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67
INTRODUCTION
Prostigmatic mites (Acari: Acariformes: Pros-
tigmata) permanently associated with birds are po-
tentially very species-rich groups supposedly in-
cluding about 5000 species (Kethley and Johnston
1975; Moss 1979), but nearly 10 % (about 500
species) of these mites are recognized to date.
These mites occupy a great diversity of microhab-
itats on the host body: skin, quills, intracutaneous
layers, and respiratory tract, and are very interest-
ing in the global evolutionary sense as well as in
several parasitological aspects, including host-
parasite relationships.
This ecological group includes mites of ve
families belonging to three phylogenetically dis-
tant superfamilies: Cheyletidae, Harpirhynchidae,
and Syringophilidae of the superfamily Cheyle-
toidea, Cloacaridae (Cloacaroidea), and Ereyneti-
dae (Tydeoidea) (Mironov and Bochkov 2009).
The absence of taxonomic revisions and identi-
cation keys for most bird-associated prostigmatans
seriously impedes their phylogenetic and ecologi-
cal investigations.
Most prostigmatic inhabitants of bird are true
parasites feeding on live host tissues, but the ma-
jority of cheyletids living in feather quills are
predators feeding on other quill-inhabiting mites
and thus should be considered as commensals or
even mutualists.
Taking into consideration the high specicity
of permanently parasitizing prostigmatans to avian
hosts and their non-random distribution on host
taxa (Fain 1994a; Bochkov 2009; Skoracki 2011),
data about these mites could be used for the valida-
tion of host phylogeny and in deciphering of the
host biogeography. Such investigations were un-
dertaken on various groups of symbionts, includ-
ing acariform mites, and often showed a high level
of phylogenetic congruence between hosts and
symbionts (Klassen 1992). Although some investi-
gations conducted on astigmatan mites did not re-
veal phylogenetically congruent pattern, they pro-
vided valuable results for the host biogeography
(Bochkov and OConnor 2005). At the same time,
as a taxonomically poorly studied group, the pros-
tigmatans permanently associated with birds have
never been involved in this kind of investigations.
The data concerning systematics, phylogeny
and host-parasite relationships of acariform mites
permanently associated with mammals and birds
were recently revised (Proctor 2003; Bochkov
2009, 2010), but bird-associated prostigmatans
were not considered in these works.
In this paper we provide the family level re-
view of bird-associated prostigmatans, keys to all
genera, and check-lists of their species along with
veried host records. The data on phylogeny, biol-
ogy and host-parasite associations of these mites
are briey discussed.
MATERIAL AND METHODS
The materials used in this work were obtained
from acarological collections housed in various
museums. Data on bird-associated prostigmatan
families including numbers of recognized species
and genera, hosts, microhabitats, and distribution
are summarized in Table 1. Idiosomal and leg seta-
A REVIEW OF PROSTIGMATA (ACARIFORMES: TROMBIDIFORMES)
PERMANENTLY ASSOCIATED WITH BIRDS
M. Skoracki
1
, S. A. Zabludovskaya
2
and A. V. Bochkov
3, 4 *
1
Department of Animal Morphology, Faculty of Biology, Adam Mickiewicz University, Umultowska
89, 61–614 Poznan, Poland
2
Schmalghausen Institute of Zoology, National Academy of Ukraine, B. Khmelnitzky str. 15, Kiev,
Ukraine
3
Zoological Institute of the Russian Academy of Sciences, Universitetskaya Emb. 1, 199034 Saint Pe-
tersburg, Russia; e-mail: prostigmata@zin.ru
4
Museum 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 veried 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
Acarina 20 (2): 67–107
© Acarina 2012
68
M. Skoracki, S.A. Zabludovskaya, A.V. Bochkov
tion of these families is represented in Tables 2–5.
Most of the examined materials are deposited
in the following collections: Adam Mickiewicz
University (Poznan, Poland); Royal Belgian Insti-
tute of Natural Sciences (Brussels, Belgium);
Royal Museum for Central Africa (Tervuren, Bel-
gium); Zoological Institute, Russian Academy of
Sciences (Saint-Petersburg, Russia).
In the taxonomic part, the gnathosomal seta-
tion follows Grandjean (1946), the leg and idio-
somal setation follows Grandjean (1939, 1944).
The universal setal nomenclature of idiosoma pro-
posed by Grandjean (1939) was adopted for Pros-
tigmata by Kethley (1990). Schemes of external
mite morphology are provided in Figs. 1 and 2.
The bird systematics follows Clements et al.
(2011). Data about all described species of bird-
associated prostigmatans are summarized in the
Appendix (deposited at Acarina web-site: http://
insects.ummz.lsa.umich.edu/acarina/).
SYSTEMATICS AND PHYLOGENY
Suborder Trombidiformes Reuter, 1909
Infraorder Prostigmata Kramer, 1887
According to the system of acariform mites
proposed by Mironov and Bochkov (2008), the
suborder Trombidiformes is separated into four
infraorders, Sphaerolichida, Bimichaelida, Oeh-
serichestida, and Prostigmata. Only the infraorder
Prostigmata includes parasitic forms.
Parasites permanently associated with birds
belong to two phylogenetically distant parvorders,
Eupodina (Ereynetidae) and Eleutherengona
(Cheyletidae, Cloacaridae, Harpirhynchidae, and
Syringophilidae).
The only possible paleontological record of
these bird-associated mites is the nding of eggs
on a feather from the Lower Cretaceous period
(northeast Brazil) (Martill and Davis 1998). Au-
thors believed that these eggs were laid by feather
mites (Astigmata: Psoroptidia). These spherical
eggs are more similar to those of Cheyletoidea,
than to the strongly elongated eggs of feather mites
(Bochkov 2008). Proctor (2003), however, sup-
posed that this feather dropped to water and these
eggs were laid by Ostracoda.
Key to families of Prostigmata — permanent
parasites of birds
1. Subcapitulum of gnathosoma distinctly devel-
oped (Fig. 1). Palps present. At least some of idio-
somal and leg setae liform .............................. 2
— Subcapitulum of gnathosoma strongly reduced
or absent. Palps absent. Idiosomal and leg setae
represented by alveoli or eshy-like projections ...
Cloacaridae Camin, Moss, Oliver et Singer, 1967
(Pneumophagus Fain et Smiley, 1989) (Fig. 10)
2. Propodonotal setae si represented by unmodi-
A B
Fig. 1. Scheme of eleutherengone gnathosoma: A — in dorsal view; B — in ventral view (from Bochkov 2009, with minor
modications).
69
A review of Prostigmata permanently associated with birds
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 modications).
A B
C
D
70
M. Skoracki, S.A. Zabludovskaya, A.V. Bochkov
ed simple setae. Genital papillae absent. Tibia of
legs I with or without solenidion φ .................... 3
— Propodonotal setae si represented by bothridia
(Fig. 11A). Genital papillae present. Tibia of legs I
with complicate ereynetal organ (sensory complex,
Fig. 12A) ............ Ereynetidae (Speleognathinae)
3. Distal part of hypostome fused with subcapitu-
lum. Peritremes situated on subcapitulum, not lin-
ear. Palpal trochanter, femur and genu not fused.
Setae scx very small, covered by lateral extensions
of propodonotum. Tibia and genu I with 1 solen-
idion each .......................................................... 4
— Distal part of hypostome not fused with subca-
pitulum. Peritremes situated at base of subcapitu-
lum, linear. Palpal trochanter, femur and genu
fused to each other. Setae scx distinctly developed,
situated dorsally. Tibia and genu I without solen-
idion ......................................... Harpirhynchidae
Dubinin, 1957 (Figs. 8, 9)
4. Subcapitulum not deeply submerged in idioso-
ma, subcapitular apodeme absent. Claw of palpal
tibia present. Palpal tibia and tarsus separated. Se-
tae 4a present. Setae a’I absent .............................
.................... Cheyletidae Leach, 1815 (Figs. 3–5)
— Subcapitulum deeply submerged in idiosoma,
subcapitular apodeme distinctly developed. Claw
of palpal tibia absent. Palpal tibia and tarsus fused.
Setae 4a absent. Setae a’I present ...............................
.... Syringophilidae Lavoipierre, 1953 (Figs. 6, 7)
Parvorder Eleutherengona Oudemans, 1909
Superfamily Cheyletoidea Leach, 1815
The supefamily Cheyletoidea includes cur-
rently ve families, Cheyletidae, Syringophilidae,
Harpirhynchidae, Psorergatidae, and Demodici-
dae; mites of the three rst families are associated
with birds (Bochkov 2009). The phylogeny of
these mites was recently reconstructed by Bochk-
ov (2002, 2008). There are two main phylogenetic
lineages in this superfamily. Lineage I includes the
families Cheyletidae and Syringophilidae, where-
as lineage II is represented by three families,
Harpirhynchidae (Psorergatidae–Demodicidae).
The monophyly of lineage I is supported by the
following synapomorphies: the peritremes are sit-
uated on the rostral part of the stylophore, distinct-
ly segmented, and arch-like or M-shaped; the dis-
tal part of the hypostomal apex is fused with the
stylophore, and the hypostomal lips are present.
Lineage II could be characterized by the fol-
lowing synapomorphies: the presence of the
strongly sclerotized pharyngeal bulb, the absence
of adoral setae ao1 and ao2, the palpal tibia-tarsus
is located on the ventral side of the trochanter-fe-
mur-genu, the absence of the eupathidia on the
palpal tarsus, the absence of the separate hyster-
onotal shield, the completely fused genital and
anal openings in females, the absence of the
aggenital setae in both sexes, the absence of setae
ft, vsI, solenidion φI, σI, coxal setae 1b (reversed
in Ophioptinae), 3b, and 4c (reversed in Harpypal-
pinae).
Family Cheyletidae Leach, 1815
Figs. 3–5
Type genus: Cheyletus Latreille, 1796
Diagnosis. Gnathosoma moderately or dis-
tinctly developed (1/5–1/3 of idisosomal length).
Subcapitulum bearing setae elc. p, n, ao1 and ao2.
Peritremes situated on rostral part of stylophore,
distinctly segmented, generally M-shaped. Distal
part of hypostome fused with stylophore. Hypos-
tomal lips present. Palps linear, consisting of 5
segments, with various projections in some para-
sitic forms. Palpal tibia with distinct claw. Palpal
tarsus strongly reduced. Palpal setation: femur
with setae d, v’, v”, genu — d, l”, tibia — d, lT, l’,
and tarsus with eupathidia acm, sul, ul’, ul”, and
solenidion ω. Eupathidia acm and sul” comb-like
in most predaceous mites and smooth in most par-
asites. Idiosoma generally rhomb-like or ovate in
outlline. Bases of trochanters inserted on ventral
side of idiosoma. In Cheyletiellini, idiosoma bear-
ing 1 pair of distinctly developed lateral scapular
lobes. Eyes absent; or 1 pair of lens-like eyes pres-
ent on propodonotum in some free-living preda-
tors. Propodonotal shield present, distinctly devel-
oped. Hysteronotal and pygidial shields present or
absent. Ventral side of idiosoma generally devoid
of shields but in males of some predatory forms
weakly sclerotized pseudosternal shield present.
Opisthosoma moderately developed. In females,
anal and genital-ovipore openings situated ven-
trally, close to posterior end of body or terminally,
adnate and covered by pair of folds. In males, ae-
deagus tube-like or comma-like and pointed api-
cally, genital and anal openings fused (or anal
opening absent). In males of free-living forms,
genital opening situated terminally or ventro ter-
minally. In parasitic males, genital opening dor-
sally displaced, and in some bird parasites, it lo-
cated in anterior part of idiosoma. Distance
between coxal elds II and III distinctly shorter
than idiosomal width (excluding tribe Bakini). In
most taxa, 3 pairs of cupules present: im, ip, and
ih. In some parasites, these cupules absent. Idio-
71
A review of Prostigmata permanently associated with birds
somal setation (maximum set): scx, vi, ve, si, se,
c1, c2, d1, d2, e1, e2, f1, f2, h1, h2, h3, ps1, ps2,
ps3, ag1, ag2, ag3, g1, g2. Setae scx covered dor-
sally by extending lateral margins of propodono-
tum. In males, 2 pairs of aggenital setae, 2 pairs of
genital setae, and 3 pairs of pseudoanal setae pres-
ent. In some taxa idiosomal chaetome neotrichous
and many neotrichous dorsal setae strongly modi-
ed. In most taxa, legs slender, consisting of 5 ar-
ticulated segments, and their tarsi having weakly
developed pretarsi bearing 2 smooth lateral claws
and empodium with tenent hairs. Pretarsi separat-
ed from respective tarsi by dorsal knob. Leg length
usually 60–70% of idiosomal length. In Metachey-
letia, legs IV lost or primordial. In free-living
forms, leg coxae distinctly bordered, and in some
cases well sclerotized. In many parasitic mites, leg
coxae represented exclusively by coxal apodemes.
Projections of leg segments present only in some
parasitic species. Maximum set of leg setae (paired
setae are in parentheses): tarsi I–IV — I, ft, (tc),
a”, (p), (u), vs, ω1; II, (tc), (p), (u), vs, ω1; III,
(tc), (p), (u), vs, ω1 (only in males); IV, (tc), (p),
(u), vs, ω1 (only in males); tibiae I–IV — I, d, (l),
(v), φ; II, d, l”, (v), φ; III, d, l”, (v) + φ (only in
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;
trochanters I–IV — I, II, IV, v; III, l, v; coxae I–IV
— I, 1a, 1b, 1c; II, 2c; III, 3a, 3b, 3c; IV, 4a, 4b,
4c. Solenidion ω1 II situated ventrally. Larva, pro-
to- and tritonymphs present; deutonymph absent.
Male moulting from protonymph.
Detailed historical reviews of the cheyletid
systematics were provided by Volgin (1969) and
Bochkov (2009). Therefore we discuss it here very
briey. The excellent taxonomic revision of the
family Cheyletidae prepared by Volgin (1969)
could be considered as a base of the modern chey-
letid systematics. According to his system, the
family consisted of ten tribes and the two subfam-
ilies Cheyletinae and Cheyletiellinae. The revision
of the family Cheyletidae prepared by Summers
and Price (1970) appeared almost simultaneously
with the Volgin’s monograph but was not so com-
prehensive. The latter work did not introduce any
principal improvements to the cheyletid system
and the tribes and subfamilies established by Vol-
gin were unceremoniously ignored by these au-
thors.
Fig. 3. Metacheletoides numidae (Cheyletidae: Cheletosoma-
tini), male in dorsal view (from Bochkov and Skoracki 2012
with minor modications).
Fig. 4. Ornithocheyletia chirovi (Cheyletidae: Ornithochey-
letiini), female in dorsal view.
72
M. Skoracki, S.A. Zabludovskaya, A.V. Bochkov
Fain et al. (1997) and Gerson et al. (1999) re-(1999) re-
vised the family at the generic level and provided
a list of all described species.
The most important synapomorphies of Chey-
letidae are the ventral situation of solenidion on
tarsi II, male moulting from protonymph, and the
absence of setae a’ of tarsus I (Bochkov 2008).
The attempt of the cladistic analysis of the
family Cheyletidae was undertaken by Bochkov
and Fain (2001). According to the result of this
analysis with some corrections (Bochkov 2004),
the family Cheyletidae includes 14 tribes most of
which were established by Volgin (1969): Ac-
aropsellini, Bakini, Cheletogenini, Cheletosoma-
tini, Chelonotini, Cheyletiini, Cheyletiellini,
“Cheyletini”, Cheletomorphini, Metacheyletiini,
Niheliini, Ornithocheyletiini, Teinocheylini, and a
unnamed tribe represented by the genus Cau-
dacheles. The relationships between these tribes
remain unresolved. The bird parasites and nidi-
colous mites belong to the tribes Cheletosomatini,
Metacheyletiini, and Ornithocheyletiini.
Key to bird-associated genera of Cheyletidae
1. Legs IV present, normally developed. Idiosoma
rhomb-like in outline ......................................... 2
— Legs IV absent or primordial. Idiosoma ovate
in outline .................. (Metacheyletiini Fain, 1980)
Metacheyletia Fain, 1972
2. Eupathidium sul” comb-like or serrate. Palpal
claw widely opened, slightly curved laterally, with
basal angle(s). At least some idiosomal setae of
females ultralong (excluding normally always
long c2 and h2). Male genital opening situated ter-
minally. Tarsi of legs I much longer than wide
(Cheletosomatini Volgin, 1969) ......................... 3
— Eupathidium sul” smooth. Palpal claw strongly
curved ventrally and without basal angles. Idiosom-
al setae of females moderately long (excluding nor-
mally always long c2 and h2). Male genital opening
situated dorsally. Tarsi of legs I slightly longer than
wide (Ornithocheyletiini Volgin, 1969) ............... 8
3. Eupathidium sul” smooth or slightly serrate.
Setae c3 present ................................................. 5
A B
Fig. 5. Metacheyletia ngaii (Cheyletidae: Metacheyletiini), female: A — in dorsal view, B — in ventral view (from Bochkov
and Skoracki 2011 with minor modications).
73
A review of Prostigmata permanently associated with birds
— Eupathidium sul” comb-like. Setae c3 absent .
............................................................................ 4
4. Coxal elds III and IV contiguous. Setae c1
situated on propodonotal shield .............................
............................... Cheletopsis Oudemans, 1904
— Coxal elds III and IV distinctly separated. Se-
tae c1 situated off propodonotal shield ..................
.................................. Eucheletopsis Volgin, 1969
5. Setae c4 absent. Female hysteronotal shield ab-
sent or situated on posterior part of opisthosoma.
Setae e1 and f1 absent ....................................... 6
— Setae c4 present. Female hysteronotal shield
present, situated between levels of setal bases d2
and e2. Setae e1 and f1 present .............................
........... Picocheyletus Bochkov et OConnor, 2003
6. Eupathidium sul” serrate. Palpal claw with 1
basal angle. Tarsal claws on all legs equal or sub-
equal in size ....................................................... 7
— Eupathidium sul” smooth. Palpal claw with
several basal angles. Tarsal claws on legs I much
smaller than claws of tarsi III–IV ..........................
.................................. Metacheletoides Fain, 1972
7. In female, small hysteronotal shield present and
setae h1 short, much shorter than c2 .....................
.............................. Cheletosoma Oudemans, 1905
— In female, hysteronotal shield absent and setae
h1 long, subequal to c2 ..........................................
............................... Cheletoides Oudemans, 1904
8. Genua III and IV with 1 and without setae, re-
spectively ......................................................... 10
— Genua III and IV with 2 and with 1–2 setae,
respectively ........................................................ 9
9. Hysteronotal shield present. Trochanters III and
IV without setae. Genu IV with 1 seta ..................
................................. Ornithocheyla Volgin, 1964
— Hysteronotal shield absent. Trochanters III and
IV with 2 and 1 setae, respectively. Genu IV with
2 setae ....................... Bakericheyla Volgin, 1966
10. Palpal femora without ventral processes.
Propodonotal shield very small. Setae c3 present. In
females, 2 pairs of aggenital setae present. Solen-
idion φI present. Trochanter IV, femora III and IV
with 1 seta each ........ Neocheyletiella Baker, 1949
— Palpal femora with ventral processes.
Propodonotal shield distinctly developed, cover-
ing part of hysteronotum. Setae c3 absent. In fe-
males, aggenital setae absent. Solenidion φI ab-
sent. Trochanter IV, femora III and IV without
setae ............................... Apodicheles Fain, 1979
Family Syringophilidae Lavoipierre, 1953
Figs. 6–7
Type genus: Syringophilus Heller, 1880
Diagnosis. Gnathosoma moderately devel-Gnathosoma moderately devel-
oped. Subcapitulum deeply inserted into idiosoma,
bearing setae elcp, n, ao1 and ao2; subcapitular
apodeme distinctly developed. Peritremes situated
on rostral part of stylophore, distinctly segmented,
generally M-shaped. Hypostomal lobes fused with
stylophore. Hypostomal lips present. In many taxa,
female hypostome bearing 1 pair of hyaline protu-
berances. Palps linear, consisting of 4 segments
(tibia and tarsus fused). Palpal tibiotarsus without
claw. Palpal setation: femur with setae d, v’,v”,
genu — d, l”, tibiotarsus — d, lT, l’, eupathidia
acm, sul, ul’, ul”, and solenidion ω. Idiosoma
strongly elongated. Bases of trochanters inserted on
ventral side of idiosoma. Eyes absent. Propodono-
tal shield present. Hysteronotal and pygidial shields
present or absent. Ventral side of idiosoma gener-
ally devoid of shields. Opisthosoma distinctly de-
veloped. Female anal and genital-ovipore openings
situated ventrally close to posterior end of body or
terminally, adnate, and covered by pair of folds. In
males, aedeagus tube-like and pointed apically,
genital and anal openings fused (or anal opening
absent) and situated dorsally. Distance between
coxal elds II and III subequal or about twice as
long as idiosomal width. Idiosomal cupules absent.
Idiosomal setation (maximum set): scx, vi, ve, si, se,
c1, c2, d1, d2, e2, f1, f2, h1, h2, ps1, ps2, ag1, ag2,
ag3, g1, g2. Setae scx covered dorsally by extend-
ing lateral margins of propodonotum. In males, 2–3
pairs of aggenital setae, 2 pairs of genital setae, and
2 pairs of pseudoanal setae present. In some taxa,
aggenital setae neotrichous. Legs slender, consist-
ing of 5 articulated segments; their tarsi having
weakly developed pretarsi bearing 2 smooth lateral
claws and empodium with tenet hairs. Maximum
set of leg setae (paired setae are in parentheses):
tarsi I–IV — I, ft, (tc), (a), (p), (u), vs, ω1; II, (tc),
(p), (u), vs, ω1; III, (tc), (p), (u), IV, (tc), (p), (u);
tibiae I–IV — I, d, (l), v, φ; II, d, (l), v, φ; III, d, (l);
IV, d, (l); genua I–IV — I, d, l’, σ; II, d, l’; III and IV
l’, femora I–IV — I and II, d, v, III and IV d; tro-
chanters I–IV, v; III; coxae I–IV — I, 1a, 1b, 1c; II,
2c; III, 3a, 3b, 3c; IV, 4b, 4c. Solenidion ω1II situ-
ated dorsally. Larva, proto- and tritonymphs pres-
ent; deutonymph absent. Male moulting from
tritonymph.
The detailed discussion of syringophilid sys-
tematics is provided in the monograph by Skoracki
(2011). Therefore, only the principal works are
considered below. The most important review for
syringophilid systematics was prepared by Keth-
ley (1970). In the work following this revision,
74
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.
75
A review of Prostigmata permanently associated with birds
Fig. 7. Picobia heeri (Syringophilidae: Picobiinae), female: A — in dorsal view; B — in ventral view.
Johnston and Kethley (1973) proposed the origi-
nal variant of syringophilid system based on the
result of their phenetic analysis. This analysis con-
rmed all syringophilid genera established by
Kethley (1970) and allowed the division of this
family into two unequal subfamilies, Syringophili-
nae and Picobiinae. The third syringophilid sub-
family — Lobatinae was created by Casto (1977)
for the monotypic genus Cuculiphilus. This genus
was subsequently included in the subfamily Pico-
biinae (Fain et al. 2000). From the beginning of
XXI century, the biodiversity of syringophilids
has been studied mainly by A.V. Bochkov and M.
Skoracki (with collaborators). To date, this family
includes 278 species of 53 genera versus 24 spe-
cies of 15 genera in Kethley´s review (1970). The
A B
76
M. Skoracki, S.A. Zabludovskaya, A.V. Bochkov
modern comprehensive revision of Palaearctic Sy-
ringophilidae was recently prepared by Skoracki
(2011).
The most distinctive synapomorphies of Sy-
ringophilidae are the subcapitulum deeply inserted
into the idiosoma; four segmented linear palps,
widely separated coxae I–II and III–IV, and the
absence of setae 4a (Bochkov 2008).
Any works on the phylogeny of this family
are absent.
Key to genera of Syringophilidae
1. Tibiotarsus of palps truncate on distal margin.
Prorals setae p’ and p” of legs I–IV with two min-
ute tines, rod-like. Physogastric forms of females
present. Juvenile stages with chaetotaxy reduced
to small spinose structures. Body feathers as main
habitat. (Picobiinae Johnston et Kethley, 1973) ....
.......................................................................... 48
— Tibiotarsus of palps rounded on distal margin.
Prorals setae p ’ and p” of legs I–IV multiserrate,
fan-like. Physogastric forms of females absent.
Setae of body and legs in juveniles well devel-
oped. Flight feathers as main habitat. (Syringo-
philinae Lavoipierre, 1953) ............................... 2
2. Setae vi present ............................................ 12
— Setae vi absent .............................................. 3
3. Setae dFII present .......................................... 9
— Setae dFII absent .......................................... 4
4. Setae dFIII and dFIV absent .......................... 5
— Setae dFIII and dFIV present and replaced ven-
trally .................................... Terratosyringophilus
Bochkov et Perez 2002
5. Setae ve situated anterior to level of setae si.
Pocket-like structures absent. Stylophore rounded
posteriorly .......................................................... 6
— Setae ve and si situated at same transverse lev-
el. Pocket-like structures in anterior part of pro-
podonotum present. Stylophore constricted poste-
riorly .............................................. Psittaciphilus
Fain, Bochkov et Mironov, 2000
6. Hypostomal apex ornamented by 2 pairs of me-
dian nger-like protuberances. Apodemes I and II
parallel and fused to each other ......................... 7
— Hypostomal apex smooth. Apodemes I diver-
gent, not fused to apodemes II .......... Meitingsunes
Glowska et Skoracki, 2010
7. Setal pattern of propodonotal region arranged
2–1–2 ................................................................. 8
— Setal pattern of propodonotal region arranged
2–3 ................. Neoperisterophila Skoracki, 2005
8. Legs I and II subequal in size ..... Peristerophila
Kethley, 1970
— Legs I 1.5 times longer than legs II .......................
........ Castosyringophilus Bochkov et Perez, 2002
9. Setae dGII absent. Apodemes I divergent .........
................ Galliphilopsis Skoracki et Sikora 2004
— Setae dGII present. Apodemes I parallel .... 10
10. Setae l’RI and l’RII present ....................... 11
— Setae l’RI and l’RII absent ............. Aulonastus
Kethley, 1970
11. Two pairs of pseudanal setae (ps1 and ps2)
present ................ Neoaulonastus Skoracki, 1999
— One pair of pseudanal setae present (ps2 ab-
sent) ............... Krantziaulonastus Skoracki, 2011
12. Legs with full complement of setae ........... 29
— Some of leg setae absent ............................. 13
13. Setae dGII present ...................................... 18
— Setae dGII absent ........................................ 14
14. Setae vsI present ......................................... 15
— Setae vsI absent ...................... Philoxanthornea
Kethley, 1970
15. Setae l’GIV absent ..................................... 17
— Setae l’GIV present ..................................... 16
16. Two pairs of pseudanal setae present ..................
............................ Syringophiloidus Kethley, 1970
— One pair of pseudanal setae present (ps2 ab-
sent) .......... Betasyringophiloidus Skoracki, 2011
17. Apodemes I parallel fused to apodemes II ........
.. Apodisyringophilus Skoracki et OConnor, 2010
— Apodemes I distinctly divergent, not fused to
apodemes II ..... Apodisyringiana Skoracki, 2005
18. Setae dFII present ...................................... 20
— Setae dFII absent ........................................ 19
19. Setae dTIII and dTIV present. Propodonotal
shield divided longitudinally ........ Ascetomylla
Kethley, 1970
— Setae dTIII and dTIV absent. Propodonotal
shield entire .................... Fritschisyringophilus
Bochkov, Fain et Skoracki, 2004
20. Setae dTIII present ..................................... 22
— Setae dTIII absent ....................................... 21
21. Setae vsII present. Apodemes I parallel .............
....... Neoaulobia Fain, Bochkov et Mironov, 2000
— Setae vsII absent. Apodemes I divergent .......
..................... Cuculisyringophilus Skoracki, 2008
22. Setae vsI present ........................................ 24
— Setae vsI absent ........................................... 23
23. Apodemes I parallel. Stylophore constricted
posteriorly. Claws strongly recurved ..................
................. Bochkovia Skoracki et OConnor, 2010
— Apodemes I divergent. Stylophore rounded
posteriorly. Claws typically opened ..................
............................. Paraniglarobia Skoracki, 2011
24. Setae vsII absent, l’GIV present ................. 25
— Setae vsII present, l’GIV absent ........................
..... Neosyringophilopsis Skoracki et Sikora, 2005
77
A review of Prostigmata permanently associated with birds
25. Lateral hypostomal teeth absent ................ 26
— Lateral hypostomal teeth present ......................
..................................... Stibarokris Kethley, 1970
26. Two pairs of genital setae present (g1, g2).
Apodemes I not fused to apodemes II ............. 27
— One pair of genital setae present (g2 absent).
Apodemes I fused to apodemes II ........................
................ Procellariisyringophilus Kethley, 1970
27. Apodemes I parallel. Stylophore without large
tip on posterior margin .................................... 28
— Apodemes I divergent. Stylophore with large
tip on posterior margin ............ Ciconichenophilus
Skoracki et OConnor, 2010
28. Claws small, with basal angle, typically opened
..................................... Niglarobia Kethley, 1970
— Claws large, without basal angle, strongly re-
curved ........................................... Phalarophilus
Skoracki, Bochkov et OConnor, 2011
29. Two or 3 pairs of aggenital setae present ... 31
— Aggenital series with 4–9 pairs of setae ..... 30
30. Coxal elds I and II similar in size and shape.
Supernumerary setae vsIII absent. Legs I thicker
than II–IV ..................... Torotrogla Kethley, 1970
— Coxal elds I and II disimilar in size and shape.
Supernumerary setae vsIII present. Legs I–IV sub-
equal ......................... Trypetoptila Kethley, 1970
31. Three pairs of aggenital setae (ag1–3) present
.......................................................................... 34
— Two pairs of aggenital setae present (setae ag2
absent) .............................................................. 32
32. Apodemes I not fused to apodemes II. Claws
typically opened .............................................. 33
— Apodemes I fused to apodemes II. Claws
strongly recurved .... Kethleyana Kivganov, 1995
33. Hysteronotal shield not fused to pygidial
shield. Coxal elds of legs I and II are similar in
size and shape, apodemes of legs III and IV well
developed ................................ Picisyringophilus
Skoracki et OConnor, 2010
— Hysteronotal shield fused to pygidial shield.
