Berlesia multisetosa, Lindquist & Oconnor & Shaw & Sidorchuk, 2020

Lindquist, Evert E., Oconnor, Barry M., Shaw, Matthew D. & Sidorchuk, Ekaterina A., 2020, Review of the genera Berlesia Canestrini, 1884, and Katydiseius Fain & Lukoschus 1983, the subfamily Katydiseiinae Fain & Lukoschus, 1983, and their family group relationships (Acari: Mesostigmata: Gamasina), with description of three new species parasitic on gryllacridid crickets (Orthoptera), Zootaxa 4857 (1), pp. 5-70 : 18-26

Berlesia multisetosa new species

( Figures 7–12 )

Diagnosis

Adult: dorsal shield entirely smooth, female’s with irregular indication of lateral incisions at level of setae s6; vertex setae j1, mid-dorsals (j4–j6, z5, J1–J3) and J4 of moderate length (e.g., j4 nearly as long as interval between j4 and j5); dorsal shield setae j3 and z4 slightly longer, but shorter than diagonal interval between them, and shorter than transverse interval between j3; setae z1 and J2 present. Dorsolateral areas of idiosoma with setae of r–R and S series hypertrichous. Soft ventral cuticle hypertrichous, with about 15 pairs of opisthogastric setae. Leg I genu and tibia each with 11 setae, pd-3 and av-2 absent. Tarsus IV with seta pv-1 bluntly spine-like and asymmetrically shorter than attenuated av-1. Peritremes embellished with internal cellular structures.

Description

Idiosomatic dorsum. Adult female ( Fig. 7A ). Idiosoma 625–700 long, 412–432 wide between insertions of setae r3; dorsal shield 520–530 long, greatest width 290–320 at level between insertions of s4 (n=2), unornamented except for subdermal sigillar patterns; shield with irregularly formed lateral incisions at level of setae s6. Dorsal shield with usually 10 to 14 pairs (5 to 8 podonotal, 4 to 6 opisthonotal) of smooth setae of which any of z2, s5, J4 or Z4 asymmetrically on/off shield margin; of shield setae, j3 (60–65), z4 (62–75), s4 (62–75), s5 (60–70) of similar lengths; j4 (45–65), j5 (40–60), j6 (31–53), z5 (33–52), J1–J4 somewhat shorter but not tiny (22–40), with z4 and s4 (62–75), s5 (60–67), Z1 (57–70) and Z4 (60–72) similarly longer. Soft dorsal peripheral cuticle with setae j1 (47–52), z1 (40–43), z2 (54) anteriorly, s6 (60–65) and a hypertrichy of 12–15 r -setae (50–62) anterolaterally, and Z2 (57–65), S1–S4 (55–70), S5 (75) and hypertrichy of 10–15 R -setae (52–65) posterolaterally. Dorsal shield with complement of nine pairs of discernible pore-like structures (four podonotal, five opisthonotal), of which three pairs (one podonotal, two opisthonotal) superficially appear secretory (gland pores) and six pairs (three podonotal, three opisthonotal) non-secretory (poroids).

Adult male ( Fig. 9A ). Idiosoma 580–612 long, 370–437 wide between insertions of setae r3; dorsal shield 520– 532 long, greatest width 337–365 at level between insertions of s6 (n=3), unornamented, with irregular but entire lateral margins, which widened to hold setae j1, z1, s3 anteriorly and Z2, S2 laterally. Dorsal shield with usually 23 to 25 pairs (14 or 15 podonotal, 9 or 10 opisthonotal) of smooth setae of which s6, Z2, Z4 often asymmetrically off shield margin; relative setal lengths much as in female, except J1–J4 shorter (20–30) compared to j4–j6 (37–57); other setal lengths j1 (35–42), j3 (58–68), z1 (25–32), z2 (55–62), z4 (60–65), s4–s6 (57–67), about eight r -setae (40–52), Z1–Z2 (60–65), Z3 (60–72), Z4–Z5 (65–77), S1–S3 (62–68), S4 (65–72), S5 (68–85), about ten R -setae (52–68). Dorsal shield with complement of eleven pairs of discernible pore-like structures (4 podonotal, 7 opisthonotal), most of which are difficult to discern whether secretory (gland pores) or non-secretory (poroids).

