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Four new species records of Antennoseius and Anystipalpus (Acari: Mesostigmata: Ascidae) phoretic on Carabidae beetles from France

Faraji, Farid1; Dehelean, Stefan-Bogdan2; Vuyk, Mike3 and Bakker, Frank4

1MITOX Consultants/Eurofins, Science Park 408, 1098 XH Amsterdam, The Netherlands and Institute for Biodiversity and Ecosystem Dynamics, University of Amsterdam, Science Park 904, 1098 XH Amsterdam, The Netherlands
2MITOX Consultants/Eurofins, Science Park 408, 1098 XH Amsterdam, The Netherlands
3MITOX Consultants/Eurofins, Science Park 408, 1098 XH Amsterdam, The Netherlands
4MITOX Consultants/Eurofins, Science Park 408, 1098 XH Amsterdam, The Netherlands

2017 - Volume: 57 Issue: 2 pages: 337-353

DOI: 10.1051/acarologia/20164159
ZooBank LSID: 363F78B5-C0AC-471B-A658-EC47C6912F10

Keywords

Ascidae Carabidae France new species records Antennoseius Ansystipalpus

Abstract

Antennoseius pannonicus Willmann, A. pseudospinosus Eidelberg, Ansystipalpus livshitsi (Eidelberg) and A. percicola Berlese, collected from carabid beetles in southwest France, are new species records of ascid mites of France. The two species of Antennoseius are re-described and new host associations are provided. An attempt has been made to describe the spermathecal apparatus of these two genera for the first time.

Introduction

According to the catalogue of Moraes et al. (2016), Antennoseius includes 61 described species mainly inhabiting soil, litter, moss, humus, salt marshes and nests of rodents, birds and ants (Lindquist and Walter 1989; Karg 1993; Gwiazdowicz 2007; Beaulieu et al. 2008; Gwiazdowicz and Haitlinger 2010; Gwiazdowicz and Halliday 2010; Kazemi and Moraza 2013; Trach 2013). Not much is known about the feeding habits of these mites. The two noticeable studies are the works by Lindquist and Walter (1989) and Beaulieu et al. (2008) finding nematodes, Collembola and astigmatic mites as prey sources.

A few species of Antennoseius have been shown to have two female morphological morphs: smooth (phoretic) and granular (free-living) (Lindquist and Walter 1989; Beaulieu et al. 2008). This phenomenon might reveal more synonymies in this group of mites. Lindquist and Moraza (2009) re-described Anystipalpus and by describing two new species considered four members in that genus. Some species of Antennoseius Berlese, 1916 and Anystipalpus Berlese, 1911 are known from adult females phoretic under elytra of carabid beetles and the tegmina of labidurid earwigs (Lindquist and Moraza, 2009; Table 1).

Table 1. Species of Antennoseius Berlese, 1916 and Anystipalpus Berlese, 1911 associated with carabid beetles.

This paper intends to re-describe two species of Antennoseius and report two species of Anystipalpus, which have been found during a study in southwest France in 2014-15. The data provided in this study refer only to the phoretic morph of these four species. More data are given on the frequency and abundance of these four species.

Materials and methods

The study was conducted in an arable field in Villeneuve de Mezin, Lot-et-Garonne, southwest France sown with winter wheat. The carabid beetles were captured using funnel pitfall traps modified after Barber (1931) in late 2014 and 2015 during 12 sampling dates. In total 76 carabid species were identified using the work of Hurka (1996) and Freude et al. (2004). All individuals were checked for mites under the elytra and wings. Sixteen out of 76 species of carabid had mites associated with them.

Mite specimens were cleared in a mixture of Nesbitt and lactophenol solutions 1:1, and mounted in modified Hoyer's medium as described by Faraji and Bakker (2008). Drawings were made with the aid of a camera lucida (drawing tube) attached to an Olympus phase contrast microscope. The setal notations for the idiosoma follow Lindquist & Evans (1965), and leg chaetotaxy follows Evans (1963). Ranges of measurements are given in micrometers. The voucher specimens of mites are deposited in the Acari collection of Mitox Consultants, Amsterdam Science Park.

