spe- family — Lobatinae was created by Casto (1977) cies of 15 genera in Kethley´s review (1970). The

75 M. Skoracki, S.A. Zabludovskaya, A.V. Bochkov modern comprehensive revision of Palaearctic Sy- — Legs I 1.5 times longer than legs II ...... ringophilidae was recently prepared by Skoracki ...... Castosyringophilus Bochkov et Perez, 2002 (2011). 9. Setae dGII absent. Apodemes I divergent ...... The most distinctive synapomorphies of Sy- ...... Galliphilopsis Skoracki et Sikora 2004 ringophilidae are the subcapitulum deeply inserted — Setae dGII present. Apodemes I parallel .... 10 into the idiosoma; four segmented linear palps, 10. Setae l’RI and l’RII present ...... 11 widely separated coxae I–II and III–IV, and the — Setae l’RI and l’RII absent ...... Aulonastus absence of setae 4a (Bochkov 2008). Kethley, 1970 Any works on the phylogeny of this family 11. Two pairs of pseudanal setae (ps1 and ps2) are absent. present ...... Neoaulonastus Skoracki, 1999 — One pair of pseudanal setae present (ps2 ab- Key to genera of Syringophilidae sent) ...... Krantziaulonastus Skoracki, 2011 1. Tibiotarsus of palps truncate on distal margin. 12. Legs with full complement of setae ...... 29 Prorals setae p’ and p” of legs I–IV with two min- — Some of leg setae absent ...... 13 ute tines, rod-like. Physogastric forms of females 13. Setae dGII present ...... 18 present. Juvenile stages with chaetotaxy reduced — Setae dGII absent ...... 14 to small spinose structures. Body feathers as main 14. Setae vsI present ...... 15 habitat. (Picobiinae Johnston et Kethley, 1973) .... — Setae vsI absent ...... Philoxanthornea ...... 48 Kethley, 1970 — Tibiotarsus of palps rounded on distal margin. 15. Setae l’GIV absent ...... 17 Prorals setae p’ and p” of legs I–IV multiserrate, — Setae l’GIV present ...... 16 fan-like. Physogastric forms of females absent. 16. Two pairs of pseudanal setae present ...... Setae of body and legs in juveniles well devel- ...... Syringophiloidus Kethley, 1970 oped. Flight feathers as main habitat. (Syringo- — One pair of pseudanal setae present (ps2 ab- philinae Lavoipierre, 1953) ...... 2 sent) ...... Betasyringophiloidus Skoracki, 2011 2. Setae vi present ...... 12 17. Apodemes I parallel fused to apodemes II ...... Apodisyringophilus Skoracki et OConnor, 2010 — Setae vi absent ...... 3 — Apodemes I distinctly divergent, not fused to 3. Setae dFII present ...... 9 apodemes II ..... Apodisyringiana Skoracki, 2005 — Setae dFII absent ...... 4 18. Setae dFII present ...... 20 4. Setae dFIII and dFIV absent ...... 5 — Setae dFII absent ...... 19 — Setae dFIII and dFIV present and replaced ven- 19. Setae dTIII and dTIV present. Propodonotal trally ...... Terratosyringophilus shield divided longitudinally ...... Ascetomylla Bochkov et Perez 2002 Kethley, 1970 5. Setae ve situated anterior to level of setae si. — Setae dTIII and dTIV absent. Propodonotal Pocket-like structures absent. Stylophore rounded shield entire ...... Fritschisyringophilus posteriorly ...... 6 Bochkov, Fain et Skoracki, 2004 — Setae ve and si situated at same transverse lev- 20. Setae dTIII present ...... 22 el. Pocket-like structures in anterior part of pro­ — Setae dTIII absent ...... 21 podonotum present. Stylophore constricted poste- 21. Setae vsII present. Apodemes I parallel ...... riorly ...... Psittaciphilus ...... Neoaulobia Fain, Bochkov et Mironov, 2000 Fain, Bochkov et Mironov, 2000 — Setae vsII absent. Apodemes I divergent ...... 6. Hypostomal apex ornamented by 2 pairs of me- ...... Cuculisyringophilus Skoracki, 2008 dian finger-like protuberances. Apodemes I and II 22. Setae vsI present ...... 24 parallel and fused to each other ...... 7 — Setae vsI absent ...... 23 — Hypostomal apex smooth. Apodemes I diver- 23. Apodemes I parallel. Stylophore constricted gent, not fused to apodemes II ...... Meitingsunes posteriorly. Claws strongly recurved ...... Glowska et Skoracki, 2010 ...... Bochkovia Skoracki et OConnor, 2010 7. Setal pattern of propodonotal region arranged — Apodemes I divergent. Stylophore rounded 2–1–2 ...... 8 posteriorly. Claws typically opened ...... — Setal pattern of propodonotal region arranged ...... Paraniglarobia Skoracki, 2011 2–3 ...... Neoperisterophila Skoracki, 2005 24. Setae vsII absent, l’GIV present ...... 25 8. Legs I and II subequal in size ..... Peristerophila — Setae vsII present, l’GIV absent ...... Kethley, 1970 ..... Neosyringophilopsis Skoracki et Sikora, 2005

76 A review of Prostigmata permanently associated with birds

25. Lateral hypostomal teeth absent ...... 26 37. Setae f2 long (longer than c1). Setae si and se — Lateral hypostomal teeth present ...... situated at same level ...... 38 ...... Stibarokris Kethley, 1970 — Setae f2 short (2–3 times shorter than c1). Setae 26. Two pairs of genital setae present (g1, g2). si situated anterior to level of setae se ...... Apodemes I not fused to apodemes II ...... 27 ...... Chenophila Kethley, 1970 — One pair of genital setae present (g2 absent). 38. Claws broadly open ...... Selenonycha Apodemes I fused to apodemes II ...... Kethley, 1970 ...... Procellariisyringophilus Kethley, 1970 — Claws strongly recurved ...... Creagonycha 27. Apodemes I parallel. Stylophore without large Kethley, 1970 tip on posterior margin ...... 28 39. Apodemes I divergent ...... 40 — Apodemes I divergent. Stylophore with large — Apodemes I parallel .... Aulobia Kethley, 1970 tip on posterior margin ...... Ciconichenophilus 40. Apodemes I not fused to apodemes II ...... 42 Skoracki et OConnor, 2010 — Apodemes I fused to apodemes II ...... 41 28. Claws small, with basal angle, typically opened 41. Propodonotal shield divided longitudinally ...... Niglarobia Kethley, 1970 ...... Crotophagisyringophilus Skoracki, 2008 — Claws large, without basal angle, strongly re- — Propodonotal shield entire .... Syringophilopsis curved ...... Phalarophilus Kethley, 1970 Skoracki, Bochkov et OConnor, 2011 42. Stylophore constricted posteriorly ...... 44 29. Two or 3 pairs of aggenital setae present ... 31 — Stylophore rounded posteriorly ...... 43 — Aggenital series with 4–9 pairs of setae ..... 30 43. Dorsal setae of idiosoma ornamented ...... 30. Coxal fields I and II similar in size and shape...... Blaszakia Skoracki et Sikora, 2008 Supernumerary setae vsIII absent. Legs I thicker — Dorsal setae of idiosoma smooth ...... than II–IV ...... Torotrogla Kethley, 1970 ...... Bubophilus Philips et Norton, 1978 — Coxal fields I and II disimilar in size and shape. 44. Apodemes I strongly divergent. Setae se lo- Supernumerary setae vsIII present. Legs I–IV sub- cated anterior to the level of setae c1 ...... 47 equal ...... Trypetoptila Kethley, 1970 — Apodemes I slightly divergent. Setae se located 31. Three pairs of aggenital setae (ag1–3) present posterior to level of setae c1 ...... 45 ...... 34 45. Setae f2 situated close to the bases of setae f1 . — Two pairs of aggenital setae present (setae ag2 ...... 46 absent) ...... 32 — Setae f2 situated distinctly anterior to the level 32. Apodemes I not fused to apodemes II. Claws of setae f1 ...... Charadriphilus Bo- typically opened ...... 33 chkov et Chystiakov, 2001 — Apodemes I fused to apodemes II. Claws 46. Each lateral branch of peritremes reduced to 3 strongly recurved .... Kethleyana Kivganov, 1995 chambers. Setae se and c1 situated at same trans- 33. Hysteronotal shield not fused to pygidial verse level. In males 2 pairs of aggenital setae pres- shield. Coxal fields of legs I and II are similar in ent and setae se situated distinctly anterior to level of size and shape, apodemes of legs III and IV well setae c1 ... Pteroclidisyringophilus Skoracki, 2011 developed ...... Picisyringophilus — Each lateral branch with 7–12 chambers. Setae Skoracki et OConnor, 2010 se situated distinctly posterior to level of setae c1. — Hysteronotal shield fused to pygidial shield. In males 3 pairs of aggenital setae present and se- Coxal fields of legs I and II are dissimilar in size tae se and c1 situated at same transverse level ...... and shape. Apodemes of legs III and IV absent ...... Kalamotrypetes Casto, 1980 ...... Mironovia Chirov et Kravtsova, 1995 47. Setae si and se situated at same level. All ter- 34. Lateral hypostomal teeth absent ...... 35 minal setae long ...... Megasyringophilus — Lateral hypostomal teeth present ...... Fain, Bochkov et Mironov, 2000 ...... Colinophilus Kethley, 1970 — Setae si set distinctly anterior to level of setae 35. Peritremes M-shaped ...... 36 se, setae f1 and f2 long, setae h1 and h2 short ...... — Peritremes U-shaped ...... Syringophilus ...... Tinamiphilopsis Skoracki et Sikora, 2004 Heller, 1880 48. End of hysterosoma rounded, without opistho- 36. Coxal fields I and II dissimilar in size and somal lobes. Movable cheliceral digit edentate .... 49 shape ...... 39 — End of hysterosoma with pair of opisthosomal — Coxal fields I and II similar in size and shape .. lobes. Movable cheliceral digit dentate, each with ...... 37 3 teeth ...... Calamincola (Casto, 1978)

77 M. Skoracki, S.A. Zabludovskaya, A.V. Bochkov

49. Bases of setae vi set anterior to level of setae anal and genital-ovipore openings fused and situ- ve. Apodemes I divergent, without thorn-like scle- ated ventrally close to posterior end of body or rotization in middle part. Genital lobes absent. terminally, covered by pair of folds. In males, ae- Physogastric female campanulliform or bulb- deagus tube-like or comma-like and pointed api- shaped outline ...... 50 cally; genital and anal openings fused (or anal — Bases of setae vi and ve set at same transverse opening absent), dorsally displaced. Distance be- level. Apodemes I parallel, thorn-like sclerotiza- tween coxal fields II and III distinctly shorter tion in middle part. Genital lobes present or ab- than idiosomal width. Idiosomal cupules absent. sent. Physogastric female worm-shaped in outline Idiosomal setation (maximum set observed in ...... Picobia Haller, 1878 Harpypalpinae): scx, vi, ve, si, se, c2, d1, d2, e2, 50. Solenidia phi on tibia of leg I absent. Hypos- f1, f2, h1, ps1–3, g1. Setae scx situated dorsally, tomal apex truncate. Peritremes with clearly or distinctly developed. In males, setae c1 and 2 weakly visible chambers in short lateral branches. pairs of pseudoanal setae present. In females of Propodonotal shield entire and shirt-like. Pygidial Harpirhynchinae, idiosoma bearing only setae shield present. Bases of setae 1a–1a not coalesced. scx, vi, ve, si, se, c2, h1, ps1; in harpirhynchine Physogastric females bulb-shaped outline ...... 51 males, 2–3 pairs of pseudoanal setae present. — Solenidia phi on tibia of leg I present. Hypos- Legs shortened; legs I and II consisting of 5 ar- tomal apex rounded. Peritremes with clearly visi- ticulated segments, their tarsi having weakly de- ble chambers in lateral branches. Propodonotal veloped pretarsi bearing 2 smooth lateral claws shield divided longitudinally into 2 lateral shields, and empodium with tenet hairs. In Harpypalpi- bearing bases of setae vi, ve, si and se. Pygidial nae, legs III and IV normally developed, similar shield absent. Bases of setae 1a–1a coalesced. with anterior legs. In Harpirhynchinae legs III Physogastric females campanulliform outline and IV strongly reduced, 1–3 segmented. Maxi- ...... Columbiphilus Kivganov et Sharafat, 1995 mum set of leg setae in Har­pypalpinae (paired 51. One pair of genital setae and 2 pairs of pseuda- setae are in parentheses): tarsi I–IV — I, II, (tc), nal setae present ...... Rafapicobia Skoracki, 2011 p”, (a), (u), ω1; III, IV (tc), (a), (u); tibiae I–IV — Genital setae absent, 2 pairs of pseudanal setae — I, II, d, (l), (v); III, IV, d, (v); genua I–IV — I, present ...... Neopicobia Skoracki, 2011 II, d, (l),(v); III, IV, without setae; femora I–IV Family Harpirhynchidae Dubinin, 1957 — I, II, d, v; III, IV, v; trochanters I–IV — v; coxae I–IV — I, 1a, 1c; II, 2c; III, 3a, 3c; IV, 4c. Figs. 8–9 Solenidion ω1 II situated dorsally. Larva, proto- Type genus: Harpirhynchus Latreille, 1796 and tritonumphs present; deutonymph absent. Diagnosis. Gnathosoma moderately develdevel-- Male moulting from protonymph. oped. Subcapitulum bearing setae elc. p, n, and This family was revised by Fain (1976, 1994b, m. Peritremes situated at base of stylophore, seg- 1995) who described most of its taxa. Before this mented only in distal parts, straight. Hypostomal revision only a few particular works concerning lobes not fused with stylophore. Hypostomal lips these mites were published. Most important of absent. Palps linear, consisting of 3 articulated them was a paper by Fritsch (1954) concerning segments — trochanter-femur-genu, tibia and ru- harphirhynchids from Germany. dimentary tarsus. Palpal tibia replaced on ventral Within the content of harpirhynchids, Fain surface of trochanter-femur-genu and bearing (1972) established a new subfamily Harpypalpi- distinct paraxial claw-like seta. Palpal tarsus nae for a single genus Harpypalpus with normally strongly reduced. Palpal setation: femur with se- developed legs III and IV. Later, Lombert and tae d, v (only with seta d in Harpypalpinae), genu Moss (1983) described Harpypalpoides, the sec- — d, l”, tibia — d, lT, and tarsus with 1 seta. Id- ond genus in this subfamily. iosoma flattened dorso-ventrally and rounded Presently, A.V. Bochkov along with collabo- outline or sucker-like. Bases of trochanters in- rators works on the systematics of these mites. serted ventro-laterally. Eyes absent. Propodono- The monophyly of Harpirhynchidae is sup- tal shield present, distinctly developed, fused ported by three unique synapomorphies: the with remnants of hysteronotal shield. Ventral side strongly reduced, membranous palpal tarsus, seta of idiosoma devoid of shields in Harpirhynchi- l”G grouped together with dG in the apical part of nae and bearing genital-anal shield in Harpypal- the palpal throchanter-femur-genu, and modified pinae. Opisthosoma weakly developed. Female setae l’T of the palpal tibia (Bochkov 2008).

78 A review of Prostigmata permanently associated with birds

A B

Fig. 8. Harpirhynchus dusbabeki (Harpirhynchidae: Harpirhynchinae), female: A — in dorsal view; B — in ventral view; C — palpal apex (from Bochkov and Literak 2006, with minor modifications).

Bochkov et al. (1999) analysed cladistic rela- tionships of the harpirhynchid genera. In the result of this study, the family Harpirhynchidae has in- corporated the family Ophioptidae, which includes parasites of some snakes belonging to the super- family Colubroidea and was subdivided into three subfamilies: Harpirhynchinae (Harpypalpinae- Ophioptinae). Later on, Bochkov (2008) reconsid- ered the relationships between harpirhynchid sub- families using multiple outgoups and reached the same conclusions. The sister relationships between Harpypalpinae and Ophioptinae are supported by the following unique synapomorphies: subcapitu- lar setae m and n are situated at the same trans- verse level, setae v’F of the palpal femur are ab- sent, the female vulva edged by the sclerotized structures, setae scx are absent in immatures, the genital setae are spur-like and surrounded into the idiosomal cuticle, in immature instars, seta l”G of the palpal genu is absent, seta dG of the palpal fe- mur is comb-like, most prodorsal setae are absent, the hysterosomal setae are dislocated on the ven- Fig. 9. Harpypalpoides sp. (Harpirhynchidae: Harpypalpi- tral side of the idiosoma, the anal opening is ab- nae), female in dorsal view. sent, immatures are apode.

79 M. Skoracki, S.A. Zabludovskaya, A.V. Bochkov

Key to genera of Harpirhynchidae 8. Body not sacciform but either dome-shaped or (based on Fain [1995]) wider than long and narrowed posteriorly ...... 10 — Body elongate, sacciform ...... 9 1. Most hysteronotal setae absent. Legs III and IV 9. Legs I–II moderately reduced. Legs I with 2–4 shortened, 1–2 segmented. Pretarsi III and IV ab- articulated segments. Legs II with 2–3 articulated sent … (Harpirhynchinae Dubinin, 1957) ...... 3 segments...... Neharpyrhynchus Fain, 1972 — Most hysteronotal setae present. Legs III and — Legs I–II strongly reduced. Legs I with only 1 IV normally developed, consisting of 5 articulate very short segment. Legs II similar to legs I or ab- segments. Pretarsi III and IV present, each bearing sent ...... Metharpyrhynchus Fain, 1972 empodium and 2 claws. (Harpypalpinae Fain, 10. Female: Idiosoma wider than long, with 2 1972) ...... 2 broad lateral lobes. Dorsal shield poorly devel- 2. Setae se situated at level of seta si bases. Setae oped. Palpal seta l”G absent ...... 11 d2 present ...... Harpypalpus Dubinin, 1957 — Female: Body dome-shaped, with 2 postero- — Setae se situated distinctly posterior to level of lateral lobes. Dorsal shield distinctly developed. seta si bases. Setae d2 absent...... Harpypalpoides Palpal seta l”G present ...... Anharpyrhynchus Lombert et Moss, 1983 Fain, 1972 3. Setae dG and dF not filiform or rodlike, but rigid 11. Female: Setae ve and si modified into very and shorter than palp tibia, with generally strong short spinelets. Lateral lobes situated in anterior pectinations. Setae l”G similar to other palpalae or third of idiosoma. Legs I and II ventral; bearing rodlike and shortly pectinate ...... 4 claws and empodium ...... Harpyrhynchiella — Setae dG, l”G, and dF of palps filiform, shortly Fain, 1972 pectinate, longer than palp tibia...... — Setae ve and si normal and pectinate. Lateral ...... Trichorhynchiella Fain, 1954 lobes situated in middle part of idiosoma. Legs I 4. Female: Idiosoma rounded, elongate or wider and II marginal, without claws and empodium ...... than long. Legs I–II reduced in size, with maxi- ...... Cypsharpirhynchus Fain, 1995 mum of 4 articulate segments ...... 6 — Female: Idiosoma circular or subcircular. Legs Superfamily Cloacaroidea Camin, Moss, Oliver et Singer, 1967 I and II well developed, with 5 articulated seg- ments ...... 5 The superfamily Cloacaroidea has been re- 5. In females, setae dG, l”G, and dF of palps sim- cently established for two families, Cloacaridae ilar in shape but l”G slightly shorter and narrower and Epimyodicidae (Bochkov and OConnor 2008). than dG and dF. In males, genital opening situated This superfamily is characterized by the following in anterior quarter of dorsum ...... Harpirhynchus synapomorphies: the gnathosoma is shifted ven- Megnin, 1877 trally, the chelicerae are devoid of digits and setae, — In females, setae dG, l”G, and dF of palps dif- the subcapitular setation is completely lost, the ferent in size and shape. In males, genital opening coxal apodemes are longer than half of the body situated in posterior third of dorsum ...... width, in adults, coxal apodemes I are fused into a ...... Harpyrhynchoides Fain, 1972 sternum, and females, genital and anal openings 6. Idiosoma variable in shape, but not rounded or are completely fused to each other. circular ...... 8 The relationships of Cloacaridae and — Body rounded or circular ...... 7 Epimyodicidae with other prostigmatic mites are 7. Dorsal shield small. Legs I and II slightly re- not clear and, therefore, this superfamily is con- duced in size, with only 4 free segments (genu and sidered as incertae sedis within Eleutherengona. femur fused) and inserted marginally; tarsi I–II These mites are strongly modified for endopara- with claws and empodium. Legs III normally sitic mode of life and have lost many morphologi- shaped. Legs IV absent. Propodonotal setae cal structures present in other Prostigmata. The smooth. Setae vi, si, and se short and smooth ...... distinctly separated chelicerae of Cloacaroidea ...... Perharpyrhynchus Fain, 1972 obviously prevent their inclusion in the Cheyle- — Dorsal shield very large. Legs I and II strongly toidea, because all cheyletoid families have com- reduced in size, lacking claws and empodium; legs pletely fused chelicerae with stylet-like movable I slightly ventral, legs II marginal. Legs III and IV cheliceral digits. Such characters, as the fusion of very small, not segmented. Setae vi, si, and se pec- the anal and genital openings and the dorsal posi- tinate ...... Ralliharpirhynchus Fain, 1972 tion of an aedeagus in males indicate the affinity

80 A review of Prostigmata permanently associated with birds of these two families with the parvorder Eleutheren- onto ventral side of idiosoma due to coxal flatten- gona, but are not helpful in clarifying their rela- ing in these mites. Ventral condyles distinctly tionships to other lineages within the parvorder. smaller than dorsal condyles. Idiosomal setae Family Cloacaridae Camin, Moss, Oliver et strongly reduced in size and number. In Cloacari- Singer, 1967 nae, female propodonotum with 1–2 pairs of al- veoli. Opisthosoma of cloacarine females bearing Fig. 10 2–4 pairs of setae. Setation of male propodonotum Type genus: Cloacarus Camin, Moss, Oliver similar to that of females; opisthosoma with 1 pair et Singer, 1967 of terminal, fleshy setae. In females ofPneumoph - Diagnosis. Gnathosoma positioned ventrally, agus propodonotum with 1 pair of alveoli, opist- strongly reduced, oriented perpendicular to idio- hosoma with 3 pairs of short spine-like setae situ- somal axis, retracted into idiosoma, and surround- ated dorsally, terminally and ventrally. Pair of ed ventrally by idiosomal wall. In Cloacarinae, alveoli flanking genitor-anal opening dorsally. In remnants of posterior subcapitulum represented males, only 1 pair of short spine-like setae present, by distinctly sclerotized subcapitular ring, open- situated dorsally on opisthosoma. ing anteriorly and located immediately under dor- Legs I–IV with 3 articulated segments, very sal surface of idiosoma. Pair of strongly sclero- short; rudimentary segment inserted proximal to tized, fang-like chelicerae present ventrally. In apical segment. Two basal leg segments devoid of Pneumophaginae, female gnathosoma devoid of setae. Trochanters largest leg segments. Posterior posterior remnants of subcapitulum (subcapitular margins of all trochanters with large triangular ring) and internal median apodemes. In males, projection articulated with respective coxal apo- gnathosoma completely absent. Idiosoma ellipti- demes. In most cloacarids reduced tibiae with 1 cal and distinctly flattened dorso-ventrally. spine-like seta, but devoid of setae in Pneumopha- Propodonotal shield present and occupying most gus, except for 1 seta on tibia I. Tarsi broadly of propodonotal surface, distinctly ornamented in rounded in cloacarines; in Pneumophagus tarsi II– Cloacarinae, and weakly sclerotized and devoid of IV flattened laterally. Claws absent on all tarsi. In ornamentation in Pneumophaginae. Opisthosoma Cloacarinae tarsi I and III each with 8 setae (in distinctly shorter than metapodosoma; opistho- Caminacarus chrysemys Pence and Casto, 1975, somal striations absent. In females, anal and geni- tarsi III with 9 setae), tarsi II — 9 setae, and tarsi tal openings fused to each other forming single IV — 7 setae. Short globose solenidion on tarsi opening situated ventrally, terminally or dorsally. I–III situated ventro-terminally. In Pneumopha- In females of some genera, lateral walls of genito- gus; tarsi I–IV bearing 5–9–12–12 setae, respec- anal opening elongated and modified into 1 pair of tively. Solenidion absent on all tarsi. Larva un- short, finger-like projections. In Chelonacarus, known. Only 1 nymphal stage known. genito-anal opening covered ventrally by pair of The monophyly of the Cloacaridae is strongly triangular sclerotized folds. In males, aedeagus supported by the following synapomorphies: the present and genital opening situated dorsally on gnathosoma is inserted perpendicular to the idio- anterior part of body. Coxae completely fused with somal axis, the subcapitulum is strongly reduced, ventral surface of idiosoma; only their anterior the palps are completely lost, the opisthosoma is margins represented by distinct coxal apodemes. shortened and distinctly shorter than the metapo- Coxal apodemes strongly elongated, longer than dosoma, the idiosomal striation is absent, there is half of idiosomal width. In adults, coxal apodemes a sclerotized ventro-marginal sclerotized frame I fused forming sternum. In Cloacarinae, sternum bordering the lateral margins of the idiosoma, the with internal median keel, at least in anterior half. legs are inserted laterally and are strongly short- Keel continuing anteriorly as median apodeme ened and thickened, the femora and genua of all with bifurcate apex. Apices fused anteriorly with the legs are completely fused, the leg trochanters subcapitular remnants and associated with scler- have distinct basal projection bearing glenoid cav- ites, interpreted here as sigmoid pieces (= capitu- ities, and the pretarsal claws are absent. lar apophyses). In females of Pneumophagus, an- The family Cloacaroidea includes two sub- terior parts of coxal apodemes I fused with families, Cloacarinae and Pneumophaginae with anterior part of subcapitular remnants. Proximal one species. The latter, bird-associated subfamily parts of coxal apodemes I–IV with 2 condyles at was described by Fain and Smiley (1989), and is trochanteral articulation. Dorsal condyle shifted characterized by the following synapomorphies:

81 M. Skoracki, S.A. Zabludovskaya, A.V. Bochkov

A B

Fig. 10. Pneumophagus bubonis (Cloacaridae: Pneumophaginae): A — female in ventral view; B — male in ventral view (from Fain and Summers 1989, with minor modifications). the gnathosoma of the male is absent, the anterior acterized by such autapomorphies as the presence branches of coxal apodemes I surround the female of the ereynetal organ and the double genital gnathosoma, the coxal apodemes are fused to each discs. other proximally forming a ventral idiosomal Family Ereynetidae Oudemans, 1931 frame, tibiae II–IV are devoid of setae, solenidia are absent on all leg tarsi, tarsi II–IV are flattened Figs. 11–12 laterally, and most of their setae are arranged in a Type genus: Ereynetes Berlese, 1883 row (Bochkov and OConnor 2008). Diagnosis. Chelicerae bases fused to each other and with subcapitulum forming gnathosomal Parvorder Eupodina Krantz, 1978 capsule. Movable cheliceral digit short, linear and Superfamily Tydeoidea Kramer, 1877 stylet-like. Palps linear non-raptorial, composed The Tydeoidea includes four families: Tydei- of 4 segments (trochanter, femuro-genu, tibia, tar- dae, Triophtydeidae, Iolinidae and Ereynetidae sus) in Ereynetinae and reduced to 1–3 segments (Andre and Fain 2000; Walter et al. 2009). This in Lawrencarinae and Speleognathinae. Claw-like superfamily has a worldwide distribution, and its structures correspond to strong ventral setae (v) members are associated with a wide range of hab- and observed only in Ereynetinae. Palpal setation, itats, from soil to the nasal cavities of mammals. femur-genu with 2 pairs of dorsal setae dF and dG Highly specialized blood-sucking bird endopara- (in Ereynetinae, absent in other subfamilies), tibi- sites are found only in the family Ereynetidae ae with 2 dorsal setae dT and lT in Ereynetinae, 1 (subfamily Speleognathinae) (Fain and van Goe­ or without setae in other subfamilies, tarsus with them 1978, 1986; Domrow 1965, 1969). The phy- setae d, l’, l”, pζ, v, in Ereynetinae, reduced to 2–3 logeny of the Tydeoidea, based on morphological setae (d, l’, l”) in Lawrencarinae and Speleognathi- characters, was reconstructed by Andre and Fain nae, tarsal solenidion ω present in Ereynetinae and (2000). According to their results, the Ereynetidae present or absent in other subfamilies. Hypostomal was derived from an ancestral tydeids and is char- apex with 1–2 pairs of subcapitular setae (n, m).

82 A review of Prostigmata permanently associated with birds

A B Fig. 11. Boydaia sturni (Ereynetidae: Speleognathinae), female: A — in dorsal view; B — in ventral view.

A B

Fig. 12. Boydaia sturni (Ereynetidae: Speleognathinae), details of female: A — ereynetidal organ (e.o.); B — tarsus I in dorsal view.