Idiosomatic venter. Adult female ( Fig. 7B ). Tritosternum base 23–25 long and 23–26 wide at its base, with paired ciliate laciniae fused for about 5–7 of entire length (ca 100). Presternal area with a delineated, lightly sclerotized, streaky region without clear platelets. Sternal shield mid-length (98–100) much less than narrowest width (148–155) between legs II, faintly scabrous; sternal setae st1, st2 attenuated, of similar length (55–67). Sternal setae st3, st4 similarly attenuated (50–60), with poroid iv3 on soft cuticle. Endopodal strips rudimentary alongside coxae III–IV. Epigynal shield smooth, its rounded hyaline anterior margin reaching or slightly overlapping posterior edge of sternal shield, its lateral margins slightly widened at insertions of attenuated setae st5 (57–65), its posterior margin slightly convex; shield 205–223 long, 60–63 at narrowest width between legs IV, 77–85 at widest level at setae st5. Postgenital groove undeveloped, without elements of platelets. Opisthosomatic venter with two pairs of weak metapodal platelets, lateral pair 30–32 x 8–12, inner pair 17–20 x 7–10. Anal shield subrhomboidal, lightly lineated; shield greatest width (128) slightly less than length (132–147, including cribrum 20), cribrum width 75; postanal seta (60–70) about 1.5 longer than paranal setae (37–50) inserted in line with posterior margin of anus. Soft cuticle with ca 15 pairs of opisthogastric setae of similar lengths (43–57) flanking anal shield, JV5 not distinguishable posterolaterally; of four or five pairs of pore-like structures on soft cuticle, paranal gland pores well separated from lateral margins of anal shield. Peritrematal shield and peritreme extending barely to mid-level of coxae III; peritreme with embellished internal chambers ( Fig. 7F ), its length 45, including stigmatal diameter of ca 22; peritrematal shield continuous posteriorly with short exopodal strip extending partly around posterior margin of coxa IV. Exopodal elements absent alongside coxae II–IV. Spermathecal apparatus with simple, elongate (90–95) tubulus leading from solenostome between bases of legs III–IV to an unpaired sacculus ( Fig. 7G ).

Adult male ( Fig. 9B ). Tritosternum formed as in female, with laciniae fused for only about one-tenth of entire length (ca 100). Presternal region with two pairs of small platelets anterior to genital opening. Sternitigenital shield length 245–255, narrowest widths 115–125 at level between coxae II, 100–107 between coxae III; posterior margin width 62–80; shield unornamented, narrowly separated from anterior edge of endopodal strips beside coxae IV; alveoli of setae st5 on corners of irregularly shaped posterior margin of shield; setae st1–st5 similarly attenuated as in female, st1 and st5 (53–67) slightly longer than st2–st4 (50–65). Paragenital poroids on soft cuticle flanking posterolateral corners of sternitigenital shield. Postgenital groove not developed. Inguinal region with five or six pairs of sigilla indicating unusual muscular development connected with deeply retractable bases of chelicerae. Anal shield similar in rhomboid form, slight ornamentation as in female, its anterior and lateral apices more irregular; shield longer (130–135, including cribrum ca 15–20 long) than wide (103–112, cribrum width 60–62); postanal seta (57–62) ca 1.5 to nearly twice as long as paranal setae (30–40). Soft cuticle with about 15 pairs of opisthogastric setae flanking anal shield; these setae gradually slightly lengthening posteriorly, those in JV1-JV2-ZV1-ZV2 area 43–48, those mid-posteriorly 45–62, and posterior-most JV5 (62–75); of five pairs of pore-like structures evident on soft cuticle, paranal gland pores well separated from lateral margins of anal shield. Short but embellished peritremes (40–45), and peritrematal shield with brief poststigmatal exopodal extension as in female ( Figs 9J , 20B ).

