ACAROLOGIA A quarterly journal of acarology, since 1959 Publishing on all aspects of the Acari All information: http://www1.montpellier.inra.fr/CBGP/acarologia/ [email protected] Acarologia is proudly non-profit, with no page charges and free open access Please help us maintain this system by encouraging your institutes to subscribe to the print version of the journal and by sending us your high quality research on the Acari. Subscriptions: Year 2017 (Volume 57): 380 € http://www1.montpellier.inra.fr/CBGP/acarologia/subscribe.php Previous volumes (2010-2015): 250 € / year (4 issues) Acarologia, CBGP, CS 30016, 34988 MONTFERRIER-sur-LEZ Cedex, France The digitalization of Acarologia papers prior to 2000 was supported by Agropolis Fondation under the reference ID 1500-024 through the « Investissements d’avenir » programme (Labex Agro: ANR-10-LABX-0001-01) Acarologia is under free license and distributed under the terms of the Creative Commons-BY-NC-ND which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original author and source are credited. REMARKABLE SPECIES OF HISTIOSTOMA (HISTIOSTOMATIDAE, ASTIGMATA) FROM JAPAN WITH A PAIR OF NEWLY IDENTIFIED VENTRAL COXAL ORGANS AND SEXUAL DIMORPHISM OF CHEMICALS COMPONENTS KAZUMI TAGAMI 1 AND YASUMASA KUWAHARA 2 (Accepted November 2004) HISTIOSTOMA, HISTIOSTOMATIDAE, 3-HYDROXYBENZENE1,2-DICARBOALDEHYDE, NEW ORGAN. SuMMARY: Histiostoma globotuberum sp. n. was collected from Kyoto and Tsukuba, Japan. This species showed one pair of ring- or disk-like structures (coxal tuber) on the anterior-lateral part of coxa I of the deutonymph, adult male and female. Hexane extracts (possibly the opisthonotal gland extract) of the species consisted of 3-hydroxybenzene-1 ,2-dicarboaldehyde as a major component, in addition to geranial which commonly occurs in histiostomatid mites. Sexual dimorphism was observed in hydrocarbon profiles of mite extracts. RESUME: Histiostoma globotuberum sp.n. a ete recolte de Kyoto et Tsukuba, Japon. Cette espece montre une paire d'anneau ou de disque (tubercule coxal) a la partie anterieure de la coxa I de la deutonymphe de l'adulte male et femelle. L'extrait par l'hexane (possible extrait de la glande opisthonotale) est du 3-hydroxybenzene-1 ,2-dicarboaldehyde principalement avec du gerenial qui est · commun aux Histiostomatidae. Le dimorphisme sexuel a ete observe sur le profil des hydrcarbones des extraits des acariens. Small sized species of mites belonging to the genus Histiostoma are poorly investigated in Japan. They are found hosted on insects, in soil or litter samples, decaying vegetables, mushrooms, fruits and animal feces. MATERIAL AND METHODS In July 1996 and March 2000, two strains of histiostomatid mites were collected from the university campuses of Kyoto and Tsukuba from organic soil samples (ea. 500 g) enriched with moistened dried yeast on 5.5 cm paper disks. The mites were reared to obtain sufficient material for investigation. Subsequently, the morphology of specimens was studied at Tsukuba University and chemistry at Kyoto University. The Tsukuba strain was maintained through this study, but the former strain had been lost within one year after rearing began. Morphological studies Live specimens of all stages were cleared with Keifer's solution (KEIFER, 1953) in 1. Laboratory of Environmental Health, Division of Health and Sport Sciences, Graduate School of Human Comprehensive Science, University of Tsukuba, Tsukuba, Ibaraki 305-8574, Japan. 2. Laboratory of Chemical Ecology, Division of Applied Life Sciences, Graduate School of Agriculture, Kyoto University, Sakyo-ku, Kyoto, 606-8502, Japan Acaro/ogia, 2005. XLV, I : 83-98. 