The number of carabid beetles carrying at least one mite of the genera Antennoseius or Anystipalpus was recorded at different sampling dates. To know about the abundance of mite species of these two genera, all mites were mounted.

Taxonomy
Antennoseius Berlese
Antennoseius Berlese, 1916: 303.

Type species: Antennoseius delicatus Berlese, 1916, by original designation.

For the genus diagnosis see Moraes et al. (2016).

Antennoseius (Antennoseius) pannonicus Willmann, 1951 (Figures 1-4, 8A)

Antennoseius pannonicus Willmann, 1951: 109.

Antennoseius pannonicus. Athias-Henriot, 1961: 461; Ryke, 1962: 662; Karg, 1971: 298; 1977: 4; 1993: 305; Farrier & Hennessey, 1993: 23.

Antennoseius (Antennoseius) pannonicus. Bregetova, 1977: 248; Beaulieu et al., 2008: 47; Lindquist & Moraza, 2009: 34; Moraes et al., 2016: 73.

Female — five specimens measured.

Figure 1. Antennoseius (Antennoseius) pannonicus, adult female: A – Idiosoma, dorsal view; B – Idiosoma, ventral view.

Figure 2. Antennoseius (Antennoseius) pannonicus, adult female: A – Subcapitulum; B – Tectum; C – Chelicera, lateral paraxial view.

Figure 3. Antennoseius (Antennoseius) pannonicus, adult female: A – Leg I (excluding tarsus); B – Leg II; C – Leg III; D – Leg IV.

Figure 4. Pterostichus madidus (Fabricius) carrying some Antennoseius (Antennoseius) pannonicus under the elytra.

Dorsal idiosoma (Figure 1A) — Idiosoma oval, 565 (530 – 590) long; dorsal shield divided and reticulate over entire surface; podonotal shield 294 (288 – 300) long and 240 (238 – 243) wide at j5 level, with 19 pairs of dorsal setae (r2 seta on the shield) and 7 pairs of pore-like structures (with no distinction made between poroids and glandular pores); j1 short 10 (9-10), j2, j3, j4 and j5 modified, short and spinelike 15 – 18, z1 20 – 21; opisthonotal shield 249 (235 – 263) long and 233 (230 – 238) wide at J2 level, with 15 pairs of dorsal setae and 8 pairs of pore-like structures; J2 35 (34 – 38), Z5 35 (32 – 38); lateral soft integument with 4 pairs of r setae (r3-6), r3 27 (25 – 28) and 21 – 22 pairs of R – UR setae, dorsal idiosomal setae smooth, except Z5 and most R – UR setae are slightly barbed.

Peritreme — Extending anteriorly to level of seta s1, at venter, anterior to coxa I (Figures 1A & 1B). Peritrematal-exopodal plate nearly smooth with three to four longitudinal lines near posterior end of stigma, widened and notched posteriorly; pores on peritrematal-exopodal plate not visible.

Ventral idiosoma (Figure 1B) — All ventral setae smooth; tritosternum 108 (105 – 110) long with paired laciniae, free for about three-fourths of total length and pilose, subrectangular base 16 – 18 long and 12 – 13 wide medially; two pairs of subtriangular sclerotized presternal plates; sternal shield 145 (138 – 149) long along midline, 111 (110 – 112) wide at st2 level; with 2 pairs of sternal setae (st3 off the shield) and three pairs of lyrifissures; st1 35 (33 – 38) long; with a prominent brownish crown-shape configuration between st1 setae, posterior part of sternal shield between lyrifissures iv2 and iv3 weakly sclerotized and laterally eroded; epigynal shield 58 (55 – 60) at greatest width, evenly rounded posteriorly with setae st5 on lateral margins; paragenital poroids iv5 on soft cuticle; endopodal plates weakly developed (except for region between coxae I and II, where fused with sternal shield), posterior section represented by one pair of strips between coxae III and IV parallel to epyginal shield and the other pairs positioned transversely between coxae II and III; anal shield weakly reticulate and pear-shaped 109 (102 – 115) long and 69 (65 – 73) of greatest width, with three circumanal setae and gland pores gv3 on shield margin; ten pairs of setae (JV1-5 and ZV1-5) on soft cuticle laterad of anal shield; with two pairs of metapodal plates, the smaller pair posteriad of peritrematal plate, the larger pair oval (about 28x10).

