P1;

BULLETIN DE L'INSTITUT ROYAL DES SCIENCES NATURELLES DE BELGIQUE ENTOMOLOGIE, 69: 35-86, 1999 BULLETIN VAN HET KONINKLIJK BELGISCH INSTITUUT VOOR NATUURWETENSCHAPPEN ENTOMOLOGIE, 69: 35-86, 1999 1 G Further observations on the Cheyletidae (), with a key to the genera of the Cheyletinae and a list of all known species in the family

by U. GERSON, A. FAIN and R.L. SMILEY

Summary the Philippines; GERSON (1994) for Australia] were pub­ lished. However, no recent revisions or summations of A revised key to separate the subfamilies of the family Cheyletidae is the family are at hand, nor is a key to all described genera provided and the subfamily Chelonotinae is newly defined. The genera of the Cheyletidae are defined, their habitats noted, and all known available to interested students. Furthermore, the relevant species are listed. A key to separate the genera of the Cheyletinae is literature is scattered in many, sometimes hard-to-get presented. Problems relating to some ofthe genera are discussed but no periodicals, placing yet another onus on the students. In new taxa are proposed. a previous paper (FAIN et al., 1997) we studied. the solenidiotaxy and chaetotaxy of the Cheyletidae and provided a key to its eight subfamilies. We also revised Introduction seven of these subfamilies and included keys to all their genera. The present paper deals with the largest subfam- The family Cheyletidae LEACH, 1815, may roughly be ily, the Cheyletinae, includes briefdiagnoses ofall genera separated into two assemblages, the free living predators and provides a key for them. We also list all species in the and the associates, of which many are parasites. , family. Although the dividing line between these groups is in- \ SUMMERS and PRICE (1970) remarked on the large determinate, the obviously parasitic taxa can be recog- r,umber of monospecific genera in the family, believing nized by bizalTe modifications to their mouthparts and }hat this may reflect on the small sample of the world legs and by their reduced, or enhanced, setation (hypo- cheyletid fauna which had been collected. Their hypoth­ trichy or neotrichy). Animal-associated species occur on / esis is strongly supported by two developments. Addi- birds, mammals or , and may damage birds, tional species in many of those genera are now known, cats and dogs. Further, some induce allergies and papular and most ofthe genera,described in the intervening period dermatitis in humans (BRONS"':I~K a~d DE K~EK, 1976; are I~onospecific. MaIl\of those ~ener~ were collected KEH et al., 1987). Most paraSItIC, hIghly-denved forms off vertebrates, suggestllig that theIr umque body forms have been assigned to seven subfamilies (FAIN et al., may represent specific adaptations. Most radiation, how- 1997). The predatory forms possess a more generalized ever, seems to have taken place in the less specialized body. They occur on plants, in the soil, in stored products genera, e.g. Cheyletlls with over 65 named species, Hemi- and vertebrate nests, feeding on many and small cheyletia with ca 35. insects. A few cheyletids are considered to be biological The Cheyletidae are best recognized by their fused control ~gents, and one, Cheyletus eruditus (SCHRANK), is chelicerae and the robust gnathosoma with a palpal Etlin!}used in commerce (GERSON and SMILEY, 1990). The thumb-claw complex (Fig la). The palpal tarsus often -'Cheyletidae are cosmopolitah in distribution.' carries two comb-Eke (Fig lb) and/or two sickle-like The modern taxonomic edifice of the Cheyletidae has setae as weU as a minute solenidion (Fig lc). The palpti- been erected by BAKER (1949), VOLGIN (1969; English bial claws (with or without teeth) are usually oriented 1987), and o/Y SUMMERS and PRICE (1970). "horizontally. The palpfemur is the largest segment and is robust construct, bJ6adened and deepened during the elbowed in midsection. The segmented peritremes are lUt1erveninQ' years, has remained vigorous to this day. located on the stylophore; their individual segments are UIylJ\1EItS and PRICE (1970) counted ca 50 genera and called links. SUMMERS and PRICE (1970) named the region 190 species inthe family; the total has since risen behind the peritremes "tegmen" and the area in front of 75 genera and to more than 400 species. Of them "protegmen" (Figs la and 35). The dorsum of the are the dozen vertebrate-associated genera Cheyletidae may, or may not, carry a pair ofanterior eyes by FAIN (for their revision, sye FAIN et al., (Figs 27a; 31; 45b). The dorsum is usually covered by one regional treatments [TsENG (1977) for or more plates or shields (Fig 1a), which bear setae n~ ._ (1988 and former papers) for showing a bewildering diversity of form. Some are sim- 36 U. GERSON et at. pIe, slender (Fig l6b; 17; 21; 23), others are lanceolate or us to present the first-ever figure ofD. polylepis (Fig. 45) spatulate (Fig la); sometimes they occur in the shape of and to provide additional data on both genera. Dubini­ fans (fan-like or shell-like, Figs 12; 14; 28; 51), some niola has well-developed eyes. The peritremes are appear squamiform (Figs 26 and 55), and there may also slightly arched, with four links on each side. All tarsi be setae which look like the horns of stags (Figs 15, 49 carry a unique, large and sehllose, branched seta and two and 53). In most cases the shape of the dorsal setae is claws, those on leg I are minute. There are five pairs of relatively uniform for each species, although they may be delicately-setose genital setae, two pairs of similar anal heteromorphic or dissimilar (Figs 15; 26; 48). Tarsus I setae, and two extra pairs of apparently-neotrichous, carries a solenidion (col) which is often accompanied by a post-anal, squamate setae. Thewkachela (formerly placed guard seta (Fig le). Solenidia (without guard setae and in the subfamily Chelonotinae, along with Chelonotus, much reduced in size) also occur on tarsus Il; tibiae I and Muricheyla and Pl'Omuricheyla) is similar to the latter Il (

- Tarsi IV with empodia but without claws; dorsum 2. (1) Palpal tarsus with 1-2 comb-like setae. .. .. 5 with three median shields; idiosoma distally with - Palpal tal;SUS without comblike setae 3 lobe-like projections...... Teinocheylinae 3. (2) Palpal claw with teeth; body ovoid;' dorsum FAIN; only one genus, Teinocheylus FAIN (Fig. 63) with one dorsal median shield and several ultralong 4. (3) Idiosoma without contiguous dorsal shields cov­ setae 4 ering or overlapping part of the ventral hysterosoma - Palpal claw edentate; body fusiform; without dor- ...... 5 sal median shields, without ultralong setae . - Idiosoma with very large contiguous dorsal shields ·...... Atarsacheylus THEWKE (Fig. 17) covering or overlapping part of the ventral hystero- 4. (3) Palpal claw with a single tooth; palpal tarsus soma Chelonotinae VOLGIN; with a single stiff seta bearing vestigial teeth ..... only one genus, Chelonotus BERLESE (Fig. 64) ·...... Cheletoides OUDEMANS (Figs 3-4) 5. (4) Gnathosoma modified, with ventral basal hooks - Palpal claw with 2-7 basal teeth; stiffseta on palpal and/or lateral hooks on palpal segments, or palpal tarsus without vestigial teeth ...... segments reduced; solenidion 01 on genu I replaced · Metacheletoides FAIN (Fig. 2) by a stellate seta 6 5. (2) Tarsi III-IV without strong dorsal triangular pro- - Gnathosoma unmodified; without basal hooks (ex­ cesses 7 cept in Apodicheles, Ornithocheyletiinae, which has - Tarsi III-IV with strong dorsal triangular pro- ventral hooks at the base of the gnathosoma); palpal cesses 6 segments normal, never reduced; genu I with soleni- 6. Palpal tarsus with 1 comb-like seta . dion crI 7 ·...... Muricheyla FAIN (Fig. 10) 6. (5) Gnathosoma with very large lateral hook-like - Palpal tarsus with 2 comb-like setae . processes; palpi small, narrow, their tibiae and tarsi ·...... Promuricheyla FAIN (Fig. 9) fused, forming a short segment with a thick comb­ 7. (5) Palpal tarsus with 2 comb-like setae " 13 like seta; idiosoma and legs without processes; dor­ - Palpal tarsus with 1 comb-like seta 8 sum without neotrichial setae .. .. Criokerontinae 8. (7) Coxae I-ll without lateral or posterior lobes. 9 SMILEY; only one genus, Criokeron VOLGIN (Fig. 79) - Coxae I-ll with lateral or posterior lobes . - Gnathosoma not as above; palpal tibial claw ·...... Thewkachela IDE and KETHLEY (Fig. 8) strongly hooked; without comb-like setae; palpi, 9. (8) Dorsum with a propodosomal and a hysteroso- idiosoma and legs with processes; dorsum may mal shield 11 bear neotrichial setae . - Dorsum with only with a propodosomal shield ...... Niheliinae SMILEY (Figs 72-75) · " JO .." 7. (5) Gnathosoma poorly developed, but palpal claw 10. (9) Coxae III and IV separated; solenidion CDl lack- strongly curved; no combclike setae and/or eyes " ing Eucheletopsis VOLGIN (Fig. 5) ...... Ornithocheyletiinae SMILEY (Figs 66-71) - Coxae III and IV contiguous; solenidion CDl pre- - Gnathosoma well developed; palpal claw at most sent " Cheletopsis OUDEMANS (Fig. 6) moderately curved, often with comb-like setae and/or 11. (9) Hysteronotal shield large, on entire hysterono­ eyes...... Cheyletinae VOLGIN tum, bearing regular and neotrichous setae '" 12 - Hysteronotal shield small, restricted to suranal (*) The genera Alliea YUNKER (Fig. 19) and Thl)lonomy­ area, bearing only 2 pairs of slender setae . cheyla FAIN were excluded from the above and the fol­ ·...... Cheletosoma OUDEMANS (Fig. 7) lowing keys. The former because only a male (for both 12. (11) Lateral and median dorsal setae dissimilar; A. laruei YUNKER and A. prasadi CORPUZ-RAROS) and anus ventral; ilU1er comb-like seta of palpal tarsus an incomplete female (without gnathosoma, for A. laruei) replaced by a small spiculate seta . were available, the latter because the female is un­ · Camincheyletus SMILEY and WHITAKER (Fig. 13) known. - All dorsal setae similar; anus on a caudal lobe; inner comb-like seta of palpal tarsus replaced by a small smooth seta .. Subfamily CheyIetinae FAIN ·...... Caudacheles GERSON (Fig. 14) 13. (7) Body ovoid, peritremes arched forward or with The Cheyletinae is the largest subfamily in the family short transverse arms, curving and descending back- Cheyletidae. Its members occur on plants, in the soil, in wards !vert .J.M.., ..s1~'V~) " 14 bee hives, in the nests ofrodents and birds, on the bodies - Body elongated, pentremes M-shaped . of arthropods and commonly occur in stored food. The ...... /vIe&-w.. Bak YUNKER (Fig. 18) following key was constructed only to facilitate the se­ 14. (13) Dorsum with 2A-shields, one on propodosoma, paration ofthe genera, and does not reflect any perceived other on hysterosoma, peritremes arched or with relationships. short transverse links 15 - Dorsum with only 1 shield, on propodosoma; peri­ 1. With one pair of eyes ...... 18 tremes not arched but with short transverse links .. - Without eyes 2 ·...... Cheletonella WOMERSLEY (Fig. 11) 38 U. GERSON et al.

15. (14) Tarsus I with claws; palpal claw with at least 1 - Humeral setae ultralong . tooth, peritremes not arched forward, with short ·...... Acaropsellina SUMMERS (Fig. 29) transverse links, descending backwards ...... 16 ~ 29. (18) Dorsum with at least one shield 30 - Tarsus I without claws; palpal claws toothless; - Dorsum without shields .'. peritremes arched fOlward, curving backwards ... ·...... Chelacheles BAKER (*) (Fig. 21) ·...... Eutogenes BAKER (Fig. 12) 30. (29) Dorsum with 2 or 3 shields ...... 32 16. (15) Peritremes M-shaped or forming an inverted - Dorsum with 1 shield '. .. 31 U; lateral dorsal setae slender to narrowly-spatu- 31. (30) Palpal claw toothed only along basal half; dor- late 17 sal setae mostly rodlike . - Peritremes usually with 3 short transverse links; ·...... Philippicheyla CORJ'UZ-RAROS (Fig. 31) lateral dorsal setae fan-like . - Palpal claw toothed along entire inner margin; · Eucheyletia BAKER (Fig. 15) dorsal setae mostly fanlike . 17. (16) Posterior peritremallink abmply bent inwards; ·...... Cheletacarus VOLGIN (Fig. 32) tibial claw with 4-7 teeth . 32. (30) Dorsum with 2 shields 36 ·...... Hylopecheyla FAIN (Fig. 16) - Dorsum with 3 shields...... 33 - Posterior peritremal link not abmply bent turned 33. (32) Palpal claw toothed 34 inwards; tibial claw with 1-4 teeth . - Palpal claw edentate . ·...... Cheyletus LATREILLE (Fig. 1) · Aegyptocheyla YOUSEF (Fig. 33) j.' 18. (1) Palpal tarsus with 2 comb-like setae 29 34. (33) Each hysteronotal shield with 1-2 setae.. 35 - Palpal tarsus with 1 comb-like seta 19 - Each hysteronotal shield with 5-7 setae . 19. (18) Dorsum with at least 1 shield 21 ·...... Oudell1ansicheyla VOLGIN (Fig. 34) - Dorsum without shields, entirely striated 20 35. (34) All tarsi with long solenidia (subequal in length 20. (19) Body ovoid, coxae Il and III separated by to seglnent) . less than body width . ·...... Paracheyletiella KUZNETZOV (Fig. 36) ·...... Chelachecaropsis ATTIAH (Fig. 20) - Only tarsi 1-Il with solenidia;. if solenidia present - Body fusiform, coxae II and III separated by on tarsi Il-IV, then shorter than width of segment . about body width . ·...... Cheletoll1ill1us OUDEMANS (Fig. 35) ·...... Chelacheles BAKER (*) (Fig. 21) 36. (32) Palpal claw with 1 or more teeth...... 40 21. (19) Dorsum with propodosomal and hysteronotal - Palpal claw edentate ,.... 37 shields '...... 22 37. (36) With sickle-like setae; median dorsal setae si­ - Dorsum only with a propodosomal shield . milar to lateral setae ...... 38 ·...... Chelacaropsis BAKER (Fig. 22) - Without sickle-like setae; median dorsal setae dis- 22. (21) Median hysteronotal shield large, bears several similar to lateral setae . setae 24 ·. (: .. Colull1bicheyla THEWKE and ENNS (Fig. 37) - Median hysteronotal shield small, without setae 38. (37) Leg I with claws :...... 39 · 23 - Leg I without claws . 23. (22) Dorsal setae spatulate to fan-like; humerals ·...... Chiapacheylus DE LEoN (Fig. 38) similar to other dorsal setae . 39. (38) Dorsal body and tegmen with coarse reticula- ·...... Cheletonata WOMERSLEY (Fig. 25) tion Ker MUMA (Fig. 40) - Dorsal setae slender; humeral setae ultralong ... - Dorsal body and tegmen without reticulation ... ·...... Paracaropsis VOLGIN (Fig. 23) · Pavlovskicheyla VOLGIN (Fig. 39) 24. (22) All tarsi with empodia ;md claws 25 40. (36) Palpal claw with more than 3 teeth ... .. 46 - All tarsi with empodia but' without claws . - Palpal claw with 1-3 teeth .. .'...... '41 ·. Parall1icrocheyla OLIVIER and THERON (Fig. 28) 41. (40) Leg I shorter (ca 70%) than idiosoma ... 43 25. (24) Body ovoid, coxae II and III separated by less - Leg I as long as idiosoma or longer ...... 42 than body width '...... 26 42. (41) Dorsal shields separated; claws on leg I -Body fusiform, coxae Il and III separated by more minute or absent '.. than body width " . .. '.'...... Cheletoll1orpha OUDEMANS (Fig. 41) ·. Neochelacheles SMILEY and WILLIAMS (Fig. 27) - Dorsal shields indeterminately separated; claws on 26. (25) Claws on legs Il-IV with smooth hooks.. 27 leg I normal ...... Nodele MUMA (Fig. 42) - Claws on legs II-IV with a basal pi'ocess . 43. (41) Hysteronotal shield covers most of hysteroso­ · Neoacaropsis VOLGIN (Fig. 24) ma; peritremes forming an inverted U...... 44 27. (26) Claws on all tarsi ofregular size; dorsal lateral - Hysteronotal shield confined to opisthosomal re- and median setae similar 28 gion; peritremes M-shaped . - Claws on all tarsi minute or absent; dorsal and · ,. .. Cheletophyes OUDEMANS (Fig. 43) lateral setae dissimilar . 44. (43) With a §rn~l~'ge basal tooth on palpal claw · Microcheyla VOLGIN (Fig. 26) · " 45 28. (27) Humeral setae similar to dorsal setae . - With 2 basal teeth on palpal claw . ·...... Acaropsella VOLGIN (Fig. 30) · Polycheyletus VAIVANIJKUL (Fig. 44) Further observations on the Cheyletidae 39