Postcheliceral stigmata obscure. Idiosoma weakly setae: ag1–5, ps1–2, g1–5. Neotrichy of setae ps sclerotized, sac-like. Reticulate ornamentation of observed only in Lawrencarus eweri (Lawrencari- prodorsum present or absent. Eyes present or ab- nae). In males of Ereynetinae 1–3 pairs of eugeni- sent. Propodonotal and hysteronotal shields pres- tal setae (eu) present, in Lawrencarinae and Spe- ent in some ereynetines and absent in most of spe- leognathinae 1 pair of setae eu present or absent. leognathines and Lawrencarinae. Ventral side of Genital papillae 2 pairs, well developed in Ereynet- idiosoma without plates. Distance between coxal inae, or reduced in Lawrencarinae and Speleog- fields II and III distinctly shorter than idiosomal nathinae. Legs long and massive, covered by re- width. Idiosomal setation (maximum in Ereyneti- ticulate ornamentation. Coxal fields of legs I–IV nae): vi, ve, si (= anterior sensilla), se, c1, c2, d1, distinctly bordered and well sclerotized. Coxae e1, f1, f2 (= posterior sensilla), h1. Pair of poste- setation (maximum set) I–IV: I 1a, 1b, 1c; II 2c; rior sensilla usually present in free-living species III 3a, 3b, 3c, 3d; IV 4a, 4b, 4c. Setae 3d and 4c and absent in most parasities. Cupules im and ia absent in Lawrencarinae and Speleognathinae. present or absent. Genital opening situated poste- Trochanters I–IV — I–III l’; IV without setae. rior to legs IV, longitudinal. Genital region with Femora I–IV — I with 4–7 setae; II (d), (v); III v,

83 M. Skoracki, S.A. Zabludovskaya, A.V. Bochkov d, l; IV (d), (v). Femora I, IV divided on basi- and 2. Setae on trochanters I and II present. Solenidion telofemur in Ereynetinae, not in Lawrencarinae on tarsi of palps present. Femur IV with at least 2 and Speleognathinae. Genua I–IV — I (d) (l); II setae. Genu IV with at least 3 setae ...... (d), (l); III d, (l); IV d, (l). Tibiae I–IV — I d, (l), ...... Boydaia Womersley, 1953 (v); II d, (v), III d, (v); IV d (v). Tibiae I with in- — Setae on trochanters I and II absent. Solenidion ternal sensory complex (ereynetal organ) where on tarsi of palps absent. Femur IV with 1 seta. solenidion of tibia I (φI) recessed into inverted Genu IV with 2 setae ...... Coboydaia Fain, 1971 sac-like structure with narrow duct that opens 3. Tarsi I–IV without striated setae ...... 5 near distal margin of segment. Tarsi I–IV — I — Tarsi I–IV with striated setae ...... 4 with 12 setae, solenidion ω present or absent; II 4. Palps with 1 segment. Eye lenses absent. Num- with 9 setae, III and IV with 8 setae. Tarsi I–IV ber of coxal setae I–IV 1–1–1–1, trochanters with- with pair of large claws and with setiform median out setae ...... Aureliania Fain, 1958 empodium. In development 4 immature instars: — Palps with 3 segments. One pair of prominent larva, protonymph, deutonymph, tritonymph. All eye lenses present. Number of coxal setae I–IV nymphs mobile in ontogenesis of Ereynetinae and 2–1–1–0, trochanteral setae 1–1–0–0 ...... Lawrencarinae, while in Speleognathinae all ...... Ophthalmognathus Dubinin, 1957 nymphs calyptostatic. 5. Apical or ventro-apical setae of tarsi I–IV nor- The taxonomic content of the family Ereyneti- mally developed and barbed ...... 6 dae was reviewed by Fain (1957). He included the — Four apical and 2 ventro-apical setae of tarsi family Speleognathidae erected by Womersley I–IV rod-like microsetae ...... Psittaboydaia (1936) to the Ereynetidae as a subfamily together Fain, 1985 with the newly created Lawrencarinae. 6. All tarsal setae of legs I–IV barbed ...... 10 Within the Ereynetidae, the Speleognathinae — Only 6–8 apical or ventro-apical setae of tarsi form a monophyletic group. Its monophyly is sup- I–IV barbed, other setae smooth ...... 7 ported by such synapomorphy as calyptostatic 7. Dorsal and ventral setae of idiosoma dentate. nymphs. The subfamily Lawrencarinae is the sis- Dorsal shield absent ...... 9 ter group to Speleognathinae (Andre and Fain — Ventral idiosoma with filiform setae mixed with 2000). striated or dentate setae. Dorsal shield variable ... 8 In 1985, Fain divided the subfamily Speleog- 8. Dorsal shield present. Pair of eye lenses present nathinae into five tribes: Boydaiini, Trispeleog- ...... Neastrida Fain, 1962 nathini, Speleognathini, Speleochirini, and Para­ — Dorsal shield and eye lenses absent ...... speleognathopsini. Among them, representatives ...... Neoboydaya Fain, 1958 of the two first tribes are associated with birds. 9. Palps with 3 segments. One pair of eye lenses The tribe Boidaiini with the genera, Boydaia and present. Claws of leg I stronger than claws II–IV. Coboydaia is characterized by the absence of dor- Hypostome with 1 pair of setae. Five pairs of gen- sal shields and eye lenses and by the presence of ital setae present. Number of coxal setae I–IV 3-segmented palps. The tribe Trispeleognathini in- 2–1–1–1 ...... Trispeleognathus Fain, 1958 cludes 14 genera of nasal parasites of non-passer- — Palps with 1 segment. Eye lenses absent. Claws iform birds. These genera are characterized by the of leg I–IV subequal. Hypostome with 2 pairs of polymorphic chaetotaxy and 1–3 segmented palps. setae. Two pairs of genital setae present. Number Additionally, eye lenses and dorsal shields are of coxal setae I–IV 2–0–1–0 ...... Pterniboydaia present in some species. Fain, 1985 10. Dorsal and ventral setae of idiosoma either Key to bird-associated genera of dentate or dorsal setae mainly piliform or narrow Speleognathinae (Ereynetidae ) with striated base ...... 12 1. Chaetotaxy strongly variable, 2 or 3 different — Dorsal and ventral setae of idiosoma either ex- types of setae present in same species. Tibiae I–IV clusively or predominantly barbed ...... 11 with 5–3(2)–3(2)–3(2). Dorsal shield and eye 11. Dorsal shield well developed. Palps with 3 lenses present or absent. Palps with 1–3 segments. segments. Eyes either flat or with prominent lenses (Trispeleognathini Fain, 1980) ...... 3 ...... Speleognathopsis Cooreman, 1954 — All setae barbed. Tibiae I–IV with 5–3–3(2)–3(2) — Dorsal shield absent. Palps with 2 segments. or 4–2–2–2. Dorsal shield or eye lenses absent. Eyes with prominent lenses ...... Metaboydaia Palps with 3 segments (Boydaiini Fain, 1985) ..... 2 Fain, 1962

84 A review of Prostigmata permanently associated with birds

12. Dorsal setae of idiosoma either exclusively or tion of the idiosoma and legs, and the male genital predominantly dentate ...... 14 opening located dorsally. Their legs are inserted — Dorsal setae of idiosoma either exclusively or laterally; ambulacra are well developed, at least, predominantly filiform ...... 13 on the anterior legs (Fig. 13C). Many representa- 13. Dorsal shield absent. Palps with 1 segment. Se- tives of cheyletids spin the group (Bakericheyla) tae vi and sce absent. Hypostomal setae absent ..... or individual (Ornithocheyletia) cobweb covers...... Meropiboydaia Fain, 1985 Such cobweb cover protects mites and their eggs, — Dorsal shield present. Palps with 3 segments. stabilises the temperature and humidity regime Setae vi and sce present. Hypostomal setae present and prevents elimination from the host body (Aki- ...... Astrida Fain, 1955 mov and Gorgol 1990). 14. Claws of legs III and IV subequal in sizes ... 15 Harpirhynchine females of some skin-inhab- — Claws either of tarsi III and IV or only of tarsi iting genera, for example, Neharpyrhynchus, are III unequal and modified; one or both abruptly quite different morphologically from other skin- bent in their apical parts ...... Ralliboydaia inhabiting mites. Their females are attached at the Fain, 1962 base of calamus of the body covert feathers on the 15. Dorsal shield absent. Palps with 3 segments. neck or head of avian host. They are oriented par- Eyes with prominent lenses ...... Phoenicopteriella allel to the length of the shaft, with mouthparts Fain, 1970 inserted in the host skin; their idiosoma is sacci- –Large dorsal shield present. Palps with 1 seg- form and the eggs are laid in a string; the female ment. Eyes absent ...... Picinyssus Fain, 1969 opisthosoma, thus, serves as the “stalk” or site of attachment for the eggs (Moss et al. 1979; Boch- RELATIONSHIPS WITH HOSTS kov and Literak 2011) (Fig. 13D). Microhabitats on the host body The bird skin almost devoid of glands is not so Amongst parasitic acariform mites perma- beneficial substrate as the mammalian skin, there- nently associated with their hosts, the external fore, mites of the intracutaneous morphoecotype morphological adaptations to parasitism were ana- like Demodicidae and Psorergatidae are absent lyzed in feather mites (Psoroptidia) by Mironov among bird-associated prostigmatans. Among (1987) and in mites associated with mammals them, however, mites of the skin-burrowing mor- (Bochkov 2007). According to several external phoecotype (analogous to the knemidocoptid mor- morphological criteria (structures of the gnatho- phoecotype in feather mites) belonging to the fam- soma, idiosoma, legs, and setation) these authors ily Harpirhynchidae are present. In this family, the have recognized different morpho-ecological tendency of transition from the skin surface (ec- types (morphoecotypes); representatives of each toparasitism) to intradermal parasitism (endopara- of them are associated with a particular habitat on sitism) is widely observed and this switches inde- the host body. Prostigmatan mites permanently as- pendently happened within several harpirhynchid sociated with birds, being represented by both genera (for example Harpyrhynchoides) (Fain ecto- and endoparasities, occupy wide spectrum of 1995). Harpirhynchid females accumulate in the microhabitats on the host body: skin, subcutane- feather follicles, where they develop subcutaneous ous layers, nasal cavities, feather quills, and even cysts (Fain 1995; Literak et al. 2005); many of bronchi (Table 1). Most of these mites belong to them are strongly morphologically modified. At morphoecotypes recognized in feather mites the same time, harpirhynchid males belonging to (Mironov 1987). skin-burrowing species do not undergo strong mor- Skin inhabiting prostigmatans of the families phological modifications. Females of the subfamily Cheyletidae (Ornithocheyletiini) and Harpirhyn­ Harpypalpinae, forming the subcutaneous cysts chidae (many genera of Harpirhynchinae) corre- (Fig. 13B), are even less specialized than ectopara- spond to the epidermoptoid morphoecotype sitic mites of the subfamily Harpirhynchinae, be- (Mironov 1987), which is analogous to the skin cause their posterior legs are not reduced, and their morphoecotype of mammal-associated mites (Bo- opisthosomal setation is more reach than in harpi- chkov 2007). Prostigmatan mites referred to this rhynchins. In some genera of Harpirhynchinae, morphoecotype have the terminally situated gna- cyst-inhabiting species (Harpirhynchus or Hapy- thosoma, which can possess various apophyses rhynchoides) also do not significantly differ from used for attaching, the idiosoma is strongly flat- ectoparasitic (skin-inhabiting) representatives of tened dorso-ventrally, distinctly developed seta- the family, whereas species of the genera Anharpy-

85 M. Skoracki, S.A. Zabludovskaya, A.V. Bochkov

A B

C D

Fig. 13. A — dissected quill with syringophilid mites (SEM photo); B — cyst formed by harpirhynchid mites (photo by I. Lit- erak); C — Harpirhynchoides spp. in dorsal view (SEM photo); D — female of Neharpyrhynchus sp. attached to bird feather (from Bochkov and Literak 2011, with minor modifications). rhynchus, Cypsharpirhynchus, Harpyrhynchiella, Legs are relatively short. In contrast to feather and Trichorhynchiella are strongly modified being mites which take harborage within quills, many habitually similar with some skin-burrowing mites idiosomal and some leg setae of these mites are of the astigmatan subfamily Teinocoptinae (Sar- long or ultralong and control, thus, a significant coptoidea: Sarcoptidae). Their gnathosoma is re- space around the mite. Mites of the subfamily Sy- placed to the ventral idiosomal surface, the idioso- ringophilinae are mostly associated with the quills ma is sacciform, the anterior legs are shortened and of flight feather, whereas the absolute majority of the posterior legs are strongly reduced or lost; the picobiins inhabit quills of the body feathers (John- body and leg setation is very poor. ston and Kethley 1973). Females of the latter sub- The parasitic prostigmatans inhabiting quills family are characterized by the physogastry — an correspond to the dermoglyphid morphoecotype extensive enlargement of the female hysterosoma (Mironov 1987). Among prostigmatan mites this in the process of feeding (Skoracki et al. 2001; morphoecotype is represented by mites of the fam- Skoracki 2011) — it is unknown for other quill ily Syringophilidae and cheyletid mites of the tribe mites. Metacheyletiini. The idiosoma of these mites is The predatory mites inhabiting quills belong strongly elongated (Syringophilidae) or egg- to the tribe Cheletosomatini (Cheyletidae). These shaped (Metacheyletia) and weakly sclerotized. mites do not differ habitually from nidicolous

86 A review of Prostigmata permanently associated with birds cheyletids. Even mites of the cheletosomatine ge- ly omitting a stage of the skin ectoparasitism. Quill nus Picocheyletus transferred from a predation to predators and parasites switched to the life in the parasitism do not differ morphologically from feather quills originate directly from the nest pre- their predatory relatives. It should be noted, how- dation (Bochkov et al. 2008); nasal mites (Spe- ever, that some idiosomal and leg setae of these leognathinae) also originated directly from the mites are ultralong, as in parasitic prostigmatans free-living forms omitting the skin parasitism; in- inhabiting quills. terstitial mites (Pneumophaginae) supposedly The nasal prostigmatan parasites of birds are switched to birds from turtles and the origin of represented by the subfamily Speleognathinae cloacarid parasitism is still not clear (Bochkov and (Ereynetidae). These mites belong to the respira- OConnor 2008); only intracutaneous mites (some tory morphoecotype established by Bochkov Harpirhynchidae) originated from ectoparasitic (2007) and are not significantly modified in com- forms inhabiting the skin surface. parison to free-living relatives. Their palps are This complicated picture of colonization of 1–3-segmented (four-segmented palps in free-liv- the avian body by prostigmatan mites could be ex- ing ereynetids), the idiosoma is slightly flattened plained by the absence of the direct phylogenetic dorso-ventrally and in some species bears the eyes, links between these mites. Since all bird-associat- the body and leg setae are relatively short, and the ed prostigmatans are true parasites feeding on the legs are well developed, inserted laterally, and live tissues or predators, they did not colonize the bear empodium and paired claws; in males, an ae- feather vexillums (proctophyllodid morphoeco- deagus is absent. An interesting adaptation of par- type) or the down feathers (analgid morphoeco- asitic ereynetids is the presence of calyptostatic type); nor they did occupy the quill walls inhabited nymphs. In these mites, the active stages are only by some feather mites (Laminosioptidae). larva and adults (Andre and Fain 2001). The interstitial bird parasites among prostig- Host specificity and evolution on birds matans are represented by a single species Pneu- Bird systematics. All extant birds (Neorni- mophagus bubonis (Cloacaridae) from tissues thes) are split into two infraclasses: Palaeognathae around bronchi of Bubo virginianus (Strigiformes: (two extant orders, ratites and tinamous), and Strigidae) from USA (Fain and Smiley 1989). This Neognathae with two cohorts, Galloanserae (land- is a typical representative of the interstitial mor- fowls and waterfowls) and Neoaves (other neog- phoecotype. Among acariform mites permanently naths). The cohort Galloanserae includes two or- associated with vertebrates this morphoecotype is ders, Galliformes and Anseriformes. The cohort represented by the families Epimyodicidae and Neoaves includes 30 orders (Clements 2011). Tax- Cloacaridae (Bochkov 2007). The body of these onomic accounts of all bird orders are provided in mites has a slightly elongated and streamlined Table 6. form; in females, the gnathosoma is almost com- Hosts scpecificity. Most prostigmatans are pletely reduced and represented only by chelicerae monoxenous or narrowly oligoxenous parasites (absent in males), the legs are thickened and short- and only 8.7–30.5% of species in various groups ened and their segments are telescopic, almost all are associated with host of one family, 1.7–20.6% body setae are absent, and the leg setation is — with hosts of one order and only 1.1–3.4% — strongly reduced; the coxal apodemes are hyper- with hosts of various orders. All data about pros- trophied. Thus, cloacarid mites are almost ideally tigmatan host specificity are summarized in Table adapted for moving in friable interstitial tissues. 7. Many species records, however, are solitary and Thus, bird-associated prostigmatans are rep- the level of host specificity could be overestimated resented by four main morphoecotypes: skin (epi- for some species. The most strictly specific symbi- dermoptid), skin-burrowing (knemidocoptid), onts are parasitic Cheyletidae and Syringophili- quill (dermoglyphid), and interstitial. dae, 83% and 70.5% of monoxenous species, re- In feather mites and mammal-associated Ac- spectively; the less specific symbionts are ariformes, the skin-inhabiting forms belong to the Ereynetidae and Harpirhynchidae — 37% and ancestral morphoecotype and most other morpho- 62.7%, respectively. It is interesting that some spe- ecotypes are its derivations (Mironov 1987; Boch- cies in the families, represented generally by the kov 2007). In prostigmatans, the colonization of highly specific parasites, are associated with wide different microhabitats on the bird body has been spectrum of hosts, for example, Bakericheyla realized by free-living mites independently, most- chanayi (Cheyletidae) (Akimov and Gorgol 1990;

87 M. Skoracki, S.A. Zabludovskaya, A.V. Bochkov see also Appendix deposited at Acarina web-site: Johnston 1975; Moss et al. 1979). At the same http://insects.ummz.lsa.umich.edu/acarina/). In the time, it could be stated, based on distribution of time of global cataclysms, such generalists ensure these mites, that at least 50% of their extant genera the survival of the group when most hosts with are known to date. For example, among 110 extant specialized parasites undergo extinction. passerine families, prostigmatans are recorded on At the generic level the host specificity of hosts from more than 50 families (Appendix). prostigmatan mites is less marked but still signifi- Mites of the family Cheyletidae are known cant. In the Cheyletidae, four genera (36%) para- from birds of 12 orders, Syringophilidae — from sitize birds from one family, the genus Apodiche- 21 orders, Harpirhynchidae — from 16 orders and les is associated exclusively with Apodiformes, Ereynetidae — from 18 orders (Table 6). Moss and six other genera (54%) are associated with and Wojcik (1978) also listed Caprimulgiformes birds of two or even more orders. In the Harpi- as hosts of harpirhynchids but did not provide the rhynchidae, four genera (33%) are associated with full data; therefore these records were not taken hosts of one family, four (33%) — with hosts of here into account. one order, and four (33%) — with hosts of various As it is expected, representatives of all pros- orders. In Syringophilidae, 39 genera (74%) para- tigmatan families, excluding Cloacaridae, are re- sitize birds of one order and only 14 genera (26%) corded mostly from hosts of species-rich orders, are associated with birds of two or even five (Pico- i.e. Apodiformes, Charadriformes, Columbifor­ bia) various orders. Finally, in the Ereynetidae, mes, Coraciiformes, Cuculiformes, Galliformes, eight genera (50%) are associated with one bird Gruiformes, Passeriformes, Pelecaniformes, Pici- family, and eight (50%) — with birds of various formes, Psittaciformes, and Strigiformes. orders (Appendix). The only mites of the family Syringophilidae Most prostigmatan families and subfamilies are known to date from palaeognathous birds of the are cosmopolite (Table 1), excluding Pneumocop- order Tinamiformes; whereas other prostigmatans tinae (Cloacaridae), which is still known from a are still recorded exclusively from Neognathae. single record (Fain and Smiley 1989). Within the Neognathae, hosts of the more ar- To date, prostigmatans are known from birds chaic Galloanserae clade are parasitized by most from 25 of 34 orders belonging to neognathous as prostigmatan families, excluding Cheyletidae and well as to paleognathous (Tinamiformes) birds Cloacaridae on Anseriformes, but the absolute (Table 6). These mites are absent on birds of the majority of mite species are associated with hosts orders Cariamiformes, Eurypygiformes, Gavii- of the clade Neoaves and with passerines within formes, Mesitornithiformes, Otidiformes, Pha- this clade. ethoniformes, Sphenisciformes, Struthioniformes, Thus, birds were colonized independently by and Trogoniformes. Their absence on Struthioni- mites of several prostigmatan phylogenetic lines formes and Sphenisciformes is explainable the ex- — three groups of cheyletids (Ornithocheyletiini, ternal morphological modifications of these hosts Metacheyletiini, and Cheletosomatini), Syringo- (especially feathers), whereas birds of the other philidae, Harpirhynchidae, Ereynetidae (Speleog- orders are insufficiently explored (or not explored nathinae), and Cloacaridae (Pneumocoptinae). at all) for the presence of these mites. However, As it was the above mentioned, the superfam- with a high probability they could also be infected ily Cheyletoidea, including most bird permanent by prostigmatans. Prostigmatans are characterized symbionts, is represented by two main phyloge- by a relatively low prevalence and many of unex- netic lineages: Cheyletidae–Syringophilidae (I), plored hosts are rare and poorly represented in and Harpirhynchidae (Demodicidae–Psorergati- collections. Moreover, most of museum bird spec- dae) (II) (Bochkov 2008). imens are preserved as dry skins and unavailable The ancestor of lineage I undoubtedly was a for examination on prostigmatans because tech- predator preying on other arthropods. Most part of niques of their collecting can damage the skins. the apomorphic modifications of these mites con- The number of described prostigmatans rep- cerns the gnathosomal structures and, probably, resents only a small fraction of their actual biodi- serves for more effective preying. At the same versity because their expected number is estimated time, features of the idiosoma and legs in these as 5000 species (instead of 500 presently known mites are relatively archaic and weakly modified, species) based on the data about host specificity like those in the other predatory family Raphig- and number of their potential hosts (Kethley and nathidae. The family Syringophilidae originated

88 A review of Prostigmata permanently associated with birds from the common ancestor with Cheyletidae, ventrally in its basal part, exposing the hollow in- which preyed in������������������������������������ bird nests. Syringophilids devel- terior. The mites occupied spaces between septae, oped some adaptations to the parasitic mode of one mite per inter-septal space. In addition, from life in feather quills (the stylophore surrounded by one to four eggs were present in some inter-septal the idiosoma and the elongate body); they have spaces. It should be noted that no other mites or also lost some structures, which serve for preying remains of mites were observed inside the quills and are useless for permanent parasites: the thumb- that might serve as a prey. Some examined speci- claw complex, cupules, and the strong sclerotiza- mens contained dark inclusions, similar to hema- tion of the shields. In these mites, some idiosomal tite inclusions, which usually are visible in blood- setae (4a and ps3) have undergone a reduction in feeding cheyletids of the genus Bakericheyla parallel to parasitic cheyletoids of lineage II. In (Ornilhocheyletiini). Therefore, it can be con- Cheyletidae, the parasitism on birds originated in- cluded that these mites are true parasites on the dependently in three phylogenetic lineages (tribes) avian host. In this case we see transition to para- (Bochkov and Fain 2001; Bochkov 2004). sitism happened literally in “our presence”. Family Cheyletidae. This family is reprerepre-- Among other representatives of the tribe, species sented by the predators and permanent parasites of of the genus Cheletoides are associated with gal- the terrestrial vertebrates, birds and small mam- liform birds (Gallifor­mes). A single species of the mals. To date, it includes more 380 species in 72 genus Eu­­che­­letop­sis is recorded from tree swifts genera and is the most species-rich grouping com- of the genus Hemiprocne (Apodiformes: paring to other families of the superfamily Chey- Hemiprocnidae), and two species of the genus letoidea (Volgin 1969; Summers and Price 1970; Cheletosoma are associated with hosts of the or- Bochkov 2004). ders Gruiformes and Cuculiformes. The genus Cheyletids have a worldwide distribution and Metacheyletoides includes species from galliform are known from a wide spectrum of biotopes. They (Galliformes) and cuculiform (Cuculiformes) occur in a soil, vegetable debris, in nests of in- birds. Mites of the genus Cheletopsis are associ- sects, mammals and birds (Volgin 1969). This ated exclusively with charadriiform birds (Char- family includes almost all intermediate stages be- adriiformes) (Bochkov et al. 2002; Bochkov and tween free-living predators and highly specialized OConnor 2003). Its representatives show variable permanent parasites of vertebrates, birds and specificity to their hosts; some species are limited mammals (Bochkov and Fain 2001; Bochkov to hosts of one genus, whereas others occur on 2004). The obligatory phoresy on certain insects is birds of several charadriiform families. typical for some predatory cheyletids (Volgin The tribe Ornithocheyletiini (four genera) is 1969; Klimov et al. 2006). There are several gen- represented by ectoparasites inhabiting the skin eral directions in the cheyletid evolution: predator- surface of birds. Mites of the genus Neocheyle- dendrophils, predator-entomophils, forms being tiella are mono- or oligoxenous parasites associ- phoretic on insects and living in their nests, nidi- ated with birds of four orders: Passeriformes, colous predators in vertebrate nests, parasites of Pelecaniformes, Columbiformes, and Strigi- birds and mammals (Volgin 1969; Bochkov formes. Species of the genus Bakericheyla are 2004). known from birds if three orders, Apodiformes, Among bird-associated cheyletids, mites of Coraciiformes, and Passeriformes. Most of these the tribe Cheletosomatini (six genera) include species are highly specific to their hosts and dis- predators inhabiting the feather quills and prey tributed in Africa, but Bakericheyla chanayi, how- there on parasitic astigmatic mites and mites of ever, is known from birds of various families and the family Syringophilidae. All representatives of orders and has a cosmopolitan distribution (Aki- this tribe are morphologically similar to Ac- mov and Gorgol 1990). The genus Ornitocheyletia aropsellina-like mites and probably originated is represented by parasites of six bird orders: from the common ancestor inhabited bird nests. Columbiformes, Cuculiformes, Galliformes, Pas- The monotypic genus Picocheyletus was recently seriformes, Piciformes, and Psittaciformes. Orni- described from quills of Tricholaema hirsuta fla- tocheyletia species are mostly mono- or oligoxe- vipunctata from Gabon (Bochkov and OConnor nous parasites. Three species known in the genus 2003). These mites were collected from quills of Apodicheles are restricted to swifts from South- the primary and secondary wing feathers. In each East Asia and Africa (Apodiformes: Apodidae). infested feather, the rachis was naturally split Thus, mites of the tribe Ornithocheyletiini are

89 M. Skoracki, S.A. Zabludovskaya, A.V. Bochkov known from representatives of several bird orders, tion of syringophilid species is determined by belonging to the two major phyletic lines, Gal- thickness of the quill walls and mite capability to loanserae (excluding Anseriformes) and Neoaves. pierce it. It can be hypothesized that the ancestor of this The second important evolution aspect in sy- tribe parasitized on a common ancestor of these ringophilid evolution on their hosts is a distribu- superorders. tion of these mites on various types of quills. The The tribe Metacheyletiini includes the only wing-feathers, i.e. primaries and secondaries, are genus Metacheyletia with five species inhabiting probably the ancestral type of the syringophilid the feather quills (Bochkov and Skoracki 2011). habitat. The majority of representatives of the Three of these species parasitize parrots from the family, including the most archaic ones, were as- Old and New Worlds, and two species were re- sociated with quills of these feathers. The subfam- corded from quills of passeriform and cuculiform ily Picobiinae represented mostly by the dwellers birds in the Tropical Africa. The biology of Me- of the body covert feathers was probably originat- tacheyletia species is still unknown. Atyeo et al. ed in wing quills, because representatives of the (1984) believed that these mites are predators be- archaic genus Calamincola occupy these micro- cause their chelicerae are too short to pierce the habitats (Casto 1977). In comparison to syringo- quill wall. Moreover, in all cases, these cheyletids philines, picobiines are much more morphologi- were collected together with other mites in those cally specialized and possess some advantages, quills. Bochkov and Fain (2001), however, had ror instance heterosomy, which, probably, allowed another opinion and suggested that these mites are them to use more effectively small quills of the parasites. According to these authors, the morpho- body coverts. Thus, picobiines avoided competi- logical characters of Metacheyletia species are tions with other syringophilids and formed the similar to those of other parasitic cheyletids: legs second line distributed in parallel with syringophi- I–III are short, legs IV are reduced, setae of the lines on the same hosts. Unfortunately, the biodi- palpal tarsi are short and smooth, the gnathosoma versity of this group is yet poorly studied in com- is relatively small. It is suggested that these mites parison to Syringophilinae. use for feeding the openings pierced by the adult Among 48 genera of the subfamily Syringo- syringophilids. This hypothesis is supported by philinae, 38 genera are restricted to particular avi- the presence of syringophilids in all cases where an orders, whereas most of remaining mite genera these mites were found. occur on birds of several orders. Family Syringophilidae. The syringophilid Phylogenetic lineage II of Cheyletoidea is ex- ancestors were, probably, initially predators on clusively represented by permanent parasites of other mites inhabiting feather quills. According to vertebrates which can be characterized by the the estimation based on the “molecular clock”, the unique palpal structure. All three families of this divergence between cheyletids and syringophilids lineage demonstrate a tendency towards intracuta- could take place approximately 180–185 million neous parasitism (a part of Harpirhynchidae, De- years ago in the Early Jurassic (Dabert et al. 2011). modicidae and Psorergatidae) (Bochkov 2009). Thus, syringophilids were, probably, associated Skin-inhabiting Harpirhynchidae are relatively even with the bird ancestors — theropod dino- slow-moving ectoparasites, which are very often saurs, many of which possessed feathers (Mayr et embedded in the host epidermis. By this reason, al. 2005). the reduction tendencies prevailed in the evolution The host-parasite relationships of syringo- of this mite lineage. philid were analyzed by Kethley and Johnston It is presumed from the wide distribution of (1975). According to data of these authors the these mites on birds (Harpirhynchidae) and mam- phylogenetic pattern of these mites and birds are mals (Psorergatidae–Demodicidae), that the com- strongly incongruent at the level of mite genera mon ancestor of this branch could occur on the and bird orders. This incongruence could also be common ancestor of birds and mammals. Transi- explained by the horizontal syringophilid switches tions from birds, which are hosts of the most ar- on phylogenetically distant hosts. Kethley and chaic representatives of line II (Harpypalpinae), Johnston (1975) considered host switches as the onto mammals, however, can not be excluded dur- main modus in the evolution of this family and ing the early phase of mammal evolution (Bochk- even proposed a new term for it — resource track- ov 2008). A possible reason for the absence of ing. According to their hypothesis, host distribu- cheyletoids on recent reptiles (excluding snakes)