Gnathosoma — Tectum (Figure 2B) with anterior margin serrate and with three short projections, dorsal surface with a transverse line of denticles; deutosternum with a smooth transverse line between h3 followed by 7 denticulate transverse lines, denticulate lines 2 – 6 with 12 – 14 denticles, line 7 with 9 – 10 denticles and curved, proximalmost line longer with 25 – 30 denticles (Figure 2A); palpcoxal setae (pc) 27 – 29, hypostomal setae h1 25 – 26, h2 10 – 12, and h3 26 – 27, all smooth; corniculi horn-like; internal malae slightly fimbriated laterally; fixed cheliceral digit with setiform pilus dentilis and a row of about 10 teeth, movable digit tridentate (Figure 2C); palp apotele two-tined.

Legs (Figure 3) — Lengths, Leg I 607 (600 – 610), leg II 420 (410 – 430), leg III 410 (400 – 420), leg IV 557 (550 – 560); most of leg setae smooth except some setae lightly barbed, dorsal seta of trochanter I slightly thickened and barbed, ventral setae of trochanters I-IV slightly thickened; setation of legs I – II – III – IV: coxae 2 – 2 – 2 – 1, trochanters 6 – 5 – 5 – 5, femora 12 – 11 – 6 – 6, genua 13 (2 3/1 3/2 2) – 11(2 3/1 2/1 2) – 9 (2 2/1 2/1 1) – 9 (2 2/1 3/0 1), tibiae 13 (2 3/1 3/2 2) – 10 (2 3/1 2/1 1) – 8 (2 2/1 1/1 1) – 10 (2 2/1 2/1 2), tarsus II-IV 3 3/2 3/2 3+mv,md.

Spermathecal Apparatus — This species has apparently type A complex (laelapid) of sperm access system (Figure 8A); sacculus vestibules finger-shaped with relatively wider duct, situated between coxae III and IV.

Male — Not phoretic and not found.

Specimens examined — 23 June 2015: 8♀♀ on Harpalus rufipes (Degeer); 20♀♀ on Poecilus cupreus (Linne); 3♀♀ on Pterostichus madidus (Fabricius); 23 Sep. 2015: 3♀♀ on Calathus fuscipes (Goeze); 12♀♀ on Harpalus calceatus (Duftschmid); 1♀ on Nebria salina Fairmaire et Laboulbene; 18♀♀ on Pterostichus madidus (Fabricius); 3♀♀ on Scybalicus oblongiusculus (Dejean); 04 Aug. 2015: 3♀♀ on Harpalus distinguendus (Duftschmid); 1♀ on Pterostichus melanarius (Illiger), all collected by Bogdan Dehelean in Villeneuve de Mezin, Lot-et-Garonne, South West France.

Remarks — The prominent brownish crown-shape configuration on the sternal shield of A. pannonicus is a unique feature that allows identification of this species by just using a dissecting microscope. Descriptions of legs, tectum and hypostome, which were lacking in the original description by Willmann (1951) are provided here. Figure 4 shows the position of mites under the elytra of a carabid beetle.

Antennoseius (Antennoseius) pseudospinosus Eidelberg, 1990. (Figures 5 – 7, 8B)

Antennoseius (Antennoseius) pseudospinosus, Eidelberg, 1990: 78; Moraes et al., 2016: 73.

Female — five specimens measured.