45. (44) Dorsum with up to 21 pairs of setae; median 60. (59) Dorsal setae mostly fan-like; humeral setae on dorsal and lateral setae dissimilar . dorsum .....'.. Hel11icheyletia VOLGIN (Fig. 61) ·...... Hoffinannita PELAEZ (Fig. 48) - Dorsal setae rod-like; humeral setae on pleuroven- - Dorsum with more than 30 pairs of setae; dorsal trally displaced platelets . median and lateral setae similar . . .. Laeliocheyletia SUMMERS and PRICE (Fig. 62) · " Dubininiola VOLGIN (Fig. 45) 46: (40) Gnathosoma not overhung by propodosoma (*) Due to different definitions ofChelacheles, the genus · 47 is keyed out twice; see remarks to this genus. - Gnathosoma partly covered by overhanging propo- dosoma : Sal11sinakia VOLGIN (Fig. 46) 47. (46) Inner sickle-like seta on palpal tarsus normal, The genera and species of the Cheyletidae seta-like...... 49 (*species of uncertain standing; - Inner sickle-like s~ta on palpal tarsus inflated subspecies not listed) · 48 48. (47) Posterior links of peritremes straight, devoid I. Genus Acaropsella VOLGIN 1962 of vesicular chamber . ( f. J .3,,) ·...... NeoeucheylaRADFoRD (Fig. 52) Type species: Neoacaropsis rohdendorji VOLGIN 1962 - Posterior links of peritremes straight or looped, terminating at vesicular chambers . Diagnosis: Eyes present; palpal tarsus with 1 comb-like. ·...... CunlifJella VOLGIN (Fig. 47) seta and 2 sickle-like setae; palpal claw with more 49. (47) Claws on leg I normal, similar to claws on other than 3 teeth; peritremes with at least 3 links; body legs .... " ..... : ... '...... 53 ovoid; dorsum with a propodosomal and a hysteronotal - Claws on leg I either minute, smaller than other shield, both with legs claws, or absent ...... 50 II and III width; all legs 50. (49) Claws on leg I minute 52 shorter than body; all tarsi with claws and empo- - Claws on leg I absent ...;...... 51 dia. 51. (50) Tarsus I with 2 conspicuous terminal setae ...... Cheletogenes OUDEMANS (Fig. 51) Other species - Tarsus I with 4 conspicuous terminal setae ..... A. aegyptiaca (WAFA and SOLIMAN) 1968 · Prosocheyla VOLGIN (Fig. 50) A. jilipina CORPUZ-RAROS 1988 52. (50) Leg I shOlter (ca 70%) than idiosoma . A. kinshasensis FAIN 1972 · Paracheyletia VOLGIN (Fig. 49) A. konoi TSENG 1977 - Leg I as long as idiosoma or longer . A. kulagini (RHODENDOIti') 1940 ·...... Mexecheles DE LEON (Fig. 53) A. nobilis RASOOL et al., 1980 53. (49) All anterior prodorsal setae similar .. . .. 54 A. schl11idtl11anni PRICE 1972 "'~ - Anterior prodorsal setae dissimilar ...... ~ A. volgini (GERSON) 1967 ~7J I' ·...... Grallacheles DE LEON (Fig. 54) '(' A. 'l1-Wh."{o..AA1.r-.v>V. X( "'- B"", et; "1' '111- 54. (53) Dorsal lateral setae similar to dorsal median Habitat: Stored products, birddnests, soil. Cosmopoli- setae 56 tan. <' - Dorsal lateral setae dissimilar to dorsal median setae 55 55. (54) Palpal claw toothed only along basal half 2. Genus Aca,ropsellina SUMMERS 1976 E • ...... •COf';.e,.~.,.6-.• P. h'H.'"(,Lu ..'(F'Ig.. 5tl.) (= Acaropsls MOQUIl'l-TANDON 1862) - Palpal claw toothed along entire length ' .. (, '. ( ~t· <-9 ) · K~(.'~'iiY~ ~ (Fig. 55) Type species: Acaropsis sollers RHODENDORF 1940 56. (54) Leg I shorter than idiosoma (ca 70%); no con­ centric striae around dorsal setae...... 57 Diagnosis: Eyes present; palpal tarsus with 1 comb-like - Leg I as long as idiosoma; some dorsal setae seta and 2 sickle-like setae; palpal claw with more than 3 surrounded by concentric striae . teeth; peritremes with more than 3 links; body ovoid; ·...... Cheletophanes OUDEMANS (Fig. 57) dorsum with a propodosomal and a hysteronotal shield, 57. (56) Hysterosoma rounded posteriorly 58 both with slender to spatulate setae; ~. - Hysterosoma tapering posteriorly . 1~~ll~t.~~~~~('·; legs II and III separated y ess a · Lepidocheyla VOLGIN (Fig. 58) ; all legs shorter than body; all tarsi with smooth 58. (57) Anus not placed on a caudal lobe 59 claws an'cl empodia. Anus placed on a caudal lobe ...... Anthribicheyla THEWKE (Fig. 59) Other species Peritremes forming an inverted U...... 60 A, anarsia SUMMERS 1976 I:'eriltrelnes M-shaped . A. clal110 QAYYUM and CHAUDHRI* 1979 Tutacheyla CORPuz-RAROS (Fig. 60) A. docta (BERLESE) 1886 40 . U. GERSON et al.

4.n~~~ft--- r -++--'!V-==*!-- pte 9 Pf -~---+-~te9

e

~--'l~d;-:;2~----rr;--I--· hs

f

Fig. 1 - Cheyletlls misol1l1ei FAIN and LUKOSCHUS. Female, dorsum (a); palpal tarsus and tibia, dorsal (b) and ventral (c); gnathosoma, ventral (d); tarsus I, dorso-lateral (e); genito-anal region (t). Abbreviations: proponotal shield (ps); hysteronotal shield (hs); palpal tarsus (pt); palpal tibia (pti); comb-like setae (cs); palpal claw (Pcl); sickle-like setae (ss); tegmen (teg); protegmen (pteg); palpal femur (pt); rostrum (1'); solenidion omega of tarsus I: (w); guard seta (gs) (from FAIN and LUKOSCHUS, 1981a). t

A. opsis RASOOL et al., * 1980 A. S1l111111ersi SMILEY and WHITAKER 1981 A. orbis QAYYUM and CHAUDHRI* 1979 A. tyrophaglls ELBADRY* 1969 A. philippinensis (CoRPuz-RAROS and SOTTO) 1977 A. vitrlls AHEER, AKBAR and CHAUDHRI* 1991 A. platessa AHEER et al., * 1991 A. porta QAYYUM and CHAUDHRI* 1979 Habitat: Stored products, plants, flying squirrels (Glallc­ A. shorkotiensis AKBAR, et al. * 1988 0111YS). Cosmopolitan. Further observations on the Cheyletidae 41

,j . Genus Aegyptocheyla YOUSEF 1978 Habitat: Anthribid beetles. USA. ({';;} 33) Type species: Aegyptocheyla $ummersi YOUSEF 1978 C. Genus Apodicheles FAIN 1979 Diagnosis: Eyes present; pa1pal tarsus with 2 comb-like (-&.:-* 1'1 ) setae and 2 sickle-like setae; pa1pa1 claw edentate; peri­ Type species: Apodicheles cypsiurus FAIN 1979 tremes with more than 3 links; body ovoid; dorsum with a single propodosoma1 shield and a pair of hysteronota1 Diagnosis: Eyes absent; without comb-like setae but with shields, placed side-by-side, all with fan-like setae; hum­ 1-2 sickle-like setae; tibial claw edentate; each palpal era1s alike; legs II and III separated by less than body femur bears 2 pairs of retrorse, ventral processes; peri­ width; all legs shorter than body; all tarsi with smooth tremes M-shaped, with more than 3 broad links; body claws and empodia. ovoid, dorsum with an ill-defined, finely striated shield which extends from legs II to beyond legs IV; dorsal setae Habitat: Plant. Egypt. slender, barbed; some, including humerals, ultralong; anal setae reduced to one pair or missing; legs II and III separated by less than body width; all legs shorter than i.f. Genus Alliea YUNKER 1960 body; coxae III and IV with 1 seta; all tarsi with smooth (~if·19) claws and empodia; tibia I without solenidion; coxae III Type species: Alliea laruei YUNKER 1960 and IV with 1 seta.

Diagnosis: Eyes absent; gnathosoma not seen; body Other species ovoid, dorsum with large propodosomal and hysteronotal A. apus FATN 1979 shields, each bearing 16 pairs ofsquamatiform or fan-like A. hetempalpus MEGNIN* 1878 setae; humerals similar; legs II and III separated by less than body width; all legs shorter than body; all tarsi with Habitat: Birds (swifts). Africa. smooth claws and empodia, claws on tarsus I slightly smaller. ? Genus Atarsacheylus THEWKE 1980 Habitat: Rats, stored food. U.S.A., The Philippines. (~J /1 ) Type species: Atarsacheylus vichii THEWKE 1980 Other species A. prasadi CORPuz-RAROS 1998 Diagnosis: Eyes absent; pa1pa1 tarsus absent (thus lacking comb-like and sickle-like setae); palpal claw edentate; Remarks: The above diagnosis is based only on a single peritremes with more than 3 links; body fusiform, legs II damaged female specimen, and the lack ofthe gnathoso­ and III separated by more than body width; dorsum ma did not allow us to place this genus in the key. devoid of shields, all dorsal setae slender, barbed; hum­ Nevertheless, there are sufficient characters to separate era1s alike; all legs shorter than body; all tarsi with Alliea from two other cheletine genera, namely Euto­ smooth claws and empodia. genes and Caudacheles, which also lack eyes and bear numerous fan-like setae on their extensive dorsal shields. Habitat: Tree bark. USA. Tarsus I of Eutogenes is devoid of claws and bears four long setae, whereas the anus ofCaudacheles is borne on a projecting caudal lobe. The only other named species, A. 8 Genus Bak YUNKER 1961 prasadi, is likewise known only as a male, and like A. {f:J. t8 j laruei, it bears an inflated seta on the palpal tarsus. Type species: Bak sanctaehelenae YUNKER 1961

Diagnosis: Eyes absent; pa1pa1 tarsus with 2 comb-like " ... Genus Allthribicheyla THEWKE 1980 setae and 2 sickle-like setae; pa1pa1 claw with 1-3 teeth; CR.} '''1) peritremes M-shaped, with more than 3 links; body fusi­ Type species: Anthribicheyla bocld THEwKE 1980 form, legs II and III separated by more than body width; dorsum with one propodosoma1 shield, hysteronotum Diagnosis: Eyes present; palpal tarsus with 2 comb-like with several platelets; all dorsal setae slender or barbed; setae and 2 sickle-like setae; palpal claw with more than 3 humera1s similar; all legs shorter than body; all tarsi with teeth; peritremes with more than 3 links; body ovoid, smooth claws and empodia. dorsum with a propodosomal and a hysteronotal shield, both with spatulate, barbed setae; humerals similar; anus Other species on stalked caudal lobe; legs II and III separated by less B. deleoni YUNKER 1961 than body width; all legs shorter than body; all tarsi with B. elongatus PATXOT and GOFF 1985 smooth claws and empodia. B. jit/'catus GERSON and FAIN 1991 42 U. GERSON et al.

2

a

------~ -~ a2 al

3 a

c

Figs 2-4 - Fig. 2. Metacheletoides Ilumidae FAIN. Female, dorsum (a) and venter (b); palpal tarsus and tibia, ventral (c) and dorsal (d) (this and the next Fig. from FAIN, 197ge). Fig. 3. Cheletoides chirullduellsis FAIN. Female,dorsum (a); palpal tarsus and tibia, ventral (b) and dorsal (c). Fig. 4. Cheletoides ullcinatus (Heller). Female, dorsum (a) (from OUDEMANS, 1906) (a); palpal tarsus and tibia, ventral (b) and dorsal (c) (from FAIN, 197ge). Further observations on the Cheyletidae 43

b

Figs 5-7 - Fig. 5. ElIcheletopsis major (TROUESSART). Female, dorsum (a) and venter (b); palpal tarsus and tibia, dorsal (c); apex oftal'SUS I (d) (this and the next two Figs from OUDEMANS, 1906). Fig. 6. Cheletopsis l1ol'lleri (Poppe). Female, dorsum (a) and venter (b); palpal tarsus and tibia, dorsal (c); apex oftarsus I (d); a dorsal seta (e). Fig. 7. Cheletosoma tyral1l1l1s OUDEMANS. Female, dorsum (a) and venter (b); palpal tarsus and tibia, dorsal (c); apex of tarsus I (d).

B. ligysnttatus FLECHTMANN 1979(**) (**) May be an illegal name because published only in a B. micidus SUMMERS and PRICE 1970 thesis.. Dzarkensis THEWKE and ENNS 1974 payatus CORPUZ-RAROS and SOTTO 1977 Habitats: Decomposing plants, soil, bee hive debris; cos­ truncatus CORPuz-RAROS and SOTTO 1977 mopolitan. .t~ Cofr-2..,fWu.-j .2"""" . (J <:J)...S ~ ur-R. "-"k.l 02.0" 0 44 U. GERSON et al.

\~.. , -e ..JV

c d

Figs 8-10 - Fig. 8. Thewkachela ratufi IOE and KETHLEY. Female, dorsum (a); leg 1 (b); coxae I-IT (c); palpal tarsus and tibia, venter (d) (f1'Om IOEand KETHLEY, 1977). Fig. 9. Pl'Ol11uricheyla lukoschusi FAIN. Female, dorsum (a); palpal tarsus and tibia, dorsal (b); ventl'al (c); tarsus 1 (d); tarsus IV in lateral view (e) (this and the next Fig. f1'Om FAIN, 19791). Fig. 10. Muricheyla sicista FAIN. Female, dorsum (a); palpal tarsus and tibia, dorsal (b); leg I, dorsal (c); tarsus III, lateral (d); leg IV, lateral (e). Further observations on the Cheyletidae 45

1 Genus Bakerichey[a VOLGIN 1966 If 1.. Genus Caudache[es GERSON 1968 (~}j (,~- 0g ) (~J .Ilf) Type species: chanayi BERLESE and TROUES­ Type species: Caudache!es khayae GERSON 1968 SARt 1889 Diagnosis: Eyes absent; palpal tarsus with 1 comb-like Diagnosis: Eyes absent; palpal tarsus devoid ofcomb-like seta, other seta short, smooth, and 2 sickle-like setae; setae but with 1-2 sickle-like setae; tibial claw edentate; palpal claw edentate; peritremes with 2-5 links; body no retrorse processes on palpal femora; peritremes with ovoid, legs II and III separated by less than body more than 3 links; body ovoid, legs II and III separated by width; dorsum carries a propodosomal and a hystero­ less than body width; dorsum with a delicate propodoso­ notal shield, both neotrichous; all dorsal setae similar, mal shield which may be discernible only by more deli­ fan-like; humerals alike; anus on caudal lobe; all legs cate .striae; dorsal setae slender, some, including hum­ shorter than body; all tarsi with smooth claws and empo­ erals, ultralong; all legs shorter than body; coxae III and dia. IV contiguous, coxa IV with 2 setae; tibia I with soleni­ dion; all tarsi with smooth claws and empodia. Other species C. lieni TSENG 1977 ,th'Z..~t<,-d..~ ~ I-~ 1~'i1 Other species c. Lv:, 7L'w,,- ..R-, B, aji'icana FAIN 1979 Habitat: Plants. Israel, Taiwan. B, benoW FAIN 1980 B.faini (LAWRENCE) 1959 B. subquadrata (LAWRENCE) 1959 ~~ Genus Che[acaropsis BAKER 1949 B. transvaalica (LAWRENCE) 1959 ( /c'9-' 2 2- ) Type species: Che!acaropsis 1110!Lei BAKER 1949 Habitat: Birds. Cosmopolitan. Diagnosis: Eyes present; palpal tarsus with 1 comb-like seta, other seta similar to the 2 sickle-like setae; palpal 10 Genus Bichey[etiella FAIN 1972 claw with 3 teeth; peritremes with more than 3 links; ( ~(~ ?-1-) body ovoid, legs II and III separated by less than body Type species: Bicheyletiella r0711erolagi FAIN 1972 width; idiosoma carries only a propodosomal shield; all dorsal setae spatulate-barbed, humerals similar or slightly Diagnosis: Eyes absent; palpal tarsus without comb-like longer; all legs shorter than body, all tarsi with smooth setae but with 2 rodlike setae, one forked terminally, other claws and empodia. rod-like; palpal claw edentate; peritremes with more than 3 links; body ovoid; legs II and III separated by less than Other species body width; dorsum with a propodosomal and a wider C. apus FAIN 1972 than long hysteronotal shield; all dorsal setae slender, C. reticulata SOLIMAN 1975 ( nude or barbed; humerals alike; all tarsi without claws C. nvandana FAIN 1972=-= but bear feather-like empodia; tibia I lacks solenidion cp. C. terrestris CORPuz-RAROS and SOTTO 1977.,.,.,... Habitat: On R0711erolagus diazi. Mexico. Habitat: Glaucomys; nests of rodents and birds; food stores; soil. USA, Asia, Africa.

1'1 Genus Call1illchey[etus SMILEY and WHITAKER 1981 Remarks: The concept of this genus was changed by (f-\J 13) LEKPRAYOON and SMILEY (1986), who examined the types Type species: Ca711incheyletus glauco711ys SMILEY and of Chelacaropsis moorei and noted that, contrary to the WHITAKER 1981 original description (BAKER, 1949), the female bears a propodosomal shield. This suggests that the four other Diagnosis: Eyes absent; palpal tarsus with 1 comb-like species presently placed in Chelacaropsis should be ac­ seta, other seta short, spiculate, and 2 sickle-like setae; commodated elsewhere. palpal claw with 1 tooth; peritremes with more than 3 links; body ovoid, legs II and III separated by less than body width; with a propodosomal and a hysteronotal 1'1 Genus Che!achecaropsis ATTlAH 1973 shield that cover most of dorsum, with dissimilar setae: C~J <0) laterals spatulate, medians squamate; more than 10 pairs of Type species: Chelachecaropsis bakeri ATTIAH 1973. neotrichous setae on hysteronotum; humerals like lateral setae, on separate pleuroventral platelets; all legs shorter Diagnosis: Eyes present; palpal tarsus with 1 comb-like than body; all tarsi with smooth claws andempodia. seta, other seta similar to the 2 sickle~like setae; palpal claw with more than 3 teeth; peritremes with more than Habitat: Nest of flying squirrels, Glauco711ys, USA. 3 links; body ovoid, legs II and III separated by less 46 U. GERSON et al.

Figs 11-13 - Fig. 11. Cheletonella vespertilionis WOMERSLEY. Female, dorsum (this and the next Fig. from SUMMERS and PRICE, 1970). Fig. 12. Eutogenesfoxi BAKER. Female, dorsum. Fig. 13. Camincheyletus glaucomys SMILEY and WHITAKER. Female, dorsum (a) and venter (b) (from SMILEY and WHITAKER, 1981). Further observations on the Chey1etidae 47 than body width; idiosoma striated, without shields; all hysteronotal shield a single propodosomal shield bearing dorsal setae slender to barbed, humerals ultralong; all spatulate or lanceolate setae; humerals similar; all legs legs shorter than body; all tarsi with smooth claws and shorter than body; all tarsi with smooth claws and empo­ empodia. dia.