Coxal elds of legs I and II are dissimilar in size
and shape. Apodemes of legs III and IV absent
.................. Mironovia Chirov et Kravtsova, 1995
34. Lateral hypostomal teeth absent ................ 35
— Lateral hypostomal teeth present ........................
.................................. Colinophilus Kethley, 1970
35. Peritremes M-shaped ................................. 36
— Peritremes U-shaped .................. Syringophilus
Heller, 1880
36. Coxal elds I and II dissimilar in size and
shape ................................................................ 39
— Coxal elds I and II similar in size and shape ..
.......................................................................... 37
37. Setae f2 long (longer than c1). Setae si and se
situated at same level ....................................... 38
— Setae f2 short (2–3 times shorter than c1). Setae
si situated anterior to level of setae se ...................
.................................... Chenophila Kethley, 1970
38. Claws broadly open ................... Selenonycha
Kethley, 1970
— Claws strongly recurved ............ Creagonycha
Kethley, 1970
39. Apodemes I divergent ................................ 40
— Apodemes I parallel .... Aulobia Kethley, 1970
40. Apodemes I not fused to apodemes II ........ 42
— Apodemes I fused to apodemes II ............... 41
41. Propodonotal shield divided longitudinally .....
............. Crotophagisyringophilus Skoracki, 2008
— Propodonotal shield entire .... Syringophilopsis
Kethley, 1970
42. Stylophore constricted posteriorly ............ 44
— Stylophore rounded posteriorly .................. 43
43. Dorsal setae of idiosoma ornamented ...............
...................... Blaszakia Skoracki et Sikora, 2008
— Dorsal setae of idiosoma smooth ......................
...................... Bubophilus Philips et Norton, 1978
44. Apodemes I strongly divergent. Setae se lo-
cated anterior to the level of setae c1 ............... 47
— Apodemes I slightly divergent. Setae se located
posterior to level of setae c1 ............................ 45
45. Setae f2 situated close to the bases of setae f1 .
.......................................................................... 46
— Setae f2 situated distinctly anterior to the level
of setae f1 ............................ Charadriphilus Bo-
chkov et Chystiakov, 2001
46. Each lateral branch of peritremes reduced to 3
chambers. Setae se and c1 situated at same trans-
verse level. In males 2 pairs of aggenital setae pres-
ent and setae se situated distinctly anterior to level of
setae c1 ... Pteroclidisyringophilus Skoracki, 2011
— Each lateral branch with 7–12 chambers. Setae
se situated distinctly posterior to level of setae c1.
In males 3 pairs of aggenital setae present and se-
tae se and c1 situated at same transverse level ......
................................ Kalamotrypetes Casto, 1980
47. Setae si and se situated at same level. All ter-
minal setae long ..................... Megasyringophilus
Fain, Bochkov et Mironov, 2000
— Setae si set distinctly anterior to level of setae
se, setae f1 and f2 long, setae h1 and h2 short .........
............ Tinamiphilopsis Skoracki et Sikora, 2004
48. End of hysterosoma rounded, without opistho-
somal lobes. Movable cheliceral digit edentate .... 49
— End of hysterosoma with pair of opisthosomal
lobes. Movable cheliceral digit dentate, each with
3 teeth ....................... Calamincola (Casto, 1978)
78
M. Skoracki, S.A. Zabludovskaya, A.V. Bochkov
49. Bases of setae vi set anterior to level of setae
ve. Apodemes I divergent, without thorn-like scle-
rotization in middle part. Genital lobes absent.
Physogastric female campanulliform or bulb-
shaped outline .................................................. 50
— Bases of setae vi and ve set at same transverse
level. Apodemes I parallel, thorn-like sclerotiza-
tion in middle part. Genital lobes present or ab-
sent. Physogastric female worm-shaped in outline
............................................. Picobia Haller, 1878
50. Solenidia phi on tibia of leg I absent. Hypos-
tomal apex truncate. Peritremes with clearly or
weakly visible chambers in short lateral branches.
Propodonotal shield entire and shirt-like. Pygidial
shield present. Bases of setae 1a–1a not coalesced.
Physogastric females bulb-shaped outline ....... 51
— Solenidia phi on tibia of leg I present. Hypos-
tomal apex rounded. Peritremes with clearly visi-
ble chambers in lateral branches. Propodonotal
shield divided longitudinally into 2 lateral shields,
bearing bases of setae vi, ve, si and se. Pygidial
shield absent. Bases of setae 1a–1a coalesced.
Physogastric females campanulliform outline
.......... Columbiphilus Kivganov et Sharafat, 1995
51. One pair of genital setae and 2 pairs of pseuda-
nal setae present ...... Rafapicobia Skoracki, 2011
— Genital setae absent, 2 pairs of pseudanal setae
present ...................... Neopicobia Skoracki, 2011
Family Harpirhynchidae Dubinin, 1957
Figs. 8–9
Type genus: Harpirhynchus Latreille, 1796
Diagnosis. Gnathosoma moderately devel-Gnathosoma moderately devel-
oped. Subcapitulum bearing setae elc. p, n, and
m. Peritremes situated at base of stylophore, seg-
mented only in distal parts, straight. Hypostomal
lobes not fused with stylophore. Hypostomal lips
absent. Palps linear, consisting of 3 articulated
segments — trochanter-femur-genu, tibia and ru-
dimentary tarsus. Palpal tibia replaced on ventral
surface of trochanter-femur-genu and bearing
distinct paraxial claw-like seta. Palpal tarsus
strongly reduced. Palpal setation: femur with se-
tae d, v (only with seta d in Harpypalpinae), genu
— d, l”, tibia — d, lT, and tarsus with 1 seta. Id-
iosoma attened dorso-ventrally and rounded
outline or sucker-like. Bases of trochanters in-
serted ventro-laterally. Eyes absent. Propodono-
tal shield present, distinctly developed, fused
with remnants of hysteronotal shield. Ventral side
of idiosoma devoid of shields in Harpirhynchi-
nae and bearing genital-anal shield in Harpypal-
pinae. Opisthosoma weakly developed. Female
anal and genital-ovipore openings fused and situ-
ated ventrally close to posterior end of body or
terminally, covered by pair of folds. In males, ae-
deagus tube-like or comma-like and pointed api-
cally; genital and anal openings fused (or anal
opening absent), dorsally displaced. Distance be-
tween coxal elds II and III distinctly shorter
than idiosomal width. Idiosomal cupules absent.
Idiosomal setation (maximum set observed in
Harpypalpinae): scx, vi, ve, si, se, c2, d1, d2, e2,
f1, f2, h1, ps1–3, g1. Setae scx situated dorsally,
distinctly developed. In males, setae c1 and 2
pairs of pseudoanal setae present. In females of
Harpirhynchinae, idiosoma bearing only setae
scx, vi, ve, si, se, c2, h1, ps1; in harpirhynchine
males, 2–3 pairs of pseudoanal setae present.
Legs shortened; legs I and II consisting of 5 ar-
ticulated segments, their tarsi having weakly de-
veloped pretarsi bearing 2 smooth lateral claws
and empodium with tenet hairs. In Harpypalpi-
nae, legs III and IV normally developed, similar
with anterior legs. In Harpirhynchinae legs III
and IV strongly reduced, 1–3 segmented. Maxi-
mum set of leg setae in Har pypalpinae (paired
setae are in parentheses): tarsi I–IV — I, II, (tc),
p”, (a), (u), ω1; III, IV (tc), (a), (u); tibiae I–IV
— I, II, d, (l), (v); III, IV, d, (v); genua I–IV — I,
II, d, (l),(v); III, IV, without setae; femora I–IV
— I, II, d, v; III, IV, v; trochanters I–IV — v;
coxae I–IV — I, 1a, 1c; II, 2c; III, 3a, 3c; IV, 4c.
Solenidion ω1 II situated dorsally. Larva, proto-
and tritonumphs present; deutonymph absent.
Male moulting from protonymph.
This family was revised by Fain (1976, 1994b,
1995) who described most of its taxa. Before this
revision only a few particular works concerning
these mites were published. Most important of
them was a paper by Fritsch (1954) concerning
harphirhynchids from Germany.
Within the content of harpirhynchids, Fain
(1972) established a new subfamily Harpypalpi-
nae for a single genus Harpypalpus with normally
developed legs III and IV. Later, Lombert and
Moss (1983) described Harpypalpoides, the sec-
ond genus in this subfamily.
Presently, A.V. Bochkov along with collabo-
rators works on the systematics of these mites.
The monophyly of Harpirhynchidae is sup-
ported by three unique synapomorphies: the
strongly reduced, membranous palpal tarsus, seta
l”G grouped together with dG in the apical part of
the palpal throchanter-femur-genu, and modied
setae l’T of the palpal tibia (Bochkov 2008).
79
A review of Prostigmata permanently associated with birds
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-
tral side of the idiosoma, the anal opening is ab-
sent, immatures are apode.
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 modications).
Fig. 9. Harpypalpoides sp. (Harpirhynchidae: Harpypalpi-
nae), female in dorsal view.
80
M. Skoracki, S.A. Zabludovskaya, A.V. Bochkov
Key to genera of Harpirhynchidae
(based on Fain [1995])
1. Most hysteronotal setae absent. Legs III and IV
shortened, 1–2 segmented. Pretarsi III and IV ab-
sent … (Harpirhynchinae Dubinin, 1957) ......... 3
— Most hysteronotal setae present. Legs III and
IV normally developed, consisting of 5 articulate
segments. Pretarsi III and IV present, each bearing
empodium and 2 claws. (Harpypalpinae Fain,
1972) .................................................................. 2
2. Setae se situated at level of seta si bases. Setae
d2 present ............... Harpypalpus Dubinin, 1957
— Setae se situated distinctly posterior to level of
seta si bases. Setae d2 absent. ...... Harpypalpoides
Lombert et Moss, 1983
3. Setae dG and dF not liform or rodlike, but rigid
and shorter than palp tibia, with generally strong
pectinations. Setae l”G similar to other palpalae or
rodlike and shortly pectinate ............................... 4
— Setae dG, l”G, and dF of palps liform, shortly
pectinate, longer than palp tibia. ........................
............................... Trichorhynchiella Fain, 1954
4. Female: Idiosoma rounded, elongate or wider
than long. Legs I–II reduced in size, with maxi-
mum of 4 articulate segments ............................ 6
— Female: Idiosoma circular or subcircular. Legs
I and II well developed, with 5 articulated seg-
ments ................................................................. 5
5. In females, setae dG, l”G, and dF of palps sim-
ilar in shape but l”G slightly shorter and narrower
than dG and dF. In males, genital opening situated
in anterior quarter of dorsum ......... Harpirhynchus
Megnin, 1877
— In females, setae dG, l”G, and dF of palps dif-
ferent in size and shape. In males, genital opening
situated in posterior third of dorsum .......................
............................... Harpyrhynchoides Fain, 1972
6. Idiosoma variable in shape, but not rounded or
circular ................................................................ 8
— Body rounded or circular .............................. 7
7. Dorsal shield small. Legs I and II slightly re-
duced in size, with only 4 free segments (genu and
femur fused) and inserted marginally; tarsi I–II
with claws and empodium. Legs III normally
shaped. Legs IV absent. Propodonotal setae
smooth. Setae vi, si, and se short and smooth .......
.............................. Perharpyrhynchus Fain, 1972
— Dorsal shield very large. Legs I and II strongly
reduced in size, lacking claws and empodium; legs
I slightly ventral, legs II marginal. Legs III and IV
very small, not segmented. Setae vi, si, and se pec-
tinate ................... Ralliharpirhynchus Fain, 1972
8. Body not sacciform but either dome-shaped or
wider than long and narrowed posteriorly ....... 10
— Body elongate, sacciform ............................. 9
9. Legs I–II moderately reduced. Legs I with 2–4
articulated segments. Legs II with 2–3 articulated
segments. ............... Neharpyrhynchus Fain, 1972
— Legs I–II strongly reduced. Legs I with only 1
very short segment. Legs II similar to legs I or ab-
sent ...................... Metharpyrhynchus Fain, 1972
10. Female: Idiosoma wider than long, with 2
broad lateral lobes. Dorsal shield poorly devel-
oped. Palpal seta l”G absent ............................ 11
— Female: Body dome-shaped, with 2 postero-
lateral lobes. Dorsal shield distinctly developed.
Palpal seta l”G present ............. Anharpyrhynchus
Fain, 1972
11. Female: Setae ve and si modied into very
short spinelets. Lateral lobes situated in anterior
third of idiosoma. Legs I and II ventral; bearing
claws and empodium ......... Harpyrhynchiella
Fain, 1972
— Setae ve and si normal and pectinate. Lateral
lobes situated in middle part of idiosoma. Legs I
and II marginal, without claws and empodium ......
............................. Cypsharpirhynchus Fain, 1995
Superfamily Cloacaroidea Camin, Moss,
Oliver et Singer, 1967
The superfamily Cloacaroidea has been re-
cently established for two families, Cloacaridae
and Epimyodicidae (Bochkov and OConnor 2008).
This superfamily is characterized by the following
synapomorphies: the gnathosoma is shifted ven-
trally, the chelicerae are devoid of digits and setae,
the subcapitular setation is completely lost, the
coxal apodemes are longer than half of the body
width, in adults, coxal apodemes I are fused into a
sternum, and females, genital and anal openings
are completely fused to each other.
The relationships of Cloacaridae and
Epimyodicidae with other prostigmatic mites are
not clear and, therefore, this superfamily is con-
sidered as incertae sedis within Eleutherengona.
These mites are strongly modied for endopara-
sitic mode of life and have lost many morphologi-
cal structures present in other Prostigmata. The
distinctly separated chelicerae of Cloacaroidea
obviously prevent their inclusion in the Cheyle-
toidea, because all cheyletoid families have com-
pletely fused chelicerae with stylet-like movable
cheliceral digits. Such characters, as the fusion of
the anal and genital openings and the dorsal posi-
tion of an aedeagus in males indicate the afnity
81
A review of Prostigmata permanently associated with birds
of these two families with the parvorder Eleutheren-
gona, but are not helpful in clarifying their rela-
tionships to other lineages within the parvorder.
Family Cloacaridae Camin, Moss, Oliver et
Singer, 1967
Fig. 10
Type genus: Cloacarus Camin, Moss, Oliver
et Singer, 1967
Diagnosis. Gnathosoma positioned ventrally,
strongly reduced, oriented perpendicular to idio-
somal axis, retracted into idiosoma, and surround-
ed ventrally by idiosomal wall. In Cloacarinae,
remnants of posterior subcapitulum represented
by distinctly sclerotized subcapitular ring, open-
ing anteriorly and located immediately under dor-
sal surface of idiosoma. Pair of strongly sclero-
tized, fang-like chelicerae present ventrally. In
Pneumophaginae, female gnathosoma devoid of
posterior remnants of subcapitulum (subcapitular
ring) and internal median apodemes. In males,
gnathosoma completely absent. Idiosoma ellipti-
cal and distinctly attened dorso-ventrally.
Propodonotal shield present and occupying most
of propodonotal surface, distinctly ornamented in
Cloacarinae, and weakly sclerotized and devoid of
ornamentation in Pneumophaginae. Opisthosoma
distinctly shorter than metapodosoma; opistho-
somal striations absent. In females, anal and geni-
tal openings fused to each other forming single
opening situated ventrally, terminally or dorsally.
In females of some genera, lateral walls of genito-
anal opening elongated and modied into 1 pair of
short, nger-like projections. In Chelonacarus,
genito-anal opening covered ventrally by pair of
triangular sclerotized folds. In males, aedeagus
present and genital opening situated dorsally on
anterior part of body. Coxae completely fused with
ventral surface of idiosoma; only their anterior
margins represented by distinct coxal apodemes.
Coxal apodemes strongly elongated, longer than
half of idiosomal width. In adults, coxal apodemes
I fused forming sternum. In Cloacarinae, sternum
with internal median keel, at least in anterior half.
Keel continuing anteriorly as median apodeme
with bifurcate apex. Apices fused anteriorly with
subcapitular remnants and associated with scler-
ites, interpreted here as sigmoid pieces (= capitu-
lar apophyses). In females of Pneumophagus, an-
terior parts of coxal apodemes I fused with
anterior part of subcapitular remnants. Proximal
parts of coxal apodemes I–IV with 2 condyles at
trochanteral articulation. Dorsal condyle shifted
onto ventral side of idiosoma due to coxal atten-
ing in these mites. Ventral condyles distinctly
smaller than dorsal condyles. Idiosomal setae
strongly reduced in size and number. In Cloacari-
nae, female propodonotum with 1–2 pairs of al-
veoli. Opisthosoma of cloacarine females bearing
2–4 pairs of setae. Setation of male propodonotum
similar to that of females; opisthosoma with 1 pair
of terminal, eshy setae. In females of Pneumoph-
agus propodonotum with 1 pair of alveoli, opist-
hosoma with 3 pairs of short spine-like setae situ-
ated dorsally, terminally and ventrally. Pair of
alveoli anking genitor-anal opening dorsally. In
males, only 1 pair of short spine-like setae present,
situated dorsally on opisthosoma.
Legs I–IV with 3 articulated segments, very
short; rudimentary segment inserted proximal to
apical segment. Two basal leg segments devoid of
setae. Trochanters largest leg segments. Posterior
margins of all trochanters with large triangular
projection articulated with respective coxal apo-
demes. In most cloacarids reduced tibiae with 1
spine-like seta, but devoid of setae in Pneumopha-
gus, except for 1 seta on tibia I. Tarsi broadly
rounded in cloacarines; in Pneumophagus tarsi II–
IV attened laterally. Claws absent on all tarsi. In
Cloacarinae tarsi I and III each with 8 setae (in
Caminacarus chrysemys Pence and Casto, 1975,
tarsi III with 9 setae), tarsi II — 9 setae, and tarsi
IV — 7 setae. Short globose solenidion on tarsi
I–III situated ventro-terminally. In Pneumopha-
gus; tarsi I–IV bearing 5–9–12–12 setae, respec-
tively. Solenidion absent on all tarsi. Larva un-
known. Only 1 nymphal stage known.
The monophyly of the Cloacaridae is strongly
supported by the following synapomorphies: the
gnathosoma is inserted perpendicular to the idio-
somal axis, the subcapitulum is strongly reduced,
the palps are completely lost, the opisthosoma is
shortened and distinctly shorter than the metapo-
dosoma, the idiosomal striation is absent, there is
a sclerotized ventro-marginal sclerotized frame
bordering the lateral margins of the idiosoma, the
legs are inserted laterally and are strongly short-
ened and thickened, the femora and genua of all
the legs are completely fused, the leg trochanters
have distinct basal projection bearing glenoid cav-
ities, and the pretarsal claws are absent.
The family Cloacaroidea includes two sub-
families, Cloacarinae and Pneumophaginae with
one species. The latter, bird-associated subfamily
was described by Fain and Smiley (1989), and is
characterized by the following synapomorphies:
82
M. Skoracki, S.A. Zabludovskaya, A.V. Bochkov
the gnathosoma of the male is absent, the anterior
branches of coxal apodemes I surround the female
gnathosoma, the coxal apodemes are fused to each
other proximally forming a ventral idiosomal
frame, tibiae II–IV are devoid of setae, solenidia
are absent on all leg tarsi, tarsi II–IV are attened
laterally, and most of their setae are arranged in a
row (Bochkov and OConnor 2008).
Parvorder Eupodina Krantz, 1978
Superfamily Tydeoidea Kramer, 1877
The Tydeoidea includes four families: Tydei-
dae, Triophtydeidae, Iolinidae and Ereynetidae
(Andre and Fain 2000; Walter et al. 2009). This
superfamily has a worldwide distribution, and its
members are associated with a wide range of hab-
itats, from soil to the nasal cavities of mammals.
Highly specialized blood-sucking bird endopara-
sites are found only in the family Ereynetidae
(subfamily Speleognathinae) (Fain and van Goe-
them 1978, 1986; Domrow 1965, 1969). The phy-
logeny of the Tydeoidea, based on morphological
characters, was reconstructed by Andre and Fain
(2000). According to their results, the Ereynetidae
was derived from an ancestral tydeids and is char-
acterized by such autapomorphies as the presence
of the ereynetal organ and the double genital
discs.
Family Ereynetidae Oudemans, 1931
Figs. 11–12
Type genus: Ereynetes Berlese, 1883
Diagnosis. Chelicerae bases fused to each
other and with subcapitulum forming gnathosomal
capsule. Movable cheliceral digit short, linear and
stylet-like. Palps linear non-raptorial, composed
of 4 segments (trochanter, femuro-genu, tibia, tar-
sus) in Ereynetinae and reduced to 1–3 segments
in Lawrencarinae and Speleognathinae. Claw-like
structures correspond to strong ventral setae (v)
and observed only in Ereynetinae. Palpal setation,
femur-genu with 2 pairs of dorsal setae dF and dG
(in Ereynetinae, absent in other subfamilies), tibi-
ae with 2 dorsal setae dT and lT in Ereynetinae, 1
or without setae in other subfamilies, tarsus with
setae d, l’, l”, pζ, v, in Ereynetinae, reduced to 2–3
setae (d, l’, l”) in Lawrencarinae and Speleognathi-
nae, tarsal solenidion ω present in Ereynetinae and
present or absent in other subfamilies. Hypostomal
apex with 1–2 pairs of subcapitular setae (n, m).
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 modications).
83
A review of Prostigmata permanently associated with birds
Postcheliceral stigmata obscure. Idiosoma weakly
sclerotized, sac-like. Reticulate ornamentation of
prodorsum present or absent. Eyes present or ab-
sent. Propodonotal and hysteronotal shields pres-
ent in some ereynetines and absent in most of spe-
leognathines and Lawrencarinae. Ventral side of
idiosoma without plates. Distance between coxal
elds II and III distinctly shorter than idiosomal
width. Idiosomal setation (maximum in Ereyneti-
nae): vi, ve, si (= anterior sensilla), se, c1, c2, d1,
e1, f1, f2 (= posterior sensilla), h1. Pair of poste-
rior sensilla usually present in free-living species
and absent in most parasities. Cupules im and ia
present or absent. Genital opening situated poste-
rior to legs IV, longitudinal. Genital region with
setae: ag1–5, ps1–2, g1–5. Neotrichy of setae ps
observed only in Lawrencarus eweri (Lawrencari-
nae). In males of Ereynetinae 1–3 pairs of eugeni-
tal setae (eu) present, in Lawrencarinae and Spe-
leognathinae 1 pair of setae eu present or absent.
Genital papillae 2 pairs, well developed in Ereynet-
inae, or reduced in Lawrencarinae and Speleog-
nathinae. Legs long and massive, covered by re-
ticulate ornamentation. Coxal elds of legs I–IV
distinctly bordered and well sclerotized. Coxae
setation (maximum set) I–IV: I 1a, 1b, 1c; II 2c;
III 3a, 3b, 3c, 3d; IV 4a, 4b, 4c. Setae 3d and 4c
absent in Lawrencarinae and Speleognathinae.
Trochanters I–IV — I–III l’; IV without setae.
Femora I–IV — I with 4–7 setae; II (d), (v); III v,
A
B
Fig. 11. Boydaia sturni (Ereynetidae: Speleognathinae), female: A — in dorsal view; B — in ventral view.
B
A
Fig. 12. Boydaia sturni (Ereynetidae: Speleognathinae), details of female: A — ereynetidal organ (e.o.); B — tarsus I in dorsal
view.
84
M. Skoracki, S.A. Zabludovskaya, A.V. Bochkov
d, l; IV (d), (v). Femora I, IV divided on basi- and
telofemur in Ereynetinae, not in Lawrencarinae
and Speleognathinae. Genua I–IV — I (d) (l); II
(d), (l); III d, (l); IV d, (l). Tibiae I–IV — I d, (l),
(v); II d, (v), III d, (v); IV d (v). Tibiae I with in-
ternal sensory complex (ereynetal organ) where
solenidion of tibia I (φI) recessed into inverted
sac-like structure with narrow duct that opens
near distal margin of segment. Tarsi I–IV — I
with 12 setae, solenidion ω present or absent; II
with 9 setae, III and IV with 8 setae. Tarsi I–IV
with pair of large claws and with setiform median
empodium. In development 4 immature instars:
larva, protonymph, deutonymph, tritonymph. All
nymphs mobile in ontogenesis of Ereynetinae and
Lawrencarinae, while in Speleognathinae all
nymphs calyptostatic.
The taxonomic content of the family Ereyneti-
dae was reviewed by Fain (1957). He included the
family Speleognathidae erected by Womersley
(1936) to the Ereynetidae as a subfamily together
with the newly created Lawrencarinae.
Within the Ereynetidae, the Speleognathinae
form a monophyletic group. Its monophyly is sup-
ported by such synapomorphy as calyptostatic
nymphs. The subfamily Lawrencarinae is the sis-
ter group to Speleognathinae (Andre and Fain
2000).
In 1985, Fain divided the subfamily Speleog-
nathinae into ve tribes: Boydaiini, Trispeleog-
nathini, Speleognathini, Speleochirini, and Para-
speleognathopsini. Among them, representatives
of the two rst tribes are associated with birds.
The tribe Boidaiini with the genera, Boydaia and
Coboydaia is characterized by the absence of dor-
sal shields and eye lenses and by the presence of
3-segmented palps. The tribe Trispeleognathini in-
cludes 14 genera of nasal parasites of non-passer-
iform birds. These genera are characterized by the
polymorphic chaetotaxy and 1–3 segmented palps.
Additionally, eye lenses and dorsal shields are
present in some species.
Key to bird-associated genera of
Speleognathinae (Ereynetidae )
1. Chaetotaxy strongly variable, 2 or 3 different
types of setae present in same species. Tibiae I–IV
with 5–3(2)–3(2)–3(2). Dorsal shield and eye
lenses present or absent. Palps with 1–3 segments.
(Trispeleognathini Fain, 1980) .......................... 3
— All setae barbed. Tibiae I–IV with 5–3–3(2)–3(2)
or 4–2–2–2. Dorsal shield or eye lenses absent.
Palps with 3 segments (Boydaiini Fain, 1985) ..... 2
2. Setae on trochanters I and II present. Solenidion
on tarsi of palps present. Femur IV with at least 2
setae. Genu IV with at least 3 setae .......................
.................................... Boydaia Womersley, 1953
— Setae on trochanters I and II absent. Solenidion
on tarsi of palps absent. Femur IV with 1 seta.
Genu IV with 2 setae ........ Coboydaia Fain, 1971
3. Tarsi I–IV without striated setae .................... 5
— Tarsi I–IV with striated setae ........................ 4
4. Palps with 1 segment. Eye lenses absent. Num-
ber of coxal setae I–IV 1–1–1–1, trochanters with-
out setae ............................ Aureliania Fain, 1958
— Palps with 3 segments. One pair of prominent
eye lenses present. Number of coxal setae I–IV
2–1–1–0, trochanteral setae 1–1–0–0 .................
....................... Ophthalmognathus Dubinin, 1957
5. Apical or ventro-apical setae of tarsi I–IV nor-
mally developed and barbed .............................. 6
— Four apical and 2 ventro-apical setae of tarsi
I–IV rod-like microsetae ................. Psittaboydaia
Fain, 1985
6. All tarsal setae of legs I–IV barbed .............. 10
— Only 6–8 apical or ventro-apical setae of tarsi
I–IV barbed, other setae smooth ........................ 7
7. Dorsal and ventral setae of idiosoma dentate.
Dorsal shield absent ........................................... 9
— Ventral idiosoma with liform setae mixed with
striated or dentate setae. Dorsal shield variable ... 8
8. Dorsal shield present. Pair of eye lenses present
............................................ Neastrida Fain, 1962
— Dorsal shield and eye lenses absent ..................
........................................ Neoboydaya Fain, 1958
9. Palps with 3 segments. One pair of eye lenses
present. Claws of leg I stronger than claws II–IV.
Hypostome with 1 pair of setae. Five pairs of gen-
ital setae present. Number of coxal setae I–IV
2–1–1–1 ................. Trispeleognathus Fain, 1958
— Palps with 1 segment. Eye lenses absent. Claws
of leg I–IV subequal. Hypostome with 2 pairs of
setae. Two pairs of genital setae present. Number
of coxal setae I–IV 2–0–1–0 ........... Pterniboydaia
Fain, 1985
10. Dorsal and ventral setae of idiosoma either
dentate or dorsal setae mainly piliform or narrow
with striated base ............................................. 12
— Dorsal and ventral setae of idiosoma either ex-
clusively or predominantly barbed ................... 11
11. Dorsal shield well developed. Palps with 3
segments. Eyes either at or with prominent lenses
...................... Speleognathopsis Cooreman, 1954
— Dorsal shield absent. Palps with 2 segments.
Eyes with prominent lenses ........... Metaboydaia
Fain, 1962
85
A review of Prostigmata permanently associated with birds
12. Dorsal setae of idiosoma either exclusively or
predominantly dentate ...................................... 14
— Dorsal setae of idiosoma either exclusively or
predominantly liform ..................................... 13
13. Dorsal shield absent. Palps with 1 segment. Se-
tae vi and sce absent. Hypostomal setae absent .....
................................... Meropiboydaia Fain, 1985
— Dorsal shield present. Palps with 3 segments.
Setae vi and sce present. Hypostomal setae present
................................................. Astrida Fain, 1955
14. Claws of legs III and IV subequal in sizes ... 15
— Claws either of tarsi III and IV or only of tarsi
III unequal and modied; one or both abruptly
bent in their apical parts ................ Ralliboydaia
Fain, 1962
15. Dorsal shield absent. Palps with 3 segments.
Eyes with prominent lenses ...... Phoenicopteriella
Fain, 1970
–Large dorsal shield present. Palps with 1 seg-
ment. Eyes absent ............. Picinyssus Fain, 1969
RELATIONSHIPS WITH HOSTS
Microhabitats on the host body
Amongst parasitic acariform mites perma-
nently associated with their hosts, the external
morphological adaptations to parasitism were ana-
lyzed in feather mites (Psoroptidia) by Mironov
(1987) and in mites associated with mammals
(Bochkov 2007). According to several external
morphological criteria (structures of the gnatho-
soma, idiosoma, legs, and setation) these authors
have recognized different morpho-ecological
types (morphoecotypes); representatives of each
of them are associated with a particular habitat on
the host body. Prostigmatan mites permanently as-
sociated with birds, being represented by both
ecto- and endoparasities, occupy wide spectrum of
microhabitats on the host body: skin, subcutane-
ous layers, nasal cavities, feather quills, and even
bronchi (Table 1). Most of these mites belong to
morphoecotypes recognized in feather mites
(Mironov 1987).
