Julus colchicus Lohmander, 1936

Figs 1B, 2B, 3B–D, 4B–D, 5, 6C–I, 14B, 15B, 16B, 17B, Map 1

Julus colchicus Lohmander, 1936: 77 (D).

Julus colchicus — Lang, 1959: 1791 (M); Kobakhidze, 1964: 190 (R); 1965: 393 (R); Golovatch, 1977: 49 (D, R); Lokšina and Golovatch, 1979: 386 (M); Talikadze, 1984: 143 (M); Enghoff, 2006: 181 (R); Chumachenko, 2012: 452 (R); 2016: 409 (R); Zuev, 2014: 350 (R); Korobushkin et al., 2016: 25 (R); Kokhia and Golovatch, 2018: 40 (M).

Material examined. 2 males, 30 females, 3 juveniles (ZMUM), Russia, Krasnodar Prov., Severskaya Distr., 3–

10 km S of Ubinskaya, Quercus, Fagus, Carpinus etc. forest, 300–450 m a.s.l., litter and under bark, 3–4.VII.1986; 2 males, 22 females (ZMUM), Mostovskoy Distr., Shedok N of Psebai, karst funnel, wet Fagus, Acer, Quercus etc. forest, litter and under bark, 650 m a.s.l., 4.VIII.1986; 2 males, 1 female, 4 juveniles (ZMUM), Caucasian Nature Reserve, Pslukh ca 20 km E of Krasnaya Polyana, Fagus, Quercus, Abies, Picea etc. forest, 1000 m a.s.l., litter, 12.VIII.1986; 2 males, 5 females, 2 juveniles (ZMUM), same locality, 1 8–20.V.1985; 1 male, 6 females, 2 juveniles (ZMUM), Sochi, ca 8 km S of Khosta, Cave “Our Lady”, Buxus, Fagus, Acer etc. forest near entrance, litter and under stones, 16.V.1985; 1 male, 1 female (ZMUM), same locality, Taxus and Buxus forest, litter, 28.X.1981; 3 males, 5 females, 1 juvenile (ZMUM), same locality, Lazarevskoye, Saloniki, deciduous forest, litter, 28.X.1981, all leg. S. Golovatch; 5 males, 25 females, 4 juveniles (ZMUM), Belorechensky Distr., Dolgogusevsky, Belaya River valley, litter, 7.VI.2009, leg. L. Rybalov and V. Semenov; 2 males, 4 females, 7 juveniles (ZMUM), 5.5 km NE of Krasnaya Polyana, lower flow of Achipse River, 600 m a.s.l., 43°43’10”N, 40°15’25”E, leg. K. Makarov and A. Matalin; 6 males, 6 females, 4 juveniles (AE), near Adler, I.2008; 8 males, 8 females, 1 juvenile (AE), Caucasian Nature Reserve, near Khosta, Taxus and Buxus grove, VI.2006; 17 males, 11 females, 2 juveniles (AE), same locality, V.2006, all leg. Yu. Chumachenko; 2 females, 1 juvenile (SMNG), Khosta, Taxus and Buxus forest, 43.5317°N, 39.8772°E, 80 m a.s.l., 07.VIII.2012, leg. F. Walther; 1 male, 1 female (SMNG), Utrish, Ostrya, Juniperus, Acer campestre, Quercus, Cornus mas, Pistacia etc. coppice, 44.7111°N, 37.4825°E, 11.V.2004; 1 female, 1 juvenile (SMNG), Mezmai, valley with Buxus, under bark, 44.1902°N, 39.9677°E, 16.V.2004, all leg. K. Voigtländer.

2 males (ZMUM), Russia, Adygea Republic, Sakhrai near Novoprokhladnoe, S of Maikop, 700 m a.s.l.,