86a small Syracuse dish for 30-120 min. Following clearing, a drop of gum chloral was added to the dish and mixed thoroughly. The specimen was then transferred to a drop of fresh gum chloral on a glass slide for a few minutes to acclimate, and then gradually added to the final mounting solution, which was diluted with the cleaning solution into which the specimen had been immersed. Finally, the specimens were mounted on another glass slide with fresh gum chloral, covered with a cover slip and heated overnight to eliminate excess water before sealing at sooc on a hot plate. In this paper, nomenclature for idiosomal setae followed GRIFFITHS et al. (1990) and for leg setae GRANDJEAN (1939). Scanning Electron Microscopy (SEM): SEM observations were performed with JSM 6700F NT (JEOL, Tokyo). Detailed steps of preparation were as reported in SuzuKI et al. 1995. Chemical analysis Hexane extracts of mites (50 females and males, separately) were prepared using a conical-bottomed test tube (8 mm i.d. x 5 cm in length, hand made). Mites collected by a needle were transferred into the tube, and dipped in hexane (4 tLl) for three minutes. The resulting extract, after removal of mite bodies by a micro-syringe (1 0 tLl), was subjected to gas chromatography coupled with mass spectrometry (GC/MS), to obtain gas chromatographic (GC) profiles, and to identify compounds present in the extract. GC/MS was carried out by a Hewlett Packard HP-5890 GC/MS operated at 70 eV, using an HP-5 capillary column (0.25 mm i.d. x 30 m, 0.25 fLm in film thickness, Hewlett Packard) with He as the carrier gas at a 1.23 ml/min flow rate in the split-less mode at a temperature programmed to change from 60°C (2 min) to 290°C at a raising rate of 10°C/min and then held for 5 min. RESULTS Morphology More than 100 structures and some RLI index (ratio of length or distance of structure to the idiosomallength, Kurosa, 1987 and 2002) were used for the following descriptions (TABLE 1). Histiostoma globotuberum sp. n. (FIGS. 1-19) FEMALE (FIGS. 1-9) Body trapezoidal or vaseshaped. Idiosomal length 340-392 fLm, width 196300 fLm (n=6). Gnathosoma Chelicerae (FIG. 1); fixed digit with membranous finger-like distal portion, a median region with 3 large well developed teeth, movable digit with 14 teeth ending in a distinct bristle. Dorsal chelicera! seta medium in length. Each palp (FIG. 2 and 4) reflexed outward with short apical palpal solenidion (~)and eupathidial seta (u/') ( < 30 tLm). Palpal membrane poorly developed. Ventral subcapitular setae (m) filiform. Dorsal view (FIG. 3) Cuticle covered with fine microtrichae. Anterior propodosoma with a distinct sclerite extending posteriorly above legs I base level. A wing shaped and chelated pit located on the posterior margin of the propodosoma in almost all specimens. Two pairs of distinct protuberances on the posterior hysterosoma form the basement of the oil gland, d.£ and g_L (anterior protuberance) and b_z, b_£, b_1 andj£ setae (posterior protuberance). Setae ri and ve filiform. Scapular setae filiform, internal scapular setae (~) short, positioned anterio-medial to little longer external scapular setae (se). Sejugal furrow distinct. Hysterosomal setae filiform, subequal in length containing propodosomal setae. Opisthosomal gland (gjg_) well developed located between setae d2 and e2. Three pairs of cupules (ia, im, and iJl.). A pair of vestigial setal alveoli (va) described by Bongers et al. (1985) were present at upper third between el and hl. Well developed and pear-shaped copulatory opening with duct and bursa copulatrix (FIG. 5). Ventral view (FIG. 4) Anterior apodemes (coxa I) fused medially and V-shaped. Anterior and posterior papillae well screlotized and oval in shape. One pair of ring like screlotized structures on the posteriorlateral part of coxa I. These structures have never been recorded in the 460 known species of this family and are not mentioned in the review of astigmatid mites by Griffiths et al. (1990). We propose to call this structure