90 A review of Prostigmata permanently associated with birds may be explained by the peculiarities of their skin It looks like a paradox that mites of the more structure and molting. Squamata lose the entire archaic subfamily are associated with the highly external dermal layer or large pieces of it during evolved hosts. On the contrary, the immature instars the molt (Landmann 1986); moreover, they have a of the Harpypalpinae are highly derived in their multi-layer keratinized epithelium, and skin glands morphology, because they have lost legs and are are absent. The high probability of loss of mites characterized by a quite specialized idiosomal seta- during reptile molting seems to have prevented the tion (Bochkov et al. 1999). Apparently in this case primary settling of cheyletoid parasites on these we have two principally different pathways of the hosts. These mites are absent also on crocodilians morphological evolution in these two subfamilies probably because of their aquatic mode of life. In (Lombert and Moss 1983). In the Harpirhynchinae birds, the skin has undergone significant evolu- the progressive characters are represented in adults, tionary changes comparable to that in of mam- whilst in Harpypalpinae they are developed in im- mals. This probably allowed some cheyletoid matures. Relationships of Passeriformes with other mites of the family Harpirhynchidae to transfer to higher Neoaves are not clear (Kurochkin 1993). It the intradermal parasitism in capsules that is simi- is possible that this order represents some earlier lar to that developed by representatives of Psorer- separated branch. Certain parasitological data sup- gatidae (Literak et al. 2005). port the hypothesis of the early origin of Passeri- Family Harpirhynchidae. This family in- formes. Thus, the rather archaic representatives of cludes three subfamilies, two subfamilies, Harpi- the families Rhinonyssidae (Mesostigmata) and rhynchinae and Harpypalpinae are represented by Ereynetidae (Prostigmata) are associated with Pas- bird parasites and the subfamily Ophioptinae — seriformes (Moss 1979). The feather mite family by ectoparasites of snakes from the superfamily Proctophyllodidae, mostly restricted to the passer- Colubroidea. The monophyly of this family is ines, is one of most evolved feather mite families strongly approved (Bochkov 2008), and hypothe- but is also characterized by certain archaic features sis about the independent origin of the subfamilies (Mironov 1998). If one admits the hypothesis of Harpirhynchinae and Harpypalpinae from cheyle- earlier origin of Passeriformes the separate phylo- toid-like ancestors, proposed by Moss (1979), is genetic position of the Harpypalpinae could be eas- groundless. ily explained by their coevolution with passerines. Mites of the subfamily Harpirhynchinae are The subfamily Ophioptinae is a sister lineage known exclusively from Neognathae (Galloanser- to the Harpypalpinae (Bochkov et al. 1999). It was ae and Neoaves). Mites of the most archaic genus hypothesized that harpypalpine-like ancestor of Harpirhynchoides are known from 14 bird orders Ophioptinae switched to parasitism on snakes (Fain 1994b). Mites of the genera Neharpirhynchus from passerines. The possibility of this host shift and Metharpirhynchus parasitize passerines. In was originally proposed by Kethley (after: Lombert addition, the first genus is associated with hum- and Moss 1983). Certain snakes feed on nestlings ming birds (Apodiformes: Trochilidae), and the and adult birds. Most of this prey is small passer- second genus — with Picidae (Piciformes) (Fain ine birds. The subfamily Ophioptinae is associated 1995). Mites of the monotypic genus Anharpi- exclusively with Colubridae and Elapidae (Boch- rhynchus are associated with passerine and pici- kov et al. 1999). These two snake families are form birds. Another six genera are represented by closely related and represent a group of higher a few species which are associated with hosts of snakes (Rieppel 1988). one order. Mites of two genera, Harpirhynchus, Family Cloacaridae (Pneumophaginae). and Trichorhynchiella are associated with passer- The record of Pneumophagus bubonis from the ines; mites of the genera Cypsharpirhynchus and lungs of a single specimen of Bubo virginianus by Harpyrhynchiella are known from Apodidae Fain and Smiley (1989) is still enigmatic. As dis- (Apodiformes); the genus Perharpirhynchus is as- cussed above, these mites exhibiting general cloa- sociated with hosts of the order Charadriiformes; carid morphology are clearly distinguished from the mites of the genus Ralliharpirhynchus are the Cloacarinae, and their placement into a sepa- known only from Rallidae (Gruiformes) (Fain rate subfamily has a strong morphological sup- 1995). port. The distinctions between pneumophagines Mites of the subfamily Harpypalpinae are and cloacarines are stronger than these between known from hosts of 11 families, all of them be- cloacarines parasitizing two chelonian orders long to higher passerines. Pleurodyra and Cryptodyra. Such morphological

91 M. Skoracki, S.A. Zabludovskaya, A.V. Bochkov distinctions suggest a long history of pneumaphag- The tribe Trispeleognathini includes the re- ine parasitism on non-turtle hosts. We believe maining 14 genera of bird-associated speleog- therefore, that this record is not likely to be the nathines. All of them parasitize non passerine result of a host switch from some as yet unexam- birds, with one exception — Speleognathopsis ined turtle species that happened within the onychognathi (Fain, 1964). Most genera are as- lifespan of that individual bird. On the other hand, sociated with hosts of a single order: the genus the knowledge of the diversity and host distribu- Trispeleognathus is associated with Columbi- tion of pneumophagines is so far limited to this formes, Aureliania — with Strigiformes, Oph- single record. Therefore we can not hypothesize thalmognathus — with Pelecaniformes, Psitta- whether this parasitism has an ancient origin, and boydaia — with Psittaciformes, Neastrida — with birds have inherited cloacarids from their common Gruiformes, Neoboydaia — mostly with Char- ancestor with turtles or, that seems much more adriiformes and one species is recorded from probable, it was the result of an ancient host shift Podicipediformes, Pterniboydaia — with Galli- caused by predation by some birds on live or dead formes, Meropiboydaya — with Coraciiformes, turtles (Bochkov and OConnor 2008). and Picinyssus — with Piciformes. Five genera Family Ereynetidae (Speleognathinae). are, however, associated with hosts of various or- This family is represented by three subfamilies. ders: Astrida — with Caprimulgiformes, Coracii- Mites of the subfamily Ereynetinae are predators formes, and Strigiformes, Metaboydaia — with inhabiting humid microhabitats or facultative par- Coraciiformes and Pelecaniformes, Ralliboydaia asites of mollusc lungs (snails). Mites of the sub- — Gruiformes and Cuculiformes, Phoenicopteri- family Lawrencarinae are endoparasites of amphi­ ella — with Anseriformes and Phoenicopteri- bian nasal cavities. The subfamily Speleo­ ­gnathinae formes, and Speleognathopsis — with Acipitri- consists of nasal endoparasites of mammals and formes, Galliformes, Charadriiformes, and birds (Andre and Fain 2000). It could be suggested Passeriformes. that ereynetine-like ancestor of the subfamily Biology Lawrencarinae switched to parasitism on amphib- ians from their prey — snails or directly from the Data about the biology of bird-associated soil. In the absence of phylogenetic hypothesis for prostigmatans are very fragmental and far from speleognathine mites, two hypotheses of their host complete; almost nothing is known about the biol- relationships could be proposed. According to the ogy of Pneumophaginae and Speleognathinae. first hypothesis, amniotic vertebrates received All prostigmatans permanently associated these mites from amphibians or directly from the with birds possess piercing chelicerae and feed on soil. Monophyly of the subfamily Speleognathi- live host (or prey) tissues. The exception are males nae is well grounded, therefore, such transition of Pneumophagus bubonis, their chelicerae are should have happened on one of these two groups. lost but oral opening is still present (Bochkov and This host switch must be very ancient because the OConnor 2008). There are two types of food spe- wide distribution of these mites on avian and cialization in skin-inhabiting prostigmatans: lym- mammal orders. An alternative hypothesis sug- phophagy (most prostigmatans) and haematophagy gests the initial parasitism of these mites on a (Bakericheyla) (Akimov and Gorgol 1990). common ancestor of Anamnia and Amniota and Females of these mites lay eggs on (in) the extinction of these mites on modern reptiles. host body. Normally their life-cycle includes ac- The subfamily Speleognathinae is separated tive larva, two nymphal stages and adults, females into five tribes, two of them are associated with and males; no cases of parthenogenesis were re- birds and other three — with mammals (Fain corded. The immatures of Pneumophagines, how- 1985). ever, are still unknown (Bochkov and OConnor Two genera of the tribe Boydaiini, Boydaia 2008) and all nymphal stages of Speleognathinae and Coboydaia are mostly associated with passer- are calyptostatic (Andre and Fain 2000). The life- ines. Among 40 species of the first genus, only cycle of prostigmatans studied in this relationship four species are, probably, secondary associated is short. In free-living cheyletids (Cheyletus) the with non passerine hosts — Caprimulgiformes, time of life-cycle critically depends from tempera- Galliformes, Psittaciformes, and Falconiformes. ture and takes from 10 to 80 days (Akomov and Among five Coboydaia spp., only one is associat- Gorgol 1990). In syringophilids, it takes about 40 ed with humming birds. days (Kethley 1971). In symbiotic cheyletids and

92 A review of Prostigmata permanently associated with birds harpirhynchids these characteristics should prob- mesticated birds (Rebrassier and Martin 1932; ably be similar. Schwabe 1956; Hwang 1959; Gritschenko 1973; Prostigmatans, being permanent symbionts, Schmaschke et al. 2003; Principato et al. 2005) colonize new hosts by two main ways — from Skoracki (2011) found no evidences that quill mites parents to nestlings and in the time of contacts be- induce the pathological changes in the structure of tween adults (particularly during molt of gregari- skin in wild birds, even during heavy infestations. ous birds or at breeding time). The first way is es- Casto’s (1974) observations also indicate that the pecially characteristic for syringophilids (Skoracki exit channels and many small holes produced dur- 2011). Is worth to note that in syringophilid mites ing feeding activity of syringophilids, do not seem only fertilized females disperse from one host to to weaken the structure of the feather. another, whereas the males never leave the feather The role of prostigmatan mites as vectors of quills (Kethley 1971). bird diseases is almost unstudied. It is only known, The population ecology of bird-associated that syringophilids could be potential vectors of prostigmatans is weakly studied. They are charac- bacterium Anaplasma phagocytophilum — an ob- terized by relatively low prevalence index (IP). ligate intracellular pathogen (Skoracki et al. For parasitic cheyletids (Bakericheyla chanayi 2006). and Ornithocheyletia sp.) it was estimated for dif- ACKNOWLEDGEMENTS ferent seasons and hosts as 0.4–8% (Akimov and Gorgol 990). According to Kethley (1971) and We thank Dr S.V. Mironov (Zoological Insti- Skoracki (2011) , IP in syringophilids is very low tute RAS, St. Petersburg) for critical review of the for the wild, non-social, and separately nesting MS. The English of this MS was improved by hosts, for example, IP for Lanius excubitor para- B. Nattress. We thank Dr. I. Literak for the photo sitized by Syringophiloidus weiszi is 3.5% of harpirhynchid cyst. This research was support- (N=508). In the same time IP for Hirundo rustica ed by the Russian Foundation for Basic Research infected with Syringophiloidus hirundus is 17% (Grant No 10-04-00160_a) and by the NCN grant (N=208). The data about syringophilid IP on wild (2011/01/B/NZ8/01749). In the frames of this birds has been recently summarized and discussed project visit of AVB to the Adam Mickiewicz Uni- by Skoracki et al. (2010). The highest IP was versity, Poznan, Poland was supported by the spe- shown for social and domestic birds, e.g. IP for cial grant from administration of this university. Passer domesticus infested by Syringophiloidus REFERENCES minor is 82% (N=492), or IP for Gallus gallus do- Akimov, A.I. and Gorgol, V.T. 1999. Khishchnye i mesticus is 75% (N=1.500). In various species of paraziticheskie kleshchi cheyletidy [Predaceous the genus Neharpyrhynchus demonstrated on the and parasitic mites-cheyletids]. Naukova Dumka, European birds 2–14% IPs (Martinu et al. 2008) Kiev, 120 pp. [In Russian] and 15–21% on birds in Peru (Literak et al. 2012); Amaral, V. 1963. The occurenceof the nasal mite mites of the genus Harpirhynchus dusbabeki from Trispeleognathus striatus (Crossley, 1952), Fain, Panurus biarmicus in Europe showed IP 7.4% 1962 (Ereynetidae Oudemans, 1931) from Lepto- (N=378) (Literak et al. 2005). tila verreauxi (Aves: Columbiform). Arquivos do The damage caused by prostigmatans to their Instituto Biologico Sao Paulo, 30: 91. Andre, H.M. and Fain, A. 2000. Phylogeny, ontogeny avian hosts is markedly varied. Most of these mites and adaptive radiation in the superfamily do not cause a huge loss of bird condition. Some Tydeoidea (Acari: Actinedida), with a reappraisal cyst-forming harpirhynchids are considered as of morphological characters. Zoological Journal harmful parasites (Moss 1979). At the same time of the Linnean Society, 130: 405–448. Henry et al. (2004) did not find a markedly nega- Atyeo, W.T., Kethley, J.B. and Perez, T.M. 1984. Pae- tive impact of the lesions by harpirhynchids on the domorphosis in Metacheyletia (Acari: Cheyleti- health status of birds infected. According to Lit- dae), with the description of a new species. Jour- erak et al. (2005), the traumatic effect from harpi- nal of Medical Entomology, 21:125–131. rhynchid cysts was also not too serious and it Baker, E.W. 1949. A review of the mites of the family Cheyletidae in the United States National Muse- seems that birds with cysts survived for a long pe- um. Proceedings of the United States National riod. Museum, 99: 267–320. In contrast to some reports about the clinical Banks, N. 1905. Descriptions of some new mites. Pro- signs of feather’s picking caused by syringophilid ceedings of entomological Society of Washington, mites (mostly Syringophilus bipectinatus) in do- 7: 133–142.

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(Acari:Cheyletidae), a new species land. Parasitological Research, 108: 715–722. of parasitic mite from the bank swallow from Kh- Bochkov, A.V. and Skoracki, M. 2011. A new cheyletid irgizia. Acarina, 6: 35–36. mite Metacheyletia ngaii n. sp. (Acariformes: Chirov, P.A. and Kravtsova, N.T. 1995. [A new genus Cheyletidae) from quills of Corythaixodes leuco- and new species of mites of the family Syringo- gaster (Musophagidae) from Tanzania. Acarolo- philidae]. Parazitologiya, 29: 370–379. [In Rus- gia, 51: 93–97. sian with English summary] Bochkov, A.V. and Skoracki, M. 2012. New and poorly Clark, G.M. 1958. One new previously unreported spe- known predaceous cheyletids (Acariformes: Chey- cies of nasal mite (Acarina, Speleognathidae) from letidae) from feather quills. International Journal Noth American birds. Proceedings of the Helmint- of Acarology, 38: 486–496. hological Society of Washington, 25: 78–86. Bochkov, A.V., Skoracki, M., Hendricks, S.A., and Clark, G.M. 1960. 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R.A.N., Zoologichesky institute, logichesky institut. 65pp. [In Russian] St.Petersburg. 65 pp. [In Russian] Landmann, L. 1986. The skin of reptiles: epidermis and Mironov, S.V. and Bochkov, A.V. 2009. [Modern hy- dermis. In: Bereiter-Hahn J., Matolsty A.G., Rich- potheses about macrophylogeny of acariform

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Some mite parasites of Australian lung naturwissenschaftlich. Verein, Bremen, 10: birds. Proceedings of the Linnean Society of New 205–240. South Wales, 90: 190–217. Porter, J.C. and Strandtmann, R. 1952. Nasal mites of Moss, W.W., Oliver, J.H. Jr. and Nelson, B.C. 1968. the English sparrow. Texas Journal of Science, 4: Karyotypes and developmental stages of Harpy- 393–399. rhynchus novoplumaris sp. n. (Cheyletoidea: Har- Proctor, H.C. 2003. Feather mites (Acari: Astigmata): pyrhynchidae), a parasite of North American birds. Ecology, behavior and evolution. Annual Review Journal of Parasitology, 54: 377–392. of Entomology, 48: 185–209. Moss, W.W. and Wojcik, J.F. 1978. Numerical taxo- Principato, M., Lisi, F., Moretta, I., Samra, N., and Puc- nomic studies of the mite family Harpyrhynchidae cetti, F. 2005. The alterations of plumage of para- (Acari: Cheylctoidea): the higher taxa. Annals of sitic origin. 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Inter- Syringophilidae) parasitic on birds (Aves) in Eng- nationalen Symposiums über die Erkrankungen land. Zootaxa, 2133: 49–54. der Zoo- und Wildtiere, 41: 127–133. Nitzsch, C.L. 1818. Acarina. In: “Ersch & Gruber”. Schwabe, O. 1956. A quill mite of poultry—a case re- Allgemeine Encyclopedic Wissenschaft, 1: 250– port. Journal of the American Veterinary & Medi- 251. cal Association, 129: 481–482. Oudemans, A.C. 1904. Acarologische Aanteckeningen. Sikora, B., Fajfer, M. and Skoracki, M. 2011. Quill XIII. Entomologische Berichten Netherlands En- mites (Acari: Syringophilidae) from mimid birds tomologie, 1: 169–174. (Aves: Mimidae). Zootaxa, 3027: 29–38. Oudemans, A.C. 1905. Acarologische Aanteekingen. Sikora, B., Fajfer, M., Kavetska, K., and Skoracki, M. XV. Entomol. Bericht. Nederl 1: 207–210. 2012. Three new species of quill mites (Acari: Sy- Oudemans, A.C. 1906. Revision des Cheletines. Me- ringophilidae) parasitizing the wrens (Aves: Trog- moires de la Sociéte Zoologique de France, 19: lodytidae). Zootaxa, 3167: 57–65. 36–144. Smiley, R.L. 1970. A review of the family Cheyletiel- Oudemans, A.C. 1939. Über Phthiriasis und über ihren lidae (Acarina). Annals of the Entomological Soci- Erzeuger, Harpyrynchus tabescentium (Berthold ety of America, 63: 1056–1078. 1845). Zeitschrift für Parasitenkunde, 11, 145– Sixl, W. 1972. Nasale Milben einheimischer Vogel und 198. Kleinsauger. II Mitteilungen Naturwissenschaftli- Pence, D.B. 1973a. The nasal mites of birds from Loui- cher Verein fur Steirmark, 102: 173–174. siana VII. The Ereynetidae (Speleognathinae). Skoracki, M. 1999. New genus and species of Syringo- Journal of Parasitology, 59: 364–368. philidae (Acari: Prostigmata) from Eurasian Reed- Pence, D.B. 1973b. Nasal mites of birds of Louisiana Warbler, Acrocephalus scirpaceus (Sylviidae: VIII. Additional Records. Journal of Parasitology, Passeriformes). Genus, 10: 155–162. 59: 878–880. Skoracki, M. 2002a. Three new species of quill mites Pence, D.B. 1975. Keys, Species and Host list, and bib- of the genus Aulonastus Kethley, 1970 (Acari, liography for nasal Mites of North American birds Prostigmata, Syringophilidae) from passerine (Acarina: Rhinonyssidae, Turbinoptidae, Speleog- birds. Acta Parasitologica, 47: 300–305.

100 A review of Prostigmata permanently associated with birds

Skoracki, M. 2002b. Three new species of the ectopara- Skoracki, M., Bochkov, A.V. and Sikora, B. 2004a. sitic mites of the genus Syringophiloidus Kethley, New records of the cheyletoid mites (Acari: Chey- 1970 (Acari: Syringophilidae) from passeriform letoidea) from passeriform birds in Poland. Bel- birds from Slovakia. Folia Parasitologica, 49: gian Journal of Entomology, 6: 83–90. 305–313. Skoracki, M., Bochkov, A.V. and Wauthy, G. 2004b. Skoracki, M. 2004a. A review of quill mites of the ge- Revision of the quill mites of the genus Picobia nus Syringophiloidus Kethley, 1970 (Acari: Pros- Haller, 1878 (Acari: Syringophilidae) with notes tigmata: Syringophilidae) parasitizing quills of on their host-parasite relationships. Insect System- passeriform birds, with descriptions of four new atics and Evolution, 35: 155–176. species. Genus, 15: 281–300. Skoracki, M. and Dabert, J. 1999a. A new species of the Skoracki, M. 2004b. Quill mites of the genus Syringo- genus Syringophilopsis Kethley, 1970 (Acari: philopsis (Acari, Syringophilidae) from passeri- Prostigmata: Syringophilidae) from the Tree Pipit form birds of Poland with descriptions of five new Anthus trivialis (Passeriformes: Motacillidae). Ac- species. Acta Parasitologica, 49: 45–62. arina, 7: 89–92. Skoracki, M. 2004c. New data on systematics of the Skoracki, M. and Dabert, J. 1999b. New species of Sy- quill mites of the genus Torotrogla Kethley, 1970 ringophilidae (Acari: Prostigmata) from African (Acari: Syringophilidae). Belgian Journal of En- birds. Genus, 10: 523–527. tomology, 6: 303–314. Skoracki, M. and Dabert, J. 2000. Syringophilopsis al- Skoracki, M. 2005a. A new genus of ectoparasitic mites bicollisi, a new species of the quill mite of the of the family Syringophilidae (Acari: Cheyletoidea) family Syringophilidae Lavoipierre, 1953 (Acari: from the Treeswifts (Apodiformes: Hemiprocni- Prostigmata). Acarina, 8: 59–63. dae). Acta Parasitologica, 50: 336–343. Skoracki, M. and Dabert, J. 2001a. Two new species of Skoracki, M. 2005b. A new genus and species of quill the parasitic mites of the genus Aulobia Kethley, mites (Acari: Syringophilidae) from a king bird- 1970 (Acari, Syringophilidae) from African birds. of-paradise Cicinnurus regius (L.). Systematic Acta Parasitologica, 46: 208–215. Parasitology, 60: 155–158. Skoracki, M. and Dabert, J. 2001b. The quill mites of Skoracki, M. 2005c. A review of the quill mites (Acari: the genus Syringophilopsis Kethley, 1970 (Acari, Syringophilidae) parasitizing parrots (Aves: Psitt- Syringophilidae) from African birds. Acarina, 9: aciformes) with description of three new species. 105–112. Acarina, 13: 127–136. Skoracki, M. and Dabert, J. 2002. A review of parasitic Skoracki, M. 2008a. A new genus of syringophilid mites of the family Syringophilidae (Acari, Pros- mites (Acari: Cheyletoidea: Syringophilidae) from tigmata) from African birds, with descriptions of cuculiform birds (Aves: Cuculiformes). Folia four new species. Acta Parasitologica, 47: 137– Parasitologica, 55: 155–158. 146. Skoracki, M. 2008b. Crotophagisyringophilus n. g. Skoracki, M., Dabert, J. and Ehrnsberger, R. 2000. A (Acari: Syringophilidae), a new genus of parasitic new quill mite (Acari: Syringophilidae) from the quill mites. Systematic Parasitology, 71: 75–80. Blackbird. Osnabrucker Naturwissenschaftliche Skoracki, M. 2008c. Description of two new quill mite Mitteilungen, 26: 191–198. species (Acari: Syringophilidae). Acarina, 16: Skoracki, M., Dabert, J. and Schmaschke, R. 2006. Ob- 45–50. servations on the quill mites (Acari: Syringophili- Skoracki, M. 2011. Quill mites (Acari: Syringophili- dae) from charadriiform birds. Zootaxa, 1156: dae) of the Palaearctic region. Zootaxa, 2840: 51–64. 1–415. Skoracki, M., Flannery, M.E. and Spicer, G.S. 2008. Skoracki, M., Antczak, M., Riegert, J., Fainova, D., and Quill mites of the genus Syringophilopsis Kethley, Mikes, V. 2009a. New species and new records of 1970 (Acari: Syringophilidae) from North Ameri- quill mites (Acari: Syringophilidae) inhabiting Af- can birds. Folia Parasitologica, 55: 291–300. rican passerines (Aves: Passeriformes). Acta Zoo- Skoracki, M., Flannery, M.E. and Spicer, G.S. 2009b. logica Academiae Scientiarum Hungaricae, 55: New data on systematics of the quill mites of the 123–137. genus Syringophiloidus Kethley, 1970 (Acari: Sy- Skoracki, M. and Bochkov, A.V. 2002. A new quill mite ringophilidae) from North American birds. Acta species Bubophilus asiobius sp. n. (Acari: Syrin- Parasitologica, 54: 64–72. gophilidae) from the Long-eared Owl Asio otus Skoracki, M. and Glowska, E. 2008a. Quill mites (Ac- (Strigiformes: Strigidae). Genus, 13: 141–144. ari: Syringophilidae) associated with columbiform Skoracki, M. and Bochkov, A.V. 2010. Quill mites from birds. Genus, 19: 151–160. Kazakhstan. Zootaxa, 2546: 52–68. Skoracki, M. and Glowska, E. 2008b. Two new species Skoracki, M., Bochkov, A. V. and OConnor, B. M. of the quill mite genus Aulobia Kethley, 1970 (Ac- 2011. Notes on quill mites (Acariformes, Syringo- ari, Syringophilidae) associated with sunbirds philidae) from aquatic birds in North America. (Passeriformes, Nectariniidae). Acta Parasitologi- Acta Parasitologica, 56: 325–330. ca, 53: 179–185.

