(Figs 38–55)
Halozercon karacholana Wiśniewski et al., 1992: 180
Material examined: NORTH ALTAI: 63 females, 15 males, Shebalino District, Cherginskii range, Belok Mukhor- Cherga Mt., 900 m a.s.l., 51 ̊21ʹ N, 85 ̊90ʹ E, Pinus sibirica - Larix sibirica forest, in litter, 8 August 2013, leg. I.I. Marchenko; 43 females, 8 males, 2 deutonymphs, 1 protonymph, same data, 21 July 2011, leg. I.I. Marchenko; 8 females, 2 males, Maima District, Malaja Sinyuha Mt., 1190 m a.s.l., 51 ̊49ʹ N, 85 ̊21ʹ E, Betula sp.–– Abies sibirica - Pinus sibirica forest, in litter, 28 August 2016, leg. I.I. Marchenko; NORTH-EAST ALTAI: 21 females, 14 males, 1 deutonymph, Turochak District, Altyn-Too Range, Archa Mt., 1700 m, 51 ̊32ʹ N, 87 ̊26ʹ E, alpine tundra with Betula rotundifolia, in litter and moss, 9 August, 2002, leg. E.V. Sleptsova; 35 females, 13 males, same geographical data, 14 August 2007, leg. I.I. Marchenko; 3 females, same geographical region, 900 m a.s.l., Abies sibirica –– Picea obovata –– Pinus sibirica forest, in litter, 21 July 2010, leg. I.I. Marchenko; 13 females, 7 males, 9 deutonymphs, 1 protonymph, same geographical region, 1100 m a.s.l., Pinus sibirica forest, in litter, 26 June 2006, leg. N.V. Vladimirova; 2 females, same geographical region, Evrechala Mt., 1800 m, 51 ̊29ʹ N, 87 ̊24ʹ E, under Pinus sibirica, in litter, 12 August 2002, leg. E.V. Sleptsova; 32 females, 10 males, 9 deutonymphs, 5 protonymphs, Turochak District, environs of Verh-Biisk village, 390 m, 52 ̊03ʹ N, 87 ̊05ʹ E Betula sp.–– Populus tremula forest, in litter, 9 August 2006, leg. I.I. Marchenko; 6 females, 3 males, 2 deutonymphs, Turochak District, environs of Kebezen village, 450 m a.s.l., 51 ̊54ʹ N, 87 ̊06ʹ E Pinus sylvestris –– Betula sp. forest, in litter, 8 August 2006, leg. I.I. Marchenko; 5 females, 2 males, Solton Ditrict, environs of Saidyp village, 500 m a.s.l., 52 ̊34ʹ N, 86 ̊34ʹ E Betula sp.–– Populus tremula forest, in litter, 22 August, 2006, leg. I.I. Marchenko; WEST ALTAI: 2 males, Ust- Koksa District, Krasnaya Mt., 1700 m a.s.l., 50 ̊05ʹ N, 85 ̊11ʹ E, Picea obovata –– Abies sibirica –– Pinus sibirica forest, in litter, 23 July 2015, leg. I.I. Marchenko; CENTRAL ALTAI: 76 females, 18 males, 1 deutonymph, Seminskii Range, Sarlyk Mt., 2000 m a.s.l., 51 ̊04ʹ N, 85 ̊44ʹ E, Pinus sibirica forest and alpine tundra, in litter and moss, 6 July 2013 and 13 June 213, leg. I.I. Marchenko; 13 females, 2 males, 3 deutonymphs, Chemal district, near of Karakol lakes, 1300 m a.s.l., 51 ̊30ʹ N, 86 ̊23ʹ E, Pinus sibirica –– Abies sibirica forest, in litter, 28 July 2012, leg. I.I. Marchenko; 31 females, 4 males, Severo-Chuiskii Range, Aktru Mt., 1600 m a.s.l., 50 ̊03ʹ N, 87 ̊40ʹ E, Pinus sibirica forest, in litter, 21 July 2013, leg. I.I. Marchenko; 12 females, 9 males, 4 larvae, Kuraiskii Range, 1840 m a.s.l., 50 ̊19ʹ N, 87 ̊41ʹ E, Abies sibirica –– Pinus sibirica –– Larix sibirica forest with moss, in moss, 26 August 2017, leg. I.I. Marchenko; 1 female, 1 deutonymph, Ulagan District, environs of Saratan village, along Baschkaus river, 1500 m a.s.l., 50 ̊25ʹ N, 80 ̊27ʹ E, Larix sibirica –– Picea obovata –– Pinus sibirica forest, in litter, 5 July 2015, leg. I.I. Marchenko; 8 females, 1 male, Kosch-Agach District, Yuzhno-Chuiskii Range, near of Tara river, 2250 