Skin inhabiting prostigmatans of the families
Cheyletidae (Ornithocheyletiini) and Harpirhyn-
chidae (many genera of Harpirhynchinae) corre-
spond to the epidermoptoid morphoecotype
(Mironov 1987), which is analogous to the skin
morphoecotype of mammal-associated mites (Bo-
chkov 2007). Prostigmatan mites referred to this
morphoecotype have the terminally situated gna-
thosoma, which can possess various apophyses
used for attaching, the idiosoma is strongly at-
tened dorso-ventrally, distinctly developed seta-
tion of the idiosoma and legs, and the male genital
opening located dorsally. Their legs are inserted
laterally; ambulacra are well developed, at least,
on the anterior legs (Fig. 13C). Many representa-
tives of cheyletids spin the group (Bakericheyla)
or individual (Ornithocheyletia) cobweb covers.
Such cobweb cover protects mites and their eggs,
stabilises the temperature and humidity regime
and prevents elimination from the host body (Aki-
mov and Gorgol 1990).
Harpirhynchine females of some skin-inhab-
iting genera, for example, Neharpyrhynchus, are
quite different morphologically from other skin-
inhabiting mites. Their females are attached at the
base of calamus of the body covert feathers on the
neck or head of avian host. They are oriented par-
allel to the length of the shaft, with mouthparts
inserted in the host skin; their idiosoma is sacci-
form and the eggs are laid in a string; the female
opisthosoma, thus, serves as the “stalk” or site of
attachment for the eggs (Moss et al. 1979; Boch-
kov and Literak 2011) (Fig. 13D).
The bird skin almost devoid of glands is not so
benecial substrate as the mammalian skin, there-
fore, mites of the intracutaneous morphoecotype
like Demodicidae and Psorergatidae are absent
among bird-associated prostigmatans. Among
them, however, mites of the skin-burrowing mor-
phoecotype (analogous to the knemidocoptid mor-
phoecotype in feather mites) belonging to the fam-
ily Harpirhynchidae are present. In this family, the
tendency of transition from the skin surface (ec-
toparasitism) to intradermal parasitism (endopara-
sitism) is widely observed and this switches inde-
pendently happened within several harpirhynchid
genera (for example Harpyrhynchoides) (Fain
1995). Harpirhynchid females accumulate in the
feather follicles, where they develop subcutaneous
cysts (Fain 1995; Literak et al. 2005); many of
them are strongly morphologically modied. At
the same time, harpirhynchid males belonging to
skin-burrowing species do not undergo strong mor-
phological modications. Females of the subfamily
Harpypalpinae, forming the subcutaneous cysts
(Fig. 13B), are even less specialized than ectopara-
sitic mites of the subfamily Harpirhynchinae, be-
cause their posterior legs are not reduced, and their
opisthosomal setation is more reach than in harpi-
rhynchins. In some genera of Harpirhynchinae,
cyst-inhabiting species (Harpirhynchus or Hapy-
rhynchoides) also do not signicantly differ from
ectoparasitic (skin-inhabiting) representatives of
the family, whereas species of the genera Anharpy-
86
M. Skoracki, S.A. Zabludovskaya, A.V. Bochkov
rhynchus, Cypsharpirhynchus, Harpyrhynchiella,
and Trichorhynchiella are strongly modied being
habitually similar with some skin-burrowing mites
of the astigmatan subfamily Teinocoptinae (Sar-
coptoidea: Sarcoptidae). Their gnathosoma is re-
placed to the ventral idiosomal surface, the idioso-
ma is sacciform, the anterior legs are shortened and
the posterior legs are strongly reduced or lost; the
body and leg setation is very poor.
The parasitic prostigmatans inhabiting quills
correspond to the dermoglyphid morphoecotype
(Mironov 1987). Among prostigmatan mites this
morphoecotype is represented by mites of the fam-
ily Syringophilidae and cheyletid mites of the tribe
Metacheyletiini. The idiosoma of these mites is
strongly elongated (Syringophilidae) or egg-
shaped (Metacheyletia) and weakly sclerotized.
Legs are relatively short. In contrast to feather
mites which take harborage within quills, many
idiosomal and some leg setae of these mites are
long or ultralong and control, thus, a signicant
space around the mite. Mites of the subfamily Sy-
ringophilinae are mostly associated with the quills
of ight feather, whereas the absolute majority of
picobiins inhabit quills of the body feathers (John-
ston and Kethley 1973). Females of the latter sub-
family are characterized by the physogastry — an
extensive enlargement of the female hysterosoma
in the process of feeding (Skoracki et al. 2001;
Skoracki 2011) — it is unknown for other quill
mites.
The predatory mites inhabiting quills belong
to the tribe Cheletosomatini (Cheyletidae). These
mites do not differ habitually from nidicolous
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 modications).
87
A review of Prostigmata permanently associated with birds
cheyletids. Even mites of the cheletosomatine ge-
nus Picocheyletus transferred from a predation to
parasitism do not differ morphologically from
their predatory relatives. It should be noted, how-
ever, that some idiosomal and leg setae of these
mites are ultralong, as in parasitic prostigmatans
inhabiting quills.
The nasal prostigmatan parasites of birds are
represented by the subfamily Speleognathinae
(Ereynetidae). These mites belong to the respira-
tory morphoecotype established by Bochkov
(2007) and are not signicantly modied in com-
parison to free-living relatives. Their palps are
1–3-segmented (four-segmented palps in free-liv-
ing ereynetids), the idiosoma is slightly attened
dorso-ventrally and in some species bears the eyes,
the body and leg setae are relatively short, and the
legs are well developed, inserted laterally, and
bear empodium and paired claws; in males, an ae-
deagus is absent. An interesting adaptation of par-
asitic ereynetids is the presence of calyptostatic
nymphs. In these mites, the active stages are only
larva and adults (Andre and Fain 2001).
The interstitial bird parasites among prostig-
matans are represented by a single species Pneu-
mophagus bubonis (Cloacaridae) from tissues
around bronchi of Bubo virginianus (Strigiformes:
Strigidae) from USA (Fain and Smiley 1989). This
is a typical representative of the interstitial mor-
phoecotype. Among acariform mites permanently
associated with vertebrates this morphoecotype is
represented by the families Epimyodicidae and
Cloacaridae (Bochkov 2007). The body of these
mites has a slightly elongated and streamlined
form; in females, the gnathosoma is almost com-
pletely reduced and represented only by chelicerae
(absent in males), the legs are thickened and short-
ened and their segments are telescopic, almost all
body setae are absent, and the leg setation is
strongly reduced; the coxal apodemes are hyper-
trophied. Thus, cloacarid mites are almost ideally
adapted for moving in friable interstitial tissues.
Thus, bird-associated prostigmatans are rep-
resented by four main morphoecotypes: skin (epi-
dermoptid), skin-burrowing (knemidocoptid),
quill (dermoglyphid), and interstitial.
In feather mites and mammal-associated Ac-
ariformes, the skin-inhabiting forms belong to the
ancestral morphoecotype and most other morpho-
ecotypes are its derivations (Mironov 1987; Boch-
kov 2007). In prostigmatans, the colonization of
different microhabitats on the bird body has been
realized by free-living mites independently, most-
ly omitting a stage of the skin ectoparasitism. Quill
predators and parasites switched to the life in the
feather quills originate directly from the nest pre-
dation (Bochkov et al. 2008); nasal mites (Spe-
leognathinae) also originated directly from the
free-living forms omitting the skin parasitism; in-
terstitial mites (Pneumophaginae) supposedly
switched to birds from turtles and the origin of
cloacarid parasitism is still not clear (Bochkov and
OConnor 2008); only intracutaneous mites (some
Harpirhynchidae) originated from ectoparasitic
forms inhabiting the skin surface.
This complicated picture of colonization of
the avian body by prostigmatan mites could be ex-
plained by the absence of the direct phylogenetic
links between these mites. Since all bird-associat-
ed prostigmatans are true parasites feeding on the
live tissues or predators, they did not colonize the
feather vexillums (proctophyllodid morphoeco-
type) or the down feathers (analgid morphoeco-
type); nor they did occupy the quill walls inhabited
by some feather mites (Laminosioptidae).
Host specicity and evolution on birds
Bird systematics. All extant birds (Neorni-
thes) are split into two infraclasses: Palaeognathae
(two extant orders, ratites and tinamous), and
Neognathae with two cohorts, Galloanserae (land-
fowls and waterfowls) and Neoaves (other neog-
naths). The cohort Galloanserae includes two or-
ders, Galliformes and Anseriformes. The cohort
Neoaves includes 30 orders (Clements 2011). Tax-
onomic accounts of all bird orders are provided in
Table 6.
Hosts scpecicity. Most prostigmatans are
monoxenous or narrowly oligoxenous parasites
and only 8.7–30.5% of species in various groups
are associated with host of one family, 1.7–20.6%
— with hosts of one order and only 1.1–3.4% —
with hosts of various orders. All data about pros-
tigmatan host specicity are summarized in Table
7. Many species records, however, are solitary and
the level of host specicity could be overestimated
for some species. The most strictly specic symbi-
onts are parasitic Cheyletidae and Syringophili-
dae, 83% and 70.5% of monoxenous species, re-
spectively; the less specic symbionts are
Ereynetidae and Harpirhynchidae — 37% and
62.7%, respectively. It is interesting that some spe-
cies in the families, represented generally by the
highly specic parasites, are associated with wide
spectrum of hosts, for example, Bakericheyla
chanayi (Cheyletidae) (Akimov and Gorgol 1990;
88
M. Skoracki, S.A. Zabludovskaya, A.V. Bochkov
see also Appendix deposited at Acarina web-site:
http://insects.ummz.lsa.umich.edu/acarina/). In the
time of global cataclysms, such generalists ensure
the survival of the group when most hosts with
specialized parasites undergo extinction.
At the generic level the host specicity of
prostigmatan mites is less marked but still signi-
cant. In the Cheyletidae, four genera (36%) para-
sitize birds from one family, the genus Apodiche-
les is associated exclusively with Apodiformes,
and six other genera (54%) are associated with
birds of two or even more orders. In the Harpi-
rhynchidae, four genera (33%) are associated with
hosts of one family, four (33%) — with hosts of
one order, and four (33%) — with hosts of various
orders. In Syringophilidae, 39 genera (74%) para-
sitize birds of one order and only 14 genera (26%)
are associated with birds of two or even ve (Pico-
bia) various orders. Finally, in the Ereynetidae,
eight genera (50%) are associated with one bird
family, and eight (50%) — with birds of various
orders (Appendix).
Most prostigmatan families and subfamilies
are cosmopolite (Table 1), excluding Pneumocop-
tinae (Cloacaridae), which is still known from a
single record (Fain and Smiley 1989).
To date, prostigmatans are known from birds
from 25 of 34 orders belonging to neognathous as
well as to paleognathous (Tinamiformes) birds
(Table 6). These mites are absent on birds of the
orders Cariamiformes, Eurypygiformes, Gavii-
formes, Mesitornithiformes, Otidiformes, Pha-
ethoniformes, Sphenisciformes, Struthioniformes,
and Trogoniformes. Their absence on Struthioni-
formes and Sphenisciformes is explainable the ex-
ternal morphological modications of these hosts
(especially feathers), whereas birds of the other
orders are insufciently explored (or not explored
at all) for the presence of these mites. However,
with a high probability they could also be infected
by prostigmatans. Prostigmatans are characterized
by a relatively low prevalence and many of unex-
plored hosts are rare and poorly represented in
collections. Moreover, most of museum bird spec-
imens are preserved as dry skins and unavailable
for examination on prostigmatans because tech-
niques of their collecting can damage the skins.
The number of described prostigmatans rep-
resents only a small fraction of their actual biodi-
versity because their expected number is estimated
as 5000 species (instead of 500 presently known
species) based on the data about host specicity
and number of their potential hosts (Kethley and
Johnston 1975; Moss et al. 1979). At the same
time, it could be stated, based on distribution of
these mites, that at least 50% of their extant genera
are known to date. For example, among 110 extant
passerine families, prostigmatans are recorded on
hosts from more than 50 families (Appendix).
Mites of the family Cheyletidae are known
from birds of 12 orders, Syringophilidae — from
21 orders, Harpirhynchidae — from 16 orders and
Ereynetidae — from 18 orders (Table 6). Moss
and Wojcik (1978) also listed Caprimulgiformes
as hosts of harpirhynchids but did not provide the
full data; therefore these records were not taken
here into account.
As it is expected, representatives of all pros-
tigmatan families, excluding Cloacaridae, are re-
corded mostly from hosts of species-rich orders,
i.e. Apodiformes, Charadriformes, Columbifor-
mes, Coraciiformes, Cuculiformes, Galliformes,
Gruiformes, Passeriformes, Pelecaniformes, Pici-
formes, Psittaciformes, and Strigiformes.
The only mites of the family Syringophilidae
are known to date from palaeognathous birds of the
order Tinamiformes; whereas other prostigmatans
are still recorded exclusively from Neognathae.
Within the Neognathae, hosts of the more ar-
chaic Galloanserae clade are parasitized by most
prostigmatan families, excluding Cheyletidae and
Cloacaridae on Anseriformes, but the absolute
majority of mite species are associated with hosts
of the clade Neoaves and with passerines within
this clade.
Thus, birds were colonized independently by
mites of several prostigmatan phylogenetic lines
— three groups of cheyletids (Ornithocheyletiini,
Metacheyletiini, and Cheletosomatini), Syringo-
philidae, Harpirhynchidae, Ereynetidae (Speleog-
nathinae), and Cloacaridae (Pneumocoptinae).
As it was the above mentioned, the superfam-
ily Cheyletoidea, including most bird permanent
symbionts, is represented by two main phyloge-
netic lineages: Cheyletidae–Syringophilidae (I),
and Harpirhynchidae (Demodicidae–Psorergati-
dae) (II) (Bochkov 2008).
The ancestor of lineage I undoubtedly was a
predator preying on other arthropods. Most part of
the apomorphic modications of these mites con-
cerns the gnathosomal structures and, probably,
serves for more effective preying. At the same
time, features of the idiosoma and legs in these
mites are relatively archaic and weakly modied,
like those in the other predatory family Raphig-
nathidae. The family Syringophilidae originated
89
A review of Prostigmata permanently associated with birds
from the common ancestor with Cheyletidae,
which preyed in bird nests. Syringophilids devel-in bird nests. Syringophilids devel-
oped some adaptations to the parasitic mode of
life in feather quills (the stylophore surrounded by
the idiosoma and the elongate body); they have
also lost some structures, which serve for preying
and are useless for permanent parasites: the thumb-
claw complex, cupules, and the strong sclerotiza-
tion of the shields. In these mites, some idiosomal
setae (4a and ps3) have undergone a reduction in
parallel to parasitic cheyletoids of lineage II. In
Cheyletidae, the parasitism on birds originated in-
dependently in three phylogenetic lineages (tribes)
(Bochkov and Fain 2001; Bochkov 2004).
Family Cheyletidae. This family is repre-This family is repre-
sented by the predators and permanent parasites of
the terrestrial vertebrates, birds and small mam-
mals. To date, it includes more 380 species in 72
genera and is the most species-rich grouping com-
paring to other families of the superfamily Chey-
letoidea (Volgin 1969; Summers and Price 1970;
Bochkov 2004).
Cheyletids have a worldwide distribution and
are known from a wide spectrum of biotopes. They
occur in a soil, vegetable debris, in nests of in-
sects, mammals and birds (Volgin 1969). This
family includes almost all intermediate stages be-
tween free-living predators and highly specialized
permanent parasites of vertebrates, birds and
mammals (Bochkov and Fain 2001; Bochkov
2004). The obligatory phoresy on certain insects is
typical for some predatory cheyletids (Volgin
1969; Klimov et al. 2006). There are several gen-
eral directions in the cheyletid evolution: predator-
dendrophils, predator-entomophils, forms being
phoretic on insects and living in their nests, nidi-
colous predators in vertebrate nests, parasites of
birds and mammals (Volgin 1969; Bochkov
2004).
Among bird-associated cheyletids, mites of
the tribe Cheletosomatini (six genera) include
predators inhabiting the feather quills and prey
there on parasitic astigmatic mites and mites of
the family Syringophilidae. All representatives of
this tribe are morphologically similar to Ac-
aropsellina-like mites and probably originated
from the common ancestor inhabited bird nests.
The monotypic genus Picocheyletus was recently
described from quills of Tricholaema hirsuta a-
vipunctata from Gabon (Bochkov and OConnor
2003). These mites were collected from quills of
the primary and secondary wing feathers. In each
infested feather, the rachis was naturally split
ventrally in its basal part, exposing the hollow in-
terior. The mites occupied spaces between septae,
one mite per inter-septal space. In addition, from
one to four eggs were present in some inter-septal
spaces. It should be noted that no other mites or
remains of mites were observed inside the quills
that might serve as a prey. Some examined speci-
mens contained dark inclusions, similar to hema-
tite inclusions, which usually are visible in blood-
feeding cheyletids of the genus Bakericheyla
(Ornilhocheyletiini). Therefore, it can be con-
cluded that these mites are true parasites on the
avian host. In this case we see transition to para-
sitism happened literally in “our presence”.
Among other representatives of the tribe, species
of the genus Cheletoides are associated with gal-
liform birds (Gallifor mes). A single species of the
genus Eu che letop sis is recorded from tree swifts
of the genus Hemiprocne (Apodiformes:
Hemiprocnidae), and two species of the genus
Cheletosoma are associated with hosts of the or-
ders Gruiformes and Cuculiformes. The genus
Metacheyletoides includes species from galliform
(Galliformes) and cuculiform (Cuculiformes)
birds. Mites of the genus Cheletopsis are associ-
ated exclusively with charadriiform birds (Char-
adriiformes) (Bochkov et al. 2002; Bochkov and
OConnor 2003). Its representatives show variable
specicity to their hosts; some species are limited
to hosts of one genus, whereas others occur on
birds of several charadriiform families.
The tribe Ornithocheyletiini (four genera) is
represented by ectoparasites inhabiting the skin
surface of birds. Mites of the genus Neocheyle-
tiella are mono- or oligoxenous parasites associ-
ated with birds of four orders: Passeriformes,
Pelecaniformes, Columbiformes, and Strigi-
formes. Species of the genus Bakericheyla are
known from birds if three orders, Apodiformes,
Coraciiformes, and Passeriformes. Most of these
species are highly specic to their hosts and dis-
tributed in Africa, but Bakericheyla chanayi, how-
ever, is known from birds of various families and
orders and has a cosmopolitan distribution (Aki-
mov and Gorgol 1990). The genus Ornitocheyletia
is represented by parasites of six bird orders:
Columbiformes, Cuculiformes, Galliformes, Pas-
seriformes, Piciformes, and Psittaciformes. Orni-
tocheyletia species are mostly mono- or oligoxe-
nous parasites. Three species known in the genus
Apodicheles are restricted to swifts from South-
East Asia and Africa (Apodiformes: Apodidae).
Thus, mites of the tribe Ornithocheyletiini are
90
M. Skoracki, S.A. Zabludovskaya, A.V. Bochkov
known from representatives of several bird orders,
belonging to the two major phyletic lines, Gal-
loanserae (excluding Anseriformes) and Neoaves.
It can be hypothesized that the ancestor of this
tribe parasitized on a common ancestor of these
superorders.
The tribe Metacheyletiini includes the only
genus Metacheyletia with ve species inhabiting
the feather quills (Bochkov and Skoracki 2011).
Three of these species parasitize parrots from the
Old and New Worlds, and two species were re-
corded from quills of passeriform and cuculiform
birds in the Tropical Africa. The biology of Me-
tacheyletia species is still unknown. Atyeo et al.
(1984) believed that these mites are predators be-
cause their chelicerae are too short to pierce the
quill wall. Moreover, in all cases, these cheyletids
were collected together with other mites in those
quills. Bochkov and Fain (2001), however, had
another opinion and suggested that these mites are
parasites. According to these authors, the morpho-
logical characters of Metacheyletia species are
similar to those of other parasitic cheyletids: legs
I–III are short, legs IV are reduced, setae of the
palpal tarsi are short and smooth, the gnathosoma
is relatively small. It is suggested that these mites
use for feeding the openings pierced by the adult
syringophilids. This hypothesis is supported by
the presence of syringophilids in all cases where
these mites were found.
Family Syringophilidae. The syringophilid
ancestors were, probably, initially predators on
other mites inhabiting feather quills. According to
the estimation based on the “molecular clock”, the
divergence between cheyletids and syringophilids
could take place approximately 180–185 million
years ago in the Early Jurassic (Dabert et al. 2011).
Thus, syringophilids were, probably, associated
even with the bird ancestors — theropod dino-
saurs, many of which possessed feathers (Mayr et
al. 2005).
The host-parasite relationships of syringo-
philid were analyzed by Kethley and Johnston
(1975). According to data of these authors the
phylogenetic pattern of these mites and birds are
strongly incongruent at the level of mite genera
and bird orders. This incongruence could also be
explained by the horizontal syringophilid switches
on phylogenetically distant hosts. Kethley and
Johnston (1975) considered host switches as the
main modus in the evolution of this family and
even proposed a new term for it — resource track-
ing. According to their hypothesis, host distribu-
tion of syringophilid species is determined by
thickness of the quill walls and mite capability to
pierce it.
The second important evolution aspect in sy-
ringophilid evolution on their hosts is a distribu-
tion of these mites on various types of quills. The
wing-feathers, i.e. primaries and secondaries, are
probably the ancestral type of the syringophilid
habitat. The majority of representatives of the
family, including the most archaic ones, were as-
sociated with quills of these feathers. The subfam-
ily Picobiinae represented mostly by the dwellers
of the body covert feathers was probably originat-
ed in wing quills, because representatives of the
archaic genus Calamincola occupy these micro-
habitats (Casto 1977). In comparison to syringo-
philines, picobiines are much more morphologi-
cally specialized and possess some advantages,
ror instance heterosomy, which, probably, allowed
them to use more effectively small quills of the
body coverts. Thus, picobiines avoided competi-
tions with other syringophilids and formed the
second line distributed in parallel with syringophi-
lines on the same hosts. Unfortunately, the biodi-
versity of this group is yet poorly studied in com-
parison to Syringophilinae.
Among 48 genera of the subfamily Syringo-
philinae, 38 genera are restricted to particular avi-
an orders, whereas most of remaining mite genera
occur on birds of several orders.
Phylogenetic lineage II of Cheyletoidea is ex-
clusively represented by permanent parasites of
vertebrates which can be characterized by the
unique palpal structure. All three families of this
lineage demonstrate a tendency towards intracuta-
neous parasitism (a part of Harpirhynchidae, De-
modicidae and Psorergatidae) (Bochkov 2009).
Skin-inhabiting Harpirhynchidae are relatively
slow-moving ectoparasites, which are very often
embedded in the host epidermis. By this reason,
the reduction tendencies prevailed in the evolution
of this mite lineage.
It is presumed from the wide distribution of
these mites on birds (Harpirhynchidae) and mam-
mals (Psorergatidae–Demodicidae), that the com-
mon ancestor of this branch could occur on the
common ancestor of birds and mammals. Transi-
tions from birds, which are hosts of the most ar-
chaic representatives of line II (Harpypalpinae),
onto mammals, however, can not be excluded dur-
ing the early phase of mammal evolution (Bochk-
ov 2008). A possible reason for the absence of
cheyletoids on recent reptiles (excluding snakes)
91
A review of Prostigmata permanently associated with birds
may be explained by the peculiarities of their skin
structure and molting. Squamata lose the entire
external dermal layer or large pieces of it during
the molt (Landmann 1986); moreover, they have a
multi-layer keratinized epithelium, and skin glands
are absent. The high probability of loss of mites
during reptile molting seems to have prevented the
primary settling of cheyletoid parasites on these
hosts. These mites are absent also on crocodilians
probably because of their aquatic mode of life. In
birds, the skin has undergone signicant evolu-
tionary changes comparable to that in of mam-
mals. This probably allowed some cheyletoid
mites of the family Harpirhynchidae to transfer to
the intradermal parasitism in capsules that is simi-
lar to that developed by representatives of Psorer-
gatidae (Literak et al. 2005).
Family Harpirhynchidae. This family in-
cludes three subfamilies, two subfamilies, Harpi-
rhynchinae and Harpypalpinae are represented by
bird parasites and the subfamily Ophioptinae —
by ectoparasites of snakes from the superfamily
Colubroidea. The monophyly of this family is
strongly approved (Bochkov 2008), and hypothe-
sis about the independent origin of the subfamilies
Harpirhynchinae and Harpypalpinae from cheyle-
toid-like ancestors, proposed by Moss (1979), is
groundless.
Mites of the subfamily Harpirhynchinae are
known exclusively from Neognathae (Galloanser-
ae and Neoaves). Mites of the most archaic genus
Harpirhynchoides are known from 14 bird orders
(Fain 1994b). Mites of the genera Neharpirhynchus
and Metharpirhynchus parasitize passerines. In
addition, the rst genus is associated with hum-
ming birds (Apodiformes: Trochilidae), and the
second genus — with Picidae (Piciformes) (Fain
1995). Mites of the monotypic genus Anharpi-
rhynchus are associated with passerine and pici-
form birds. Another six genera are represented by
a few species which are associated with hosts of
one order. Mites of two genera, Harpirhynchus,
and Trichorhynchiella are associated with passer-
ines; mites of the genera Cypsharpirhynchus and
Harpyrhynchiella are known from Apodidae
(Apodiformes); the genus Perharpirhynchus is as-
sociated with hosts of the order Charadriiformes;
the mites of the genus Ralliharpirhynchus are
known only from Rallidae (Gruiformes) (Fain
1995).
Mites of the subfamily Harpypalpinae are
known from hosts of 11 families, all of them be-
long to higher passerines.
It looks like a paradox that mites of the more
archaic subfamily are associated with the highly
evolved hosts. On the contrary, the immature instars
of the Harpypalpinae are highly derived in their
morphology, because they have lost legs and are
characterized by a quite specialized idiosomal seta-
tion (Bochkov et al. 1999). Apparently in this case
we have two principally different pathways of the
morphological evolution in these two subfamilies
(Lombert and Moss 1983). In the Harpirhynchinae
the progressive characters are represented in adults,
whilst in Harpypalpinae they are developed in im-
matures. Relationships of Passeriformes with other
higher Neoaves are not clear (Kurochkin 1993). It
is possible that this order represents some earlier
separated branch. Certain parasitological data sup-
port the hypothesis of the early origin of Passeri-
formes. Thus, the rather archaic representatives of
the families Rhinonyssidae (Mesostigmata) and
Ereynetidae (Prostigmata) are associated with Pas-
seriformes (Moss 1979). The feather mite family
Proctophyllodidae, mostly restricted to the passer-
ines, is one of most evolved feather mite families
but is also characterized by certain archaic features
(Mironov 1998). If one admits the hypothesis of
earlier origin of Passeriformes the separate phylo-
genetic position of the Harpypalpinae could be eas-
ily explained by their coevolution with passerines.
The subfamily Ophioptinae is a sister lineage
to the Harpypalpinae (Bochkov et al. 1999). It was
hypothesized that harpypalpine-like ancestor of
Ophioptinae switched to parasitism on snakes
from passerines. The possibility of this host shift
was originally proposed by Kethley (after: Lombert
and Moss 1983). Certain snakes feed on nestlings
and adult birds. Most of this prey is small passer-
ine birds. The subfamily Ophioptinae is associated
exclusively with Colubridae and Elapidae (Boch-
kov et al. 1999). These two snake families are
closely related and represent a group of higher
snakes (Rieppel 1988).
Family Cloacaridae (Pneumophaginae).
The record of Pneumophagus bubonis from the
lungs of a single specimen of Bubo virginianus by
Fain and Smiley (1989) is still enigmatic. As dis-
cussed above, these mites exhibiting general cloa-
carid morphology are clearly distinguished from
the Cloacarinae, and their placement into a sepa-
rate subfamily has a strong morphological sup-
port. The distinctions between pneumophagines
and cloacarines are stronger than these between
cloacarines parasitizing two chelonian orders
Pleurodyra and Cryptodyra. Such morphological
92
M. Skoracki, S.A. Zabludovskaya, A.V. Bochkov
distinctions suggest a long history of pneumaphag-
ine parasitism on non-turtle hosts. We believe
therefore, that this record is not likely to be the
result of a host switch from some as yet unexam-
ined turtle species that happened within the
lifespan of that individual bird. On the other hand,
the knowledge of the diversity and host distribu-
tion of pneumophagines is so far limited to this
single record. Therefore we can not hypothesize
whether this parasitism has an ancient origin, and
birds have inherited cloacarids from their common
ancestor with turtles or, that seems much more
probable, it was the result of an ancient host shift
caused by predation by some birds on live or dead
turtles (Bochkov and OConnor 2008).
Family Ereynetidae (Speleognathinae).
This family is represented by three subfamilies.
Mites of the subfamily Ereynetinae are predators
inhabiting humid microhabitats or facultative par-
asites of mollusc lungs (snails). Mites of the sub-
family Lawrencarinae are endoparasites of amphi-
bian nasal cavities. The subfamily Spe leo gnathinae
consists of nasal endoparasites of mammals and
birds (Andre and Fain 2000). It could be suggested
that ereynetine-like ancestor of the subfamily
Lawrencarinae switched to parasitism on amphib-
ians from their prey — snails or directly from the
soil. In the absence of phylogenetic hypothesis for
speleognathine mites, two hypotheses of their host
relationships could be proposed. According to the
rst hypothesis, amniotic vertebrates received
these mites from amphibians or directly from the
soil. Monophyly of the subfamily Speleognathi-
nae is well grounded, therefore, such transition
should have happened on one of these two groups.
This host switch must be very ancient because the
wide distribution of these mites on avian and
mammal orders. An alternative hypothesis sug-
gests the initial parasitism of these mites on a
common ancestor of Anamnia and Amniota and
extinction of these mites on modern reptiles.
The subfamily Speleognathinae is separated
into ve tribes, two of them are associated with
birds and other three — with mammals (Fain
1985).
Two genera of the tribe Boydaiini, Boydaia
and Coboydaia are mostly associated with passer-
ines. Among 40 species of the rst genus, only
four species are, probably, secondary associated
with non passerine hosts — Caprimulgiformes,
Galliformes, Psittaciformes, and Falconiformes.
Among ve Coboydaia spp., only one is associat-
ed with humming birds.