Figure 5. Antennoseius (Antennoseius) pseudospinosus, adult female: A – Idiosoma, dorsal view; B – Idiosoma, ventral view.

Figure 6. Antennoseius (Antennoseius) pseudospinosus, adult female: A – Subcapitulum; B – Tectum; C – Chelicera, lateral paraxial view.

Figure 7. Antennoseius (Antennoseius) pseudospinosus, adult female: A – Leg I (excluding tarsus); B – Leg II; C – Leg III; D – Leg IV.

Figure 8. Female spermathecal apparatus: A – Antennoseius (Antennoseius) pannonicus; B – Antennoseius (Antennoseius) pseudospinosus; C – Anystipalpus livshitsi; D – Anystipalpus percicola, sacculus vestibules (sv).

Dorsal idiosoma (Figure 5A) — Idiosoma oval, 412 (400 – 435) long; dorsal shield divided and strongly reticulate over entire surface with some short striations especially around setae and behind J4, dorsal shield setae all setiform; podonotal shield 228 (225 – 235) long and 208 (200 – 215) wide at j5 level, with 19 pairs of dorsal setae (r2 seta on the shield), pore-like structures difficult to observe; j1 17 (15 – 18), z1 12 (11 – 13); opisthonotal shield 183 (175 – 195) long and 183 (180 – 185) wide at J2 level with 15 pairs of dorsal setae and 8 pairs of pore-like structures; J2 32 (31-33), Z5 35 (32 – 38); lateral soft integument with 4 pairs of r setae (r3-6), r3 23 (22 – 24) and 16 pairs of R – UR setae, most setae on opisthonotal shield and R – UR setae slightly barbed.

Peritreme — Extending anteriorly to level of seta z1 (Figure 5A). Peritrematal-exopodal plate reticulate with a line extending behind stigma, slightly widened and notched posteriorly.

Ventral idiosoma (Figure 5B) — All ventral setae smooth; tritosternum 90 (89 – 90) long with paired laciniae, free for about two-thirds of total length and pilose, subrectangular base 17 – 18 long and 10 – 11 wide medially; with two pairs of narrow sclerotized presternal plates; sternal shield 114 (113 – 115) long along midline (ignoring anterior concavity), 42 (40 – 45) wide at st2 level; mid anterior part with a depression, shield with two parallel longitudinal lines in the middle, posterior margin of sternal shield concave, with three pairs of sternal setae and two pairs of lyrifissures (iv3 apparently absent in all 23 mounted specimens); st1 25 (24 – 25) long, st1 and st2 slightly thicker than st3 and st4 narrower than st3; epigynal shield 42 (40 – 45) at greatest width, evenly rounded posteriorly with setae st5 on lateral margins; paragenital poroids iv5 on soft cuticle; endopodal plates weakly developed (except for region between coxae I and II, where fused with sternal shield), posterior section represented by one pair of strips between coxae III and IV parallel to epyginal shield and the other pairs positioned transversely between coxae II and III; anal shield weakly reticulate and pear-shaped 85 (78 – 88) long and 59 (55 – 63) of greatest width, with three circumanal setae and gland pores gv3 on shield margin; ten pairs of setae ( JV1-5 and ZV1-5) on soft cuticle laterad of anal shield; opisthogastric venter with two pair of metapodal plates, the smaller pair posteriad of peritrematal plate, the larger pair subcircular (about 17x13).

Gnathosoma — Tectum (Figure 6B) with anterior margin serrate and with three short projections, dorsal surface with a transverse line of denticles; deutosternum with a smooth transverse line between h3 followed by 7 denticulate transverse lines, denticulate lines with 5 – 9 denticles, line 7 curved and proximalmost line longer (Figure 6A); palpcoxal setae (pc) 18 – 20, hypostomal setae h1 25, h2 12 – 13, and h3 17 – 19, all smooth; corniculi horn-like; internal malae slightly fimbriated laterally; fixed cheliceral digit with setiform pilus dentilis and a row of about 15 – 16 teeth, movable digit tridentate (including a tiny subapical denticle)(Figure 6C); palp apotele two-tined.