m a.s.l., 49 ̊54ʹ N, 88 ̊23ʹ E, Larix sibirica forest and alpine tundra with Betula rotundifolia, in litter and moss, 17 July 2009, leg. I.I. Marchenko; KEMEROVO REGION, GORNAYA SCHORIA: 22 females, 5 males, Biiskaya Griva Mt., 750 m a.s.l., 52 ̊26ʹ N, 88 ̊17ʹ E, Betula sp. forest and Pinus sibirica forest, in litter, 28 September, leg. L.A. Trilikauskas; 39 females, 5 males, 4 deutonymphs, same geographical region, Pustag Mt., 600 m a.s.l., 52 ̊58ʹ N, 87 ̊44ʹ E, Abies sibirica –– Picea obovata –– Pinus sibirica forest, in litter, 18 June 2013, leg. I.I. Marchenko; 25 females, 8 males, 1 deutonymph, same geographical region, environs of Kuzedeevo village, 270 m a.s.l., 53 ̊22ʹ N, 87 ̊13ʹ E, Abies sibirica –– Populus tremula forest, in litter, 20 June 2013, leg. I.I. Marchenko; TUVA: 4 females, Mongun-Taiga Range, environs of Hindigtic-Kol lake, 2300 m, 50 ̊21ʹ N, 89 ̊49ʹ E, alpine tundra, in moss, 3 July 2013, leg. A. Kostyunin; KHAKASSIA: 6 females, West Sayan Range, 40 km from Abaza city, along Korasibo river, 700 m a.s.l., 52 ̊35ʹ N, 90 ̊08ʹ E, Abies sibirica –– Picea obovata –– Pinus sibirica forest, in litter, 25 June 2013, leg. I.I. Marchenko; BAIKAL REGION, BURYATIA: 8 females, 3 males, Khamar-Daban Range, Baikalskii Nature Reserve, 700 m a.s.l., 51 ̊33ʹ N, 105 ̊09ʹ E, Abies sibirica –– Picea obovata –– Pinus sibirica forest, in litter, 14 August 2014, leg. L.V. Petrozhitskaya; 5 females, 2 males, 2 deutonymphs, same geographical data, 1000 m a.s.l., alpine meadow, in soil, 13 August 2014, leg. L.V. Petrozhitskaya; 1 male, same geographical data, environs of Tankhoi village, 470 v a.s.l., along estuary of Pereemnaya river, Betula sp.–– Picea obovata –– Pinus sylvestris forest, in litter, 10 August 2014, leg. I.I. Marchenko; ZABAIKALSKII KRAI: 1 male, 2 deutonymphs, 1 protonymph, Kyra District, Sokhondinskii Nature Reserve, Verhnii Bukukun cordon, 1800 m a.s.l., 49 ̊41ʹ N, 111 ̊12ʹ E, Larix dahurica –– Pinus sibirica forest, in litter, 28 July 2016, leg. I.I. Marchenko.
Female (Figs 38–47, n=10)
Dorsal idiosoma (Fig. 38). Idiosoma suboval, 450–485 long and 270–310 wide. Anterior margin of podonotum curved ventrally to form a vertex, with two pairs of setae inserted on the ventral side of idiosoma: j1 stout, slightly pilose (15–17) and z1 (10–11) thorn-shaped, usually smooth, but in some specimens slightly pilose, located latero-posteriorly to j1 setae. Podonotal shield extensively neotrichous, with about 69–74 setae on each side (including marginal etae). Surface of podonotum ornamented by network of small tubercles. Podonotum setae of lateral regions pilose, 15–20 long, in central region—pilose or some setae smooth, 10–18 long. Four pairs of porelike structures (glands) po1–po4 are present, po1 inserted in vertex, close to marginal setae, visible from ventral side of idiosoma. Opisthonotum extensively neotrichous, with about 75–78 densely pilose setae (22–25), with network of small tubercles, with four pairs of pore-like structures (glands) Po1–Po4, glands Po4 located in posterior margin of ventri-anal shield. All dorsal setae surrounded by basal rings, particularly enlarged in opisthonotum. Marginal dorsal setae of r–R series neotrichous, inserted on tubercles, elongated, 23–25 long, slightly curved and pilose on one side.