a "ventral coxal tuber" (et). The function of this structure is still unknown. Coxal setae short; 4a located posterior-lateral of posterior papillae. Three pair of anal setae surroun- -87Stage Character IdiosomalL Idiosomal W Leg I length Leg II length Leg Ill length Leg IV length Gnathosoma L. Gnathosoma W Palp solenidion ul' m VI Dist. vi -vi ve Dist. ve- ve si Dist. si- si se Dist. se- se cl c2 c3 cp dl d2 el e2 f2 hl h2 h3 la Dist. la - la 3a Dist. 3a - 3a 3b Dist. 3b- 3b 4a Dist. 4a-4a ps3 Dist. ps3 - ps3 ps2 Dist. ps2 - ps2 psi Dist. psi -psi Sucker L. SuckerW Tarsus I L. wl Adult female 340-392 196-300 170-194 154-174 132-154 162-192 Adult male 268-288 140-158 140-144 124-132 28-32 18-21 14-18 21 -23 24-30 20-27 30-37 16-23 32-40 20-30 44-52 20-36 20-39 20-30 22-24 20-23 12-13 12-13 18-29 22-28 20-25 22-29 22-30 24-39 23-30 22-30 16-20 42-54 10-14 61-84 14-18 81-104 13-17 78-116 10-13 22-24 13-19 50-58 14-19 34-40 72-94 16-18 108-114 136-140 11-16 17-21 15-18 25-28 12-14 19-22 15-18 29-33 14-25 14-21 14-25 16-18 14-16 16-18 16-18 16-18 absent 18-21 17-20 15-17 15-16 38-40 14-16 57-59 9-11 38-49 11-12 33-38 8-10 Deutonymph 172-186 118-142 94-100 78-80 54-59 54-60 16-19 7-9 33-40 3 2-4 22-26 2-4 62 3-4 3-5 4-5 3-5 3-4 3-4 3-4 3-4 3-4 3-4 3-4 5-6 5 35-41 5 24-26 5-6 16-18 19-24 13-17 24-29 10-11 12-16 54-65 14-19 41-43 52-54 50-52 18-20 -88w2 8-14 9-11 w3 12-14 9-14 absent absent 9-12 6-9 9-10 6-8 aa ba la ra wa d Tibia I L. 13-16 10 9-12 6-8 8-14 6-7 5-6 3-4 11-13 7-9 4-6 76-86 74-76 24 22-28 20-22 16-17 26-29 7-8 <t> 13-16 13-15 hT gT Genu I L. 9-10 6-8 11-13 9-11 5-6 28-34 23-28 16-1 7 cri 12-16 13-16 cr2 10-15 9-12 8-11 9-11 2 cr cG vF Femur I L. pR Tarsus IlL. 8-11 6-7 8 7-8 50-58 43-48 15-18 7-9 7-8 64-70 48-54 34-36 w 16-21 20-23 13-14 ba wa la ra d Tibia II L. 10-13 9-10 7-9 10-14 10-12 6-7 12-15 10-12 9-10 9-11 8-11 5-6 32-41 32-36 8 20-22 15 10 <t> 13-17 18-20 11-13 hT gT Genu II L. 8-10 9-12 7 10-12 7-8 6-8 26-30 22-25 14-15 cr 12-14 12-13 8-10 cG mG vF Tarsus Ill L. d e 10-12 9-11 8-9 7-10 6-8 5-8 5-9 7-9 8-9 56-68 41-45 35-40 9-11 9-10 6-8 9-11 9-12 13-14 6-7 12 w Tibia III L. 13-16 7-8 19-22 18-20 <t> 14-29 12-15 15-19 kT Genu Ill L. sR Tarsus IV L. d e r 8-12 9-11 6-8 20-28 21-24 6-8 5-8 8-10 74-90 52-56 42-45 9-12 8-9 4-5 17-20 9-11 9-14 10-14 -89w 16-20 9-18 Tibia IV L. 20-26 20-22 <I> 13-20 10-14 kT Genu IVL. 9-11 9-12 30-32 24-26 wF 8-9 9-11 12-14 7-8 6-7 5-7 TABLE 1. Measurements of Histiostoma globotuberum sp. n. (range, fJ.ffi) ding the anal slit, 121!_ and 121J short filiform. 121?_ longer and located laterally. A pair of cupules (ih) on the posterior margin. Legs (FIGS. 3, 4, 6 -9) Legs slender, with all five segments free. Chaetotaxy: tarsi 13-12-10-10; tibiae 2-2-1-1; genua 2-2-0-0; femora 1-1-0-1; trochanters 1-1-1-0. Solenidiotaxy: tarsi 1-1-0-0; tibiae 3-1-1-1; genua 2-1-0-0. Famulus absent. Apical tarsus I situated 6 setae and 1 solenidion surrounding a claw. Tibia I excavated dorso-apically around common base of tibial solenidia Q!.L and Q!.?_· Solenidion Q!.J rod shaped. Most setae spine-like. Dorsa-apical tarsal setae d. I-11 elongated, filiform; d. III-IV supine shaped; trochanter setae .f!..B short spine, sR spine. MALE (FIGS. 10-11) Body box-like, wellsclerotized; idiosomallength 268-288 fLm, width 140158 fLm. Gnathosoma similar to that of female. Dorsal view (FIG. 10) ldiosoma sclerotized more than the female. Propodosomal shield sclerotized; propodosomal setae filiform shorter than 20 fLm. Sejugal furrow visible. Hysterosomal posterior protuberance seen in female was also distinct, however anterior protuberance was not recognizable. Hysterosomal setae filiform shorter than 25 fLm. Hysterosomal setaefl absent. Cupules: ia not visible; iJl_ moved to posterior margin; im and ih same as female. Opisthosomal gland (g]g) opening between setae