Legs (Figure 7) — Lengths, Leg I 385 (380 – 390), leg II 300 (290 – 310), leg III 273 (270 – 280), leg IV 350 (345 – 360); all leg setae smooth except some setae lightly barbed, dorsal seta of trochanter I slightly thickened and barbed apically; setation of legs I – II – III – IV: coxae 2 – 2 – 2 – 1, trochanters 6 – 5 – 5 – 5, femora 12 – 11 – 6 – 6, genua 13 (2 3/2 3/1 2) – 11(2 2/1 3/1 2) – 9 (2 2/1 2/1 1) – 9 (2 2/1 2/1 1), tibiae 13 (2 3/1 3/2 2) – 10 (2 3/1 2/1 1) – 8 (2 2/1 1/1 1) – 10 (2 2/1 2/1 2), tarsus II-IV 3 3/2 3/2 3+mv,md.

Spermathecal Apparatus — Sacculus vestibules of this species is short and comma-shaped with narrow duct (Figure 8B). The sacculus vestibules stays most of the time attached to the margin of coxa III. Male — Not phoretic and not found.

Specimens examined — 13 April 2015: 7♀♀ on Poecilus cupreus (Linne); 27 April 2015: 4♀♀ on Poecilus cupreus (Linne); 23 Sep. 2015 2♀♀ on Calathus fuscipes (Goeze); 16 Nov. 2015: 2♀♀ on Nebria brevicollis (Fabricius), 8♀♀ on Nebria salina Fairmaire et Laboulbene, all collected by Bogdan Dehelean in Villeneuve de Mezin, Lot-et-Garonne, South West France.

Remarks — The original description of A. pseudospinosus provided by Eidelberg (1990) is poor with a mistake. He mentioned podonotal shield with 21 pairs of setae. His figure clearly shows r4 and r5 on the shield. The specimens collected in this study however have 19 pairs of setae (setae r4 and r5 off the shield). Dr. Viacheslav Trach confirmed the presence of 19 pairs of podonotal setae on his collected specimens from Ukraine (Pers. Comm. with the senior author).

Anystipalpus Berlese

Anystipalpus Berlese, 1911: 184.

Type species: Anystipalpus percicola Berlese, 1911, by original designation. For the genus diagnosis see Lindquist & Moraza (2009) and Moraes et al. (2016).

Anystipalpus livshitsi (Eidelberg, 1989)

Antennoseius livshitsi Eidelberg, 1989: 74.

Anystipalpus livshitsi. Lindquist & Moraza, 2009: 3; Moraes et al., 2016: 76.

Antennoseius ukrainicus Sklyar, 1994: 484 (Synonymy by Lindquist & Moraza, 2009: 18)._Antennoseius (Antennoseius) ukrainicus. Beaulieu et al., 2008: 52.

Specimens examined — 27 April 2015: 1♀ Poecilus cupreus (Linne); 04 Aug. 2015: 1♀ on Harpalus rufipes (Degeer); 23 Sep. 2015: 6♀♀ on Brachinus crepitans (Linnaeus), 8♀♀ on Harpalus rufipes (Degeer), 27♀♀ on Poecilus cupreus (Linne), 2♀♀ on Pterostichus macer (Marsham); 16 Nov. 2015: 2♀♀ on Pterostichus madidus (Fabricius), 2♀♀ on Pterostichus melanarius (Illiger), all collected by Bogdan Dehelean in Villeneuve de Mezin, Lot-et-Garonne, South West France.

Spermathecal Apparatus — Sacculus vestibules sack-shaped, relatively larger and more sclerotized than the other three species found in this study (Figure 8C).

Remarks — Lindquist and Moraza (2009) gave a complete re-description for this species.

Anystipalpus percicola Berlese, 1911

Anystipalpus percicola Berlese, 1911: 185.