The tribe Trispeleognathini includes the re-
maining 14 genera of bird-associated speleog-
nathines. All of them parasitize non passerine
birds, with one exception — Speleognathopsis
onychognathi (Fain, 1964). Most genera are as-
sociated with hosts of a single order: the genus
Trispeleognathus is associated with Columbi-
formes, Aureliania — with Strigiformes, Oph-
thalmognathus — with Pelecaniformes, Psitta-
boydaia — with Psittaciformes, Neastrida — with
Gruiformes, Neoboydaia — mostly with Char-
adriiformes and one species is recorded from
Podicipediformes, Pterniboydaia — with Galli-
formes, Meropiboydaya — with Coraciiformes,
and Picinyssus — with Piciformes. Five genera
are, however, associated with hosts of various or-
ders: Astrida — with Caprimulgiformes, Coracii-
formes, and Strigiformes, Metaboydaia — with
Coraciiformes and Pelecaniformes, Ralliboydaia
— Gruiformes and Cuculiformes, Phoenicopteri-
ella — with Anseriformes and Phoenicopteri-
formes, and Speleognathopsis — with Acipitri-
formes, Galliformes, Charadriiformes, and
Passeriformes.
Biology
Data about the biology of bird-associated
prostigmatans are very fragmental and far from
complete; almost nothing is known about the biol-
ogy of Pneumophaginae and Speleognathinae.
All prostigmatans permanently associated
with birds possess piercing chelicerae and feed on
live host (or prey) tissues. The exception are males
of Pneumophagus bubonis, their chelicerae are
lost but oral opening is still present (Bochkov and
OConnor 2008). There are two types of food spe-
cialization in skin-inhabiting prostigmatans: lym-
phophagy (most prostigmatans) and haematophagy
(Bakericheyla) (Akimov and Gorgol 1990).
Females of these mites lay eggs on (in) the
host body. Normally their life-cycle includes ac-
tive larva, two nymphal stages and adults, females
and males; no cases of parthenogenesis were re-
corded. The immatures of Pneumophagines, how-
ever, are still unknown (Bochkov and OConnor
2008) and all nymphal stages of Speleognathinae
are calyptostatic (Andre and Fain 2000). The life-
cycle of prostigmatans studied in this relationship
is short. In free-living cheyletids (Cheyletus) the
time of life-cycle critically depends from tempera-
ture and takes from 10 to 80 days (Akomov and
Gorgol 1990). In syringophilids, it takes about 40
days (Kethley 1971). In symbiotic cheyletids and
93
A review of Prostigmata permanently associated with birds
harpirhynchids these characteristics should prob-
ably be similar.
Prostigmatans, being permanent symbionts,
colonize new hosts by two main ways — from
parents to nestlings and in the time of contacts be-
tween adults (particularly during molt of gregari-
ous birds or at breeding time). The rst way is es-
pecially characteristic for syringophilids (Skoracki
2011). Is worth to note that in syringophilid mites
only fertilized females disperse from one host to
another, whereas the males never leave the feather
quills (Kethley 1971).
The population ecology of bird-associated
prostigmatans is weakly studied. They are charac-
terized by relatively low prevalence index (IP).
For parasitic cheyletids (Bakericheyla chanayi
and Ornithocheyletia sp.) it was estimated for dif-
ferent seasons and hosts as 0.4–8% (Akimov and
Gorgol 990). According to Kethley (1971) and
Skoracki (2011) , IP in syringophilids is very low
for the wild, non-social, and separately nesting
hosts, for example, IP for Lanius excubitor para-
sitized by Syringophiloidus weiszi is 3.5%
(N=508). In the same time IP for Hirundo rustica
infected with Syringophiloidus hirundus is 17%
(N=208). The data about syringophilid IP on wild
birds has been recently summarized and discussed
by Skoracki et al. (2010). The highest IP was
shown for social and domestic birds, e.g. IP for
Passer domesticus infested by Syringophiloidus
minor is 82% (N=492), or IP for Gallus gallus do-
mesticus is 75% (N=1.500). In various species of
the genus Neharpyrhynchus demonstrated on the
European birds 2–14% IPs (Martinu et al. 2008)
and 15–21% on birds in Peru (Literak et al. 2012);
mites of the genus Harpirhynchus dusbabeki from
Panurus biarmicus in Europe showed IP 7.4%
(N=378) (Literak et al. 2005).
The damage caused by prostigmatans to their
avian hosts is markedly varied. Most of these mites
do not cause a huge loss of bird condition. Some
cyst-forming harpirhynchids are considered as
harmful parasites (Moss 1979). At the same time
Henry et al. (2004) did not nd a markedly nega-
tive impact of the lesions by harpirhynchids on the
health status of birds infected. According to Lit-
erak et al. (2005), the traumatic effect from harpi-
rhynchid cysts was also not too serious and it
seems that birds with cysts survived for a long pe-
riod.
In contrast to some reports about the clinical
signs of feather’s picking caused by syringophilid
mites (mostly Syringophilus bipectinatus) in do-
mesticated birds (Rebrassier and Martin 1932;
Schwabe 1956; Hwang 1959; Gritschenko 1973;
Schmaschke et al. 2003; Principato et al. 2005)
Skoracki (2011) found no evidences that quill mites
induce the pathological changes in the structure of
skin in wild birds, even during heavy infestations.
Casto’s (1974) observations also indicate that the
exit channels and many small holes produced dur-
ing feeding activity of syringophilids, do not seem
to weaken the structure of the feather.
The role of prostigmatan mites as vectors of
bird diseases is almost unstudied. It is only known,
that syringophilids could be potential vectors of
bacterium Anaplasma phagocytophilum — an ob-
ligate intracellular pathogen (Skoracki et al.
2006).
ACKNOWLEDGEMENTS
We thank Dr S.V. Mironov (Zoological Insti-
tute RAS, St. Petersburg) for critical review of the
MS. The English of this MS was improved by
B. Nattress. We thank Dr. I. Literak for the photo
of harpirhynchid cyst. This research was support-
ed by the Russian Foundation for Basic Research
(Grant No 10-04-00160_a) and by the NCN grant
(2011/01/B/NZ8/01749). In the frames of this
project visit of AVB to the Adam Mickiewicz Uni-
versity, Poznan, Poland was supported by the spe-
cial grant from administration of this university.
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Skoracki, M., Hromada, M. and Kuczynski, L. 2001a.
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Skoracki, M., Hromada, M. and Tryjanowski, P. 2001b.
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Skoracki, M., Hromada, M. and Wamiti, W. 2011b. A
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Skoracki, M., Lontkowski, J. and Stawarczyk, T. 2010d.
New taxa of the parasitic quill mites associated
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Skoracki, M. and Magowski, W. 2001. Two new spe-
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103
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Table 1.
Summary of taxa number, distribution and host associations of Prostigmata permanently associated with
birds
Family
(subfamily or tribe)
Species n Genera n Host orders
Type of
symbiosis
Family Cheyletidae Leach, 1815
Tribe Cheletosomatini Volgin, 1969 23 6
Apodiformes, Charadriiformes,
Cucu liformes, Galliformes, Gruiformes,
Piciformes
Predators and
parasites
(Picocheyletus)
Tribe Ornithocheyletiini Volgin, 1969 54 4
Apodiformes, Columbiformes,
Cora ciiformes, Cuculiformes, Galli-
formes, Passeriformes, Pelecaniformes,
Piciformes, Psittaciformes, Strigiformes
Parasites
Tribe Metacheyletiini Fain, 1980 5 1
Cuculiformes, Passeriformes, Psit taci-
formes
?Parasites
Family Syringophilidae Lavoipierre, 1953
Subfamily Syringophilinae
Lavoipierre, 1953
234 48
Accipitriformes, Anseriformes,
Apo di formes, Charadriiformes,
Ciconiiformes, Columbiformes,
Coraciiformes, Cuculiformes, Galbuli-
formes, Galliformes, Gruiformes,
Passeriformes, Pelecaniformes,
Phoenicopteriformes, Piciformes,
Procellariiformes, Psittaciformes,
Pte ro cliformes, Strigiformes, Suli-
formes, Tinamiformes
Parasites
Subbfamily Picobiinae
Johnston et Kethley, 1973
44 5
Charadriiformes, Columbiformes,
Coraciiformes, Cuculiformes, Galli-
formes, Passeriformes, Piciformes,
Psittaciformes, Pterocliformes
Parasites
104
M. Skoracki, S.A. Zabludovskaya, A.V. Bochkov
Family Harpirhynchidae Dubinin, 1956
Subfamily Harpirhynchinae
Dubinin, 1956
65 10
Accipitriformes, Anseriformes,
Apodiformes, Charadriiformes,
Ciconiiformes, Coliiformes, Columbi-
formes, Cuculiformes, Galliformes,
Gruiformes, Passeriformes, Pelecani-
formes, Piciformes, Procellariiformes,
Psittaciformes, Strigiformes
Parasites
Subfamily Harpypalpinae Fain, 1972 10 2 Passeriformes Parasites
Family Cloacaridae Camin, Moss, Oliver et Singer, 1967
Subfamily Pneumophaginae
Fain et Smiley, 1989
1 1 Strigiformes Parasites
Family Ereynetidae Oudemans, 1931
Subfamily Speleognathinae Fain, 1957
Tribe Boydaiini Fain, 1985 45 2
Apodiformes,Caprimulgiformes,
Falconiformes, Passeriformes, Psittaci-
formes
Parasites
Tribe Trispeleognathini Fain, 1985 28 14
Accipitriformes, Anseriformes,
Caprimulgiformes, Charadriiformes,
Columbiformes, Coraciiformes,
Cuculiformes, Galliformes, Gruiformes,
Passeriformes, Pelecaniformes,
Phoenicopteriformes, Piciformes,
Podicipediformes, Psittaciformes,
Strigiformes
Parasites
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
105
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)
Cloacaridae (Pneumophaginae)
1 pair of propodonotal alveoli (0); 1 pair of alveoli anking anal-genital opening (0), 3
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
(Pneumophaginae)
5 9 12 12
Ereynetidae
(Speleognathinae)
12, ω
8 7 7
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
(Pneumophaginae)
1 – – –
Ereynetidae
(Speleognathinae)
d, (l), (v) d, l, v d, l, v d, l, v
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
(Speleognathinae)
(d), (l) (d), (l) d (l) d, (l)
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 – –
106
M. Skoracki, S.A. Zabludovskaya, A.V. Bochkov
Harpirhynchiane
d, v d, v – –
Harpypalpinae
d, v d, v v v
Cloacaridae
(Pneumophaginae)
– – – –
Ereynetidae
(Speleognathinae)
7 (d), (v) d, (v) d (v)
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
(Speleognathinae)
l’/1a, 1b, 1c l’/2c –/3a, 3b –/4a, 4b
() — pair of setae; (M) — solenidion of male
Table 6.
Distribution of Prostigmata permanently associated with birds among host orders
Host order (N of extant species)
Mite family
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
107
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-specicity among Prostigmata permanently associated with birds
Number of mites specic to host order, family, genus and species respectively.
Mite family (N of species)
Host specicity
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%)
Appendix
i1
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
(Berlese, 1889)
Turdus merula
Turdidae Passeriformes Europe Berlese 1889
Ornithocheyletia aitkeni
Fain, 1972
Turdus fumigatus
Turdidae Passeriformes South America Fain 1972, 1981
Ornithocheyletia
dubinini Volgin, 1984
Sturnus vulgaris
Sturnidae Passeriformes
Europe, North
America
Volgin 1964;
Smiley 1977;
Fain 1981;
Skoracki et al.
2004
Ornithocheyletia eulabes
Fain, 1881
Gracula religiosa
Sturnidae Passeriformes
European Zoo
(captive)
Fain 1981
Ornithocheyletia
lamprocolius Fain, 1972
Lamprotornis
chloropterus
Sturnidae Passeriformes Africa Fain 1972, 1981
Ornithocheyletia wauthy
Fain et Bochkov, 2002
Lammprocolius sp.
Sturnidae Passeriformes Africa
Fain and
Bochkov 2002
Ornithocheyletia volgini
Smiley, 1970
Petrochelidon
pyrrhonota
Hirundinidae Passeriformes North America
Smiley 1970;
Fain 1981
Ornithocheyletia
lukoschusi Smiley, 1970
Hirundo rustica
Hirundinidae Passeriformes Europe
Smiley 1970;
Fain 1981
Ornithocheyletia
mironovi Bochkov et
Chirov, 1998
Riparia riparia
Hirundinidae Passeriformes Asia
Chirov and
Bochkov 1998
Ornithocheyletia
phylloscopi Bochkov,
Mironov et Gorgol, 1994
Phylloscopus
trochilus
Sylviidae Passeriformes Europe
Bochkov et al.
1994
Ornithocheyletia
leiothrix Fain, 1972
Leiothrix lutea
Leiothrichidae Passeriformes
European Zoo
(captive)
Fain 1972, 1981
Ornithocheyletia
garrulax Fain, 1972
Garrulax
leucolophus
Leiothrichidae Passeriformes
European Zoo
(captive)
Fain 1972, 1981
Ornithocheyletia lepidus
Fain, 1981
Garrulax
leucolophus
Leiothrichidae Passeriformes
European Zoo
(captive)
Fain 1981
Ornithocheyletia
granatina Fain, 1972
Uraeginthus
ianthinogaster
Estrildidae Passeriformes
European Zoo
(captive)
Fain 1972, 1981
Ornithocheyletia orioli
Fain et Bochkov, 2003
Oriolus oriolus
Oriolidae Passeriformes Africa
Fain and
Bochkov 2002
“
Pachyphantes
superciliosa
Ploceidae Passeriformes Africa
Fain and
Bochkov 2002
“
Lybius sp.
Lybiidae Piciformes Africa
Fain and
Bochkov 2002
“
Indicator indicator
Indicatoridae Piciformes Africa
Fain and
Bochkov 2002
“
Chrysococcyx
klaasi
Cuculidae Cuculiformes Africa
Fain and
Bochkov 2002
M. Skoracki, S. Zabludovskaya and A.V. Bochkov
i2
Ornithocheyletia
canadensis (Banks,
1909)
Picus viridis
Picidae Piciformes North America
Banks 1909;
Fain 1981
Ornithocheyletia gersoni
Smiley, 1970
Dendrocopos
pubescens
Picidae Piciformes North America
Smiley 1970;
Fain 1981
Ornithocheyletia hallae
Smiley, 1970
Columba livia
Columbidae Columbiformes
Europe, Africa,
North America
Smiley 1970,
1977; Fain 1981
Ornithocheyletia hallae
similis Fain, 1972
Chalcophaps
indica
Columbidae Columbiformes
European Zoo
(captive)
Fain 1972, 1981
Ornithocheyletia
geopeliae Fain, 1981
Geopelia striata
Columbidae Columbiformes
European Zoo
(captive)
Fain 1981
Ornithocheyletia
columbigallinae Fain et
Bochkov, 2002
Columbina sp.
Columbidae Columbiformes South America
Fain and
Bochkov 2002
Ornithocheyletia
lawrenceae Smiley, 1970
Parakeet
Psittacidae Psittaciformes North America Smiley 1970
“
Platycercus
elegans
Psittacidae Psittaciformes Australia
Womersley 1941;
Fain 1981
Ornithocheyletia
psittaculae Fain, 1972
Psittacula krameri
Psittacidae Psittaciformes
European Zoo
(captive)
Fain 1972, 1981
Ornithocheyletia psittaci
Fain, 1972
Psittacus erithacus
Psittacidae Psittaciformes
European Zoo
(captive)
Fain 1972, 1981
Ornithocheyletia psittaci
poicephali Fain, 1972
Poicephalus
senegalus
Psittacidae Psittaciformes Africa Fain 1981
Ornithocheyletia smileyi
Fain, 1972
Myiopsitta
monachus
Psittacidae Psittaciformes
European Zoo
(captive)
Fain 1972, 1981
Ornithocheyletia
argentinensis Fain, 1972
Nandayus nenday
Psittacidae Psittaciformes
European Zoo
(captive)
Fain 1972, 1981
“
Forpus passerinus
Psittacidae Psittaciformes
European Zoo
(captive)
Fain 1972, 1981
Ornithocheyletia
francolini Fain, 1972
Francolinus
natalensis
Phasianidae Galliformes Africa Fain 1972, 1981
Neocheyletiella Baker, 1949
Neocheyletiella rohweri
Baker, 1949
Sitta pygmaea
Sittidae Passeriformes North America Baker 1949
Neocheyletiella
microrhyncha (Berlese et
Trouessart, 1889)
Hirundo rustica
Hirundinidae Passeriformes
Europe, North
America
Berlese and
Trouessart 1889 ;
Smiley 1970
“
Delichon urbica
Hirundinidae Passeriformes Europe Volgin 1969
“
Riparia riparia
Hirundinidae Passeriformes Europe Volgin 1969
“
Petrochelidon
pyrrhonota
Hirundinidae Passeriformes North America Smiley 1970
“
Cecropis
abyssinica
Hirundinidae Passeriformes Africa Fain 1980a
“
Psalidoprocne
albiceps
Hirundinidae Passeriformes Africa Fain 1980a
Neocheyletiella
smallwoodae Baker,
1949
Leucosticte
australis
Fringillidae Passeriformes Australia Baker 1949
Neocheyletiella media
Fain, 1972
Leiothrix lutea
Timaliidae Passeriformes Asia Fain 1972, 1980a
Neocheyletiella siva
Fain, 1972
Minla
cyanouroptera
Timaliidae Passeriformes
European Zoo
(captive)
Fain 1972, 1980a
Neocheyletiella
amandavae Fain, 1972
Amandava
amandava
Leiothrichidae Passeriformes
European Zoo
(captive)
Fain 1972, 1980a
Appendix
i3
Neocheyletiella
megaphallos Lawrence,
1959
Estrilda
erythronotos
Estrildidae Passeriformes Africa Lawrence 1959a
Neocheyletiella
lonchurae Fain et
Bochkov, 2002
Lonchura sp.
Estrildidae Passeriformes Africa
Fain and
Bochkov 2002
Neocheyletiella pittae
Fain, 1972
Pitta megarhyncha
Pittidae Passeriformes
European Zoo
(captive)
Fain 1972, 1980a
Neocheyletiella artami
Domrow, 1966
Artamus
cyanopterus
Artamidae Passeriformes Australia
Domrow 1966,
1970
Neocheyletiella avicola
Fain, 1972
Ara sp.
Psittacidae Psittaciformes
European Zoo
(captive)
Fain 1972, 1980a
“
Agapornis sheri
Psittacidae Psittaciformes
European Zoo
(captive)
Fain 1980a
“
Erythrura prasina
Estrildidae Passeriformes
European Zoo
(captive), North
America
(captive)
Smiley 1977 ;
Fain 1980a
Neocheyletiella
nectarinia Bochkov et
OConnor, 2003
Cinnyris jugularis
Nectariniidae Passeriformes Asia
Bochkov and
OConnor 2003
Neocheyletiella aegithali
Bochkov et OConnor,
2003
Aegithalos
concinnus
Aegithalidae Passeriformes Asia
Bochkov and
OConnor 2003
Neocheyletiella queleae
Fain et Bochkov, 2002
Quelea quelea
Ploceidae Passeriformes Africa
Fain and
Bochkov 2002
“
Streptopelia
semitorquata
Columbidae Columbiformes Africa
Fain and
Bochkov 2002
Neocheyletiella ardeola
Bochkov et Ochoa, 2005
Ardeola ralloides
Ardeidae Pelecaniformes Africa
Bochkov and
Ochoa 2005
Neocheyletiella athene
Fain et Bochkov, 2002
Athene brama
Strigidae Strigiformes
European Zoo
(captive)
Fain and
Bochkov 2002
Bakericheyla Volgin, 1966
Bakericheyla chanayi
(Berlese et Trouessart,
1889)
Motacilla alba
Motacillidae Passeriformes Europe
Berlese and
Trouessart 1889
“
Fringilla coelebs
Fringillidae Passeriformes Europe
Baker 1949;
Volgin 1966
“
Fringilla
montifringilla
Fringillidae Passeriformes Europe
Akimov and
Gorgol 1990
“
Spinus spinus
Fringillidae Passeriformes Europe
Akimov and
Gorgol 1990
“
Carduelis carduelis
Fringillidae Passeriformes Europe
Akimov and
Gorgol 1990
“
Passer montanus
Passeridae Passeriformes Europasia
Akimov and
Gorgol 1990
“
Passer domesticus
indicus
Passeridae Passeriformes Asia
Akimov and
Gorgol 1990
“
Hirundo rustica
Hirundinidae Passeriformes Europe
Akimov and
Gorgol 1990
“
Delichon urbica
Hirundinidae Passeriformes Europe Volgin 1969
“
Motacilla ava
Motacillidae Passeriformes Europe
Akimov and
Gorgol 1990
“
Anthus trivialis
Motacillidae Passeriformes Europe
Akimov and
Gorgol 1990
“
Oenanthe picata
Muscicapidae Passeriformes Asia Volgin 1969
M. Skoracki, S. Zabludovskaya and A.V. Bochkov
i4
Bakericheyla chanayi
(Berlese et Trouessart,
1889)
Cyanoptila
cyanomelana
Muscicapidae Passeriformes Asia Volgin 1969
“
Muscicapa striata
Muscicapidae Passeriformes Europe Volgin 1969
“
Turdus pilaris
Turdidae Passeriformes Europe
Akimov and
Gorgol 1990
“
Sylvia atricapilla
Sylviidae Passeriformes Europe
Akimov and
Gorgol 1990
“
Sylvia borin
Sylviidae Passeriformes Europe
Akimov and
Gorgol 1990
“
Phylloscopus
trochilus
Sylviidae Passeriformes Europe
Akimov and
Gorgol 1990
“
Parus major
Paridae Passeriformes Europe Volgin 1969
“
Paroaria
nigrogenis
Emberizidae Passeriformes
European Zoo
(captive),
Fain 1972, 1980b
“
Emberiza citrinella
Emberizidae Passeriformes Europe
Akimov and
Gorgol 1990
“
Zonotrichia
leucophrys
Emberizidae Passeriformes North America
Furman and
Sousa 1969
“
Zonotrichia
atricapilla
Emberizidae Passeriformes North America
Furman and
Sousa 1969
“
Merops apiaster
Meropidae Coraciiformes Asia Volgin 1969
Bakericheyla faini
(Lawrence, 1959)
Cossypha dichroa
Muscicapidae Passeriformes Africa Lawrence 1959a
Bakericheyla subquadra-
ta (Lawrence, 1959)
Merops pusillus
Meropidae Coraciiformes Africa Lawrence 1959a
Bakericheyla transvaa-
lica (Lawrence, 1959)
Merops pusillus
Meropidae Coraciiformes Africa Lawrence 1959a
“
Merops persicus
Meropidae Coraciiformes Africa Lawrence 1959a
“
Merops apiaster
Meropidae Coraciiformes Africa, Asia
Lawrence 1959a;
Volgin 1969;
Fain 1980b
“
Merops persicus
Meropidae Coraciiformes Asia Volgin 1969
“
Merops nubicoides
Meropidae Coraciiformes Africa Fain 1980b
“
Merops
bullockoides
Meropidae Coraciiformes Africa Fain 1980b
Bakericheyla (Apodicheyla) Fain, 1979
Bakericheyla africana
Fain 1979
Cypsiurus parvus
Apodidae Apodiformes Africa Fain 1979a
Bakericheyla benoiti
Fain, 1980
Merops bulocki
Meropidae Coraciiformes Africa Fain 1980b
Apodicheles Fain, 1979
Apodicheles cypsiurus
Fain, 1979
Cypsiurus parvus
Apodidae Apodiformes Africa Fain 1979a
Apodicheles apus Fain,
1979
Apus caffer
Apodidae Apodiformes Africa Fain 1979a
Apodicheles collocalia
Bochkov et OConnor,
2003
Collocalia
esculenta
Apodidae Apodiformes Asia
Bochkov and
OConnor 2003
Metacheyletiini Fain, 1980
Metacheyletia Fain, 1972
Metacheyletia obesa
Fain, 1972
Psittacula krameri
Psittacidae Psittaciformes Africa Fain 1972, 1980c
Appendix
i5
Metacheyletia longise-
tosa Atyeo, Kethley et
Perez, 1984
Amazona nschi
Psittacidae Psittaciformes North America Atyeo et al. 1984
Metacheyletia amazonae
Bochkov et OConnor,
2003
Amazona
amazonica
Psittacidae Psittaciformes South America
Bochkov and
Oconnor 2003
Metacheyletia degene-
rata Fain et Bochkov,
2003
Serinus
mozambicus
Fringillidae Passeriformes Africa
Fain and
Bochkov 2003
Metacheyletia ngaii
Bochkov et Skoracki,
2011
Corythaixoides
leucogaster
Musophagidae Cuculiformes Africa
Bochkov and
Skoracki 2011
Cheletosomatini Volgin, 1969
Cheletosoma Oudemans, 1905
Cheletosoma tyrannus
Oudemans, 1905
Aramus guarauna
Aramidae Gruiformes South America Oudemans 1905
Cheletosoma tauraco
Bochkov et Scoracki,
2012
Tauraco
leucolophus
Musophagidae Cuculiformes Africa
Bochkov and
Skoracki 2012
Cheletoides Oudemans, 1904
Cheletoides uncinatus
Heller, 1880
Pavo cristatus
Phasianidae Galliformes
Europe
(domesticated)
Heller 1880 ;
Oudemans 1906
Cheletoides chirunduen-
sis Fain, 1972
Numida meleagris
Numididae Galliformes Africa Fain 1972, 1979b
Metacheletoides Fain, 1972
Metacheletoides
numidae Fain, 1972
Numida meleagris
Numididae Galliformes Africa
Fain 1972,
1979b; Bochkov
and Skoracki
2012
Metacheletoides
akanyaruensis Fain,
1972
Numida meleagris
Numididae Galliformes Africa Fain 1972, 1979b
Metacheletoides crinifer
Fain, 1979
Crinifer zonurus
Musophagidae Cuculiformes Africa Fain 1979b
“
Crinifer piscator
Musophagidae Cuculiformes Africa Fain 1979b
“
Corythaixoides
leucogaster
Musophagidae Cuculiformes Africa Fain 1979b
Metacheletoides
gisagarensis Fain, 1979
Crinifer zonurus
Musophagidae Cuculiformes Africa Fain 1979b
“
Crinifer piscator
Musophagidae Cuculiformes Africa Fain 1979b
Cheletopsis Oudemans, 1904
Cheletopsis norneri
(Poppe, 1888)
Sterna hirundo
Laridae
Charadrii-
formes
Eurasia
Poppe 1888 ;
Oudemans 1904,
1906 ; Volgin
1969 ; Mironov
et al. 1991
“
Sterna repressa
Laridae
Charadrii-
formes
Africa
Bochkov et al.
2002
“
Phaetusa simplex
Laridae
Charadrii-
formes
South America
Bochkov et al.
2002
“
Gelochelidon
nilotica
Laridae
Charadrii-
formes
Eurasia, North
America
Volgin 1969;
Mironov et al.
1991
“
Chlidonias hybrida
Laridae
Charadrii-
formes
Africa
Bochkov et al.
2002
M. Skoracki, S. Zabludovskaya and A.V. Bochkov
i6
Cheletopsis norneri
(Poppe, 1888)
Tringa totanus
Scolopacidae
Charadrii-
formes
Asia
Mironov et al.
1991; Bochkov
et al. 2002
Cheletopsis thalasseus
Bochkov et OConnor,
2003
Thalasseus
sandvicensis
Laridae
Charadrii-
formes
North America
Bochkov and
OConnor 2003
Cheletopsis larosterna
Bochkov et Skoracki,
2012
Larosterna inca
Laridae
Charadrii-
formes
South America
Bochkov and
Skoracki 2012
Cheletopsis basilica
Oudemans, 1904
Tringa totanus
Scolopacidae
Charadrii-
formes
Europe Oudemans 1904
“
Charadrius
hiaticula
Charadriidae
Charadrii-
formes
Europe
Bochkov et al.
2002
“
Calidris melanotos
Scolopacidae
Charadrii-
formes
North America
Bochkov et al.
2002
“
Chlidonias niger
Laridae
Charadrii-
formes
North America
Bochkov et al.
2002
Cheletopsis animosa
Oudemans, 1904
Tringa totanus
Scolopacidae
Charadrii-
formes
Europe Oudemans 1904
“
Sterna hirundo
Laridae
Charadrii-
formes
North America
Bochkov et al.
2002
Cheletopsis anax
Oudemans, 1904
Tringa totanus
Scolopacidae
Charadrii-
formes
Europe Oudemans 1904
Cheletopsis mariae
Mironov, Bochkov et
Chirov, 1991
Actitis hypoleucos
Scolopacidae
Charadrii-
formes
Eurasia
Mironov et al.
1991 ; Bochkov
et al. 2002
Cheletopsis impavida
Oudemans, 1904
Tringa totanus
Scolopacidae
Charadrii-
formes
Eurasia
Oudemans 1904;
Mironov et al.
1991
“
Tringa stagnatilis
Scolopacidae
Charadrii-
formes
Europe
Bochkov et al.
2002
“
Calidris ferruginea
Scolopacidae
Charadrii-
formes
Europe
Bochkov et al.
2002
“
Calidris minuta
Scolopacidae
Charadrii-
formes
Asia
Mironov et al.
1991
“
Calidris temminckii
Scolopacidae
Charadrii-
formes
Asia
Mironov et al.
1991
“
Calidris rucollis
Scolopacidae
Charadrii-
formes
Asia
Mironov et al.
1991
“
Calidris pusilla
Scolopacidae
Charadrii-
formes
North America
Bochkov et al.
2002
“
Calidris
himantopus
Scolopacidae
Charadrii-
formes
North America
Bochkov et al.
2002
“
Nycticryphes
semicollaris
Rostratulidae
Charadrii-
formes
North America
Bochkov et al.
2002
Cheletopsis daberti
Kivganov et Bochkov,
1994
Tringa glareola
Scolopacidae
Charadrii-
formes
Europa
Kivganov and
Bochkov 1994 ;
Bochkov et al.
2002
“
Tringa nebularia
Scolopacidae
Charadrii-
formes
Europe
Bochkov et al.
2002
“
Calidris temminckii
Scolopacidae
Charadrii-
formes
Europe
Bochkov et al.
2002
“
Calidris
himantopus
Scolopacidae
Charadrii-
formes
North America
Bochkov et al.
2002
Appendix
i7
Cheletopsis charadrii
Mironov, Bochkov et
Chirov, 1991
Charadrius dubius
Charadriidae
Charadrii-
formes
Asia
Mironov et al.
1991
Cheletopsis limnodromi
Bochkov, Fain et Dabert,
2002
Limnodromus
griseus
Scolopacidae
Charadrii-
formes
North America
Bochkov et al.
