subfamilies, Ewing, 1944

Stekolnikov, Alexandr A., 2018, Taxonomy and distribution of African chiggers (Acariformes, Trombiculidae), European Journal of Taxonomy 395, pp. 1-233 : 10-16

Key to subfamilies , genera and subgenera of African trombiculid larvae

1. Scutum with 2 AM setae, with or without anteromedian process (nasus), fsp = 6.6.6, fSt = 0.2, sensilla flagelliform (subfamily Leeuwenhoekiinae ) ........................................................................ 7

– Scutum without AM setae, fsp = 7.6.6, sensilla expanded, fusiform or clavate (subfamily Gahrliepiinae ) ................................................................................................................................... 6

– Scutum with 1 AM seta (absent in Afrotrombicula quadriscutala (Taufflieb, 1965) only), without nasus, fsp = 7.7.7 (7.6.6 or 6.6. 6 in some rare cases) (subfamily Trombiculinae ) ..........................11

– Scutum with 2 AM setae or 1 AM + nasus, fsp = 7.7.7 (subfamily Apoloniinae ) ............................ 2

2. Sensilla expanded, palpal claw 2-pronged, PL setae on scutum, genualae II and III present, microgenuala II present .................................................................................................................... 3

– Sensilla flagelliform, palpal claw with 3–4 prongs, PL setae extrascutal (peniscutum), genualae II and III absent, microgenuala II absent .............................................................................................. 4

3. Two genualae I, scutum with biconvex posterior margin, fCx = 1.2.1 ............................................... ................................................................... Afracarella Vercammen-Grandjean & Kolebinova, 1968

– Three genualae I, scutum with very prominent and rounded posterior margin, fCx = 1.1.1 .............. ............................................................................................................. Sauracarella Lawrence, 1949

4. Two anterior and two posterior sternal setae; fCx = 1.1.1; galealae branched ................................... ....................................................................................................................... Afropolonia Goff, 1983

– Two sternal setae between coxae I, two sternal setae between coxae II, and many setae between coxae III; humeroventral setae situated between coxae II and III; fCx = 1.2.1 or 1.1.1; galealae branched or nude ............................................................................................................................... 5

5. Eyes 2 + 2, nasus present (absent in Straelensia monosetosa ), microgenuala I present, genuala I absent, palpal tarsus with 4 branched setae and nude subterminala ................................................... ................................................................... Straelensia Vercammen-Grandjean & Kolebinova, 1968

– Eyes 1 + 1 or absent, nasus absent, microgenuala I and genuala I present, palpal tarsus with 5 branched setae and nude subterminala ...................................... Vargatula Brennan & Yunker, 1966

6. Scutum with posterior margin extending far beyond level of PL setae to include two or more pairs of dorsal idiosomal setae ....................................................................... Gahrliepia Oudemans, 1912

– Scutum with posterior margin extending beyond level of PL setae to include one pair of dorsal idiosomal setae ....................................................................................... Schoengastiella Hirst, 1915

– Scutum subpentagonal, with posterior margin angulate or rounded, with four setae only (2 AL and 2 PL) .................................................................................................................... Walchia Ewing, 1931

7. Cheliceral blade with large apical cap bearing numerous lateral teeth, scutum in shape of crescent, much wider than long, with nasus, sensillary bases posterior to PL, dorsal idiosomal setae sometimes expanded ........................................................................................................................................... 8

– Cheliceral blade with dorsal subapical tooth and ventral teeth or with dorsal and ventral rows of teeth, scutum subpentagonal or trapezoidal, with nasus ................................................................... 9

– Cheliceral blade long, with large, recurved ventral row of teeth and dorsal teeth or hooks, scutum wide, sometimes striplike, without nasus, legs very long (Ip = 1000–2550). Parasites of bats ......... ...................................................................................................................... Whartonia Ewing, 1944

8. PL and dorsal idiosomal setae expanded, blunt-tipped, with very long expanded branches .............. .......................................................................................................... Austrombicula Lawrence, 1949 – PL not expanded, dorsal idiosomal setae without giant branches ....... Austracarus Lawrence, 1949

9. Palpal tarsus with 7 branched setae, tracheae and stigmae present, onychotriches present, number of branched setae on leg femur I, II, and III – 6.6.5. Parasites of reptiles .............................................. ............................................................................................ Matacarus Vercammen-Grandjean, 1956

– Palpal tarsus with 6 branched setae, leg claws often with 2 conspicuous onychotriches ............... 10

