Brachydesmus (Haplobrachidesmus) kalischewskyi Lignau, 1915

Golovatch, Sergei, Evsyukov, Aleksandr & Reip, Hans, 2016, The millipede family Polydesmidae in the Caucasus (Diplopoda: Polydesmida), Zootaxa 4085 (1), pp. 1-51 : 21-45

Brachydesmus (Haplobrachidesmus) kalischewskyi Lignau, 1915

Figs 11 – 27 , Map 2

Brachydesmus kalischewskyi Lignau, 1915: 370 (D).

Brachydesmus (Haplobrachydesmus) (sic!) kalischewskyi— Lohmander, 1936: 18 (R. D); Attems, 1940: 137 (D); Lang, 1959: 1790 (M); Lokšina and Golovatch, 1979: 384 (M).

Brachydesmus kalischewskyi— Kobakhidze, 1965: 390 (R); Golovatch, 1981a: 110 (R); Talikadze, 1984: 143 (R); Enghoff, 2006: 192 (R); Chumachenko, 2012: 452 (R); Barjadze et al., 2015: 21 (R).

Brachydesmus karawajewi Lohmander, 1928: 530 (D), syn. n.

Brachydesmus (Haplobrachydesmus) (sic!) karawajewi— Lohmander, 1936: 28 (R, D).

Brachydesmus (Haplobrachydesmus) karawajewi— Attems, 1940: 136 (D); Lokšina and Golovatch, 1979: 384 (M).

Brachydesmus karawajewi— Lang, 1959: 1790 (M); Kobakhidze, 1965: 390 (M); Talikadze, 1984: 143 (R); Chumachenko, 2012: 452 (R); Zuev, 2014: 348 (R, D).

Brachydesmus karawaewi (sic!) — Bababekova, 1996: 90 (R).

Brachydesmus (Haplobrachydesmus) (sic!) ferrugineus— Lohmander, 1936: 22 (D), syn. n.

Brachydesmus (Haplobrachydesmus) ferrugineus— Attems, 1940: 137 (D); Lang, 1959: 1790 (M); Lokšina and Golovatch, 1979: 384 (M).

Brachydesmus ferrugineus— Kobakhidze, 1965: 390 (R); Ceuca, 1975: 91 (D); Golovatch, 1981a: 110 (R); 1981b: 426 (D); Talikadze, 1984: 143 (R); Loginova, 1993: 151 (R); 1995: 103 (R); Enghoff and Moravvej, 2005: 69 (R); Enghoff, 2006: 192 (R); Kokhia, 2011: 28, 50, 80 (R); Barjadze et al., 2015: 21 (R).

Brachydesmus (Haplobrachydesmus) (sic!) talyschanus Lohmander, 1936: 26 (D), syn. n.

Brachydesmus (Haplobrachydesmus) talyschanus— Attems, 1940: 137 (D); Lang, 1959: 1790 (M); Lokšina and Golovatch, 1979: 384 (M).

Brachydesmus talyschanus— Kobakhidze, 1965: 391 (R); Rakhmanov, 1972: 116 (R); Samedov et al., 1972: 1245 (R); Rasulova and Rakhmanov, 1973: 520 (R); Talikadze, 1984: 143 (R); Bababekova, 1996: 90 (R).

Brachydesmus (Haplobrachydesmus) bidentatus Golovatch, 1976: 1568 (D), syn. n.

Brachydesmus bidentatus— Lokšina and Golovatch, 1979: 384 (M); Bababekova, 1996: 90 (R).