2002
Cheletopsis prosobonia-
lis Bochkov, Fain et
Dabert, 2002
Prosobonia
cancellata
Scolopacidae
Charadrii-
formes
Australia
Bochkov et al.
2002
Cheletopsis rhynchops
Bochkov, Fain et Dabert,
2002
Rynchops
avirostris
Laridae
Charadrii-
formes
Africa
Bochkov et al.
2002
“
Rynchops niger
Laridae
Charadrii-
formes
North America
Bochkov et al.
2002
Species inquirrenda
Cheletopsis magnanima
Oudemans, 1904
Tringa avipes
Scolopacidae
Charadrii-
formes
South America
Oudemmans
1904; Bochkov
et al. 2002
Picocheyletus Bochkov et OConnor, 2003
Picocheyletus tricholae-
mae Bochkov et
OConnor, 2003
Tricholaema
hirsuta
Lybiidae Piciformes South America
Bochkov and
OConnor 2003
Eucheletopsis Volgin, 1969
Eucheletopsis major
Trouessart, 1893
Hemiprocne
mystacea
Hemiprocnidae Apodiformes Australasia
Trouessart 1893;
Oudemans 1906
Harpirhynchidae Dubinin, 1957
Harpirhynchinae Dubinin, 1957
Harpirhynchus Megnin, 1877
Harpirhynchus nidulans
(Nitzsch, 1818)
Chloris chloris
Fringillidae Passeriformes
Europe, North
America
Nitzsch 1818;
Fain 1995; Moss
1979
“
Coccothraustes
coccothraustes
Fringilidae Passeriformes
Europe, North
America
Fritsch 1954;
Literak et al.
2005; Moss 1979
“
Fringilla coelebs
Fringillidae Passeriformes Europe Fritsch 1954
“
Corvus corone
Corvidae Passeriformes
Unknown
locality
Moss 1979
“
Locustella
lanceolata
Locustellidae Passeriformes North America Moss 1979
Harpirhynchus galeridae
Fain, Bochkov et
Mironov, 1999
Galerida cristata
Alaudidae Passeriformes Asia
Moss 1979; Fain
et al. 1999
“
Alauda arvensis
Alaudidae Passeriformes
Unknown
locality
Moss 1979
Harpirhynchus dusbabe-
ki Bochkov et Literak,
2006
Panurus biarmicus
Panuridae Passeriformes Europe
Henry et al.
2004; Literak et
al. 2005;
Bochkov and
Literak 2006
Harpirhynchus quasi-
modo Bochkov et
Mertins, 2010
Molothrus ater
Icteridae Passeriformes North America
Bochkov and
Mertins 2010
Harpyrhynchoides Fain, 1972
Harpyrhynchoides
squamosus (Fain, 1972)
Psittacula
cyanocephala
Psittacidae Psittaciformes
European Zoo
(captive)
Fain 1972, 1994
M. Skoracki, S. Zabludovskaya and A.V. Bochkov
i8
Harpyrhynchoides
kakatoe (Fain, 1972)
Cacatua sp.
Cacatuidae Psittaciformes
European Zoo
(captive)
Fain 1972, 1994
“
Cacatua galerita
Cacatuidae Psittaciformes Australia Domrow 1991
Harpyrhynchoides
psittaculae (Fain, 1972)
Psittacula
alexandri
Psittacidae Psittaciformes
European Zoo
(captive)
Fain 1972, 1994
“
Psittacula
cyanocephala
Psittacidae Psittaciformes
European Zoo
(captive)
Fain 1972, 1994
Harpyrhynchoides
psittaci Fain, 1972
Psittacus erithacus
Psittacidae Psittaciformes
European Zoo
(captive)
Fain 1972, 1994
“
Poicephalus
senegalus
Psittacidae Psittaciformes
European Zoo
(captive)
Fain 1972, 1994
Harpyrhynchoides
amazonae (Fain, 1972)
Amazona aestiva
Psittacidae Psittaciformes
European Zoo
(captive)
Fain 1972, 1994
Harpyrhynchoides
lawrencei (Fain, 1972)
Nandayus nenday
Psittacidae Psittaciformes
European Zoo
(captive)
Fain 1972, 1994
Harpirhynchoides
rosellacinus (Lawrence,
1959)
Platycercus
eximius
Psittacidae Psittaciformes Australia Lawrence 1959c
“
Glossopsitta
concinna
Psittacidae Psittaciformes Australia Domrow 1991
“
Trichoglossus
chlorolepidotus
Psittacidae Psittaciformes Australia
Filippich and
Domrow 1985;
Domrow 1991
“
Trichoglossus
haematodus
moluccanus
Psittacidae Psittaciformes Australia Domrow 1991
“
Lathamus discolor
Psittacidae Psittaciformes Australia Fain 1994b
Harpyrhynchoides
metropeliae (Fain, 1972)
Metriopelia
ceciliae
Columbidae Columbiformes
European Zoo
(captive)
Fain 1972, 1994
Harpyrhynchoides
modestus (Fain, 1976)
Metriopelia
ceciliae
Columbidae Columbiformes
European Zoo
(captive)
Fain 1972, 1994
Harpyrhynchoides
capitatus (Fain, 1976)
Columbina
talpacoti
Columbidae Columbiformes
European Zoo
(captive)
Fain 1972, 1994
Harpyrhynchoides
coxatus Fain, 1972
Columbina
talpacoti
Columbidae Columbiformes
European Zoo
(captive)
Fain 1972, 1994
“
Columbina
cruziana
Columbidae Columbiformes
European Zoo
(captive)
Fain 1972, 1994
“
Columbina
squammata
Columbidae Columbiformes
European Zoo
(captive)
Fain 1972, 1994
Harpyrhynchoides
columbae (Fain, 1972)
Columba livia
Columbidae Columbiformes Europe Fain 1972, 1994
“
Columba palumbus
Columbidae Columbiformes Europe Fain 1972, 1994
Harpyrhynchoides oenae
(Fain, 1972)
Oena capensis
Columbidae Columbiformes
European Zoo
(captive)
Fain 1972, 1994
Harpyrhynchoides oenae
lamorali (Fain, 1972)
Turtur tympanistria
Columbidae Columbiformes
European Zoo
(captive)
Fain 1972, 1994
Harpyrhynchoides
coturnix Fain, 1972
Coturnix
delegorguei
Phasianidae Galliformes
European Zoo
(captive)
Fain 1972, 1994
Harpyrhynchoides
alectoris (Fain, 1972)
Alectoris graeca
Phasianidae Galliformes
European Zoo
(captive)
Fain 1972, 1994
“
Alectoris rufa
Phasianidae Galliformes Europe Fain et al. 1999
Harpyrhynchoides
numidae Lawrence, 1959
Numida meleagris
Numididae Galliformes Africa Lawrence 1959b
Harpyrhynchoides
capellae (Fritsch, 1954)
Gallinago
gallinago
Scolopacidae
Charadrii-
formes
Europe Fritsch 1954
Appendix
i9
Harpyrhynchoides
capellae (Fritsch, 1954)
Lymnocryptes
minimus
Scolopacidae
Charadrii-
formes
Europe Fain et al. 1999
Harpyrhynchoides
herodius (Boyd, 1968)
Ardea herodias
Ardeidae Pelecaniformes North America Boyd 1968
“
Ardea cinerea
Ardeidae Pelecaniformes Europe Fain 1994b
“
Butorides virescens
Ardeidae Pelecaniformes North America Boyd 1968
Harpyrhynchoides
leptoptilus (Fain, 1976)
Leptoptilos
crumeniferus
Ciconiidae Ciconiiformes
European Zoo
(captive)
Fain 1972, 1994
Harpyrhynchoides
tracheatus (Fritsch,
1954)
Buteo buteo
Accipitridae Accipitriformes Europe
Fritsch 1954;
Fain 1994b
Harpyrhynchoides tyto
(Fain, 1972)
Tyto alba
Tytonidae Strigiformes Europe Fain 1972, 1994
Harpyrhynchoides asio
(Fain, 1972)
Asio otus
Strigidae Strigiformes Europe Fain 1972, 1994
Harpyrhynchoides
anatum (Fain, 1976)
Anas querquedula
Anatidae Anseriformes
European Zoo
(captive)
Fain 1972, 1994
Harpyrhynchoides
cristagalli (Berlese et
Trouessart, 1889)
Colius striatus
Coliidae Coliiformes Africa
Berlese and
Trouessart 1889;
Lawrence 1959c
“
Colius colius
Coliidae Coliiformes Africa Fain 1994b
Harpyrhynchoides
zumpti (Fain, 1972)
Eremopterix
leucotis
Alaudidae Passeriformes Africa Fain 1972, 1994
“
Junco hyemalis
Emberizidae Passeriformes North America
Bochkov and
Galloway 2004
“
Diuca diuca
Emberizidae Passeriformes
European Zoo
(captive)
Fain 1972, 1994
“
Quiscalus quiscula
Icteridae Passeriformes North America
Bochkov and
Galloway 2001
Harpyrhynchoides
alaudinus Bochkov,
2000
Alauda arvensis
Alaudidae Passeriformes Europe
Megnin 1877;
Oudemans 1939;
Bochkov 2000a
Harpyrhynchoides
rubeculinus (Cerny et
Sixl, 1971)
Erithacus rubecula
Muscicapidae Passeriformes Europe
Cerny and Sixl
1971; Fain et al.
1999; Bochkov
and Literak 2008
Harpyrhynchoides
kirgizorum Fain,
Bochkov et Mironov,
1999
Rhodospiza
obsoleta
Fringillidae Passeriformes Asia Fain et al. 1999
Harpyrhynchoides
vulgaris Bochkov et Gal-
loway, 2004
Carpodacus
purpureus
Fringillidae Passeriformes North America
Bochkov and
Galloway 2004
“
Setophaga ruticilla
Parulidae Passeriformes North America
Bochkov and
Galloway 2004
Harpyrhynchoides
parazumpti Fain,
Bochkov et Mironov,
1999
Corvus monedula
Corvidae Passeriformes Europe Fain et al. 1999
Harpyrhynchoides
pectinifer (Lawrence,
1959)
Campethera
abingoni
Picidae Piciformes Africa Lawrence 1959a
“
Campethera
cailliauti
Picidae Piciformes Africa Fain 1994b
Harpyrhynchoides
vercammeni (Lawrence,
1959)
Centropus grillii
Cuculidae Cuculiformes Africa Lawrence 1959a
M. Skoracki, S. Zabludovskaya and A.V. Bochkov
i10
Harpyrhynchoides (Pseudoharpirhynchus) Fain, Bochkov et Mironov, 1999
Harpirhynchoides
agapornis (Fain, 1972)
Agapornis
roseicollis
Psittacidae Psittaciformes
European Zoo
(captive)
Fain 1972, 1994
“
Agapornis
pullarius
Psittacidae Psittaciformes
European Zoo
(captive)
Fain 1994b
“
Agapornis
nigrigenis
Psittacidae Psittaciformes
European Zoo
(captive)
Fain 1994b
Harpyrhynchoides
cylindripalpus (Fritsch,
1954)
Fringilla coelebs
Fringillidae Passeriformes Europe Fritsch 1954
“
Passerina ciris
Cardinalidae Passeriformes
European Zoo
(captive)
Fain 1994b
Harpirhynchoides sp. Pufnus gravis
Procellariidae
Procellarii-
formes
North America Foster et al. 1996
Neharpyrhynchus Fain, 1972
Neharpyrhynchus
plumaris (Fritsch, 1954)
Fringilla coelebs
Fringillidae Passeriformes Europe
Fritsch 1954;
Martinu et al.
2008; Bochkov
2000b
“
Muscicapa striata
Muscicapidae Passeriformes Europe Fritsch 1954
Neharpyrhynchus baile
Bochkov, Literak et
Capek, 2007
Turdus leucomelas
Turdidae Passeriformes South America
Bochkov et al.
2007
Neharpyrhynchus turdi
Martinu, Dusbabek et
Literak, 2008
Turdus merula
Turdidae Passeriformes Europe
Martinu et al.
2008
Neharpyrhynchus
chlorospingus Bochkov
et Literak, 2011
Chlorospingus
pileatus
Emberizidae Passeriformes South America
Bochkov and
Literak 2011
Neharpyrhynchus
hippolae Bochkov, 2000
Hippolais icterina
Acrocephalidae Passeriformes Europe Bochkov 2000b
Neharpyrhynchus
mironovi Bochkov et
Literak, 2011
Dacnis cayana
Thraupidae Passeriformes South America
Bochkov and
Literak 2011
Neharpyrhynchus
tangara Bochkov et
Literak, 2011
Tangara cayana
Thraupidae Passeriformes South America
Bochkov and
Literak 2011
“
Thraupis episcopus
Thraupidae Passeriformes South America
Literak et al.
2012
Neharpyrhynchus
novoplumaris (Moss,
Oliver et Nelson, 1968)
Certhia familiaris
Certhiidae Passeriformes North America Moss et al. 1968
“
Cardinalis
cardinalis
Cardinalidae Passeriformes North America Moss et al. 1968
“
Campylorhynchus
brunneicapillus
Troglodytidae Passeriformes North America Moss 1979
“
Spizella passerina
Emberizidae Passeriformes North America Moss 1979
“
Amphispiza
bilineata
Emberizidae Passeriformes North America Moss 1979
“
Melozone fusca
Emberizidae Passeriformes North America Moss 1979
Neharpyrhynchus pari
Martinu, Dusbabek et
Literak, 2008
Parus major
Paridae Passeriformes Europe
Martinu et al.
2008
“
Cyanistes
caeruleus
Paridae Passeriformes Europe
Martinu et al.
2008
Appendix
i11
Neharpyrhynchus pari
Martinu, Dusbabek et
Literak, 2008
Poecile montanus
Paridae Passeriformes Europe
Martinu et al.
2008
“
Poecile palustris
Paridae Passeriformes Europe
Moss 1979;
Martinu et al.
2008
“
Baeolophus bicolor
Paridae Passeriformes North America Moss 1979
Neharpyrhynchus
pilirostris (Berlese et
Trouessart, 1889)
Passer domesticus
Passeridae Passeriformes
Europe, Africa,
North America
Berlese and
Trouessart 1889;
Fritsch 1954;
Lawrence 1959a;
Fain 1995;
Martinu et al.
2008
“
Aegithalos
caudatus
Aegithalidae Passeriformes
Unknown
locality
Moss 1979
Neharpyrhynchus
schoenobaenus Martinu,
Dusbabek et Literak,
2008
Acrocephalus
schoenobaenus
Acrocephalidae Passeriformes Europe
Martinu et al.
2008
Neharpyrhynchus spinus
Martinu, Dusbabek et
Literak, 2008
Spinus spinus
Fringillidae Passeriformes Europe
Martinu et al.
2008
“
Carduelis
cannabina
Fringillidae Passeriformes Europe Fritsch 1954
Neharpyrhynchus
trochilinus (Fain, 1972)
Hummingbird Trochilidae Apodiformes South America Fain 1972, 1995
“
Chrysolampis
mosquitus
Trochilidae Apodiformes South America Fain 1995
“
Panterpe insignis
Trochilidae Apodiformes South America
Bochkov and
Literak 2011
“
Eugenes fulgens
Trochilidae Apodiformes South America
Bochkov and
Literak 2011
“
Amazilia lactea
Trochilidae Apodiformes South America
Bochkov and
Literak 2011;
Literak et al.
2012
“
Amazilia
chionogaster
Trochilidae Apodiformes South America
Literak et al.
2012
Metharpyrhynchus Fain, 1972
Metharpyrhynchus
macrophallus Fain, 1972
Lonchura
oryzivora
Estrildidae Passeriformes
European Zoo
(captive)
Fain 1972, 1995
Metharpyrhynchus
macrophallus Fain, 1972
Lonchura
oryzivora
Estrildidae Passeriformes
European Zoo
(captive)
Fain 1972, 1995
Metharpyrhyncus mossi
Fain, 1995
Prinia subava
Cisticolidae Passeriformes Africa Fain 1995
Metharpyrhynchus
namibiensis Fain, 1995
Philetairus socius
Ploceidae Passeriformes Africa Fain 1995
Metharpyrhynchoides
jynx Fain, 1972
Jynx rucollis
Picidae Piciformes Africa Fain 1995
Perharpyrhynchus Fain, 1972
Perharpyrhynchus
jacana Fain, 1972
Jacana spinosa
intermedia
Jacanidae
Charadrii-
formes
European Zoo
(captive)
Fain 1972, 1995
Perharpyrhynchus
recurvirostra Fain, 1976
Recurvirostra
avosetta
Recurvirostridae
Charadrii-
formes
European Zoo
(captive)
Fain 1976, 1995
Ralliharpirhynchus Fain, 1972
M. Skoracki, S. Zabludovskaya and A.V. Bochkov
i12
Ralliharpirhynchus
porphyrio (Fain, 1972)
Porphyrio
porphyrio
Rallidae Gruiformes
European Zoo
(captive)
Fain 1972, 1995
Ralliharpirhynchus
limnocorax (Fain, 1972)
Amaurornis
avirostra
Rallidae Gruiformes Africa Fain 1972, 1995
Harpyrhynchiella Fain, 1972
Harpyrhynchiella
reductus (Fritsch, 1954)
Apus apus
Apodidae Apodiformes Europe
Fritsch 1954;
Fain 1972, 1995
Cypsharpirhynchus Fain, 1995
Cypsharpirhynchus
cypsiuri (Fain, 1972)
Cypsiurus parvus
Apodidae Apodiformes Africa Fain 1972, 1995
“
Apus afnis
Apodidae Apodiformes Africa Fain 1972, 1995
Anharpyrhynchus Fain, 1972
Anharpyrhynchus
monstrosus (Fritsch,
1954)
Garrulus
glandarius
Corvidae Passeriformes Europe
Fritsch 1954;
Fain 1972, 1995
“
Corvus monedula
Corvidae Passeriformes
Unknown
locality
Moss 1979
“
Cyanocitta cristata
Corvidae Passeriformes
Unknown
locality
Moss 1979
“
Cyanocitta stelleri
Corvidae Passeriformes
Unknown
locality
Moss 1979
“
Phylidonyris
novaehollandiae
Meliphagidae Passeriformes
Unknown
locality
Moss 1979
“
Cissomela
pectoralis
Meliphagidae Passeriformes
Unknown
locality
Moss 1979
“
Manorina avigula
Meliphagidae Passeriformes
Unknown
locality
Moss 1979
“
Lichmera
indistincta
Meliphagidae Passeriformes
Unknown
locality
Moss 1979
“
Colaptes auratus
Picidae Piciformes
Unknown
locality
Moss 1979
Trichorhynchiella Fain, 1995
Trichorhynchiella
paddae Fain, 1995
Lonchura
oryzivora
Estrildidae Passeriformes
European Zoo
(captive)
Fain 1995
Harpypalpinae Fain, 1972
Harpypalpus Dubinin, 1957
Harpypalpus holopus
(Berlese et Trouessart,
1889)
Passer domesticus
Passeridae Passeriformes Europe
Berlese and
Trouessart 1889;
Berlese 1894;
Fain 1972; Fain
1999
“
Luscinia svecica
Muscicapidae Passeriformes
Unknown
locality
Moss 1979
Harpypalpus longipis
(Fritsch, 1954)
Troglodytis
troglodytis
Troglodytidae Passeriformes Europe Fritsch 1954
“
Parus major
Paridae Passeriformes Europe
Moss 1979;
Skoracki et al.
2004
“
Cyanistes
caeruleus
Paridae Passeriformes
Unknown
locality
Moss 1979
“
Spinus spinus
Fringillidae Passeriformes
Unknown
locality
Moss 1979
“
Pyrrhula pyrrhula
Fringillidae Passeriformes
Unknown
locality
Moss 1979
“
Nucifraga
caryocatactes
Corvidae Passeriformes
Unknown
locality
Moss 1979
Appendix
i13
Harpypalpus serini Fain,
1972
Serinus
mozambicus
Fringillidae Passeriformes
European Zoo
(captive)
Fain 1972; Fain
et al. 1999
Harpypalpus dubinini
Fain, 1972
Ploceus
intermedius
Ploceidae Passeriformes
European Zoo
(captive)
Fain 1972; Fain
et al. 1999
Harpypalpus tiarae Fain,
Bochkov et Mironov,
1999
Tiaris canorus
Emberizidae Passeriformes
European Zoo
(captive)
Fain et al. 1999
Harpypalpus spermestes
Fain, Bochkov et
Mironov, 1999
Spermestes
cucullatus
Estrildidae Passeriformes Africa Fain et al. 1999
Harpypalpoides Lombert et Moss, 1983
Harpypalpoides
lukoschusi Lombert et
Moss, 1983
Turdus merula
Turdidae Passeriformes Europe
Lombert and
Moss 1983;
Skoracki et al.
2004
Harpypalpoides lesickii
Skoracki, Bochkov et
Sikora, 2004
Coccothraustes
coccothraustes
Fringillidae Passeriformes Europe
Skoracki et al.
2004
Harpypalpoides
hirundinus (Fain, 1972)
Psalidoprocne sp.
Hirundinidae Passeriformes Africa
Fain 1972; Fain
et al. 1999
Harpypalpoides
namibiensis Fain,
Bochkov et Mironov,
1999
Emberiza
impetuani
Emberizidae Passeriformes Africa Fain 1999
Species inquirenda
Harpirhynchus brevis
Ewing, 1911
Coccothraustes
vespertinus
Fringillidae Passeriformes North America Ewing 1911
Harpirhynchus longipi-
lus Banks, 1905
Loxia sp.
Fringillidae Passeriformes North America Banks 1905
Cloacaridae Camin, Moss, Oliver et Singer, 1967
Pneumophaginae Fain et Smiley, 1989
Pneumophagus Fain et Smiley, 1989
Pneumophagus bubonis
Fain et Smiley, 1989
Bubo virginianus
Strigidae Strigiformes North America
Fain and Smiley
1989
Syringophilidae Lavoipierre, 1953
Syringophilinae Lavoipierre, 1953
Apodisyringiana Skoracki, 2005
Apodisyringiana
haszprunari Skoracki,
2005
Hemiprocne
comata
Hemiprocnidae Apodiformes Asia Skoracki 2005a
Apodisyringiana
mystaceae Skoracki,
2005
Hemiprocne
mystacea
Hemiprocnidae Apodiformes Australia Skoracki 2005a
Apodisyringophilus Skoracki et OConnor, 2010
Apodisyringophilus
collocalius Skoracki et
OConnor, 2010
Collocalia
esculenta
Apodidae Apodiformes Asia
Skoracki and
OConnor 2010
Ascetomylla Kethley, 1970
Ascetomylla gallinula
Kethley, 1970
Gallinula
chloropus
Rallidae Gruiformes Africa Kethley 1970
Ascetomylla porzanae
(Bochkov et Galloway,
2004)
Porzana carolina
Rallidae Gruiformes North America
Bochkov and
Galloway 2004
Aulobia Kethley, 1970
M. Skoracki, S. Zabludovskaya and A.V. Bochkov
i14
Aulobia anthreptes
Skoracki et Glowska,
2008
Anthreptes
malacensis
Nectariniidae Passeriformes Asia
Skoracki and
Glowska 2008b
Aulobia cardueli
Skoracki, Hendricks et
Spicer, 2010
Carduelis psaltria
Fringillidae Passeriformes North America
Skoracki et al.
2010a
“
Carduelis carduelis
Fringillidae Passeriformes Europe Skoracki 2011
“
Carduelis spinus
Fringillidae Passeriformes Europe, Asia
Skoracki and
Bochkov 2010
“
Carduelis amea
Fringillidae Passeriformes Europe Skoracki 2011
“
Carduelis citrinella
Fringillidae Passeriformes Europe Skoracki 2011
“
Carduelis
avirostris
Fringillidae Passeriformes Europe Skoracki 2011
Aulobia cisticolae
Skoracki et Sikora, 2003
Cisticola erythrops
Cisticolidae Passeriformes Africa
Skoracki and
Sikora 2003
“
Cisticola cantans
Cisticolidae Passeriformes Africa
Skoracki and
Sikora 2003
“
Cisticola lateralis
Cisticolidae Passeriformes Africa
Skoracki and
Sikora 2003
“
Cisticola chubbi
Cisticolidae Passeriformes Africa
Skoracki et al.
2009a
Aulobia dendroicae
Kethley, 1970
Setophaga
coronata
Parulidae Passeriformes North America Kethley 1970
“
Setophaga
nigrescens
Parulidae Passeriformes North America
Skoracki et al.
2010a
Aulobia erythroptera
Skoracki et Dabert, 2001
Prinia erythroptera
Cisticolidae Passeriformes Africa
Skoracki and
Dabert 2001a
Aulobia leucostictus
Skoracki, 2011
Leucosticte arctoa
Fringillidae Passeriformes Europe Skoracki 2011
Aulobia nectariniae
Skoracki et Glowska,
2008
Cinnyris jugularis
Nectariniidae Passeriformes Australia
Skoracki and
Glowska 2008b
“
Leptocoma sericea
Nectariniidae Passeriformes Australia
Skoracki and
Glowska 2008b
Aulobia stachyris
(Bochkov, Mironov et
Skoracki, 2001)
Stachyridopsis
pyrrhops
Timaliidae Passeriformes Asia
Bochkov et al.
2001
Aulobia sylviae Bochkov
et Mironov, 1998
Sylvia curruca
Sylviidae Passeriformes Europe, Asia
Bochkov and
Mironov 1998;
Glowska et al.
2007; Skoracki
and Bochkov
2010
“
Sylvia atricapilla
Sylviidae Passeriformes Europe
Bochkov and
Mironov 1998
“
Sylvia nisoria
Sylviidae Passeriformes Europe Skoracki 2011
Aulobia sylviettae (Fain,
Bochkov et Mironov,
2000)
Sylvietta whytti
Macrosphenidae Passeriformes Africa Fain et al. 2000
Aulobia virens Skoracki
et Dabert, 2001
Sylvietta virens
aviventris
Macrosphenidae Passeriformes Africa
Skoracki and
Dabert 2001a
Aulonastus Kethley, 1970
Aulonastus albus
Skoracki, 2002
Motacilla alba
Motacillidae Passeriformes Europe Skoracki 2002a
Aulonastus anthus
Skoracki, 2011
Anthus trivialis
Motacillidae Passeriformes Europe Skoracki 2011
Appendix
i15
Aulonastus anthus
Skoracki, 2011
Anthus cervinus
Motacillidae Passeriformes Europe Skoracki 2011
Aulonastus emberizicus
Skoracki, Hendricks et
Spicer, 2010
Ammodramus
savannarum
Emberizidae Passeriformes North America
Skoracki et al.
2010b
“
Zonotrichia
atricapilla
Emberizidae Passeriformes North America
Skoracki et al.
2010b
“
Passerculus
sandwichensis
Emberizidae Passeriformes North America
Skoracki et al.
2010b
Aulonastus euphagus
Skoracki, Hendricks et
Spicer, 2010
Euphagus
cyanocephalus
Icteridae Passeriformes North America
Skoracki et al.
2010b
Aulonastus fringillus
Skoracki, 2011
Fringilla coelebs
Fringillidae Passeriformes Europe Skoracki 2011
Aulonastus henicorhina
Sikora et Skoracki, 2012
Henicorhina
leucosticta
Troglodytidae Passeriformes South America Sikora et al. 2012
Aulonastus lanius
Skoracki, 2011
Lanius senator
Laniidae Passeriformes Europe Skoracki 2011
Aulonastus loxius
Skoracki, 2011
Loxia curvirostra
Fringillidae Passeriformes Europe Skoracki 2011
Aulonastus lusciniae
Skoracki, 2002
Luscinia luscinia
Muscicapidae Passeriformes Europe Skoracki 2002a
Aulonastus pipili
Kethley, 1970
Pipilo
erythrophthalmus
Emberizidae Passeriformes North America Kethley 1970
Aulonastus pirangus
Skoracki, Hendricks et
Spicer, 2010
Piranga
ludoviciana
Cardinalidae Passeriformes North America
Skoracki et al.
2010b
Aulonastus prunellae
Bochkov et Mironov,
1999
Prunella modularis
Prunellidae Passeriformes Europe
Bochkov and
Mironov 1999
Aulonastus sturnellus
Skoracki, Hendricks et
Spicer, 2010
Sturnella magna
Icteridae Passeriformes North America
Skoracki et al.
2010b
Betasyringophiloidus Skoracki, 2011
Betasyringophiloidus
motacillae (Bochkov et
Mironov, 1998)
Motacilla ava
Motacillidae Passeriformes Europe
Bochkov and
Mironov 1998
Betasyringophiloidus
phoenicurus Skoracki,
2011
Phoenicurus
ochruros
Muscicapidae Passeriformes Europe Skoracki 2011
Betasyringophiloidus
saxicolus Skoracki, 2011
Saxicola rubetra
Muscicapidae Passeriformes Europe Skoracki 2011
Betasyringophiloidus
schoeniclus (Skoracki,
2002)
Emberiza
schoeniclus
Emberizidae Passeriformes Europe, Asia
Skoracki 2002b,
2004a, 2011;
Skoracki and
Bochkov 2010
Blaszakia Skoracki et Sikora, 2008
Blaszakia tauracos
Skoracki et Sikora, 2008
Tauraco livingstoni
Musophagidae Cuculiformes Africa
Skoracki and
Sikora 2008
“
Tauraco schalowi
Musophagidae Cuculiformes Africa
Skoracki and
Sikora 2008
Blaszakia rossae
Skoracki et Sikora, 2008
Musophaga rossae
Musophagidae Cuculiformes Africa
Skoracki and
Sikora 2008
Bochkovia Skoracki et OConnor, 2010
Bochkovia phalaropi
Skoracki et OConnor,
2010
Phalaropus
fulicarius
Scolopacidae
Charadrii-
formes
North America
Skoracki and
OConnor 2010
M. Skoracki, S. Zabludovskaya and A.V. Bochkov
i16
Bubophilus Philips et Norton, 1978
Bubophilus ascalaphus
Philips et Norton, 1978
Bubo virginianus
Strigidae Strigiformes North America
Philips and
Norton 1978
“
Bubo africanus
Strigidae Strigiformes Africa
Skoracki and
Dabert 2002
“
Tyto alba afnis
Tytonidae Strigiformes Africa
Skoracki and
Dabert 2002
Bubophilus asiobius
Skoracki et Bochkov,
2002
Asio otus
Strigidae Strigiformes Europe
Skoracki and
Bochkov 2002
Bubophilus aluconis
Nattress et Skoracki,
2009
Strix aluco
Strigidae Strigiformes Europe
Nattress and
Skoracki 2009
“
Asio otus
Strigidae Strigiformes Europe
Skoracki et al.