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Skoracki, M. and Glowska, E. 2008c. Two new species Skoracki, M. and Magowski, W. 2001. Two new spe- of the genus Picobia Haller (Acari: Syringophili- cies of quill mites of the genus Picobia (Acari: dae) from Australian and Indonesian passeriform Prostigmata: Syringophilidae) from passerine birds. New Zealand Journal of Zoology, 35: 281– birds of Poland. Acarina, 9: 113–119. 286. Skoracki, M., Magowski, W. and Dabert, J. 2001c. Pi- Skoracki, M., Glowska, E. and Sikora, B. 2008. Four cobia polonica sp. n. (Acari: Prostigmata: Syrin- new species of the quill mite genus Picobia Haller gophilidae), a new species of quill mite from the (Acari: Syringophilidae) parasitizing birds in the domestic hen, Gallus gallus domesticus (Aves: Australian region. Zootaxa, 1961: 58–68. Phasianidae). Folia Parasitologica, 48: 154–158. Skoracki, M. and Hebda, G. 2004. Quill mites (Acari: Skoracki, M., Michalik, J., Skotarczak, B., Rymaszew- Syringophilidae) from Aegithalos caudatus (Pas- ska, A., Sikora, B., Hofman, T., Wodecka, B. and seriformes: Aegithalidae). Zootaxa, 691: 1–6. Sawczuk, M. 2006. First detection of Anaplasma Skoracki, M., Hendricks, S.A. and Spicer, G.S. 2010a. phagocytophilum in quill mites (Acari: Syringo- Systematics of the ectoparasitic quill mites of the philidae) parasitizing passerine birds in Poland. genus Aulobia Kethley, 1970 (Acari: Syringophil- Microbes and Infection: 8, 303–307. idae) with the description of a new species. Zoot- Skoracki, M., Møller, A.P. and Tryjanowski, P. 2003. A axa, 2399: 31–41. new species of parasitic mites of the genus Syrin- Skoracki, M., Hendricks, S.A. and Spicer, G.S. 2010b. gophiloidus Kethley, 1970 (Acari: Syringophili- Four new species of Aulonastus Kethley, 1970 (Ac- dae) from the Barn Swallow Hirundo rustica Lin- ari: Syringophilidae) from North American passer- naeus, 1758. Parasite, 10: 17–20. ines. Systematic Parasitology, 76: 131–144. Skoracki, M. and OConnor, B. 2010. New taxa of quill Skoracki, M., Hendricks, S.A. and Spicer, G.S. 2010c. mites (Acari: Cheyletoidea: Syringophilidae). New species of parasitic quill mites of the genus Zootaxa, 2341: 1–32. Picobia (Acari: Syringophilidae: Picobiinae) from Skoracki, M. and Sikora, B. 2002. New ectoparasitic North American birds. Journal of Medical Ento- mites of the family Syringophilidae (Acari: Pros- mology, 47: 727–742. tigmata: Cheyletoidea) associated with birds from Skoracki, M., Hendricks, S.A. and Spicer, G.S. 2011a. Argentina. Zootaxa, 27: 1–8. Systematics of the genus Syringophilopsis Keth- Skoracki, M. and Sikora, B. 2003. Quill mites (Acari: ley, 1970 (Acari: Prostigmata: Syringophilidae) Prostigmata: Syringophilidae) from African pas- with description of three new species from North seriform birds. Zootaxa, 129: 1–10. American passerines. Zootaxa, 2793: 1–22. Skoracki, M. and Sikora, B. 2004a. Tinamiphilopsis el- Skoracki, M., Hromada, M. and Kuczynski, L. 2001a. egans gen. nov. et sp. nov., a first record of the Torotrogla lullulae, a new species of the family quill mites (Acari, Syringophilidae) from tinamou Syringophilidae Lavoipierre, 1953 (Acari: Pros- birds (Tinamiformes, Tinamidae). Acta Parasito- tigmata: Cheyletoidea). Genus, 12: 87–92. logica, 49: 348–352. Skoracki, M., Hromada, M. and Tryjanowski, P. 2001b. Skoracki, M. and Sikora, B. 2004b. A new genus and Description of a new species of quill mite Syringo- four new species of quill mites (Acari: Prostig- philoidus weiszii sp. n. (Acari, Prostigmata, Syrin- mata: Syringophilidae) from phasianid birds (Gal- gophilidae) from Great Grey Shrike Lanius excu- liformes: Phasianidae). Parasite, 11: 379–386. bitor. Acta Parasitologica, 46: 30–34. Skoracki, M. and Sikora, B. 2005. Neosyringophilop- Skoracki, M., Hromada, M. and Wamiti, W. 2011b. A sis, a new genus of the subfamily Syringophilinae new species and new host records of syringophilid (Acari: Syringophilidae). Zootaxa, 1052: 21–28. mites (Acari: Syringophilidae) from passerines Skoracki, M. and Sikora, B. 2008. Blaszakia, a new ge- from Kenya. Zootaxa, 2922: 34–40. nus of quill mites (Acari: Syringophilidae) parasit- Skoracki, M., Kavetska, K., Ozminski, M., and Sikora, izing Turacos (Aves: Musophagiformes). Annales B. 2012a. The morphology of juvenile stages of Zoologici, 58: 327–332. Bubophilus aluconis Nattress and Skoracki, 2009 Skoracki, M. and Sikora, B. 2011. Quill mites (Acari: (Acari: Syringophilidae) with a description of a Syringophilidae) associated with galliform birds male. Annals of Parasitology, 58: 37–48. (Aves: Galliformes). Zootaxa, 2966: 13–30. Skoracki, M. and Kiljan, G. 2002. Picobia paludicola Skoracki, M., Sikora, B. and Ozminski, M. 2012b. A sp. n. a new species of quill mite (Acari: Prostig- new quill mite species (Acari: Syringophilidae) mata: Syringophilidae) from the Aquatic Warbler parasitising tinamous (Aves: Tinamiformes). Sys- Acrocephalus paludicola (Passeriformes: Sylvii- tematic Parasitology, 81: 109–113. dae). Zootaxa, 91: 1–6. Skoracki, M. and Solarczyk, P. 2012. New picobiin Skoracki, M., Lontkowski, J. and Stawarczyk, T. 2010d. mites (Acari: Syringophilidae: Picobiinae) associ- New taxa of the parasitic quill mites associated ated with woodcreeper birds (Passeriformes: Den- with accipitrid birds indicating close relationship drocolaptidae). Zootaxa, 3406: 59–66. of falconid birds to Psittaci-Columbi clade. Jour- Skoracki, M., Tryjanowski, P. and Hromada, M. 2002. nal of Natural History, 44: 1203–1214. Two new species of the genus Syringophilopsis

102 A review of Prostigmata permanently associated with birds

Kethley, 1970 (Acari: Syringophilidae) parasitiz- lished by the Zoological Institute of the Academy ing quills of true shrikes (Aves: Laniidae). Para- of Sciences of the USSR], 101, 432 pp. [In Rus- site, 9: 11–16. sian] Smiley, R.L. 1977. Further studies on the family Chey- Walter, D.E., Lindquist, E.E, Smith, I.M. Cook, D.R., letiellidae (Acarina). Acarologia, 19: 225–241. and Krantz, G.W. 2009. Order Trombidiformes. Spicer, G.S. 1977. New host records for some nasal In: Krantz G.W. and Walter, D.E. (Eds). A manual bird mites (Acari: Rhinonyssinae, Speleognathi- of Acarology. 3rd Ed. Texas Tech University, Lub- nae). Journal of Medical Entomology, 14: 498. bock, Texas, 233–420. Spicer, G.S. 1987. Prevalence and host-parasite list of Womersley, H. 1936. On a new family of Acarina with some nasal mites from birds (Acarina: Rhininys- description of a new genus and species. Annals sidae, Speleognathidae). Journal of Parasitology, and Magazine of Natural History, 18: 312–315. 73: 259–264. Womersley, H. 1941. Notes on the Cheyletidae (Acari- Summers, F.M. and Price, D.W. 1970. Review of the na, Trombidoidea) of Australia and New Zealand, mite family Cheyletidae. University of California with descriptions of new species. Records of the publications in Entomology, 61: 1–152. South Australian Museum, 7: 51–64. Volgin, V.I. 1964. [On the taxonomy of predatory mites Zabludovskaya, S.A. 1998. Ereynetid mites (Acari- of the family Cheyletidae VI. The genus Orni- formes: Tydeoidea: Ereynetidae) paraziting the thocheyletia Volgin gen. nov.]. Zoologischesky respiratory ways of birds (Aves: Anseriformes, zhurnal, 43: 28–36. [In Russian] Charadriiformes, Passeriformes) of the European Volgin, V.I. 1969. Kleshschi semeystva Cheyletidae — Siberian subregion of the Palearctic. Acarina, mirovoy fauny [Mites of the family Cheyletidae 6: 47–50. of the world]. Opredeliteli po faune SSSR, izda- Zabludovskaya, S.A. 2002. On studying of the fauna of vaemie Zoologicheskim institutom Akademii Ereynetid mites (Ereynetidae, Trombidiformes). nauk SSSR [Keys to the fauna of the USSR pub- Vestnik zoologii, 36: 93–94.

Table 1. Summary of taxa number, distribution and host associations of Prostigmata permanently associated with birds

Family Type of Species n Genera n Host orders (subfamily or tribe) symbiosis Family Cheyletidae Leach, 1815 Apodiformes, Charadriiformes, Predators and Tribe Cheletosomatini Volgin, 1969 23 6 Cucu­liformes, Galliformes, Gruiformes, parasites Piciformes (Picocheyletus) Apodiformes, Columbiformes, Cora­ciiformes, Cuculiformes, Galli- Tribe Ornithocheyletiini Volgin, 1969 54 4 Parasites formes, Passeriformes, Pelecaniformes, Piciformes, Psittaciformes, Strigiformes Cuculiformes, Passeriformes, Psittaci­ ­ Tribe Metacheyletiini Fain, 1980 5 1 ?Parasites formes Family Syringophilidae Lavoipierre, 1953 Accipitriformes, Anseriformes, Apo­di­formes, Charadriiformes, Ciconiiformes, Columbiformes, Coraciiformes, Cuculiformes, Galbuli- Subfamily Syringophilinae formes, Galliformes, Gruiformes, 234 48 Parasites Lavoipierre, 1953 Passeriformes, Pelecaniformes, Phoenicopteriformes, Piciformes, Procellariiformes, Psittaciformes, Pte­ro­cliformes, Strigiformes, Suli- formes, Tinamiformes Charadriiformes, Columbiformes, Subbfamily Picobiinae Coraciiformes, Cuculiformes, Galli- 44 5 Parasites Johnston et Kethley, 1973 formes, Passeriformes, Piciformes, Psittaciformes, Pterocliformes

103 M. Skoracki, S.A. Zabludovskaya, A.V. Bochkov

Family Harpirhynchidae Dubinin, 1956 Accipitriformes, Anseriformes, Apodiformes, Charadriiformes, Ciconiiformes, Coliiformes, Columbi- Subfamily Harpirhynchinae 65 10 formes, Cuculiformes, Galliformes, Parasites Dubinin, 1956 Gruiformes, Passeriformes, Pelecani- formes, Piciformes, Procellariiformes, Psittaciformes, Strigiformes Subfamily Harpypalpinae Fain, 1972 10 2 Passeriformes Parasites Family Cloacaridae Camin, Moss, Oliver et Singer, 1967 Subfamily Pneumophaginae 1 1 Strigiformes Parasites Fain et Smiley, 1989 Family Ereynetidae Oudemans, 1931 Subfamily Speleognathinae Fain, 1957 Apodiformes,Caprimulgiformes, Tribe Boydaiini Fain, 1985 45 2 Falconiformes, Passeriformes, Psittaci- Parasites formes Accipitriformes, Anseriformes, Caprimulgiformes, Charadriiformes, Columbiformes, Coraciiformes, Cuculiformes, Galliformes, Gruiformes, Tribe Trispeleognathini Fain, 1985 28 14 Parasites Passeriformes, Pelecaniformes, Phoenicopteriformes, Piciformes, Podicipediformes, Psittaciformes, Strigiformes

Table 2. Setation of gnathosoma

Family/Subfamily Setae Cheyletidae elc. p, n, ao1, ao2 Syringophilinae elc. p, n, ao1, ao2 Picobiinae elc. p, n, ao1, ao2 Harpirhynchinae elc. p, m, n Harpypalpinae elc. p, m, n Cloacaridae (Pneumophaginae) – Ereynetidae (Speleognathinae) n (sc1), m (sc2)

Table 3. Setation of palps

Species Femur Genu Tibia Tarsus Cheyletidae d, v’,v” d, l” d, lT, l’ acm, sul, ul’, ul”, ω Syringophlinae d, v’,v” d, l” d, lT, l’ acm, sul, ul’, ul”, ω Picobiinae d, v’,v” d, l” d, lT, l’ acm, sul, ul’, ul”, ω Harpirhynchinae d, v d, l” d, lT 1 seta Harpypalpinae d d, l” d, lT 1 seta Cloacaridae (Pneumophaginae) – – – – Ereynetidae (Speleognathinae) – – d d, l’, l”, ω

104 A review of Prostigmata permanently associated with birds

Table 4. Setation of idiosoma

Species Setae Cheyletidae scx, vi, ve, si, se, c1, c2, d1, d2, e1, e2, f1, f2, h1, h2, ps1–3, ag1–3 (2), g1–2 (2) Syringophilinae scx, vi, ve, si, se, c1, c2, d1, d2, e2, f1, f2, h1, h2, ps1–2, ag1–3 (3), g1–2 (2) Picobiinae scx, vi, ve, si, se, c1, c2, d1, d2, e1, f1, f2, h1, h2, ps1–2, ag1–3 (3), g1–2 (2) Harpirhynchinae scx, vi, ve, si, se, c2, h1, g1 (3) Harpypalpinae scx, vi, ve, si, se, c1, c2, d1, d2, e2, f1, f2, h1, ps1–3 (2), g1 (0) 1 pair of propodonotal alveoli (0); 1 pair of alveoli flanking anal-genital opening (0), 3 Cloacaridae (Pneumophaginae) pairs of opisthosomal spine like setae (1) Ereynetidae (Speleognathinae) vi, ve, si, se, c1, c2, d1, e1, f1, f2, h1, ag1–2 (2), g1–3 (3), ps1–2 (2), (eu) () — number of setae in male

Table 5. Setation of legs

Family Tarsus I Tarsus II Tarsus III Tarsus IV Cheyletidae ft,(tc), (p), a”, (u), vs, ω (tc), (p),(u), vs, ω (tc), (p),(u), vs, ω (M) (tc), (p),(u), vs, ω (M) Syringophilinae ft, (tc),(p), ω (a),(u), vs (tc),(p),(u), vs, ω (tc),(p),(u) (tc),(p),(u) Picobiinae ft, (tc),(p), ω (a),(u), vs (tc),(p),(u), vs, ω (tc),(p),(u) (tc),(p),(u) Harpirhynchinae (tc),(p), (a),(u), ω (tc), p”,(a),( u), ω 8 or less 8 or less Harpypalpinae (tc), p”, (a),(u), ω (tc), p”, (a), (u), ω (tc), (a),(u) (tc), (a),(u) Cloacaridae 5 9 12 12 (Pneumophaginae) Ereynetidae 12, ω 8 7 7 (Speleognathinae) Family Tibia I Tibia II Tibia III Tibia IV Cheyletidae d, (l), (v), φ d, l”, (v) d, l”, (v)+ φ (M) d, l”, (v) Syringophilinae d, (l), v, φ d, (l), v d, (l) d, (l) Picobiinae d, (l), v, φ d, (l), v d, (l) d, (l) Harpirhynchinae d, (l), (v) d, (l), (v) – – Harpypalpinae d, (l), (v) d, (l), (v) d, (v) d, (v) Cloacaridae 1 – – – (Pneumophaginae) Ereynetidae d, (l), (v) d, l, v d, l, v d, l, v (Speleognathinae) Family Genu I Genu II Genu III Genu IV Cheyletidae d, l’, σ d, l’ d, l’ d, l’ Syringophilinae d, l’, σ d, l’ l’ l’ Picobiinae d, l’, σ d, l’ l’ l’ Harpirhynchinae d, l’, (v) d, l’, (v) – – Harpypalpinae d, (l),( v) d, (l),(v) – – Cloacaridae – – – – (Pneumophaginae) Ereynetidae (d), (l) (d), (l) d (l) d, (l) (Speleognathinae) Family Femur I Femur II Femur III Femur IV Cheyletidae d, v d, v d, v d, v Syringophilinae d, v d, v d d Picobiinae d, v d, v – –

105 M. Skoracki, S.A. Zabludovskaya, A.V. Bochkov

Harpirhynchiane d, v d, v – – Harpypalpinae d, v d, v v v Cloacaridae – – – – (Pneumophaginae) Ereynetidae 7 (d), (v) d, (v) d (v) (Speleognathinae) Family Trochanter/ coxa I Trochanter/ coxa II Trochanter/ coxa III Trochanter/ coxa IV Cheyletidae v/1a, 1b, 1c v/2c l, v/3a,3b,3c v/4a, 4b, 4c Syringophilinae v/1a, 1b, 1c v/2c v/3a, 3b, 3c v/4b, 4c Picobiinae v/1a, 1b, 1c v/2c –/3a, 3b, 3c –/4b, 4c Harpirhynchinae v/1a, 1c v/– –/3a –/– Harpypalpinae v/1a, 1c v/2c v/3a, 3c v/4c Cloacaridae – – – – (Pneumophaginae) Ereynetidae l’/1a, 1b, 1c l’/2c –/3a, 3b –/4a, 4b (Speleognathinae) () — pair of setae; (M) — solenidion of male

Table 6. Distribution of Prostigmata permanently associated with birds among host orders

Mite family Host order (N of extant species) Cheyletidae Harpirhynchidae Syringophilidae Cloacaridae Ereynetidae Palaeognathae Struthioniformes Tinamiformes Neognathae Galloanserae Anseriformes Galliformes Neoaves Charadriiformes Gaviiformes Podicipediformes Procellariiformes Sphenisciformes Phaethontiformes Ciconiiformes Suliformes Pelecaniformes Phoenicopteriformes Accipitriformes Falconiformes Otidiformes Mesitornithiformes Cariamiformes Eurypygiformes Gruiformes Pterocliformes Columbiformes

106 A review of Prostigmata permanently associated with birds

Psittaciformes Cuculiformes Galbuliformes Strigiformes Caprimulgiformes Apodiformes Coraciiformes Piciformes Trogoniformes Coliiformes Passeriformes Bold — mites unknown

Table 7. Host-specificity among Prostigmata permanently associated with birds Number of mites specific to host order, family, genus and species respectively.

Host specificity Mite family (N of species) To species To genus To family To order To orders Cheyletidae (bird-associated, parasites — 59) 49 (83%) 1 (1.7%) 6 (10.2%) 1 (1.7%) 2 (3.4%) Cheyletidae (bird-associated, predators — 23) 15 (65.2%) 2 (8.7%) 2 (8.7%) 4 (17.4%) – Syringophilidae (278) 196 (70.5%) 43 (15.5%) 30 (10.8%) 6 (2.2%) 3 (1.1%) Harpirhynchidae (bird-associated — 75) 47 (62.7%) 7 (9.3%) 11 (14.7%) 9 (12%) 1 (1.3%) Cloacaridae (bird-associated — 1) 1 (100%) – – – – Ereynetidae (bird-associated — 73) 27 (37%) 8 (11%) 22 (30.1%) 14 (19.2%) 2 (2.7%)

107 Appendix

Appendix. List of Prostigmata species permanently parasitizing birds and their hosts Geographical references: The European Region does not include Turkey, which is considered here under the Asiatic Region. In the Australian Region, Oceania does not include Eastern Timor and the Indonesian islands, which are considered under the Asiatic Region.The South American Region includes Central America and islands of the Caribbean Sea. A few records containing in our opinion doubtfully determined mite species are not included. Bird systematic follows Clements et al. 2011.

Mite species Host species Host family Host order Locality References Cheyletidae Leach, 1815 Ornithocheyletini Volgin, 1969 Ornithocheyletia Volgin, 1964 Ornithocheyletia pinguis Turdus merula Turdidae Passeriformes Europe Berlese 1889 (Berlese, 1889) Ornithocheyletia aitkeni Turdus fumigatus Turdidae Passeriformes South America Fain 1972, 1981 Fain, 1972 Volgin 1964; Smiley 1977; Ornithocheyletia Europe, North Sturnus vulgaris Sturnidae Passeriformes Fain 1981; dubinini Volgin, 1984 America Skoracki et al. 2004 Ornithocheyletia eulabes European Zoo Gracula religiosa Sturnidae Passeriformes Fain 1981 Fain, 1881 (captive) Ornithocheyletia Lamprotornis Sturnidae Passeriformes Africa Fain 1972, 1981 lamprocolius Fain, 1972 chloropterus Ornithocheyletia wauthy Fain and Lammprocolius sp. Sturnidae Passeriformes Africa Fain et Bochkov, 2002 Bochkov 2002 Ornithocheyletia volgini Petrochelidon Smiley 1970; Hirundinidae Passeriformes North America Smiley, 1970 pyrrhonota Fain 1981 Ornithocheyletia Smiley 1970; Hirundo rustica Hirundinidae Passeriformes Europe lukoschusi Smiley, 1970 Fain 1981 Ornithocheyletia Chirov and mironovi Bochkov et Riparia riparia Hirundinidae Passeriformes Asia Bochkov 1998 Chirov, 1998 Ornithocheyletia Phylloscopus Bochkov et al. phylloscopi Bochkov, Sylviidae Passeriformes Europe trochilus 1994 Mironov et Gorgol, 1994 Ornithocheyletia European Zoo Leiothrix lutea Leiothrichidae Passeriformes Fain 1972, 1981 leiothrix Fain, 1972 (captive) Ornithocheyletia Garrulax European Zoo Leiothrichidae Passeriformes Fain 1972, 1981 garrulax Fain, 1972 leucolophus (captive) Ornithocheyletia lepidus Garrulax European Zoo Leiothrichidae Passeriformes Fain 1981 Fain, 1981 leucolophus (captive) Ornithocheyletia Uraeginthus European Zoo Estrildidae Passeriformes Fain 1972, 1981 granatina Fain, 1972 ianthinogaster (captive) Ornithocheyletia orioli Fain and Oriolus oriolus Oriolidae Passeriformes Africa Fain et Bochkov, 2003 Bochkov 2002 Pachyphantes Fain and “ Ploceidae Passeriformes Africa superciliosa Bochkov 2002 Fain and “ Lybius sp. Lybiidae Piciformes Africa Bochkov 2002 Fain and “ Indicator indicator Indicatoridae Piciformes Africa Bochkov 2002 Chrysococcyx Fain and “ Cuculidae Cuculiformes Africa klaasi Bochkov 2002

i1 M. Skoracki, S. Zabludovskaya and A.V. Bochkov

Ornithocheyletia Banks 1909; canadensis (Banks, Picus viridis Picidae Piciformes North America Fain 1981 1909) Ornithocheyletia gersoni Dendrocopos Smiley 1970; Picidae Piciformes North America Smiley, 1970 pubescens Fain 1981 Ornithocheyletia hallae Europe, Africa, Smiley 1970, Columba livia Columbidae Columbiformes Smiley, 1970 North America 1977; Fain 1981 Ornithocheyletia hallae Chalcophaps European Zoo Columbidae Columbiformes Fain 1972, 1981 similis Fain, 1972 indica (captive) Ornithocheyletia European Zoo Geopelia striata Columbidae Columbiformes Fain 1981 geopeliae Fain, 1981 (captive) Ornithocheyletia Fain and columbigallinae Fain et Columbina sp. Columbidae Columbiformes South America Bochkov 2002 Bochkov, 2002 Ornithocheyletia Parakeet Psittacidae Psittaciformes North America Smiley 1970 lawrenceae Smiley, 1970 Platycercus Womersley 1941; “ Psittacidae Psittaciformes Australia elegans Fain 1981 Ornithocheyletia European Zoo Psittacula krameri Psittacidae Psittaciformes Fain 1972, 1981 psittaculae Fain, 1972 (captive) Ornithocheyletia psittaci European Zoo Psittacus erithacus Psittacidae Psittaciformes Fain 1972, 1981 Fain, 1972 (captive) Ornithocheyletia psittaci Poicephalus Psittacidae Psittaciformes Africa Fain 1981 poicephali Fain, 1972 senegalus Ornithocheyletia smileyi Myiopsitta European Zoo Psittacidae Psittaciformes Fain 1972, 1981 Fain, 1972 monachus (captive) Ornithocheyletia European Zoo Nandayus nenday Psittacidae Psittaciformes Fain 1972, 1981 argentinensis Fain, 1972 (captive) European Zoo “ Forpus passerinus Psittacidae Psittaciformes Fain 1972, 1981 (captive) Ornithocheyletia Francolinus Phasianidae Galliformes Africa Fain 1972, 1981 francolini Fain, 1972 natalensis Neocheyletiella Baker, 1949 Neocheyletiella rohweri Sitta pygmaea Sittidae Passeriformes North America Baker 1949 Baker, 1949 Neocheyletiella Berlese and Europe, North microrhyncha (Berlese et Hirundo rustica Hirundinidae Passeriformes Trouessart 1889 ; America Trouessart, 1889) Smiley 1970 “ Delichon urbica Hirundinidae Passeriformes Europe Volgin 1969 “ Riparia riparia Hirundinidae Passeriformes Europe Volgin 1969 Petrochelidon “ Hirundinidae Passeriformes North America Smiley 1970 pyrrhonota Cecropis “ Hirundinidae Passeriformes Africa Fain 1980a abyssinica Psalidoprocne “ Hirundinidae Passeriformes Africa Fain 1980a albiceps Neocheyletiella Leucosticte smallwoodae Baker, Fringillidae Passeriformes Australia Baker 1949 australis 1949 Neocheyletiella media Leiothrix lutea Timaliidae Passeriformes Asia Fain 1972, 1980a Fain, 1972 Neocheyletiella siva Minla European Zoo Timaliidae Passeriformes Fain 1972, 1980a Fain, 1972 cyanouroptera (captive) Neocheyletiella Amandava European Zoo Leiothrichidae Passeriformes Fain 1972, 1980a amandavae Fain, 1972 amandava (captive)

i2 Appendix

Neocheyletiella Estrilda megaphallos Lawrence, Estrildidae Passeriformes Africa Lawrence 1959a erythronotos 1959 Neocheyletiella Fain and lonchurae Fain et Lonchura sp. Estrildidae Passeriformes Africa Bochkov 2002 Bochkov, 2002 Neocheyletiella pittae European Zoo Pitta megarhyncha Pittidae Passeriformes Fain 1972, 1980a Fain, 1972 (captive) Neocheyletiella artami Artamus Domrow 1966, Artamidae Passeriformes Australia Domrow, 1966 cyanopterus 1970 Neocheyletiella avicola European Zoo Ara sp. Psittacidae Psittaciformes Fain 1972, 1980a Fain, 1972 (captive) European Zoo “ Agapornis fisheri Psittacidae Psittaciformes Fain 1980a (captive) European Zoo (captive), North Smiley 1977 ; “ Erythrura prasina Estrildidae Passeriformes America Fain 1980a (captive) Neocheyletiella Bochkov and nectarinia Bochkov et Cinnyris jugularis Nectariniidae Passeriformes Asia OConnor 2003 OConnor, 2003 Neocheyletiella aegithali Aegithalos Bochkov and Bochkov et OConnor, Aegithalidae Passeriformes Asia concinnus OConnor 2003 2003 Neocheyletiella queleae Fain and Quelea quelea Ploceidae Passeriformes Africa Fain et Bochkov, 2002 Bochkov 2002 Streptopelia Fain and “ Columbidae Columbiformes Africa semitorquata Bochkov 2002 Neocheyletiella ardeola Bochkov and Ardeola ralloides Ardeidae Pelecaniformes Africa Bochkov et Ochoa, 2005 Ochoa 2005 Neocheyletiella athene European Zoo Fain and Athene brama Strigidae Strigiformes Fain et Bochkov, 2002 (captive) Bochkov 2002 Bakericheyla Volgin, 1966 Bakericheyla chanayi Berlese and (Berlese et Trouessart, Motacilla alba Motacillidae Passeriformes Europe Trouessart 1889 1889) Baker 1949; “ Fringilla coelebs Fringillidae Passeriformes Europe Volgin 1966 Fringilla Akimov and “ Fringillidae Passeriformes Europe montifringilla Gorgol 1990 Akimov and “ Spinus spinus Fringillidae Passeriformes Europe Gorgol 1990 Akimov and “ Carduelis carduelis Fringillidae Passeriformes Europe Gorgol 1990 Akimov and “ Passer montanus Passeridae Passeriformes Europasia Gorgol 1990 Passer domesticus Akimov and “ Passeridae Passeriformes Asia indicus Gorgol 1990 Akimov and “ Hirundo rustica Hirundinidae Passeriformes Europe Gorgol 1990 “ Delichon urbica Hirundinidae Passeriformes Europe Volgin 1969 Akimov and “ Motacilla flava Motacillidae Passeriformes Europe Gorgol 1990 Akimov and “ Anthus trivialis Motacillidae Passeriformes Europe Gorgol 1990 “ Oenanthe picata Muscicapidae Passeriformes Asia Volgin 1969

i3 M. Skoracki, S. Zabludovskaya and A.V. Bochkov

Bakericheyla chanayi Cyanoptila (Berlese et Trouessart, Muscicapidae Passeriformes Asia Volgin 1969 cyanomelana 1889) “ Muscicapa striata Muscicapidae Passeriformes Europe Volgin 1969 Akimov and “ Turdus pilaris Turdidae Passeriformes Europe Gorgol 1990 Akimov and “ Sylvia atricapilla Sylviidae Passeriformes Europe Gorgol 1990 Akimov and “ Sylvia borin Sylviidae Passeriformes Europe Gorgol 1990 Phylloscopus Akimov and “ Sylviidae Passeriformes Europe trochilus Gorgol 1990 “ Parus major Paridae Passeriformes Europe Volgin 1969 Paroaria European Zoo “ Emberizidae Passeriformes Fain 1972, 1980b nigrogenis (captive), Akimov and “ Emberiza citrinella Emberizidae Passeriformes Europe Gorgol 1990 Zonotrichia Furman and “ Emberizidae Passeriformes North America leucophrys Sousa 1969 Zonotrichia Furman and “ Emberizidae Passeriformes North America atricapilla Sousa 1969 “ Merops apiaster Meropidae Coraciiformes Asia Volgin 1969 Bakericheyla faini Cossypha dichroa Muscicapidae Passeriformes Africa Lawrence 1959a (Lawrence, 1959) Bakericheyla subquadra­ Merops pusillus Meropidae Coraciiformes Africa Lawrence 1959a ta (Lawrence, 1959) Bakericheyla transvaa­ Merops pusillus Meropidae Coraciiformes Africa Lawrence 1959a lica (Lawrence, 1959) “ Merops persicus Meropidae Coraciiformes Africa Lawrence 1959a Lawrence 1959a; “ Merops apiaster Meropidae Coraciiformes Africa, Asia Volgin 1969; Fain 1980b “ Merops persicus Meropidae Coraciiformes Asia Volgin 1969 “ Merops nubicoides Meropidae Coraciiformes Africa Fain 1980b Merops “ Meropidae Coraciiformes Africa Fain 1980b bullockoides Bakericheyla (Apodicheyla) Fain, 1979 Bakericheyla africana Cypsiurus parvus Apodidae Apodiformes Africa Fain 1979a Fain 1979 Bakericheyla benoiti Merops bulocki Meropidae Coraciiformes Africa Fain 1980b Fain, 1980 Apodicheles Fain, 1979 Apodicheles cypsiurus Cypsiurus parvus Apodidae Apodiformes Africa Fain 1979a Fain, 1979 Apodicheles apus Fain, Apus caffer Apodidae Apodiformes Africa Fain 1979a 1979 Apodicheles collocalia Collocalia Bochkov and Bochkov et OConnor, Apodidae Apodiformes Asia esculenta OConnor 2003 2003 Metacheyletiini Fain, 1980 Metacheyletia Fain, 1972 Metacheyletia obesa Psittacula krameri Psittacidae Psittaciformes Africa Fain 1972, 1980c Fain, 1972

i4 Appendix

Metacheyletia longise­ tosa Atyeo, Kethley et Amazona finschi Psittacidae Psittaciformes North America Atyeo et al. 1984 Perez, 1984 Metacheyletia amazonae Amazona Bochkov and Bochkov et OConnor, Psittacidae Psittaciformes South America amazonica Oconnor 2003 2003 Metacheyletia degene­ Serinus Fain and rata Fain et Bochkov, Fringillidae Passeriformes Africa mozambicus Bochkov 2003 2003 Metacheyletia ngaii Corythaixoides Bochkov and Bochkov et Skoracki, Musophagidae Cuculiformes Africa leucogaster Skoracki 2011 2011 Cheletosomatini Volgin, 1969 Cheletosoma Oudemans, 1905 Cheletosoma tyrannus Aramus guarauna Aramidae Gruiformes South America Oudemans 1905 Oudemans, 1905 Cheletosoma tauraco Tauraco Bochkov and Bochkov et Scoracki, Musophagidae Cuculiformes Africa leucolophus Skoracki 2012 2012 Cheletoides Oudemans, 1904 Cheletoides uncinatus Europe Heller 1880 ; Pavo cristatus Phasianidae Galliformes Heller, 1880 (domesticated) Oudemans 1906 Cheletoides chirunduen­ Numida meleagris Numididae Galliformes Africa Fain 1972, 1979b sis Fain, 1972 Metacheletoides Fain, 1972 Fain 1972, Metacheletoides 1979b; Bochkov Numida meleagris Numididae Galliformes Africa numidae Fain, 1972 and Skoracki 2012 Metacheletoides akanyaruensis Fain, Numida meleagris Numididae Galliformes Africa Fain 1972, 1979b 1972 Metacheletoides crinifer Crinifer zonurus Musophagidae Cuculiformes Africa Fain 1979b Fain, 1979 “ Crinifer piscator Musophagidae Cuculiformes Africa Fain 1979b Corythaixoides “ Musophagidae Cuculiformes Africa Fain 1979b leucogaster Metacheletoides Crinifer zonurus Musophagidae Cuculiformes Africa Fain 1979b gisagarensis Fain, 1979 “ Crinifer piscator Musophagidae Cuculiformes Africa Fain 1979b Cheletopsis Oudemans, 1904 Poppe 1888 ; Oudemans 1904, Cheletopsis norneri Charadrii- Sterna hirundo Laridae Eurasia 1906 ; Volgin (Poppe, 1888) formes 1969 ; Mironov et al. 1991 Charadrii- Bochkov et al. “ Sterna repressa Laridae Africa formes 2002 Charadrii- Bochkov et al. “ Phaetusa simplex Laridae South America formes 2002 Volgin 1969; Gelochelidon Charadrii- Eurasia, North “ Laridae Mironov et al. nilotica formes America 1991 Charadrii- Bochkov et al. “ Chlidonias hybrida Laridae Africa formes 2002