Ventral idiosoma. (Figs 39–41). Base of tritosternum 24–26 long and 13–15 wide, paired pilose laciniae free from each other along entire length, 55–60 long, basal ends of laciniae inserted into the basal segments, as in cups (Fig. 39). Presternal (jugular) platelets indistinct, weakly sclerotised, with pair of St1 setae (18–20) (Fig. 40). First pair of sternal platelets suboval, irregular, 18–25 long and 15–18 wide, with pair of St2 setae (18–20) and pair of lyrifissures iv2; with a folded surface. Median area between sternal platelets ornamented by rounded tubercles. Second pair of sternal platelets suboval, irregular, with folded surface, 23–28 long and 18–20 wide, with two pairs of setae St3, St4 (15–18) and one pair of lyrifissures iv3, inserted anteriorly to St3 setae. Genital shield 63–70 long and 37–50 wide at level of St5 setae, irregularly-shaped, with individual shape in each specimen; expanded anteriorly and posteriorly; anterior margin with fine folding; with pair of genital setae St5 (15–18) and lyrifissures iv5 on lateral edges of shield or in soft cuticle laterad of shield. Subtriangular membrane surrounding the genital shield (Fig. 41). Genital shield with complex form: one median unpaired triangular cap and two pairs of fine sclerites—short anterior pair and elongate posterior pair. Endo-, exopodal and metapodal shields absent. Adgenital gland pores gv2 are multiple, dispersed over the surface: one pair of glands opening located in soft cuticle posterolaterad to genital shield, three other pairs of openings located in ventri-anal shield. Peritrematal shields fused anteriorly and with dorsal shield laterally; strong sclerotised, postero-lateral ends are drawn back in the oblique angle; with festoon reticulation along entire length. Five pairs of pore-like structures inserted in peritrematal shield: enlarged gp1, gp2 and small ip1–ip3. Peritremes 85–105 long, slightly curved or straight, reaching anterior margins of coxae III or level of posterior 1/3 part of coxae II; with internal cell structure. Arch of vertex with two dorsal setae j1 and z1. Ventri-anal shield broad, 135–150 long and 238–275 wide; fused to opisthonotal shield around the outer edge; ornamented by small tubercles and festoon reticulation; with 7–8 pairs of smooth opisthogasric setae (15–18) inserted asymmetrically, including Jv1 and Zv1 at anterior margin, or Jv1 located in soft cuticle in some specimens; and with 8–9 pilose opisthonotal setae (18–20) with enlarged basal rings at postero-lateral margins of shield. The most posterior marginal row with elongated and smooth setae (23–25) on raised tubercles duplicates opisthonotal rows of setae. Anal area with slightly pilose pre-anal setae and smooth post-anal seta (11–13), anal opening 23-25 long; with two lyrifissures on each valve; cribrum located posteriorly of post-anal seta. Pair of glands gv3 located antero-laterad to para-anal setae.
Gnathosoma. (Figs 42–47) Fixed digit of chelicera 45–48 long, with five teeth in addition to apical hook and leaf-shaped pilus dentilis (Figs 42, 43); movable digit the same length (45–48) with three teeth in additional to apical hook. Chelicera with long dorsal seta (16–18), lateral (antiaxial) and dorsal (paraxial) lyrifissures and arthrodial corona. Epistome subtriangular, with irregularly serrated edges and smooth pointed median projection (Fig. 44). Corniculi 25–26 long and 11–12 wide. Internal malae slightly longer than the corniculi with complex three-layer structure: acute elongated anterior projection with short lateral process; serrated structure on intermediate layer and pilose basal structure (Fig. 45). Deutosternal groove with 7–8 transverse denticulate rows (Figs 45, 46), with three paired transverse lines, the pair of arched transverse rows of large denticles located at the level of pc setae. Setae h1–h3 simple: h1 (34–35) longest, seta h2 (14–15) shorter then h3 (30–32); pc (15–16) serrated. Dorsal side of gnathosoma with pair of protruding wedge-shaped structures between palpal coxa and trochanter, lateral to epistome, connected with inner longitudinal chord (Fig. 47). Palpal chaetotaxy 2–5–6–12 –15, palp with five free segments; trochanter with seta al2 long and pilose in the distal third (27–32) and short, smooth al1 (13–15) (Fig. 45); palp genu with setae al1 and al2 pilose in distal third; palp apotele two-tined.