Other species Other species C. stigmaeoides BARILO 1989 C. gryphus SUMMERS and PRICE 1970 C. novemdentis MEYER 1988 Habitat: Rice mill, soil. Egypt; Uzbekistan. C. ploceus GUPTA and PAUL 1985 C. rugosus (WOMERSLEY) 1941

le; Genus Chelacheles BAKER 1958 Habitat: Trees, associated with scale insects (Homoptera: (-(-.'g .2.1) Cocoidea) or beetles; birds' nests. Cosmopolitan. Type species: Chelacheles strabismus BAKER 1958. Remarks: SUMMERS and PRICE (1970) considered the Diagnosis: Eyes present; palpal tarsus with 2 comb-like placement of C. rugosus in Cheletacarus as provisional, setae; one ofthe comb-like setae may be similar to the 2 because part ofthe type specimen was damaged. Another sickle-like setae; palpal claw with 3 or more teeth; peri­ reason for reconsidering the status of this species is the tremes with more than 3 links; body fusiform, legs II and form of the two anterior propodosomal setae, which are III separated by about body width; dorsum striated, with­ dissimilar to other dorsals and much longer. out shields; all dorsal setae slender, may be barbed, humerals same shape but ultralong; all legs shorter than body; all tarsi with smooth claws and empodia. Iq Genus Cheletogenes OUDEMANS 1905 (-#...~ .:,-, ) Other species Type species: Cheylus ornatus CANESTRINI and FANZAGO C. alexandrinus HASSAN and GOMAA 1981 1876. C. bacchusi BOCHKOV, HAUSTOV and KUZNETZOV, 1999 C. baiwanganae CORPUZ-RAROS and SOTTO 1977 Diagnosis: Eyes present; palpal tarsus with 2 comb-like C. bipanus SUMMERS and PRICE 1970 setae and 2 sickle-like setae; palpal claw with more than 3 C. humilis RASOOL et al., 1980 teeth; peritremes with more than 3 links; body ovoid, legs C. lanceolatus TSENG 1977 II and III separated by less than body width; dorsum with C. michalskii SAMSINAK 1962 a propodosomal and a hysteronotal shield, both with C. peritremaculatus THEWKE 1974 similar fan-like setae; humerals alike; all legs shorter than C. robustus CORPuz-RAROS 1998 body; tarsus I without claws and empodia, bearing 2 conspicuous tenninal setae; tarsi II-IV with smooth claws Habitat: Stored products, bark beetle galleries, chicken and empodia. feathers. Europe, USA, Asia. Other species Remarks: The genus was defined (BAKER, 1958) as hav­ C. aceriai KHAN* 1970 ing 2 comb-like setae, a practice followed by CORPUZ­ C. carinatus AHEER et al., 1992 RAROS (1998). But SUMMERS and PRICE (1970) stated that C. dissitus AKBAR et al., 1988 the inner comb-like seta may be "comblike or plain", C. iconis AHEER et al., 1992 and the relevant seta of their bipamls has only "several C. meihuashanense iN and Lru 1994 exceedingly fine barbs on its convext curvature". TSENG C. monosetosus TSENG 1977 (1977) defined Chelacheles as having Olily 1 comb-like C. petiginis QAYYUM and CHAUDHRI 1977 seta. The significance of this character remains unre­ C. sagacis AHEER et al., 1992 solved. C. scaber QAYYUM and CHAUDHRI 1977 C. vulgatus RASOOL and CHAUDHRI 1979 C. }vaitei GERSON 1994

(0 Genus Cheletacarus VOLGIN 1961 Habitat; Plants. Cosmopolitan. (hr· 3 :L) Type species: Cheletacarus raptor VOLGIN 1961 13 Genus Cheletoides OUDEMANS 1904 Diagnosis: Eyes present; palpal tarsus with 2 comb-like (i-''Y' 3~Y) setae and 2 sickle-like setae; palpal claw with more than 3 Type species: Syringophilus uncinatus HelIer 1880 teeth, placed along most of palpal claw; peritremes with lUore than 3 links; body ovoid, legs II and III separated by Diagnosis: Eyes absent; palpal tarsus without comb-like less than body width; dorsum with a propodosomal but no setae, but bearing a stiffseta canying vestigial teeth and 2 48 U. GERSON et al.

16

Figs 14-16 - Fig. 14. Calldacheles khayae GERSON. Female, dorsum (from GERSON, 1968). Fig. 15. ElIcheyletia bishoppi BAKER. Female, dorsum (from SUMMERS and PRICE, 1970). Fig. 16. Hylopecheyla lIlalayi FAIN and NADCHATRAM. Female, dorsum (a); venter (b) (from FAIN and NADCHATRAM, 1980). Further observations on the Cheyletidae 49

Figs 17-19 - Fig. 17. Atal'sacheyllls vichii THEWKE. Female, dorsum (redrawn by AF. from THEWKE, 1980). Fig. 18. Bak sanctaehelenae YUNKER. Female, dorsum (from SUMMERS and PRICE, 1970). Fig. 19. Alliea lal'lIei YUNKER. Female, dorsum without gnathosoma (a); tarsus I (b) (from YUNKER, 1960).

sickle-like setae; palpal claw with a single tooth; peri­ I~. Genus Cheleto11l;11lUS OUDElVIANS 1904 tremes with more than 3 links; body ovoid, legs II and III (-?'J-. 3 !J) separated by about body width; dorsum only with a Type species: Cheletes berlesei OUDEMANS 1904 propodosomal shield, dorsal setae slender, barbed; sev­ eral, including humerals, ultralong; all legs shorter than Diagnosis: Eyes present; palpal tarsus with 2 comb-like body; all tarsi with smooth claws and empodia. setae and 2 sickle-like setae; palpal claw with more than 3 teeth; peritremes with more than 3 links; body ovoid, legs Other species II and III separated by less than body width; dorsum with Cheletoides chirllndllensis FAIN 1979. a propodosomal and 2 hysteronotal shields, placed side­ by-side, all bearing similar, lanceolate to nanowly fan­ Habitat: Birds. Europe, Africa. like dorsal setae; each hysteronotal shield with 1-2 setae; 50 U. GERSON et al.

Figs 20-22 - Fig. 20. Chelachecaropsis bakel'i ATTIAH. Female, dorsum (a); left eye (b); palpal tarsus and tibia (c) (from ATTIAH, 1973). Fig. 21. Chelacheles strabislllus BAKER. Female, dorsum (a); palpal tarsus and tibia (b) and tarsus I (from BAKER, 1958). Fig. 22. Chelacal'Opsis lIlool'ei BAKER. Female, dorsum (a) (from LEKPRAYOON and SMILEY, 1986); gnathosoma, dorsal (b) (from BAKER, 1949). humerals alike; all legs shorter than body, all tarsi with C. bisetosus TSENG 1977 smooth claws and empodia. C.cambio AHEER, AKBAR and CHAUDHRl 1994 C. cantor RASOOL, CHAUDHRl and AKBAR 1980 Other species C. citrosinensis PATXOT and GOFF 1985 C. binus TSENG 1973 C. daltoniensis CORPUZ-RAROS 1998 Further observations on the Cheyletidae 51

C. duosetosus MUMA 1964 than body, with smooth claws and empodia, claws on C. heredis QAYYUM and CHAUDHRI 1979 tarsus I minute. C. larmae AHEER et al., 1994 C. minutus SOLIMAN 1977 Habitat: Bird's nest. Australia. C. zamia AHEER et al., 1994 c. ~cJ::a AHe&,,, id- ~ //993 C. ;/;;'l-"-~ IfIiGG'I'C t4- ~ /992 Habitat: Plants, soil. Cosmopolitan. t<. Genus Cheletollella WOlVIERSLEY 1941 C"'-'I' /1'1 } Type species: Cheletonella vespertilionis WOMERSLEY ~o Genus Chelet011t01pha OUDEMANS 1904 1941 (= Achelet011l01pha VOLGIN 1969) (~{j-.l+I) Diagnosis: Eyes absent; palpal tarsus with 2 comb-like Type species: Acarus lepidopterorum SHAw 1794 setae and 2 sickle-like setae; palpal claw with 2-4 teeth; peritremes forming an inverted V or M-shaped, with more Diagnosis: Eyes present; palpal tarsus with 2 comb-like than 3 links; body ovoid, legs II and III separated by less setae and 2 sickle-like setae; palpal claw with 1 large than body width; dorsum with a propodosomal but no basal tooth; peritremes with more than 3 links; body hysteronotal shield, dorsal setae lanceolate or fan-like; ovoid, legs II and III separated by less than body width; humerals similar or dissimilar to other dorsal setae; all dorsum with a propodosomal and a hysteronotal shield; legs shorter than body, all tarsi with smooth claws and lateral and humerals setae rod-like, barbed, medians dis­ empodia. similar, staghorn-like; legs I longer than body, their tarsi with empodia, with or without minute claws; other legs Other species shorter; their tarsi with smooth claws and empodia. C. caucasica VOLGIN 1955 C. hoffinannae SMILEY 1996 Other species C. juglandis XIA et al., 1999 C. bakeri LAWRENCE 1954 C. pilosa TSENG 1977 C. dolosus AHEER et al., 1997 C. obrutus QAYYUM and CHAUDHRI 1977 Habitat: Bat guano, bird nests, soil. Australia, Russia, C. opacus QAYYUM and CHAUDHRI 1977 China. C. orientalis OUDEMANS 1928 C. tenerUl11 QAYYUM and CHAUDHRI 1977 Remarks: An application of our criteria for separating genera in the family Cheyletidae suggests that C. pilosa Habitat: Moths, stored products, plants. Cosmopolitan. and C. juglandis should be placed elsewhere. The peri­ tremes of both are M-shaped, their dorsal setae are lan­ Remarks: VOLGIN (1969) created Acheletomorpha gen. ceolate-barbed and the humerals are ultralong, dissimilar novo for C. bakeri, because its median dorsal and lateral to the dorsals; these characters are in contrast to the setae are similar. SUMMERS and PRICE (1970) considered inverted V-shaped peritremes, fan-like dorsals and simi­ this difference to be specific rather thal1 generic, thereby lar humerals in the type species. returning C. bakeri to CheletomOlpha. Tln'ee species, namely C. dolosus, C. obrutus, and C. opacus, bear paired hysteronotal shields, which suggests that they should be ,<3 Genus Cheletoplullles OUDElVIANS 1904 placed elsewhere. Alternatively, one or more of the ex­ (r=l.}n. ") l! amined specimens ofthese taxa were not adults, because Type species: Cheyletus l11ontandoni BERLESE and the nympl,ls ofmany cheyletids bear2 hysteronotal shields. TROUESSART 1889 We suggest that the position of some of the described species of Cheletol1101pha should be reconsidered. Diagnosis: Eyes present; palpal tarsus with 2 comb-like setae and 2 sickle-like setae; palpal claw with more than 3 teeth; peritremes M-shaped, with more than 3 links; body -<..\ Genus Cheletollata WOlVIERSLEY 1955 ovoid, legs II and III separated by less than body width; (-6v}.2.~ ) dorsum with a propodosomal and a contiguous hystero­ Type species: Cheletonata milesi WOMERSLEY 1955 notal shield; both shields bear similar spatulate setae surrounded by concentric striae, humeral setae alike, Diagnosis: Eyes present; palpal tarsus with 1 comb-like displaced pleuroventrally; leg I subequal in length to seta and 2 sickle-like setae; palpal claw with 4 basal teeth; idiosoma; all tarsi with smooth claws and empodia. peritremes forming an inverted V, with more than 3 linkS; body ovoid, legs II and III separated by less than body Other species: width; dorsum with a propodosomal and a hysteronotal C. peregrinus BERLESE* 1921 shield, latter only a nude, small median platelet; dorsal setae spatulate to fan-like, humerals alike; all legs shorter lfabitat: Bugs. Eastern Europe. 52 U. GERSON et al.

Figs 23-26 - Fig. 23. Paracaropsis travisi (BAKER). Female, dorsum (from SUMMERS and PRICE, 1970). Fig. 24. Neoacaropsis granlllatlls VOLGIN. Female, dorsum (from VOLGIN, 1969). Fig. 25. Cheletonata milesi WOMERSLEY. Female, dorsum (from SUMMERS and PRICE, 1970). Fig. 26. Microcheyla parvllla VOLGIN. Female, dorsum (from VOLGIN, 1969).

-<"I Genus Clteletopll)les OUDEMANS 1914 peritremes M-shaped, with more than 3 bulbous links; (1f:g.. '-d ) body ovoid, legs II and III separated by less than body Type species: Cheletophyes vitzthumi OUDEMANS 1914 width; dorsum with a propodosomal and a hysteronotal shield, latter restricted to opisthosomal region; both with Diagnosis: Eyes present; palpal tarsus with 2 comb-like similar, rod-like, barbed setae; humerals similar; all legs setae and 2 sickle-like setae; palpal claw with 2-3 teeth; shorterthan body, all tarsi with smooth claws and empodia. Further observations on the Cheyletidae 53

Other species Habitat: Bird, from shaft of wing feathers. Tropical C.apicola FAIN, LUKOSCHUS and NADCHATRAM 1980 America C. clavipilis FAIN, LUKOSCHUS and NADCHATRAM 1980 C. deodikari PUTATUNDA and KAPIL 1988 C. eckerti SUMMERS and PRICE 1970 .q Genus Chelol/otus BERLESE 1893 C. hamaji PUTATUNDA and KAPIL 1988 (-&:~.(,if) C. haJ)lanaensis PUTATUNDA and KAPIL 1988 Type species: Chelonotus selenirhynchus BERLESE 1893 C. indiacus SMILEY and WHITAKER 1981 C. newtoni PUTATUNDA and KAPIL 1988 Diagnosis: Eyes absent; palpal tarsus with 1 comb-like C. orientalis PUTATUNDA and KAPIL 1988 seta and a conic spine and 2 sickle-like setae; palpal C. panamensis KLOMPEN et al., 1984 claw with 1 large tooth; peritremes with more than 3 C. ruttneri PUTATUNDA and KAPIL 1988 links; palpal femur very large, wider than long; body C. shendei PUTATUNDA and KAPIL 1988 ovoid; legs II and III separated by less than body width; C. xylocopae RAMARAJU and MOHANASUNDARAM 1999 dorsum with contiguous propodosomal and hysteronotal shields which cover the entire idiosoma and extend pleu­ Habitat: On carpenter bees (Xylocopinae). South-east rally; dorsal setae mostly slender, none ultralong, med­ Asia. ians apparently missing; humerals similar, displaced pleuroventrally; all legs shorter than body; coxae II and III almost adjoining; all tarsi with smooth claws and ~) Genus Cheletopsis OUDEMANS 1904 empodia. (~}(;) Type species: Cheyletus nomeri Poppe 1888 Habitat: Squirrels and a tree shrew (Tupaia). South-east Asia. . Diagnosis: Eyes absent; palpal tarsus with 1 comb-like seta and 2 sickle-like setae; palpal claw with 1-2 teeth; peritreme with more than 3 links; body fusiform, legs II ~\s' Genus Cheyletia HALLER 1884 and III separated by less than body width; dorsum with a (-t$;' 5"') single, propodosomal shield; all setae slender, finely Type species: Acarus squamosus DE GEER 1776 barbed, several, usually including humerals, ultralong; all legs shorter than body, coxae III and IV contiguous; Diagnosis: Eyes present; palpal tarsus with 2 comb-like all tarsi with smooth claws and empodia; tarsus I with setae and 2 sickle-like setae; palpal claw with more than 3 solenidion. teeth, restricted to its basal part; peritremes with more than 3 links, posteriorly straight; body ovoid, legs II and Other species III separated by less than body width; dorsum with a C. animosa OUDEMANS 1904 propodosomal and a hysteronotal shield; both bear dis­ C. anax OUDEMANS 1904 similar setae, laterals fan-lilee, medians squamate; hum­ C. basilica OUDEMANS 1904 erals resemble lateral setae, pleuroventrally displaced; all C. charadrii MIRONOV et al., 1991 tarsi with smooth claws and empodia. C. daberti K1VGANOV and BOCHKOV 1994 C. impavida OUDEMANS 1904 Other species: C. magnanima OUDEMANS 1904 C. americana VOLGIN* 1969 C. mariae MIRONOV et al., 1991 C. aradiphila VOLGIN 1966 C. laureata HALLER* 1884 Habitat: Birds. Europe, South America. C. papillijera VOLGIN 1955

Habitat: Bugs, scale insects, birds. Europe, USA. .z (, Genus Cheletosoma OUDEMANS 1905 ('?tr ':f ) Remarks: We follow SUMMERS and PRICE (1970) in con­ Type species: Cheletosoma tyrannus OUDEMANS 1905 sidering squamosa to be the type species of Cheyletia, and agree with them that the concept of this genus "re­ Diagnosis: Eyes absent; palpal tarsus with 1 comb-like mains nebulous". Three different species, from three seta and 2 siclde-like setae; palpal claw edentate (but different habitats, seem to have been described under nymph II with one tooth); peritremes forming an inverted squamosa. One (off a bug) has remained with the U with more than 3 links; body ovoid, legs II and III specific name, and a second (from a bird) was renamed separated by less than body width; dorsum with a propo­ papillijera VOLGIN. The third (described by BAKER, 1949, dosomal and a small suranal hysteronotal shield bearing 2 from scale insects) was renamed americana by VOLGIN pairs of slender setae unequal in length; other dorsals (1969), a decision challenged by SUMMERS and PRICE slender, humerals ultralong; all legs shorter than body, (1970). VOLGIN (1969) considered lal/reata to be the type all tarsi with smooth claws and empodia. species, thereby negating BAKER'S (1949) decision to 54 U. GERSON et al.

c c

d

Figs 27-28 ~ Fig. 27. Neochelacheles messersmithi SMILEY and WILUAMS. Female, dorsum (a); leg I (b); genito-anal region (c) (fromSMILEY and WILUAMS, 1972). Fig. 28. Paramicrocheyla spillula OUVIER and THERON. Female, dorsum (a); venter (b); a dorsal seta (c); genito~anal region (d) (from OUVIER and THERON, 1989).

synonymize it with squamosa. A better understanding of with more than 3 broad links; body ovoid; legs II and Cheyletia is thus hindered by our deficient concept ofthe III separated by less than body width; dorsum bears.a type species. single wide propodosomal shield, with 4-5 pairs ofsetae; dorsal setae slender, nude or barbed; humerals alike; all tarsi without claws but with feather-like empodia; tibia I z, Genus Cheyletiella CANESTRlNI 1886 without solenidion