– Palpal tarsus with 5 branched setae, tracheae and stigmae absent, onychotriches absent, multiple mastisetae present on leg III ................................................ Mastalacarus Goff & Lukoschus, 1983

– Palpal tarsus with 4 branched setae, AM setae with one accessory branch, tracheae and stigmae absent, onychotriches absent ........................................................................ Tateracarus Goff, 1983

10. Cheliceral blade with ventral row of denticles and dorsal teeth, tracheae and stigmae present, sensilla usually branched .................................................................................... Acomatacarus Ewing, 1942 – Dorsal teeth on cheliceral blade absent, tracheae and stigmae absent, sensilla usually nude ............ ................................................................................................................. Hyracarus Lawrence, 1949

11. Sensilla expanded, fusiform to globose (tribe Schoengastiini ) ...................................................... 12 – Sensilla flagelliform, usually branched (tribe Trombiculini ) .......................................................... 33

12. Scutum wide, crescent-shaped, with rounded or concave posterior margin, sensillary bases posterior to PL, AL and PL setae approximate to each other, eyes absent, two or more pairs of humeral setae, scutal and idiosomal setae covered with long thin barbs, galeal setae branched ................................ ........................................................................................ Brunehaldia Vercammen-Grandjean, 1960

– Scutum not crescent-shaped ........................................................................................................... 13

13. Tibiala III absent, palpal tarsus with 3–5 branched setae and sometimes nude subterminala ........ 14 – Tibiala III present, palpal tarsus with 4–7 branched setae and sometimes nude subterminala ...... 20

14. Three mastitarsalae, 3 mastitibialae, and 1 mastifemorala present, genualae II and III absent .......... ............................................................................................................ Gerbillicula Kolebinova, 1984 – Mastisetae absent, genualae II and III usually present ................................................................... 15

15. Cheliceral blade with one dorsal tooth and large hook, posterior scutal margin prominent, sensillary bases posterior to PL, eyes absent ................................. Tauffliebiella Vercammen-Grandjean, 1960

– Cheliceral blade with tricuspid cap, sometimes with ventral row of small teeth or dorsal serration, eyes usually present ........................................................................................................................ 16

16. Palpal claw usually divided by more than 3 prongs (4–12), distance between sensillary bases almost the same as distance from sensillary base to lateral scutal margin, cheliceral blade sometimes with ventral row of small teeth, palpal tarsus with 4 branched setae, always 4 humeral setae .................. ............................................................................... Cheladonta Lipovsky, Crossley & Loomis, 1955

– Palpal claw always 3-pronged, sensillary bases situated far apart, closer to lateral scutal margin than to each other (genus Schoutedenichia ). .......................................................................................... 17

17. Palpal tarsus with 3 branched setae, fCx = 1.1.9, Ip = 485 ................................................................ ............................................................. Schoutedenichia (Trisetichia) Vercammen-Grandjean, 1958

– Palpal tarsus with 5 branched setae, cheliceral blade with dorsal serration ....................................... ............................................................. Schoutedenichia (Pentachia) Vercammen-Grandjean, 1958

– Palpal tarsus with 4 branched setae ................................................................................................ 18

– Palpal tarsus with 4 branched setae and nude subterminala, scutum small, as wide as long, SD> AW, fPp usually N/N/NNN, galealae nude, fsp = 7.7.7 or 7.6.6, fCx = 1.1.(2–7). Intranasal parasites of mammals ............................................... Schoutedenichia (Nasichia) Vercammen-Grandjean, 1958

18. One genuala I, galealae always branched, PL setae sometimes foliate, Ip = 670–760 ....................... ........................................................... Schoutedenichia (Platytrichia) Vercammen-Grandjean, 1960 – Two genualae I ................................................................................................................................ 19

19. AL> PL> AM .............................. Schoutedenichia (Brennanichia) Vercammen-Grandjean, 1960 – PL> AL ....................... Schoutedenichia (Schoutedenichia) Jadin & Vercammen-Grandjean, 1954

20. Palpal tarsus with 4–5 branched setae ............................................................................................ 21 – Palpal tarsus with 6–7 branched setae and sometimes nude subterminala ..................................... 25

21. Cheliceral blade with one large dorsal hook, palpal claw 2-pronged (axial prong internal), galeal setae nude, scutum small, trapezoidal, elongated, longer than width, 3 genualae I ........................... ........................................................................... Holubicula Daniel & Vercammen-Grandjean, 1985 – Cheliceral blade with tricuspid cap, palpal claw 3-pronged ........................................................... 22