i5 M. Skoracki, S. Zabludovskaya and A.V. Bochkov

Mironov et al. Cheletopsis norneri Charadrii- Tringa totanus Scolopacidae Asia 1991; Bochkov (Poppe, 1888) formes et al. 2002 Cheletopsis thalasseus Thalasseus Charadrii- Bochkov and Bochkov et OConnor, Laridae North America sandvicensis formes OConnor 2003 2003 Cheletopsis larosterna Charadrii- Bochkov and Bochkov et Skoracki, Larosterna inca Laridae South America formes Skoracki 2012 2012 Cheletopsis basilica Charadrii- Tringa totanus Scolopacidae Europe Oudemans 1904 Oudemans, 1904 formes Charadrius Charadrii- Bochkov et al. “ Charadriidae Europe hiaticula formes 2002 Charadrii- Bochkov et al. “ Calidris melanotos Scolopacidae North America formes 2002 Charadrii- Bochkov et al. “ Chlidonias niger Laridae North America formes 2002 Cheletopsis animosa Charadrii- Tringa totanus Scolopacidae Europe Oudemans 1904 Oudemans, 1904 formes Charadrii- Bochkov et al. “ Sterna hirundo Laridae North America formes 2002 Cheletopsis anax Charadrii- Tringa totanus Scolopacidae Europe Oudemans 1904 Oudemans, 1904 formes Cheletopsis mariae Mironov et al. Charadrii- Mironov, Bochkov et Actitis hypoleucos Scolopacidae Eurasia 1991 ; Bochkov formes Chirov, 1991 et al. 2002 Oudemans 1904; Cheletopsis impavida Charadrii- Tringa totanus Scolopacidae Eurasia Mironov et al. Oudemans, 1904 formes 1991 Charadrii- Bochkov et al. “ Tringa stagnatilis Scolopacidae Europe formes 2002 Charadrii- Bochkov et al. “ Calidris ferruginea Scolopacidae Europe formes 2002 Charadrii- Mironov et al. “ Calidris minuta Scolopacidae Asia formes 1991 Charadrii- Mironov et al. “ Calidris temminckii Scolopacidae Asia formes 1991 Charadrii- Mironov et al. “ Calidris ruficollis Scolopacidae Asia formes 1991 Charadrii- Bochkov et al. “ Calidris pusilla Scolopacidae North America formes 2002 Calidris Charadrii- Bochkov et al. “ Scolopacidae North America himantopus formes 2002 Nycticryphes Charadrii- Bochkov et al. “ Rostratulidae North America semicollaris formes 2002 Kivganov and Cheletopsis daberti Charadrii- Bochkov 1994 ; Kivganov et Bochkov, Tringa glareola Scolopacidae Europa formes Bochkov et al. 1994 2002 Charadrii- Bochkov et al. “ Tringa nebularia Scolopacidae Europe formes 2002 Charadrii- Bochkov et al. “ Calidris temminckii Scolopacidae Europe formes 2002 Calidris Charadrii- Bochkov et al. “ Scolopacidae North America himantopus formes 2002

i6 Appendix

Cheletopsis charadrii Charadrii- Mironov et al. Mironov, Bochkov et Charadrius dubius Charadriidae Asia formes 1991 Chirov, 1991 Cheletopsis limnodromi Limnodromus Charadrii- Bochkov et al. Bochkov, Fain et Dabert, Scolopacidae North America griseus formes 2002 2002 Cheletopsis prosobonia­ Prosobonia Charadrii- Bochkov et al. lis Bochkov, Fain et Scolopacidae Australia cancellata formes 2002 Dabert, 2002 Cheletopsis rhynchops Rynchops Charadrii- Bochkov et al. Bochkov, Fain et Dabert, Laridae Africa flavirostris formes 2002 2002 Charadrii- Bochkov et al. “ Rynchops niger Laridae North America formes 2002 Species inquirrenda Oudemmans Cheletopsis magnanima Charadrii- Tringa flavipes Scolopacidae South America 1904; Bochkov Oudemans, 1904 formes et al. 2002 Picocheyletus Bochkov et OConnor, 2003 Picocheyletus tricholae­ Tricholaema Bochkov and mae Bochkov et Lybiidae Piciformes South America hirsuta OConnor 2003 OConnor, 2003 Eucheletopsis Volgin, 1969 Eucheletopsis major Hemiprocne Trouessart 1893; Hemiprocnidae Apodiformes Australasia Trouessart, 1893 mystacea Oudemans 1906 Harpirhynchidae Dubinin, 1957 Harpirhynchinae Dubinin, 1957 Harpirhynchus Megnin, 1877 Nitzsch 1818; Harpirhynchus nidulans Europe, North Chloris chloris Fringillidae Passeriformes Fain 1995; Moss (Nitzsch, 1818) America 1979 Fritsch 1954; Coccothraustes Europe, North “ Fringilidae Passeriformes Literak et al. coccothraustes America 2005; Moss 1979 “ Fringilla coelebs Fringillidae Passeriformes Europe Fritsch 1954 Unknown “ Corvus corone Corvidae Passeriformes Moss 1979 locality Locustella “ Locustellidae Passeriformes North America Moss 1979 lanceolata Harpirhynchus galeridae Moss 1979; Fain Fain, Bochkov et Galerida cristata Alaudidae Passeriformes Asia et al. 1999 Mironov, 1999 Unknown “ Alauda arvensis Alaudidae Passeriformes Moss 1979 locality Henry et al. Harpirhynchus dusbabe­ 2004; Literak et ki Bochkov et Literak, Panurus biarmicus Panuridae Passeriformes Europe al. 2005; 2006 Bochkov and Literak 2006 Harpirhynchus quasi­ Bochkov and modo Bochkov et Molothrus ater Icteridae Passeriformes North America Mertins 2010 Mertins, 2010 Harpyrhynchoides Fain, 1972 Harpyrhynchoides Psittacula European Zoo Psittacidae Psittaciformes Fain 1972, 1994 squamosus (Fain, 1972) cyanocephala (captive)

i7 M. Skoracki, S. Zabludovskaya and A.V. Bochkov

Harpyrhynchoides European Zoo Cacatua sp. Cacatuidae Psittaciformes Fain 1972, 1994 kakatoe (Fain, 1972) (captive) “ Cacatua galerita Cacatuidae Psittaciformes Australia Domrow 1991 Harpyrhynchoides Psittacula European Zoo Psittacidae Psittaciformes Fain 1972, 1994 psittaculae (Fain, 1972) alexandri (captive) Psittacula European Zoo “ Psittacidae Psittaciformes Fain 1972, 1994 cyanocephala (captive) Harpyrhynchoides European Zoo Psittacus erithacus Psittacidae Psittaciformes Fain 1972, 1994 psittaci Fain, 1972 (captive) Poicephalus European Zoo “ Psittacidae Psittaciformes Fain 1972, 1994 senegalus (captive) Harpyrhynchoides European Zoo Amazona aestiva Psittacidae Psittaciformes Fain 1972, 1994 amazonae (Fain, 1972) (captive) Harpyrhynchoides European Zoo Nandayus nenday Psittacidae Psittaciformes Fain 1972, 1994 lawrencei (Fain, 1972) (captive) Harpirhynchoides Platycercus rosellacinus (Lawrence, Psittacidae Psittaciformes Australia Lawrence 1959c eximius 1959) Glossopsitta “ Psittacidae Psittaciformes Australia Domrow 1991 concinna Filippich and Trichoglossus “ Psittacidae Psittaciformes Australia Domrow 1985; chlorolepidotus Domrow 1991 Trichoglossus “ haematodus Psittacidae Psittaciformes Australia Domrow 1991 moluccanus “ Lathamus discolor Psittacidae Psittaciformes Australia Fain 1994b Harpyrhynchoides Metriopelia European Zoo Columbidae Columbiformes Fain 1972, 1994 metropeliae (Fain, 1972) ceciliae (captive) Harpyrhynchoides Metriopelia European Zoo Columbidae Columbiformes Fain 1972, 1994 modestus (Fain, 1976) ceciliae (captive) Harpyrhynchoides Columbina European Zoo Columbidae Columbiformes Fain 1972, 1994 capitatus (Fain, 1976) talpacoti (captive) Harpyrhynchoides Columbina European Zoo Columbidae Columbiformes Fain 1972, 1994 coxatus Fain, 1972 talpacoti (captive) Columbina European Zoo “ Columbidae Columbiformes Fain 1972, 1994 cruziana (captive) Columbina European Zoo “ Columbidae Columbiformes Fain 1972, 1994 squammata (captive) Harpyrhynchoides Columba livia Columbidae Columbiformes Europe Fain 1972, 1994 columbae (Fain, 1972) “ Columba palumbus Columbidae Columbiformes Europe Fain 1972, 1994 Harpyrhynchoides oenae European Zoo Oena capensis Columbidae Columbiformes Fain 1972, 1994 (Fain, 1972) (captive) Harpyrhynchoides oenae European Zoo Turtur tympanistria Columbidae Columbiformes Fain 1972, 1994 lamorali (Fain, 1972) (captive) Harpyrhynchoides Coturnix European Zoo Phasianidae Galliformes Fain 1972, 1994 coturnix Fain, 1972 delegorguei (captive) Harpyrhynchoides European Zoo Alectoris graeca Phasianidae Galliformes Fain 1972, 1994 alectoris (Fain, 1972) (captive) “ Alectoris rufa Phasianidae Galliformes Europe Fain et al. 1999 Harpyrhynchoides Numida meleagris Numididae Galliformes Africa Lawrence 1959b numidae Lawrence, 1959 Harpyrhynchoides Gallinago Charadrii- Scolopacidae Europe Fritsch 1954 capellae (Fritsch, 1954) gallinago formes

i8 Appendix

Harpyrhynchoides Lymnocryptes Charadrii- Scolopacidae Europe Fain et al. 1999 capellae (Fritsch, 1954) minimus formes Harpyrhynchoides Ardea herodias Ardeidae Pelecaniformes North America Boyd 1968 herodius (Boyd, 1968) “ Ardea cinerea Ardeidae Pelecaniformes Europe Fain 1994b “ Butorides virescens Ardeidae Pelecaniformes North America Boyd 1968 Harpyrhynchoides Leptoptilos European Zoo Ciconiidae Ciconiiformes Fain 1972, 1994 leptoptilus (Fain, 1976) crumeniferus (captive) Harpyrhynchoides Fritsch 1954; tracheatus (Fritsch, Buteo buteo Accipitridae Accipitriformes Europe Fain 1994b 1954) Harpyrhynchoides tyto Tyto alba Tytonidae Strigiformes Europe Fain 1972, 1994 (Fain, 1972) Harpyrhynchoides asio Asio otus Strigidae Strigiformes Europe Fain 1972, 1994 (Fain, 1972) Harpyrhynchoides European Zoo Anas querquedula Anatidae Anseriformes Fain 1972, 1994 anatum (Fain, 1976) (captive) Harpyrhynchoides Berlese and cristagalli (Berlese et Colius striatus Coliidae Coliiformes Africa Trouessart 1889; Trouessart, 1889) Lawrence 1959c “ Colius colius Coliidae Coliiformes Africa Fain 1994b Harpyrhynchoides Eremopterix Alaudidae Passeriformes Africa Fain 1972, 1994 zumpti (Fain, 1972) leucotis Bochkov and “ Junco hyemalis Emberizidae Passeriformes North America Galloway 2004 European Zoo “ Diuca diuca Emberizidae Passeriformes Fain 1972, 1994 (captive) Bochkov and “ Quiscalus quiscula Icteridae Passeriformes North America Galloway 2001 Harpyrhynchoides Megnin 1877; alaudinus Bochkov, Alauda arvensis Alaudidae Passeriformes Europe Oudemans 1939; 2000 Bochkov 2000a Cerny and Sixl Harpyrhynchoides 1971; Fain et al. rubeculinus (Cerny et Erithacus rubecula Muscicapidae Passeriformes Europe 1999; Bochkov Sixl, 1971) and Literak 2008 Harpyrhynchoides kirgizorum Fain, Rhodospiza Fringillidae Passeriformes Asia Fain et al. 1999 Bochkov et Mironov, obsoleta 1999 Harpyrhynchoides Carpodacus Bochkov and vulgaris Bochkov et Gal- Fringillidae Passeriformes North America purpureus Galloway 2004 loway, 2004 Bochkov and “ Setophaga ruticilla Parulidae Passeriformes North America Galloway 2004 Harpyrhynchoides parazumpti Fain, Corvus monedula Corvidae Passeriformes Europe Fain et al. 1999 Bochkov et Mironov, 1999 Harpyrhynchoides Campethera pectinifer (Lawrence, Picidae Piciformes Africa Lawrence 1959a abingoni 1959) Campethera “ Picidae Piciformes Africa Fain 1994b cailliauti Harpyrhynchoides vercammeni (Lawrence, Centropus grillii Cuculidae Cuculiformes Africa Lawrence 1959a 1959)

i9 M. Skoracki, S. Zabludovskaya and A.V. Bochkov

Harpyrhynchoides (Pseudoharpirhynchus) Fain, Bochkov et Mironov, 1999 Harpirhynchoides Agapornis European Zoo Psittacidae Psittaciformes Fain 1972, 1994 agapornis (Fain, 1972) roseicollis (captive) Agapornis European Zoo “ Psittacidae Psittaciformes Fain 1994b pullarius (captive) Agapornis European Zoo “ Psittacidae Psittaciformes Fain 1994b nigrigenis (captive) Harpyrhynchoides cylindripalpus (Fritsch, Fringilla coelebs Fringillidae Passeriformes Europe Fritsch 1954 1954) European Zoo “ Passerina ciris Cardinalidae Passeriformes Fain 1994b (captive) Procellarii- Harpirhynchoides sp. Puffinus gravis Procellariidae North America Foster et al. 1996 formes Neharpyrhynchus Fain, 1972 Fritsch 1954; Neharpyrhynchus Martinu et al. Fringilla coelebs Fringillidae Passeriformes Europe plumaris (Fritsch, 1954) 2008; Bochkov 2000b “ Muscicapa striata Muscicapidae Passeriformes Europe Fritsch 1954 Neharpyrhynchus baile Bochkov et al. Bochkov, Literak et Turdus leucomelas Turdidae Passeriformes South America 2007 Capek, 2007 Neharpyrhynchus turdi Martinu et al. Martinu, Dusbabek et Turdus merula Turdidae Passeriformes Europe 2008 Literak, 2008 Neharpyrhynchus Chlorospingus Bochkov and chlorospingus Bochkov Emberizidae Passeriformes South America pileatus Literak 2011 et Literak, 2011 Neharpyrhynchus Hippolais icterina Acrocephalidae Passeriformes Europe Bochkov 2000b hippolae Bochkov, 2000 Neharpyrhynchus Bochkov and mironovi Bochkov et Dacnis cayana Thraupidae Passeriformes South America Literak 2011 Literak, 2011 Neharpyrhynchus Bochkov and tangara Bochkov et Tangara cayana Thraupidae Passeriformes South America Literak 2011 Literak, 2011 Literak et al. “ Thraupis episcopus Thraupidae Passeriformes South America 2012 Neharpyrhynchus novoplumaris (Moss, Certhia familiaris Certhiidae Passeriformes North America Moss et al. 1968 Oliver et Nelson, 1968) Cardinalis “ Cardinalidae Passeriformes North America Moss et al. 1968 cardinalis Campylorhynchus “ Troglodytidae Passeriformes North America Moss 1979 brunneicapillus “ Spizella passerina Emberizidae Passeriformes North America Moss 1979 Amphispiza “ Emberizidae Passeriformes North America Moss 1979 bilineata “ Melozone fusca Emberizidae Passeriformes North America Moss 1979 Neharpyrhynchus pari Martinu et al. Martinu, Dusbabek et Parus major Paridae Passeriformes Europe 2008 Literak, 2008 Cyanistes Martinu et al. “ Paridae Passeriformes Europe caeruleus 2008

i10 Appendix

Neharpyrhynchus pari Martinu et al. Martinu, Dusbabek et Poecile montanus Paridae Passeriformes Europe 2008 Literak, 2008 Moss 1979; “ Poecile palustris Paridae Passeriformes Europe Martinu et al. 2008 “ Baeolophus bicolor Paridae Passeriformes North America Moss 1979 Berlese and Trouessart 1889; Neharpyrhynchus Fritsch 1954; Europe, Africa, pilirostris (Berlese et Passer domesticus Passeridae Passeriformes Lawrence 1959a; North America Trouessart, 1889) Fain 1995; Martinu et al. 2008 Aegithalos Unknown “ Aegithalidae Passeriformes Moss 1979 caudatus locality Neharpyrhynchus schoenobaenus Martinu, Acrocephalus Martinu et al. Acrocephalidae Passeriformes Europe Dusbabek et Literak, schoenobaenus 2008 2008 Neharpyrhynchus spinus Martinu et al. Martinu, Dusbabek et Spinus spinus Fringillidae Passeriformes Europe 2008 Literak, 2008 Carduelis “ Fringillidae Passeriformes Europe Fritsch 1954 cannabina Neharpyrhynchus Hummingbird Trochilidae Apodiformes South America Fain 1972, 1995 trochilinus (Fain, 1972) Chrysolampis “ Trochilidae Apodiformes South America Fain 1995 mosquitus Bochkov and “ Panterpe insignis Trochilidae Apodiformes South America Literak 2011 Bochkov and “ Eugenes fulgens Trochilidae Apodiformes South America Literak 2011 Bochkov and Literak 2011; “ Amazilia lactea Trochilidae Apodiformes South America Literak et al. 2012 Amazilia Literak et al. “ Trochilidae Apodiformes South America chionogaster 2012 Metharpyrhynchus Fain, 1972 Metharpyrhynchus Lonchura European Zoo Estrildidae Passeriformes Fain 1972, 1995 macrophallus Fain, 1972 oryzivora (captive) Metharpyrhynchus Lonchura European Zoo Estrildidae Passeriformes Fain 1972, 1995 macrophallus Fain, 1972 oryzivora (captive) Metharpyrhyncus mossi Prinia subflava Cisticolidae Passeriformes Africa Fain 1995 Fain, 1995 Metharpyrhynchus Philetairus socius Ploceidae Passeriformes Africa Fain 1995 namibiensis Fain, 1995 Metharpyrhynchoides Jynx ruficollis Picidae Piciformes Africa Fain 1995 jynx Fain, 1972 Perharpyrhynchus Fain, 1972 Perharpyrhynchus Jacana spinosa Charadrii- European Zoo Jacanidae Fain 1972, 1995 jacana Fain, 1972 intermedia formes (captive) Perharpyrhynchus Recurvirostra Charadrii- European Zoo Recurvirostridae Fain 1976, 1995 recurvirostra Fain, 1976 avosetta formes (captive) Ralliharpirhynchus Fain, 1972

i11 M. Skoracki, S. Zabludovskaya and A.V. Bochkov

Ralliharpirhynchus Porphyrio European Zoo Rallidae Gruiformes Fain 1972, 1995 porphyrio (Fain, 1972) porphyrio (captive) Ralliharpirhynchus Amaurornis Rallidae Gruiformes Africa Fain 1972, 1995 limnocorax (Fain, 1972) flavirostra Harpyrhynchiella Fain, 1972 Harpyrhynchiella Fritsch 1954; Apus apus Apodidae Apodiformes Europe reductus (Fritsch, 1954) Fain 1972, 1995 Cypsharpirhynchus Fain, 1995 Cypsharpirhynchus Cypsiurus parvus Apodidae Apodiformes Africa Fain 1972, 1995 cypsiuri (Fain, 1972) “ Apus affinis Apodidae Apodiformes Africa Fain 1972, 1995 Anharpyrhynchus Fain, 1972 Anharpyrhynchus Garrulus Fritsch 1954; monstrosus (Fritsch, Corvidae Passeriformes Europe glandarius Fain 1972, 1995 1954) Unknown “ Corvus monedula Corvidae Passeriformes Moss 1979 locality Unknown “ Cyanocitta cristata Corvidae Passeriformes Moss 1979 locality Unknown “ Cyanocitta stelleri Corvidae Passeriformes Moss 1979 locality Phylidonyris Unknown “ Meliphagidae Passeriformes Moss 1979 novaehollandiae locality Cissomela Unknown “ Meliphagidae Passeriformes Moss 1979 pectoralis locality Unknown “ Manorina flavigula Meliphagidae Passeriformes Moss 1979 locality Lichmera Unknown “ Meliphagidae Passeriformes Moss 1979 indistincta locality Unknown “ Colaptes auratus Picidae Piciformes Moss 1979 locality Trichorhynchiella Fain, 1995 Trichorhynchiella Lonchura European Zoo Estrildidae Passeriformes Fain 1995 paddae Fain, 1995 oryzivora (captive) Harpypalpinae Fain, 1972 Harpypalpus Dubinin, 1957 Berlese and Harpypalpus holopus Trouessart 1889; (Berlese et Trouessart, Passer domesticus Passeridae Passeriformes Europe Berlese 1894; 1889) Fain 1972; Fain 1999 Unknown “ Luscinia svecica Muscicapidae Passeriformes Moss 1979 locality Harpypalpus longipis Troglodytis Troglodytidae Passeriformes Europe Fritsch 1954 (Fritsch, 1954) troglodytis Moss 1979; “ Parus major Paridae Passeriformes Europe Skoracki et al. 2004 Cyanistes Unknown “ Paridae Passeriformes Moss 1979 caeruleus locality Unknown “ Spinus spinus Fringillidae Passeriformes Moss 1979 locality Unknown “ Pyrrhula pyrrhula Fringillidae Passeriformes Moss 1979 locality Nucifraga Unknown “ Corvidae Passeriformes Moss 1979 caryocatactes locality

i12 Appendix

Harpypalpus serini Fain, Serinus European Zoo Fain 1972; Fain Fringillidae Passeriformes 1972 mozambicus (captive) et al. 1999 Harpypalpus dubinini Ploceus European Zoo Fain 1972; Fain Ploceidae Passeriformes Fain, 1972 intermedius (captive) et al. 1999 Harpypalpus tiarae Fain, European Zoo Bochkov et Mironov, Tiaris canorus Emberizidae Passeriformes Fain et al. 1999 (captive) 1999 Harpypalpus spermestes Spermestes Fain, Bochkov et Estrildidae Passeriformes Africa Fain et al. 1999 cucullatus Mironov, 1999 Harpypalpoides Lombert et Moss, 1983 Lombert and Harpypalpoides Moss 1983; lukoschusi Lombert et Turdus merula Turdidae Passeriformes Europe Skoracki et al. Moss, 1983 2004 Harpypalpoides lesickii Coccothraustes Skoracki et al. Skoracki, Bochkov et Fringillidae Passeriformes Europe coccothraustes 2004 Sikora, 2004 Harpypalpoides Fain 1972; Fain Psalidoprocne sp. Hirundinidae Passeriformes Africa hirundinus (Fain, 1972) et al. 1999 Harpypalpoides namibiensis Fain, Emberiza Emberizidae Passeriformes Africa Fain 1999 Bochkov et Mironov, impetuani 1999 Species inquirenda Harpirhynchus brevis Coccothraustes Fringillidae Passeriformes North America Ewing 1911 Ewing, 1911 vespertinus Harpirhynchus longipi­ Loxia sp. Fringillidae Passeriformes North America Banks 1905 lus Banks, 1905 Cloacaridae Camin, Moss, Oliver et Singer, 1967 Pneumophaginae Fain et Smiley, 1989 Pneumophagus Fain et Smiley, 1989 Pneumophagus bubonis Fain and Smiley Bubo virginianus Strigidae Strigiformes North America Fain et Smiley, 1989 1989 Syringophilidae Lavoipierre, 1953 Syringophilinae Lavoipierre, 1953 Apodisyringiana Skoracki, 2005 Apodisyringiana Hemiprocne haszprunari Skoracki, Hemiprocnidae Apodiformes Asia Skoracki 2005a comata 2005 Apodisyringiana Hemiprocne mystaceae Skoracki, Hemiprocnidae Apodiformes Australia Skoracki 2005a mystacea 2005 Apodisyringophilus Skoracki et OConnor, 2010 Apodisyringophilus Collocalia Skoracki and collocalius Skoracki et Apodidae Apodiformes Asia esculenta OConnor 2010 OConnor, 2010 Ascetomylla Kethley, 1970 Ascetomylla gallinula Gallinula Rallidae Gruiformes Africa Kethley 1970 Kethley, 1970 chloropus Ascetomylla porzanae Bochkov and (Bochkov et Galloway, Porzana carolina Rallidae Gruiformes North America Galloway 2004 2004) Aulobia Kethley, 1970

i13 M. Skoracki, S. Zabludovskaya and A.V. Bochkov

Aulobia anthreptes Anthreptes Skoracki and Skoracki et Glowska, Nectariniidae Passeriformes Asia malacensis Glowska 2008b 2008 Aulobia cardueli Skoracki et al. Skoracki, Hendricks et Carduelis psaltria Fringillidae Passeriformes North America 2010a Spicer, 2010 “ Carduelis carduelis Fringillidae Passeriformes Europe Skoracki 2011 Skoracki and “ Carduelis spinus Fringillidae Passeriformes Europe, Asia Bochkov 2010 “ Carduelis flamea Fringillidae Passeriformes Europe Skoracki 2011 “ Carduelis citrinella Fringillidae Passeriformes Europe Skoracki 2011 Carduelis “ Fringillidae Passeriformes Europe Skoracki 2011 flavirostris Aulobia cisticolae Skoracki and Cisticola erythrops Cisticolidae Passeriformes Africa Skoracki et Sikora, 2003 Sikora 2003 Skoracki and “ Cisticola cantans Cisticolidae Passeriformes Africa Sikora 2003 Skoracki and “ Cisticola lateralis Cisticolidae Passeriformes Africa Sikora 2003 Skoracki et al. “ Cisticola chubbi Cisticolidae Passeriformes Africa 2009a Aulobia dendroicae Setophaga Parulidae Passeriformes North America Kethley 1970 Kethley, 1970 coronata Setophaga Skoracki et al. “ Parulidae Passeriformes North America nigrescens 2010a Aulobia erythroptera Skoracki and Prinia erythroptera Cisticolidae Passeriformes Africa Skoracki et Dabert, 2001 Dabert 2001a Aulobia leucostictus Leucosticte arctoa Fringillidae Passeriformes Europe Skoracki 2011 Skoracki, 2011 Aulobia nectariniae Skoracki and Skoracki et Glowska, Cinnyris jugularis Nectariniidae Passeriformes Australia Glowska 2008b 2008 Skoracki and “ Leptocoma sericea Nectariniidae Passeriformes Australia Glowska 2008b Aulobia stachyris Stachyridopsis Bochkov et al. (Bochkov, Mironov et Timaliidae Passeriformes Asia pyrrhops 2001 Skoracki, 2001) Bochkov and Mironov 1998; Aulobia sylviae Bochkov Glowska et al. Sylvia curruca Sylviidae Passeriformes Europe, Asia et Mironov, 1998 2007; Skoracki and Bochkov 2010 Bochkov and “ Sylvia atricapilla Sylviidae Passeriformes Europe Mironov 1998 “ Sylvia nisoria Sylviidae Passeriformes Europe Skoracki 2011 Aulobia sylviettae (Fain, Bochkov et Mironov, Sylvietta whytti Macrosphenidae Passeriformes Africa Fain et al. 2000 2000) Aulobia virens Skoracki Sylvietta virens Skoracki and Macrosphenidae Passeriformes Africa et Dabert, 2001 flaviventris Dabert 2001a Aulonastus Kethley, 1970 Aulonastus albus Motacilla alba Motacillidae Passeriformes Europe Skoracki 2002a Skoracki, 2002 Aulonastus anthus Anthus trivialis Motacillidae Passeriformes Europe Skoracki 2011 Skoracki, 2011