d.l_andgL. Ventral view (FIG. 11) Y-shaped sternum formed by medial junction of Apodemes I. Apodeme 11 free. Posterior apodeme of leg 11 and anterior apodeme of leg Ill fused together, pointed inside and free medially. Ventral coxal tuber (et) same as on female. Anterior papillae oval, situated medially on apodeme IV and 3b; posterior papillae anterior to genital organ; the distance between the posterior papillae shorter than the anterior one. Coxal seta (3a) anterior-laterally to anterior genital papillae; stouter than that of female. Three pairs of anal setae (.];M_L,121l and 121J) present, 121!_ and 121?_ close to the anal slit, of subequallength and filiform, short and filiform_m:J is close to anterior part of the anal slit. Hysterosomal setae b_J is short and in ventra-posterior position. Legs (FIGS. 10-11) Tarsus of Legs I-IV stouter than those of the female. Leg chaetotaxy and solenidiotaxy same as in female. Apical tarsal solenidion Q!.J with cocked apices. Male solenidion !I.?_ shorter than female, solenidion Q!.!_ I longer. Famulus (~) absent. Setae ra and la leg 11 and sR leg Ill stouter than those of female. DEUTONYMPH (FIGS. 12-17) Body: ovoid, widest in region 1/5 posterior of sejugal; idiosomallength 172186 fLm, width 118-142 fLm. Gnathosoma (FIG 13) Gnathosoma reduced. Subcapitulum cylindrical, length 16-19 fLm, width 7-9 fLm. Palpal supracoxal setae (ul') dorsolateral, subapical (3 !Lm); palpal solenidia 33-44 !Lm in length. Dorsal view (FIG. 12) Propodosoma triangular in shape with a single faint transverse striation and medial curved end; setae sei and see short (2-4 !Lm). Hysterosoma striated anteriorly, smooth cuticle; hysterosomal setae mostly minute except stouter b_J. Ovoid ia between !l.j_ and 92, opisthosomal gland between dl and f?_, iJl_ invisible, probably hidden by some structure. Ventral view (FIG. 13) Apodemes I fused together at posterior to subcapitular remnant, forming a long sternum free posteriorly; apodemes 11 extending posterior-medially and end freely; apodemes Ill not fused medially. Apodemes IV extending forward, fused with median apodeme. Genital opening small and oval with two pairs of genital papillae. Genital setae (gg) minute, filiform, positioned posteriorlaterally to genital opening. Ventral coxal tuber (et) " :~\••...•i . 5. . -z.:~~ ' \0 0 4 3 FJm,: 1-5. Histiostoma globotuberum Tagami and Kuwahara, sp. n., female; habitus. 1. chelicera; 2. pulp; 3. dorsum; 4. venter; 5. copulatory opening and bursa copulatrix. Scale: 200 I-'m for FIGS. 3-4, 50 I-'m for FIGS. I, 2 and 5. -91- 50 ~m u~ s Fras. 6-9. Histiostoma globotuberum Tagami and Kuwahara, sp. n., female; legs. 6. leg I; 7. leg II; 8. leg Ill; 9. leg IV. 1OOJlfll \0 N 3a~ @~b 10 FIGS. 11 10-11. Histiostoma globotuberum Tagami and Kuwahara, sp. n., male; habitus. 10. dorsum; 11. venter. -93- 100 !!111 -- 13 FIGS. 12- 13 · Histiostoma globotuberum Tagam1. an d Kuwahara, sp. n., deutonymPh·' habitus. 12. dorsum,. 13 . venter. -94- 50 e J..Lm e ··c__) 1d FIGS. 14-17. Histiostoma globotuberum Tagami and Kuwahara, sp. n., deutonymphallegs (left). 14. leg I; 15. leg II; 16. leg Ill; 17. leg IV. -95- FIG I 8. Histiostoma globotuberum Tagami and Kuwahara, sp. n., coxal tuber (SEM microscopy). A. deutonymph. B. adult male. also situated anterio-lateral of coxal plate I. Sucker plate large and wide (40 flm (H) x 55 flm (W)), ovoid in shape with striate margin. Legs (FIGS. 14-17) Total length of distal4 segments moderate in all legs; tarsus I length (50-52 flm) sub equal to the combined length of tibio-tarsus HI (5459 flm) an <;I IV (54-60 flm), shorter than the combined length of leg II (78-80 flm). Chaetotaxy of tibia-genufemur I-IV same as in female. Setae vFlegl,g_,r_andsR leg Ill, and g_, r_ and .11: leg IV filiform. Distal tarsal setae g_ leg I spoon-shaped; g_ leg II knife- or leaf-shaped; g_ leg Ill (12 flm) and IV (17-20 flm) needle-shaped. Famulus £thick, hair-like and shorter (1 0 flm) than Qll. (18-20 flm). Tibial solenidion Q leg I thin and minute triangle(2-3 flm). SolenidionQl(13-14 flm)legiilarge rod and Q (8-1 0 flm) leg II rod shaped. SEM SEM microscopy of coxal tuber of the deutonymph is shown in FIG. 18 (A). A simple circular view was observed by the ordinal microscopic obser~ vation. However, the SEM figure showed a smooth globular tuberance. The view of adult male is shown in FIG. 18 (B). A circular pit, quite different from the surrounding surface material, was usually covered with a mass of bacteria. The granular surface showed some prints of bacteria when the material was fixed with ethyl-alcohol, the bacteria were sonicated away before SEM observation. The organs of female adults were similar in shape as that of male. The surface is not shown, because the process to remove the covering bacteria was not completely successful. Chemical analysis Gas liquid chromatogram was shown in FIG. 19. Total nine peaks (peak b, f, a, e, c, i, d, hand g, in decreasing order together with 9 unknown peaks) were observed in the female extract, while that of males composed of 6 peaks (i, a, f, h, g and b, in decreasing order with an unknown peak). By comparing their GC retention times and mass spectra with those of authentic compounds and/or known mite components (KuwAHARA, 1999), peak a was identified as geranial [(E)-3,7-dimethyl-2,6octadienal] (Kuwwahara et al., 1991 ), likewise peak b as 3-hydroxybenzene-1,2-dicarbaldehyde (SAKATA & KUWAHARA, 2001), peak c as tridecane, peak d as pentadecene, peak e as pentadecane, peak f as heptadecene, peak h as diethylhexyl phthalate, peak i as squalene, peak gas dibutyl phthalate. Among known compounds listed above, two phthalates (peak g and h) and squalene (peak i) were contaminants derived from diet and environment (Kuwahara, 1999), and furthermore, there is no unknown peaks corresponding to those detected among 52 species of Astigmata as ones to be studied, indicative of contaminants. As a result, total six compounds were identified as mite components. Recorder response of the male extracts was less than one-fifth of the females. Three hydrocarbons -96- b 90 80 A a f 70 60 . e* 50 Q) en c 0 40 a. en Q) ~ * c 30 ~ Q) ""0 ~ 0 0 20 d Q) * 0:::: 10 0 \1 I * *I I I I I I I I I .... I I I 1 r * g ** * l T ·1 I r I I T I h l T I T I T B 10 (\ 1"-... I I I I I I 1 f b l* l I T . a 20 V - . 1 10 h g l I 20 Retention time (m in) FIG. 19. Typical gas liquid chromatogram of females (A) and males (B) of Histiostoma globotuberum Tagami and Kuwahara, sp. n .. ; Analytical conditions: see the text. a:geranial [(E)-3,7-dimethyl-2,6-octadienal), b: 3-hydroxybenzene-1 ,2-dicarbaldehyde, c: tridecane, d: pentadecene, e: pentadecane, f: heptadecene, g: diethylhexyl phthalate, h: squalene, i: dibutyl phthalate. *: structures unknown. Peaks i and g were contamination and peak h (squalene) originated from dried yeast used as food. 97 (tridecane, pentadecene and pentadecane) were not detectable in males, indicative of sexual dimorphism. Type materials Holotype (adult female) and paratype (deutonymphs, adult males and females) and boucher specimens are deposited in the collection of the National Science Museum (Nat. Hist.), Tokyo (NSMT), Japan. Live specimens reared at Tsukuba University will be available for Japanese local researchers without authorized permission. However, requests from abroad require authorized permission from the local government for shipment up to 1 year after publication of this paper. _ Etymology This species is named from a specific structure on the deutonymphal coxal tuber (et) SEM photograph. The Japanese name "HARA-IBOHIGEDANI" is also after the shape and the position of the et. REMARKS Out of about 190 species of the