2012a
Castosyringophilus Bochkov et Perez, 2002
Castosyringophilus
claravis Skoracki et
Glowska, 2008
Claravis pretiosa
Columbidae Columbiformes South America
Skoracki and
Glowska 2008a
Castosyringophilus forpi
Bochkov et Perez, 2002
Forpus
cyanopygius
Psittacidae Psittaciformes North America
Bochkov and
Perez 2002
Castosyringophilus
mucuya (Casto, 1980)
Columbina
passerina
Columbidae Columbiformes North America
Casto 1980a;
Bochkov and
Perez 2002
“
Columbina
squammata
Columbidae Columbiformes South America
Bochkov and
Fain 2003
“
Columbina
talpacoti
Columbidae Columbiformes South America
Skoracki and
Glowska 2008a
“
Geophaps
plumifera
Columbidae Columbiformes Australia
Bochkov and
Fain 2003
“
Metriopelia
melanoptera
Columbidae Columbiformes South America
Skoracki and
Glowska 2008a
“
Streptopelia
decaocto
Columbidae Columbiformes Asia
Glowska et al.
2007
“
Brotogeris
versicolurus
Psittacidae Psittaciformes South America
Bochkov and
Fain 2003
“
Trichoglossus
haematodus
Psittacidae Psittaciformes Asia
Bochkov and
Fain 2003
“
Psilopsiagon
aymara
Psittacidae Psittaciformes South America
Bochkov and
Fain 2003
Charadriphilus Bochkov et Chystiakov, 2001
Charadriphilus
gallinago (Bochkov et
Mironov, 1998)
Gallinago
gallinago
Scolopacidae
Charadrii-
formes
Europe
Bochkov and
Mironov, 1998;
Skoracki et al.
2006
Charadriphilus
lyudmilae Bochkov et
Chystiakov, 2001
Scolopax rusticola
Scolopacidae
Charadrii-
formes
Europe
Bochkov and
Chystiakov 2001
Charadriphilus paragua-
iae (Skoracki et Sikora,
2002)
Gallinago
paraguaiae
Scolopacidae
Charadrii-
formes
South America
Skoracki and
Sikora 2002
Charadriphilus ralli
Skoracki et Bochkov,
2010
Rallus aquaticus
Rallidae Gruiformes Asia
Skoracki and
Bochkov 2010
Charadriphilus vanelli
Bochkov, Fain et
Skoracki, 2004
Vanellus chilensis
Charadriidae
Charadrii-
formes
South America
Bochkov et al.
2004
Appendix
i17
Chenophila Kethley, 1970
Chenophila branta
Kethley, 1970
Branta canadensis
Anatidae Anseriformes North America Kethley 1970
Chenophila kanduli
Bochkov, 1998
Anas querquedula
Anatidae Anseriformes Europe
Bochkov and
Mironov 1998
Ciconichenophilus Skoracki et OConnor, 2010
Ciconichenophilus
phoeniconaias Skoracki
et OConnor, 2010
Phoenicopterus
minor
Phoenicopteri-
dae
Phoenicopteri-
formes
Africa
Skoracki and
OConnor 2010
Colinophilus Kethley, 1973
Colinophilus wilsoni
Kethley, 1973
Colinus virginianus
Odontophoridae Galliformes North America Kethley 1973
“
Callipepla
squamata
Odontophoridae Galliformes North America Casto 1976
Corvitorotroglus Skoracki et Bochkov, 2010
Corvitorotroglus alpha
Skoracki et Bochkov,
2010
Corvus frugilegus
Corvidae Passeriformes Asia
Skoracki and
Bochkov 2010
Creagonycha Kethley, 1970
Creagonycha lara
Kethley, 1970
Larus delawarensis
Laridae
Charadrii-
formes
North America Kethley 1970
Creagonycha sternae
Kivganov, 1995
Sterna albifrons
Laridae
Charadrii-
formes
Europe
Kivganov and
Sharafat 1995
Creagonycha totani
(Oudemans, 1904)
Tringa totanus
Scolopacidae
Charadrii-
formes
Europe
Oudemans
1904,1906;
“
Tringa glareola
Scolopacidae
Charadrii-
formes
Europe
Skoracki et al.
2006
“
Calidris temminckii
Scolopacidae
Charadrii-
formes
Africa, Europe
Skoracki and
Dabert 2002;
Skoracki et al.
2006
“
Calidris
himantopus
Scolopacidae
Charadrii-
formes
North America
Skoracki et al.
2006
“
Calidris minuta
Scolopacidae
Charadrii-
formes
Europe, Asia
Bochkov and
Mironov 1998;
Skoracki et al.
2006
Crotophagisyringophilus Skoracki, 2008
Crotophagisyringophilus
io Skoracki, 2008
Crotophaga ani
Cuculidae Cuculiformes South America Skoracki, 2008b
Cuculisyringophilus Skoracki, 2008
Cuculisyringophilus
crotophaginus Skoracki,
2008
Guira guira
Cuculidae Cuculiformes South America Skoracki 2008a
“
Crotophaga
sulcirostris
Cuculidae Cuculiformes
North and South
America
Skoracki 2008a
Cuculisyringophilus
chirovi (Bochkov et
Mironov, 1998)
Cuculus canorus
Cuculidae Cuculiformes Asia
Bochkov and
Mironov 1998
Fritschisyringophilus Bochkov, Fain et Skoracki, 2004
Fritschisyringophilus
lonchurae Bochkov, Fain
et Skoracki, 2004
Lonchura
punctulata
Estrildidae Passeriformes Asia
Bochkov et al.
2004
Galliphilopsis Skoracki et Sikora, 2004
Galliphilopsis bochkovi
Skoracki et Sikora, 2004
Tragopan sp.
Phasianidae Galliformes Asia
Skoracki and
Sikora 2004b
M. Skoracki, S. Zabludovskaya and A.V. Bochkov
i18
Galliphilopsis colinus
Skoracki et Sikora, 2011
Colinus cristatus
Odontophoridae Galliformes South America
Skoracki and
Sikora 2011
Galliphilopsis francoli-
nus Skoracki et Sikora,
2004
Francolinus
levaillantoides
Phasianidae Galliformes South Africa
Skoracki and
Sikora 2004b
Galliphilopsis lophurus
Skoracki et Sikora, 2004
Lophura
leucomelanos
Phasianidae Galliformes Asia
Skoracki and
Sikora 2004b
Galliphilopsis szeptyckii
Skoracki et Sikora, 2011
Coturnix pectoralis
Phasianidae Galliformes Australia
Skoracki and
Sikora 2011
“
Coturnix coturnix
Phasianidae Galliformes Asia
Skoracki and
Sikora 2011
“
Coturnix chinensis
Phasianidae Galliformes Asia
Skoracki and
Sikora 2011
“
Alectoris barbara
Phasianidae Galliformes Europe
Skoracki and
Sikora 2011
Kalamotrypetes Casto, 1980
Kalamotrypetes
colinastes Casto, 1980
Colinus virginianus
Odontophoridae Galliformes North America Casto 1980b
Kalamotrypetes cracidus
Skoracki et Sikora, 2011
Penelope sp.
Cracidae Galliformes South America
Skoracki and
Sikora 2011
Kethleyana Kivganov, 1995
Kethleyana gelocheli-
doni Kivganov, 1995
Gelochelidon
nilotica
Laridae
Charadrii-
formes
Europe
Kivganov and
Sharafat 1995
Krantziaulonastus Skoracki, 2011
Krantziaulonastus
buczekae (Skoracki,
2002)
Sturnus vulgaris
Sturnidae Passeriformes Europe
Skoracki 2002a,
2011
Krantziaulonastus
galbulicus (Skoracki,
2008)
Jacamaralcyon
tridactyla
Galbulidae Galbuliformes South America Skoracki 2008c
Krantziaulonastus
oryzivorus Skoracki,
2011
Lonchura
oryzivora
Estrildidae Passeriformes Asia Skoracki 2011
Krantziaulonastus
lonchurus Skoracki,
2011
Lonchura
punctulata
Estrildidae Passeriformes Asia Skoracki 2011
Krantziaulonastus
yoyomi Glowska et
Skoracki, 2011
Dinemellia
dinemelli
Ploceidae Passeriformes Africa
Glowska and
Skoracki 2011b
Megasyringophilus Fain, Bochkov et Mironov, 2000
Megasyringophilus
aquilus Skoracki,
Lontkowski et Stawarc-
zyk, 2010
Aquila rapax
Accipitridae Accipitriformes Africa
Skoracki et al.
2010d
“
Aquila pomarina
Accipitridae Accipitriformes Europe
Skoracki et al.
2010d
“
Accipiter nisus
Accipitridae Accipitriformes Asia
Skoracki and
Bochkov 2010
Megasyringophilus
cyanocephala Fain,
Bochkov et Mironov,
2000
Psittacula
cyanocephala
Psittacidae Psittaciformes Asia Fain et al. 2000
“
Psittacula eupatria
Psittacidae Psittaciformes Asia
Bochkov and
Fain 2003
“
Psittacula krameri
Psittacidae Psittaciformes Asia
Bochkov and
Fain 2003
Appendix
i19
Megasyringophilus
dubinini Bochkov et
Fain, 2003
Trichoglossus
ornatus
Psittacidae Psittaciformes Asia
Bochkov and
Fain 2003
Megasyringophilus eos
Skoracki, 2005
Eos bornea
Psittacidae Psittaciformes Asia Skoracki 2005c
Megasyringophilus
geoffroyus Skoracki,
2005
Geoffroyus
geoffroyi
Psittacidae Psittaciformes Australia Skoracki 2005c
Megasyringophilus
kethleyi Fain, Bochkov
et Mironov, 2000
Aratinga jandaya
Psittacidae Psittaciformes South America Fain et al. 2000
“
Aratinga pertinax
Psittacidae Psittaciformes South America
Bochkov and
Fain 2003
“
Brotogeris
versicolurus
Psittacidae Psittaciformes South America
Bochkov and
Fain 2003
Megasyringophilus
platycercus Bochkov et
Fain, 2003
Platycercus
eximius
Psittacidae Psittaciformes Australia
Bochkov and
Fain 2003
Megasyringophilus
rhynchopsittae Bochkov
et Perez, 2002
Rhynchopsitta
pachyrhyncha
Psittacidae Psittaciformes North America
Bochkov and
Perez, 2002
Megasyringophilus
trichoglossus Fain,
Bochkov et Mironov,
2000
Trichoglossus sp.
Psittacidae Psittaciformes South America Fain et al. 2000
“
Trichoglossus
euteles
Psittacidae Psittaciformes Asia Skoracki 2005c
“
Trichoglossus
chlorolepidotus
Psittacidae Psittaciformes Australia Skoracki 2005c
Meitingsunes Glowska et Skoracki, 2010
Meitingsunes aldwelles
Glowska et Skoracki,
2010
Geotrygon frenata
Columbidae Columbiformes South America
Glowska and
Skoracki 2010
Meitingsunes columbicus
Skoracki, 2011
Columba oenas
Columbidae Columbiformes Asia Skoracki 2011
“
Columba livia
Columbidae Columbiformes Europe
“
Columba palumbus
Columbidae Columbiformes Europe
Meitingsunes zenadou-
rae (Clark, 1964)
Zenaida macroura
Columbidae Columbiformes North America Clark 1964b
“
Zenaida asiatica
Columbidae Columbiformes North America Casto 1976
“
Zenaida auriculata
Columbidae Columbiformes South America
Skoracki and
Sikora 2002
“
Columba livia
Columbidae Columbiformes
North America,
Africa
Casto 1976
Bochkov and
Mironov 1998
Meitingsunes tympanis-
tria Skoracki et Dabert,
2002
Turtur tympanistria
Columbidae Columbiformes Africa
Skoracki and
Dabert 2002
Mironovia Chirov et Kravtsova, 1995
Mironovia phasiani
Chirov et Kravtsova,
1995
Phasianus
colchicus
Phasianidae Galliformes Asia
Chirov and
Kravtsova 1995
Mironovia coturnae
Bochkov, Fain et
Skoracki, 2004
Coturnix coturnix
Phasianidae Galliformes Europe
Bochkov et al.
2004
Mironovia rouloul
Skoracki et Sikora, 2004
Rollulus rouloul
Phasianidae Galliformes Asia
Skoracki and
Sikora 2004b
M. Skoracki, S. Zabludovskaya and A.V. Bochkov
i20
Mironovia lagopus
Bochkov et Skirnisson,
2011
Lagopus muta
Phasianidae Galliformes Europe
Bochkov and
Skirnisson 2011
“
Lagopus lagopus
Phasianidae Galliformes Europe
Skoracki and
Sikora 2011
Neoaulobia Fain, Bochkov et Mironov, 2000
Neoaulobia agapornis
Fain, Bochkov et
Mironov, 2000
Agapornis
nigrigenis
Psittacidae Psittaciformes Africa Fain et al. 2000
“
Agapornis scheri
Psittacidae Psittaciformes Africa
Bochkov and
Fain 2003
“
Agapornis
personatus
Psittacidae Psittaciformes Africa
Bochkov and
Fain 2003
“
Agapornis
roseicollis
Psittacidae Psittaciformes Africa
Bochkov and
Fain 2003
“
Agapornis taranta
Psittacidae Psittaciformes Africa
Bochkov and
Fain 2003
Neoaulobia aratingae
Fain, Bochkov et
Mironov, 2000
Aratinga jandaya
Psittacidae Psittaciformes South America Fain et al. 2000
Neoaulobia krafti
Skoracki, 2005
Cacatua
tenuirostris
Cacatuidae Psittaciformes Australia Skoracki 2005c
Neoaulobia mexicana
Bochkov et Perez, 2002
Aratinga
canicularis
Psittacidae Psittaciformes North America
Bochkov and
Perez 2002
“
Aratinga pertinax
Psittacidae Psittaciformes South America
Bochkov and
Fain 2003
Neoaulobia mironovi
Bochkov et Perez, 2002
Amazona nschi
Psittacidae Psittaciformes North America
Bochkov and
Perez 2002
Neoaulobia psittaculae
Fain, Bochkov et
Mironov, 2000
Psittacula
cyanocephala
Psittacidae Psittaciformes Asia Fain et al. 2000
Neoaulobia puylaerti
(Skoracki et Dabert,
1999)
Poicephalus
senegalus versteri
Psittacidae Psittaciformes Africa
Skoracki and
Dabert 1999b
“
Loriculus pusillus
Psittacidae Psittaciformes Asia Skoracki 2005c
“
Loriculus
philippens
Psittacidae Psittaciformes Asia Skoracki 2005c
Neoaulonastus Skoracki, 2004
Neoaulonastus aegitha-
los Skoracki, 2011
Aegithalos
caudatus
Aegithalidae Passeriformes Europe Skoracki 2011
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
scirpaceus
Acrocephalidae Passeriformes Europe Skoracki 1999a
“
Acrocephalus
paludicola
Acrocephalidae Passeriformes Europe Skoracki 2011
“
Acrocephalus
dumetorum
Acrocephalidae Passeriformes Asia Skoracki 2011
“
Acrocephalus
arundinaceus
Acrocephalidae Passeriformes Europe Skoracki 2011
“
Acrocephalus
palustris
Acrocephalidae Passeriformes Africa
Skoracki et al.
2011b
Neoaulonastus caligatus
Skoracki, 2011
Hippolais caligata
Acrocephalidae Passeriformes Europe Skoracki 2011
Appendix
i21
Neoaulonastus chrysoco-
laptes Skoracki et OCon-
nor, 2010
Chrysocolaptes
lucidus
Picidae Piciformes Asia
Skoracki and
OConnor 2010
Neoaulonastus picidus
Skoracki, 2011
Picus canus
Picidae Piciformes Europe Skoracki 2011
“
Dendrocopos
leucotos
Picidae Piciformes Europe Skoracki 2011
“
Dendrocopos
major
Picidae Piciformes Europe Skoracki 2011
Neoaulonastus remizus
Skoracki, 2011
Remiz pendulinus
Remizidae Passeriformes Europe Skoracki 2011
Neoaulonastus riparius
Skoracki, 2011
Riparia riparia
Hirundinidae Passeriformes Europe Skoracki 2011
Neoaulonastus zosterops
Skoracki, Antczak et
Riegert, 2009
Zosterops
senegalensis
Zosteropidae Passeriformes Africa
Skoracki et al.
2009a
Neoperisterophila Skoracki, 2005
Neoperisterophila
regiusi Skoracki, 2005
Cicinnurus regius
Paradisaeidae Passeriformes Australia Skoracki 2005b
Neosyringophilopsis Skoracki et Sikora, 2005
Neosyringophilopsis
acanthizus Skoracki et
Sikora, 2005
Acanthiza inornata
Acanthizidae Passeriformes Australia
Skoracki and
Sikora 2005
Neosyringophilopsis
aegithali (Bochkov,
Mironov et Skoracki,
2001)
Aegithalos
caudatus
Aegithalidae Passeriformes Europe
Bochkov et al.
2001; Skoracki
2004b; Nattress
and Skoracki
2009
Neosyringophilopsis
garrulaxi (Bochkov,
Mironov et Skoracki,
2001)
Garrulax afnis
Leiothrichidae Passeriformes Asia
Bochkov et al.
2001
Neosyringophilopsis
locustellus (Skoracki,
2004)
Locustella naevia
Locustellidae Passeriformes Europe Skoracki 2004b
“
Locustella
luscinioides
Locustellidae Passeriformes Europe Skoracki 2004b
Neosyringophilopsis
lonchurus Skoracki,
2008
Spermestes
cucullatus
Estrildidae Passeriformes Africa Skoracki 2008c
Neosyringophilopsis
phylloscopi (Bochkov,
Mironov et Skoracki,
2001)
Phylloscopus
collybita
Phylloscopidae Passeriformes Europe
Bochkov et al.
2001
“
Phylloscopus
trochilus
Phylloscopidae Passeriformes Europe Skoracki 2004b
Neosyringophilopsis
troglodytis (Fritsch,
1958)
Troglodytes
troglodytes
Troglodytidae Passeriformes Europe
Fritsch 1958;
Nattress and
Skoracki 2007
“
Troglodytes aedon
Troglodytidae Passeriformes North America
Bochkov and
Galloway 2001
Niglarobia Kethley, 1970
Niglarobia cursoriae
Skoracki, Dabert et
Schmaschke, 2006
Cursorius
temminckii
Glareolidae
Charadrii-
formes
Africa
Skoracki et al.
2006
Niglarobia ereuneti
Kethley, 1970
Calidris pusilla
Scolopacidae
Charadrii-
formes
North America Kethley 1970
M. Skoracki, S. Zabludovskaya and A.V. Bochkov
i22
Niglarobia helleri
(Oudemans, 1904)
Tringa ochropus
Scolopacidae
Charadrii-
formes
Europe
Oudemans 1904,
1906; Kethley
1970
Tringa avipes
Scolopacidae
Charadrii-
formes
South America
Skoracki and
Sikora 2002
Tringa stagnatilis
Scolopacidae
Charadrii-
formes
Europe
Skoracki et al.
2006
Arenaria interpres
Scolopacidae
Charadrii-
formes
North America
Skoracki et al.
2006
Niglarobia rhinoptili
Fain, Bochkov et
Mironov, 2000
Smutsornis
africanus
Glareolidae
Charadrii-
formes
Africa Fain et al. 2000
Niglarobia trouessarti
(Oudemans, 1904)
Aramus guarauna
Aramidae Gruiformes South America
Oudemans 1904,
1906; Kethley
1970
Paraniglarobia Skoracki, 2011
Paraniglarobia skorackii
(Bochkov et Galloway,
2004)
Tringa
melanoleuca
Scolopacidae
Charadrii-
formes
North America
Bochkov and
Galloway 2004
Paraniglarobia
calidridis (Bochkov et
Mironov, 1998)
Calidris minuta
Scolopacidae
Charadrii-
formes
Asia
Bochkov and
Mironov 1998
Peristerophila Kethley, 1970
Peristerophila accip-
itridicus Skoracki,
Lontkowski et Stawarc-
zyk, 2010
Accipiter nisus
Accipitridae Accipitriformes Asia
Skoracki and
Bochkov 2010
“
Terathopius
ecaudatus
Accipitridae Accipitriformes Africa
Skoracki et al.
2010d
Peristerophila columbae
(Hirst, 1920)
Columba livia
Columbidae Columbiformes
North America,
Europe, Asia
Kethley 1970
Bochkov and
Mironov 1998;
Nattress and
Skoracki 2009;
Skoracki 2011
“
Zenaida auriculata
Columbidae Columbiformes South America
Skoracki and
Sikora 2002
“
Buteo jamaicensis
Accipitridae Accipitriformes North America Casto 1976
Phalarophilus Skoracki, Bochkov et OConnor, 2011
Phalarophilus fulicarius
Skoracki, Bochkov et
OConnor, 2011
Phalaropus
fulicarius
Scolopacidae
Charadrii-
formes
North America
Skoracki et al.
2011c
Philoxanthornea Kethley, 1970
Philoxanthornea anoa
Kethley, 1970
Anous tenuirostris
Laridae
Charadrii-
formes
North America Kethley 1970
Philoxanthornea clarki
Kivganov, 1995
Sternula albifrons
Laridae
Charadrii-
formes
Europe
Kivganov and
Sharafat 1995
“
Sterna hirundo
Laridae
Charadrii-
formes
Europe
Skoracki et al.
2006
“
Larus canus
Laridae
Charadrii-
formes
Europe
Bochkov and
Mironov 1998
Philoxanthornea
dubinini Bochkov et
Mironov, 1998
Gelochelidon
nilotica
Laridae
Charadrii-
formes
Asia
Bochkov and
Mironov 1998
“
Chlidonias
leucopterus
Laridae
Charadrii-
formes
Africa
Skoracki et al.
2006
Appendix
i23
Picisyringophilus Skoracki et OConnor, 2010
Picisyringophilus kratos
Skoracki et OConnor,
2010
Picoides pubescens
Picidae Piciformes North America
Skoracki and
OConnor 2010
Procellariisyringophilus (Kethley, 1970)
Procellariisyringophilus
bulwerius (Kethley,
1970)
Bulweria bulwerii
Procellariidae
Procellarii-
formes
North America Kethley 1970
Psittaciphilus Fain, Bochkov et Mironov, 2000
Psittaciphilus amazonae
Fain, Bochkov et
Mironov, 2000
Amazona
amazonica
Psittacidae Psittaciformes South America Fain et al. 2000
“
Amazona aestiva
Psittacidae Psittaciformes South America
Bochkov and
Fain 2003
Psittaciphilus fritschi
Fain, Bochkov et
Mironov, 2000
unidentied parrot
Psittacidae Psittaciformes
unknown
locality
Fain et al. 2000
Pteroclidisyringophilus Skoracki, 2011
Pteroclidisyringophilus
re (Skoracki et OConnor,
2010)
Pterocles
senegallus
Pteroclidae Pterocliformes Africa
Skoracki and
OConnor 2010
Selenonycha Kethley, 1970
Selenonycha baltoda
Kethley, 1970
Charadrius
wilsonia
Charadriidae
Charadrii-
formes
North America Kethley 1970
Selenonycha chardriifor-
micus Skoracki, 2011
Larus genei
Laridae
Charadrii-
formes
Africa Skoracki 2011
“
Charadrius
placidus
Charadriidae
Charadrii-
formes
Asia Skoracki 2011
“
Chroicocephalus
ridibundus
Laridae
Charadrii-
formes
Europe Skoracki 2011
“
Philomachus
pugnax
Scolopacidae
Charadrii-
formes
Europe Skoracki 2011
Stibarokris Kethley, 1970
Stibarokris dastychi
Glowska et Skoracki,
2011
Phalacrocorax
carbo
Phalacrocoraci-
dae
Suliformes
unknown
locality
Glowska and
Skoracki 2011a
Stibarokris langei
Bochkov et Mironov,
1999
Ciconia ciconia
Ciconiidae Ciconiiformes Europe
Bochkov and
Mironov 1999;
Skoracki 2011
Stibarokris phalacrus
Kethley, 1970
Phalacrocorax
auritus
Phalacrocoraci-
dae
Suliformes North America Kethley 1970
Stibarokris phoenicona-
ias Skoracki et OConnor,
2010
Phoenicopterus
minor
Phoenicopteri-
dae
Phoenicopteri-
formes
Africa
Skoracki and
OConnor 2010
“
Phoenicopterus
ruber
Phoenicopteri-
dae
Phoenicopteri-
formes
European Zoo
(captive)
Glowska and
Skoracki 2011a
Syringophiloidus Kethley, 1970
Syringophiloidus
agelaius Bochkov,
Skoracki, Hendricks et
Spicer, 2011
Agelaius
phoeniceus
Icteridae Passeriformes North America
Bochkov et al.
2011
Syringophiloidus
artamus Skoracki, 2004
Artamus
leucorynchus
Artamidae Passeriformes Asia Skoracki 2004a
“
Artamus fuscus
Artamidae Passeriformes unknown Skoracki 2004a
Syringophiloidus
bombycillae Skoracki,
2002
Bombycilla
garrulus
Bombycillidae Passeriformes Europe
Skoracki 2002b;
Skoracki 2004a
M. Skoracki, S. Zabludovskaya and A.V. Bochkov
i24
Syringophiloidus carolae
Skoracki, Flannery et
Spicer, 2009
Cardinalis
cardinalis
Cardinalidae Passeriformes North America
Skoracki et al.
2009b
“
Melanerpes
formicivorus
Picidae Piciformes North America
Skoracki et al.
2009b
Syringophiloidus
carpodaci Bochkov et
Apanaskevich, 2001
Carpodacus
erythrinus
Fringillidae Passeriformes Europe, Asia
Bochkov and
Apanaskevich
2001; Skoracki
2011
Syringophiloidus
coccothraustes Skoracki,
2011
Coccothraustes
coccothraustes
Fringillidae Passeriformes Europe Skoracki 2011
Syringophiloidus
cypsiuri Fain, Bochkov
et Mironov, 2000
Cypsiurus parvus
Apodidae Apodiformes Africa Fain et al. 2000
Syringophiloidus daberti
Bochkov, Fain et
Skoracki, 2004
Passerina ciris
Cardinalidae Passeriformes North America
Bochkov et al.
2004
Syringophiloidus
delichonum Bochkov,
2001
Delichon urbicum
Hirundinidae Passeriformes Europe Bochkov 2001
Syringophiloidus
dendrocittae Fain,
Bochkov et Mironov,
2000
Dendrocitta
vagabunda
Corvidae Passeriformes Asia Fain et al. 2000
Syringophiloidus
erythrurus Skoracki,
2004
Erythrura trichroa
Estrildidae Passeriformes Europe Skoracki 2004a
Syringophiloidus
glandarii (Fritsch, 1958)
Garrulus
glandarius
Corviidae Passeriformes Europe
Fritsch 1958;
Bochkov 2001
“
Pica pica
Corviidae Passeriformes Asia Skoracki 2004a
“
Corvus monedula
Corviidae Passeriformes Asia
Skoracki and
Bochkov 2010
“
Corvus frugilegus
Corviidae Passeriformes Asia
Skoracki and
Bochkov 2010
Syringophiloidus
graculae Fain, Bochkov
et Mironov, 2000
Gracula religiosa
Sturnidae Passeriformes Asia Fain et al. 2000
Syringophiloidus
hirundinis Skoracki,
Moller et Tryjanowski,
2003
Hirundo rustica
Hirundinidae Passeriformes Europe
Skoracki et al.
2003
Syringophiloidus
jackowiaki Bochkov,
Skoracki, Hendricks et
Spicer, 2011
Poecile
carolinensis
Paridae Passeriformes North America
Bochkov et al.
2011
Syringophiloidus klimovi
Skoracki et Bochkov,
2010
Chloris chloris
Fringillidae Passeriformes Asia
Skoracki and
Bochkov 2010
Syringophiloidus
microcerculus Sikora et
Skoracki, 2012
Microcerculus
marginatus
Troglodytidae Passeriformes South America Sikora et al. 2012
Syringophiloidus minor
(Berlese, 1887)
Passer domesticus
Passeridae Passeriformes
North America,
Europe, Asia
Kethley 1970;
Skoracki 2011
“
Passer
hispaniolensis
Passeridae Passeriformes Asia
Glowska et al.
2007
“
Passer montanus
Passeridae Passeriformes Europe
Bochkov and
Mironov 1998
Appendix
i25
Syringophiloidus
molothrus Skoracki,
Flannery et Spicer, 2009
Molothrus ater
Icteridae Passeriformes North America
Skoracki et al.
2009b
Syringophiloidus
montanus Skoracki,
2002
Passer montanus
Passeridae Passeriformes Europe Skoracki 2002b
Syringophiloidus oriolus
Skoracki, 2004
Oriolus oriolus
Oriolidae Passeriformes Europe Skoracki 2004a
“
Oriolus chinensis
Oriolidae Passeriformes Asia Skoracki 2011
Syringophiloidus
parapresentalis
Skoracki, 2011
Turdus merula
Turdidae Passeriformes Europe, Asia 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,
2011
Petronia petronia
Passeridae Passeriformes Europe Skoracki 2011
Syringophiloidus
philomelosus Skoracki,
2011
Turdus philomelos
Turdidae Passeriformes Asia Skoracki 2011
Syringophiloidus
presentalis Chirov et
Kravtsova, 1995
Sturnus vulgaris
Sturnidae Passeriformes Europe, Asia
Chirov and
Kravtsova 1995;
Skoracki 2011
Syringophiloidus seiuri
(Clark, 1964)
Seiurus aurocapilla
Parulidae Passeriformes North America
Clark 1964b;
Bochkov and
Galloway 2001
“
Helmitheros
vermivorum
Parulidae Passeriformes North America Clark 1964b
“
Melospiza melodia
Emberizidae Passeriformes North America Clark 1964b
“
Quelea erythrops
Ploceidae Passeriformes Africa
Skoracki and
Dabert 2002
“
Ploceus baglafecht
Ploceidae Passeriformes Africa
Skoracki and
Dabert 2002
“
Ploceus
superciliosus
Ploceidae Passeriformes Africa
Skoracki and
Dabert 2002
“
Melospiza lincolnii
Emberizidae Passeriformes North America
Bochkov et al.
2011
“
Oreothlypis
rucapilla
Parulidae Passeriformes North America
Bochkov et al.
2011
Syringophiloidus serini
Bochkov, Fain et
Skoracki, 2004
Serinus
mozambicus
Fringillidae Passeriformes Africa
Bochkov et al.