i14 Appendix

Aulonastus anthus Anthus cervinus Motacillidae Passeriformes Europe Skoracki 2011 Skoracki, 2011 Aulonastus emberizicus Ammodramus Skoracki et al. Skoracki, Hendricks et Emberizidae Passeriformes North America savannarum 2010b Spicer, 2010 Zonotrichia Skoracki et al. “ Emberizidae Passeriformes North America atricapilla 2010b Passerculus Skoracki et al. “ Emberizidae Passeriformes North America sandwichensis 2010b Aulonastus euphagus Euphagus Skoracki et al. Skoracki, Hendricks et Icteridae Passeriformes North America cyanocephalus 2010b Spicer, 2010 Aulonastus fringillus Fringilla coelebs Fringillidae Passeriformes Europe Skoracki 2011 Skoracki, 2011 Aulonastus henicorhina Henicorhina Troglodytidae Passeriformes South America Sikora et al. 2012 Sikora et Skoracki, 2012 leucosticta Aulonastus lanius Lanius senator Laniidae Passeriformes Europe Skoracki 2011 Skoracki, 2011 Aulonastus loxius Loxia curvirostra Fringillidae Passeriformes Europe Skoracki 2011 Skoracki, 2011 Aulonastus lusciniae Luscinia luscinia Muscicapidae Passeriformes Europe Skoracki 2002a Skoracki, 2002 Aulonastus pipili Pipilo Emberizidae Passeriformes North America Kethley 1970 Kethley, 1970 erythrophthalmus Aulonastus pirangus Piranga Skoracki et al. Skoracki, Hendricks et Cardinalidae Passeriformes North America ludoviciana 2010b Spicer, 2010 Aulonastus prunellae Bochkov and Bochkov et Mironov, Prunella modularis Prunellidae Passeriformes Europe Mironov 1999 1999 Aulonastus sturnellus Skoracki et al. Skoracki, Hendricks et Sturnella magna Icteridae Passeriformes North America 2010b Spicer, 2010 Betasyringophiloidus Skoracki, 2011 Betasyringophiloidus Bochkov and motacillae (Bochkov et Motacilla flava Motacillidae Passeriformes Europe Mironov 1998 Mironov, 1998) Betasyringophiloidus Phoenicurus phoenicurus Skoracki, Muscicapidae Passeriformes Europe Skoracki 2011 ochruros 2011 Betasyringophiloidus Saxicola rubetra Muscicapidae Passeriformes Europe Skoracki 2011 saxicolus Skoracki, 2011 Skoracki 2002b, Betasyringophiloidus Emberiza 2004a, 2011; schoeniclus (Skoracki, Emberizidae Passeriformes Europe, Asia schoeniclus Skoracki and 2002) Bochkov 2010 Blaszakia Skoracki et Sikora, 2008 Blaszakia tauracos Skoracki and Tauraco livingstoni Musophagidae Cuculiformes Africa Skoracki et Sikora, 2008 Sikora 2008 Skoracki and “ Tauraco schalowi Musophagidae Cuculiformes Africa Sikora 2008 Blaszakia rossae Skoracki and Musophaga rossae Musophagidae Cuculiformes Africa Skoracki et Sikora, 2008 Sikora 2008 Bochkovia Skoracki et OConnor, 2010 Bochkovia phalaropi Phalaropus Charadrii- Skoracki and Skoracki et OConnor, Scolopacidae North America fulicarius formes OConnor 2010 2010

i15 M. Skoracki, S. Zabludovskaya and A.V. Bochkov

Bubophilus Philips et Norton, 1978 Bubophilus ascalaphus Philips and Bubo virginianus Strigidae Strigiformes North America Philips et Norton, 1978 Norton 1978 Skoracki and “ Bubo africanus Strigidae Strigiformes Africa Dabert 2002 Skoracki and “ Tyto alba affinis Tytonidae Strigiformes Africa Dabert 2002 Bubophilus asiobius Skoracki and Skoracki et Bochkov, Asio otus Strigidae Strigiformes Europe Bochkov 2002 2002 Bubophilus aluconis Nattress and Nattress et Skoracki, Strix aluco Strigidae Strigiformes Europe Skoracki 2009 2009 Skoracki et al. “ Asio otus Strigidae Strigiformes Europe 2012a Castosyringophilus Bochkov et Perez, 2002 Castosyringophilus Skoracki and claravis Skoracki et Claravis pretiosa Columbidae Columbiformes South America Glowska 2008a Glowska, 2008 Castosyringophilus forpi Forpus Bochkov and Psittacidae Psittaciformes North America Bochkov et Perez, 2002 cyanopygius Perez 2002 Casto 1980a; Castosyringophilus Columbina Columbidae Columbiformes North America Bochkov and mucuya (Casto, 1980) passerina Perez 2002 Columbina Bochkov and “ Columbidae Columbiformes South America squammata Fain 2003 Columbina Skoracki and “ Columbidae Columbiformes South America talpacoti Glowska 2008a Geophaps Bochkov and “ Columbidae Columbiformes Australia plumifera Fain 2003 Metriopelia Skoracki and “ Columbidae Columbiformes South America melanoptera Glowska 2008a Streptopelia Glowska et al. “ Columbidae Columbiformes Asia decaocto 2007 Brotogeris Bochkov and “ Psittacidae Psittaciformes South America versicolurus Fain 2003 Trichoglossus Bochkov and “ Psittacidae Psittaciformes Asia haematodus Fain 2003 Psilopsiagon Bochkov and “ Psittacidae Psittaciformes South America aymara Fain 2003 Charadriphilus Bochkov et Chystiakov, 2001 Bochkov and Charadriphilus Gallinago Charadrii- Mironov, 1998; gallinago (Bochkov et Scolopacidae Europe gallinago formes Skoracki et al. Mironov, 1998) 2006 Charadriphilus Charadrii- Bochkov and lyudmilae Bochkov et Scolopax rusticola Scolopacidae Europe formes Chystiakov 2001 Chystiakov, 2001 Charadriphilus paragua­ Gallinago Charadrii- Skoracki and iae (Skoracki et Sikora, Scolopacidae South America paraguaiae formes Sikora 2002 2002) Charadriphilus ralli Skoracki and Skoracki et Bochkov, Rallus aquaticus Rallidae Gruiformes Asia Bochkov 2010 2010 Charadriphilus vanelli Charadrii- Bochkov et al. Bochkov, Fain et Vanellus chilensis Charadriidae South America formes 2004 Skoracki, 2004

i16 Appendix

Chenophila Kethley, 1970 Chenophila branta Branta canadensis Anatidae Anseriformes North America Kethley 1970 Kethley, 1970 Chenophila kanduli Bochkov and Anas querquedula Anatidae Anseriformes Europe Bochkov, 1998 Mironov 1998 Ciconichenophilus Skoracki et OConnor, 2010 Ciconichenophilus Phoenicopterus Phoenicopteri- Phoenicopteri- Skoracki and phoeniconaias Skoracki Africa minor dae formes OConnor 2010 et OConnor, 2010 Colinophilus Kethley, 1973 Colinophilus wilsoni Colinus virginianus Odontophoridae Galliformes North America Kethley 1973 Kethley, 1973 Callipepla “ Odontophoridae Galliformes North America Casto 1976 squamata Corvitorotroglus Skoracki et Bochkov, 2010 Corvitorotroglus alpha Skoracki and Skoracki et Bochkov, Corvus frugilegus Corvidae Passeriformes Asia Bochkov 2010 2010 Creagonycha Kethley, 1970 Creagonycha lara Charadrii- Larus delawarensis Laridae North America Kethley 1970 Kethley, 1970 formes Creagonycha sternae Charadrii- Kivganov and Sterna albifrons Laridae Europe Kivganov, 1995 formes Sharafat 1995 Creagonycha totani Charadrii- Oudemans Tringa totanus Scolopacidae Europe (Oudemans, 1904) formes 1904,1906; Charadrii- Skoracki et al. “ Tringa glareola Scolopacidae Europe formes 2006 Skoracki and Charadrii- Dabert 2002; “ Calidris temminckii Scolopacidae Africa, Europe formes Skoracki et al. 2006 Calidris Charadrii- Skoracki et al. “ Scolopacidae North America himantopus formes 2006 Bochkov and Charadrii- Mironov 1998; “ Calidris minuta Scolopacidae Europe, Asia formes Skoracki et al. 2006 Crotophagisyringophilus Skoracki, 2008 Crotophagisyringophilus Crotophaga ani Cuculidae Cuculiformes South America Skoracki, 2008b io Skoracki, 2008 Cuculisyringophilus Skoracki, 2008 Cuculisyringophilus crotophaginus Skoracki, Guira guira Cuculidae Cuculiformes South America Skoracki 2008a 2008 Crotophaga North and South “ Cuculidae Cuculiformes Skoracki 2008a sulcirostris America Cuculisyringophilus Bochkov and chirovi (Bochkov et Cuculus canorus Cuculidae Cuculiformes Asia Mironov 1998 Mironov, 1998) Fritschisyringophilus Bochkov, Fain et Skoracki, 2004 Fritschisyringophilus Lonchura Bochkov et al. lonchurae Bochkov, Fain Estrildidae Passeriformes Asia punctulata 2004 et Skoracki, 2004 Galliphilopsis Skoracki et Sikora, 2004 Galliphilopsis bochkovi Skoracki and Tragopan sp. Phasianidae Galliformes Asia Skoracki et Sikora, 2004 Sikora 2004b

i17 M. Skoracki, S. Zabludovskaya and A.V. Bochkov

Galliphilopsis colinus Skoracki and Colinus cristatus Odontophoridae Galliformes South America Skoracki et Sikora, 2011 Sikora 2011 Galliphilopsis francoli­ Francolinus Skoracki and nus Skoracki et Sikora, Phasianidae Galliformes South Africa levaillantoides Sikora 2004b 2004 Galliphilopsis lophurus Lophura Skoracki and Phasianidae Galliformes Asia Skoracki et Sikora, 2004 leucomelanos Sikora 2004b Galliphilopsis szeptyckii Skoracki and Coturnix pectoralis Phasianidae Galliformes Australia Skoracki et Sikora, 2011 Sikora 2011 Skoracki and “ Coturnix coturnix Phasianidae Galliformes Asia Sikora 2011 Skoracki and “ Coturnix chinensis Phasianidae Galliformes Asia Sikora 2011 Skoracki and “ Alectoris barbara Phasianidae Galliformes Europe Sikora 2011 Kalamotrypetes Casto, 1980 Kalamotrypetes Colinus virginianus Odontophoridae Galliformes North America Casto 1980b colinastes Casto, 1980 Kalamotrypetes cracidus Skoracki and Penelope sp. Cracidae Galliformes South America Skoracki et Sikora, 2011 Sikora 2011 Kethleyana Kivganov, 1995 Kethleyana gelocheli­ Gelochelidon Charadrii- Kivganov and Laridae Europe doni Kivganov, 1995 nilotica formes Sharafat 1995 Krantziaulonastus Skoracki, 2011 Krantziaulonastus Skoracki 2002a, buczekae (Skoracki, Sturnus vulgaris Sturnidae Passeriformes Europe 2011 2002) Krantziaulonastus Jacamaralcyon galbulicus (Skoracki, Galbulidae Galbuliformes South America Skoracki 2008c tridactyla 2008) Krantziaulonastus Lonchura oryzivorus Skoracki, Estrildidae Passeriformes Asia Skoracki 2011 oryzivora 2011 Krantziaulonastus Lonchura lonchurus Skoracki, Estrildidae Passeriformes Asia Skoracki 2011 punctulata 2011 Krantziaulonastus Dinemellia Glowska and yoyomi Glowska et Ploceidae Passeriformes Africa dinemelli Skoracki 2011b Skoracki, 2011 Megasyringophilus Fain, Bochkov et Mironov, 2000 Megasyringophilus aquilus Skoracki, Skoracki et al. Aquila rapax Accipitridae Accipitriformes Africa Lontkowski et Stawarc- 2010d zyk, 2010 Skoracki et al. “ Aquila pomarina Accipitridae Accipitriformes Europe 2010d Skoracki and “ Accipiter nisus Accipitridae Accipitriformes Asia Bochkov 2010 Megasyringophilus cyanocephala Fain, Psittacula Psittacidae Psittaciformes Asia Fain et al. 2000 Bochkov et Mironov, cyanocephala 2000 Bochkov and “ Psittacula eupatria Psittacidae Psittaciformes Asia Fain 2003 Bochkov and “ Psittacula krameri Psittacidae Psittaciformes Asia Fain 2003

i18 Appendix

Megasyringophilus Trichoglossus Bochkov and dubinini Bochkov et Psittacidae Psittaciformes Asia ornatus Fain 2003 Fain, 2003 Megasyringophilus eos Eos bornea Psittacidae Psittaciformes Asia Skoracki 2005c Skoracki, 2005 Megasyringophilus Geoffroyus geoffroyus Skoracki, Psittacidae Psittaciformes Australia Skoracki 2005c geoffroyi 2005 Megasyringophilus kethleyi Fain, Bochkov Aratinga jandaya Psittacidae Psittaciformes South America Fain et al. 2000 et Mironov, 2000 Bochkov and “ Aratinga pertinax Psittacidae Psittaciformes South America Fain 2003 Brotogeris Bochkov and “ Psittacidae Psittaciformes South America versicolurus Fain 2003 Megasyringophilus Platycercus Bochkov and platycercus Bochkov et Psittacidae Psittaciformes Australia eximius Fain 2003 Fain, 2003 Megasyringophilus Rhynchopsitta Bochkov and rhynchopsittae Bochkov Psittacidae Psittaciformes North America pachyrhyncha Perez, 2002 et Perez, 2002 Megasyringophilus trichoglossus Fain, Trichoglossus sp. Psittacidae Psittaciformes South America Fain et al. 2000 Bochkov et Mironov, 2000 Trichoglossus “ Psittacidae Psittaciformes Asia Skoracki 2005c euteles Trichoglossus “ Psittacidae Psittaciformes Australia Skoracki 2005c chlorolepidotus Meitingsunes Glowska et Skoracki, 2010 Meitingsunes aldwelles Glowska and Glowska et Skoracki, Geotrygon frenata Columbidae Columbiformes South America Skoracki 2010 2010 Meitingsunes columbicus Columba oenas Columbidae Columbiformes Asia Skoracki 2011 Skoracki, 2011 “ Columba livia Columbidae Columbiformes Europe “ Columba palumbus Columbidae Columbiformes Europe Meitingsunes zenadou­ Zenaida macroura Columbidae Columbiformes North America Clark 1964b rae (Clark, 1964) “ Zenaida asiatica Columbidae Columbiformes North America Casto 1976 Skoracki and “ Zenaida auriculata Columbidae Columbiformes South America Sikora 2002 Casto 1976 North America, “ Columba livia Columbidae Columbiformes Bochkov and Africa Mironov 1998 Meitingsunes tympanis­ Skoracki and tria Skoracki et Dabert, Turtur tympanistria Columbidae Columbiformes Africa Dabert 2002 2002 Mironovia Chirov et Kravtsova, 1995 Mironovia phasiani Phasianus Chirov and Chirov et Kravtsova, Phasianidae Galliformes Asia colchicus Kravtsova 1995 1995 Mironovia coturnae Bochkov et al. Bochkov, Fain et Coturnix coturnix Phasianidae Galliformes Europe 2004 Skoracki, 2004 Mironovia rouloul Skoracki and Rollulus rouloul Phasianidae Galliformes Asia Skoracki et Sikora, 2004 Sikora 2004b

i19 M. Skoracki, S. Zabludovskaya and A.V. Bochkov

Mironovia lagopus Bochkov and Bochkov et Skirnisson, Lagopus muta Phasianidae Galliformes Europe Skirnisson 2011 2011 Skoracki and “ Lagopus lagopus Phasianidae Galliformes Europe Sikora 2011 Neoaulobia Fain, Bochkov et Mironov, 2000 Neoaulobia agapornis Agapornis Fain, Bochkov et Psittacidae Psittaciformes Africa Fain et al. 2000 nigrigenis Mironov, 2000 Bochkov and “ Agapornis fischeri Psittacidae Psittaciformes Africa Fain 2003 Agapornis Bochkov and “ Psittacidae Psittaciformes Africa personatus Fain 2003 Agapornis Bochkov and “ Psittacidae Psittaciformes Africa roseicollis Fain 2003 Bochkov and “ Agapornis taranta Psittacidae Psittaciformes Africa Fain 2003 Neoaulobia aratingae Fain, Bochkov et Aratinga jandaya Psittacidae Psittaciformes South America Fain et al. 2000 Mironov, 2000 Neoaulobia krafti Cacatua Cacatuidae Psittaciformes Australia Skoracki 2005c Skoracki, 2005 tenuirostris Neoaulobia mexicana Aratinga Bochkov and Psittacidae Psittaciformes North America Bochkov et Perez, 2002 canicularis Perez 2002 Bochkov and “ Aratinga pertinax Psittacidae Psittaciformes South America Fain 2003 Neoaulobia mironovi Bochkov and Amazona finschi Psittacidae Psittaciformes North America Bochkov et Perez, 2002 Perez 2002 Neoaulobia psittaculae Psittacula Fain, Bochkov et Psittacidae Psittaciformes Asia Fain et al. 2000 cyanocephala Mironov, 2000 Neoaulobia puylaerti Poicephalus Skoracki and (Skoracki et Dabert, Psittacidae Psittaciformes Africa senegalus versteri Dabert 1999b 1999) “ Loriculus pusillus Psittacidae Psittaciformes Asia Skoracki 2005c Loriculus “ Psittacidae Psittaciformes Asia Skoracki 2005c philippens Neoaulonastus Skoracki, 2004 Neoaulonastus aegitha­ Aegithalos Aegithalidae Passeriformes Europe Skoracki 2011 los Skoracki, 2011 caudatus Neoaulonastus bisetatus Fritsch 1958; Sylvia communis Sylviidae Passeriformes Europe (Fritsch, 1958) Skoracki 2011 “ Sylvia curruca Sylviidae Passeriformes Europe, Asia Skoracki 2011 “ Sylvia atricapilla Sylviidae Passeriformes Europe Skoracki 2011 Acrocephalus “ Acrocephalidae Passeriformes Europe Skoracki 1999a scirpaceus Acrocephalus “ Acrocephalidae Passeriformes Europe Skoracki 2011 paludicola Acrocephalus “ Acrocephalidae Passeriformes Asia Skoracki 2011 dumetorum Acrocephalus “ Acrocephalidae Passeriformes Europe Skoracki 2011 arundinaceus Acrocephalus Skoracki et al. “ Acrocephalidae Passeriformes Africa palustris 2011b Neoaulonastus caligatus Hippolais caligata Acrocephalidae Passeriformes Europe Skoracki 2011 Skoracki, 2011

i20 Appendix

Neoaulonastus chrysoco­ Chrysocolaptes Skoracki and laptes Skoracki et OCon- Picidae Piciformes Asia lucidus OConnor 2010 nor, 2010 Neoaulonastus picidus Picus canus Picidae Piciformes Europe Skoracki 2011 Skoracki, 2011 Dendrocopos “ Picidae Piciformes Europe Skoracki 2011 leucotos Dendrocopos “ Picidae Piciformes Europe Skoracki 2011 major Neoaulonastus remizus Remiz pendulinus Remizidae Passeriformes Europe Skoracki 2011 Skoracki, 2011 Neoaulonastus riparius Riparia riparia Hirundinidae Passeriformes Europe Skoracki 2011 Skoracki, 2011 Neoaulonastus zosterops Zosterops Skoracki et al. Skoracki, Antczak et Zosteropidae Passeriformes Africa senegalensis 2009a Riegert, 2009 Neoperisterophila Skoracki, 2005 Neoperisterophila Cicinnurus regius Paradisaeidae Passeriformes Australia Skoracki 2005b regiusi Skoracki, 2005 Neosyringophilopsis Skoracki et Sikora, 2005 Neosyringophilopsis Skoracki and acanthizus Skoracki et Acanthiza inornata Acanthizidae Passeriformes Australia Sikora 2005 Sikora, 2005 Bochkov et al. Neosyringophilopsis 2001; Skoracki aegithali (Bochkov, Aegithalos Aegithalidae Passeriformes Europe 2004b; Nattress Mironov et Skoracki, caudatus and Skoracki 2001) 2009 Neosyringophilopsis garrulaxi (Bochkov, Bochkov et al. Garrulax affinis Leiothrichidae Passeriformes Asia Mironov et Skoracki, 2001 2001) Neosyringophilopsis locustellus (Skoracki, Locustella naevia Locustellidae Passeriformes Europe Skoracki 2004b 2004) Locustella “ Locustellidae Passeriformes Europe Skoracki 2004b luscinioides Neosyringophilopsis Spermestes lonchurus Skoracki, Estrildidae Passeriformes Africa Skoracki 2008c cucullatus 2008 Neosyringophilopsis phylloscopi (Bochkov, Phylloscopus Bochkov et al. Phylloscopidae Passeriformes Europe Mironov et Skoracki, collybita 2001 2001) Phylloscopus “ Phylloscopidae Passeriformes Europe Skoracki 2004b trochilus Neosyringophilopsis Fritsch 1958; Troglodytes troglodytis (Fritsch, Troglodytidae Passeriformes Europe Nattress and troglodytes 1958) Skoracki 2007 Bochkov and “ Troglodytes aedon Troglodytidae Passeriformes North America Galloway 2001 Niglarobia Kethley, 1970 Niglarobia cursoriae Cursorius Charadrii- Skoracki et al. Skoracki, Dabert et Glareolidae Africa temminckii formes 2006 Schmaschke, 2006 Niglarobia ereuneti Charadrii- Calidris pusilla Scolopacidae North America Kethley 1970 Kethley, 1970 formes

i21 M. Skoracki, S. Zabludovskaya and A.V. Bochkov

Oudemans 1904, Niglarobia helleri Charadrii- Tringa ochropus Scolopacidae Europe 1906; Kethley (Oudemans, 1904) formes 1970 Charadrii- Skoracki and Tringa flavipes Scolopacidae South America formes Sikora 2002 Charadrii- Skoracki et al. Tringa stagnatilis Scolopacidae Europe formes 2006 Charadrii- Skoracki et al. Arenaria interpres Scolopacidae North America formes 2006 Niglarobia rhinoptili Smutsornis Charadrii- Fain, Bochkov et Glareolidae Africa Fain et al. 2000 africanus formes Mironov, 2000 Oudemans 1904, Niglarobia trouessarti Aramus guarauna Aramidae Gruiformes South America 1906; Kethley (Oudemans, 1904) 1970 Paraniglarobia Skoracki, 2011 Paraniglarobia skorackii Tringa Charadrii- Bochkov and (Bochkov et Galloway, Scolopacidae North America melanoleuca formes Galloway 2004 2004) Paraniglarobia Charadrii- Bochkov and calidridis (Bochkov et Calidris minuta Scolopacidae Asia formes Mironov 1998 Mironov, 1998) Peristerophila Kethley, 1970 Peristerophila accip­ itridicus Skoracki, Skoracki and Accipiter nisus Accipitridae Accipitriformes Asia Lontkowski et Stawarc- Bochkov 2010 zyk, 2010 Terathopius Skoracki et al. “ Accipitridae Accipitriformes Africa ecaudatus 2010d Kethley 1970 Bochkov and Peristerophila columbae North America, Mironov 1998; Columba livia Columbidae Columbiformes (Hirst, 1920) Europe, Asia Nattress and Skoracki 2009; Skoracki 2011 Skoracki and “ Zenaida auriculata Columbidae Columbiformes South America Sikora 2002 “ Buteo jamaicensis Accipitridae Accipitriformes North America Casto 1976 Phalarophilus Skoracki, Bochkov et OConnor, 2011 Phalarophilus fulicarius Phalaropus Charadrii- Skoracki et al. Skoracki, Bochkov et Scolopacidae North America fulicarius formes 2011c OConnor, 2011 Philoxanthornea Kethley, 1970 Philoxanthornea anoa Charadrii- Anous tenuirostris Laridae North America Kethley 1970 Kethley, 1970 formes Philoxanthornea clarki Charadrii- Kivganov and Sternula albifrons Laridae Europe Kivganov, 1995 formes Sharafat 1995 Charadrii- Skoracki et al. “ Sterna hirundo Laridae Europe formes 2006 Charadrii- Bochkov and “ Larus canus Laridae Europe formes Mironov 1998 Philoxanthornea Gelochelidon Charadrii- Bochkov and dubinini Bochkov et Laridae Asia nilotica formes Mironov 1998 Mironov, 1998 Chlidonias Charadrii- Skoracki et al. “ Laridae Africa leucopterus formes 2006

i22 Appendix

Picisyringophilus Skoracki et OConnor, 2010 Picisyringophilus kratos Skoracki and Skoracki et OConnor, Picoides pubescens Picidae Piciformes North America OConnor 2010 2010 Procellariisyringophilus (Kethley, 1970) Procellariisyringophilus Procellarii- bulwerius (Kethley, Bulweria bulwerii Procellariidae North America Kethley 1970 formes 1970) Psittaciphilus Fain, Bochkov et Mironov, 2000 Psittaciphilus amazonae Amazona Fain, Bochkov et Psittacidae Psittaciformes South America Fain et al. 2000 amazonica Mironov, 2000 Bochkov and “ Amazona aestiva Psittacidae Psittaciformes South America Fain 2003 Psittaciphilus fritschi unknown Fain, Bochkov et unidentified parrot Psittacidae Psittaciformes Fain et al. 2000 locality Mironov, 2000 Pteroclidisyringophilus Skoracki, 2011 Pteroclidisyringophilus Pterocles Skoracki and re (Skoracki et OConnor, Pteroclidae Pterocliformes Africa senegallus OConnor 2010 2010) Selenonycha Kethley, 1970 Selenonycha baltoda Charadrius Charadrii- Charadriidae North America Kethley 1970 Kethley, 1970 wilsonia formes Selenonycha chardriifor­ Charadrii- Larus genei Laridae Africa Skoracki 2011 micus Skoracki, 2011 formes Charadrius Charadrii- “ Charadriidae Asia Skoracki 2011 placidus formes Chroicocephalus Charadrii- “ Laridae Europe Skoracki 2011 ridibundus formes Philomachus Charadrii- “ Scolopacidae Europe Skoracki 2011 pugnax formes Stibarokris Kethley, 1970 Stibarokris dastychi Phalacrocorax Phalacrocoraci- unknown Glowska and Glowska et Skoracki, Suliformes carbo dae locality Skoracki 2011a 2011 Stibarokris langei Bochkov and Bochkov et Mironov, Ciconia ciconia Ciconiidae Ciconiiformes Europe Mironov 1999; 1999 Skoracki 2011 Stibarokris phalacrus Phalacrocorax Phalacrocoraci- Suliformes North America Kethley 1970 Kethley, 1970 auritus dae Stibarokris phoenicona­ Phoenicopterus Phoenicopteri- Phoenicopteri- Skoracki and ias Skoracki et OConnor, Africa minor dae formes OConnor 2010 2010 Phoenicopterus Phoenicopteri- Phoenicopteri- European Zoo Glowska and “ ruber dae formes (captive) Skoracki 2011a Syringophiloidus Kethley, 1970 Syringophiloidus agelaius Bochkov, Agelaius Bochkov et al. Icteridae Passeriformes North America Skoracki, Hendricks et phoeniceus 2011 Spicer, 2011 Syringophiloidus Artamus Artamidae Passeriformes Asia Skoracki 2004a artamus Skoracki, 2004 leucorynchus “ Artamus fuscus Artamidae Passeriformes unknown Skoracki 2004a Syringophiloidus Bombycilla Skoracki 2002b; bombycillae Skoracki, Bombycillidae Passeriformes Europe garrulus Skoracki 2004a 2002 i23 M. Skoracki, S. Zabludovskaya and A.V. Bochkov