Fagus and Alnus forest, litter, 7.VII.1986, leg. S. Golovatch; 1 female (SMNG), Sachrai, Novopchladnoe, slope,

mixed forest, 44.1363°N, 40.2880°E, 31.VIII.2005, leg. R. Bischof; 7 juveniles (SMNG), S of Novoprokhladnoe, Bzhebs River valley, 44.1365°N, 40.3179°E, under logs, 19.VIII.2005; 1 juvenile (SMNG), same locality, 30.VIII.2005;2 males, 2 females (SMNG), SE of Novoprokhladnoe, Kuna Valley, wet Fagus orientalis forest, litter, 44.1131°N, 40.3214°E, 30.VIII.2005; 1 female (SMNG), Ust’-Sakhrai, Kuna River valley, Fagus orientalis etc. forest, under logs, 44.2086°N, 40.2816°E, 19.V.2004; 2 female, 4 juveniles (SMNG), SE of Novoprokhladnoe, 44.0866°N, 40.4183°E, 23.V.2004; 1 male (SMNG), same locality, 44.1000°N, 40.3666°E, 26.V.2004; 1 male (SMNG), same locality, Petasites, Heracleum etc. meadow, 44.0999°N, 40.3998°E, 22–26.V.2004; 1 male (SMNG), E of Novoprokhladnoe, Sachrai Valley, Fagus orientalis etc. forest, 44.1365°N, 40.3179°E, 19– 21.V.2004, all leg. K. Voigtländer; 4 males, 6 females (AE), Caucasian Nature Reserve, near Guzeripl, Fagus and Abies forest, 1000 m a.s.l., V.2009; 1 male, 1 female (AE), same locality, 700 m a.s.l., 21.V.2009; 4 males, 11 females (AE), same locality, 1000 m a.s.l., 23.VI.–24.VII.2009; 2 males, 7 females (AE), same locality, 8.VII.– 24.VII.2009; 14 males, 17 females (AE), same locality, VI.2009, all leg. Yu. Chumachenko; 2 males, 5 females (ZMUM), same locality, forest, summer 1976, leg. B. Striganova; 3 males, 2 females (AE), Caucasian Nature Reserve, NE slope of Mt Oshten, 2150 m a.s.l., 17.VII.2004; 5 males, 6 females (AE), near Maikop, Quercus forest, pitfall trapping, 14.IV.–01.V.2009; 7 males, 4 females (AE), near Maikop, 44°20.724’N, 40°11.368’E, 16.III.–1.IV.2011; 3 males, 5 females (AE), same locality, 15.IV.–2.V.2011; 2 males, 5 females (AE), same locality, 15.V.–4.VI.2011; 1 male, 2 females (AE), same locality, 4.VI.–16.VI.2011; 2 males, 8 females, 1 juvenile (AE), same locality, 16.VI.–15.VII.2011; 12 females, 1 juvenile (AE), same locality, 15.VII.–29.VII.2011; 7 females (AE), same locality, 29.VII.–16.VIII.2011; 1 male, 6 females (AE), same locality, 16.IX.–2.X.2011; 3 males, 4 females, 2 juveniles (AE), same locality, 1.X.–17.X.2011, all leg. Yu. Chumachenko; 1 male (AE), 6 km NNE of Guzeripl, mouth of Kisha River, mixed forest, 11.VI.2013, leg. R. Zuev; 1 male, 1 female (AE), Guzeripl, mixed forest, 3.VI.2012, leg.?

3 males, 3 females (ZMUM), Russia, Karachaevo-Cherkessia, Teberda Nature Reserve, Canyon Gonachkhir, Pass Klukhor, 2760–2850 m a.s.l., alpine meadow, under stones, 2.VIII.1986, leg. S. Golovatch ; 1 male (AE), Caucasian Nature Reserve, near Zakan, 30.VI.–9.VII.2009, leg. Yu. Chumachenko .

1 male (ZMUM), Russia, Stavropol Prov., Georgievsk, Quercus, Acer and Fagus forest, litter and under bark, 7.VI.1985, leg. S. Golovatch; 1 male, 3 females, 1 juvenile (AE), Kochubeevsky Distr., near Nevinnomyssk, floodplain of Bolshoy Zelenchuk River, 44.5970°N, 41,9252°E, 340 m a.s.l., 06.XI.2017, leg. R. Zuev.