Gnathosoma . Adult female. Gnathotectum anterior margin obtusely smoothly triangular, with a blunt, medial anterior projection ( Figs 7H, J ). Cheliceral shaft, excluding basal section, 155–158 long; fixed digit with blunt apex, vestige of pilus dentilis, and a vestigial tooth midway on paraxial face ( Fig. 7E ); movable digit (35) with three coarse teeth flared laterally ( Figs 7D, E ). Subcapitulum with 11 to 13 rows of deutosternal denticles, each similarly narrow, with a single denticle, or any row occasionally with two contiguous denticles ( Fig. 7C ); subcapitular setae hp1 (60–63) and slightly longer hp3 (75–80) similarly attenuated, hp2 (20–25) and capitular setae pc (22–27) similarly short, but pc strongly bulbiform basally. Corniculi somewhat hyaline, blade-like, bluntly pointed apically, moderately short (35–38), closely adjacent to similarly short tubular salivary styli (ca 37–40) and longer, slender, finely fimbriated internal malae (87–90). Palp length 150–162; palptrochanter setae dissimilar in length, v1 22–25, v2 38–52; palpus with femoral seta al-1 and genual setae al-1, al-2 blunt, not spatulate; palpfemoral seta d (30–32) slightly spine-like, shorter than femur length; palptarsal apotele two-tined.

Adult male. Gnathotectum with simple, triangular anterior margin much as in female, but sides acutely angular, tapering to single median projection ( Fig. 9E ). Cheliceral shaft, excluding short basal section 120–125 long; fixed digit straight, short (30–42) more slender and pointed than in female; movable digit bluntly pointed beyond apex of fixed digit, reduced, amalgamated into spermatodactyl base which extends anteroventrally; spermatodactyl greatly elongated, sinuous, gradually tapering, 425–465 long, without elaborations ( Figs 9C, D ). Unlike female, corniculi shorter (22–25), more truncated apically, greatly exceeded by elongated salivary styli (63–70), and capitular setae pc nearly setiform, slightly bulbous basally. Form and length of other gnathosomatic structures much as in female: internal malae (120–130); setae hp1 (62–63) and hp3 (65–77) much longer than hp2 (18–20) and pc (23–25); deutosternum with 11–12 rows of mostly contiguously doubled denticles ( Fig. 9F ). Palp length 160–165; palpfemoral seta d (25–30) similar in form as in female.

Legs. Adult female ( Figs 8 A–H). Legs I (425–450) nearly as long as legs IV (460–470), and about 0.8 as long as dorsal shield; other leg lengths (excluding pretarsi): II (375–380), III (385–400). Leg I pretarsus (37–38) to apex of pulvillus, tarsus (115–125) about 1.7–2.1 longer than each of the similarly long femur (67–70), genu (53–63), and tibia (60–65). Legs II–IV with tarsus/tibia length ratios about 1.8–2.3. Complement of setae on leg segments holotrichous for femora II-III-IV (11-6-6) as presented for the genus, but femur I with 11 setae, lacking pv-2; chaetotactic formulae for genua I-II-III-IV: 11 (2-3/1, 2/1-2), 11 (2-3/1, 2/1-2), 9 (2-2/1, 2/1-1), 10 (2-2/1, 3/1-1); tibiae: 11 (2-3/1, 2/1-2), 10 (2-2/1, 2/1-2), 8 (2-1/1, 2/1-1), 10 (2-1/1, 3/1-2); genu I and tibia I without pd-3 and av-2; tibia III without pl-2. Tarsus I with seta md (55–60) longer than other setae on basal half of segment ( Fig. 8B ); dorsal subapical cluster of sensory structures with a cluster of three, tightly adjacent, tiny, pointed setae surrounded by three solenidia, one of which biramous ( Fig. 8C ). Tarsi II–IV with a finely serrated ventroapical ridge basal to a short, rounded process ( Figs. 8E, H ), and with dorsoapical setae (d-1) (12–15) nearly half as long as pretarsus to base of claws (ca 25). Coxal seta av on leg I and pv on legs II, III strongly bulbiform basally, like subcapitular pc; coxa IV seta v setiform. Trochanters I–IV with basal seta pv-2 attenuated (50–55), and III–IV ad slightly spinelike. Tarsus IV with apical seta pv-1 asymmetrically shorter, bluntly spine-like, than attenuated av-1 ( Fig. 8H ). Legs I–IV each with particular setae enlarged, spine-like: I Fe ad-1 (65), pd-1 (45); I Ge ad-1 (50–60), pd-1 (55–58); II Fe ad-1 (57–63), pd-1 (47–48); II Ge ad-1 (50–52), pd- 1 (47–48); II Ti pd-1 not differentiated; III Fe ad-1 (52–54); III Ge ad-1 (57–62), pd-1 (50–52); III Ti ad (42), pd-1 (30); IV Fe ad-1 (57–62), pv not differentiated; IV Ge ad-1 (66–68), pd-1 (62–65); IV Ti ad-1 (53–60), pd-1 (55–60). Relative lengths of other leg setae as shown in Figs 8A, D , F–H.