22. Scutum with reduced posterior angles (peniscutum), PL setae extrascutal ........................................ ............................................................................................................ Trisetica Traub & Evans, 1950 – PL setae situated on scutum ............................................................................................................ 23

23. Tarsala I in distal position (level of subterminala), sensillary bases situated close to each other ...... ........................................................................................................................ Helenicula Audy, 1954 – Tarsala I situated clearly posterior to level of subterminala ........................................................... 24

24. Scutum with cuticular striations around sensillary bases, fPp = B/B/NNB, galealae nude, mastitarsalae and mastitibialae ciliated in basal part sometimes present ................................................................. ....................................................................................... Ornithogastia Vercammen-Grandjean, 1960

– Scutum without cuticular striations, galealae branched or nude, mastisetae absent ........................... ......................................................................................................... Susa Audy & Nadchatram, 1960

25. Palpal tarsus with 6 branched setae (6B), scutum with anterolateral shoulders, mastitarsala III usually present ............................................................................................................................................. 26

– Palpal tarsus with 6 branched setae and nude subterminala (6BS), or 7B, or 7BS, scutum without anterolateral shoulders .................................................................................................................... 27

26. Scutum subquadrate .......................................................................... Ascoschoengastia Ewing, 1946 – Scutum very long, with posterior margin extending far beyond level of PL setae (to level of 2 nd row of dorsal setae), sensillary bases situated far anterior to PL and close to lateral scutal margins ........ .............................................................................................. Elianella Vercammen-Grandjean, 1956

27. Cheliceral blade serrate along its dorsal edge or having large dorsal and lateral hooks, palpal tarsus with 7 branched setae and nude subterminala, galealae nude, scutum subpentagonal, sensilla globose ............................................................................................................................................ 28

– Cheliceral blade with tricuspid cap only, scutum trapezoidal ........................................................ 29

28. Cheliceral blade usually serrate on their dorsal edge, 2–3 genualae I ................................................ ........................................................................................................... Schoengastia Oudemans, 1910 – Cheliceral blade with 2 large dorsal hooks and 1 lateral hook, 4–5 genualae I. Parasites of batrachians .......................................................................................... Endotrombicula Ewing, 1931

29. Scutum with cuticular striations around sensillary bases, sensillary bases situated far anterior to PL, sensilla pyriform or globose. Parasites of birds ................................. Neoschoengastia Ewing, 1929 – Scutum without cuticular striations or striated on its posterior half or margins ............................. 30

30. Eyes absent, legs with expanded (lanceolate) claws and empodia, tarsi with supplementary bars or semi-bars. Parasites of bats ...................................................................... Riedlinia Oudemans, 1914 – Eyes 2 + 2, leg claws unexpanded .................................................................................................. 31

31. Tarsala I long and slender, twice as long as tarsala II, scutum with concave lateral margins, sensilla fusiform, mastitarsala usually present. Parasites of bats .................................................................... ..................................................................... Trombigastia Vercammen-Grandjean & Brennan, 1957

– Tarsala I normal, scutum wider than long, with convex or biconvex posterior margin, sensilla globose to fusiform, sensillary bases often situated far apart (closer to lateral scutal margins than to each other), mastitarsala and mastitibiala sometimes present (genus Herpetacarus ) .................... 32

32. Palpal tarsus with 7 branched setae, sensilla globose or claviform, sensillary bases situated far apart ...................................................... Herpetacarus (Abonnencia) Vercammen-Grandjean, 1960

– Palpal tarsus with 6 branched setae and nude subterminala, sensilla fusiform, sensillary bases situated not too far apart, mastisetae absent. Parasites of reptiles ................................................................... ................................................................. Herpetacarus (Cricacarus) Vercammen-Grandjean, 1966

– Palpal tarsus with 7 branched setae and nude subterminala, sensilla fusiform, PL always longest scutal setae. Parasites of reptiles ......... Herpetacarus (Herpetacarus) Vercammen-Grandjean, 1960

33. Palpal tarsus with 5 branched setae, 3 genualae I. Parasites of bats ............................................... 34 – Palpal tarsus with 6B, 7B or 7BS ................................................................................................... 35

34. Scutum subquadrate or subpentagonal, 2 genualae III (one genuala can be regarded as mastigenuala) ............................................................. Sasatrombicula Vercammen-Grandjean, 1960