Syringophiloidus carolae Cardinalis Skoracki et al. Skoracki, Flannery et Cardinalidae Passeriformes North America cardinalis 2009b Spicer, 2009 Melanerpes Skoracki et al. “ Picidae Piciformes North America formicivorus 2009b Bochkov and Syringophiloidus Carpodacus Apanaskevich carpodaci Bochkov et Fringillidae Passeriformes Europe, Asia erythrinus 2001; Skoracki Apanaskevich, 2001 2011 Syringophiloidus Coccothraustes coccothraustes Skoracki, Fringillidae Passeriformes Europe Skoracki 2011 coccothraustes 2011 Syringophiloidus cypsiuri Fain, Bochkov Cypsiurus parvus Apodidae Apodiformes Africa Fain et al. 2000 et Mironov, 2000 Syringophiloidus daberti Bochkov et al. Bochkov, Fain et Passerina ciris Cardinalidae Passeriformes North America 2004 Skoracki, 2004 Syringophiloidus delichonum Bochkov, Delichon urbicum Hirundinidae Passeriformes Europe Bochkov 2001 2001 Syringophiloidus dendrocittae Fain, Dendrocitta Corvidae Passeriformes Asia Fain et al. 2000 Bochkov et Mironov, vagabunda 2000 Syringophiloidus erythrurus Skoracki, Erythrura trichroa Estrildidae Passeriformes Europe Skoracki 2004a 2004 Syringophiloidus Garrulus Fritsch 1958; Corviidae Passeriformes Europe glandarii (Fritsch, 1958) glandarius Bochkov 2001 “ Pica pica Corviidae Passeriformes Asia Skoracki 2004a Skoracki and “ Corvus monedula Corviidae Passeriformes Asia Bochkov 2010 Skoracki and “ Corvus frugilegus Corviidae Passeriformes Asia Bochkov 2010 Syringophiloidus graculae Fain, Bochkov Gracula religiosa Sturnidae Passeriformes Asia Fain et al. 2000 et Mironov, 2000 Syringophiloidus hirundinis Skoracki, Skoracki et al. Hirundo rustica Hirundinidae Passeriformes Europe Moller et Tryjanowski, 2003 2003 Syringophiloidus jackowiaki Bochkov, Poecile Bochkov et al. Paridae Passeriformes North America Skoracki, Hendricks et carolinensis 2011 Spicer, 2011 Syringophiloidus klimovi Skoracki and Skoracki et Bochkov, Chloris chloris Fringillidae Passeriformes Asia Bochkov 2010 2010 Syringophiloidus Microcerculus microcerculus Sikora et Troglodytidae Passeriformes South America Sikora et al. 2012 marginatus Skoracki, 2012 Syringophiloidus minor North America, Kethley 1970; Passer domesticus Passeridae Passeriformes (Berlese, 1887) Europe, Asia Skoracki 2011 Passer Glowska et al. “ Passeridae Passeriformes Asia hispaniolensis 2007 Bochkov and “ Passer montanus Passeridae Passeriformes Europe Mironov 1998

i24 Appendix

Syringophiloidus Skoracki et al. molothrus Skoracki, Molothrus ater Icteridae Passeriformes North America 2009b Flannery et Spicer, 2009 Syringophiloidus montanus Skoracki, Passer montanus Passeridae Passeriformes Europe Skoracki 2002b 2002 Syringophiloidus oriolus Oriolus oriolus Oriolidae Passeriformes Europe Skoracki 2004a Skoracki, 2004 “ Oriolus chinensis Oriolidae Passeriformes Asia Skoracki 2011 Syringophiloidus parapresentalis Turdus merula Turdidae Passeriformes Europe, Asia Skoracki 2011 Skoracki, 2011 “ Turdus pilaris Turdidae Passeriformes Europe Skoracki 2011 “ Turdus iliacus Turdidae Passeriformes Europe Skoracki 2011 “ Turdus atrogularis Turdidae Passeriformes Asia Skoracki 2011 Syringophiloidus petronicus Skoracki, Petronia petronia Passeridae Passeriformes Europe Skoracki 2011 2011 Syringophiloidus philomelosus Skoracki, Turdus philomelos Turdidae Passeriformes Asia Skoracki 2011 2011 Syringophiloidus Chirov and presentalis Chirov et Sturnus vulgaris Sturnidae Passeriformes Europe, Asia Kravtsova 1995; Kravtsova, 1995 Skoracki 2011 Clark 1964b; Syringophiloidus seiuri Seiurus aurocapilla Parulidae Passeriformes North America Bochkov and (Clark, 1964) Galloway 2001 Helmitheros “ Parulidae Passeriformes North America Clark 1964b vermivorum “ Melospiza melodia Emberizidae Passeriformes North America Clark 1964b Skoracki and “ Quelea erythrops Ploceidae Passeriformes Africa Dabert 2002 Skoracki and “ Ploceus baglafecht Ploceidae Passeriformes Africa Dabert 2002 Ploceus Skoracki and “ Ploceidae Passeriformes Africa superciliosus Dabert 2002 Bochkov et al. “ Melospiza lincolnii Emberizidae Passeriformes North America 2011 Oreothlypis Bochkov et al. “ Parulidae Passeriformes North America ruficapilla 2011 Syringophiloidus serini Serinus Bochkov et al. Bochkov, Fain et Fringillidae Passeriformes Africa mozambicus 2004 Skoracki, 2004 Syringophiloidus sialius Skoracki et al. Skoracki, Flannery et Sialia mexicana Turdidae Passeriformes North America 2009b Spicer, 2009 Bochkov et al. “ Spizella breweri Emberizidae Passeriformes North America 2011 Syringophiloidus Euphonia stawarczyki Skoracki, Fringillidae Passeriformes South America Skoracki 2004a cyanocephala 2004 Syringophiloidus tarnii Skoracki and Pteroptochos tarnii Rhinocryptidae Passeriformes South America Skoracki et Sikora, 2002 Sikora 2002 Syringophiloidus Thryothorus Skoracki et al. thryothorus Skoracki, Troglodytidae Passeriformes North America ludovicianus 2009b Flannery et Spicer, 2009

i25 M. Skoracki, S. Zabludovskaya and A.V. Bochkov

Syringophiloidus weiszii Skoracki et al. Skoracki, Hromada et Lanius excubitor Laniidae Passeriformes Europe 2001b Tryjanowski, 2001 Syringophiloidus xanthocephalus Xanthocephalus Bochkov et al. Bochkov, Skoracki, Icteridae Passeriformes North America xanthocephalus 2011 Hendricks et Spicer, 2011 Syringophiloidus zonotrichia Bochkov, Zonotrichia Bochkov et al. Emberizidae Passeriformes North America Skoracki, Hendricks et albicollis 2011 Spicer, 2011 Syringophilopsis Kethley, 1970 Syringophilopsis Acrocephalus acrocephali Skoracki, Acrocephalidae Passeriformes Europe Skoracki 1999a scirpaceus 1999 Acrocephalus “ Acrocephalidae Passeriformes Europe, Africa Skoracki 2011 schoenobaenus Acrocephalus “ Acrocephalidae Passeriformes Europe Skoracki 2011 palustris Syringophilopsis Skoracki and albicollisi Skoracki et Merops albicollis Meropidae Coraciiformes Africa Dabert 2000 Dabert, 2000 Skoracki and Syringophilopsis Dabert 1999a blaszaki Skoracki et Anthus trivialis Motacillidae Passeriformes Europe Bochkov and Dabert, 1999 Galloway 2001 Bochkov and Syringophilopsis borini Mironov 1999; Bochkov et Mironov, Sylvia borin Sylviidae Passeriformes Europe, Asia Skoracki 2004b, 1999 2011 Syringophilopsis Hippolais caligata Acrocephalidae Passeriformes Europe Skoracki 2011 caligatus Skoracki, 2011 Syringophilopsis certhiae Skoracki, Skoracki et al. Certhia americana Certhiidae Passeriformes North America Hendricks et Spicer, 2011a 2011 Syringophilopsis Skoracki and corvinae Skoracki et Corvinella corvina Laniidae Passeriformes Africa Sikora 2003 Sikora, 2003 Syringophilopsis Setophaga Bochkov and dendroicae Bochkov et Parulidae Passeriformes North America coronata Galloway 2001 Galloway, 2001 Skoracki et al. “ Setophaga graciae Parulidae Passeriformes North America 2008 Skoracki et al. “ Cardellina pusilla Parulidae Passeriformes North America 2008 Syringophilopsis dicruri Skoracki et al. Skoracki, Hromada et Dicrurus adsimilis Dicruridae Passeriformes Africa 2011b Wamiti, 2011 Syringophilopsis Agelaius Icteridae Passeriformes North America Kethley 1970 elongatus (Ewing, 1911) phoeniceus Euphagus “ Icteridae Passeriformes North America Clark 1964b carolinus Euphagus Skoracki et al. “ Icteridae Passeriformes North America cyanocephalus 2008 Quiscalus “ Icteridae Passeriformes North America Casto 1976 mexicanus

i26 Appendix

Clark 1964b Syringophilopsis Quiscalus quiscula Icteridae Passeriformes North America Bochkov and elongatus (Ewing, 1911) Galloway 2001 Pheucticus “ Cardinalidae Passeriformes North America Casto 1976 melanocephalus Syringophilopsis emberizae Fain, Sicalis luteola Emberizidae Passeriformes Africa Fain et al. 2000 Bochkov et Mironov, 2000 Syringophilopsis Empidonax Skoracki et al. empidonax Skoracki, Tyrannidae Passeriformes North America hammondi 2008 Flannery et Spicer, 2008 Empidonax Skoracki et al. “ Tyrannidae Passeriformes North America wrightii 2008 Fritsch 1958; Bochkov and Syringophilopsis Mironov 1998; Fringilla coelebs Fringillidae Passeriformes Europe, Asia fringillae (Fritsch, 1958) Skoracki 2004b; Skoracki and Bochkov 2010 Syringophilopsis Garrulus garrulus Skoracki et Corviidae Passeriformes Africa Skoracki 2011 glandarius Dabert, 2002 Skoracki 2004b; Syringophilopsis Hirundo rustica Hirundinidae Passeriformes Europe, Asia Skoracki and hirundus Skoracki, 2004 Bochkov 2010 Syringophilopsis Catharus hylocichlae (Clark, Turdidae Passeriformes North America Clark 1964b fuscescens 1964) Syringophilopsis icteri Bochkov and Bochkov et Galloway, Icterus galbula Icteridae Passeriformes North America Galloway 2001 2001 Skoracki et al. “ Icterus bullocki Icteridae Passeriformes North America 2011a Syringophilopsis Skoracki 2004b; kazmierski Skoracki, Ficedula hypoleuca Muscicapidae Passeriformes Europe 2011 2004 Skoracki 2004b, “ Ficedula parva Muscicapidae Passeriformes Europe 2011 Phoenicurus Glowska et al. “ Muscicapidae Passeriformes Asia phoenicurus 2007 Bochkov et al. Syringophilopsis 2000; Skoracki kirgizorum Bochkov, 2004b; Nattress Chloris chloris Fringillidae Passeriformes Europe, Asia Mironov et Kravtsova, and Skoracki 2000 2007; Glowska et al. 2007 Carduelis Glowska et al. “ Fringillidae Passeriformes Asia cannabina 2007 Skoracki 2004b; “ Carduelis carduelis Fringillidae Passeriformes Europe, Asia 2011 Rhodospiza Bochkov et al. “ Fringillidae Passeriformes Asia obsoleta 2000 Skoracki et al. “ Linurgus olivaceus Fringillidae Passeriformes Africa 2009a Syringophilopsis kristini Skoracki et al. Skoracki, Tryjanowski et Lanius minor Laniidae Passeriformes Europe 2002; Skoracki Hromada, 2002 2011

i27 M. Skoracki, S. Zabludovskaya and A.V. Bochkov

Syringophilopsis Lagonosticta Skoracki and lagonostictus Skoracki et Estrildidae Passeriformes Africa senegala Dabert 2002 Dabert, 2002 Lagonosticta Skoracki and “ Estrildidae Passeriformes Africa rufopicta Dabert 2002 Syringophilopsis Skoracki and melittophagi Skoracki et Merops bulocki Meropidae Coraciiformes Africa Dabert 2001b Dabert, 2001 Syringophilopsis Margarops mimidus Sikora, Fajfer et Mimidae Passeriformes North America Sikora et al. 2011 fuscatus Skoracki, 2011 Syringophilopsis Muscicapa muscicapus Skoracki, Muscicapidae Passeriformes Asia Skoracki 2011 dauurica 2011 Syringophilopsis nitens Skoracki and Malimbus nitens Ploceidae Passeriformes Africa Skoracki et Dabert, 2001 Dabert 2001b Skoracki and “ Quelea erythrops Ploceidae Passeriformes Africa Sikora 2003 Skoracki et al. “ Ploceus vitellinus Ploceidae Passeriformes Africa 2011b Syringophilopsis Nucifraga nucifragus Skoracki, Corviidae Passeriformes Europe Skoracki 2011 caryocatactes 2011 Syringophilopsis passericus Skoracki, Passer domesticus Passeridae Passeriformes Asia Skoracki 2011 2011 Clark 1964b; Syringophilopsis Passerina cyanea Cardinalidae Passeriformes North America Skoracki et al. passerinae (Clark, 1964) 2011a Skoracki et al. “ Passerina ciris Cardinalidae Passeriformes North America 2011a Syringophilopsis Skoracki et al. polioptilus Skoracki, Polioptila caerulea Polioptilidae Passeriformes North America 2008 Flannery et Spicer, 2008 Syringophilopsis rusticus Hirundo rustica Hirundinidae Passeriformes Europe Skoracki 2004b Skoracki, 2004 Syringophilopsis sialiae Skoracki et al. Skoracki, Flannery et Sialia mexicana Turdidae Passeriformes North America 2008 Spicer, 2008 Syringophilopsis sittae Skoracki et al. Skoracki, Hendricks et Sitta carolinensis Sittidae Passeriformes North America 2011a Spicer, 2011 Skoracki 2004b, Syringophilopsis 2011; Skoracki spinolettus Skoracki, Anthus spinoletta Motacillidae Passeriformes Europe, Asia and Bochkov 2004 2010 Syringophilopsis sturnellus Skoracki, Skoracki et al. Sturnella neglecta Icteridae Passeriformes North America Hendricks et Spicer, 2011a 2011 Chirov and Kravtsova 1995; Syringophilopsis sturni Bochkov and Chirov et Kravtsova, Sturnus vulgaris Sturnidae Passeriformes Europe, Asia Mironov 1998; 1995 Skoracki 2004b, 2011 Syringophilopsis turdi Fritsch 1958; Turdus pilaris Turdidae Passeriformes Europe (Fritsch, 1958) Skoracki 2004b

i28 Appendix

Bochkov and Syringophilopsis turdi Turdus philomelos Turdidae Passeriformes Europe Galloway 2001; (Fritsch, 1958) Skoracki 2004b Skoracki and “ Turdus atrogularis Turdidae Passeriformes Asia Bochkov 2010 “ Turdus iliacus Turdidae Passeriformes Europe Skoracki 2011 Skoracki et al. “ Turdus migratorius Turdidae Passeriformes North America 2008 Skoracki and “ Turdus albicollis Turdidae Passeriformes South America Sikora 2003 Syringophilopsis tyranni Bochkov and Bochkov et Galloway, Tyrannus tyrannus Tyrannidae Passeriformes North America Galloway 2004 2004 Skoracki et al. “ Tyrannus verticalis Tyrannidae Passeriformes North America 2008 Skoracki et al. “ Myiarchus crinitus Tyrannidae Passeriformes North America 2008 Myiarchus Skoracki et al. “ Tyrannidae Passeriformes North America cinerascens 2008 Syringophilopsis Pycnonotus Skoracki et al. veselovsky Skoracki, Pycnonotidae Passeriformes Africa barbatus 2009a Antczak et Riegert, 2009 Chlorocichla Skoracki et al. “ Pycnonotidae Passeriformes Africa flaviventris 2011b Skoracki et al. “ Eurillas latirostris Pycnonotidae Passeriformes Africa 2011b Syringophilopsis yosefi Skoracki et al. Skoracki, Tryjanowski et Lanius sp. Laniidae Passeriformes Africa 2002 Hromada, 2002 Syringophilus Heller, 1880 Syringophilus bipectina­ Gallus gallus Phasianidae Galliformes Worldwide Skoracki 2011 tus Heller, 1880 “ Alectoris rufa Phasianidae Galliformes Europe Skoracki 2011 Ptilopachus Skoracki and “ Phasianidae Galliformes Africa petrosus Sikora 2003 Syringophilus numidae Numida meleagris Numididae Galliformes Africa Bochkov 2000c Bochkov, 2000 Terratosyringophilus Bochkov et Perez, 2002 Terratosyringophilus Skoracki and geotrygonus Skoracki et Geotrygon linearis Columbidae Columbiformes South America Glowska 2008 Glowska, 2008 Terratosyringophilus Bochkov and pioni Bochkov et Perez, Pionus senilis Psittacidae Psittaciformes North America Perez 2002 2002 Terratosyringophilus Zenaida asiatica Columbidae Columbiformes North America Casto 1979 longisoma (Casto, 1979) “ Zenaida macroura Columbidae Columbiformes North America Casto 1979 Tinamiphilopsis Skoracki et Sikora, 2004 Tinamiphilopsis ariconte Rhynchotus Skoracki et al. Skoracki, Sikora et Tinamidae Tinamiformes South America rufescens 2012b Ozminski, 2012 Skoracki et al. “ Nothura boraquira Tinamidae Tinamiformes South America 2012b Skoracki et al. “ Nothura minor Tinamidae Tinamiformes South America 2012b

i29 M. Skoracki, S. Zabludovskaya and A.V. Bochkov

Tinamiphilopsis elegans Skoracki and Eudromia elegans Tinamidae Tinamiformes South America Skoracki et Sikora, 2004 Sikora, 2004a Torotrogla Kethley, 1970 Torotrogla aphelocoma Aphelocoma Bochkov et al. Bochkov, Flannery et Corvidae Passeriformes North America californica 2009 Spicer, 2009 Torotrogla calcarius Calcarius Calcariidae Passeriformes Europe Skoracki 2004c Skoracki, 2004 lapponicus Torotrogla cardinalis Cardinalis Bochkov et al. Bochkov, Flannery et Cardinalidae Passeriformes North America cardinalis 2009 Spicer, 2009 Torotrogla cardueli Bochkov and Bochkov et Mironov, Spinus spinus Fringillidae Passeriformes Europe Mironov 1999; 1999 Skoracki 2004c Carduelis Skoracki 2004c, “ Fringillidae Passeriformes Europe cannabina 2011 “ Carduelis carduelis Fringillidae Passeriformes Europe Skoracki 2004c “ Serinus canaria Fringillidae Passeriformes Europe Skoracki 2011 Loxia curvirostra Fringillidae Passeriformes Europe Skoracki 2011 “ Loxia leucoptera Fringillidae Passeriformes Europe Skoracki 2011 Loxia “ Fringillidae Passeriformes Europe Skoracki 2011 pytyopsittacus Torotrogla cocco­ Coccothraustes Bochkov et al. thraustes Bochkov, Fringillidae Passeriformes North America vespertinus 2009 Flannery et Spicer, 2009 Torotrogla cyanocitta Bochkov et al. Bochkov, Flannery et Cyanocitta stelleri Corvidae Passeriformes North America 2009 Spicer, 2009 Torotrogla gaudi Bochkov and Bochkov et Mironov, Fringilla coelebs Fringillidae Passeriformes Europe Mironov 1998 1998 Fringilla “ Fringillidae Passeriformes Europe Skoracki 2011 montifringilla “ Pyrrhula pyrrhula Fringillidae Passeriformes Europe Skoracki 2011 Torotrogla lullulae Skoracki et al. Skoracki, Hromada et Lullula arborea Alaudidae Passeriformes Europe 2001a Kuczynski, 2001 Eremophila “ Alaudidae Passeriformes Asia Skoracki 2011 alpestris Torotrogla lusciniae Luscinia Muscicapidae Passeriformes Europe Skoracki 2004c Skoracki, 2004 megarhynchos “ Luscinia svecica Muscicapidae Passeriformes Europe Skoracki 2004c Skoracki et al. Torotrogla merulae 2000; Bochkov Skoracki, Dabert et Turdus merula Turdidae Passeriformes Europe, Asia et al. 2009; Ehrnsberger, 2000 Glowska et al. 2007 “ Turdus viscivorus Turdidae Passeriformes Europe Skoracki 2004c Skoracki 2004c; “ Turdus philomelos Turdidae Passeriformes Europe, Asia Glowska et al. 2007 “ Turdus torquatus Turdidae Passeriformes Europe Skoracki 2011 Torotrogla mima Mimus polyglottos Mimidae Passeriformes North America Kethley 1970 Kethley, 1970 “ Mimus triurus Mimidae Passeriformes South America Sikora et al. 2011 “ Mimus patagonicus Mimidae Passeriformes Africa Sikora et al. 2011

i30 Appendix

Torotrogla modularis Nattress and Nattress et Skoracki, Prunella modularis Prunellidae Passeriformes Europe Skoracki 2007; 2007 Skoracki 2011 Torotrogla piranga Piranga Bochkov et al. Bochkov, Flannery et Cardinalidae Passeriformes North America ludoviciana 2009 Spicer, 2009 Torotrogla pycnonotus Pycnonotus Pycnonotidae Passeriformes Asia Skoracki 2011 Skoracki, 2011 xanthopygos Skoracki 2004c; Torotrogla rubeculi Erithacus rubecula Muscicapidae Passeriformes Europe Bochkov et al. Skoracki, 2004 2009 Torotrogla villosa Phainopepla nitens Ptilogonatidae Passeriformes North America Kethley 1970 (Hancock, 1895) Trypetoptila Kethley, 1970 Trypetoptila casmerodia Ardea alba Ardeidae Pelecaniformes North America Kethley 1970 Kethley, 1970 Species inquirenda Syringophilopsis Liu 1988; Passer domesticus Passeridae Passeriformes Asia hunanensis Liu, 1988 Skoracki 2011 Picobiinae Johnston et Kethley, 1973 Calamincola Casto, 1977 Calamincola lobata Crotophaga Cuculidae Cuculiformes North America Casto 1977 (Casto, 1977) sulcirostris Columbiphilus Kivganov et Sharafat, 1995 Columbiphilus alectoris Fain et al. 2000; (Fain, Bochkov et Alectoris sp. Phasianidae Galliformes Africa Skoracki 2011 Mironov, 2000) Skoracki and “ Alectoris rufa Phasianidae Galliformes Europe Sikora 2011 Columbiphilus khush­ Kivganov and alkhani Kivganov et Columba livia Columbidae Columbiformes Asia Sharafat 1995 Sharafat, 1995 Columbiphilus polonica Skoracki et al. (Skoracki, Magowski et Gallus gallus Phasianidae Galliformes Europe 2001; Skoracki Dabert, 2001) 2011 Skoracki and “ Gallus sonneratii Phasianidae Galliformes Asia Sikora 2011 Columbiphilus pteroclesi Pterocles Skoracki and (Skoracki et OConnor, Pteroclidae Pterocliformes Africa senegallus OConnor 2010 2010) Pterocles Skoracki and “ Pteroclidae Pterocliformes Africa coronatus OConnor 2010 Neopicobia Skoracki, 2011 Neopicobia anthi Anthus trivialis Motacillidae Passeriformes Europe Fritsch 1958 (Fritsch, 1958) “ Anthus cervinus Motacillidae Passeriformes Europe Skoracki 2011 Neopicobia cardinalis Cardinalis Skoracki et al. (Skoracki, Hendricks et Cardinalidae Passeriformes North America cardinalis 2010c Spicer, 2010) Neopicobia carpodacus Carpodacus Skoracki et al. (Skoracki, Hendricks et Fringillidae Passeriformes North America purpureus 2010c Spicer, 2010) Neopicobia epthianura Epthianura Skoracki et al. (Skoracki, Glowska et Meliphagidae Passeriformes Australia aurifrons 2008 Sikora, 2008)

i31 M. Skoracki, S. Zabludovskaya and A.V. Bochkov

Neopicobia glossopsitta Glossopsitta Skoracki et al. (Skoracki, Glowska et Psittacidae Psittaciformes Australia porphyrocephala 2008 Sikora, 2008) Neopicobia modularis Skoracki and (Skoracki et Magowski, Prunella modularis Prunellidae Passeriformes Europe Magowski 2001 2001) Prunella “ Prunellidae Passeriformes Asia Skoracki 2011 atrogularis Neopicobia locustella Skoracki et al. (Skoracki, Bochkov et Locustella naevia Locustellidae Passeriformes Europe 2004b Wauthy, 2004) Locustella “ Locustellidae Passeriformes Europe Skoracki 2011 luscinioides Neopicobia pyrrholae­ Pyrrholaemus Skoracki and mus (Skoracki et Acanthizidae Passeriformes Australia sagittatus Glowska 2008c Glowska, 2008) Neopicobia troglodytes Skoracki et al. (Skoracki, Hendricks et Troglodytes aedon Troglodytidae Passeriformes North America 2010c Spicer, 2010) Neopicobia zumpti Streptopelia Columbidae Columbiformes Africa Lawrence 1959a (Lawrence, 1959) capicola Streptopelia “ Columbidae Columbiformes Africa Lawrence 1959a senegalensis Bochkov et al. “ Columba livia Columbidae Columbiformes North America 2005 Picobia Haller, 1878 Picobia biarmicus Skoracki et al. Skoracki, Bochkov et Panurus biarmicus Panuridae Passeriformes Europe 2004b Wauthy, 2004 Picobia brotogeris Fain, Brotogeris Bochkov et Mironov, Psittacidae Psittaciformes South America Fain et al. 2000 jugularis 2000 inc. sedis Picobia caudati Aegithalos Skoracki and Aegithalidae Passeriformes Europe Skoracki et Hebda, 2004 caudatus Hebda 2004 Picobia cissa Skoracki, Skoracki et al. Bochkov et Wauhty, Cissa chinensis Corviidae Passeriformes Asia 2004b 2004 Picobia cetti Skoracki, Cettia cetti Cettidae Passeriformes Asia Skoracki 2011 2011 Picobia chloris Bochkov et al. Bochkov, Mironov et Chloris chloris Fringillidae Passeriformes Asia 2000 Kravtsova, 2000 Picobia corcoracus Corcorax Skoracki et al. Skoracki, Glowska et Corcoracidae Passeriformes Australia melanorhamphos 2008 Sikora, 2008 Picobia currucae Skoracki and Skoracki et Magowski, Sylvia curruca Sylviidae Passeriformes Europe, Asia Magowski 2001; 2001 Skoracki 2011 Glowska et al. “ Sylvia hortensis Sylviidae Passeriformes Asia 2007 “ Sylvia nisoria Sylviidae Passeriformes Asia Skoracki 2011 Picobia dinemellia Dinemellia Glowska and Glowska et Skoracki, Ploceidae Passeriformes Africa dinemelli Skoracki 2011b 2011 Fritsch 1956; Picobia dryobatis Dendrocopos Picidae Piciformes Europe Skoracki et al. (Fritsch, 1958) major 2004b

i32 Appendix

Picobia dryobatis Dendrocopos Picidae Piciformes Europe Skoracki 2011 (Fritsch, 1958) minor hortorum Dendrocopos “ Picidae Piciformes Europe Skoracki 2011 leucotos Skoracki et al. Picoides North America, “ Picidae Piciformes 2010c; Skoracki tridactylus Europe 2011 Skoracki et al. “ Picoides scalaris Picidae Piciformes North America 2010c Picobia dziabaszewskii Glowska et al. Glowska, Dragun-Dami- Garrulax formosus Leiothrichidae Passeriformes Asia 2012 an et Dabert, 2012 Picobia eremophila Eremophila Alaudidae Passeriformes Africa Skoracki 2011 Skoracki, 2011 bilopha Picobia galerida Galerida cristata Alaudidae Passeriformes Europe Skoracki 2011 Skoracki, 2011 Picobia heeri Haller, Picus canus Picidae Piciformes Europe Skoracki 2011 1878 “ Picus viridis Picidae Piciformes Europe Skoracki 2011 Picobia indonesiana Skoracki and Skoracki et Glowska, Aplonis panayensis Sturnidae Passeriformes Asia Glowska 2008c 2008 Picobia lemi Skoracki, Manucodia Skoracki et al. Paradisaeidae Passeriformes Australia Glowska et Sikora, 2008 chalybatus 2008 Picobia leucophaeus Leucophaeus Charadrii- Skoracki et al. Skoracki, Hendricks et Laridae North America atricilla formes 2010c Spicer, 2010 inc. sedis Picobia oritis Skoracki, Skoracki et al. Cyanomitra oritis Nectarinidae Passeriformes Africa Antczak et Riegert, 2009 2009 Cyanomitra Skoracki et al. “ Nectarinidae Passeriformes Africa olivacea 2011 Picobia paludicola Acrocephalus Skoracki and Acrocephalidae Passeriformes Europe Skoracki et Kiljan, 2002 paludicola Kiljan 2002 Picobia phoeniculi Fain, Phoeniculus Bochkov et Mironov, Phoeniculidae Coraciiformes Africa Fain et al. 2000 purpureus 2000 Picobia poicephali Poicephalus Skoracki and Skoracki et Dabert, 2002 Psittacidae Psittaciformes Africa senegalus Dabert 2002 inc. sedis Picobia pycnonoti Pycnonotus Glowska et al. Glowska, Skoracki et Pycnonotidae Passeriformes Asia xanthopygos 2007 Khourly, 2007 Picobia riparius Riparia riparia Hirundinidae Passeriformes Europe Skoracki 2011 Skoracki, 2011 Picobia sturni Skoracki, Skoracki et al. Bochkov et Wauthy, Sturnus vulgaris Sturnidae Passeriformes Europe 2004b 2004 “ Sturnus cineraceus Sturnidae Passeriformes Asia Skoracki 2011 Picobia troglodytidus Henicorhina Troglodytidae Passeriformes North America Sikora et al. 2012 Sikora et Skoracki, 2012 leucophrys Rafapicobia Skoracki, 2011 Rafapicobia zirnitra Saxicola rubetra Muscicapidae Passeriformes Europe Skoracki 2011 Skoracki, 2011 Rafapicobia toxostoma Toxostoma Sikora, Fajfer et Mimidae Passeriformes North America Sikora et al. 2011 curvirostre Skoracki, 2011

i33 M. Skoracki, S. Zabludovskaya and A.V. Bochkov

Rafapicobia dendroco­ Dendrocolaptes Skoracki and laptesi Skoracki et Funariidae Passeriformes South America platyrostris Solarczyk 2012 Solarczyk, 2012 Dendrocolaptes Skoracki and “ Funariidae Passeriformes South America picumnus Solarczyk 2012 Rafapicobia lepidoco­ Lepidocolaptes Skoracki and laptesi Skoracki et Funariidae Passeriformes South America affinis Solarczyk 2012 Solarczyk, 2012 Lepidocolaptes Skoracki and “ Funariidae Passeriformes South America souleyetii Solarczyk 2012 Ereynetidae Oudemans, 1931 Speleognathinae Fain, 1957 Boydaiini Fain, 1985 Boydaia Womersley, 1953 Boyd 1948; Fain1961a; Fain 1963b; Domrow Eurasia, Boydaia sturni (Boyd, 1969; Sixl 1972; Sturnus vulgaris Sturnidae Passeriformes Australia, North 1948) Pence 1973a, America 1975; Zablu- dovskaya 1998; Knee et al. 2008 “ Acridotheres tristis Sturnidae Passeriformes Australia Domrow 1969 Ampeliceps “ Sturnidae Passeriformes Europe Fain 1971 coronatus Zabludovskaya “ Pastor roseus Sturnidae Passeriformes Asia 1998 Porter and “ Passer domesticus Ploceidae Passeriformes North America Strandtmann 1952 Fain and Aitken Boydaia agelaii Fain et Chrysomus Icteridae Passeriformes South America 1968; Fain and Aitken, 1968 icterocephalus Lukoschus 1972 Fain and Aitken Agelaius “ Icteridae Passeriformes North America 1968; Pence phoeniceus 1973 Chrysomus European Zoo Fain and Aitken “ Icteridae Passeriformes ruficapillus (captive) 1968 Dusbabek and “ Agelaius humeralis Icteridae Passeriformes South America Cerny 1970 Dives Dusbabek and “ Icteridae Passeriformes South America atroviolaceus Cerny 1970 Fain and Hyland “ Molothrus ater Icteridae Passeriformes North America 1975 Quiscalus Fain and Hyland “ Icteridae Passeriformes North America mexicanus 1975 Fain and Hyland “ Spiza americana Cardinalidae Passeriformes North America 1975 Boydaia americana Fain, European Zoo Passerina caerulea Cardinalidae Passeriformes Fain 1963b 1963 (captive) Cyanocompsa Fain and Aitken “ Cardinalidae Passeriformes South America cyanoides 1970 European Zoo Fain 1963b; “ Piranga rubra Cardinalidae Passeriformes (captive) Pence 1975 Ramphocelus Fain and Aitken “ Thraupidae Passeriformes South America carbo 1968