Material examined. 3 males, 2 females (ZMUM), RUSSIA, Krasnodar Prov., Tuapse Distr., 15 km SE of Novomikhailovskii, Psebe, deciduous forest, litter, under bark and stones, 29.X.1981; 1 female (ZMUM), Krasnodar Prov., Sochi, Dagomys, Quercus forest, litter, 28.X.1981; 1 male, 1 female (ZMUM), Krasnodar Prov., Sochi, Lazarevskoye, Saloniki, deciduous forest, litter, 28.X.1981; 3 males (ZMUM), Krasnodar Prov., Sochi, Khosta, Taxus and Buxus forest, litter, 28.X.1981, all leg. S. Golovatch; 1 male, 2 females (ZMUM), same locality, Taxus and Buxus forest, III.2006, leg. Yu. Chumachenko; 2 females, 1 subadult female (ZMUM), Krasnodar Prov., Sochi, 5.5 km NE of Krasnaya Polyana, lower course of Achipse River, ca 600 m a.s.l., N43°43’10”, E40°15’25”, 19 – 23.VIII.2014, leg. K. Makarov and A. Matalin; 2 males, 5 females (ZMUM), Krasnodar Prov., Aibga Mt. Range, sources of Tikhaya Rechka River, 1700 – 1900 m a.s.l., N43.638º, E40.2481º, 17.06.1997, leg. I. Belousov; 1 male (ZMUM), Krasnodar Prov., Tuapse Distr., Georgievskoe Forestry, Castanea forest, 6.V.1954, leg. K. Arnoldi, det. H. Lohmander, det. C. Dziadosz, 1968; 3 juv. (ZMUM), Krasnodar Prov., Caucasian Nature Reserve, Pslukh, ca 20 km E of Krasnaya Polyana, 1000 m a.s.l., Fagus , Quercus , Abies , Picea etc. forest, litter, 11 – 12.VIII.1986; 5 males, 3 females, 3 juv. (ZMUM), Krasnodar Prov., Goryachiy Klyuch, 12 km SW of Fanagoriyskoye, Cave Fanagoriyskaya, 19.V.1983; 7 juv. (ZMUM), Krasnodar Prov., Mostovskoy Distr., Yaroslavskaya, 300 m a.s.l., Quercus , Acer, Corylus etc. scrub, 4.VIII.1986, all leg. S. Golovatch; 1 female (ZMUM), Krasnodar Prov., Tuapse Distr., Georgievskoye Forestry, Kazachye, forest, 7.V.1954, leg. M. Ghilarov; 1 female (ZMUM), Adygea, Maikop, garden, V.2011; 1 male, 1 male fragment (without segments 14 – 19), 1 male fragment (without head and segments 1 – 10) (ZMUM), Adygea, Caucasian Nature Reserve, Pasture Abago S of Guzeripl, 1800 m a.s.l., subalpine meadow, pitfall traps, 3 – 29.VIII.2009, all leg. Yu. Chumachenko; 1 male, 2 females, 1 juv. (SMNG 34137), Adygea, Koryto, 22.VIII.2005; 1 male (SMNG 34144), Adygea, Polyana Shostakova, 25.VIII.2005; 3 males (SMNG 34158), Adygea, Sakhrai, Kuna Valley, 30.VIII.2005, all leg. K. Voigtländer. 