C. katangae FAIN 1972 C. nigripes MOLA* 1907 C. strandtmanni SMILEY 1970 C. parwnsetosus KARPELLES 1884 C. yasguri SMILEY 1965 C. patagiatus* NORDENSKIOLD 1900 C. ftf&Q . ~~k.sv C. phantosis AKBAR and AHEER 1994 Habitat: Cats, dogs, rabbits. Cosmopolitan. C. philippinensis CORPuz-RAROS 1988 C. pluridens FAIN and NADCHATRAM 1980 C. polymOlphus VOLGIN 1949 :So Genus Cheyletlls LATREILLE 1796 C. praedibundus RHODENDORF 1940 C-?-j.. /I) C. promptus OUDEMANS 1904 Type species: Acarus eruditus SCHRANK 1781 C. pseudomalaccensis FAIN 1982 C. punctulatus FAIN and LUKOSCHUS 1981 Diagnosis: Eyes absent; palpal tarsus with 2 comb-like C. pyriformis BANKS 1904 setae and 2 sickle-like setae; palpal claw with 1-4 teeth; C. rapax OUDEMANS 1903 peritremes M-shaped, with more than 3 links, posterior C. rohdendorfi ZACHVATKIN 1949 link straight; body ovoid, legs II and III separated by C. nvandae FAIN 1972 less than body width; dorsum with a propodosomal and C. saevus OUDEMANS * 1904 a hysteronotal shield, both with slender to spatulate C. schneideri OUDEMANS 1904 barbed lateral setae; humerals may be longer; median C. spatiosus QAYYUM and CHAUDHRI 1977 setae, if present, usually small or dissimilar; all legs C. strenuus OUDEMANS 1904 shorter than body, all tarsi bear smooth claws and empo­ C. tenuipilis FAIN et al., 1980 dia. C. trouessarti OUDEMANS 1903 C. truculentus VOLGIN 1949 Other species C. trux RHODENDORF 1940 C. acarophagus ZAHER and SOLIMAN 1967 C. tutela QAYYUM and CHAUDHRI 1977 C. acer OUDEMANS 1904 C. ugandanus LAWRENCE 1954 C. alacer OUDEMANS 1904 C. venator VITZTHUM 1920 C. allactaga FAIN and LUKOSCHUS 1981 C. vivatus QAYYUM and CHAUDHRI 1977 C. attiahi YOUSEF and ISSA 1972 C. vorax OUDEMANS 1903 C. audax OUDEMANS 1904 C. vvahndoensis AKBAR and AHEER 1994 C. aversor RHODENDORF 1940 C. woodrofJei JEFFREY 1979 C. avidus QAYYUM and CHAUDHRI 1977 C. zumpti FAIN 1972 C. ayyazi AKBAR et al., 1993 C. baloghi VOLGIN 1969 Habitat: Stored food, soil, plants, nests. Cosmopolitan. C. baridos AKBAR et al., 1988 C. bidentatus FAIN and NADCHATRAM 1980 Remarks. This is the largest and most problematic genus C. burmiticus COCKERELL * 1917 in the family. VOLGIN (1969) keyed out 29 species, of C. cacahuamilpensis BAKER 1949 which 13 were known only as males (some hetero­ C. carnifex ZACHVATKIN 1935 morphic), 8 only as females and both genders were C. clavispinus* BANKS 1902 described for the remaining 8 species. SUMMERS and C. crassus QAYYUM and CHAUDHRI 1977 PRICE (1970) listed 36 species but provided keys for C. desitus QAYYUM and CHAUDHRI 1977 9 only. Problems in species determination include C. digitarsus SUGIMOTO* 1942 loss of type material, inadequate descriptions, the need C. egypticus ELBADRY 1969 to match up males and females, some variation in C. ferox TRouEssART 1885 shape and length of various setae and a sometimes C. fortis OUDEMANS 1904 broad definition of the genus. Thus we agree with SUM­ C. furibundus RHODENDORF 1940 MERS and PRICE (1970) that a complete revision of Chey­ C. gerbillicola FAIN and LUKOSCHUS 1981 letus may require the combined efforts of acarologists C. hendersoni BAKER 1949 from different parts of the world. A preliminary effort C. infensus AKBAR et al., 1993 towards understanding the extent of variation within 5 C. intrepidus OUDEMANS 1903 common species of Cheyletus was made by SUMMERS et C. kuznetzovi BOCHKOV and KHAUSTOV, 1999 al. (1972). C. legendrei FAIN 1982 C. linsdalei BAKER 1949 C. mafekingensis FAIN 1982 Jl Genus Chiapacheyllls DE LEON 1962 C. malaccensis OUDEMANS 1903 (~~JiI) C. malayensis CUNLIFFE 1962 Type species: Chiapacheylus edentatus DE LEON 1962 C. misonnei FAIN and LUKOSCHus,1981 C. mortelmansi FAIN 1972 Diagnosis: Eyes present; palpal tarsus with 2 comb-like C. nidicolus FAIN 1972 setae and 2 sickle-like setae; palpal claw edentate; peri- 56 U. GERSON et al.

Figs 29-32 - Fig. 29. Acal'opsellina sol/el's (RHODENDORF). Female, dorsum (a); leg I (b) (from SUMMERS and PRICE, 1970). Fig. 30. Acal'Opsel/a I'Ohdend01:fi (VOLGIN). Female, dorsum (from VOLGIN, -1969). Fig. 31. Philippicheylajilipina Corpus-RAROS. Female, dorsum (from Corpus-RAROS, 1972). Fig. 32. Cheletacal'lIs raptol' VOLGIN. Female, dorsum (from VOLGIN, 1969). Further observations on the Cheyletidae 57 tremes with more than 3 links; body ovoid, legs Il and 3V- Genus CUllliflel/a VOLGIN 1969 III separated by iess than body width; dorsum with a (= Bothrocheyla VOLGIN 1964) propodosomal and a hysteronotal shield, with similar (~c'J It'f- ) fan-like setae, some neotrichous; humerals similar to Type species: Neoellcheyla tllberclllicoxa VOLGIN 1964. lateral setae; all legs shorter than body; all tarsi with empodia; tarsus I without claws but tarsi Il-IV with Diagnosis: Eyes present; palpal tarsus with 2 comb-like smooth claws. setae, 1 sickle-like seta and an inflated seta; palpal claw with more than 3 teeth; peritremes with more than 3 links, Other species posteriorly straight or looped around a vesicular chamber; C. desertorum ZAHER and SOL/MAN* 1967 body ovoid, legs Il and III separated by less than body C. macrOC07'l1eus ZAHER and SOL/MAN* 1967 width; dorsum with a propodosomal and a hysteronotal shield, both covered by fan-like to squamate setae; lat­ Habitat: Plants. Mexico. erals usually differ from medians, and latter may differ amongst themselves; humerals displaced to pleuroventral Remarks: In their definition ofChiapacheylus, ZAHER and position, similar to lateral setae; all tarsi with smooth SOL/MAN (1967) stated that this genus had all legs with claws and empodia. claws and "pulvillus". This incorrect definition suggests that C. desertorum and C. maCI'OC07'l1eus may belong Other species: elsewhere. C. panamensis (BAKER) 1949 C. variegata BARILO 1985 C. whartoni (BAKER) 1949 3z. Genus Columbicheyla THEWKE and ENNS 1972 ( -?J -n) Habitat: Soil. Eastern Europe, USA. Type species: Columbicheyla macroflabellata THEWKE and ENNS 1972 Remarks: The definition of the genus is based on the inflated tarsal seta along with the vesicular chamber Diagnosis: Eyes present; palpal tarsus with 2 comb-like at the posterior part of the peritremes. BOCHKOV and setae, no sickle-like setae; palpal claw edentate; peri­ MIRONOV (1997) made an effort to elucidate the rela­ tremes with more than 3 links; body ovoid, legs II tionships of ClInlifJella, and the closely related Neoeu­ and III separated by less than body width; dorsum cheyla, by applying cladistic methods. As a result with a propodosomal and a hysteronotal shield, dorsal they resurrected BothrocheylaVoLGIN 1964 and re-as­ setae dissimilar, laterals and humerals fan-like, med­ signed species currently referable to ClI111ifJella and to ians squamate; all tarsi with smooth claws and empo­ Neoellcheyla. However, these authors used characters dia. that are variable within genera, a practice that detracts from their use for intergeneric analysis. Further, the 2

Habitat: Tree bark, North America.g most distinctive characters separating CunlifJella and ~, ~oiJl.U: Q offr..i c, L...;.. k""" .tk

Diagnosis: Eyes absent; gnathosoma with large lateral hook-like processes; palpal tarsus fused with pa1pa1 .?5'Genus Dubillilliola VOLGIN 1969 tibia, with 1 comb-like seta and 1-2 sickle-like setae; (+~ 1/-5') peritreme with more than 3 broad links, posteriorly Type species: Dubininiola polylepis VOLGIN 1969 convulated; body ovoid, legs Il and III separated by less than body width; dorsum with a propodosomal shield Diagnosis: Eyes present; palpal tarsus with 2 comb-like and a hysteronotal shield; all setae slender, humerals setae and 2 sickle-like setae; palpal claw with 1 large on pleuroventral platelets, similar to lateral setae; all tooth; peritremes with more than 3 links; body ovoid, legs legs shorter than body; solenidion cr on genu I replaced II and III separated by less than body width; dorsum with by a stellate seta; all tarsi with smooth claws and empo­ a propodosomal and a hysteronotal shield, each with dia. many (17 pairs together) neotrichous squamiform setae; venter with 2 pairs ofapparently neotrichous squamiform Other species setae, placed beyond the anals;humerals similar to lateral C. thailandicus FAIN and LUKOSCHUS 1985. setae; all legs shorter than body, all tarsi with a large branched ventral seta, and smooth claws and empodia; Habitat:TlIpaia glis. Africa, South-east Asia. claws on tarsus I minute. 58 U. GERSON et al.

Figs 33-36 - Fig. 33. Aegyptocheyla sUlIllllersi YOUSEF. Female, dorsum (a); genito-anal region (b); (from YOUSEF, 1978). Fig. 34. Oudelllansicheyl(l denlllarki (YUNKER). Female, dorsum (a); antero-dorsal'seta (b) (this and the next Fig. from SUMMERS and PRICE, 1970). Fig. 35. Cheletolllilllus berlesei (OUDEMANS). Female, dorsum. Fig. 36. Paracheyletiella volgini KUZNETZOV. Female, dorsum (from KUZNETZOV, 1977). Further observations on the Cheyletidae 59

Figs ,37-40 - Fig. 37. Columbicheyla macroflabellata THEWKE and ENNs.Female, dorsum (redrawn by A.F. from a paratype). Fig. 38. Chiapacheylus edentatus DE LEON. Female, dorsum (a); palpus (b); leg I (c) (from SUMMERS and PRICE, 1970). Fig. 39. Pavlovskicheyla semenovi (RHODENDORF). Female, dorsum (from VOLGIN, 1969). Fig. 40. Ker palmatus MUMA. Female, dorsum (from SUMMERS and PRICE, 1970). 60 U. GERSON et al.

Figs 41-44 - Fig. 41. CheletolllOlpha lepidopterorlllll (SHAW). Female, dorsum (this and the next Fig. from SUMMERS and PRICE, 1970). Fig. 42. Nodele calalllondin MUMA. Female, dorsum. Fig. 43. Cheletophyes vitzthllllli (OUDEMANS). Female, dorsum (from OUDEMANS, 1914). Fig. 44. Polycheyletlls boonkongae VAIVANIJKUL. Female, dorsum (from VAIVA­ NIJKUL,1979).

Habitat: Rodents. Turkemenistan. genus we could not treat herein (see above). The exam­ ination of D. polylepis indicated that the 2 genera are Remarks: Some salient features of this genus were very different. In contrast to Alliea, DlIbininiola has noted above. We formerly (FAIN et al., 1997) believed eyes, a large setulose and branched ventral seta on all Dubininiola to be very close to Alliea YUNKER, a tarsi, and 2 pairs ofsquamate setae beyond the anal setae. Further observations on the Cheyletidae 61

3(,. Genus ElIcheletopsis VOLGIN 1969 and similar lateral and median setae. SUMMERS and PRICE . . C-?;J ~) (1970) returned I'eticulata to Eucheyletia, arguing that the Type species: Cheletopsis major OUDEMANS 1904 of the genus will thus be simplified. The issue requires re-evaluation. Diagnosis: Eyes absent; palpal tarsus with one comb-like seta and two sickle-like setae; palpal claw with a single tooth; peritreme with more than 3 links; body ? ~ Genus ElIcheyletiella VOLGIN 1969 ovoid, legs II and III separated by less than body width; C-?:3 .18) dorsum only with a propodosomal shield; all setae, in­ Type species: Cheyletiella ochotonae VOLGIN 1960 cluding humerals, slender, ultralong; all legs shorter than body, coxae III and IV clearly separated; all tarsi Diagnosis: Eyes absent; palpal tarsus without comb-like with smooth claws and empodia; tarsus I without soleni­ and sickle-like setae; palpal claw edentate; peritremes dion CD. with more than 3 broad links; body ovoid; legs II and III separated by less than body width; female dorsum Habitat: Bird, New Guinea. bears only a propodosomal shield, as long or longer than wide; dorsal setae slender, mostly barbed; humerals si­ milar; all tarsi without claws but with feather-like empo­ dia; tibia I without solenidion Wv t9'i9 Diagnosis: Eyeti absent; palpal tarsus with two comb-like ~ ,.- 1~~~ I~l :~ I~I setae and two sickle-like setae; palpal claw with 2-4 Habitat: Rabbits (Lagomorpha), field mice. Cosmopoli- teeth; peritremes forming an inverted D, with more than tan. 3 links; body ovoid, legs 11 and III separated by less than body width; dorsum with a propodosomal and a hyster­ onotal shield, both with dissimilar dorsal setae, laterals :?g Genus ElItogenes BAKER 1949 and humerals fan-like, medians staghorn-like or squa­ ( icJ-·I~) mate; all legs shorter than body, all tarsi with smooth Type species: Eutogenes foxi BAKER 1949. claws and empodia. Diagnosis: Eyes absent; palpal tarsus with two comb-like Other species setae and two sickle-like setae; palpal claw edentate; E. asiatica VnLGIN 1955 peritremes with more than 3 links; body ovoid, legs E. bakeri VOLGIN 1963 II and III separated by less than body width, leg I E. bothl'Ophilia VOLGIN 1963 almost as .long as body, other legs shorter; dorsum E. eoa VOLGIN 1963 with a propodosomal and a hysteronotal shield, both E. flabellifera (MICHAEL) 1878 with similar dorsal fan-like setae; humerals alike; leg I E. fitnisciuri FAIN 1972 without claws and empodia, bearing four conspicuous E. hardyi BAKER 1949 terminal setae; tarsi II-IV with smooth claws and empo­ E. hmpyia (RHODENDORF) 1940 dia. E. kivuensis FAIN 1972 E. nidicola DEFINADO and KHATNG-FIELDS 1976 Other species E. nintoda CORPUZ-R<\ROS* 1988 E. africanus WAFA and SOLIMAN 1968 E. oregonensis SMILEY and WHITAKER 1981 E. bakeri CORPuz-RAROS 1998 E. pavlovskyi VOLGIN 1963 E. citri GERSON 1967 E. reticulata CUNLIFFE 1962 E. C07'l1utus CORPuz-RAROS 1998 E. sibirica VOLGIN 1963 E. fi'ater VOLGIN 1958 E. sinensis VOLGIN 1963 E. makilingiensis CORPuz-RAROS 1998 E. tanzaniensis FAIN 1972 E. narashinoensis HARA and HANADA 1960 E. taurica VOLGIN 1963 E. onoi (SHIBA) 1976 E. womersleyi VOLGIN 1963 E. pinicola THEWKE and ENNS 1972 E. punctata ZAHER and SOLIMAN 1965 Habitat: Soil, stored food, nests of small mammals. Cos­ E. quadrisetatus (BERLESE) 1913 mopolitan. E. reticularis OLIVIER and THERON 1988 E. vicinus SUMMERS and PRICE 1970 Remarks: VOLGIN (1969) erected Zachvatkiniola gen. novo for E. reticulata, due to its strong dorsal reticulation Habitat: Plants, soil. Cosmopolitan. 62 U. GERSON et al.

Fig. 45 - Dubininiola polylepis VOLGIN, 1969. Ho10type female, venter Ca) and dorsum Cb); tarsal claw Cc); branched seta on tarsus I (drawn by A.F.; scale line 5011m).

lto Genus Ga!agoche!es FAIN 1979 II and III separated by less than body width; idiosoma (~'~'1-3) with retrorse lateral processes between legs II and Ill; Type species: Cheletiella le17luricola LAWRENCE 1948 dorsum with a propodosomal shield and a hysteronotal shield; all setae, including humerals, slender to spine­ Diagnosis: Eyes absent; palpal tarsus with several un­ like; all legs shorter than body; coxa I with a lateral even rod-like setae but no comb-like and sickle-like process; legs 1-I1 with retrorse ventral processes on tarsi, setae; palpal claw edentate, hooked; palpal femur with genua and femora; solenidion 01 on genu I replaced by a lateral and ventral processes; base of gnathosoma with a stellate seta; all tarsi with smooth claws and empodia. pair of retrose processes; peritreme with more than 3 broad links, much expanded anteriorly; body ovoid, legs Habitat: Lemurs. Africa. Further observations on the Cheyletidae 63

b d

b

Figs 46-48 - Fig. 46. Samsinakia volgini (FAIN). Female, dorsum (a); gnathosoma dorsal (b) and ventral (c); leg I (d); genito-anal region (e) (from FAIN 1980c). Fig. 47. CllnlifJella tllberclllicoxa VOLGIN. Female, dorsum (from VOLGIN, 1969). Fig. 48. Hoffinannita mexicana PELAEZ. Female, dorsum (a); leg I (tarsus and tibia) (b); palpal tarsus and tibia, ventral (c) (from PELAEZ, 1962).

4' Genus Grallaclteles DE LEON 1962 dorsum bears a propodosomal and a hysteronotal shield; (~J n) anterior propodosomal setae fan-like, second pair and Type species: Grallacheles bakeri DE LEON 1962 posterior setae narrowly fan-like, all other dorsals rod­ like, barbed; humerals alike; all legs shorter than body, all Diagnosis: Eyes present; palpal tarsus with two comb­ tarsi with smooth claws and empodia. like setae and two sickle-like setae; palpal claw with more 1han 3 teeth; peritremes with more then 3 links; body Other species ovoid, legs II and III separated by less than body width; G. nan!engensis XIA et al., 1997. 64 U. GERSON et al.