i34 Appendix

Boydaia americana Fain, Fain and Aitken Tangara gyrola Thraupidae Passeriformes South America 1963 1970 Fain and Aitken “ Coereba flaveola Coerebidae Passeriformes South America 1968 Sporophila Fain and Aitken “ Emberizidae Passeriformes South America americana 1970 Fain and Aitken “ Volatinia jacarina Emberizidae Passeriformes South America 1968 Boydaia bradornis Fain, Bradornis pallidus Muscicapidae Passeriformes Africa Fain 1956b, 1971 1956 griseus Zabludovskaya “ Luscinia luscinia Muscicapidae Passeriformes Europe 1998 Boydaia buphagi Fain, Buphagus Buphagidae Passeriformes Africa Fain 1961a 1961 africanus Lamprotornis “ Sturnidae Passeriformes Africa Fain 1963b chloropterus Boydaia cecropis Fain, Cecropis Hirundinidae Passeriformes Africa Fain 1969, 1971 1969 abyssinica Boydaia cinnyris Fain, Cinnyris cupreus Nectariniidae Passeriformes Africa Fain 1969, 1971 1969 Chalcomitra “ Nectariniidae Passeriformes Africa Fain 1969, 1971 senegalensis Boydaia cyanerpes Fain, European Zoo Cyanerpes cyaneus Thraupidae Passeriformes Fain 1963b 1963 (captive) Boydaia cyanerpes Fain and Hyland hylocichla Fain et Catharus ustulatus Turdidae Passeriformes North America 1975 Hyland, 1975 Dusbabek and Boydaia faini Dusbabek Setophaga cerulea Parulidae Passeriformes South America Cerny 1970; et Cerny, 1970 Pence 1975 Boydaia formicarii Fain Fain and Aitken Formicarius colma Formicariidae Passeriformes South America et Aitken, 1970 1970 Fain and Aitken “ Formicarius analis Formicariidae Passeriformes South America 1970 Fain 1963a; Fain Africa, Europe, and Hyland Boydaia hirundoae Fain, North America, 1975; Domrow, Hirundo rustica Hirundinidae Passeriformes 1956 Australia 1969; Sixl 1972; (introduced) Pence 1973a, 1975 Stelgidopteryx Fain and Aitken “ Hirundinidae Passeriformes South America ruficollis 1968 Fain and Hyland “ Progne subis Hirundinidae Passeriformes South America 1970 “ Hirundo neoxena Hirundinidae Passeriformes Australia Domrow 1965 “ Petrochelidon ariel Hirundinidae Passeriformes Australia Domrow 1969 Cecropis “ Hirundinidae Passeriformes Africa Zumpt 1961 abyssinica “ Hirundo aethiopica Hirundinidae Passeriformes Africa Fain et al. 1977 Boydaia indica Fain, European Zoo Acridotheres tristis Sturnidae Passeriformes Fain 1969 1969 (captive) European Zoo “ Gracupica contra Sturnidae Passeriformes Fain 1969 (captive) European Zoo “ Sturnia malabarica Sturnidae Passeriformes Fain 1969 (captive)

i35 M. Skoracki, S. Zabludovskaya and A.V. Bochkov

Boydaia indica Fain, Temenuchus European Zoo Sturnidae Passeriformes Fain 1969 1969 pagodarum (captive) Lamprotornis “ Sturnidae Passeriformes Africa Fain 1971 chloropterus Lamprotornis European Zoo “ Sturnidae Passeriformes Fain 1971 chalybaeus (captive) Lamprotornis European Zoo “ Sturnidae Passeriformes Fain 1971 superbus (captive) Boydaia jordani Van Van Endhoven Turdus pilaris Turdidae Passeriformes Europe Eyndhoven, 1955 1955 Clark 1967; “ Turdus migratorius Turdidae Passeriformes North America Pence 1975 Zabludovskaya “ Turdus viscivorus Turdidae Passeriformes Europe 1998 Boydaia laticoxa Fain et Glyphorynchus Fain and Aitken Funariidae Passeriformes South America Aitken, 1969 spirurus 1969, 1970 Boydaia laticoxa Fain and Aitken philydori Fain et Aitken, Philydor pyrrhodes Furnariidae Passeriformes South America 1970 1970 Boydaia locustellae Locustella Locustellidae Passeriformes Europe Fain 1970 Fain, 1970 luscinioides Boydaia loxiae Fain, Loxia curvirostra Fringillidae Passeriformes Europe Fain 1963a 1963 Fain and Hyland “ Icterus galbula Icteridae Passeriformes North America 1975 Boydaia maluri Malurus amabilis Maluridae Passeriformes Australia Domrow 1969 Domrow, 1969 Malurus “ Maluridae Passeriformes Australia Domrow 1969 melanocephalus Boydaia mimi Fain et Fain and Hyland Mimus gilvus Mimidae Passeriformes South America Hyland, 1970 1970 Pence 1973a, “ Mimus polyglottos Mimidae Passeriformes North America 1975 Boydaia morenoi Dusbabek and Mimus polyglottos Mimidae Passeriformes South America Dusbabek et Cerny, 1970 Cerny 1970 Boydaia myzomelae Myzomela Meliphagidae Passeriformes Australia Domrow 1969 Domrow, 1969 sanguinolenta Xanthotis “ Meliphagidae Passeriformes Australia Domrow 1991 flaviventer Boydaia nectarinia Fain, Cyanomitra Fain 1958b, Nectariniidae Passeriformes Africa 1958 verticalis 1963b Boydaia pheucticola Pheucticus Pence and Costa Cardinalidae Passeriformes North America Pence et Costa, 1976 melanocephalus 1976 “ Spiza americana Cardinalidae Passeriformes North America Spicer 1987 Boydaia psalidoprocnei Psalidoprocne Hirundinidae Passeriformes Africa Fain 1956b, 1971 Fain, 1956 albiceps Pence 1973a, Tachycineta “ Hirundinidae Passeriformes North America 1975; Knee et al. bicolor 2008 Zabludovskaya “ Riparia riparia Hirundinidae Passeriformes Asia 1998 Boydaia pycnonoti Fain, Pycnonotus Fain 1956b; Fain Pycnonotidae Passeriformes Africa 1956 barbatus et al. 1977 Pycnonotus “ Pycnonotidae Passeriformes Asia Fain 1963b goiavier

i36 Appendix

Boydaia pycnonoti Fain, Pycnonotus Pycnonotidae Passeriformes Asia Fain 1963b, 1971 1956 leucogenys Clark 1960; Boydaia quiscali Clark, Quiscalus quiscula Icteridae Passeriformes North America Pence 1973a, 1960 1975 Quiscalus Pence 1973a, “ Icteridae Passeriformes North America mexicanus 1975 Pence 1973a, “ Molothrus ater Icteridae Passeriformes North America 1975 Agelaius Pence 1973a, “ Icteridae Passeriformes North America phoeniceus 1975 Fain and Aitken “ Saltator maximus Thraupidae Passeriformes South America 1970 Saltator Fain and Hyland “ Thraupidae Passeriformes South America coerulescens 1970 Dusbabek and Boydaia rosickyi Passerina cyanea Cardinalidae Passeriformes South America Cerny 1970; Dusbabek et Cerny 1970 Pence 1975 Boydaia saxicolae Fain, Saxicola torquatus Muscicapidae Passeriformes Africa Fain 1969 1969 axillaris Myrmecocichla “ Muscicapidae Passeriformes Africa Fain 1969 nigra Monticola “ Muscicapidae Passeriformes Africa Fain 1969 angolensis Boydaia sinensis Fain et Fain and Bafort Leiothrix lutea Leiothrichidae Passeriformes Asia Bafort, 1963 1963 Boydaia spatulata Fain, Cercotrichas Fain, 1955, Muscicapidae Passeriformes Africa 1955 hartlaubi 1956b Pycnonotus Fain 1956a, “ Pycnonotidae Passeriformes Africa barbatus 1956b, 1971 Cossypha Fain 1956a, “ Muscicapidae Passeriformes Africa natalensis 1956b, 1971 Cossypha Fain, 1956b, “ Muscicapidae Passeriformes Africa polioptera 1963b Pogonocichla Fain 1956a, “ Muscicapidae Passeriformes Africa stellata 1956b, 1971 Fain 1956a, “ Cercomela tractrac Muscicapidae Passeriformes Africa 1956b, 1971 Fain 1956a, “ Chloropeta similis Acrocephalidae Passeriformes Africa 1956b, 1971 Schistolais Fain 1956a, “ Cisticolidae Passeriformes Africa leucopogon 1956b, 1971 Fain 1956a, “ Melaniparus niger Paridae Passeriformes Africa 1956b, 1971 Fain 1956a, “ Macronyx croceus Motacillidae Passeriformes Africa 1956b, 1971 European Zoo “ Tiaris canorus Emberizidae Passeriformes Fain 1958a (captive) European Zoo “ Tangara sp. Thraupidae Passeriformes Fain 1958a (captive) Cissomela “ Meliphagidae Passeriformes Australia Domrow 1969 pectoralis Philemon “ Meliphagidae Passeriformes Australia Domrow 1969 corniculatus Philemon “ Meliphagidae Passeriformes Australia Domrow 1969 citreogularis

i37 M. Skoracki, S. Zabludovskaya and A.V. Bochkov

Boydaia spatulata Fain, Cyssomela Meliphagidae Passeriformes Australia Domrow 1969 1955 pectoralis Pence 1973c, “ Sialia sialis Turdidae Passeriformes North America 1975 Pycnonotus Zabludovskaya “ Pycnonotidae Passeriformes Africa barbatus 2002 Boydaia synallaxis Fain, Synallaxis rutilans Furnariidae Passeriformes South America Fain, 1969 Hyland et Aitken, 1969 Synallaxis “ Furnariidae Passeriformes South America Fain, 1969 gujanensis Synallaxis “ Furnariidae Passeriformes South America Fain, 1969 erythrothorax Fain and Hyland “ Empidonax sp. Tyrannidae Passeriformes South America 1970 Boydaia tyrannis Ford, Tyrannus tyrannus Tyrannidae Passeriformes North America Ford 1959 1959 Myiophobus Fain and Aitken “ Tyrannidae Passeriformes South America fasciatus 1968 Brooks and Strandtmann “ Tyrannus verticalis Tyrannidae Passeriformes North America 1960; Pence 1975 Fain and Aitken “ Elaenia flavogaster Tyrannidae Passeriformes South America 1968 Pyrocephalus Fain and Hyland “ Tyrannidae Passeriformes South America rubinus 1970 Myiarchus Pence and Casto “ Tyrannidae Passeriformes North America cinerascens 1976 Dusbabek and Tyrannus Cerny 1970; Fain “ Tyrannidae Passeriformes South America dominicensis et al. 1975; Pence 1975 Pachyramphus Fain and Hyland “ Tityridae Passeriformes South America aglaiae 1970 Fain and Aitken “ Manacus manacus Pipridae Passeriformes South America 1968, 1970 Fain and Aitken “ Pipra pipra Pipridae Passeriformes South America 1970c Fain and Aitken “ Pipra fasciicauda Pipridae Passeriformes South America 1970 Fain and Aitken “ Lepidothrix iris Pipridae Passeriformes South America 1970 Boydaia zosteropis Fain, Zosterops Zosteropidae Passeriformes Africa Fain 1963b, 1971 1963 senegalensis “ Zosterops lateralis Zosteropidae Passeriformes Australia Domrow 1969 Boydaia zumpti Fain, Eurillas latirostris Fain 1955, Pycnonotidae Passeriformes Africa 1955 eugenius 1956a, 1956b European Zoo “ Bleda syndactylus Pycnonotidae Passeriformes Fain 1971 (captive) Schoenicola “ Locustellidae Passeriformes Africa Fain 1958a, 1971 brevirostris Berlese and Trouessart 1889; Boydaia crassipes Europe, Porter and (Berlese et Trouessart, Passer domesticus Passeridae Passeriformes Australia, North Strandtmann, 1889) America 1952; Domrow 1969

i38 Appendix

Boydaia crassipes (Berlese et Trouessart, Carduelis carduelis Fringillidae Passeriformes Australia Domrow 1969 1889) Australia “ Motacilla flava Fringillidae Passeriformes Domrow 1969 (introduced) “ Lalage sueurii Campephagidae Passeriformes Australia Domrow 1991 Boydaia psittaci Fain et Pionites European Zoo Psittacidae Psittaciformes Fain 1963b Mortelmans, 1959 melanocephalus (captive) Pionites “ Psittacidae Psittaciformes South America Fain 1963b leucogaster Bolborhynchus European Zoo “ Psittacidae Psittaciformes Fain 1963b lineola (captive) European Zoo “ Forpus passerinus Psittacidae Psittaciformes Fain 1971 (captive) Brotogeris European Zoo “ Psittacidae Psittaciformes Fain 1971 jugularis (captive) Brotogeris European Zoo “ Psittacidae Psittaciformes Fain 1971 versicolurus (captive) Psilopsiagon European Zoo “ Psittacidae Psittaciformes Fain 1971 aymara (captive) Boydaia aratingae Fain, European Zoo Aratinga jandaya Psittacidae Psittaciformes Fain 1963b, 1971 1963 (captive) Boydaia falconis Fain, Falco cuvierii Falconidae Falconiformes Africa Fain 1956b 1956 “ Falco tinnunculus Falconidae Falconiformes Africa Fain 1963b “ Falco berigora Falconidae Falconiformes Australia Domrow 1969 Pence and Casto “ Falco sparverius Falconidae Falconiformes North America 1976 Boydaia podargi Fain et Podargus Caprimulgi- Fain and Podargidae Australia Lukoschus, 1979 strigoides formes Lukoschus 1979 Boydaia colini Clark, Clark 1958; Fain Colinus virginianus Odontophoridae Galliformes North America 1958 and Hyland 1975 European Zoo Fain 1963b; Boydaia clarki Fain, Callipepla Odontophoridae Galliformes (captive); North Pence and Casto 1963 squamata America 1976 Coboydaia Fain, 1985 Coboydaia clavata Fain 1955, Ploceus cucullatus Ploceidae Passeriformes Africa (Fain, 1955) 1956a, 1963b Fain 1956a, “ Ploceus xanthops Ploceidae Passeriformes Africa 1963b, 1971 Fain 1956a, “ Ploceus pelzelni Ploceidae Passeriformes Africa 1963b, 1971 Fain. 1956a, 1960, 1963b, “ Quelea quelea Ploceidae Passeriformes Africa 1971; Fain et al. 1977 Fain 1956a, “ Ploceus baglafecht Ploceidae Passeriformes Africa 1963b, 1971 “ Ploceus vitellinus Ploceidae Passeriformes Africa Fain et al. 1977 “ Ploceus cucullatus Ploceidae Passeriformes Africa Fain et al. 1977 Euplectes “ Ploceidae Passeriformes Africa Fain et al. 1977 hordeaceus “ Euplectes ardens Ploceidae Passeriformes Africa Fain et al. 1977 Euplectes “ Ploceidae Passeriformes Africa Fain et al. 1977 macroura

i39 M. Skoracki, S. Zabludovskaya and A.V. Bochkov

Fain 1956a, 1960, 1963b, Coboydaia clavata Quelea erythrops Ploceidae Passeriformes Africa 1971; Fain et al. (Fain, 1955) 1977; Mwase and Baker 2006 “ Vidua chalybeata Viduidae Passeriformes Africa Fain 1963b, 1971 Coboydaia amandavae Sporaeginthus European Zoo Estrildidae Passeriformes Fain 1962a (Fain, 1962) subflavus (captive) Amandava “ Estrildidae Passeriformes Asia Fain 1963b amandava European Zoo “ Erythrura prasina Estrildidae Passeriformes Fain1970 (captive) Malurus “ Maluridae Passeriformes Australia Domrow 1969 melanocephalus Coboydaia nigra (Fain, Serinus sulphura­ Fain 1955, Fringillidae Passeriformes Africa 1955) tus shelleyi 1956a, 1956b Serinus Fain 1956b, “ Fringillidae Passeriformes Africa citrinelloides 1963b, 1971 Fain 1963b, Serinus “ Fringillidae Passeriformes Africa 1971; Fain et al. mozambicus 1977 Fain, 1963b, “ Serinus burtoni Fringillidae Passeriformes Africa 1971 Fain 1962b, “ Carduelis carduelis Fringillidae Passeriformes Europe 1963b, 1971; Sixl 1972 Carpodacus Fain and Hyland “ Fringillidae Passeriformes North America mexicanus 1975 Carpodacus Zabludovskaya “ Fringillidae Passeriformes Eurasia erythrinus 1998 Fringilla Zabludovskaya “ Fringillidae Passeriformes Asia montifringilla 1998 Sixl 1972; “ Fringilla coelebs Fringillidae Passeriformes Eurasia Zabludovskaya 1998 “ Passer griseus Passeridae Passeriformes Africa Fain 1963b, 1971 Zabludovskaya “ Passer montanus Passeridae Passeriformes Asia 1998 Passer Zabludovskaya “ Passeridae Passeriformes Asia ammodendri 1998 Passer Zabludovskaya “ Passeridae Passeriformes Asia hispaniolensis 1998 Fain 1963b, “ Emberiza rutila Emberizidae Passeriformes Africa, Asia 1971; Zablu- dovskaya 1998 Zabludovskaya “ Emberiza tristrami Emberizidae Passeriformes Asia 1998 Emberiza Zabludovskaya “ Emberizidae Passeriformes Asia spodocephala 1998 Zabludovskaya “ Emberiza pusilla Emberizidae Passeriformes Asia 1998 Zabludovskaya “ Emberiza rustica Emberizidae Passeriformes Asia 1998 Zabludovskaya “ Emberiza citrinella Emberizidae Passeriformes Asia 1998

i40 Appendix

Coboydaia nigra (Fain, Zabludovskaya Emberiza aureola Emberizidae Passeriformes Asia 1955) 1998 Fain and Hyland “ Spizella passerina Emberizidae Passeriformes North America 1975 Fain and Aitken “ Euphonia violacea Fringillidae Passeriformes South America 1968 “ Anthus spinoletta Motacillidae Passeriformes North America Pence 1973 Zabludovskaya “ Anthus trivialis Motacillidae Passeriformes Eurasia 1998 Coboydaia nigra icteri Fain and Hyland Icterus spurius Icteridae Passeriformes South America (Fain et Hyland, 1970) 1970 Fain and Hyland Icterus galbula Icteridae Passeriformes North America 1970 Fain 1969, 1971; Coboydaia nigra Motacilla flava Motacillidae Passeriformes Africa, Eurasia Zabludovskaya motacillae (Fain, 1969) 1998 “ Motacilla aguimp Motacillidae Passeriformes Africa Fain 1969, 1971 “ Motacilla capensis Motacillidae Passeriformes Africa Fain 1969, 1971 “ Anthus trivialis Motacillidae Passeriformes Europe Fain 1963a, 1971 Acrocephalus “ Acrocephalidae Passeriformes Europe Fain 1963b, 1971 arundinaceus Acrocephalus “ Acrocephalidae Passeriformes Europe Fain 1963b, 1971 scirpaceus Zabludovskaya “ Motacilla alba Motacillidae Passeriformes Asia 1998 Coboydaia sturnellae Sturnella magna Icteridae Passeriformes North America Clark 1960 (Clark, 1960) Coboydaia (Apodiboydaia) Fain, 1985 Fain, 1958a, Coboydaia trochila European Zoo Chlorestes notata Trochilidae Apodiformes 1961a, 1963b, (Fain, 1958) (captive) 1971 Trispeleognathini Fain, 1985 Trispeleognathus Fain, 1958 North America, Trispeleognathus striatus Crossley 1952; Columba livia Columbidae Columbiformes Australia (Crossley, 1952) Domrow 1969 (introduced) Streptopelia “ Columbidae Columbiformes Africa Fain 1956a semitorquata Columbina Fain and Aitken “ Columbidae Columbiformes South America talpacoti 1968 “ Leptotila verreauxi Columbidae Columbiformes South America Amaral 1963 Aureliania Fain, 1958 Fain 1955; Africa, Aureliania aureliani Dusbabek and Tyto alba Tytonidae Strigiformes Australia, South (Fain, 1955) Cerny 1970; America Domrow 1991 Ophthalmognathus Dubinin, 1957 Ophthalmognathus Ardeola idae Ardeidae Pelecaniformes Africa Fain 1955, 1956a shoutedeni (Fain, 1955) “ Ardea cinerea Ardeidae Pelecaniformes Europe Dubinin 1957 Nycticorax “ Ardeidae Pelecaniformes Africa Fain 1955, 1956a nycticorax Nycticorax Domrow 1969, “ Ardeidae Pelecaniformes Australia caledonicus 1991

i41 M. Skoracki, S. Zabludovskaya and A.V. Bochkov

Ophthalmognathus Nycticorax Australia Fain and Goff tenorioae Fain et Goff, Ardeidae Pelecaniformes nycticorax (Hawaii) 1980 1980 Psittaboydaia Fain, 1985 Psittaboydaia psittacu­ Psittacula krameri Psittacidae Psittaciformes Africa Fain 1962c lae (Fain, 1962) Aprosmictus “ Psittacidae Psittaciformes Australia Domrow 1991 erythropterus Platycercus “ Psittacidae Psittaciformes Australia Domrow 1991 adscitus Psittaboydaia amazona Amazona European Zoo Fain and (Fain et Lukoschus, Psittacidae Psittaciformes amazonica (captive) Lukoschus, 1972 1972) Agapornis Australia “ Psittacidae Psittaciformes Domrow 1969 roseicollis (captive) Psittaboydaia (Trichoglossiella) Fain, 1985 European Zoo Fain 1970; Psittaboydaia tricho­ Trichoglossus Psittacidae Psittaciformes (captive); Domrow 1969, glossi (Fain, 1970) haematodus Australia 1991 Psitteuteles Domrow 1969, “ Psittacidae Psittaciformes Australia versicolor 1991 Neastrida Fain, 1962 Neastrida parrae (Fain, Actophilornis Jacanidae Gruiformes South Africa Fain 1956d 1956) africanus Neoboydaia Fain, 1958 Fain 1956a, Africa, Europe, Neoboydaia philomachi Philomachus Charadrii- 1956c; Domrow Charadriidae Australia (Fain, 1956) pugnax formes 1969; Zablu- (introduced) dovskaya 1998 Charadrii- Fain 1963b; “ Tringa glareola Scolopacidae Africa, Australia formes Domrow 1969 Charadrii- Fain and Aitken “ Actitis macularius Scolopacidae South Africa formes 1968 Charadrii- Domrow 1969, “ Limosa lapponica Scolopacidae Australia formes 1991 Neoboydaia philomachi Charadrii- Fain and Hyland eroliae Fain et Hyland, Calidris minutilla Scolopacidae South America formes 1970 1970 Charadrii- Fain and Hyland “ Actitis macularius Scolopacidae South America formes 1970 Charadrii- Fain and Hyland “ Arenaria interpres Scolopacidae North America formes 1975 Limnodromus Charadrii- Fain and Hyland “ Scolopacidae North America griseus formes 1975 Charadrii- Fain and Hyland “ Tringa solitaria Scolopacidae North America formes 1975 Tringa Charadrii- Clark 1964a; “ Scolopacidae North America melanoleuca formes Pence 1973a Charadrii- “ Tringa flavipes Scolopacidae North America Clark 1964a formes Charadrii- “ Calidris melanotos Scolopacidae North America Clark 1964a formes Pence 1973a; Gallinago Charadrii- North America “ Scolopacidae Zabludovskaya gallinago formes Asia 1998

i42 Appendix

Neoboydaia philomachi Limnodromus Charadrii- eroliae Fain et Hyland, Scolopacidae North America Pence 1973a scolopaceus formes 1970 Neoboydaia philomachi Thalasseus Charadrii- Fain and Hyland thalasseus Fain et Laridae North America maximus formes 1975 Hyland, 1975 Charadrii- Fain and Hyland “ Sterna hirundo Laridae North America formes 1975 Neoboydaia galachrisiae Charadrii- Glareola cinerea Glareolidae Africa Fain 1961b Fain, 1961 formes Charadrii- “ Glareola nuchalis Glareolidae Africa Fain 1964b formes Neoboydaia colymbi­ Podiceps Podicipedi- Podicipedidae North America Clark 1964a formi Clark, 1964 nigricollis formes Podilymbus Podicipedi- “ Podicipedidae North America Pence 1973a podiceps formes Tachybaptus Podicipedi- Domrow 1969, “ Podicipedidae Australia ruficollis formes 1991 Pterniboydaia Fain, 1985 Pterniboydaia pternistis Francolinus afer Phasianidae Galliformes Africa Fain 1955 (Fain, 1955) Francolinus “ Phasianidae Galliformes Africa Fain 1963 swainsonii “ Perdix perdix Phasianidae Galliformes Europe Fain 1962b Speleognathopsis Cooreman, 1954 Speleognathopsis galli North America, Cooreman 1954; Gallus gallus Phasianidae Galliformes Cooreman, 1954 Australia Domrow 1969 Africa, Australia Fain 1956a; “ Numida meleagris Numididae Galliformes (introduced) Domrow 1969 Speleognathopsis benoiti Vanellus Charadrii- Charadriidae Africa Fain 1955, 1956a Fain, 1955 crassirostris formes Charadrii- “ Vanellus senegallus Charadriidae Africa Fain 1956a formes Charadrii- Domrow 1969, “ Vanellus miles Charadriidae Australia formes 1991 Elseyornis Charadrii- Domrow 1969, “ Charadriidae Australia melanops formes 1991 Erythrogonys Charadrii- Domrow 1969, “ Charadriidae Australia cinctus formes 1991 Smutsornis Charadrii- “ Glareolidae South Africa Fain et al. 1966 africanus formes Speleognathopsis Charadrius Charadrii- European Zoo charadricola (Fain, Charadriidae Fain 1964a pecuarius formes (captive) 1964) Speleognathopsis Onychognathus onychognathi (Fain, Sturnidae Passeriformes Africa Fain 1964a walleri 1964) Speleognathopsis Domrow 1969, accipitris (Domrow, Accipiter fasciatus Accipitridae Accipitriformes Australia 1991 1969) Accipiter “ Accipitridae Accipitriformes Australia Domrow 1969 cirrocephalus “ Aquila audax Accipitridae Accipitriformes Australia Domrow 1969 Haliaeetus “ Accipitridae Accipitriformes Australia Domrow 1969 leucogaster

i43 M. Skoracki, S. Zabludovskaya and A.V. Bochkov

Speleognathopsis Haliastur accipitris (Domrow, Accipitridae Accipitriformes Australia Domrow 1969 sphenurus 1969) “ Milvus migrans Accipitridae Accipitriformes Australia Domrow 1969 “ Elanus axillaris Accipitridae Accipitriformes Australia Domrow 1969 Speleognathopsis wai Fain, Vercammen- Rhinoptilus Charadrii- Glareolidae South America Fain et al. 1966 Grandjean et Wagner, africanus formes 1966 Metaboydaia Fain, 1962 Metaboydaia poffei Scopus umbretta Scopidae Pelecaniformes Africa Fain 1955 (Fain, 1955) “ Coracias caudatus Coraciidae Coraciiformes Africa Fain 1955 Meropiboydaia Fain, 1985 Meropiboydaia merops Merops apiaster Meropidae Coraciiformes Africa Fain 1955, 1956a (Fain, 1956) Domrow 1969, “ Merops ornatus Meropidae Coraciiformes Australia 1991 Astrida Fain, 1955 Astrida caprimulgi Fain, Caprimulgi- Caprimulgus fossii Caprimulgidae Africa Fain 1956a 1955 formes “ Bubo africanus Strigidae Strigiformes Africa Fain 1956b Glaucidium “ Strigidae Strigiformes Africa Fain 1956b perlatum “ Otus senegalensis Strigidae Strigiformes Africa Fain 1956b “ Otus spilocephalus Strigidae Strigiformes Asia Fain 1963b Astrida (Cerylonyssus) Fain et Aitken, 1970 Astrida chlorocerylei Fain and Aitken Chloroceryle aenea Alcediniidae Coraciiformes South America Fain et Aitken, 1970 1970 Ralliboydaia Fain, 1962 Ralliboydaia lateralli Laterallus Rallidae Gruiformes South America Fain 1962a Fain, 1962 melanophaius Fulica americana Rallidae Gruiformes North America Pence 1973a Ralliboydaia coccyzae Coccyzus Cuculidae Cuculiformes North America Pence 1973a (Pence, 1973) americanus Ralliboydaia porphyrio­ Porphyrio Domrow 1965b, Rallidae Gruiformes Australia nis (Domrow, 1965) porphyrio 1969 Picinyssus Fain, 1969 Picinyssus buccanodon Buccanodon Lybiidae Piciformes Africa Fain 1969 Fain, 1969 duchaillui Phoenicopteriella Fain, 1970 Phoenicopteriella Phoenicopterus Phoenicopteri- Phoenicopteri- European Zoo Fain 1970 mirabilis Fain, 1970 ruber dae formes (captive) Phoenicopteriella Dendrocygna Anatidae Anseriformes Africa Fain 1955, 1956a womersleyi (Fain, 1955) viduata Sarkidiornis “ Anatidae Anseriformes North America Fain 1956a melanotos “ Aythya affinis Anatidae Anseriformes North America Clark 1958 “ Anas strepera Anatidae Anseriformes North America Clark 1958 Clark 1958; North America, “ Anas acuta Anatidae Anseriformes Zabludovskaya Asia 1998 Fain and Hyland “ Anas discors Anatidae Anseriformes North America 1975

i44 Appendix

Phoenicopteriella Zabludovskaya Anas crecca Anatidae Anseriformes Europe womersleyi (Fain, 1955) 1998 Zabludovskaya “ Anas querquedula Anatidae Anseriformes Europe 1998

i45