Adult male ( Figs 10 A–D). Legs similar in relative dimensions to those of female; lengths I 415–430, II 360– 378, III 375–400, IV 440–470. Legs I and IV without dimorphism in thickness of segments, legs II and III thickersegmented than in female. Leg I with form and relative sizes of setae similar to those of female, except femur I with 12 setae, including pv-2, and with av slightly bulbiform. Legs I–IV retain attenuated form of basal seta pv-2 of trochanter. Legs II–IV with particular setae dimorphically modified: leg II Fe av spine-like; Ge av, Ti av, and tarsus mv, av-2, av-1, pv-1 bulbiform; leg III coxal pv simple; Tr ad markedly arched; Fe al less spine-like, and tarsus mv, av-2, pv-2, av-1, pv-1 bulbiform; leg IV Tr ad markedly arched, av, pv-1 thickened, more spine-like; Fe pd, pl more spine-like; Ge pd-2, pd-3 and Ti pl-1 larger, more spine-like; tarsus av-2, pv-2, av-1, pv-1 bulbiform. Lengths of spine-like setae similarly enlarged as in female: I Fe ad-1 (48–62), pd-1 (37–45); I Ge ad-1 (45–52), pd-1 (52–60); II Fe ad-1 (55–65), pd-1 (42–49); II Ge ad-1 (38–45), pd-1 (50–57); II Ti pd-1 not differentiated; III Fe ad-1 (40–50); III Ge ad-1 (43–53), pd-1 (47–50); III Ti ad (35), pd-1 (27); IV Fe ad-1 (57–70), pv not differentiated; IV Ge ad-1 (67–70), pd-1 (67–75); IV Ti ad-1 (55–65), pd-1 (60–67).

Deutonymph. Idiosomatic dorsum ( Fig. 11A ). Idiosoma 475–500 long, 362–365 wide between insertions of setae r3 (n=3); dorsal shields weakly sclerotized, unornamented, faintly divided by transverse strip of soft cuticle between setae j6 and J1. Podonotal shield 210–225 long, ca 215–225 wide at its greatest width at level of setae s5, with ca 12 pairs of setae (j1, j3–j6, z1, z2, z4. z5, s4–s6); opisthonotal shield ca 175–195 long and 175–205 at widest level near setae Z2, with nine pairs of setae (J1–J4, Z1–Z5). Soft cuticle with ca 25 pairs of dorsolateral setae, including uncertain homologies of the s -and S -series among hypertrichous elements of the marginal r–R -series ( Fig. 11A ). Form and relative lengths of dorsal setae more homologous than in adult, many of them slightly barbed as shown in Fig. 11A ; other than z1 clearly shortest (18–28), setal lengths range from j1 (35–40) to Z5 longest (65). Peritreme similarly short (length 37–38, including stigma diameter 15–17) and embellished as in adult ( Fig. 11D ).

Idiosomatic venter ( Fig. 11B ). Tritosternum similar in form to that of adult, with laciniae fused for only about 5–10 of entire length (87–92 excluding base). Presternal area devoid of ornamentation. Intercoxal shield 182–200 long, weakly sclerotized, bearing five pairs of setae and three pairs of poroids, seta st5 sometimes barely off shield edge; setae st1–st4 (43–50) slightly shorter than st5 (52–57); paragenital poroids on soft cuticle flanking posterior edge of shield. Opisthogaster with oboval, smooth anal shield; length 95–100, greatest width 75–80; shield with postanal seta (45–50) about 1.3 longer and stouter than paranal setae (35–37). Soft cuticle with hypertrichous complement and relative lengths of setae flanking anal shield much as in adult, with ca 15 pairs of slender opisthogastric setae flanked by 5–10 pairs of thicker R -marginal setae of similar lengths (40–47), the more posterolateral elements of them thicker, blunter.