2 females (AE), Krasnodar Prov., Khadyzhensk, 15.VI.2007, leg. A. Evsyukov and D. Khisametdinova; 4 females (ZMUM), same locality, 1350 – 1400 m a.s.l., Abies and Fagus forest, litter, under bark and stones, 24 – 26.V.1985; 5 females (ZMUM), same locality, 1700 – 1850 m a.s.l., Abies and Fagus forest, Rhododendron thicket, litter, 24 – 26.V.1985; 2 females (ZMUM), Krasnodar Prov., Caucasian Nature Reserve, Mt Aishkho-2 E of Pslukh, ca 28 km E of Krasnaya Polyana, S slope, 1600 m a.s.l., Abies , Picea , Fagus , Betula etc. forest, litter, 12.VIII.1986; 1 male, 1 female (ZMUM), Karachaevo-Cherkessia, ca 30 km S of Kurdjinovo, Laba Valley, 4 km N of Damkhurts, 1050 – 1100 m a.s.l., Fagus , Acer , Picea etc. forest, litter and under bark, 4.VIII.1986; 1 juv. (ZMUM), Karachaevo-Cherkessia, Teberda Nature Reserve, above Teberda, Mt Malaya Khatipara, 2400 m a.s.l., subalpine meadow, under stones, 29 – 30.V.1985; 1 female (ZMUM), same locality, Kizgich Canyon, N of Arkhyz, 1550 – 1600 m a.s.l., Abies , Picea , Pinus , Fagus , Betula etc. forest, litter, under bark and stones, 5.VI.1985; 27 males, 1 female, 3 juv. (ZMUM), Karachaevo-Cherkessia, Lower Teberda S of Karachaevsk, 1000 m a.s.l., Quercus , Fagus , Acer etc. forest, litter, 3.VIII.1986, all leg. S. Golovatch; 2 females (ZMUM), Karachaevo-Cherkessia, Teberda, ca 1200 – 2000 m a.s.l., VI – VII.1908, leg. A.P. Zolotarev; 31 juv. (ZMUM), Kabardino-Balkaria, Lower Chegem, 800 m a.s.l., Fagus forest, litter, 14.VII.1986; 18 males, 2 females (ZMUM), North Ossetia, Novo-Georgievskoye near Mozdok, Quercus , Acer , Alnus , Fraxinus etc. forest, floodplain of Terek River, litter, logs, under stones, 27.V.1982; 1 male, 7 females, 6 juv. (ZMUM), North Ossetia, S of Ordzhonikidze (= Vladikavkaz), between Chmi and Baltik, Quercus and Alnus scrub on slope, litter and under stones 2.VI.1982; 12 males, 5 females (ZMUM), North Ossetia, S of Alagir, 200 m a.s.l., Acer , Fagus etc. forest, litter, 17.X.1987; 3 males, 4 females, 9 juv. (ZMUM), Ingushetia, Assa Valley, ca 9 km SSW of Muzhichi, 800 m a.s.l., Fagus , Alnus , Carpinus etc. forest, litter, under bark and stones, 15.VII.1986; 2 females, 2 juv. (ZMUM), Chechnya, 5 km W of Shaami-Yurt, ca 26 km W of Groznyi, Acer forest, litter, 6.VI.1982; 1 female (ZMUM), Dagestan, Chiragchay Valley, Sardarkent between Kasumkent and Khiv, 650 m a.s.l., Crataegus, Rosa , Carpinus , Juglans etc. scrub, litter, 24.X.1987; 1 male (ZMUM), Dagestan, Samur Valley, Garakh ca 35 km SW of Magaramkent, 700 – 800 m a.s.l., Crataegus , Quercus , Acer, Rosa etc. scrub, litter, 23.X.1987, all leg. S. Golovatch; 10 males, 8 females (ZMUM), Dagestan, Kaytag Distr., Karatsan, 6.X.2012, leg. P. Gamilova.