1 male, 3 females, 1 juvenile (ZMUM), Abkhazia, Lake Ritsa, 950–1100 m a.s.l., Fagus, Abies, Picea, Acer etc. forest, litter, under bark and stones, 13–14.VIII.1986; 19 males, 14 females, 15 juveniles (ZMUM), Myussera Nature Reserve, 20–130 m a.s.l., mixed deciduous forest ( Castanea, Alnus etc.), litter, under bark and stones, 8– 10.IV.1983; 1 male (ZMUM), same locality and dates; 1 male, 9 females (ZMUM), Sukhumi Distr., Bzyb River Valley, Pskhu, 700–900 m a.s.l., Fagus, Quercus, Castanea etc. forest, litter, under bark and stones, 15– 16.VIII.1986; 5 males, 2 females, 1 juvenile (ZMUM), environs of Sukhumi, near Cave Kelassuri, litter, 11.IV.1983, all leg. S. Golovatch; 1 male (ZMUM), near Sukhumi, under stones, 22.IV.1971, leg. N. Zalesskaja; 2 males, 3 females (ZMUM), Abkhazia, Sukhumi Distr., Yashtukha, tree nursery, 29.III.1995, leg. A. Markossian; 1 male (ZMUM), Abkhazia, Sukhum, 42°58.4’N, 41°04.04’E, ca 50 m a.s.l., litter, Eucalyptus, tree shaking etc., 7.XII.2003, leg. Yu. Marusik and G. Antipova.

5 males, 2 females, 1 juvenile (ZMUM), Georgia, Borzhomi Nature Reserve, Banishkevi Valley, 800–900 m a.s.l., Picea, Fagus and Carpinus forest, litter, logs, under stones, 12 and 16.V.1983; 1 male, 8 females, 3 juveniles (ZMUM), Ajaria, Khulo Distr., 3 km W of Danisparauli, deciduous forest, litter, 10.X.1981; 1 male, 3 females (ZMUM), Ajaria, Batumi Botanical Garden, litter, 13.X.1978; all leg S. Golovatch; 2 males, 1 juvenile (ZMUM), Ajaria, Kintrish Nature Reserve, Zeraboseli, 450–600 m a.s.l., 1–3.VI.1981, leg. S. Golovatch and J. Martens; 1 male (SMNG), Khashuri, Akhaldaba, Nedzvi Managemened Reserve, Nedzvistskali River, 41.9082°N, 43.5151°E, humid Picea forest, 01.VI.2011; 1 male (SMNG), Akhaltsikhe to Borjomi, S Chobiskhevi, Chobiskhevi River, 41.7651°N, 43.3265°E, Fagus forest, 01.VI.2011; 1 male (SMNG), Sairme Gorge, 41.9569°N 42.7692°E, mixed forest, leaf litter ( Fagus orientalis and Castanea sativa), 14.VII.2013, all leg. L. Mumladze; 1 male, 1 female (SMNG), Samzche-Dschawachetien, Abastumani Observatory on Mt Kanobili, 41.7544°N, 42.8202°E, 1580 m a.s.l., 28.X.2009; 1 male, 3 females, 1 juvenile (SMNG), Shida Kartli, Ormotsi, Tana valley 16.6 km upstream Didi Ateni, 41.8611°N, 43.9444°E, 1110 m a.s.l., 16.IX.2011; 1 male (SMNG), Samtskhe-Javakheti, 500 m W of Tsaghveri, Gujarula River valley, 41.8078°N, 43.4672°E, 1010 m a.s.l., 17.IX.2011; 1 male (SMNG), Kvemo Kartli, SW of Kojori, 41.6472°N, 44.6936°E, 1200 m a.s.l., 06.X.2011; 1 male (SMNG), Guria, SW of K’vabgha, 41.9131°N, 42.4000°E, 590 m a.s.l., 26.IX.2011; 1 male (SMNG), Ajaria, Q’orolistavi towards Mt Mta Mt’irala, 41.6494°N, 41.7731°E, 730 m a.s.l., 01.X.2012; 1 female, 1 juvenile (SMNG), Mtskheta-Mtianeti, Darial gorge at 8–131 km, 42.7039°N, 44.6264°E, 1480 m.a.s.l., 10.VIII.2014, all leg. F. Walther; 1 male, 1 female (ZMUM), Tskhaltubo, near Cave Belaya (= Tetri), deciduous forest on limestone rock, litter and under stones, 26.X.1981; 1 male (ZMUM), Kutaisi Distr., Sataplia Nature Reserve near Tskhaltubo, forest, litter and under stones, 25.X.1981, all leg. S. Golovatch; 1 male, 1 female (ZMUM), same locality, IV.1973, leg. D. Krivolutsky; 1 male, 1 female (ZMUM), Okriba, Mukhura ca 15 km E of Tkibuli, 700–800 m a.s.l., Castanea, Fagus, Carpinus etc. forest, litter, under bark and stones, 7–9.V.1987; 4 males, 8 females, 4 juveniles (ZMUM), Surami Mt. Ridge, Pass Djvari between Gomi and Sachkhere, 850 m a.s.l., Alnus, Fagus, Rhododendron etc. forest, litter, 7.V.1987; 1 male (ZMUM), Kakhetia, Babaneuri Nature Reserve, ca 16 km ENE of Akhmeta, near Babaneuri, 500 m a.s.l., 4– 5.V.1987, all leg. S. Golovatch and K. Eskov.