Legs. Lengths: I 285–310, II 225–245, III 220–232, IV 275–300 µ. Chaetotaxy of legs I–IV: coxae––2, 2, 2, 1; trochanters––6, 5, 5, 5, femora––13 (2 5/5 1), 11 (1 5/3 2), 6 (2 2/2 0), 6 (2 2/2 0); genua––13 (2 5/3 3), 11 (1 5/3 2) or 12 (1 5/3 3), 10 (2 4/3 1), 10 (2 4/3 1); tibiae––14 (2 6/4 2), 10 (1 4/3 2), 9 (2 4/2 1), 10 (2 5/2 1); tarsi–– II–IV 18 (3 7/5 3), 18 (3 7/5 3), 18 (3 7/5 3). All legs with pair of sclerotised claws and pulvillus with five lobes. Pretarsus of II–IV legs with ambulacral stalk, legs I with sitting claws. All legs ornamented by rows of tubercles, many setae at dorsal side on raised bases, some of them pilose. Coxae I–IV are closely grouped together; coxae I–II with splitting on dorsal side, coxae III–IV on anterolateral side. Coxae II–III with antero-dorsal spines: coxae II with large sharp spine, coxae III with small one.
Male. (Figs. 48, 49, n=10).
Dorsal idiosoma. Dorsal shield suboval in shape, 440–455 long and 255–300 wide. Ornamentation, porotaxy and chaetotaxy similar to that female.
Ventral idiosoma (Fig. 48). Base of tritosternum 23–25 long and 10–12 wide, laciniae pilose, 55–60 long. Presternal (jugular) platelets weakly sclerotised, with pair of St1 setae. First sternal platelet entire, 25–33 long and 45–50 wide, with pair of St2 setae and pair of iv2 lyrifissures, entire, ornamented with folds and grooves. Second sternal platelet entire, 25–27 long on median line and 37–45 wide at level of St3 setae; subchordate shape, ornamented by folds; with two pairs of setae St3, St4 and one pairs of lyrifissures iv 3 in antero-lateral margins; surrounds the genital opening. Genital opening located at level of coxae III, with pair of eugenital setae, covering by two platelets. Third sternal platelet entire, triangular shape, 25–30 long and 30–50 wide, with pair of St5 setae and pair of iv5 lyrifissures. Measurements of St1–St5 setae 11–17 µ. Endo-, exopodal and metapodal shields absent. Peritrematal shields similar to that female, with five pairs of pore-like structures: enlarged gp1, gp2 and small ip1- ip3. Peritremes similar to that female. Arch of vertex with two dorsal setae: slightly pilose j1 (15–16) and smooth thorn-shape z1 (10–11). Ventri-anal shield broad, 140–150 long and 250–265 wide, fused to opisthonotal shield around the outer edge; ornamented by network of small tubercles; with 6–7 pairs of opisthogasric smooth setae inserted asymmetrically (13–17) and 12–15 pairs of opisthonotal pilose setae (20–25) inserted posteriorly; with transverse split along the junction of opisthonotal shield to ventral side in some specimens. Adgenital gland pores gv2 are multiple, dispersed over the surface: one pair of glands located in soft cuticle posteriorly of coxae IV, other 1–3 pairs of openings located in ventri-anal shield. The most posterior marginal row with elongated smooth setae (22–25) on raised tubercles duplicates opisthonotal rows of setae as in female. Anal area with slightly pilose preanal setae and smooth post-anal seta 10–11 long, anal opening 23–24 long; with two lyrifissures on each valve; cribrum presented. Pair of glands gv3 located antero-laterad of para-anal setae.
Gnathosoma. Male chelicera lacking spermatodactyl. Fixed digit of chelicera 40–42 long, with two large basal teeth and 5–6 small distal teeth in addition to apical hook and leaf-shaped pilus dentilis (Fig. 49). Movable digit of chelicera 40–42 long, tridentate in addition to apical hook, with arthrodial corona at a base of digit. Chelicera with long dorsal seta, lateral (antiaxial) and dorsal (paraxial) lyrifissures. Epistome, corniculi, internal malae, hypostomal and palp structures as in female.
Legs. Lengths: I 255–285, II 225–240, III 215–235, IV 280-290 µ. Chaetotaxy and morphology of legs as in female. Leg II without sexual differences.
Deutonymph (Figs 50–51, n=10).
Dorsal idiosoma (Fig. 50). Dorsal shield suboval shape, 375–430 long and 240–275 wide. Anterior margin of podonotum curved ventrally to form a vertex, with pairs of setae j1 and z1. Dorsal shields extensively neotrichous, with network of small tubercles and porotaxy as in adults. Podonotal shield with about 67–69 setae on each side, opisthonotal shield with about 47–51 setae on each side.