1 female (ZISP), SOUTH OSSETIA, near Kvaisi, under stones, 1700 m a.s.l., 19.X.1956, leg. K. Gorodkov; 1 male, 1 female (ZMUM), 15 km E of Kvaisi, Surami Mt. Ridge, Pass Ertso, ca 1000 m a.s.l., deciduous brush, along stream, litter and stones, 20.X.1981; 4 males, 2 females (ZMUM), Lower Roki S of Pass Rokskii, 1500 m a.s.l., Fagus , Betula etc. forest, litter, 17.X.1987, all leg. S. Golovatch.

2 males, syntypes of B. ferrugineus ( ZISP), ABKHAZIA, “W Transcaucasia, Olginskoe, Tsebelda, 20.V.2012, leg. V. Lindholm”; 1 male, syntype of B. ferrugineus (gonopods missing) ( ZISP), “Black Sea Gouvernement, Novyi Afon, 12.V.1912, leg. V. Lindholm”; 1 female ( ZISP), “ 1 female juv., Polydesmus sp., det. C. Dziadosz, 1967”, Gagra, Mamsdyshkha, ca 2000 m, VIII.1911, leg. V. Isaev?; 1 female ( ZMUM), Lake Ritsa, 1 800 m a.s.l., Abies and Fagus forest, litter, 13.IX.1985, leg. I. Ushakov; 2 males, 1 female, 1 juv. ( ZMUM), Lake Ritsa, 950 – 1100 m a.s.l., Fagus , Abies , Picea , Acer etc. forest, litter, under bark and stones, 13 – 14.VIII.1986, leg. S. Golovatch; 1 male, 1 juv. ( SMNG 14685), Transcaucasus, Riza-See (= Lake Ritsa), 20.VI.1984, leg. Dunger; 1 female ( ZMUM), Sukhumi Distr., near Cave Kelassuri, litter, 11.IV.1983; 1 male ( ZMUM), same locality, near Verkhnyaya Kelassuri, deciduous bush on rock, slope, litter, 27.X.1981, all leg. S. Golovatch; 6 males ( ZMUM), Shroma near Sukhumi, 27.IV.1971, leg. N. Zalesskaya; 1 male ( ZMUM), Sukhumi, I.1986, leg. V. Dolin; 1 juv. ( ZMUM), Sukhumi Distr., Lake Amtkel, ca 16 km N of Tsebelda, 550 m a.s.l., Alnus forest, litter and under bark, 19.VIII.1986; 1 juv. ( ZMUM), Sukhumi Distr., Tsebelda, 300 m a.s.l., Carpinus , Acer and Buxus scrub, litter, 19.VIII.1986; 2 males, 1 juv. ( ZMUM), Sukhumi Botanical Garden, 9.X.1978, leg. S. Golovatch; 1 female, 1 juv. ( ZMUM), 2 km N of Duripsh N of Gudauta, Fagus log, under bark, 8.IX.1985, leg. I. Ushakov; 1 female ( ZMUM), Shroma near Sukhumi, 2.V.1971, leg. N. Zalesskaya; 1 male ( ZMUM), Sukhumi, Canyon Alekseevskoe, in house, 3.XII.1953; 1 female ( ZMUM), same locality, 8.X.1953, all leg. E. Borutzky; 1 male, 1 female ( ZMUM), Sukhumi, University Campus, N42°58,4’ E41°04,04’, ca 50 m a.s.l., Encopytus, tree shaking, litter, 7.XII.2003, leg. Yu. Marusik and G. Antonova; 1 male ( ZMUM), Myussera Nature Reserve, 1.II.1979, leg. L. Malinina.

