The millipede genus Julus Linnaeus, 1758 in the Caucasus (Diplopoda: Julida: Julidae) Author Evsyukov, Aleksandr Author Golovatch, Sergei Author Reip, Hans S. text Zootaxa 2018 2018-08-20 4461 1 89 117 journal article 28990 10.11646/zootaxa.4461.1.7 1fbcced0-ec5a-4afe-bcb1-496990e850c9 1175-5326 1459872 3FACE488-20DC-46A4-A511-026B7F2DBEA2 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. FIGURE 1. Antennae in Caucasian species of Julus , lateral views. A: J. alexandrae Evsyukov, 2016 . B: J. colchicus Lohmander, 1936 . C: J. jedryczkowskii Golovatch, 1981 . D: J. kubanus Verhoeff, 1921 . E: J. lignaui Verhoeff, 1910 . F: J. lindholmi Lohmander, 1936 . G: J. dagestanus sp. n. H: J. khostensis sp. n. Scale bar: 0.3 mm. 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.? FIGURE 2. Gnathochilaria in Caucasian species of Julus , ventral views. A: J. alexandrae Evsyukov, 2016 . B: J. colchicus Lohmander, 1936 . C: J. jedryczkowskii Golovatch, 1981 . D: J. kubanus Verhoeff, 1921 . E: J. lignaui Verhoeff, 1910 . F: J. lindholmi Lohmander, 1936 . G: J. subalpinus Lohmander, 1936 . H: J. dagestanus sp. n. I: J. khostensis sp. n. Scale bars: 0.3 mm. 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. FIGURE 3. Male legs 1 in Julus alexandrae Evsyukov, 2016 ( A ), J. colchicus Lohmander, 1936 ( B–D ), J. jedryczkowskii Golovatch, 1981 ( E–F ), J. kubanus Verhoeff, 1921 ( G–I ), J. lignaui Verhoeff, 1910 ( J–M ), J. lindholmi Lohmander, 1936 ( N– P ), J. subalpinus Lohmander, 1936 ( Q–R ), J. dagestanus sp. n. ( S ) and J. khostensis sp. n. ( T ). A: from the Rostov-on-Don Region. B: from Krasnaya Polyana, Krasnodar Province. C: from Sakhrai, Adygea Republic. D: syntype from Gagry, Abkhazia, after Lohmander (1936). E: from North Ossetia. F: paratype from North Ossetia, after Golovatch (1981). G: from Maikop, Adygea Republic. H: from Pskhu, Abkhazia. I: from Aksaut, Karachaevo-Cherkessia, after Lohmander (1936). J: from Krasnaya Polyana, Krasnodar Province. K: from Teberda, Karachaevo-Cherkessia. L: from Mestia, Svanetia, Georgia. M: from Teberda, Karachaevo-Cherkessia. N: from Svanetia, Georgia. O: from near Teberda, Karachaevo-Cherkessia. P: holotype from Olginskoe, Abkhazia, after Lohmander (1936). Q: from North Ossetia. R: syntype from Mount Djuga, Krasnodar Province, after Lohmander (1936). S: holotype from Dagestan. T: holotype from Khosta, Krasnodar Province. Oral views B–O ; caudal views A, P . Abbreviations : cx, coxa; te, telopodite remnant. Scale bars: 0.3 mm ( A–C, E, G, H, J–O, Q, S, T ), 0.2 mm ( F ), not to scale ( D, I, P, R ). 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.