Gnathosoma . Gnathotectum, chelicerae, palpal structures, and subcapitulum much as in female, except setal pair pc simple. Cheliceral shaft, excluding basal section, 132–138 long; movable digit 28–35 long. Subcapitulum with 11 to 13 rows of deutosternal denticles formed as in female; subcapitular setae hp1 (45–50) and slightly longer hp3 (52–58) similarly attenuated, hp2 (13–15) and capitular setae pc (15–18) similarly short. Corniculi (25–30) formed as in female, exceeded by elongated tubular salivary styli (45–50) and even longer, slender, finely fimbriated internal malae (70–75). Palpus length 127–133; palptrochanter setae v1 12–15, v2 30–34; palpgenual seta d (22–23) slightly spine-like.

Legs. Leg lengths, excluding pretarsi, I (325–350), II (278–312), III (287–315), IV (335–360). Leg setation with normal deutonymphal complement of setae as given for adult female, with occasional variabilities. Tarsi II to IV with ventroapical process and pretarsal structures formed as in female; apical dorsosetal processes (d-1) relatively short (9–10, about 0.6 as long as pretarsus to base of claws 13–15) as in female; form and relative lengths of setae much as in female, with particular setae enlarged, spine-like: I Fe ad-1 (45–50), pd-1 (30–33); I Ge ad-1 (38–45), pd-1 (37–43); II Fe ad-1 (50–53), pd-1 (32–33); II Ge ad-1 (35–38), pd-1 (34–35); III Fe ad-1 (45–50); III Ge ad-1 (42–43), pd-1 (37–40); IV Fe ad-1 (47–52); IV Ge ad-1 (45–20), pd-1 (40–48); IV Ti ad-1 (40–47), pd-1 (36–42). Tarsus IV with asymmetry in lengths of setae (v-1) as in female. Claws of all legs strongly formed (length ca 13), projecting freely from pulvillus (as in Fig. 19E ), in contrast to adult and protonymph.

Protonymph. Idiosomatic dorsum ( Fig. 12A ). Idiosoma 370 long, 268 wide at widest level of setae r3, r5 (n = 1), with weakly sclerotized dorsal shields; podonotal shield 200 long and 178 at widest level between setae s4, with ten pairs of setae (j1, j3–j6, z1, z2, z4, z5, s4); pygidial shield ca 80 long and 152 at widest level between setae Z4, with five pairs of setae (J3, J4, Z4, Z5, S 5). Soft cuticle with ca 12 pairs of podonotal setae, indicating some hypertrichy among s -and r -series, and 12–14 pairs of opisthonotal setae including J1, J2, Z1–Z3 and some hypertrichy among S -and R -series). Form and relative lengths of dorsal setae more homologous than in adult, many of them stout, blunt, but smooth as shown in Fig. 12A ; on podonotal shield, other than z1 clearly shortest (15), setal lengths range from z2 (20) to j6 slightly longest (28); on pygidial shield, setal lengths range from J4 (12) to Z5 (23); setal lengths on soft cuticle of similar range length (12–20). Peritreme short (length 32, including stigma diameter 13), with slight indication of embellishment as formed in adult.

Idiosomatic venter ( Fig. 12B ). Tritosternum formed much as in adult, with laciniae fused for only about 8 of total length (72). Presternal area devoid of ornamentation. Sternal shield not clearly delineated, distance between bases of setae st1 to st3 ca 105, with setae st1–st3 (35–40) and poroids iv1, iv2; setae st5 short (20), inserted closely anterior to opisthogastric setae, and flanked by paragenital poroids (one of which asymmetrically formed into minute seta, Fig. 12B ). Anal shield smooth, oval, length ca 58 including cribrum, width 55 (cribrum strip width 35); postanal seta (37) about 1.3 longer than paranal setae (28). Soft opisthogastric integument with four pairs of setae, JV1 (37), JV2 (30), ZV2 (22), JV5 shortest (15).