1 male, syntype of B. ferrugineus (ZISP), GEORGIA, “Tiflis Gouvernement, Canyon Banis-khevi-tskhali [near Borzhomi], 13.VI.1909, leg. L. Berg”; 1 male (ZMUM), Lanchkhuti Distr., upper reaches of Supsa River, 15.X.1983; 2 juv. (ZMUM), Chokhatauri Distr., Bakhmaro ca 40 km SSW of Nabeglavi, 2000 m a.s.l., IV.1984; 3 females (ZMUM), Ajaria, Kintrish Nature Reserve, Zeraboseli, Khekhpara Valley, 12.X.1984, all leg. E. Kvavadze; 21 males, 13 females (ZMUM), same locality, 450–600 m a.s.l., deciduous forest, litter and under stones, 13.X.1981; 3 females (ZMUM), same locality, 800 m a.s.l., Rhododendron thicket, litter, 13.X.1981, all leg. S. Golovatch; 5 males, 6 females (ZMUM), same locality, 450 – 600 m a.s.l., deciduous forest, 1 – 3.VI.1981; 2 males, 1 female (ZMUM), same locality, 600 – 700 m a.s.l., Rhododendron thicket, 2.VI.1981, all leg. S. Golovatch and J. Martens; 17 juv. (ZMUM), Ajaria, between Kobuleti and Ureki, seashore Pinus forest, litter, 5 m a.s.l., N41º55’58.1", E41º45’53.0", 21.07.2012, leg. Y.M. Marusik; 5 males, 1 female, 1 juv. (ZMUM), Ajaria, Keda Distr., Magutseti, Platanus forest, litter, 9.X.1981; 9 males, 2 females, 2 juv. (ZMUM), Ajaria, Khulo, 900 m a.s.l., Quercus , Abies , Alnus etc. forest, litter, 11.X.1981; 4 juv. (ZMUM), Ajaria, E of Khulo, 950 m a.s.l., deciduous forest on rock, near spring, litter, 10.X.1981; 1 juv. (ZMUM), Ajaria, 6 km W of Khulo, 800 m a.s.l., deciduous forest on rocky slope, litter, 11.X.1981, all leg. S. Golovatch; 1 male, 1 female (ZMUM), Ajaria, Tsikhisdziri N of Batumi, 100 m a.s.l., gardens, 30.V.1981; 7 males, 20 females (ZMUM), Ajaria, Batumi Botanical Garden, 20 – 150 m a.s.l., 30.V. – 7.VI.1981, all leg. S. Golovatch and J. Martens; 3 males, 1 juv. (ZMUM), Ajaria, Meskheti Mt. Range, Pass Goderdzi, 1800 m a.s.l., Picea forest, under stones, litter, umder bark, 2.IX.1999, leg. L. Penev; 1 juv. (ZMUM), Ajaria, E of Pass Goderdzi, ca 1620 m a.s.l., Abies and Pinus forest, N41º38’42.6", E42º35’45.5", 22.07.2012, leg. Y. Marusik; 6 males, 12 females (ZMUM), 15 km W of Adigeni, 1500 – 1700 m a.s.l, Abies , Picea , Fagus , Acer etc. forest, litter, logs, under stones, 14 – 15.V.1983, leg. S. Golovatch; 4 females (ZMUM), W of Adigeni, 1500 – 1700 m a.s.l., Abies , Picea , Fagus etc. forest, 21 – 22.VI.1978, leg. V. Dolin; 13 females (ZMUM), Borzhomi Nature Reserve, Baniskhevi Valley, 800–900 m a.s.l., Picea , Fagus , Carpinus etc. forest, litter, under bark and stones, 12 and 16.V.1983; 1 male (ZMUM), S of Bakuriani, Pinus and Fagus forest, 1750 m a.s.l., litter and logs, 13.V.1983, all leg. S. Golovatch; 1 female (ZISP), Bakuriani, Bakuriani River valley, 11.V.1986, leg. L. Zhiltsova; 6 males, 5 females (ZMUM), Mukhura, 14 km E of Tkibuli, Quercus , Fagus , Castanea etc. forest, litter and under stones along spring, 23.X.1981, leg. S. Golovatch; 1 female (ZMUM), 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, leg. S. Golovatch and K. Eskov; 8 juv. (ZMUM), ca 10 km E of Tkibuli, Mukhura, 1010 m a.s.l., Carpinus and Fagus forest with Buxus thickets, N42°20’05,8”, E43°02’38,5”, 23.VII.2012, leg. Yu. Marusik; 1 male, 1 female, 3 juv. (ZMUM), Svanetia, ca 40 km W of Mestia, Kherkhvashi E of Nakra (= Naki), 1200 – 1700 m a.s.l., Quercus , Fagus , Carpinus , Picea , Abies etc. forest, litter and under bark, 21.VIII – 21.IX.1986; 4 males, 1 female (ZMUM), Svanetia, Mestia, 1500 m a.s.l., Betula