Figs 49-52 - Fig. 49. Paracheyletia pyriformis (BANKS). Female, dorsum (a); leg I (b) (this and the next two Figs are from SUMMERS and PRICE, 1970). Fig. 50. Prosocheyla oaklandia (BAKER). Female, dorsum (a); leg I (b). Fig. 51. Cheletogenes O1'11atlls (CANESTRINI'and FANZAGO). Female, dorsum. Fig. 52. Neoellcheyla loricata (BERLESE). Female, dorsum (from BERLrSE, 1913). Further observations on the Cheyletidae 65

Habitat: Plants; stored food. USA, Pacific Islands, East Habitat. Plants, soil. Cosmopolitan. A~ia, Israel. Remarks: The present concept of Hemicheyletia, the second largest genus in the family, is unsatisfactory. It l[~ Genus Hemicheyletia VOLGIN 1969 contains one group whose median and lateral dorsal setae (= Dendrocheyla VOLGIN 1969; are similar and another with dissimilar setae. The hyster­ = Andrecheyla VOLG~N 1969) onotal shield may be reduced in members of both groups (Peg- (;; I) (H. volgini, with dissimilar dorsal setae, has an almost Type species: Paracheyletia bakeri EHARA 1962 obsolete, nude hysteronotal shield). VOLGIN (1969) tried to address the problem by restricting Hemicheyletia to Diagnosis: Eyes present; palpal tarsus with a two comb­ species with similar dorsal setae, creating Dendl'Ocheyla like setae and two sickle-like setae; palpal claw with more gen. novo for taxa with dissimilar dorsals, and adding than 3 teeth; peritremes with more than 3 links; body Andrecheyla gen. novo for H. scutellata, which bears a ovoid, legs II and III separated by less than body width; small hysteronotal shield. SUMMERS and PRICE (1970) did dorsum with ~m:~jg~pdilJly~11Q!illshield; not accept this arrangement and rejected Dendrocheyla lateral and humeral setae spatulate to fan-like, medians and Andrecheyla, a decision followed by most subsequent similar or dissimilar, in latter cases staghorn-like; all legs authors. However, as the number of species assigned to shorter than body, all tarsi with smooth claws and empo­ Hemicheyletia has almost trebled since then, it is time for dia. a new effort, utilizing more characters..

Other species H. anarbora (DE LEON) 1967 43. Genus Hoffma1l1dta PELAEZ 1962 H. arecana DYING 1972 (= Myrmicocheyla VOLGIN 1963) H. asiatica VOLGIN 1978 L -?'S Ifg ) H. athenae CORPuz-RAROS 1988 Type species: Hojjinannita mexicana PELAEZ 1962 H. bregetovae (VOLGIN) 1969 H. chlli TSENG 1977 Diagnosis: Eyes present; palpal tarsus with two comb­ H. congensis (c'UNLIFFE) 1962 like setae and two sickle-like setae; palpal claw with a H. cordove,;sis (DE LEON) 1962 single large tooth; peritremes forming an inverted U, with H. darwinia SUMMERS and PRICE 1970 more than 3 links; body ovoid, legs II and III separated by H. granllla SUMMERS and PRICE 1970 less than body width; dorsum with a propodosomal and a H. hissariensis MATHtJR and MATHUR 1981 hysteronotal shield, both with fan-like lateral and hum­ H. indica GUPTA 1991 erals setae and boatlike median setae; all legs shorter than H. kllreatollensis GOFF 1982 ;:; ~~ body, all tarsi with smooth claws and empodia. H. kysenyiensis THEWKE and ENNS 1979 H. lacinia RASOOL and CHAUDHRI 1979 Other species H. laguncllla RASOOL and CHAUDHRI 1979 H. clavipes (VOLGIN) 1963 H. lanceolata CORPuz-RAROS 1998 H. rimandoi CORPuz-RAROS 1972 -:r~ H. leytensis CORPuz-RAROS 1988 1-1 ,onwn<:-<.v4 L.' - .u- J A 9 n H. makilingensis CORPuz-RARos 1972 Habitat: Invertebrates: scorpion, millipede, tree bugs. H. mexicai1C1 THEWKE and ENNS 1979 Mexico, Eastern Europe, The Philippines. H. morii EHARA and GHAN! IBRAIDM 1988 H. newyorkensis DEFINADO and KHAING-FIELDS 1976 Remarks: Although the three species assigned to HofJ­ H. omissa TSENG 1977 mannita appear to be very similar, they actually make up H. pllsillifolillm LIN, PEN and CHEN 1994 two groups. The type species bears a pair ofeyes whereas H. reticlllata JEFFREY and CAMPBELL 1975 the other two taxa lack this character. An examination of H. rostella SUMMERS and PRICE 1970 the types and an evaluation ofadditional characters would H. scitllla CORPUZ-RAROS 1972 be needed to decide the issue H. sClltellata (DE LEON) 1962 H. serrula SUMMERS and PRICE 1970 H. transversa CORPUZ-RAROS 1972 41/ .Genus Hylopecheyla FAIN 1972 H. tropica (SHIBA) 1976 (f.'J" /(,,) H. tumidus QAYYUM and CHAUDHRI Type species: Hylopecheyla bunguranensis FAIN 1972 H. uichancoi CORPuz-RAROS 1972 H. vescus QAYYUM and CHAUDHRI Diagnosis: Eyes absent; palpal tarsus with two comb-like H. volgini (CUNLIFFE) 1962 setae and two sickle-like setae; palpal claw with more H. wellsi (BAKER) 1949 than 3 teeth; peritremes M-shaped, with more than 31inks, H. lindquisti THEWKE and ENNS 1979 posterior link abruptly bent inwards; body ovoid, legs II 19 H. l;vellsina (DE LEON) 1967 and III separated by less than body width; dorsum with a 66 U. GERSON et al.

56 b

Figs 53-56 - Fig. 53. Mexecheles cunlifJei DE LEON. Female, dorsum (this and the following two Figs from SUMMERS and PRICE, 1970). Fig. 54. Grallacheles bakeri DE LEON. Female, dorsum. Fig. 55. Hypopicheyla elongata VOLGIN. Female, dorsum (a); Tarsus I (b) Fig. 56. Cheyletia papillijera VOLGIN. Female, dorsum (from VOLGIN, 1969). lfVlY>l "-~ ~P,'V'

Other species Diagnosis: Eyes present; palpal tarsus with two comb­ like setae and two sickle-like setae; palpal claw with more Hylopecheyla malayi FAIN and NADCHATRAM 1980 than 3 teeth, placed along its entire length; peritremes with 3-4 links, posteriorly straight; body ovoid, legs II Habitat: Squirrels (Sciuridae). South Asia. and III separated by less than body width; dorsum with a Further observations on the Cheyletidae 67 propodosomal and a hysteronotal shield, contiguous; both propodosomal and a contiguous hysteronotal shield; both shields bear dissitnilar setae, laterals fan-like, medians bearing fan-like setae, humerals similar; hysteronotal squamate; humerals like lateral setae, pleuroventrally shield tapering posteriorly; all legs shorter than idiosoma; displaced; all tarsi with smooth claws and empodia. all tarsi with smooth claws and empodia.

Other species: Habitat: Old manure. Eastern Europe. H mirabilis (VOLGIN) 1955

Habitat: Bugs, beetles, soil. Eastern Europe, Asia, USA. If 4 Genus Metacheletoides FAIN 1972 (~t ~) Type species: Metacheletoides numidae FAIN 1972 ~, Genus J(er MUMA 1964 L-0sf}''fo) Diagnosis: Eyes absent; palpal tarsus without comb-like Type species: Ker palmatus MUMA 1964 setae, bearing a smooth stiff seta and two sickle-like setae; palpal claw with more than one tooth; peritremes Diagnosis: Eyes present; palpal tarsus with two comb­ with more than 3 links; body ovoid, legs II and m like setae and two sickle-like setae; palpal claw edentate; separated by less than body width; dorsum only with a peritremes with 3-4 links; body ovoid, legs II and m propodosomal shield, dorsal setae slender, barbed; sev­ separated by less than body width; dorsum with a propo­ eral, including humerals, ultralong; claws on tarsus I dosomal and a hysteronotal shield, both coarsely reticu­ minute, other tarsi with smooth claws and empodia. lated and with similar, spatulate to fan-like setae, hum­ eraIs alike; all legs shorter than body, all tarsi with Other species smooth claws and empodia. M akanyaruensis (FAIN) 1972 M crinijer FAIN 1979 Other species M gisagarensis FAIN 1979 K. acidalia AHEER et al., 1997 K. bakeri ZAHER and SOLIMAN 1967 Habitat: Birds. Africa. K. caeterus BARILO 1986 K. mercedesae CORPuz-RAROS 1998 K. pintoriensis CORPuz-RAROS 1998 :;0 Genus Metacheyletia FA~N 1972 C{,.·~. c" J Habitat: Soil, food stores, bird's nest. USA, Asia. Type species: Metacheyletia obesa FAIN 1972

Diagnosis: Eyes absent; palpal tarsus with two sickle-like it,. Genus Laeliocheyletia SUMMERS and PRICE 1970 setae and two setae that lackdentitions; palpal claw with {-?-.~ 62.. ) a single basal tooth; peritremes with more than 3 links; Type species: Laeliocheyletia teretis SUMMERS and PRICE body ovoid, legs II and m separated by less than body 1970 width; dorsum only with a small propodosomal shield; all setae slender; humerals similar; all legs shorter than body; Diagnosis: Eyes present; palpal tarsus with two comb­ tarsi I-m with smooth claws and empodia; leg IV vesti­ like setae and two sickle-like setae; palpal claw with more gial or absent. than 3 teeth; peritremes with more than 3 links; body ovoid, legs II and m separated by less than body width; Other species dorsum with propodosomal and hysteronotal shields, both Metacheyletia longisetosa ATYEO et al., 1984 with similar, rod-like, barbed setae; humerals alike but displaced onto ventral platelets; all legs shorter than Habitat: Parrots. Africa, South-East Asia, Mexico. body, all tarsi with smooth claws and empodia.

Habitat: Tenebrionidae (Coleoptera). Central America. . 51 Genus Mexecheles DE LEON 1962 (= Acarocheyla VOLGIN 1965) (FvJ 53) '1B Genus Lepidocheyla VOLGIN 1963 Type species: Mexecheles cunlijJei DE LEON 1962 ((-Vj n) Type species: Lepidocheyla gracilis VOLGIN 1963 Diagnosis: Eyes present; palpal tarsus with two comb­ like setae and two sickle-like setae; palpal claw with more Diagnosis: Eyes present; palpal tarsus with two comb­ than 3 teeth; peritremes with more than 3 links; body like setae and two sickle-like setae; palpal claw with more ovoid; legs II and m separated by less than body width; than 3 teeth; peritremes with 3-4 links; body ovoid, legs II dorsum with a propodosomal and a hysteronotal shield, and m separated by less than body width; dorsum with a both with dissimilar dorsal setae: laterals and humerals 68 U. GERSON et al.

d

c

e

Figs 57-58 - Fig. 57. Cheletophanes I/lontandoni (BERLESE and TROUESSART). Female, dorsum (a) and venter (b); a dorsal seta (c); palpus venter (d) and dorsal (e) (frpm OUDEMANS, 1906). Fig. 58. Lepidocheyla gracilis VOLGIN. Female, dorsum (a) and venter (b) (from VOLGIN, 1969). Further observations on the Cheyletidae 69

62

Figs 59-62 - Fig. 59. Anthribicheyla bocki THEWKE. Female, dorsum (redrawn by A.F. from the holotype). Fig, 60. TlItacheyla robllsta CORPuz-RAROS. Female, dorsum (from CORPUZ-RAROS, 1972). Fig. 61. Hemicheyletia bakeri (EHARA). Female, dorsum (from EHARA, 1962). Fig. 62. Laeliocheyletia teretis SUMMERS and PRICE. Female, dorsum (a); gnathosoma (b); a dorsal seta (c) (from SUMMERS and PRICE, 1970). 70 U. GERSON et al.

63

Figs 63-64 - Fig. 63. Teinocheylus longissimus FAIN. Female, dorsum (a) and venter (b); palpus venter (c) and dorsal (d); gnathosoma dorsal (e); leg I (t) and II (g) (from FAIN, 1974). Fig. 64. Chelonotus selenirhynchus BERLESE. Female, dorsum (a) and venter (b) (from DOMROW, 1960). lanceolate to strap-like, medians staghorn-like; leg I with Habitat: Plants, house dust, nests. USA, England, Aus­ minute claws and empodia, subequal in length to body or tralia. longer; other legs shorter than body; tarsi II-IV with smooth claws and empodia. ~-l, Genus Microcheyla VOLGIN 1966 Other species' (f,'}.1.G) M aztecorllm DE LEON 1962 Type species: Microcheyla parvlIla VOLGIN 1966 M hawaiiensis (BAKER) 1949 M impolitlls (SMILEY and MOSER) 1970 Eyes present; with a single comb-like seta, other seta M marshalli (BAKER) 1949 scimitar-like, and two sickle-like setae; palpal claw with M pannetts SUMMERS and PRICE 1970 more than 3 teeth; peritremes with 3-4 links; body ovoid, M virginiensis BAKER 1949 legs II and III separated by less than body width; dorsum M. votandinii JEFFREY 1975 with a propodosomal and a hysteronotal shield; lateral Further observations on the Cheyletidae 71 setae fan-like, medians dissimilar, squamate; humeral cesses". This suggests that the species should be placed setae displaced to a pleuroventral platelet, similar to elsewhere. laterals; all legs shorter than body, all tarsi with minute claws and empodia (see FAIN et. al. 1997). )l) Genus Neochelacheles SMILEY and WILLIAMS 1972 Other species Cf·iJ- •.2 1- ) M bengalensis GUPTA and PAUL 1992 Type species: Neochelacheles messersmithi SMILEY and M. granifera KUZNETZOV 1977 WILLIAMS 1972 M. ozkani Kot; and AYYILDIZ 1995 Diagnosis: Eyes present; palpal tarsus with one comb-like Habitat: Plants, soil.'Tadzhekistan, Russia, Turkey, USA. seta and two sickle-like setae; palpal claw with more than 3 teeth; peritremes with more than 3 links; body Remarks: VOLGIN (1969) described the type species as fusiform, legs II and III separated by more than body bearing minute claws, whereas SUMMERS and PRICE width; dorsum with a propodosomal and a hysteronotal (1970) emphasized the lack of claws on all tarsi. How­ shield; all dorsal setae spatulate-barbed, humerals like­ ever, the figures provided by KUZNETZOV (1977) and by wise; all legs shorter than body; all tarsi with smooth Kot; and AYYILDIZ (1995) indicate that their species carry claws and empodia. claws on all legs. Claws also seem to be present on leg I of M bengalensis according to the figure provided by Habitat: Tenebrionidae (Coleoptera), USA. GUPTA and PAUL (1992). These authors also wrote that M bengalensis has two comb-like setae, a feature not shared by the type species. n Genus Neocheyletiella BAKER 1949 (= 01'1litllOcheyla LAWRENCE 1959) (-to'g .1 0 ) D Genus Mudcheyla FAIN 1972 Type species: Neocheyletiella rohweri BAKER 1949 (.~v.}. ,.f" ) Type species: Muricheyla sicista FAIN 1972 Diagnosis: Eyes absent; palpal tarsus devoid ofcomb-like setae but with 1-2 sickle-like setae; tibial claw edentate; Diagnosis: Eyes absent; palpal tarsus with one comb-like no retrorse processes on palpal femora; peritremes with seta and two sickle-like setae; palpal claw with 1-2 teeth; more than 3 links; body ovoid, legs II and III separated by peritremes with more than 3 links; body ovoid; legs II and less than body width; dorsum with a propodosomal shield III separated by less than body width; dorsum with a which may be distinguished only by delicate striae; dorsal propodosomal and a hysteronotal shield; dorsal setae setae slender, a few, including humerals, may be ultra­ slender, barbed; humerals similar; all legs shorter than long; setae ic4 missing; all legs shOlier than body; all tarsi body; all tarsi with smooth claws and empodia; tarsi III with basally-inflated claws and empodia; coxae III and IV and IV with 3 dorsal conical processes each. widely separated, coxa IV with one seta; tibia I with solenidion -/ Genus Neoacaropsis VOLGIN 1962 N artami DOMROW 1965 Cf.;g 2'-1 ) N avicola FAIN 1972 Type species: Neoacaropsis granulatus VOLGIN 1962 N macronycha MEGNIN* 1878 N media FAIN 1972 Diagnosis: Eyes present; palpal tarsus with a single N megaphallos (LAWRENCE) 1959 comb-like seta; palpal claw with more than 3 teeth; N microrhyncha (BERLESE and TROUESSART) 1889 peritremes with more than 3 links; body ovoid; dorsum N pittae FAIN 1972 with propodosomal and hysteronotal shields, both with N rohweri BAKER 1949 spatulate setae; humerals similar; legs II and III separated N smallwoodae BAKER 1949 by less than body width; claws on all tarsi on swollen N siva FAIN 1972 stalks and with basal processes. N vestergaardi SMILEY* 1977

Other species: Habitat: Birds. Cosmopolitan. N levis CORPUZ-RAROS 1972 5'f Genus Neoeucheyla RADFORD 1950 Habitat: Soil, Russia. ('f;J n, ), Type species: Cheyletia loricata BERLESE 1913. Remarks: CORPuz-RARos (1972) described N levis as having "claws of legs II-IV smooth, without basal pro- Diagnosis: Eyes present; palpa1 tarsus with two comb-like 72 U. GERSON et al.