Anystipalpus percicola. Lindquist & Moraza, 2009: 14; Moraes et al., 2016: 76.

Antennoseius (Antennoseius) nataliae Eidelberg, 1990: 75 (Synonymy by Lindquist & Moraza, 2009: 14).

Antennoseius (Antennoseius) nataliae. Beaulieu et al., 2008: 52.

Specimens examined — 04 Aug. 2015: 2♀♀ on Harpalus rufipes (Degeer); 23 Sep. 2015: 2♀♀ on Scybalicus oblongiusculus (Dejean), all collected by Bogdan Dehelean in Villeneuve de Mezin, Lot-et-Garonne, South West France.

Spermathecal Apparatus — Proximal part of sacculus vestibules narrow, sclerotized and then funnel-shaped (Figure 8D).

Remarks — Lindquist and Moraza (2009) gave a complete re-description for this species.

Key* to the species of the genera Antennoseius and Anystipalpus associated with carabid beetles

1. Palpus of normal length, at most one-third as long as leg I, with palptibia less than two times as long as palptarsus; sternal shield lyrifissures iv1-iv3 present (some species without iv3, Figure 3B); tibia IV with ten setae (pl2 present) .
...... Antennoseius Berlese 2

— Palpus slender, elongated, about half as long as leg I; palptibia two to three times as long as palptarsus; sternal shield lacking lyrifissures iv1, iv3, and sometimes iv2; tibia IV with nine setae (pl2 absent)
...... Anystipalpus Berlese 3

2. Leg I with pretarsus and a pair of small to well-developed claws
...... Antennoseius (Vitzthumia) Thor 6

— Leg I lacking pretarsus and claws Antennoseius (Antennoseius) Berlese
...... 10

3. Podonotal shield with eight pairs of strongly thickened, smooth, spinelike setae; sternal lyrifissures iv1-iv3 absent
...... 4

— Podonotal shield without enlarged, modified setae; one pair of sternal lyrifissures, iv2, present
...... 5

4. Thickened dorsal setae shorter (half as long) than other podonotal setae, and with rounded, blunt tips; epigynal shield drop-shaped, its lateral margins widened posteriorly; setae pd on trochanter I, pd2 on femur I and pd1-pd3 on genu I, short, thick, bluntly spine-shaped
...... Anystipalpus livshitsi Eidelberg, 1989

— Thickened dorsal setae about as long as other podonotal setae, and with distal half of length tapered, acutely pointed; epigynal shield tongue-shaped, its lateral margins nearly parallel; setae pd on trochanter I and pd1 on femur I acutely pointed, slightly spine-shaped
...... Anystipalpus kazemii Lindquist and Moraza, 2009

5. Dorsal shield setae moderately short, J1J4 about half as long as intervals between their bases; epigynal shield with posterior margin rounded; tarsus II with three posterolateral setae modified, spine-shaped; coxa I with both setae bluntly spine-shaped; genu I with pd3 a short blunt spine, femur I with pd2 a longer pointed spine
...... Anystipalpus percicola Berlese, 1911

— Dorsal shield setae moderately long, J1J4 nearly as long as intervals between their bases; epigynal shield with posterior margin bluntly pointed; tarsus II without modified setae; coxa I with one seta (av) bluntly spinelike; genu and femur I without bluntly spinelike setae .
...... Anystipalpus labiduricola Lindquist and Moraza, 2009

6. Podonotal shield with setae strongly thickened, spine-shaped or spur-shaped; setae of coxae I and/or II spine-shaped or swollen basally
...... Antennoseius (V.) perseus Beaulieu et al., 2008

— Podonotal shield with none of the setae thicker than others; setae of coxae I and II slender, setiform
...... 7

7. Podonotal shield with at least 30 pairs of short, smooth setae, unpaired setae present (neotrichia present)
...... Antennoseius (V.) multisetus Eidelberg, 2000

