KEYS
Key to families of Prostigmata — permanent parasites of birds
1. Subcapitulum of gnathosoma distinctly developed. Palps present. At least some of idiosomal and leg setae filiform ...... 2
— Subcapitulum of gnathosoma strongly reduced
or absent. Palps absent. Idiosomal and leg setae
represented by alveoli or fleshy-like projections ...... Cloacaridae Camin, Moss, Oliver et Singer, 1967
(Pneumophagus Fain et Smiley, 1989)
2. Propodonotal setae si represented by unmodi-fied simple setae. Genital papillae absent. Tibia of
legs I with or without solenidion φ ...... 3
— Propodonotal setae si represented by bothridia. Genital papillae present. Tibia of legs I
with complicate ereynetal organ (sensory complex) ...... Ereynetidae (Speleognathinae)
3. Distal part of hypostome fused with subcapitulum. 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 solenidion each ...... 4
— Distal part of hypostome not fused with subcapitulum. Peritremes situated at base of subcapitulum, linear. Palpal trochanter, femur and genu
fused to each other. Setae scx distinctly developed,
situated dorsally. Tibia and genu I without solenidion ...... Harpirhynchidae Dubinin, 1957
4. Subcapitulum not deeply submerged in idiosoma, subcapitular apodeme absent. Claw of palpal
tibia present. Palpal tibia and tarsus separated. Se
tae 4a present. Setae a’I absent ...... Cheyletidae Leach, 1815
— 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
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 terminally. Tarsi of legs I much longer than wide ......
(Cheletosomatini Volgin, 1969) ...... 3
— Eupathidium sul” smooth. Palpal claw strongly
curved ventrally and without basal angles. Idiosomal 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
— Eupathidium sul” comb-like. Setae c3 absent
...... 4
4. Coxal fields III and IV contiguous. Setae c1
situated on propodonotal shield ...... Cheletopsis Oudemans, 1904
— Coxal fields III and IV distinctly separated. Se
tae c1 situated off propodonotal shield ...... Eucheletopsis Volgin, 1969
5. Setae c4 absent. Female hysteronotal shield absent 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 subequal 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, respectively ...... 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. Solenidion φI present. Trochanter IV, femora III and IV
with 1 seta each ...... Neocheyletiella Baker, 1949
— Palpal femora with ventral processes.
Propodonotal shield distinctly developed, covering part of hysteronotum. Setae c3 absent. In females, aggenital setae absent. Solenidion φI absent. Trochanter IV, femora III and IV without
setae ...... Apodicheles Fain, 1979
Key to genera of Syringophilidae
1. Tibiotarsus of palps truncate on distal margin.
Prorals setae p’ and p” of legs I–IV with two minute 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 developed. Flight feathers as main habitat. (Syringophilinae 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 ventrally ........ 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 level. Pocket-like structures in anterior part of propodonotum present. Stylophore constricted posteriorly ....... Psittaciphilus Fain, Bochkov et Mironov, 2000
6. Hypostomal apex ornamented by 2 pairs of median finger-like protuberances. Apodemes I and II
parallel and fused to each other ........ 7
— Hypostomal apex smooth. Apodemes I divergent, 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 absent) .......... 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 absent) .......... 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
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 recurved ........... 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 fields I and II similar in size and shape.
Supernumerary setae vsIII absent. Legs I thicker
than II–IV ......... Torotrogla Kethley, 1970
— Coxal fields I and II disimilar in size and shape.
Supernumerary setae vsIII present. Legs I–IV subequal .......... 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 fields 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 fields 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 fields I and II dissimilar in size and
shape ............ 39
— Coxal fields 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 located 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 Bochkov et Chystiakov, 2001
46. Each lateral branch of peritremes reduced to 3
chambers. Setae se and c1 situated at same transverse level. In males 2 pairs of aggenital setae present 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 setae se and c1 situated at same transverse level ............ Kalamotrypetes Casto, 1980
47. Setae si and se situated at same level. All terminal 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 opisthosomal 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)
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 sclerotization in middle part. Genital lobes present or absent. Physogastric female worm-shaped in outline
........... Picobia Haller, 1878
50. Solenidia phi on tibia of leg I absent. Hypostomal 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. Hypostomal apex rounded. Peritremes with clearly visible 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 pseudanal setae present ......... Rafapicobia Skoracki, 2011
— Genital setae absent, 2 pairs of pseudanal setae
present ............ Neopicobia Skoracki, 2011
Key to genera of Harpirhynchidae
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 filiform 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 filiform, 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 maximum of 4 articulate segments ............. 6
— Female: Idiosoma circular or subcircular. Legs
I and II well developed, with 5 articulated segments ............ 5
5. In females, setae dG, l”G, and dF of palps similar 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 different 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 reduced 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 pectinate ............ 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 absent ............. Metharpyrhynchus Fain, 1972
10. Female: Idiosoma wider than long, with 2
broad lateral lobes. Dorsal shield poorly developed. Palpal seta l”G absent ........... 11
— Female: Body dome-shaped, with 2 posterolateral lobes. Dorsal shield distinctly developed.
Palpal seta l”G present ............. Anharpyrhynchus
Fain, 1972
11. Female: Setae ve and si modified 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
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. Number of coxal setae I–IV 1–1–1–1, trochanters without 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 normally 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 filiform 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 genital 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 exclusively or predominantly barbed ............. 11
11. Dorsal shield well developed. Palps with 3
segments. Eyes either flat or with prominent lenses
........... Speleognathopsis Cooreman, 1954
— Dorsal shield absent. Palps with 2 segments.
Eyes with prominent lenses ........... Metaboydaia Fain, 1962
12. Dorsal setae of idiosoma either exclusively or
predominantly dentate .............. 14
— Dorsal setae of idiosoma either exclusively or
predominantly filiform ............ 13
13. Dorsal shield absent. Palps with 1 segment. Setae 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 modified; 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 segment. Eyes absent ............. Picinyssus Fain, 1969