.. '\ ~,,' ' ~

Figs 65-66 - Fig. 65. Metacheyletia obesa FAIN. Female, dorsum (a) and venter (b) (fro111 FAIN, 1972, 1980c). Fig. 66. Ol'l1ithocheyletia aitkeni FAIN. Female, venter (a) and dorsum (b) (from FAIN, 1981). Further observations on the Cheyletidae 73

setae, a single sickle-like seta and an inflated seta; palpal ~ Genus Nodele MUMA 1964 claw with a single to more than 3~~~!!1;_p~elit!en~ith (= Neoclleletophyes VOLGIN 1965) more than 3 links, none paired,@:illl_~1:11:Jl2.~t~li()l:yesicllThD ('V?.4J- ) c£halii~~)bodyovoid, legs Il and III separated by less than Type species: Nodele calamondin MUMA 1964 body width; dorsum with a propodosomal and a hyster­ onotal shield, both covered by fan-like to squamate setae; Diagnosis: Eyes present; palpal tarsus with two comb­ laterals usually differ from medians, and latter may differ like setae and two sickle-like setae; palpal claw with a among themselves; humerals similar to other lateral setae; single basal tooth; peritremes with more than 3 links; all tarsi with smooth claws and empodia. body ovoid, legs Il and III separated by less than body width; dorsal propodosomal and hysteronotal shields in­ Other species detenninately separated, with similar rod-like barbed N beeri THEWKE and ENNS 1972 setae; humerals alike; leg I as long as body, other legs N bulgarica (VOLGIN) 1955 shorter; all tarsi with smooth claws and empodia. N dua CORPuz-RAROS 1998 N macrOCOl'l1eus SOLIMAN 1975 Other species N maysc[ CORPuz-RAROS 1998 N coccineae THEWKE and ENNS 1968 N minuta BARILO 1986 N mu HAINES 1988 N mUlnC1i VOLGIN 1969 N philippinensis (BAKER) 1949 N Ol'l1ata WAFA and SOLlMAN 1969 N simplex WAFA and SOLlMAN 1968 N pavlovskyi VOLGIN 1964 N superba KUZNETZOV 1977 N ploceus GUPTA and PAUL* 1992 N typhosa SUMMERS and PRICE 1970 Habitat: Soil, galleries of bark beetles. USA, Russia, N. ~e",- fa..J>" M- A'1-

Habitat: Soil, moss, plants. Europe, USA. (I. ~ Genus Omithocheyletia VOLGIN 1964 ~ Remarks: The definition of NeoeuclwrlCf.. Ms based on the jv1-'-U"Q. AA'l ( +;;. .G(, ) inflatedtM5J~eta along with the(;,i\l.bse~~ f a vesicular Type species: Ol'l1ithocheyletia dubinini VOLGIN 1964 chamber at the posterior part of the entremes; see also under CunlifJella. The presence or absence of eyes in N Diagnosis: Eyes absent; palpal tarsus devoid ofcomb-like ploceus could be determined neither from its description setae but with 1-2 sickle-like setae; tibial claw edentate; (GUPTA and PAUL, 1987) nor from the accompanying no retrose processes on palpal femora; peritremes with 3­ figure. The status of this species remains unclear. 4 links; body ovoid, legs Il and III separated by less than body width; dorsum with a propodosomal and two hys­ teronotal shields: one large, covers hysteronotum, other 1)'8 Genus Nihelia DOl\'1ROW and BAKER 1960 suranal; dorsal setae slender, barbed; some, including (= Hemicheyletus LAWRENCE 1954) humerals, ultralong; all legs shorter than body; all tarsi (-f:J 'p... ) with smooth claws and empodia; coxa IV with 1 seta; Type species: Nihelia calcarata DOl'vlROW and BAKER tibia I with solenidion

68

Figs 67-69 - Fig. 67. Ornithocheyletia dubinini VOLGIN. Female, dorsum (from VOLGIN, 1969). Fig. 68. Bakericheyla be/lOiN FAIN. Female, dorsum (a); leg I (b) (from FAIN, 1980b). Fig. 69. Bakericheyla (Apodicheyla) afi'icana FAIN. Female, dorsum (a)and venter (b) (froin FAIN, 1979b).

0. phylloscopi BOCHKOV et al., 1994 0. smileyi FAIN 1972 O. pinguis (BERLESE) 1889 0. volgini SMILEY 1970 O. psittaci FAIN 1972 0. psittaculae FAIN 1972 Habitat: Birds; cosmopolitan. Further observations on the Cheyletidae 75

Figs 70-71 - Fig. 70. Neocheyletiella media FAIN. Female, dorsum (a) and venter (b) (from FAIN, 1980a). Fig. 71. Apodicl1eles cypsiurus FAIN. Female, dorsunl (a) and venter (b) (from FAIN, 1979b). 76 U. GERSON et al.

73

a

Figs 72-73 - Fig. 72. Nihelia curvidens (LAWRENCE). Female, venter (a) and dorsum (b); gnathosoma: venter, left; dorsum, right (c) (from LAWRENCE, 1948). Fig. 73. Galagocheles le111uricola (LAWRENCE). Female, venter (a) and dorsum (b); gnathosoma: venter, left; dorsum, right (c) (from LAWRENCE, 1948 and FAIN, 1979a). Further observations on the Cheyletidae 77

b( Genus Oudemallsicheyla VOLGIN 1969 of the other species are lmown, and they bear similar . ( --&v rr 1'4 ) dorsal setae. A clearer definition of the genus will thus Type species: Cheletomimus denmarki YUNKER 1961 have to await the description of additional species or of the females of named taxa. Diagnosis: Eyes present; palpal tarsus with two comb-like setae and two sickle-like setae; palpal claw with more than 3 teeth; peritremes with at least 3 links; body ovoid, legs II 61.1. Genus Parache).Iletiella KUZNETZOV 1977 and III separated by less than body width; dorsum with a ({.} 3") propodosomal and two hysteronotal shields, placed side­ Type species: Paracheyletiella volgini KUZNETZOV 1977 by-side; all with similar dorsal fan-like setae; each hyster­ onotal shield with 5-7 setae; humerals alike; all legs short­ Diagnosis: Eyes present; palpal tarsus with two comb-ilce er than body; all tarsi with smooth claws and empodia. setae and two sickle-like setae; palpal claw with more than 3 teeth; peritremes with at least 3 links; body ovoid, Other species legs II and III separated by less than body width; dorsum 0. coprosomae THEWKE and ENNS 1976 with a propodosomal and two hysteronotal shields, placed side-by-side; with rod-like, barbed dorsal setae; each Habitat: Soil, plants. USA, Australasia. hysteronotal shield with 2 setae; humerals similar to other dorsal setae; all tarsi with smoQth claws and empodia, solenidia on all tarsi subequal to segment in length. b2. Genus Paracal'opsis VOLGIN 1969 C-c"J . .:/."» Habitat: Tree holes, Russia. Type species: Acal'Opsis travisi BAKER ·1949 Remarks: The type ofthis genus is very similar to several Diagnosis: Eyes present; palpal tarsus with a single comb­ species of Cheletomimus, the only consistent difference like seta and two sickle-like setae; palpal claw with more being the length ofthe solenidia on tarsi II-IY. A further than 3 teeth; peritremes with more than 3 links; body evaluation of this character is warranted. ovoid, legs II and III separated by less. than body width; dorsum with a propodosomal and 2 minute hysteronotal shields, placed in tandem, median, devoid ofsetae; dorsal &6'Genus Paramicrocheyla OLIVIER and THERON 1989 . setae slender, humerals similar but ultralong; all tarsi with (~.~ ,2 S) smooth claws and empodia, claws on tarsus I minute. Type sp€!cies: Paramicl'Ocheyla spinula OLIVIER anti THERON 1989 Habitat: Lizard, flies, moss. USA, Europe. Diagnosis: Eyes present; palpal tarsus with a single comb­ like seta; palpal claw with more than 3 teeth; peritremes (,3 Genus Paracheyletia VOLGIN 1955 withIQ~C': than 3 links; body ovoid, legs II and III sepa­ ({i) '(0) rated by less than body width; dorsum with a propodoso­ Type species: Cheyletus pyriformis BANKS 1904 mal and a hysteronotal shield, both with fan-like setae, humerals similar; all tarsi with empodia but lack claws. Diagnosis: Eyes present; palpal tarsus with two comb-like setae and two sickle-like setae; palpal clawwithmore than Other species 3 teeth; peritremes with more than 3 links; body ovoid, Paramicl'Ocheyla ungulina OUVIER and THERON 1989. legs II and III separated by less than body widtli; dorsum with a propodosomal and a hysteronota1 shield, both with Habitat: Soil, South Africa. dissimilar setae, lateral and humerals fan-like, medians staghorn-like; all legs shorter than body, all tarsi with Remarks: The description of this genus presents two smooth claws and empodia; claws on tarsus I minute. separate problems. First, the figure of P. spinula (herein reproduced as Fig. 28) suggests that the specimen exam­ Other species ined by OLIVIER and THERON may have been a nymph. P. hortensis VOLGIN 1969 Second, P. spinula lacks claws on all legs, whereas the P. recki VOLGIN 1966 second species, P. ungulina, has tarsal claws. P. samsinaki VOLGIN 1966

Habitat: Plants, insects. Eastern Europe, USA. &G .Genus Pavlovskicheyla VOLGIN 1965 ("C-'} , 33 ) Remarks: The present concept of Paracheyletia is based Type species: Cheletophyes semenovi RHODENDORF 1940 on SUMMERS and PRICE (1970). The female of the type species has dissimilar lateral and median dorsal setae, Diagnosis: Eyes present; palpal tarsus with two comb­ whereas all those ofthe male are similar. Only the males like seta and two sickle-like setae; palpal claw edentate; 78 U. GERSON et al.

Figs 74-75 - Fig. 74. SciurGcheyla squamosa (DOMROW and BAKER). Female dorsum, (a) and venter (b) (from DOMROW and BAKER, 1963). Fig. 75. Smileycheles camerGunensis FAIN. Female dorsum, (a) and venter Cb) (from FAIN, 1979a).

peritremes with more than 3 links; body ovoid, legs II and Habitat: Stored products, manure, Tenebrionidae (Co­ III separated by less than body width; dorsum with a leoptera). Uzbekistan, USA. propodosomal and a hysteronotal shield, both smooth or punctated and with similar, lanceolate to fan-like dorsal setae; humerals similar; all legs shorter than body, all (,1- Genus Pltiliyp;cltey/(/ CORPuz-RAROS 1972 tarsi with smooth claws and empodia. l ~J-.3 i) Type species: Philippicheyla jilipina CORPuz-RAROS Other species 1972 P. philippicana CORPuz-RAROS 1998 P. platydemae THEWKE and ENNS 1975 Diagnosis: Eyes present;palpal tarsus with two comb- Further observations on the Cheyletidae 79 like setae and two sickle-like setae; palpal claw with more Diagnosis: Eyes present; palpal tarsus with two comb-like than 3 teeth, located only along basal half; peritremes setae and two sickle-like setae; palpal claw with more with more than 3 links; body ovoid, legs II and III than 3 teeth; peritremes with more than 3 links; body separated by less than body width; dorsum with a single ovoid, legs II and III separated by less than body width; propQdosoma'l shieldbearing rod-like setae; humeral se­ dorsum with a propodosomal and a hysteronotal shield, tae alike; all tarsi with smooth claws and empodia. both with similar or dissimilar fan-like or staghorn-like V""'J ~ bo ~~, setae; humerals similar to lateral setae; leg J wit~ 11 Othel' species ,claws an(eml?odia, bearing four to six conspicuous term­ P. notelaeae GERSON 1994 inal setae; tarsi II-IV with smooth claws and empodia..

Habitat: Plf?ilts. The Philippines; Australia. Other species P. acantha (SMILEY and MOSER) 1970 P. buclmeri (BAKER) 1949 G2 Genus Polyclleyletlls VAIVANIJKUL 1979 P. hepburni (LAWRENCE) 1954 C~-Jg- . Li-Lf ) P. traubi (BAKER) 1949 Type species: Polycheyletus boonkongae VAIVANIJKUL P. villosa BOCHKOV and HAUSTOV, 1999 1979 Habitat: Plants. Cosmopolitan. Diagnosis: Eyes present; palpal tarsus with two comb-like setae and two sickle-like setae; palpal claw with 2 basal Remarks: VOLGIN (1969) as well as SUMMERS and PRICE teeth; peritremes with more than 3 links; body ovoid, legs (1970) noted that Pl'Osocheyla is a heterogenous assem­ II and III separated by less than body width; dorsum with a blage. VOLGIN (1969) split Prosocheyla into two subge­ propodosomal and a hysteronotal shield, both bearing nera, Pl'Osocheyla and Reckiana; the former ch~racter­ many neotrichous, similar fan-like setae; humerals simi­ ized by a large shield which covers most of its hyster­ lar; all tarsi with smooth claws and empodia. onotunk)}'hereas the latter bears only an indeterminate, centrally-located hysteronotal shield. SUMMERS and Other species: PRICE (1970) generally agreed with VOLGIN, but did not P. batangenius (CORPUZ-RAROS and SOTTO) 1977 accept his subgenera. Upon applying the generic concepts used in the present summation, we conclude that the Habitat: Soil; Thailand, The Philippines. genus may have to be split even further. All dorsal setae of the type species, P. oaklandia, are similar, whereas Remarks: CORPUZ-RAROS (1980) redescribed P. batan­ they are quite dissimilar on the dorsum of another mem­ genius (originally placed in Oudemansicheyla). She noted ber of that group, P. traubi. The same disparity may be that it differed from the type species by having only a seen in regard to the dorsal setae of P. hepburni and single comb-like seta (among other characters), the ap­ P. buckneri, both placed in the Reckiana group. Finally, propriate seta being present but lacking dentitions. At P. acantha, which has a longitudinally-separated hyster­ present it is difficult to evaluate the significance of this onotal shield, clearly deserves separate status. However, variation. were we to formalize these concepts, Pl'Osochyla would be split into five genera, a step which appears to be premature. (,1 Genus Prolll11riclleyla FAIN 1979 (FeJ9) Type species: Promuricheyla lukoschusi FAIN 1979 Sfl Genus Salllsinakia VOLGIN 1965 (= Cryptoclleyla FAIN, 1972) Diagnosis: Eyes absent; palpal tarsus with two comb-like ( Fi:l 4-G) setae and two sickle-like setae; palpal claw with 1-2 teeth; Type species: Cheletophyes theodoridis Samimik 1959 peritremes with more than 3 links; body ovoid; legs II and III separated by less than body width; dorsum with a Diagnosis: Eyes present; palpal tarsus with two comb­ propodosomal and a hysteronotal shield; dorsal setae like seta and two sickle-like setae; palpal claw with 1-12 slender, barbed, humerals similar; all tarsi bear smooth teeth; peritremes with more than 3 links; gnathosoma claws and empodia; tarsi III and IV with 2 dorsal conical partly covered by anterior margin ofpropodosoma; body processes each. ovoid; legs II and III separated by less than body width; dorsum with strongly-appressed propodosomal and hys­ Habitat: On Nannosciurus surrutilus, The Philippines. teronotal shields; dorsal setae slender, lanceolate or fan­ like, humerals alike; all tarsi with smooth claws and empodia. (0 Genus Prosoclleyla VOLGIN 1969 (~J- ~o) Other species: Type species: Cheletogenes oaklandia BAKER 1949. S. carabae RAMARAJU and MOHANASUNDARAM 1999 80 U. GERSON et al.

Figs 76-77 ~ Fig. 76. Cheyletiella parasitivorax (EONIN). Female dorsum, (a) and venter (b); genu 1, dorsal (c) (from SMILEY, 1970). Fig. 77. Bicheyletiella I'OlIIerolagi FAIN. Female dorsum, (a) and venter (b); leg I (c); leg IV (d) (from FAIN, 1979f).

S. gonocephalum FAIN 1984 Remarks: The variable number of teeth located on the S. pagongae CORPUZ-RAROS and SOTTO 1977 tibial claw of species consigned to this genus, as well as S. trilobitus BOCHKOV and MIRONOV 1998 the diverse shape oftheir dorsal setae, would indicate that S. volgini (FAIN) 1972 Samsinakia, as presently understood, consists of more than a single genus. CORPUZ-RAROS and SOTTO (1977) Habitat: Tenebrionidae (Coleoptera). Africa, Australia, were aware of these difficulties and only reluctantly The Philippines. placed pagongae (collected from soil, an unusual habitat Further observations on the Cheyletidae 81

79

Figs 78-79 - Fig. 78. Eucheyletiella ochotonae (VOLGIN). Female, dorsum (from VOLGIN, 1969). Fig. 79. Criokeron quintus (DOMROW and BAKER). Female, dorsum (from DOMROW and BAKER, 1963).

for Samsinakia) in this genus. Most species ofSamsinakia smooth claws and empodia in the shape oftwo diverging, were collected off tenebrionid beetles, and the variations distally-expanded rods. noted may reflect adaptations to specific hosts. A further evaluation of the characters defining Samsinakia awaits Habitat: Squirrel (Sciuridae). Thailand. the description of additional species.

73 Genus Slllilel'cheles FAIN 1979 ~2. Genus Sciuroclleyla VOLGIN 1969 (fv(j·1') Cf.Jrlf) Type species: Smileycheles camerounensis FAIN 1979 Type species: Nihelia squamosa DOMROW and BAKER 1963 Diagnosis: Eyes absent; palpal tarsus absent, thus with­ out comb-like or sickle-like setae; palpal claw edentate, Diagnosis: Eyes absent; palpal tarsus minute, without hooked; palpal femur with a ventral process; peri­ comb-like or sickle-like setae; palpal claw edentate, treme with more than 3 beadlike links; body ovoid, hooked; palpal femur and coxa I each with a retrose legs 11 and III separated by less than body width; ventral process; peritreme with more than 3 beadlike dorsum with a propodosomal and a hysteronotal shield; links; body ovoid, legs II and III separated by less than all setae, including humerals similar, slender; all legs body width; dorsum with two contiguous, propodosomal shorter than body; solenidion aI on genu I replaced and hysteronotal shields; dorsal setae dissimilar, medians by a stellate seta; all tarsi with smooth claws and empo­ squamate, laterals and humerals slender; humeral setae on dia. a separate platelet; all legs shorter than body; solenidion aI on genu I replaced by a stellate seta; all tarsi with Habitat: Rodents. Africa. 82 U. GERSON et al.