— Podonotal shield with 18 – 20 pairs of setae
...... 8

8. Palptibia and palpgenu with respectively one and two strongly thickened, short, smooth, spur-like dorso-distal setae; podonotal shield with 20 pairs of setae; anterior margin of tectum smooth or with very few denticles .
...... Antennoseius ( V.) pyrophilus Beaulieu, Déchêne and Walter, 2008

— Palptibia and palpgenu with normal dorso-distal setae; anterior margin of tectum with numerous denticles
...... 9

9. Setae j1 swollen and serrate, conspicuously different in shape from surrounding setae, and about 1.5 times longer than other dorsal setae
...... Antennoseius ( V.) ovaliscutalis Eidelberg, 2000

— Setae j1 sparsely barbed and similar in shape and length to surrounding setae .
...... Antennoseius ( V.) kamalii Moraza and Kazemi, 2009

10. Podonotal shield with some setae strongly thickened, spine-shaped or spur-shaped
...... 11

— Podonotal shield with all setae setiform (j1 might distally expanded)
...... 18

11. Podonotal shield with four pairs of setae strongly thickened, spine-shaped or spur-shaped (j1 should be ignored)
...... 12

— Podonotal shield with five, six or eight pairs of setae strongly thickened, spine-shaped or spur-shaped (j1 should be ignored)
...... 14

12. Seta j5 strongly thickened and spur-like; sternal shield with a prominent brownish crown-shape configuration between st1 setae; coxae leg I and II without modified setae .
...... Antennoseius (A.) pannonicus Willmann, 1951

— Seta j5 setiform; sternal shield without a prominent brownish crown-shaped configuration between st1 setae; coxae leg I and II with one modified seta
...... 13

13. Modified setae of podonotal shield with sharp tips; opisthonotal shield with 16 pairs of setae as well as two unpaired Jx setae
...... Antennoseius ( A.) bytinskii Costa, 1969

— Modified setae of podonotal shield with blunt tips; opisthonotal shield with 15 pairs of setae
...... Antennoseius ( A.) quadrispinosus Gwiazdowicz and Haitlinger, 2010

14. Podonotal shield with eight pairs of short and thickened setae
...... Antennoseius (A.) maltzevi Eidelberg, 1994

— Podonotal shield with five or six pairs of short and thickened setae
...... 15

15. Podonotal shield with five pairs of short and thickened setae
...... 16

— Podonotal shield with six pairs of short and thickened setae
...... 17

16. z5 short and thickened and z4 setiform
...... Antennoseius (A.) masoviae Sellnick, 1943

z5 setiform and z4 short and thickened
...... Antennoseius ( A.) ponticus Trach & Makarova, 2008

17. Seta j5 setiform and s1 short and thickened; coxae legs I and II with one short and thickened seta
...... Antennoseius (A.) sabulicola Bregetova, 1977

— Seta j5 short and thickened and s1 setiform; coxae legs I and II with setiform setae .
...... Antennoseius ( A.) sharonovi Eidelberg, 1989

(= Antennoseius ( A.) vysotskajae Sklyar, 1994)

18. Coxae I and II with one short and thickened seta
...... Antennoseius ( A.) matsjuki Eidelberg, 2001

— Coxae I and II with setiform setae
...... 19

19. Anal shield with two or three pairs of setae in addition to circumanals
...... 20

— Anal shield with only circumanal setae
...... 21

20. Anal shield with two pairs of setae in addition to circumanals; dorsal body setae smooth; opisthonotal unpaired Jx setae present .
...... Antennoseius ( A.) olallae Haitlinger, 2011

— Anal shield with three pairs of setae in addition to circumanals; dorsal body setae barbed; opisthonotal unpaired Jx setae absent
...... Antennoseius ( A.) dungeri Karg, 1965