2004
Syringophiloidus sialius
Skoracki, Flannery et
Spicer, 2009
Sialia mexicana
Turdidae Passeriformes North America
Skoracki et al.
2009b
“
Spizella breweri
Emberizidae Passeriformes North America
Bochkov et al.
2011
Syringophiloidus
stawarczyki Skoracki,
2004
Euphonia
cyanocephala
Fringillidae Passeriformes South America Skoracki 2004a
Syringophiloidus tarnii
Skoracki et Sikora, 2002
Pteroptochos tarnii
Rhinocryptidae Passeriformes South America
Skoracki and
Sikora 2002
Syringophiloidus
thryothorus Skoracki,
Flannery et Spicer, 2009
Thryothorus
ludovicianus
Troglodytidae Passeriformes North America
Skoracki et al.
2009b
M. Skoracki, S. Zabludovskaya and A.V. Bochkov
i26
Syringophiloidus weiszii
Skoracki, Hromada et
Tryjanowski, 2001
Lanius excubitor
Laniidae Passeriformes Europe
Skoracki et al.
2001b
Syringophiloidus
xanthocephalus
Bochkov, Skoracki,
Hendricks et Spicer,
2011
Xanthocephalus
xanthocephalus
Icteridae Passeriformes North America
Bochkov et al.
2011
Syringophiloidus
zonotrichia Bochkov,
Skoracki, Hendricks et
Spicer, 2011
Zonotrichia
albicollis
Emberizidae Passeriformes North America
Bochkov et al.
2011
Syringophilopsis Kethley, 1970
Syringophilopsis
acrocephali Skoracki,
1999
Acrocephalus
scirpaceus
Acrocephalidae Passeriformes Europe Skoracki 1999a
“
Acrocephalus
schoenobaenus
Acrocephalidae Passeriformes Europe, Africa Skoracki 2011
“
Acrocephalus
palustris
Acrocephalidae Passeriformes Europe Skoracki 2011
Syringophilopsis
albicollisi Skoracki et
Dabert, 2000
Merops albicollis
Meropidae Coraciiformes Africa
Skoracki and
Dabert 2000
Syringophilopsis
blaszaki Skoracki et
Dabert, 1999
Anthus trivialis
Motacillidae Passeriformes Europe
Skoracki and
Dabert 1999a
Bochkov and
Galloway 2001
Syringophilopsis borini
Bochkov et Mironov,
1999
Sylvia borin
Sylviidae Passeriformes Europe, Asia
Bochkov and
Mironov 1999;
Skoracki 2004b,
2011
Syringophilopsis
caligatus Skoracki, 2011
Hippolais caligata
Acrocephalidae Passeriformes Europe Skoracki 2011
Syringophilopsis
certhiae Skoracki,
Hendricks et Spicer,
2011
Certhia americana
Certhiidae Passeriformes North America
Skoracki et al.
2011a
Syringophilopsis
corvinae Skoracki et
Sikora, 2003
Corvinella corvina
Laniidae Passeriformes Africa
Skoracki and
Sikora 2003
Syringophilopsis
dendroicae Bochkov et
Galloway, 2001
Setophaga
coronata
Parulidae Passeriformes North America
Bochkov and
Galloway 2001
“
Setophaga graciae
Parulidae Passeriformes North America
Skoracki et al.
2008
“
Cardellina pusilla
Parulidae Passeriformes North America
Skoracki et al.
2008
Syringophilopsis dicruri
Skoracki, Hromada et
Wamiti, 2011
Dicrurus adsimilis
Dicruridae Passeriformes Africa
Skoracki et al.
2011b
Syringophilopsis
elongatus (Ewing, 1911)
Agelaius
phoeniceus
Icteridae Passeriformes North America Kethley 1970
“
Euphagus
carolinus
Icteridae Passeriformes North America Clark 1964b
“
Euphagus
cyanocephalus
Icteridae Passeriformes North America
Skoracki et al.
2008
“
Quiscalus
mexicanus
Icteridae Passeriformes North America Casto 1976
Appendix
i27
Syringophilopsis
elongatus (Ewing, 1911)
Quiscalus quiscula
Icteridae Passeriformes North America
Clark 1964b
Bochkov and
Galloway 2001
“
Pheucticus
melanocephalus
Cardinalidae Passeriformes North America Casto 1976
Syringophilopsis
emberizae Fain,
Bochkov et Mironov,
2000
Sicalis luteola
Emberizidae Passeriformes Africa Fain et al. 2000
Syringophilopsis
empidonax Skoracki,
Flannery et Spicer, 2008
Empidonax
hammondi
Tyrannidae Passeriformes North America
Skoracki et al.
2008
“
Empidonax
wrightii
Tyrannidae Passeriformes North America
Skoracki et al.
2008
Syringophilopsis
fringillae (Fritsch, 1958)
Fringilla coelebs
Fringillidae Passeriformes Europe, Asia
Fritsch 1958;
Bochkov and
Mironov 1998;
Skoracki 2004b;
Skoracki and
Bochkov 2010
Syringophilopsis
garrulus Skoracki et
Dabert, 2002
Garrulus
glandarius
Corviidae Passeriformes Africa Skoracki 2011
Syringophilopsis
hirundus Skoracki, 2004
Hirundo rustica
Hirundinidae Passeriformes Europe, Asia
Skoracki 2004b;
Skoracki and
Bochkov 2010
Syringophilopsis
hylocichlae (Clark,
1964)
Catharus
fuscescens
Turdidae Passeriformes North America Clark 1964b
Syringophilopsis icteri
Bochkov et Galloway,
2001
Icterus galbula
Icteridae Passeriformes North America
Bochkov and
Galloway 2001
“
Icterus bullocki
Icteridae Passeriformes North America
Skoracki et al.
2011a
Syringophilopsis
kazmierski Skoracki,
2004
Ficedula hypoleuca
Muscicapidae Passeriformes Europe
Skoracki 2004b;
2011
“
Ficedula parva
Muscicapidae Passeriformes Europe
Skoracki 2004b,
2011
“
Phoenicurus
phoenicurus
Muscicapidae Passeriformes Asia
Glowska et al.
2007
Syringophilopsis
kirgizorum Bochkov,
Mironov et Kravtsova,
2000
Chloris chloris
Fringillidae Passeriformes Europe, Asia
Bochkov et al.
2000; Skoracki
2004b; Nattress
and Skoracki
2007; Glowska
et al. 2007
“
Carduelis
cannabina
Fringillidae Passeriformes Asia
Glowska et al.
2007
“
Carduelis carduelis
Fringillidae Passeriformes Europe, Asia
Skoracki 2004b;
2011
“
Rhodospiza
obsoleta
Fringillidae Passeriformes Asia
Bochkov et al.
2000
“
Linurgus olivaceus
Fringillidae Passeriformes Africa
Skoracki et al.
2009a
Syringophilopsis kristini
Skoracki, Tryjanowski et
Hromada, 2002
Lanius minor
Laniidae Passeriformes Europe
Skoracki et al.
2002; Skoracki
2011
M. Skoracki, S. Zabludovskaya and A.V. Bochkov
i28
Syringophilopsis
lagonostictus Skoracki et
Dabert, 2002
Lagonosticta
senegala
Estrildidae Passeriformes Africa
Skoracki and
Dabert 2002
“
Lagonosticta
rufopicta
Estrildidae Passeriformes Africa
Skoracki and
Dabert 2002
Syringophilopsis
melittophagi Skoracki et
Dabert, 2001
Merops bulocki
Meropidae Coraciiformes Africa
Skoracki and
Dabert 2001b
Syringophilopsis
mimidus Sikora, Fajfer et
Skoracki, 2011
Margarops
fuscatus
Mimidae Passeriformes North America Sikora et al. 2011
Syringophilopsis
muscicapus Skoracki,
2011
Muscicapa
dauurica
Muscicapidae Passeriformes Asia Skoracki 2011
Syringophilopsis nitens
Skoracki et Dabert, 2001
Malimbus nitens
Ploceidae Passeriformes Africa
Skoracki and
Dabert 2001b
“
Quelea erythrops
Ploceidae Passeriformes Africa
Skoracki and
Sikora 2003
“
Ploceus vitellinus
Ploceidae Passeriformes Africa
Skoracki et al.
2011b
Syringophilopsis
nucifragus Skoracki,
2011
Nucifraga
caryocatactes
Corviidae Passeriformes Europe Skoracki 2011
Syringophilopsis
passericus Skoracki,
2011
Passer domesticus
Passeridae Passeriformes Asia Skoracki 2011
Syringophilopsis
passerinae (Clark, 1964)
Passerina cyanea
Cardinalidae Passeriformes North America
Clark 1964b;
Skoracki et al.
2011a
“
Passerina ciris
Cardinalidae Passeriformes North America
Skoracki et al.
2011a
Syringophilopsis
polioptilus Skoracki,
Flannery et Spicer, 2008
Polioptila caerulea
Polioptilidae Passeriformes North America
Skoracki et al.
2008
Syringophilopsis rusticus
Skoracki, 2004
Hirundo rustica
Hirundinidae Passeriformes Europe Skoracki 2004b
Syringophilopsis sialiae
Skoracki, Flannery et
Spicer, 2008
Sialia mexicana
Turdidae Passeriformes North America
Skoracki et al.
2008
Syringophilopsis sittae
Skoracki, Hendricks et
Spicer, 2011
Sitta carolinensis
Sittidae Passeriformes North America
Skoracki et al.
2011a
Syringophilopsis
spinolettus Skoracki,
2004
Anthus spinoletta
Motacillidae Passeriformes Europe, Asia
Skoracki 2004b,
2011; Skoracki
and Bochkov
2010
Syringophilopsis
sturnellus Skoracki,
Hendricks et Spicer,
2011
Sturnella neglecta
Icteridae Passeriformes North America
Skoracki et al.
2011a
Syringophilopsis sturni
Chirov et Kravtsova,
1995
Sturnus vulgaris
Sturnidae Passeriformes Europe, Asia
Chirov and
Kravtsova 1995;
Bochkov and
Mironov 1998;
Skoracki 2004b,
2011
Syringophilopsis turdi
(Fritsch, 1958)
Turdus pilaris
Turdidae Passeriformes Europe
Fritsch 1958;
Skoracki 2004b
Appendix
i29
Syringophilopsis turdi
(Fritsch, 1958)
Turdus philomelos
Turdidae Passeriformes Europe
Bochkov and
Galloway 2001;
Skoracki 2004b
“
Turdus atrogularis
Turdidae Passeriformes Asia
Skoracki and
Bochkov 2010
“
Turdus iliacus
Turdidae Passeriformes Europe Skoracki 2011
“
Turdus migratorius
Turdidae Passeriformes North America
Skoracki et al.
2008
“
Turdus albicollis
Turdidae Passeriformes South America
Skoracki and
Sikora 2003
Syringophilopsis tyranni
Bochkov et Galloway,
2004
Tyrannus tyrannus
Tyrannidae Passeriformes North America
Bochkov and
Galloway 2004
“
Tyrannus verticalis
Tyrannidae Passeriformes North America
Skoracki et al.
2008
“
Myiarchus crinitus
Tyrannidae Passeriformes North America
Skoracki et al.
2008
“
Myiarchus
cinerascens
Tyrannidae Passeriformes North America
Skoracki et al.
2008
Syringophilopsis
veselovsky Skoracki,
Antczak et Riegert, 2009
Pycnonotus
barbatus
Pycnonotidae Passeriformes Africa
Skoracki et al.
2009a
“
Chlorocichla
aviventris
Pycnonotidae Passeriformes Africa
Skoracki et al.
2011b
“
Eurillas latirostris
Pycnonotidae Passeriformes Africa
Skoracki et al.
2011b
Syringophilopsis yose
Skoracki, Tryjanowski et
Hromada, 2002
Lanius sp.
Laniidae Passeriformes Africa
Skoracki et al.
2002
Syringophilus Heller, 1880
Syringophilus bipectina-
tus Heller, 1880
Gallus gallus
Phasianidae Galliformes Worldwide Skoracki 2011
“
Alectoris rufa
Phasianidae Galliformes Europe Skoracki 2011
“
Ptilopachus
petrosus
Phasianidae Galliformes Africa
Skoracki and
Sikora 2003
Syringophilus numidae
Bochkov, 2000
Numida meleagris
Numididae Galliformes Africa Bochkov 2000c
Terratosyringophilus Bochkov et Perez, 2002
Terratosyringophilus
geotrygonus Skoracki et
Glowska, 2008
Geotrygon linearis
Columbidae Columbiformes South America
Skoracki and
Glowska 2008
Terratosyringophilus
pioni Bochkov et Perez,
2002
Pionus senilis
Psittacidae Psittaciformes North America
Bochkov and
Perez 2002
Terratosyringophilus
longisoma (Casto, 1979)
Zenaida asiatica
Columbidae Columbiformes North America Casto 1979
“
Zenaida macroura
Columbidae Columbiformes North America Casto 1979
Tinamiphilopsis Skoracki et Sikora, 2004
Tinamiphilopsis ariconte
Skoracki, Sikora et
Ozminski, 2012
Rhynchotus
rufescens
Tinamidae Tinamiformes South America
Skoracki et al.
2012b
“
Nothura boraquira
Tinamidae Tinamiformes South America
Skoracki et al.
2012b
“
Nothura minor
Tinamidae Tinamiformes South America
Skoracki et al.
2012b
M. Skoracki, S. Zabludovskaya and A.V. Bochkov
i30
Tinamiphilopsis elegans
Skoracki et Sikora, 2004
Eudromia elegans
Tinamidae Tinamiformes South America
Skoracki and
Sikora, 2004a
Torotrogla Kethley, 1970
Torotrogla aphelocoma
Bochkov, Flannery et
Spicer, 2009
Aphelocoma
californica
Corvidae Passeriformes North America
Bochkov et al.
2009
Torotrogla calcarius
Skoracki, 2004
Calcarius
lapponicus
Calcariidae Passeriformes Europe Skoracki 2004c
Torotrogla cardinalis
Bochkov, Flannery et
Spicer, 2009
Cardinalis
cardinalis
Cardinalidae Passeriformes North America
Bochkov et al.
2009
Torotrogla cardueli
Bochkov et Mironov,
1999
Spinus spinus
Fringillidae Passeriformes Europe
Bochkov and
Mironov 1999;
Skoracki 2004c
“
Carduelis
cannabina
Fringillidae Passeriformes Europe
Skoracki 2004c,
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
pytyopsittacus
Fringillidae Passeriformes Europe Skoracki 2011
Torotrogla cocco-
thraustes Bochkov,
Flannery et Spicer, 2009
Coccothraustes
vespertinus
Fringillidae Passeriformes North America
Bochkov et al.
2009
Torotrogla cyanocitta
Bochkov, Flannery et
Spicer, 2009
Cyanocitta stelleri
Corvidae Passeriformes North America
Bochkov et al.
2009
Torotrogla gaudi
Bochkov et Mironov,
1998
Fringilla coelebs
Fringillidae Passeriformes Europe
Bochkov and
Mironov 1998
“
Fringilla
montifringilla
Fringillidae Passeriformes Europe Skoracki 2011
“
Pyrrhula pyrrhula
Fringillidae Passeriformes Europe Skoracki 2011
Torotrogla lullulae
Skoracki, Hromada et
Kuczynski, 2001
Lullula arborea
Alaudidae Passeriformes Europe
Skoracki et al.
2001a
“
Eremophila
alpestris
Alaudidae Passeriformes Asia Skoracki 2011
Torotrogla lusciniae
Skoracki, 2004
Luscinia
megarhynchos
Muscicapidae Passeriformes Europe Skoracki 2004c
“
Luscinia svecica
Muscicapidae Passeriformes Europe Skoracki 2004c
Torotrogla merulae
Skoracki, Dabert et
Ehrnsberger, 2000
Turdus merula
Turdidae Passeriformes Europe, Asia
Skoracki et al.
2000; Bochkov
et al. 2009;
Glowska et al.
2007
“
Turdus viscivorus
Turdidae Passeriformes Europe Skoracki 2004c
“
Turdus philomelos
Turdidae Passeriformes Europe, Asia
Skoracki 2004c;
Glowska et al.
2007
“
Turdus torquatus
Turdidae Passeriformes Europe Skoracki 2011
Torotrogla mima
Kethley, 1970
Mimus polyglottos
Mimidae Passeriformes North America Kethley 1970
“
Mimus triurus
Mimidae Passeriformes South America Sikora et al. 2011
“
Mimus patagonicus
Mimidae Passeriformes Africa Sikora et al. 2011
Appendix
i31
Torotrogla modularis
Nattress et Skoracki,
2007
Prunella modularis
Prunellidae Passeriformes Europe
Nattress and
Skoracki 2007;
Skoracki 2011
Torotrogla piranga
Bochkov, Flannery et
Spicer, 2009
Piranga
ludoviciana
Cardinalidae Passeriformes North America
Bochkov et al.
2009
Torotrogla pycnonotus
Skoracki, 2011
Pycnonotus
xanthopygos
Pycnonotidae Passeriformes Asia Skoracki 2011
Torotrogla rubeculi
Skoracki, 2004
Erithacus rubecula
Muscicapidae Passeriformes Europe
Skoracki 2004c;
Bochkov et al.
2009
Torotrogla villosa
(Hancock, 1895)
Phainopepla nitens
Ptilogonatidae Passeriformes North America Kethley 1970
Trypetoptila Kethley, 1970
Trypetoptila casmerodia
Kethley, 1970
Ardea alba
Ardeidae Pelecaniformes North America Kethley 1970
Species inquirenda
Syringophilopsis
hunanensis Liu, 1988
Passer domesticus
Passeridae Passeriformes Asia
Liu 1988;
Skoracki 2011
Picobiinae Johnston et Kethley, 1973
Calamincola Casto, 1977
Calamincola lobata
(Casto, 1977)
Crotophaga
sulcirostris
Cuculidae Cuculiformes North America Casto 1977
Columbiphilus Kivganov et Sharafat, 1995
Columbiphilus alectoris
(Fain, Bochkov et
Mironov, 2000)
Alectoris sp.
Phasianidae Galliformes Africa
Fain et al. 2000;
Skoracki 2011
“
Alectoris rufa
Phasianidae Galliformes Europe
Skoracki and
Sikora 2011
Columbiphilus khush-
alkhani Kivganov et
Sharafat, 1995
Columba livia
Columbidae Columbiformes Asia
Kivganov and
Sharafat 1995
Columbiphilus polonica
(Skoracki, Magowski et
Dabert, 2001)
Gallus gallus
Phasianidae Galliformes Europe
Skoracki et al.
2001; Skoracki
2011
“
Gallus sonneratii
Phasianidae Galliformes Asia
Skoracki and
Sikora 2011
Columbiphilus pteroclesi
(Skoracki et OConnor,
2010)
Pterocles
senegallus
Pteroclidae Pterocliformes Africa
Skoracki and
OConnor 2010
“
Pterocles
coronatus
Pteroclidae Pterocliformes Africa
Skoracki and
OConnor 2010
Neopicobia Skoracki, 2011
Neopicobia anthi
(Fritsch, 1958)
Anthus trivialis
Motacillidae Passeriformes Europe Fritsch 1958
“
Anthus cervinus
Motacillidae Passeriformes Europe Skoracki 2011
Neopicobia cardinalis
(Skoracki, Hendricks et
Spicer, 2010)
Cardinalis
cardinalis
Cardinalidae Passeriformes North America
Skoracki et al.
2010c
Neopicobia carpodacus
(Skoracki, Hendricks et
Spicer, 2010)
Carpodacus
purpureus
Fringillidae Passeriformes North America
Skoracki et al.
2010c
Neopicobia epthianura
(Skoracki, Glowska et
Sikora, 2008)
Epthianura
aurifrons
Meliphagidae Passeriformes Australia
Skoracki et al.
2008
M. Skoracki, S. Zabludovskaya and A.V. Bochkov
i32
Neopicobia glossopsitta
(Skoracki, Glowska et
Sikora, 2008)
Glossopsitta
porphyrocephala
Psittacidae Psittaciformes Australia
Skoracki et al.
2008
Neopicobia modularis
(Skoracki et Magowski,
2001)
Prunella modularis
Prunellidae Passeriformes Europe
Skoracki and
Magowski 2001
“
Prunella
atrogularis
Prunellidae Passeriformes Asia Skoracki 2011
Neopicobia locustella
(Skoracki, Bochkov et
Wauthy, 2004)
Locustella naevia
Locustellidae Passeriformes Europe
Skoracki et al.
2004b
“
Locustella
luscinioides
Locustellidae Passeriformes Europe Skoracki 2011
Neopicobia pyrrholae-
mus (Skoracki et
Glowska, 2008)
Pyrrholaemus
sagittatus
Acanthizidae Passeriformes Australia
Skoracki and
Glowska 2008c
Neopicobia troglodytes
(Skoracki, Hendricks et
Spicer, 2010)
Troglodytes aedon
Troglodytidae Passeriformes North America
Skoracki et al.
2010c
Neopicobia zumpti
(Lawrence, 1959)
Streptopelia
capicola
Columbidae Columbiformes Africa Lawrence 1959a
“
Streptopelia
senegalensis
Columbidae Columbiformes Africa Lawrence 1959a
“
Columba livia
Columbidae Columbiformes North America
Bochkov et al.
2005
Picobia Haller, 1878
Picobia biarmicus
Skoracki, Bochkov et
Wauthy, 2004
Panurus biarmicus
Panuridae Passeriformes Europe
Skoracki et al.
2004b
Picobia brotogeris Fain,
Bochkov et Mironov,
2000 inc. sedis
Brotogeris
jugularis
Psittacidae Psittaciformes South America Fain et al. 2000
Picobia caudati
Skoracki et Hebda, 2004
Aegithalos
caudatus
Aegithalidae Passeriformes Europe
Skoracki and
Hebda 2004
Picobia cissa Skoracki,
Bochkov et Wauhty,
2004
Cissa chinensis
Corviidae Passeriformes Asia
Skoracki et al.
2004b
Picobia cetti Skoracki,
2011
Cettia cetti
Cettidae Passeriformes Asia Skoracki 2011
Picobia chloris
Bochkov, Mironov et
Kravtsova, 2000
Chloris chloris
Fringillidae Passeriformes Asia
Bochkov et al.
2000
Picobia corcoracus
Skoracki, Glowska et
Sikora, 2008
Corcorax
melanorhamphos
Corcoracidae Passeriformes Australia
Skoracki et al.
2008
Picobia currucae
Skoracki et Magowski,
2001
Sylvia curruca
Sylviidae Passeriformes Europe, Asia
Skoracki and
Magowski 2001;
Skoracki 2011
“
Sylvia hortensis
Sylviidae Passeriformes Asia
Glowska et al.
2007
“
Sylvia nisoria
Sylviidae Passeriformes Asia Skoracki 2011
Picobia dinemellia
Glowska et Skoracki,
2011
Dinemellia
dinemelli
Ploceidae Passeriformes Africa
Glowska and
Skoracki 2011b
Picobia dryobatis
(Fritsch, 1958)
Dendrocopos
major
Picidae Piciformes Europe
Fritsch 1956;
Skoracki et al.
2004b
Appendix
i33
Picobia dryobatis
(Fritsch, 1958)
Dendrocopos
minor hortorum
Picidae Piciformes Europe Skoracki 2011
“
Dendrocopos
leucotos
Picidae Piciformes Europe Skoracki 2011
“
Picoides
tridactylus
Picidae Piciformes
North America,
Europe
Skoracki et al.
2010c; Skoracki
2011
“
Picoides scalaris
Picidae Piciformes North America
Skoracki et al.
2010c
Picobia dziabaszewskii
Glowska, Dragun-Dami-
an et Dabert, 2012
Garrulax formosus
Leiothrichidae Passeriformes Asia
Glowska et al.
2012
Picobia eremophila
Skoracki, 2011
Eremophila
bilopha
Alaudidae Passeriformes Africa Skoracki 2011
Picobia galerida
Skoracki, 2011
Galerida cristata
Alaudidae Passeriformes Europe Skoracki 2011
Picobia heeri Haller,
1878
Picus canus
Picidae Piciformes Europe Skoracki 2011
“
Picus viridis
Picidae Piciformes Europe Skoracki 2011
Picobia indonesiana
Skoracki et Glowska,
2008
Aplonis panayensis
Sturnidae Passeriformes Asia
Skoracki and
Glowska 2008c
Picobia lemi Skoracki,
Glowska et Sikora, 2008
Manucodia
chalybatus
Paradisaeidae Passeriformes Australia
Skoracki et al.
2008
Picobia leucophaeus
Skoracki, Hendricks et
Spicer, 2010 inc. sedis
Leucophaeus
atricilla
Laridae
Charadrii-
formes
North America
Skoracki et al.
2010c
Picobia oritis Skoracki,
Antczak et Riegert, 2009
Cyanomitra oritis
Nectarinidae Passeriformes Africa
Skoracki et al.
2009
“
Cyanomitra
olivacea
Nectarinidae Passeriformes Africa
Skoracki et al.
2011
Picobia paludicola
Skoracki et Kiljan, 2002
Acrocephalus
paludicola
Acrocephalidae Passeriformes Europe
Skoracki and
Kiljan 2002
Picobia phoeniculi Fain,
Bochkov et Mironov,
2000
Phoeniculus
purpureus
Phoeniculidae Coraciiformes Africa Fain et al. 2000
Picobia poicephali
Skoracki et Dabert, 2002
inc. sedis
Poicephalus
senegalus
Psittacidae Psittaciformes Africa
Skoracki and
Dabert 2002
Picobia pycnonoti
Glowska, Skoracki et
Khourly, 2007
Pycnonotus
xanthopygos
Pycnonotidae Passeriformes Asia
Glowska et al.
2007
Picobia riparius
Skoracki, 2011
Riparia riparia
Hirundinidae Passeriformes Europe Skoracki 2011
Picobia sturni Skoracki,
Bochkov et Wauthy,
2004
Sturnus vulgaris
Sturnidae Passeriformes Europe
Skoracki et al.