and Rhododendron on moraine, litter and under stones, 5 and 16.IX.1986; 1 male (ZMUM), same locality, under stones, 22.X.1979; 7 males, 3 females (ZMUM), Svanetia, ca 40 km W of Mestia, above Kherkhvashi, 1900 – 2200 m a.s.l., timberline ( Rhododendron , Picea , Abies ) and subalpine meadows, 21.IX.1986; 5 males, 5 females (ZMUM), Ambrolauri Distr., Cave Nikortsminda near Nikortsminda, 1100 m a.s.l., 40 – 50 m deep, on walls and rotten wood, 22.X.1981; 1 male (ZMUM), same locality, 1150 m a.s.l., on logs, 19.X.1987; 1 juv. (ZMUM), Tskhaltubo, near Cave Belaya, deciduous forest on rock, litter and under stones, 26.X.1981; 1 juv. (ZMUM), Kutaissi Distr., Sataplia Nature Reserve, mixed deciduous forest, 8.VIII.1974; 1 juv. (ZMUM), Orpiri NE of Kutaissi, rotten wood, 11.VIII.1974, all leg. S. Golovatch; 1 female (ZMUM), Sataplia Nature Reserve, 400 m a.s.l., Fagus and Buxus forest, litter, 5.VI.1981, leg. S. Golovatch and J. Martens; 2 females (ZMUM), Sataplia Nature Reserve, IV.1973, leg. D. Krivolutsky; 8 males, 7 females, 1 juv. (ZMUM), Oni Distr., ca 10 km NE of Shovi, Gurshevi near Pass Mamisoni, 2000 – 2200 m a.s.l., Abies , Fagus , Alnus etc. forest, litter and under stones, 21.X.1981; 3 males, 3 females (ZMUM), Oni Distr., SW of Pass Mamisoni, 2500 m a.s.l., Salix bush along spring in subalpine meadow, litter and under stones, 21.X.1981; 1 male, 4 females (ZMUM), Oni Distr., Shovi, Fagus , Alnus , Abies etc. forest, litter and under stones near spring, 21.X.1981, all leg. S. Golovatch; 2 males, 2 females (ZMUM), Algeti Nature Reserve, W of Manglisi, 1400 – 1450 m a.s.l., Fagus , Picea , Acer etc. forest, litter and under bark, 16 – 18.V.1987, leg. S. Golovatch and K. Eskov; 1 female (ZMUM), Algeti State Reserve, Manglisi, Quercus forest, 12.VI.1984, leg. E. Kvavadze; 4 males, 4 females (ZMUM), Saguramo Nature Reserve, NE of Mtskheta, Zedazeni, 1100 – 1200 m a.s.l., Fagus , Carpinus , Acer etc. forest, litter and under bark, 20.V.1987; 1 female (ZMUM), Lake Shaori, mixed forest, litter, N42º23’46.6", E43º01’47.7", ca 1180 m a.s.l., 19.07.2012, leg. Y. Marusik; 2 males, 4 females (ZMUM), Batsaro Nature Reserve, ca 20 km N of Akhmeta, 800 – 850 m a.s.l., Fagus , Castanea , Acer etc. forest, litter, 5 – 6.V.1987; 4 males, 2 females (ZMUM), Pass Magalakhari between Akhmeta and Tianeti, 1200 m a.s.l., Fagus and Carpinus forest, litter and under bark, 6.V.1987; 9 juv. (ZMUM), Lagodekhi Nature Reserve, trail to meteorological station, ca 1835 m a.s.l., Carpinus forest, N41º51’38.0", E46º20’26.5", 27.07.2012, leg. Yu. Marusik; 4 males, 4 females (ZMUM), Kartli Mt. Ridge, Pass Sabaduri ca 40 km NNE of Tbilisi, 1400 m a.s.l., Fagus forest, litter and under bark, 6.V.1987; 2 males, 1 female (ZMUM), Surami Mt. Ridge, SE of Pass Djvari, between Gomi and Sachkhere, 850 m a.s.l., Alnus , Fagus , Rhododendron etc. forest, litter, 7.V.1987; 1 male, 2 females (ZMUM), N of Kvareli, 700 – 750 m a.s.l., Fagus , Carpinus , Quercus etc. forest, litter and under bark, 4.V.1987, all leg. S. Golovatch and K. Eskov; 1 male, 1 juv. (ZMUM), Babaneuri Nature Reserve ca 16 km NE of Akhmeta, near Babaneuri, 500 m a.s.l., Fagus , Quercus , Carpinus etc. forest, litter, 4 – 5.V.1987; 1 female, 4 juv. (ZMUM), 10 km N of Djvari, 800 m a.s.l., Buxus , Fagus , Picea , Taxus etc. forest, litter, 20 – 21.VIII.1986; 1 male, 3 females, 1 juv. (ZMUM), Kazbegi, 2000 m a.s.l., Pinus and Betula