Diagnosis. Differs from other species of the genus by the following combination of characters. Main process (mp) of male leg-pair 2 long, with a ventral spike-like outgrowth (Figs 2B, 2C, 3B, 4B, 5B). Ends of flagella (fl) upturned (Figs 3C, 3D, 4D, 5E). Distal part of fl thin, with lamellae. Gonopodal promere (pr) wide, round at apex, with a triangular spike (t) in middle part (Figs 3C, 3D, 4C, 4D, 5C, 5D, 5E). Lateral outgrowth (lo) of op narrow, pointed (Figs 4E, 5E) or rounded (Figs 3D, 5D) at apex. Metazonae with shallow and weakly expressed striations (Figs 2D, 22B). Female vulvae (vu) relatively broad; ampulla 1 (am1) located slightly basal to ampulla 2 (am2); coxa 2 (l2) with a weakly developed apicomesal outgrowth (Fig. 23B).

Descriptive notes. Length of adults 19–36 mm, width 1.3–2.5 mm. Number of segments in adults from 44+1+T to 50+1+T. Body grey-brown with a thin, black, dorsal, axial line, venter lighter; legs light brown to yellow; antennae and anal valves brown (Fig. 5). Eye patches composed of 40–45 ocelli. Metazonae with superficial, weakly expressed striations (Figs 5D, 16B).

All other characters as in J. alexandrae, except as follows. Antennae in situ reaching back to body segment 4. Antennomeres 5 and 6 each with a corolla of bacilliform sensilla (Fig. 1B). Each stipes of gnathochilarium with a group of a few setae parabasally, below mentum (Fig. 2B).

Male leg-pair 1 (Figs 3B–D) with a well-developed suture between coxa (cx) and a 1-segmented telopodite remnant (te). Apex of lateral process of cx varying from acuminate (Figs 3B, 3D) to round (Fig. 3C); te round, with several thin setae. Male leg-pair 2 with three coxal processes (Figs 4B–D): a short anterolateral process (alp), a long, distally thickened main process (mp) with a distocaudal spike-like outgrowth, and a short and acuminate posterior process (pp). Penes (p) short, slightly bifurcate (Fig. 4B).

Promere (pr) broad, clearly rounded at apex, with a triangular mesal spike or tooth (t) near distal third (Figs 8C–I), flagella (fl) long and mostly relatively slender, with inconspicuous lamellae over most of their extent, curved at apex.

Mesomere (ms) rounded at apex (Figs 8D, 8F, 8H, 8I). Opisthomere (op) somewhat variable in shape: flattened (Figs 8D, 8F) or widely rounded (Figs 8H, 8I) (see also Lohmander 1936; Golovatch 1977). Lateral outgrowth (lo) of op narrow, pointed (Fig. 8H), rounded (Fig. 8D) or bifurcate (Fig. 8I) at apex.

Ventral edge of male segment 7 with small curved lamellae bordering the gonopodal aperture (Fig. 15B).

Female vulvae (vu) relatively wide; bursa (bu) with a small, rounded, apical outgrowth; ampulla 1 (am1) located slightly basal to ampulla 2 (am2); coxa 2 (12) with a weakly expressed apicomesal outgrowth (Fig. 17B).

Remarks. This is the most common and widespread species of Julus in the Caucasus, recorded from the Krasnodar and Stavropol provinces, the Adygea and Karachaevo-Cherkessia republics, Abkhazia, most of Georgia and the adjacent parts of Turkey (Lohmander 1936; Lang 1959; Kobakhidze 1964, 1965; Golovatch 1977; Lokšina and Golovatch 1979; Talikadze 1984; Enghoff 2006; Chumachenko 2012; Korobushkin et al. 2016). It inhabits mixed and deciduous forests, as well as subalpine and alpine meadows, occurring at a range of elevations from the Black Sea coast to the alpine belt.

The species shows only minor morphological variations in the structure of male coxa 1 and gonopodal opisthomere.