Gnathosoma . Gnathotectum with triangular smooth anterior margin, tapering to single blunt projection ( Fig. 12D ). Cheliceral shaft, excluding short basal section 120 long; fixed digit straight, bluntly bifid apically; movable digit (28) formed much as in female, with three teeth flared laterally ( Fig. 12C ). Subcapitulum with 12 similarly narrow rows of deutosternal denticles, most with a single denticle but basal and apical rows with two contiguous denticles ( Fig. 12E ); subcapitular setae hp1 attenuated (30), longer than hp3 (15), hp2 and capitular setae pc similar in short (12) simple form. Corniculi hyaline, blade-like, narrowly rounded apically, moderately short (25), slightly exceeded by tubular salivary styli (40). Palpus length 115, with normal protonymphal complement of setae; single palptrochanter seta 5; palpfemoral seta d (18) slightly spine-like.

Legs. Leg lengths, excluding pretarsi, I 265, II 225, III 225, IV 255. Leg chaetotaxy of protonymph typical of general holotrichous pattern presented for free-living Gamasina by Evans (1963): coxae, 2-2-2-1; trochanters, 4-4- 4-4; femora, 10 (2 2/1 2/1 2)-8 (1 2/1 2/1 1)-5 (1 2/1 1/0 0)-4 (1 2/0 1/0 0); genua, 8 (1 2/1 2/1 1)-6 (1 2/0 2/0 1)-6 (1 2/0 2/0 1)-5 (1 2/0 2/0 0); tibiae, 8 (1 2/1 2/1 1)-7 (1 1/1 2/1 1)-7 (1 1/1 2/1 1)-7 (1 1/1 2/1 1). Legs I–IV each with particular setae enlarged, spinelike, much as noted for deutonymph. Tarsi II–IV with ventroapical process, pretarsal structures and weakly developed claws formed as in female. Tarsus IV with asymmetry of setae (v-1) as in female.

Larva. No material available (adult females nymphiparous).

Type material. All specimens were collected by E.E. Lindquist or E.A. Sidorchuk from four adult crickets, identified as “ Craspedogryllacris sp. 21” by David Rentz, collected from low coastal heathland of the southwestern region of Australia (Western Australia, 33°33’S, 115°01’E, Cape Naturaliste, West Coast Road, 10 Dec. 1995 and 13 March 1999, coll. R.Hale). HOLOTYPE: nymphiparous adult female, ex Craspedogryllacris sp. 21 #3, 10 Dec. 1995, deposited in the Australian Natural Insect Collection , Canberra .

PARATYPES: one female, one male, one deutonymph with same data as holotype; one male, two deutonymphs, one protonymph, with same data as holotype except ex C. sp. 21 #4; one male, with same data as holotype except ex C. sp. 21 #2, 13 May 1999; seven deutonymphs, with same data as holotype except ex C. sp. 21 #1, 13 May 1999; of the seven deutonymphs ex cricket #1, five were on the wings, one on the abdominal venter, and one on the trochanter of leg III. One female, one male, three deutonymphs, one protonymph deposited in the CNC, Ottawa; one male, two deutonymphs deposited in the ZIN, St.- Petersburg; one male, two deutonymphs deposited in the UMMZ, Ann Arbor; one male, three deutonymphs deposited in the ANIC, Canberra.

Etymology. The specific name is a compound adjective derived from the Latin words multi-, meaning “many” or “multiple” and setosus, meaning “bristly” or “hairy”, and refers to the hypertrichous condition of the idiosoma of all instars of this species. As the gender of the generic name Berlesia should be feminine (Sauro Simoni, personal communication), the specific name is accordingly multisetosa .

Differential diagnosis. Adults, deutonymphs and protonymphs of B. multisetosa are immediately distinguished from those of other known species in being hypertrichous on the lateral and opisthogastric regions of the idiosoma. Also, those setae are of similarly moderate lengths, none being little more than twice as long as the others. Unlike B. hospitabilis and B. vorontsovi , paravertical setae z1 and setae J2 are present. On adults, the genu and tibia of leg I each have 11 setae, with only two ventral setae (av-1, pv-1), whereas those of hospitabilis and vorontsovi each have the third ventral seta (av-2) present. The asymmetrically spine-like form of subapical seta pv-1 on tarsus IV is also distinctive.