S{tf Genus TeillocheY/llS FAIN 1974 Acknowledgements . {{1.} ,b3) Type species: Teinocheylus longissimus FAIN 1974 We wish to thank Dr. H.V. DUBININA, Russian Academy ofSciences, St Petersburg, Russia, Dr. H. KLOMPEN, Acarology Laboratory, Ohio State University, Columbus, Ohio, U.S.A., Dr. RW. BROOKS, University of Diagnosis: Eyes absent; palpal tarsus without comb-like Kansas, LAWRENCE, Kansas, U.S.A. and Dr. D.G. FURTH, National setae and a single sickle-like seta; palpal claw edentate; Museum of Natural History, Washington, D.e., U.S.A., for the loan of type materia!. peritreme with more than 3 links; body fusiform, legs The following Institutions and periodicals have kindly granted us II and m separated by more than body width; dorsum copyright permission to reproduce certain published Figures, and we with a single propodosomal shield and two hysteronotal wish to thank the various Directors and Publishers for their courtesy. shields placed in tandem; all shields bear foliate setae; Figures # 11,12,15,18,25,34,38,40,41,42,49,50,51,53,54,55 and 62 (our numeration) are reproduced, with permission, from "Re­ other anterior dorsal setae, including humerals, slender view of the family Cheyletidae" by F.M. SUMMERS and D.W. or barbed, posterior setae feathered; idiosoma with PRICE, University of California Publications in Entomology, Vo!. 61, two setiferous lobes caudally; all legs shorter than 1970. Figures # 26, 32, 39, 47, 48, 56,58 and 78 (our numeration) are body, coxae m and IV clearly separated; tarsi I-m bear reproduced, with permission, from" Acarina ofthe Family Cheyletidae smooth claws and empodia; tarsus IV only with empo­ ofthe World" by V.I. VOLGIN, 1969. The permission was granted by dium. Dr. H. V. DUBININA, Russian Academy of Sciences, St Petersburg, Russia. Figures # 9, 10, 14,33,63,64,74,77, and 79 (our numeration) are Other species reproduced, with permission, from "Acarologia", Montpellier, Teinocheylus gundii FAIN et al., 1982. France. Vol. 2, 1960: p. 456-458; figs 1-2; Vo!. 5, 1963: p. 227-228, 231, figs 1-3; Vol. 10, 1968: p. 646, fig 1; Vo!. 16,1974: p. 271-273, figs 1-6; Vol. 20,1978: p. 366, fig 1; Vo!. 21,1979: p. 411, 414, fig l­ Habitat: Rodents. Africa. 15,36-37. Figures # 13, 16, 22, 44, 65, 69 and 71 (our numeration) .are reproduced, with permission, from "International Journal of Acarol­ ogy", Indira Publishing House, West Bloomfield, Michigan, U.S.A. tb Genus Thewkache/a IDE and KETHLEY 1977 Special permission granted by Dr. V. PRASAD, Editor in Chief. Vol. 5, (fJ. '8 ) 1979: p. 251, fig 1; p. 254, figs 1-2; Vo!. 6, 1980: p. 104, figs 1-3,7-8; Type species: Thewkachela ratuji IDE and KETHLEY 1977 p. 197, figs 19-22; Vo!. 7,1981: p. 110, figs 1,3; Vol. 12, 1986, p. 70, fig 2. Figures # 66 and 70 (our numeration) are reproduced, with permis­ Diagnosis: Eyes absent; palpal tarsus with one comb-like sion, from "Systematic Parasitology", Kluwer Academic Publishers, seta and two sickle-like setae, other comb-like seta Dordrecht, The Netherlands. Vo!. 2,1980: p. 29-31, figs 6, 8; Vo!. 2, 1981: p. 199, figs 26-27. modified to a thick spine; palpal claw edentate; peri­ Figures # 31 and 60 (our numeration) are reproduced, with pennis­ tremes with more than 3 links; base of gnathosoma with sion, from "The Philippine Entomologist", Los Banos, College, La­ strong hooks; body ovoid; legs II and m separated by less guna 4031, Philippines, Vo!. 2, 1972: p. 266-268, figs 11-12. Figures # 8 and 76 (our numeration) are reproduced, with permis­ than body width; dorsum with a propodosomal and a sion, from "The Annals of the Entomological Society of America", hysteronotal shield; dorsal setae short, barbed; humerals Lanham, Maryland, U.S.A., Vo!. 70,1977: 559-561, figs 2, 5, 9 and 15, similar, on separate platelets; all legs shorter than body; and Vol. 63, 1970: p. 1075-1076, figs 54-56. coxae I-II with lobes; all tarsi with smooth claws and Figure # 27 (our numeration) is reproduced, with permission, from "The .Proceedings of the Entomological Society of Washington", empodia. Washington, D.C., U.SA, Vo!. 74, 1972: Figure # 48 (our numeration) is reproduced, with permission, from Habitat: Squirrels. South-east Asia. "Anales de la Escuela Nacional de Ciencias Biologicas", Vo!. 11, 1962.

'f'? Genus Tlltacheyla CORPuz-RAROS 1972 (= l11dollesicltey/a THEWKE 1980) References ~o) CFeJ . Type species: Tutacheyla robusta CORPuz-RAROS 1972 (Except for references cited in the text, sources quoted by VOLGIN, 1969, or by SUMMERS and PRICE, 1970, are not re­ Diagnosis: Eyes present; palpal tarsus with two comb­ peated) like seta and two sickle-like setae; palpal claw with more AHEER, a.M., AKBAR, S. & CHAUDHRI, W.M., 1991. Two new than 3 teeth; peritremes strongly M-shaped, with more species of genus Acaropsis (Acarina: Cheyletidae) from Paki­ than 3 links; body ovoid, legs II and m separated by less stan. Acarologia, 32: 335-340. than body width; dorsum with a propodosomal and a AHEER, a.M., AKBAR, S. & CHAUDHRI, W.M., 1992. Three new hysteronotal shield which carry similar spatulate setae; species of the genus Cheletogenes OUDEMANS (Acarina: Chey­ humerals similar; all tarsi with smooth claws and empo­ letidae) from Pakistan. Acarologia, 33: 35-43. dia. AHEER, a.M., AKBAR, S. & CHAUDHRI, W.M., 1994. The genus Cheletoll1ill1l1s (Acarina: Cheyletidae). I. Descriptions of three Other species new species. Acarologia, 35: 345-351. T. buruensis (THEWKE) 1980 AHEER, a.M., AKBAR, S. & CHAUDHRI, W.M., 1997. New species ofthe genera Cheletoll1Olpha and Ker (Acarina: Chey­ Habitat: Plants. The Philippines; USA. letidae) from Pakistan. Acarologia, 38: 117-121. Further observations on the Cheyletidae 83

AKBAR, S. & AHEER, G.M., 1994. Mite fauna of summer BOCHKOV, AV., MIRONOV, S.V. & GORGOL, V.T., 1994. 01'­ vegetables in Punjab. Pakistan Joul'l1al of Zoology, 26: 339­ nithocheyletia phylloscopi sp. n. (: Cheyletidae), 345. new species ofparasitic mite from the willow warbler. Acarina, AKBAR, S., AHEER, G.M. & ISHTAQ, A., 1993. New predatory Russian Joul'l1al ofAcarology, 2: 73-80. mites from summer vegetables at Gujranwala. Pakistan Joul'l1al BRONSWIJK, l E.M.H., VAN & DE KREEK, EJ., 1976. Cheyle­ ofZoology, 25: 293-297. tiella (Acari: Cheyletiellidae) of dog, cat and domesticated AKBAR, S., SARWARRAHI, M. & CHAUDHRI, W.M., 1988. Three rabbit, a review. Joul'l1al ofMedical Entomology, 13: 315-327. new mite species of the family Cheyletidae from Pakistan. CHIROV, P.A & BOCHKOV, AV. 1998. Ol'l1itocheyletia miro­ Florida Entomologist, 1: 1-7. no1'i sp. n. (Acari: Cheyletidae) a new species ofparasitic mites ATTIAH, H.H., 1973. Chelachecaropsis bakeri, a new genus and of the bank swallow from Kirghizia. Acarina, Russian Joul'l1al species associated with stored food mites in Egypt (Acarina, ofAcarology, 6: 35-36. I Cheyletidae). In M. Daniel and B. Rosicky (eds), Proceedings CORPUZ-RAROS, L.A, 1972. Systematic studies of Philippine of the 3rd International Congress of Acarology, Akademia, cheyletid mites. I. Preliminary report of species mainly from Prague, pp. 349-352. Laguna. The Philippine Entomologist, 2: 247- 271.

ATYEo, W.T., KETHLEY, lB. & PEREZ, T.M., 1984. Paedomor­ CORPuz-RAROS, L.A, 1980. Systematic shldies of Philippine J( phosis in Metacheyletia (Acari: Cheyletidae), with the descrip­ cheyletid mites (Acarina). Ill. Polycheyletus batangenius (COR­ tion ofa new species. Journal ofMedical Entomology, 21: 125­ UZ-RAROS), comb. novo Kalikasan, Philippine Joul'l1al ofBiol- 131. ogy, 9: 61-68. BAKER, E.W., 1949. A review of the mites of the family CORPuz-RAROS, L.A, 1988a. Systematic studies of Philippine Cheyletidae in the United States National Museum. Proceed­ cheyletid mites (Acarina, Cheyletidae). IV. The genus Cheyle­ ings ofthe United States National Museum, 99: 267-320. tus LATREILLE. Philippine Joul'l1al ofScience, 117: 327-341. BAKER, KW., 1958. Chelacheles strabismus, a new genus and CORPUZ-RAROS, L.A, 1988b. Systematic studies of Philippine species ofmite from Porhlgal (Acarina, Cheyletidae). Proceed­ cheyletid mites (Acarina). V. New species and new records, ings ofthe Entomological Society ofWashington, 60: 234-235. with a note on the synonymy of Tutacheyja CORPuz-RAROS. x BARlLO, A.B., 1985. A new species of the genus Neoeucheyla Philippine Joul'l1al ofScience, 117: 413-427. RADFORD (Cheyletidae; ) from southern regions of CORPuz-RAROS, L.A, 1998. Twelve new species and one new Uzbekistan. Uzbekskii BioI. Zhur., 1985: 45-47 (in Russian). record of Cheyletidae (Acari) from the Philippines. 1ntel'l1a­ BARILO,AB., 1986a. New species of stigmaeid and cheyletid tional Joul'l1al ofAcarology 24: 259-290. mites (Acariformes, Stigmaeidae, Cheyletidae) from Uzbeki­ CORPUZ-RAROS, L.A & SOTTO, lM., 1977..Systematic studies X stan. Nauchnye Dold. vyssh. Shk. BioI. Nauki 1986:'25-29. of Philippine cheyletid mites (Acarina, Cheyletidae). II. New BARILO, AB., 1986b. Ker caeterus sp. n. (Prostigmata, Chey­ species and new records. Kalikasan, Philippine Journal of letidae) from Uzbekistan. Zoological Journal, 65: 298-300 (in Biology, 6: 143-170. Russian with English summary). DELFINADO, M.D. & KHAING-FIELDs, AA, 1976. Terrestrial BARlLO, A.B., 1989. New species of mites of the families mites ofNew York (Acarina). IV. Cheyletidae and Cheyletiel­ Raphignathidae, Stigmaeidae, Cheyletidae from Central Asia. lidae. Joul'l1al ofthe New York Entomological Society, 84: 189­ Zoological Joul'l1al, 68: 134-138 (in Russian with English 196. summary). Diaz PATXOT, l & GOFF, M.L., 1985. Two new species and new BERLESE, A 1913. Acari nuovi. Manipoli VII-VIII. Redia, 9: records of Cheyletidae (Acari) ~n Hawaii and a key to the 77-111. species. 1ntel'l1ational JouJ'llal o/Acarology, 11: 157-162. BOCHKOV, AV. & HAUSTOV, AA 1999. Prosocheyla villosa DOMROW, R. 1960. The genus Chelonotus BERLESE (Acarina, sp. N., a new mite species from Crimea (Prostigmata: Cheyle­ Cheyletidae). Acarologia, 2: 456-460. tidae). 10: 151-154 (Wroclaw) (in English). DOMROW, R. & BAKER, KW., 1960. Malaysian Parasites XLIV. BOCHKOV, AV. & KHAUSTOV, A.A. 1999. Description Cheyle­ A new genus of mites from a Thai mongoose (Acarina, Chey­ tus kuznetzovi sp. n. (Acarif011l1es, Cheyletidae) from Turkme­ letidae). Studies ofthe Institute ofMedical Research ofMalaya nia. Zoologicheskiy Zhul'l1al, 78: 115-116 (In Russian). 29: 194-197. BOCHKOV, AV., AA KHAUSTOV & KUZNETZOV, N.N. 1999. A DOMROW, R. & BAKER, E.W., t963. The genus Nihelia (Acar­ new mite species ofthe genus Chelacheles (Acarifol'lnes, Chey­ ina, Cheyletidae). Acarologia, 5: 225-231. letidae) from the Crimea. Zoologicheskiy Zhurnal, 78: 117-120 EHARA, S., 1962. Mites of greenhouse plants in Holdcaido, with (In Russian). a new species of Cheyletidae. Annotationes Zoologicae Japo­ BOCHKOV, AV., & MALIKOV, V.G. 1996. Eucheyletiella faini nenses, 35: 106-111. sp. n. (Acari: Cheyletidae), a new species of parasitic 'mites EHARA, S. & GHANI IBRAHIM, A, 1988. Cheyletid mites asso­ from Ochotona rufescens (Gray) (Lagomorpha, Ochotonidae). ciated with plants in the Malay Peninsula, with description of a Acarina, Russian Joul'l1al ofAcarology, 4: 43-48. new species (Acarina: Cheyletidae). Proceedings ofthe Japcl71 BOCHKOV, AV. & MIRONOV, S.V., 1997. On a taxonomy of Academy, 64B: 237-240. predatory mites of the genus Neoeucheyla RADFORD, 1950 and ELBADRY, E., 1969. Two new species of cheyletid mites from related genera (Acari: Cheyletidae). Acarina, Russian Journal milled wheat (Acarina:, Cheyletidae). Joul'I1al of Stored Pro­ ofAcarology, 5 (1-3): 73-78. ducts Research, 5: 157-167. BOCHKOV, A.V. & MIRONOV, S.V., 1998. Samsinakia trilobitus FAIN, A, 1972. Notes sur les acariens des famill1es Cheyletidae sp. n., a new cheyletid mite from South India (Acari: Cheyle­ et Harpyrhynchidae producteurs de gale chez les oiseaux ou les tidae). Entomologische Mitteilungen zoologische Museum mammiferes. Acta Zoologica et Pathologica Antverpiensia, 56: Hamburg, 12: 265-268. 37-60. 84 U. GERSON et al.

FAIN, A, 1974. Teinocheylus longissimus n.g., n.sp. a new fur­ letidae) from the nest of a carpenter bee in Malaysia. IntE?1'Ila­ mite from Pectinator spekei (Chey1etidae: ). tional Journal ofAcarology, 6: 309-312. Acarologia, 16: 271-273. . FAIN, A & NADCHATRAM, M., 1980. Cheyletid parasites of FAIN, A, 1979a. Chey1etidae (Acari, Prostigmata) parasitic on commensals in Malaysia (Acari: Cheyletidae). International Afrotropical primates, Carnivora and rodents. Revue de Zoolo­ Journal ofAcarology, 6: 191-200 gie aji'icaine, 93: 621-632. FAIN, A, SMILEY, R.L. & GERSON, u., 1997. New observations FAIN, A, 1979b. New Cheyletidae from Afrotropical swifts on the chaetotaxy and solenidiotaxy in the Cheyletidae (Acari: (Apodidae). International Journal ofAcarology, 5: 253-258. Prostigmata). Bulletin de I'Institut royal des Sciences naturelles FAIN, A, 1979c. Acariens du genre Cheyletus (Prostigmata: de Belgique, 67: 65-87 Cheyletidae) recoltes dans la region afrotropicale. International FLECHTMANN, C.H.W., 1971. Alguns Trombidiformes do Brasil Journal ofAcarology, 5: 275-284. e do Paraguai (Acari). Escola Superior de Agricultura "Luiz de Queiroz", Universidade de Sao Paolo, Piracicaba, Brasil, FAIN, A., 1979d. Idiosomal and leg chaetotaxy in the Cheyle­ pp. 63. tidae.lnternational Joul'l7al ofAcarology, 5: 305-310. GERSON, D., 1994. The Australian Cheyletidae (Acari: Prostig­ FAIN, A., 197ge. Notes on the genera Cheletoides OUDEMANS mata). Invertebrate Taxonomy, 8: 435-447. and Metacheletoides FAIN (Acarina, Cheyletidae) with descrip­ tion of three new species. Revue de Zoologie africaine, 93: GERSON, U. & FAIN, A, 1991. A new species of Bak (Acari: 1011-1025. Cheyletidae) from Thailand, with a key to species. Acarologia, 32: 17-21. FAIN, A, 1979f. Observations on chey1etid mites parasitic on mammals (Acari, Cheyletidae and Cheyletiellidae). Acarolo­ GERSON, D. & SMILEY, R.L., 1990. Acarine Biocontro1 Agents: gia, 21: 408-422. An lllustrated Key and Manual. Chapmanand Hall, pp. 174. GOFF, L.M., 1982. A new species of Hemicheyletia (Acari: FAIN, A, 1980a. Notes on some poorly known species of the Cheyletidae) from Kure Atoll, northwestern Hawaiian Islands. genus Neocheyletiella BAKER, 1949 (Acari: Cheyletidae) with a Proceedings ofthe Hawaiian Entomological Society, 24: 83-86. key to the genus. Systematic Parasitology, 2: 25-39. GUPTA, S.K., 1991. Studies on predatory prostigmatid mites of FAIN, A, 1980b. Le genre Bakericheyla VOLGIN (Acari, Chey­ northeast India with descriptions of new species and new letidae) dans la Region Afrotropicale. Description d'une espece records from India. Records ofthe Zoolological Survey ofIndia, nouvelle. Revue de Zoologique a./i'icaine, 94: 133-137. 88: 207-239. FAIN, A., 1980c. Notes on genera VOLGIN, 1965 and Samshzakia GUPTA, S.K. & PAUL, K. 1987. Some mites associated with Metacheyletia FAIN, 1972 (Acari: Cheyletidae). International birds' nests in West Bengal, with descriptions of eleven new Journal ofAcarology, 6: 103-108. species. Bulletin ofthe Zoological Survey ofIndia, 7: 1-23. FAIN, A, 1981. Revision ofthe genus Ornithocheyletia VOLGIN, GUPTA, S.K. & PAUL, K. 1992. Nest associated acarines ofIndia 1964 (Acari: Cheyletidae). Systematic Parasitology, 2: 181­ with descriptions of seven new species and notes on other 205. associates. Entomon, 17: 71-86. FAIN, A., 1982. Three new species of Cheyletus from Afrotro­ HAINES, C.P., 1988. A new species ofpredatory mite (Acarina: pical region and Madagascar (Acari, Cheyletidae). Revue de Cheyletidae) associated with bostrichid beetles on dried cassa­ Zoologie a.fi'icaine, 96: 79-89. va. Acarologia, 29: 361-375. FAIN, A, 1984. Samsinakia gonocephalum n. sp. , a new HASSAN, M.F. & GOMAA, E.A 1982. A new species ofthe genus cheyletid mite from an Afrotropical beetle Gonocephalum sim­ Chelacheles from Egypt (Acarina: Cheyletidae). The zoological plex Fab. (Acari, Cheyletidae). Revue de Zoologie africaine, Society ofEgypt, 31: 115-117. 98: 684-688 Hu, J.-D. & Hou, G., 1992. 011 the morphology of the genus FAIN, A., FELDMAN-MuHSAM, B. & MUMCUOGLU, Y, 1980. Cheyletiella and description of a new species from Xinjiang, Cheyletus tenuipilis n.sp. (Acari: Chey1etidae), nouvel acarien China (Acari: Cheyletiellidae). Acta Zootaxonomica Sinica, 17: des poussieres de maisons en Europe occidentale et en Israel. 183-188 (in Chinese with English Abstract). Bulletin et Annales de la Societe royale beIge d'Entomologie, IDE, G.S. & KETHLEY, J.B., 1977. Tl1ewkachela ratl!fi n.g. n.sp. , 116: 35-44. an unusual new cheyletid mite (Cheyletidae: Acariformes) from FAIN, A., GERRITS, P. & LUKOSCHUS, F.S., 1982. Teinocheylus giant squirrel, Ratl!fa, (Sciuridae: Rodentia) in Sabah and Thai­ gundii spec. novo from Ctenodactylus gundi (Acari, Cheyleti­ land. Annals ofthe Entomological Society ofAmerica, 70: 559­ dae). Revue de Zoologie aji'icaine, 96: 448-456. 562. FAIN, A & LUKOSCHUS, F.S., 1981a. Two new species of the JEFFREY, I.G., 1975. An undescribed species of Mexecheles genus Cheyletus LATREILLE, 1976 (Acari, Cheyletidae) from (Acarina: Cheyletidae) from a Scottish farm. Acarologia, 17: South African gerbils. Bulletin de l'lnstitut royal des Sciences 668-670. natlirelles de Belgique, 53: 1-7. JEFFREY, I.G., 1979. A new Cheyletus (Acarina: Cheyletidae) FAIN, A. & LUKOSCHUS, F.S., 1981b. Two new species of from a bat-roost. Acarologia, 21: 46-54. Cheyletus LATREILLE, 1796 (Acari, Cheyletidae, Prostigmata) JEFFREY, I.G. & CAMPBELL, lB., 1975. A new species of from gerbils. Rivista di Parassitologia, 42: 121- 125. Hemicheyletia (Acarina: Cheyletidae). Journal ofStored Pro­ FAIN, A & LUKOSCHUS, F.S., 1985. Description of a new duct Research, 11: 103-105. species in the genus Criokeron VOLGIN, 1966 and of the male KEH, B., LANE, R.S. & SHACHTER, S.P., 1987. Cheyletiella of Criokeron quintus (DOMROW and BAKER) (Acari: Cheyleti­ blakei, an ectoparasite of cats, as cause of cryptic arthropod dae). Acarologia, 26: 261-268. infestations affecting humans. Western Journal of Medicine, FAIN, A, LUKOSCHUS, F.S. & NADCHATRAM, M., 1980. Two new 146:192-194. species of Cheletophyes OUDEMANS, 1914 (Prostigmata: Chey- KETHLEY, lB., 1990. Acarina: Prostigmata (Actinedida). In: Further observations on the Cheyletidae 85