2004b
“
Sturnus cineraceus
Sturnidae Passeriformes Asia Skoracki 2011
Picobia troglodytidus
Sikora et Skoracki, 2012
Henicorhina
leucophrys
Troglodytidae Passeriformes North America Sikora et al. 2012
Rafapicobia Skoracki, 2011
Rafapicobia zirnitra
Skoracki, 2011
Saxicola rubetra
Muscicapidae Passeriformes Europe Skoracki 2011
Rafapicobia toxostoma
Sikora, Fajfer et
Skoracki, 2011
Toxostoma
curvirostre
Mimidae Passeriformes North America Sikora et al. 2011
M. Skoracki, S. Zabludovskaya and A.V. Bochkov
i34
Rafapicobia dendroco-
laptesi Skoracki et
Solarczyk, 2012
Dendrocolaptes
platyrostris
Funariidae Passeriformes South America
Skoracki and
Solarczyk 2012
“
Dendrocolaptes
picumnus
Funariidae Passeriformes South America
Skoracki and
Solarczyk 2012
Rafapicobia lepidoco-
laptesi Skoracki et
Solarczyk, 2012
Lepidocolaptes
afnis
Funariidae Passeriformes South America
Skoracki and
Solarczyk 2012
“
Lepidocolaptes
souleyetii
Funariidae Passeriformes South America
Skoracki and
Solarczyk 2012
Ereynetidae Oudemans, 1931
Speleognathinae Fain, 1957
Boydaiini Fain, 1985
Boydaia Womersley, 1953
Boydaia sturni (Boyd,
1948)
Sturnus vulgaris
Sturnidae Passeriformes
Eurasia,
Australia, North
America
Boyd 1948;
Fain1961a; Fain
1963b; Domrow
1969; Sixl 1972;
Pence 1973a,
1975; Zablu-
dovskaya 1998;
Knee et al. 2008
“
Acridotheres tristis
Sturnidae Passeriformes Australia Domrow 1969
“
Ampeliceps
coronatus
Sturnidae Passeriformes Europe Fain 1971
“
Pastor roseus
Sturnidae Passeriformes Asia
Zabludovskaya
1998
“
Passer domesticus
Ploceidae Passeriformes North America
Porter and
Strandtmann
1952
Boydaia agelaii Fain et
Aitken, 1968
Chrysomus
icterocephalus
Icteridae Passeriformes South America
Fain and Aitken
1968; Fain and
Lukoschus 1972
“
Agelaius
phoeniceus
Icteridae Passeriformes North America
Fain and Aitken
1968; Pence
1973
“
Chrysomus
rucapillus
Icteridae Passeriformes
European Zoo
(captive)
Fain and Aitken
1968
“
Agelaius humeralis
Icteridae Passeriformes South America
Dusbabek and
Cerny 1970
“
Dives
atroviolaceus
Icteridae Passeriformes South America
Dusbabek and
Cerny 1970
“
Molothrus ater
Icteridae Passeriformes North America
Fain and Hyland
1975
“
Quiscalus
mexicanus
Icteridae Passeriformes North America
Fain and Hyland
1975
“
Spiza americana
Cardinalidae Passeriformes North America
Fain and Hyland
1975
Boydaia americana Fain,
1963
Passerina caerulea
Cardinalidae Passeriformes
European Zoo
(captive)
Fain 1963b
“
Cyanocompsa
cyanoides
Cardinalidae Passeriformes South America
Fain and Aitken
1970
“
Piranga rubra
Cardinalidae Passeriformes
European Zoo
(captive)
Fain 1963b;
Pence 1975
“
Ramphocelus
carbo
Thraupidae Passeriformes South America
Fain and Aitken
1968
Appendix
i35
Boydaia americana Fain,
1963
Tangara gyrola
Thraupidae Passeriformes South America
Fain and Aitken
1970
“
Coereba aveola
Coerebidae Passeriformes South America
Fain and Aitken
1968
“
Sporophila
americana
Emberizidae Passeriformes South America
Fain and Aitken
1970
“
Volatinia jacarina
Emberizidae Passeriformes South America
Fain and Aitken
1968
Boydaia bradornis Fain,
1956
Bradornis pallidus
griseus
Muscicapidae Passeriformes Africa Fain 1956b, 1971
“
Luscinia luscinia
Muscicapidae Passeriformes Europe
Zabludovskaya
1998
Boydaia buphagi Fain,
1961
Buphagus
africanus
Buphagidae Passeriformes Africa Fain 1961a
“
Lamprotornis
chloropterus
Sturnidae Passeriformes Africa Fain 1963b
Boydaia cecropis Fain,
1969
Cecropis
abyssinica
Hirundinidae Passeriformes Africa Fain 1969, 1971
Boydaia cinnyris Fain,
1969
Cinnyris cupreus
Nectariniidae Passeriformes Africa Fain 1969, 1971
“
Chalcomitra
senegalensis
Nectariniidae Passeriformes Africa Fain 1969, 1971
Boydaia cyanerpes Fain,
1963
Cyanerpes cyaneus
Thraupidae Passeriformes
European Zoo
(captive)
Fain 1963b
Boydaia cyanerpes
hylocichla Fain et
Hyland, 1975
Catharus ustulatus
Turdidae Passeriformes North America
Fain and Hyland
1975
Boydaia faini Dusbabek
et Cerny, 1970
Setophaga cerulea
Parulidae Passeriformes South America
Dusbabek and
Cerny 1970;
Pence 1975
Boydaia formicarii Fain
et Aitken, 1970
Formicarius colma
Formicariidae Passeriformes South America
Fain and Aitken
1970
“
Formicarius analis
Formicariidae Passeriformes South America
Fain and Aitken
1970
Boydaia hirundoae Fain,
1956
Hirundo rustica
Hirundinidae Passeriformes
Africa, Europe,
North America,
Australia
(introduced)
Fain 1963a; Fain
and Hyland
1975; Domrow,
1969; Sixl 1972;
Pence 1973a,
1975
“
Stelgidopteryx
rucollis
Hirundinidae Passeriformes South America
Fain and Aitken
1968
“
Progne subis
Hirundinidae Passeriformes South America
Fain and Hyland
1970
“
Hirundo neoxena
Hirundinidae Passeriformes Australia Domrow 1965
“
Petrochelidon ariel
Hirundinidae Passeriformes Australia Domrow 1969
“
Cecropis
abyssinica
Hirundinidae Passeriformes Africa Zumpt 1961
“
Hirundo aethiopica
Hirundinidae Passeriformes Africa Fain et al. 1977
Boydaia indica Fain,
1969
Acridotheres tristis
Sturnidae Passeriformes
European Zoo
(captive)
Fain 1969
“
Gracupica contra
Sturnidae Passeriformes
European Zoo
(captive)
Fain 1969
“
Sturnia malabarica
Sturnidae Passeriformes
European Zoo
(captive)
Fain 1969
M. Skoracki, S. Zabludovskaya and A.V. Bochkov
i36
Boydaia indica Fain,
1969
Temenuchus
pagodarum
Sturnidae Passeriformes
European Zoo
(captive)
Fain 1969
“
Lamprotornis
chloropterus
Sturnidae Passeriformes Africa Fain 1971
“
Lamprotornis
chalybaeus
Sturnidae Passeriformes
European Zoo
(captive)
Fain 1971
“
Lamprotornis
superbus
Sturnidae Passeriformes
European Zoo
(captive)
Fain 1971
Boydaia jordani Van
Eyndhoven, 1955
Turdus pilaris
Turdidae Passeriformes Europe
Van Endhoven
1955
“
Turdus migratorius
Turdidae Passeriformes North America
Clark 1967;
Pence 1975
“
Turdus viscivorus
Turdidae Passeriformes Europe
Zabludovskaya
1998
Boydaia laticoxa Fain et
Aitken, 1969
Glyphorynchus
spirurus
Funariidae Passeriformes South America
Fain and Aitken
1969, 1970
Boydaia laticoxa
philydori Fain et Aitken,
1970
Philydor pyrrhodes
Furnariidae Passeriformes South America
Fain and Aitken
1970
Boydaia locustellae
Fain, 1970
Locustella
luscinioides
Locustellidae Passeriformes Europe Fain 1970
Boydaia loxiae Fain,
1963
Loxia curvirostra
Fringillidae Passeriformes Europe Fain 1963a
“
Icterus galbula
Icteridae Passeriformes North America
Fain and Hyland
1975
Boydaia maluri
Domrow, 1969
Malurus amabilis
Maluridae Passeriformes Australia Domrow 1969
“
Malurus
melanocephalus
Maluridae Passeriformes Australia Domrow 1969
Boydaia mimi Fain et
Hyland, 1970
Mimus gilvus
Mimidae Passeriformes South America
Fain and Hyland
1970
“
Mimus polyglottos
Mimidae Passeriformes North America
Pence 1973a,
1975
Boydaia morenoi
Dusbabek et Cerny, 1970
Mimus polyglottos
Mimidae Passeriformes South America
Dusbabek and
Cerny 1970
Boydaia myzomelae
Domrow, 1969
Myzomela
sanguinolenta
Meliphagidae Passeriformes Australia Domrow 1969
“
Xanthotis
aviventer
Meliphagidae Passeriformes Australia Domrow 1991
Boydaia nectarinia Fain,
1958
Cyanomitra
verticalis
Nectariniidae Passeriformes Africa
Fain 1958b,
1963b
Boydaia pheucticola
Pence et Costa, 1976
Pheucticus
melanocephalus
Cardinalidae Passeriformes North America
Pence and Costa
1976
“
Spiza americana
Cardinalidae Passeriformes North America Spicer 1987
Boydaia psalidoprocnei
Fain, 1956
Psalidoprocne
albiceps
Hirundinidae Passeriformes Africa Fain 1956b, 1971
“
Tachycineta
bicolor
Hirundinidae Passeriformes North America
Pence 1973a,
1975; Knee et al.
2008
“
Riparia riparia
Hirundinidae Passeriformes Asia
Zabludovskaya
1998
Boydaia pycnonoti Fain,
1956
Pycnonotus
barbatus
Pycnonotidae Passeriformes Africa
Fain 1956b; Fain
et al. 1977
“
Pycnonotus
goiavier
Pycnonotidae Passeriformes Asia Fain 1963b
Appendix
i37
Boydaia pycnonoti Fain,
1956
Pycnonotus
leucogenys
Pycnonotidae Passeriformes Asia Fain 1963b, 1971
Boydaia quiscali Clark,
1960
Quiscalus quiscula
Icteridae Passeriformes North America
Clark 1960;
Pence 1973a,
1975
“
Quiscalus
mexicanus
Icteridae Passeriformes North America
Pence 1973a,
1975
“
Molothrus ater
Icteridae Passeriformes North America
Pence 1973a,
1975
“
Agelaius
phoeniceus
Icteridae Passeriformes North America
Pence 1973a,
1975
“
Saltator maximus
Thraupidae Passeriformes South America
Fain and Aitken
1970
“
Saltator
coerulescens
Thraupidae Passeriformes South America
Fain and Hyland
1970
Boydaia rosickyi
Dusbabek et Cerny 1970
Passerina cyanea
Cardinalidae Passeriformes South America
Dusbabek and
Cerny 1970;
Pence 1975
Boydaia saxicolae Fain,
1969
Saxicola torquatus
axillaris
Muscicapidae Passeriformes Africa Fain 1969
“
Myrmecocichla
nigra
Muscicapidae Passeriformes Africa Fain 1969
“
Monticola
angolensis
Muscicapidae Passeriformes Africa Fain 1969
Boydaia sinensis Fain et
Bafort, 1963
Leiothrix lutea
Leiothrichidae Passeriformes Asia
Fain and Bafort
1963
Boydaia spatulata Fain,
1955
Cercotrichas
hartlaubi
Muscicapidae Passeriformes Africa
Fain, 1955,
1956b
“
Pycnonotus
barbatus
Pycnonotidae Passeriformes Africa
Fain 1956a,
1956b, 1971
“
Cossypha
natalensis
Muscicapidae Passeriformes Africa
Fain 1956a,
1956b, 1971
“
Cossypha
polioptera
Muscicapidae Passeriformes Africa
Fain, 1956b,
1963b
“
Pogonocichla
stellata
Muscicapidae Passeriformes Africa
Fain 1956a,
1956b, 1971
“
Cercomela tractrac
Muscicapidae Passeriformes Africa
Fain 1956a,
1956b, 1971
“
Chloropeta similis
Acrocephalidae Passeriformes Africa
Fain 1956a,
1956b, 1971
“
Schistolais
leucopogon
Cisticolidae Passeriformes Africa
Fain 1956a,
1956b, 1971
“
Melaniparus niger
Paridae Passeriformes Africa
Fain 1956a,
1956b, 1971
“
Macronyx croceus
Motacillidae Passeriformes Africa
Fain 1956a,
1956b, 1971
“
Tiaris canorus
Emberizidae Passeriformes
European Zoo
(captive)
Fain 1958a
“
Tangara sp.
Thraupidae Passeriformes
European Zoo
(captive)
Fain 1958a
“
Cissomela
pectoralis
Meliphagidae Passeriformes Australia Domrow 1969
“
Philemon
corniculatus
Meliphagidae Passeriformes Australia Domrow 1969
“
Philemon
citreogularis
Meliphagidae Passeriformes Australia Domrow 1969
M. Skoracki, S. Zabludovskaya and A.V. Bochkov
i38
Boydaia spatulata Fain,
1955
Cyssomela
pectoralis
Meliphagidae Passeriformes Australia Domrow 1969
“
Sialia sialis
Turdidae Passeriformes North America
Pence 1973c,
1975
“
Pycnonotus
barbatus
Pycnonotidae Passeriformes Africa
Zabludovskaya
2002
Boydaia synallaxis Fain,
Hyland et Aitken, 1969
Synallaxis rutilans
Furnariidae Passeriformes South America Fain, 1969
“
Synallaxis
gujanensis
Furnariidae Passeriformes South America Fain, 1969
“
Synallaxis
erythrothorax
Furnariidae Passeriformes South America Fain, 1969
“
Empidonax sp.
Tyrannidae Passeriformes South America
Fain and Hyland
1970
Boydaia tyrannis Ford,
1959
Tyrannus tyrannus
Tyrannidae Passeriformes North America Ford 1959
“
Myiophobus
fasciatus
Tyrannidae Passeriformes South America
Fain and Aitken
1968
“
Tyrannus verticalis
Tyrannidae Passeriformes North America
Brooks and
Strandtmann
1960; Pence
1975
“
Elaenia avogaster
Tyrannidae Passeriformes South America
Fain and Aitken
1968
“
Pyrocephalus
rubinus
Tyrannidae Passeriformes South America
Fain and Hyland
1970
“
Myiarchus
cinerascens
Tyrannidae Passeriformes North America
Pence and Casto
1976
“
Tyrannus
dominicensis
Tyrannidae Passeriformes South America
Dusbabek and
Cerny 1970; Fain
et al. 1975;
Pence 1975
“
Pachyramphus
aglaiae
Tityridae Passeriformes South America
Fain and Hyland
1970
“
Manacus manacus
Pipridae Passeriformes South America
Fain and Aitken
1968, 1970
“
Pipra pipra
Pipridae Passeriformes South America
Fain and Aitken
1970c
“
Pipra fasciicauda
Pipridae Passeriformes South America
Fain and Aitken
1970
“
Lepidothrix iris
Pipridae Passeriformes South America
Fain and Aitken
1970
Boydaia zosteropis Fain,
1963
Zosterops
senegalensis
Zosteropidae Passeriformes Africa Fain 1963b, 1971
“
Zosterops lateralis
Zosteropidae Passeriformes Australia Domrow 1969
Boydaia zumpti Fain,
1955
Eurillas latirostris
eugenius
Pycnonotidae Passeriformes Africa
Fain 1955,
1956a, 1956b
“
Bleda syndactylus
Pycnonotidae Passeriformes
European Zoo
(captive)
Fain 1971
“
Schoenicola
brevirostris
Locustellidae Passeriformes Africa Fain 1958a, 1971
Boydaia crassipes
(Berlese et Trouessart,
1889)
Passer domesticus
Passeridae Passeriformes
Europe,
Australia, North
America
Berlese and
Trouessart 1889;
Porter and
Strandtmann,
1952; Domrow
1969
Appendix
i39
Boydaia crassipes
(Berlese et Trouessart,
1889)
Carduelis carduelis
Fringillidae Passeriformes Australia Domrow 1969
“
Motacilla ava
Fringillidae Passeriformes
Australia
(introduced)
Domrow 1969
“
Lalage sueurii
Campephagidae Passeriformes Australia Domrow 1991
Boydaia psittaci Fain et
Mortelmans, 1959
Pionites
melanocephalus
Psittacidae Psittaciformes
European Zoo
(captive)
Fain 1963b
“
Pionites
leucogaster
Psittacidae Psittaciformes South America Fain 1963b
“
Bolborhynchus
lineola
Psittacidae Psittaciformes
European Zoo
(captive)
Fain 1963b
“
Forpus passerinus
Psittacidae Psittaciformes
European Zoo
(captive)
Fain 1971
“
Brotogeris
jugularis
Psittacidae Psittaciformes
European Zoo
(captive)
Fain 1971
“
Brotogeris
versicolurus
Psittacidae Psittaciformes
European Zoo
(captive)
Fain 1971
“
Psilopsiagon
aymara
Psittacidae Psittaciformes
European Zoo
(captive)
Fain 1971
Boydaia aratingae Fain,
1963
Aratinga jandaya
Psittacidae Psittaciformes
European Zoo
(captive)
Fain 1963b, 1971
Boydaia falconis Fain,
1956
Falco cuvierii
Falconidae Falconiformes Africa Fain 1956b
“
Falco tinnunculus
Falconidae Falconiformes Africa Fain 1963b
“
Falco berigora
Falconidae Falconiformes Australia Domrow 1969
“
Falco sparverius
Falconidae Falconiformes North America
Pence and Casto
1976
Boydaia podargi Fain et
Lukoschus, 1979
Podargus
strigoides
Podargidae
Caprimulgi-
formes
Australia
Fain and
Lukoschus 1979
Boydaia colini Clark,
1958
Colinus virginianus
Odontophoridae Galliformes North America
Clark 1958; Fain
and Hyland 1975
Boydaia clarki Fain,
1963
Callipepla
squamata
Odontophoridae Galliformes
European Zoo
(captive); North
America
Fain 1963b;
Pence and Casto
1976
Coboydaia Fain, 1985
Coboydaia clavata
(Fain, 1955)
Ploceus cucullatus
Ploceidae Passeriformes Africa
Fain 1955,
1956a, 1963b
“
Ploceus xanthops
Ploceidae Passeriformes Africa
Fain 1956a,
1963b, 1971
“
Ploceus pelzelni
Ploceidae Passeriformes Africa
Fain 1956a,
1963b, 1971
“
Quelea quelea
Ploceidae Passeriformes Africa
Fain. 1956a,
1960, 1963b,
1971; Fain et al.
1977
“
Ploceus baglafecht
Ploceidae Passeriformes Africa
Fain 1956a,
1963b, 1971
“
Ploceus vitellinus
Ploceidae Passeriformes Africa Fain et al. 1977
“
Ploceus cucullatus
Ploceidae Passeriformes Africa Fain et al. 1977
“
Euplectes
hordeaceus
Ploceidae Passeriformes Africa Fain et al. 1977
“
Euplectes ardens
Ploceidae Passeriformes Africa Fain et al. 1977
“
Euplectes
macroura
Ploceidae Passeriformes Africa Fain et al. 1977
M. Skoracki, S. Zabludovskaya and A.V. Bochkov
i40
Coboydaia clavata
(Fain, 1955)
Quelea erythrops
Ploceidae Passeriformes Africa
Fain 1956a,
1960, 1963b,
1971; Fain et al.
1977; Mwase
and Baker 2006
“
Vidua chalybeata
Viduidae Passeriformes Africa Fain 1963b, 1971
Coboydaia amandavae
(Fain, 1962)
Sporaeginthus
subavus
Estrildidae Passeriformes
European Zoo
(captive)
Fain 1962a
“
Amandava
amandava
Estrildidae Passeriformes Asia Fain 1963b
“
Erythrura prasina
Estrildidae Passeriformes
European Zoo
(captive)
Fain1970
“
Malurus
melanocephalus
Maluridae Passeriformes Australia Domrow 1969
Coboydaia nigra (Fain,
1955)
Serinus sulphura-
tus shelleyi
Fringillidae Passeriformes Africa
Fain 1955,
1956a, 1956b
“
Serinus
citrinelloides
Fringillidae Passeriformes Africa
Fain 1956b,
1963b, 1971
“
Serinus
mozambicus
Fringillidae Passeriformes Africa
Fain 1963b,
1971; Fain et al.
1977
“
Serinus burtoni
Fringillidae Passeriformes Africa
Fain, 1963b,
1971
“
Carduelis carduelis
Fringillidae Passeriformes Europe
Fain 1962b,
1963b, 1971;
Sixl 1972
“
Carpodacus
mexicanus
Fringillidae Passeriformes North America
Fain and Hyland
1975
“
Carpodacus
erythrinus
Fringillidae Passeriformes Eurasia
Zabludovskaya
1998
“
Fringilla
montifringilla
Fringillidae Passeriformes Asia
Zabludovskaya
1998
“
Fringilla coelebs
Fringillidae Passeriformes Eurasia
Sixl 1972;
Zabludovskaya
1998
“
Passer griseus
Passeridae Passeriformes Africa Fain 1963b, 1971
“
Passer montanus
Passeridae Passeriformes Asia
Zabludovskaya
1998
“
Passer
ammodendri
Passeridae Passeriformes Asia
Zabludovskaya
1998
“
Passer
hispaniolensis
Passeridae Passeriformes Asia
Zabludovskaya
1998
“
Emberiza rutila
Emberizidae Passeriformes Africa, Asia
Fain 1963b,
1971; Zablu-
dovskaya 1998
“
Emberiza tristrami
Emberizidae Passeriformes Asia
Zabludovskaya
1998
“
Emberiza
spodocephala
Emberizidae Passeriformes Asia
Zabludovskaya
1998
“
Emberiza pusilla
Emberizidae Passeriformes Asia
Zabludovskaya
1998
“
Emberiza rustica
Emberizidae Passeriformes Asia
Zabludovskaya
1998
“
Emberiza citrinella
Emberizidae Passeriformes Asia
Zabludovskaya
1998
Appendix
i41
Coboydaia nigra (Fain,
1955)
Emberiza aureola
Emberizidae Passeriformes Asia
Zabludovskaya
1998
“
Spizella passerina
Emberizidae Passeriformes North America
Fain and Hyland
1975
“
Euphonia violacea
Fringillidae Passeriformes South America
Fain and Aitken
1968
“
Anthus spinoletta
Motacillidae Passeriformes North America Pence 1973
“
Anthus trivialis
Motacillidae Passeriformes Eurasia
Zabludovskaya
1998
Coboydaia nigra icteri
(Fain et Hyland, 1970)
Icterus spurius
Icteridae Passeriformes South America
Fain and Hyland
1970
Icterus galbula
Icteridae Passeriformes North America
Fain and Hyland
1970
Coboydaia nigra
motacillae (Fain, 1969)
Motacilla ava
Motacillidae Passeriformes Africa, Eurasia
Fain 1969, 1971;
Zabludovskaya
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
arundinaceus
Acrocephalidae Passeriformes Europe Fain 1963b, 1971
“
Acrocephalus
scirpaceus
Acrocephalidae Passeriformes Europe Fain 1963b, 1971
“
Motacilla alba
Motacillidae Passeriformes Asia
Zabludovskaya
1998
Coboydaia sturnellae
(Clark, 1960)
Sturnella magna
Icteridae Passeriformes North America Clark 1960
Coboydaia (Apodiboydaia) Fain, 1985
Coboydaia trochila
(Fain, 1958)
Chlorestes notata
Trochilidae Apodiformes
European Zoo
(captive)
Fain, 1958a,
1961a, 1963b,
1971
Trispeleognathini Fain, 1985
Trispeleognathus Fain, 1958
Trispeleognathus striatus
(Crossley, 1952)
Columba livia
Columbidae Columbiformes
North America,
Australia
(introduced)
Crossley 1952;
Domrow 1969
“
Streptopelia
semitorquata
Columbidae Columbiformes Africa Fain 1956a
“
Columbina
talpacoti
Columbidae Columbiformes South America
Fain and Aitken
1968
“
Leptotila verreauxi
Columbidae Columbiformes South America Amaral 1963
Aureliania Fain, 1958
Aureliania aureliani
(Fain, 1955)
Tyto alba
Tytonidae Strigiformes
Africa,
Australia, South
America
Fain 1955;
Dusbabek and
Cerny 1970;
Domrow 1991
Ophthalmognathus Dubinin, 1957
Ophthalmognathus
shoutedeni (Fain, 1955)
Ardeola idae
Ardeidae Pelecaniformes Africa Fain 1955, 1956a
“
Ardea cinerea
Ardeidae Pelecaniformes Europe Dubinin 1957
“
Nycticorax
nycticorax
Ardeidae Pelecaniformes Africa Fain 1955, 1956a
“
Nycticorax
caledonicus
Ardeidae Pelecaniformes Australia
Domrow 1969,
1991
M. Skoracki, S. Zabludovskaya and A.V. Bochkov
i42
Ophthalmognathus
tenorioae Fain et Goff,
1980
Nycticorax
nycticorax
Ardeidae Pelecaniformes
Australia
(Hawaii)
Fain and Goff
1980
Psittaboydaia Fain, 1985
Psittaboydaia psittacu-
lae (Fain, 1962)
Psittacula krameri
Psittacidae Psittaciformes Africa Fain 1962c
“
Aprosmictus
erythropterus
Psittacidae Psittaciformes Australia Domrow 1991
“
Platycercus
adscitus
Psittacidae Psittaciformes Australia Domrow 1991
Psittaboydaia amazona
(Fain et Lukoschus,
1972)
Amazona
amazonica
Psittacidae Psittaciformes
European Zoo
(captive)
Fain and
Lukoschus, 1972
“
Agapornis
roseicollis
Psittacidae Psittaciformes
Australia
(captive)
Domrow 1969
Psittaboydaia (Trichoglossiella) Fain, 1985
Psittaboydaia tricho-
glossi (Fain, 1970)
Trichoglossus
haematodus
Psittacidae Psittaciformes
European Zoo
(captive);
Australia
Fain 1970;
Domrow 1969,
1991
“
Psitteuteles
versicolor
Psittacidae Psittaciformes Australia
Domrow 1969,
1991
Neastrida Fain, 1962
Neastrida parrae (Fain,
1956)
Actophilornis
africanus
Jacanidae Gruiformes South Africa Fain 1956d
Neoboydaia Fain, 1958
Neoboydaia philomachi
(Fain, 1956)
Philomachus
pugnax
Charadriidae
Charadrii-
formes
Africa, Europe,
Australia
(introduced)
Fain 1956a,
1956c; Domrow
1969; Zablu-
dovskaya 1998
“
Tringa glareola
Scolopacidae
Charadrii-
formes
Africa, Australia
Fain 1963b;
Domrow 1969
“
Actitis macularius
Scolopacidae
Charadrii-
formes
South Africa
Fain and Aitken
1968
“
Limosa lapponica
Scolopacidae
Charadrii-
formes
Australia
Domrow 1969,
1991
Neoboydaia philomachi
eroliae Fain et Hyland,
1970
Calidris minutilla
Scolopacidae
Charadrii-
formes
South America
Fain and Hyland
1970
“
Actitis macularius
Scolopacidae
Charadrii-
formes
South America
Fain and Hyland
1970
“
Arenaria interpres
Scolopacidae
Charadrii-
formes
North America
Fain and Hyland
1975
“
Limnodromus
griseus
Scolopacidae
Charadrii-
formes
North America
Fain and Hyland
1975
“
Tringa solitaria
Scolopacidae
Charadrii-
formes
North America
Fain and Hyland
1975
“
Tringa
melanoleuca
Scolopacidae
Charadrii-
formes
North America
Clark 1964a;
Pence 1973a
“
Tringa avipes
Scolopacidae
Charadrii-
formes
North America Clark 1964a
“
Calidris melanotos
Scolopacidae
Charadrii-
formes
North America Clark 1964a
“
Gallinago
gallinago
Scolopacidae
Charadrii-
formes
North America
Asia
Pence 1973a;
Zabludovskaya
1998
Appendix
i43
Neoboydaia philomachi
eroliae Fain et Hyland,
1970
Limnodromus
scolopaceus
Scolopacidae
Charadrii-
formes
North America Pence 1973a
Neoboydaia philomachi
thalasseus Fain et
Hyland, 1975
Thalasseus
maximus
Laridae
Charadrii-
formes
North America
Fain and Hyland
1975
“
Sterna hirundo
Laridae
Charadrii-
formes
North America
Fain and Hyland
1975
Neoboydaia galachrisiae
Fain, 1961
Glareola cinerea
Glareolidae
Charadrii-
formes
Africa Fain 1961b
“
Glareola nuchalis
Glareolidae
Charadrii-
formes
Africa Fain 1964b
Neoboydaia colymbi-
formi Clark, 1964
Podiceps
nigricollis
Podicipedidae
Podicipedi-
formes
North America Clark 1964a
“
Podilymbus
podiceps
Podicipedidae
Podicipedi-
formes
North America Pence 1973a
“
Tachybaptus
rucollis
Podicipedidae
Podicipedi-
formes
Australia
Domrow 1969,
1991
Pterniboydaia Fain, 1985
Pterniboydaia pternistis
(Fain, 1955)
Francolinus afer
Phasianidae Galliformes Africa Fain 1955
“
Francolinus
swainsonii
Phasianidae Galliformes Africa Fain 1963
“
Perdix perdix
Phasianidae Galliformes Europe Fain 1962b
Speleognathopsis Cooreman, 1954
Speleognathopsis galli
Cooreman, 1954
Gallus gallus
Phasianidae Galliformes
North America,
Australia
Cooreman 1954;
Domrow 1969
“
Numida meleagris
Numididae Galliformes
Africa, Australia
(introduced)
Fain 1956a;
Domrow 1969
Speleognathopsis benoiti
Fain, 1955
Vanellus
crassirostris
Charadriidae
Charadrii-
formes
Africa Fain 1955, 1956a
“
Vanellus senegallus
Charadriidae
Charadrii-
formes
Africa Fain 1956a
“
Vanellus miles
Charadriidae
Charadrii-
formes
Australia
Domrow 1969,
1991
“
Elseyornis
melanops
Charadriidae
Charadrii-
formes
Australia
Domrow 1969,
1991
“
Erythrogonys
cinctus
Charadriidae
Charadrii-
formes
Australia
Domrow 1969,
1991
“
Smutsornis
africanus
Glareolidae
Charadrii-
formes
South Africa Fain et al. 1966
Speleognathopsis
charadricola (Fain,
1964)
Charadrius
pecuarius
Charadriidae
Charadrii-
formes
European Zoo
(captive)
Fain 1964a
Speleognathopsis
onychognathi (Fain,
1964)
Onychognathus
walleri
Sturnidae Passeriformes Africa Fain 1964a
Speleognathopsis
accipitris (Domrow,
1969)
Accipiter fasciatus
Accipitridae Accipitriformes Australia
Domrow 1969,
1991
“
Accipiter
cirrocephalus
Accipitridae Accipitriformes Australia Domrow 1969
“
Aquila audax
Accipitridae Accipitriformes Australia Domrow 1969
“
Haliaeetus
leucogaster
Accipitridae Accipitriformes Australia Domrow 1969
M. Skoracki, S. Zabludovskaya and A.V. Bochkov
i44
Speleognathopsis
accipitris (Domrow,
1969)
Haliastur
sphenurus
Accipitridae Accipitriformes Australia Domrow 1969
“
Milvus migrans
Accipitridae Accipitriformes Australia Domrow 1969
“
Elanus axillaris
Accipitridae Accipitriformes Australia Domrow 1969
Speleognathopsis wai
Fain, Vercammen-
Grandjean et Wagner,
1966
Rhinoptilus
africanus
Glareolidae
Charadrii-
formes
South America Fain et al. 1966
Metaboydaia Fain, 1962
Metaboydaia poffei
(Fain, 1955)
Scopus umbretta
Scopidae Pelecaniformes Africa Fain 1955
“
Coracias caudatus
Coraciidae Coraciiformes Africa Fain 1955
Meropiboydaia Fain, 1985
Meropiboydaia merops
(Fain, 1956)
Merops apiaster
Meropidae Coraciiformes Africa Fain 1955, 1956a
“
Merops ornatus
Meropidae Coraciiformes Australia
Domrow 1969,
1991
Astrida Fain, 1955
Astrida caprimulgi Fain,
1955
Caprimulgus fossii
Caprimulgidae
Caprimulgi-
formes
Africa Fain 1956a
“
Bubo africanus
Strigidae Strigiformes Africa Fain 1956b
“
Glaucidium
perlatum
Strigidae Strigiformes Africa Fain 1956b
“
Otus senegalensis
Strigidae Strigiformes Africa Fain 1956b
“
Otus spilocephalus
Strigidae Strigiformes Asia Fain 1963b
Astrida (Cerylonyssus) Fain et Aitken, 1970
Astrida chlorocerylei
Fain et Aitken, 1970
Chloroceryle aenea
Alcediniidae Coraciiformes South America
Fain and Aitken
1970
Ralliboydaia Fain, 1962
Ralliboydaia lateralli
Fain, 1962
Laterallus
melanophaius
Rallidae Gruiformes South America Fain 1962a
Fulica americana
Rallidae Gruiformes North America Pence 1973a
Ralliboydaia coccyzae
(Pence, 1973)
Coccyzus
americanus
Cuculidae Cuculiformes North America Pence 1973a
Ralliboydaia porphyrio-
nis (Domrow, 1965)
Porphyrio
porphyrio
Rallidae Gruiformes Australia
Domrow 1965b,
1969
Picinyssus Fain, 1969
Picinyssus buccanodon
Fain, 1969
Buccanodon
duchaillui
Lybiidae Piciformes Africa Fain 1969
Phoenicopteriella Fain, 1970
Phoenicopteriella
mirabilis Fain, 1970
Phoenicopterus
ruber
Phoenicopteri-
dae
Phoenicopteri-
formes
European Zoo
(captive)
Fain 1970
Phoenicopteriella
womersleyi (Fain, 1955)
Dendrocygna
viduata
Anatidae Anseriformes Africa Fain 1955, 1956a
“
Sarkidiornis
melanotos
Anatidae Anseriformes North America Fain 1956a
“
Aythya afnis
Anatidae Anseriformes North America Clark 1958
“
Anas strepera
Anatidae Anseriformes North America Clark 1958
“
Anas acuta
Anatidae Anseriformes
North America,
Asia
Clark 1958;
Zabludovskaya
1998
“
Anas discors
Anatidae Anseriformes North America
Fain and Hyland
1975
Appendix
i45
Phoenicopteriella
womersleyi (Fain, 1955)
Anas crecca
Anatidae Anseriformes Europe
Zabludovskaya
1998
“
Anas querquedula
Anatidae Anseriformes Europe
Zabludovskaya
1998