– Scutum trapezoidal, PL setae sometimes extrascutal (peniscutum), pretarsala I paired, subterminala and parasubterminala absent, tarsalae I and II long and slender ........................................................ .................................................................................................... Grandjeana Koçak & Kemal, 2009

35. PL setae extrascutal (peniscutum) .................................................................................................. 36 – PL setae situated on scutum ............................................................................................................ 37

36. Palpal claw 2-pronged, eyes 1 + 1 ........................................................... Sauriscus Lawrence, 1949 – Palpal claw 3-pronged, eyes 2 + 2, palpal tarsus with 6 setae, galealae nude, 1 genuala I, genualae II and III absent, 2–3 mastitarsalae III and 1–2 mastitibialae III present ............................................... .................................................................................. Zumptrombicula Vercammen-Grandjean, 1967

37. Palpal tarsus with 6 branched setae ................................................................................................ 38 – Palpal tarsus with 7 branched setae and sometimes with nude subterminala ................................. 40

38. Scutum without anterolateral shoulders, scutal and dorsal idiosomal setae expanded, rod-like, 5 genualae I, 2 genualae II, fCx = 1.3.3, Ip> 1300 ............................................................................ ............................................................................. Multigniella Vercammen-Grandjean & Fain, 1957

– Scutum with anterolateral shoulders, scutal and dorsal idiosomal setae unexpanded, setiform, 2–3 genualae I, 1 genuala II, Ip = 400–900 ........................................................................................... 39

39. Scutum trapezoidal, wider than long, eyes absent, 2 genualae III ...................................................... ......................................................................................... Marcandrea Vercammen-Grandjean, 1960 – Scutum subtrapezoidal, subpentagonal or subquadrate, its length and width subequal, eyes usually 2 + 2, 1 genuala III ............................................................................. Microtrombicula Ewing, 1950

40. Genualae II and III absent, mastitarsala and mastitibiala III present, palpal claw 2-pronged ............ ............................................................................................. Blanciella Vercammen-Grandjean, 1960 – Genualae II and III present (genuala III absent in some Miyatrombicula only) ............................. 41

41. Cheliceral blade with large terminal hooks, tarsala I gigantic, much longer than tarsala II, 3–4 genualae I. Parasites of amphibians and reptiles ............. Vercammenia Audy & Nadchatram, 1957

– Cheliceral blade with tricuspid cap only, tarsala I gigantic, more than twice as long as tarsala II, 3 genuala I, galeal setae with one branch, fPp = B/N/NNB, scutum rectangular .................................. ........................................................................................ Tanautarsala Vercammen-Grandjean, 1960

– Cheliceral blade with tricuspid cap only (rarely with few dorsal teeth), tarsala I not gigantic (except for Blankaartia (Megaciella) ), 1–3 genualae I ............................................................................... 42

42. Scutum rectangular or trapezoidal, with posterior margin sinuous, concave, slightly bilobate or almost straight ................................................................................................................................. 43

– Scutum subpentagonal or subhexagonal, with more or less prominent, rounded or angulate posterior margin ............................................................................................................................................. 48

43. Mastitarsala III present, galeal setae nude, palpal claw 2-pronged, fPp = B/B/NNN or B/N/NNN, 3 genualae I, Ip = 1022–1455 .................................... Whartonacarus Vercammen-Grandjean, 1960

– Femorala III present, galeal setae branched, palpal claw 3-pronged, fPp = N/N/NNN, 2 genualae I. Parasites of bats ..................................... Oudemansidium Vercammen-Grandjean & André, 1966

– Extra genuala III or mastigenuala III present, mastifemorala III present, galeal setae nude, palpal claw 3-pronged, fPp = N/N/NNN or B/N/NNN, 2 genualae I. Parasites of bats ................................ ........................................................................................... Chiroptella Vercammen-Grandjean, 1960

– Mastisetae, extra genualae or femoralae absent .............................................................................. 44

44. Scutum trapezoidal, with anterolateral shoulders (AL setae inserted on lateral scutal margins far from anterior scutal margin), galeal setae branched or nude, 3 or 2 genualae I. Parasites of bats ............................................................................... Myotrombicula Womersley & Heaslip, 1943

– Scutum without anterolateral shoulders (AL setae inserted in anterolateral angles of scutum), galeal setae always branched, 2 genualae I ............................................................................................... 45