Ventral idiosoma (Fig. 51). Pairs of St1, St4 and St5 setae inserted in soft cuticle of sternal region; pairs of St2 and St3 setae—in weakly sclerotised sternal platelets or in soft cuticle. Peritrematal shields narrow, fused with dorsal shield laterally, with festoon reticulation along entire length. Two pairs of pore-like structures visible: ip1 and gp1. Peritremes longer than in adults, reaching level of top spine of coxae II anteriorly; with internal cell structure. One pair of adgenital gland pores gv2 located in soft cuticle posteriorly to coxae IV. Opisthogastric area with 5–6 pairs of simple setae inserted in soft cuticle asymmetrically. Anal shield 65–87 long and 50–60 wide, with pair of Jv3, pre-anal setae and post-anal seta; pair of glands gv3 located antero-laterad of para-anal setae and with cribrum; anal opening 25–30 long. Dorsal shield is curved on the ventral side posteriorly, does not connect with anal shield, with about 8–10pairs of pilose setae.
Gnathosoma. As in adults.
Legs. Lengths: I 250–287, II 200–225, III 210–240, IV 245–260. Chaetotaxy of legs as in adults.
Protonymph (Figs 52–53, n=5).
Dorsal idiosoma (Fig. 52). Dorsal idiosoma 290–315 long and 190–210 wide with weakly sclerotised podonotal and opisthonotal shields. Anterior margin of the podonotum is curved to the ventral side and forms a vertex with pairs of setae j1; with about 33–35 pilose setae on each side (including marginal setae); ornamented by network of small tubercles. Opisthonotal region with about 26–28 pilose setae on each side; usually entire or divided between J1 and J3 setae by strip of soft cuticle in some specimens. Row— J with 3–4 pairs of setae.
Ventral idiosoma (Fig. 53). Sternal setae St1–St3 and St5 inserted in soft cuticle. Peritrematal shields very narrow, fused with dorsal shield laterally or not distinct in some specimens. Pore-like structures not visible. Peritremes very short (18–22), located between anterior and posterior margins of coxae IV; with internal cell structure; with thin ducts coming from stigmata. One pair of adgenital gland pores gv2 located on soft cuticle posteriorly of coxae IV. Opisthogastric area with 4–5 pairs of smooth setae inserted in soft cuticle. Anal shield as long as wide (37–45), with pair of pilose pre-anal setae and smooth post-anal seta; anal opening 22–23 long, with pair of glands gv3 located antero-laterad of para-anal setae. Dorsal shield is curved on the ventral side posteriorly, does not connect with anal shield; with 4–5 pairs of pilose setae, with most posterior row of smooth marginal setae.
Gnathosoma. Fixed digit of chelicera with five teeth in addition to apical hook; movable digit the same length as fixed one, tridentate in addition to apical hook. Epistome, corniculi, internal malae as in adults and deutonymphs.
Legs. Lengths: I 185–190, II 160–180, III 155–180, IV 185–200. Structures of pretarsus and coxae of leg I–IV as in adults and deutonymphs.
Larva (Figs 54–55, n=4).
Dorsal idiosoma (Fig. 54). Dorsal idiosoma 207–237 long and 158–175 wide with podonotal, pygidial shields and three pairs of mesonotal sclerites. Podonotal shield with nine pairs of setae, including six pairs of j1-j6 setae; setae j1 the longest (23–25) located on anterior margin of shield; stout, pilose, inserted on tubercles. Pygidial shield with four pairs of pilose setae sitting on large tubercles. Two pair of setae in J -row located between mesonotal sclerites; two pairs of lateral setae pilose.
Ventral idiosoma (Fig. 55). Tritosternum as in adults. Sternal setae St1–St3 and St5 inserted in soft cuticle. Periremes and stigmas are absent. Pore-like structures not visible Opisthogastric area with 3–4 pairs of smooth setae inserted in soft cuticle, of which seta Jv2 the longest. Posterior margin of ventral idiosoma with three pairs of short pilose setae. Anal shield is not formed; anal opening surrounded by pair of short pilose pre-anal setae and long smooth post-anal seta.
Gnathosoma. Gnathosoma in forward position not curved in ventral side of idiosoma. Legs. Lengths: I 187–193, II 145–160, III 155–168. Structures of pretarsus and coxae of leg I–III as in adults, deutonymphs and protonymps.