forest and subalpine meadows, litter and under stones, 2 – 6.VI.1982; 1 male, 2 females (ZMUM), Kazbegi Distr., S of Gudauri near Pass Krestovyi, 1800 m a.s.l., Rhododendron bush, litter, 4.VI.1982, all leg. S. Golovatch; 1 male (SMNG), Kazbegi, N42.69º, E44.52º, 28.VII.1985, leg. B. Seifert; 1 male (SMNG), Polyana Firsova, 1400 m a.s.l., VIII./ IX.1 999, leg. O. Tietz; 1 male (HR), Tusheti, slope between Omalo and Khenako, 1810 m a.s.l., N42.3719º, E45.6444º, 4.IX.2009; 1 male (HR), Tusheti, pass between Omalo and Diklo, 2270 m a.s.l., N42.4044º, E45.6239º, 5.IX.2009; 3 males, 1 female (HR), Khevsureti, Khone Skali River valley, between Khone and Khone Chala, 1870 m a.s.l., N42.5675º, E45.2203º, 10.IX.2009; 1 female (HR), Khevsureti, Churo Valley, mouth of Khone Skali River, 1980 m a.s.l., N42.5683º, E45.2322º, 10.IX.2009; Khevsureti, Khone Skali River valley, between Khone and Khone Chala, 1870 m a.s.l., N42.5675º, E45.2203º, 10.IX.2009; 3 males, 1 female (HR), Khevsureti, ca 200 m downstream of Shatili, 1415 m a.s.l., N42.6594º, E45.1653º, 12.IX.2009; 4 males, 1 female (HR), Khevsureti, Argun River valley, between Shatili and Georgi Tsvinda, 1580 m a.s.l., N42.6397º, E45.1425º, 13.IX.2009; 2 males (HR), Shida Kartli, Ormotsi, Tana River valley, 16.6 road-km upstream of Didi Ateni, 1110 m a.s.l., N41.8611º, E43.9444º, 16.IX.2011; 1 male (HR), Samtskhe–Djavakheti, Borzhomi to Chitakhevi, junction to Mtsvane Monastery, 850 m a.s.l., N41.8039º, E43.3181º, 17.IX.2011; 1 male (HR), Samtskhe-Djavakheti, Bakuriani, 7.5 road-km to Tskhratskaro Ugheltekhili Pass, 2080 m a.s.l., N41.7067º, E43.5058º, 17.IX.2011; 1 female, 1 juv. (HR), Samtskhe-Djavakheti, Vardzia, 2 road-km from bridge to Tmogvi, N41.3823º, E43.2982º, 25.VIII.2014; 1 male (HR), Racha-Lechkhumi and Kvemo Svaneti, Khotevi, 1 road-km to Nikortsminda, 930 m a.s.l., N42.4636º, E43.1236º, 22.IX.2011; 1 male (HR), Racha-Lechkhumi and Kvemo Svaneti, Zubi, road between Tsageri and Tkhaltubo, junction to Zubi, 430 m a.s.l., N42.5761º, E42.6808º, 23.IX.2011; 1 male (HR), Ajaria, Machakhlispiri, Chorokhi River valley, 2.7 road-km upstream of Acharistskhali, 80 m a.s.l., N41.5278, E41.7211, 29.IX.2011; 3 males (HR), Ajaria, NE of Dologani, 700 m, 100 m a.s.l., N41.5525º, E41.8019º, 29.IX.2011; 1 male (HR), Mtskheta–Mtianeti, Saguramo, 1.2 road-km towards Zedazeni Monastery, 770 m a.s.l., N41.8889º, E44.7731º, 3.X.2011; 2 males, 1 female, 1 juv. (HR), Mtskheta–Mtianeti, Tskhinvali, 6 road-km to Tianeti, 1120 m a.s.l., N42.1392º, E44.8167º, 5.X.2011; 1 male (HR), Samegrelo-Zemo Svaneti, Nodashi, 1100 m a.s.l., N43.0586º, E42.4164º, 29.IX.2012; 1 male (HR), 3 males, 1 female (HR), Mtskheta–Mtianeti, Stepantsminda, Darial Gorge, 4.5 road-km to border (near Gveleti), 1470 m a.s.l., N42.7058º, E44.6267º, 6.X.2012; 3 females (HR), Mtskheta–Mtianeti, Andaki River valley 2.2 km upstream of its confluence with Argun River, N42.5006º, E45.2156º, 8.VIII.2014; 1 male (HR), Mtskheta–Mtianeti, Darial Gorge at road-km 8/131, N42.7039º, E44.6264º, 10.VIII.2014; 1 male, 1 female (HR), Mtskheta–Mtianeti, 1.1 km SW of Kobi, N42.5553, E44.4973º, 12.VIII.2014; 1 male (HR), Imereti, Nakeralos Ugheltekhili Pass, 730 m SSE, N42.3708, E43.0428º, 14.VIII.2014; 1 male (HR), Samegrelo and Zemo Svaneti, Ushguli, entrance to Enguri (= Inguri) River canyon, 2 km W of Ushguli, N42.9147º, E42.9747º, 19.VIII.2014, all leg. F. Walther.