Remarks. This is the first species of the genus Berlesia to be described with all active instars having peripherally hypertrichous idiosomas.

Figures

FIGURE 7. Berlesia multisetosa n. sp., female. A, idiosoma dorsally; B, idiosoma ventrally; C, subcapitulum; D, E, chelicerae, dorsally, in working position, E, enlarged detail; F, peritreme; G, sperm access system; H, J, variability of gnathotectum outline. A, B, G to same scale, C, D, F, H, J to same scale. Abbreviations: co, corniculus; d lf, dorsal lyrifissure; d s, dorsal seta; id, lyrifissure; gp, glandular pore; lb, labrum; mi, internal mala; p, unidentified pore-like structure; p d, pilus dentilis; ss, salivary stylus; t, tooth; td, tendon; others denote setae.

FIGURE 8. Berlesia multisetosa n. sp., female, legs.A, C, F, G, legs dorsally except separated coxae ventrally:A, leg I, subapical tarsal setae shown with circles; B, tarsus I dorsally; C, detail of sensory cluster on tarsus I; D–G, legs dorsally except separated coxae ventrally: D, leg II; E, detail of tarsus II tip ventrally; F, leg III, tarsus partly shown; G, leg IV, tarsus partly shown; H, tarsus IV ventrally. A, D–H to same scale.

FIGURE 9. Berlesia multisetosa n. sp., male. A, idiosoma dorsally; B, idiosoma ventrally, partial; C, D, chelicera and spermatodactyl, C, dorsal aspect, D, ventral; E, gnathotectum; F, subcapitulum and palpus; G, H, palp tarsus, G, ventral aspect, H, dorsal, enlarged; J, peritreme.A, B to same scale, C–G to same scale, H, J to same scale.Abbreviations: ar, arthrodial envelope; f d, fixed digit; m d, movable digit; others as in Fig. 7.

FIGURE 10. Berlesia multisetosa n. sp., male, legs, coxae detached. A, leg I ventrally, most setae of tarsal subapex, beyond transverse dotted line, omitted; B, leg II ventrally; C, leg III ventrally; D, leg IV ventrolaterally. All to same scale.

FIGURE 11. Berlesia multisetosa n. sp., deutonymph. A, idiosoma dorsally; B, idiosoma ventrally, partial; C, apical half of chelicera, ventrally, at resting position; D, variations of peritremes; E–G, variability of gnathotectum outline.A, B to same scale; C, E–G to same scale.

FIGURE 12. Berlesia multisetosa n. sp., protonymph. A, idiosoma dorsally; B, idiosoma ventrally, partial; C, right chelicerae dorsally, at resting position; D, gnathotectum; E, subcapitulum. A, B to same scale, C–E to same scale.

FIGURE 19. Berlesia hospitabilis n. sp.,details. A–C, gnathosomas ventrally: A, female, asterisks showing lateral borders of cheliceral bases inside; B, male; C, deutonymph (same as in D, J–K) with arrowed pharate female structures inside; D, copula, slightly parted in lactic acid, male artificially shrunk; E, F, deutonymphal ambulacra: E, leg II; F, leg I; G, male ambulacrum of leg I (similar in all legs of adult male, female and of protonymph); H, spermatophore; J, same copula, when first found on host’s wing; K, detail of D, area of remaining contact. Arrowheads indicate structures of pharate female (black) and her deutonymphal skin (white). Abbreviations: cxIII, cxIV, leg coxae III and IV; sd, spermatodactyl; sf, spermatophore; st, solenostome.

FIGURE 20. Berlesia multisetosa n. sp., male. A, ventral habitus, showing chelicerae fully withdrawn into idiosoma, yet spermatodactyl apices still protruding from subcapitulum; B, detail of peritreme; C–E, details of appendage apices: C, palp genua, tibiae, tarsi flanking tips of salivary stylets and internal malae; D, E, pretarsus and adjacent tarsal structures, showing reduced claws enveloped by pulvillus, D, leg II, E, leg I. Abbreviations: lb, labrum; mi, internal mala; sd, spermatodactyl; ss, salivary stylus; pv, pulvillus; others denote setae.