Dindal, D. (ed), Soil Biology Guide, John Wiley, New York, QAYYUM, H.A & CHAUDHRI, W.M., 1977b. Descriptions of six pp. 667-756. new mite species of the genus Cheyletus LATREILLE (Acarina: KHAN, A W. 1970. Some very common and highly destructive Cheyletidae) from Pakistan. Pakistan Journal of Zoology, 9: mites: Acarina in mango buds in West Pakistan. Science in 87-97. India, 7: 276-279. QAYYUM, H.A & CHAUDHRI, W.M., 1977c. Descriptions oftwo KIVOANOV, D.A & BocHKov, AI. 1994. A new mite species of new mite species ofthe genus Cheletogenes OUDEMANS (Acar­ the genus Cheletopsis (Acari: Cheyletidae). Bec. Zool., 1994: ina: Cheyletidae) from Pakistan. Pakistan Journal ofAgl'icul­ 39-43. tural Sciences, 14: 110-114. KLOMPEN, lS.H., MENDEZ, E. & LUKOSCHUS, F.S., 1984. Anew QAYYUM, H.A & CHAUDHRI, W.M., 1979a. Description of species of the genus Cheletophyes OUDEMANS, 1914 (Prostig­ Cheletomimus hel'edis, new species (Acarina: Cheyletidae) mata: Cheyletidae) from the nest of a carpenter bee in Panama. from Pakistan. Pakistan Entomologist, 1: 9-12. Acarologia, 25: 249-251. QAYYUM, H.A. & CHAUDHRI, W.M., 1979b. Mites of the genus Ko<;:, K. & AYYILDTZ, N., 1995. A new species ofMicl'ocheyla Hemicheyletia (Acarina: Cheyletidae) described from Pakistan. from Turkey (Acarina: Cheyletidae). Genus, 6: 225-228. Pakistan Journal o.fZoology, 11: 167-172. KUZNETZOV, N.N., 1977. Fauna ofmites of family Cheyletidae RAMARAJU, K. & MOHANASUNDARAM, M. 1999. Two new chey­ (Acariformes) in Crimea. Entomological Review, 56: 923-928 letid mites (Acari: Cheyletidae) from Tamil Nadu, India. Intel'­ (in Russian with English sunmlary). national Journal o.fAcarology, 25: 121-127. LAWRENCE, R.F. 1948. Some new pilicolus mites from South RASOOL, A. & CHAUDHRI, W.M., 1979a. Two new species ofthe African mamtnals. Parasitology, 39: 39-42. genus Hemicheyletia VOLGIN (Acarina: Cheyletidae) from Pa­ LEKPRAYOON, C. & SMILEY, R.L. 1986. Chelacal'opsis mool'ei kistan. Pakistan Entomologist, 1: 1-6. BAKER (Acari: Cheyletidae): redescription of the male and RASOOL, A. & CHAUDHRI, W.M., 1979b. Description of a new female. International Journal of Acarology, 12: 69-72. species of the genus Cheletogenes OUDEMANS (Cheyletidae) LIN, l & Lm, H., 1994. A new species of Cheletogenes (Acari: from Pakistan. Paldstan Entomologist, 1: 7-10. Cheyletidae) from Fujian. Entomotaxonomia, 16: 218-224. RASOOL, A., CHAUDHRI, W.M. & AKBAR, S., 1980. Studies on LIN, l, PEN, W. & CHEN, Y., 1994. A new species ofthe genus the mites of the family Cheyletidae from Pakistan. Pakistan Hemicheyletia VOLGIN from Fujian (Acari: Cheyletidae). Wuyi Entomologist, 2: 27-36. Science Journal, 11: 141-143. REOEV, S., 1974. Morphological and genetic evaluation ofmale MATHUR, S. & MATHUR, R.B., 1981. Hemicheyletia hissal'ien­ polymorphism in Cheyletus malaccensis (Cheyletidae: Acari­ sis, a new species of cheyletid mite from India. Acarologia, 7: na). Acarologia, 16: 85-93. 69-70. SHIBA, M., 1976. Taxonomic investigations on free-living Pros­ MEYER SMITH, M.K.P. & UECKERMANN, E.A, 1988. South tigmata from the Malay Peninsula. Nature Life in South East African Acari. Ill. On the mites ofthe Mountain Zebra National Asia, 7: 83-229. Park. Koedoe, 31: 1-29. SMILEY, R.L., 1970. A review of the family Cheyletiellidae MIRONOV, S.V., BocHKov, AV. & CHIROV, P.A, 1991. Mites (Acarina). Annals of the Entomological Society o.f Amel'ica, 63: 1056-1078. . of the genus Cheletopsis (Acarifortnes, Cheyletidae) from Charadriiformes birds of the Middle Asia. KYl'g Resp Ilim SMILEY, R.L., 1977. Further studies on the family Cheyletielli­ Akadem Kabal'lmJ' Khim-Teklmol Zhana BioI Ilim, 1991: 50­ dae (Acadna). Acarologia, 19: 225-241. 56. SMILEY, R.L., 1984. Two new species of Ornithocheyletia OLIVIER, P.AS. & THERoN, P.D., 1988. A new species of (Acari: Cheyletidae) from Australian birds. International JoUI'­ Eutogenes BAKER, 1949 (Acari: Cheyletidae) from South Afri­ nal ofAcal'Ology, 10: 239-250. ca. Phytophylactica, 20: 253-256. SMILEY, R.L., 1996. New species of Cheletonella (Acari: Pros­ OLIVIER, P.AS. & THERON, P.D., 1989. A new genus and tigmata: Cheyletidae) and a new key to the species. Anales species of Cheyletidae (Acari: Prostigmata) from South Africa. Instituto Biologico de Univel'sita Autonomica Mexico, Ser. Journal of the Entomological Society of Southern Afi'ica 52: Zoo1., 67: 239-244. 237-243. SMILEY, R.L. & MOSER, lC., 1970. Three cheyletids found with OUDEMANS, AC. 1906. Revision des Cheletines. Mbnoil'es de pine bark beetles (Acarina: Cheyletidae). Pl'Oceedings of the la Societe Zoologique de France, 19: 36-218. Entomological Society 0.[ Washington, 72: 229-236. OUDEMANS, AC. 1914. Acarologische Aanteekeningen LIV. SMILEY, R.L. & WHITAKER, lO., Jr., 1981. Studies on the Entomologische Berichten Nedel'lands Entomologie, 4 (78): idiosomal and leg chaetotaxy of the Cheyletidae (Acari) with 101-103. descriptions ofa new genus and four new species. International PRICE, D.W., 1972. A new species ofAcaropsella VOLOIN from Journal ofAcal'ology, 7: 109-128 the nest ofa turkey vulture (Acarina: Cheyletidae). Proceedings SMILEY, R.L. & WILLIAMS, G.L., 1972. A new genus 'and ofthe Entomological Society o.f Washington, 74: 45-49. species of Cheyletidae (Acarina). Proceedings o.f the Entomo­ PUTATUNDA, B.N. & KAPIL, R.P., 1988. Seven new species of logical Society of Washington, 74: 312-315. Cheletophyes (Acari: Prostigmata: Cheyletidae) associated SOLIMAN, Z.R., 1975. Three new species of cheyletid mites with carpenter bees in India. In: ChannaBasavanna, G.P. and from Egypt (Acari: Prostigmata) with a key to genera. Acar­ Viraktamath, C.A (eds), Progress in Acarology, Vo1. I, pp. 317­ ologia, 17: 95-102. 327. SUMMERS, F.M., 1976. A new genus for several cheyletid mites QAYYUM, H.A & CHAUDHRI, W.M., 1977a. Descriptions ofnew formerly in Acaropsis (Acarina: Cheyletidae). Proceedings o.f mites of the genus CheletomOlpha OUDEMANS (Acarina: Chey­ the Entomological Society of Washington, 78: 190-194. letidae) from Pakistan. Pakistan Journal ofZoology, 9: 71-77. SUMMERS, F.M. & PRICE, D.W., 1970. Review of the mite 86 U. GERSON et al. family Cheyletidae. University of Califo1'11ia Publications in n.sp. from Thailand (Acari: Cheyletidae). In te1'11ational Jou1'11al Entomology, 61:1-153. ofAcarology, 5: 251-252.. SUMMERS, F.M., WITT R. & REGEV, S., 1972. Evaluation of VOLGIN, V.I., 1969. Acarina of the Family Cheyletidae of the several characters by which five species of Cheyletus are dis­ World. Akademia Nauk, Leningrad, USSR, in Russian, pp. 432. tinguished (Acarina: Cheyletidae). Proceedings ofthe Entomo­ In English, 1987, Amerind Publishing Co., New Delhi, pp. 532. logical Society ofWashington, 74: 230-252. VOLGIN, V.I., 1978. New species of predatOlY cheyletid mites THEWKE, S.E., 1974. A new species of Chelacheles (Acarina: (Acariformes, Trombidiformes). Entomological Review, 57: Cheyletidae) from Missouri with a key to the lmown species. 213-217 (in Russian with English summalY). Entomological News, 85: 33-39. XiA, B., YE, R. & ZHU, Z., 1997. A new species of the genus THEWKE, S.E., 1980. Three new genera and species ofcheyletid Grallacheles (Acari: Cheyletidae) from Jiangxi, China. Sys­ mites (Acarina: Cheyletidae) from the US.A., Mexico, Panama tematic and Applied Acarology, 2: 173-175. and Indonesia. Jou1'11al of the Kansas Entomological Society, 53: 333-342. XIA, B., ZHU, Z-M. & YE, R. 1999. A new species ofthe genus THEWKE, S.E. & ENNS, W.R., 1968. A new species of predac­ Cheletonella (Acari: Cheyletidae) from China and a key to the eous mite (Acarina: Cheyletidae) from galleries ofbark beetles species. Systematic and Applied Acarology, 4: 149-151. in Missouri. Acarologia, 10: 215-219. YOUSEF, A..E.-T. A., 1978. Aegyptocheyla summersi n. gen., n. THEWKE, S.E. & ENNS, W.R., 1972. A new genus and three new sp. (Acarina: Prostigmata: Cheyletidae). Acarologia, 20: 365­ species of cheyletid mites (Acarina: Cheyletidae) from Mis­ 367. souri and Michigan. Joul'l1al of the Kansas Entomological YUNKER, C.E. 1960. Alliea laruei, n. gen., n. sp. , (Acarina: Society, 45: 450-459. Cheyletidae) from Rattus nO/1 l egicus (Erxleben) in Florida. THEWKE, S.E. & ENNS, W.R., 1974. A new species of Bak Pi'oceedings of the Helmintological Society of Washington, YUNKER (Acarina: Cheyletidae) from Missouri with a revised 27: 279-281. key to the known species. Jou1'11al ofthe Kansas Entomological ZAHER, M.A. and SOLIMAN, Z.R., 1967. The family Cheyletidae Society, 47: 42-53. in the UA.R., with a description offour new species (Acarina). THEWKE, S.E. & ENNS, W.R., 1975. A new species of Pavlovs­ Bulletin de la Societe Entomologique d'Egypte, 51: 21-26. kicheyla (Acarina: Cheyletidae) from the elytra of Platydema rufic017le (Coleoptera: Tenebrionidae) from Missouri. Acaro­ logia, 17: 671- 682. THEWKE, S.E. & ENNS, W.B.1976., Oudemansicheyla coproso­ U GERSON mae sp. novo (Acarina: Cheyletidae) from New Zealand. Jour­ Department of Entomology, nal ofthe Kansas Entomological Society, 49: 360-362. Faculty of Agricultural, THEWKE, S.E. & ENNS, W.B., 1979. Three new species of Food and Environmental Quality Sciences, Hemicheyletia VOLGIN (Acarina: Cheyletidae) with a key to Hebrew University, the known world species. Jou1'11al ofthe Kansas Entomological Rehovot 76100, Society, 52: 218-225. Israel DYING, I.-S., 1971. Cheyletid mites associated with stored products in Taiwan (I) (Acarina: Cheyletidae). Inspection Re­ A. FAIN search, Bulletin 4, pp. 36. Institut royal des Sciences Naturelles TJYING, I.-S., 1972. A new cheyletid mite Hemicheyletia are­ de Belgique, cana n. sp. from Taiwan (Acarina: Prostigmata). Plant Protec­ rue Vautier 29, tion Bulletin, 14: 8-10. B-lOOO; Bruxelles, Belgique TSENG, Y.-H., 1977. A contribution to the knowledge of For­ nlosan cheyletid mites (Acarina: Prostigmata). The Proceedings R.L. SMILEY ofthe National Science Council, 10: 213-263. Systematic Entomology Laboratory, UCHIKAWA, K. & SUZUKI, H., 1979. Cheyletiella mexicana Plant Sciences Institute, sp. novo (Acarina: Cheyletiellidae) parasitic on Romerolagus USDA, ARS, diazi (Mammalia, Leporidae). Tropical Medicine, 21: 21-27. Beltsville, MD 20705, VAIVANIJKUL, P., 1979. Polycheyletus boonkongae n.g., USA FURTHER OBSERVATIONS ON THE C!lEYLET1DAE (ACARI). WITH A KEY TO THE GENERA OF THE CllEYLETINAE AND A LIST OF ALL KNOWN SPECIES IN THE FNULY

by U.GERSON, 1\. FAIN AND R.L. SMILEY

(Bull.Inst.r.Sci.nat.Belg. 1999, vol.69 35-86)

CORRIGENDA

Page 39. Couplet 55 : should be read :

55. Palpal claw toothed along ba~al half•••• Cheyletia HaIler (Fig.56) ,Palpal claw toothed along entire length•••nypopicheyla Volgin (Fig.55)

Page 73 In the remarks to the genus Neoeucheyla, it should be read,: The definition of the genus Neoeucheyla is based on the presence in the internal part of the palpal-tarsus of a club-shaped seta (= modified sickle-shaped seta) along with the absence of a vesicular chamber in posterior part of peritreme.

Page 77 In the diagnosis of genus Paramicrocheyla : read peritreme with less than 3 links..

Page 80 In the legends of fig 76. instead of CEGNIN) ••• read (MEGNIN)

ADDENDA

Page 66 : In the legend of fig.55. add: U( from Summers and Price.1970)"

1996 : Cunlifella Volgin, 1969 : emend.by Bochkov et al.

1997 Neoeucheyla Radford, 1950. Ernend.Bochkov,A.,V. & S.V.Mlronov. 1967

1997 Caudache1es trigintaduae Lin:'Jian zhcn &, Zhang:,Yan-xuan,'Three n.sp. rind a',ncw record of Che~letinae from Fujian.' Wuyi Sci.J. 13: 131-138 (add ,OD p.45) Co1umbicheyla bicirci n. sp. (add pmp,' 57) Hoffmannita navicula n.sp. (add ou'o;65) Co1umbichey1a macroflabellata Thewke & Enns,1972 new record.

1997 Acaropsella nanchangensis Xia-Bin, Ye-Rong & Zhu-Zhimin One new species of r,enus Acaropsella , from China (Acari: Chey1eridae). Acta Araehno- 10gica sinica 6 : 143-145 (Add on p.39)

1998 Cheletomimus flecto Ahcer,G.M•• S.Akbar & W.M.Chaudri. New species of the genus Chel-, etomimus Oud. and a study of phoretic affinities of the ~pecies from Pakistan. Pakistan J.Zool.30: 125-132 Cheletomimus trema : same reference - 2 -

1999 Euchey1etie11a pusi11inus Bochkov and Mlronov. Mitt.hamb.zoo1.Mus.Inst.,96 : 141-15 A systematic review of the parasitic mite genus Euchey1etie11a Vo1gin,1969 (Prostigmata: Chey1etidae). (add on page 61) Euchey1etie11a pa1asius Bochkov & Mironov nov.spec. (add on page 61 ) Euchey1etie11a daurica Bochkov & Mironov nov.spec. (add on page 61) 2000 Neoeucheyla iranica Fain & Ardeshir> '2000Jntern.J.Acaro10gy, 26: 329-334 Notes on .~he genus Neoeuchey1a Radford,1950 (Acari: Cheyletidae) with . description ·of a new spec~es from Iran. ~ooo ~ Bak faini Corpuz-Raros._ Bak gersoni Corpuz-Raros Two new species and a new record of Bak from the Philip- pines (Acari: Cheyletidae. Intern.J.Acarol. 26: 328.