21. Seta j1 distally expanded (fan-shaped) .
...... Antennoseius (A.) bullitus Karg, 1969

— Seta j1 similar to the adjacent setae, might be shorter or barbed
...... 22

22. Opisthonotal shield with unpaired Jx setae
...... 23

— Opisthonotal shield without unpaired Jx setae
...... 24

23. Fixed digit of chelicera with twelve teeth; epigynal shield droplet-shaped, narrowest part one-third of the widest part
...... Antennoseius ( A.) matalini Eidelberg, 2001

— Fixed digit of chelicera with five teeth; epigynal shield tongue-shaped, narrowest part about two-thirds of the widest part
...... Antennoseius ( A.) similis Eidelberg, 2001

24. Sternal shield with two pairs of setae (st3 off the shield)
...... Antennoseius ( A.) calathi Fain et al., 1995

— Sternal shield with three pairs of setae
...... 25

25. Epigynal shield droplet- or spear-shaped, narrowest part less than one-third of the widest part
...... Antennoseius (A.) pseudospinosus Eidelberg, 1990

— Epigynal shield tongue-shaped, narrowest part about two-thirds of the widest part
...... 26

26. Dorsal shield setae moderately long, J1J4 nearly as long as intervals between their bases
...... Antennoseius (A.) longisetus Eidelberg, 2000

— Dorsal shield setae moderately short, J1J4 about half as long as intervals between their bases
...... Antennoseius (A.) belorussicus Eidelberg, 1990

*Arranged based on the keys provided by Lindquist & Moraza (2009) and Moraza & Kazemi (2009)

Discussion

No descriptions have been yet provided for the spermathecal apparatus of Antennoseius and Anystipalpus. Lindquist and Moraza (2009) and Kazemi and Moraza (2013) mentioned the spermathecal apparatus of the investigated species as not distinctly sclerotized structures. Our examination on the four collected species showed to some extend sclerotization allowing the description for four of the species collected in this work. We are not certain whether the sclerotized portion is a 'sacculus' or a sclerotized section of a longer major duct. For simplicity, we have used the term "sacculus vestibules" in this paper. Whatever they are called, it seems that each species has its own unique structural form of spermatheca that should be described. Given that the other structures of spermathecal apparatus are not distinctly sclerotized, more details could not be obtained with the microscope used in this study. For this reason, the description of the spermathecal apparatus presented in this paper should be considered provisional pending an extensive examination with an elaborate microscope.

Figure 9. Frequency of carabid beetles carrying mites of the Antennoseius and Anystipalpus (figure above bars indicate the number of beetles checked for mites).

The carabid beetles collected in late autumn 2014 and winter 2015 did not have any mites associated. The adult females started phoresy in 13, April 2015 with a small number of beetles carrying mites. The peak number of carabids having mites was observed on June 23, 2015, when half of the carabids had at least one mite (Figure 9). The species found most frequently in this study was A. pannonicus (74%) followed by A. livshitsi (17%). Less abundant species were A. pseudospinosus and A. percicola with 8% and 1% respectively (Figure 10).

Figure 10. Proportion of species of Antennoseius and Anystipalpus found phoretic on carabid beetles.

We have found lack of host specificity for mites selecting the carabid hosts and some beetle species were the host to more than one species of these mites.The study for the frequency of carabids carrying mites was done by not identifying the species of these two genera. Therefore, a comprehensive study is needed to determine the population dynamics of each species. That should be extended to more locations in France to have a good picture of the species involved.

Acknowledgements

We are indebted to Dr. Viacheslav Trach, Department of Zoology, I.I. Mechnikov Odessa National University, Ukraine for examining specimens identified as A. pseudospinosus in his collection. We are also thankful to Drs. E.E. Lindquist, E.A. Ueckermann, and two anonymous reviewers for their valuable comments and corrections. The mite species reported in this study were obtained during a project performed by Mitox Field Operations Southern Europe.

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Date received:
2016-06-23
Date accepted:
2016-09-28
Date published:
2017-04-04

Edited by:
Tixier, Marie-Stéphane

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© 2017 Faraji, Farid; Dehelean, Stefan-Bogdan; Vuyk, Mike and Bakker, Frank

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