genus Histiostoma, no other species has been described with a coxal tuber. Thus, this species can easily be distinguished from other species. DISCUSSION Classification Even though H. globotuberum sp. n possesses a pair of additional disks (et) on deutonymphal coxa I, this species should be classified into the genus Histiostoma. When more species with similar morphology are recorded to form a larger group, a new genus can be established from Histiostoma. Morphology The et shown in the H. globotuberum sp. n. is an interesting organ which has never been described in classifications of higher suborders of Astigmata nor the family Histiostomatidae and genus Histiostoma. However, some species of Oribatei possess two or more setae on their coxa I, but the et is not homologous to the setal organ seen by SEM. Authors anticipate further morphological, histological, functional and ecological studies with live specimens purchased from us. Additional ventral organs which have been described by VERCAMMEN-GRANDJEAN (1976, Claparede's organ) or by FAIN & ERTELD (1998, conoids) are not related to et, because conoids and et occur on the same deutonymph and et is recognized in all stages after the deutonymph instead of the Claparede's organ which appears before the deutonymphal stage. More detailed ontogenic studies are needed on this newly identified organ. Chemistry The present species contained geranial as the second major component, while its function remained obscure. Although GC profiles are available only for a limited species of Histiostomatidae, these species are characterized possessing geranial as one of the major components. Among 52 species of Astigmata including two species of Histiostomatidae (H. laboratorium and the present species), only four species possess geranial as one of major components; two histiostomatid mites, Dermatophagoides pteronyssinus and Chaetodaetylus nipponieus. In H. laboratorium, geranial has been known as the alarm pheromone (KUWWAHARA et al., 1991). In D. pteronyssinus, the compound functions as aggregation pheromone (KuwAHARA, unpublished). On the other hand, neral (geometrical isomer of geranial) is distributed more commonly among mites; more than 16 species, among which 5 species use neral as the alarm pheromone (KuwAHARA, 1999). Presence of 3-hydroxybenzene-1 ,2-dicarbaldehyde (used to be named as m/z 150) as the major component in the present females is one of the characteristics, not only among Histiostoma but also among Astigmata. Although the compound is distributed widely among species of Astigmata (more than 25 species among 52 species examined) as a minor component (Kuwahara, 1999), there is no species possessing the compound as the major. In the present species, sexual dimorphism was noticed in hydrocarbon components. It was practically true with no exeption that both sexes of astigmatid mites indicated the same GLC profile, however, it may be possible to indicate sexual dimorphism like the present species. Careful accumulation of GLC profiles should be carried out from now on to detect such possibilities among Astigmata species. REFERENCES & LUKOSCHUS FS (1985) Morphology and ontogeny of histiostomatid mites (Acari : Astigrnata) associated with cattle dung in the Netherlands. Zoo!. Verh. (223) : 1-56. BONGERS, MGH, 0CONNOR, BM 98 FAIN, A & BEAUCOURNU, JC (1973) Description de trois nouveaux hypo pes d' Anoetidae phoretiques sur les puces de mammiferes. Acarologia 15 : 514-520. FAIN, A & ERTELD, C (I 998) Description of a new species of Histiostoma Kramer, 1876 (Acari: Histiostomatidae) phoretic on the solitary bee Ha/ictus sexcinctus (Fabricus, 1775) (Hymenoptera: Apidae: Halictinae). Bull. Annls Soc. R. Beige Ent. 134: 47-57. GRANDJEAN, F (1939) La chaetotaxy des pattes chez les Acaridiae. Bull. 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