45. Palpal tarsus with 7 branched setae and nude subterminala ........................................................... 46 – Palpal tarsus with 7 branched setae, without nude subterminala .................................................... 47

46. Setae on palpal femur, genu and tibia nude (fPp = N/N/NNN). Parasites of bats, occasionally on other hosts ................................................. Willmannium Vercammen-Grandjean & Langston, 1976 – Palpal femoral seta always branched, palpal genual seta branched or nude ....................................... ................................................................................. Ericotrombidium Vercammen-Grandjean, 1966

47. Palpal femoral and genual seta usually nude, dorsal palpal tibial seta usually branched ................... ................................................. Leptotrombidium Nagayo, Miyagawa, Mitamura & Imamura, 1916 – Palpal femoral seta branched, palpal genual seta branched or nude, dorsal palpal tibial seta always nude ......................................................................... Hypotrombidium Vercammen-Grandjean, 1966

48. Scutum pentagonal, with pointed posterior margin and anterolateral shoulders, sensillary bases situated far apart and clearly anterior to PL, puncta on leg coxae arranged in longitudinal lines (genus Blankaartia ) ........................................................................................................................ 49

– Scutum without anterolateral shoulders, puncta on leg coxae distributed irregularly .................... 50

49. Galeal setae nude, rarely branched, posterior scutal margin rounded, AW nearly as large as PW, PL never as long as PW, leg tarsala I only slightly longer than leg tarsala II, Ip = 800–1100 ................. ...................................................................................... Blankaartia (Blankaartia) Oudemans, 1911 – Galeal setae branched, scutum striated, posterior scutal margin acute, PW much larger than AW, idiosomal setae slender and very long (> 70 µm), PL much longer than PW, leg tarsala I gigantic, at least twice as long as leg tarsala II, Ip = 1200–1520 .......................................................................... .................................................................... Blankaartia (Megaciella) Vercammen-Grandjean, 1960

50. Two or more additional setae present on posterior scutal margin and/or between scutal margin and eyes ................................................................................................................................................. 51 – Scutum with 5 setae only (1 AM, 2 AL and 2 PL) .......................................................................... 52

51. Scutum large, with prominent posterior margin, 4–10 PPL setae present, AL and PL situated close to each other ....................................................................................................... Heaslipia Ewing, 1944

– Scutum subpentagonal or nearly trapezoidal, with rounded posterior margin, two or more PPL or scuto-ocular setae present, AL and PL situated not close to each other ............................................. ......................................................................................................... Xinjiangsha Wen & Shao, 1984

52. Palpal claw 2-pronged, galeal setae nude, scutum subtrapezoidal, with broadly rounded posterior margin ..................................................................................................... Eutrombicula Ewing, 1938 – Palpal claw 3-pronged .................................................................................................................... 53

53. Scutum pentagonal, with prominent angulate posterior margin, palpal tarsus with 7B or 7BS, galeal setae branched or nude, fCx = 1.1.(1–9), mastitarsala I frequently present ....................................... ......................................................................... Miyatrombicula Sasa, Kawashima & Egashira, 1952

– Scutum subpentagonal, with prominent rounded posterior margin, palpal tarsus with 7BS, galeal setae branched, legs long and thin, Ip = 883–1002, 1–2 mastitarsalae III present. Parasites of reptiles and birds .......................................................... Pentidionis Vercammen-Grandjean & Loomis, 1967

– Scutum subhexagonal, wider than long, with rounded posterior margin, palpal tarsus with 7B, galeal setae nude or branched, 1–2 mastitarsalae III present or absent, mastitibiala sometimes present .................................................................. Neotrombiculoides Vercammen-Grandjean, 1960

– Scutum subpentagonal, subquadrate, subhexagonal or subtrapezoidal, with rounded posterior margin, palpal tarsus with 7BS, galeal setae branched or nude, mastisetae present or absent ....... 54

54. Galeal setae branched, fPp = B/B/NNB, scutum almost as wide as long, sensillary bases far anterior to PL, 3 genualae I, 1 mastitarsala III present or absent ..................................................................... ............................................................ Afrotrombicula Kolebinova & Vercammen-Grandjean, 1978 – Galeal setae branched or nude, scutum wider than long, sensillary bases usually at level of PL, slightly anterior or posterior of PL, 2–3 (rarely 1) genualae I, mastitarsala III usually present, additional 1–2 mastitarsalae, 1–2 mastitibialae, and 1 mastifemorala sometimes present ................. .................................................................................................................. Neotrombicula Hirst, 1925