3 males, 1 female (ZMUM), ARMENIA, Azizbekov Distr., Zangegur Mt. Ridge, Pass Voratan, subalpine meadow, 2100 – 2200 m a.s.l., under stones, 21.IV.1983; 1 male (ZMUM), near Azizbekov, 1200 m a.s.l., Artemisia grassland, under stones, 21.IV.1983; 2 males, 6 females (ZMUM), Ararat Distr., Khosrov Nature Reserve, 1450 – 1550 m a.s.l., Juniperus , Quercus , Crataegus etc. bush along river, litter and under stones, 19 – 20.IV.1983, all leg. S. Golovatch; 4 males, 2 females (ZMUM), Ararat Distr., Khosrov Nature Reserve, 2000 – 2200 m a.s.l., subalpine meadow, under stones, 11.V.1984; 5 males, 2 females, 1 juv. (ZMUM), Megri Distr., Nyuvadi, xerophytous canyon, under stones, 25.IV.1983, all leg. V. Januschev; 7 males, 4 females, 4 juv. (ZMUM), Megri Distr., 6 km N of Shvanidzor, sparse Quercus forest, 1200 – 1300 m a.s.l., litter, under stones and bark, 24.IV.1983; 8 males, 13 females, 3 juv. (ZMUM), environs of Megri, xerophytous bare canyon, 1000 m a.s.l., sparse J uniperus and Paliurus bush, under stones, 24.IV.1983; 13 males, 18 females (ZMUM), Megri Distr., SSE of Lichk, Megri River valley, 1530 m a.s.l., Quercus forest, litter, under stones and in rotten wood, 25.IV.1983; 17 males, 15 females (ZMUM), Megri Distr., above Kuris, 1500 m a.s.l., Quercus , Acer etc. forest, litter, under bark and stones, 26.IV.1983; 7 males, 4 females, 4 juv. (ZMUM), Megri Distr., 6 km N of Shvanidzor, sparse Quercus forest, 1200 – 1300 m a.s.l., litter, under stones and bark, 24.IV.1983; 23 males, 10 females (ZMUM), Kafan Distr., near Kadjaran, Megri Mt. Ridge, N of Pass Tashtun, 2000 m a.s.l., Quercus forest on steep slope, litter and logs, 27.IV.1983; 38 males, 29 females (ZMUM), Kafan Distr., Shikahoh Nature Reserve, Shikahoh, 900 – 950 m a.s.l., Quercus , Fagus , Carpinus etc. forest along stream, litter, under logs and stones, 28.IV.1983; 7 males, 7 females (ZMUM), Kafan Distr., Shikahoh Nature Reserve, Shishkert, 1700 – 1800 m a.s.l., Quercus , Fagus , Carpinus forest, litter and under stones, 29.IV.1983; 2 males, 6 females (ZMUM), same locality and data; 8 males, 1 female, 1 juv. (ZMUM), Kafan Distr., Shikahoh Nature Reserve, Tsav, 1000 m a.s.l., Quercus , Fagus , Carpinus etc. forest along stream, litter, under logs and stones, 29.IV.1983; 1 female (ZMUM), Kafan, log, under bark, 27.IV1983; 1 male (ZMUM), Kafan Distr., Shikahoh Nature Reserve, Nerkin And, old Platanus stand along river, litter, rotten wood, under stones, 30.IV.1983; 3 males, 1 female, 1 juv. (ZMUM), Legvaz, 4 km NWN of Megri, Juniperus , Quercus etc. scrub with Paliurus and Rosa, litter and under stones, 1000 m a.s.l., 24 – 25.IV.1983; 3 females (ZMUM), Stepanavan, foot of Mt Archasar, 1400 m a.s.l., Quercus forest near spring, 14.XI.1985, all leg. S. Golovatch; 1 male, 1 female (NHMG), Armenia, Jelal-Ogly (= now Stepanavan), 5.VIII.1920, leg. A. Shelkovnikov, det. H. Lohmander, vid. S. Golovatch; 4 females (ZMUM), Kirovakan (= Vanadzor), 1600 m a.s.l., Quercus , Acer , Fagus etc. forest, litter, 22.V.1987, leg. S. Golovatch and K. Eskov; 2 males (ZMUM, ex Coll. HNHM), Karashamb, Agveran, 1500 m a.s.l., shrubby hillside, under stones, 16.IX.1982, leg. O. Merkl and L. Ronkay; 28 males, 13 females, 3 juv. (ZMUM), SW of Shnokh, halfway between Alaverdi and Bagratashen, 750 m a.s.l., Carpinus forest, litter, 24.V.1987, leg. S. Golovatch and K. Eskov; 2 females (ZMUM), Dilizhan Nature Reserve, Akhnabat Valley near Salakh, 1400 – 1500 m a.s.l., Fagus , Taxus , Carpinus etc. forest, litter, under bark and stones, 17.IV.1983, leg. S. Golovatch; 1 female (ZMUM), Dilizhan Nature Reserve, Agartsyn, 1350 – 1400 m a.s.l., Fagus , Acer etc. forest, litter, 28 – 29.V.1987; 1 male, 7 females (ZMUM), W of Shamshadyn, halfway between Berd and Idjevan, 1500 – 1600 m a.s.l., Fagus , Carpinus , Acer etc. forest, litter and under bark, 26 – 27.V.1987; 6 males, 4 females (ZMUM), Berdavan ca 10 km S of Noemberian, 900 – 950 m a.s.l., Quercus , Carpinus , Acer etc. forest, litter and under stones, 24 – 25.V.1987, all leg. S. Golovatch and K. Eskov.

Male holotype of B. talyschanus (ZISP), AZERBAIJAN, “Baku Gouvernement, Lenkoran Distr., Lerik, 12.V.1912, leg. A. Kirichenko”; 13 males, 18 females (ZMUM), Mountainous Karabakh, Askeran Distr., ca 6 km WNW of Dashbulag, near Badara, Quercus , Carpinus etc. forest, 850–900 m a.s.l., litter, under bark and stones, 2.V.1983; 9 males, 5 females (ZMUM), Mountainous Karabakh, Dashalty near Shusha, 1100 – 1300 m a.s.l., Quercus , Carpinus etc. forest, litter, logs, under stones, 1.V.1983, all leg. S. Golovatch; 1 juv. (ZMUM), Mountainous Karabakh, environs of Firуuza, Domy, near Cave Glenkar, in soil, 16 – 17.VIII.1977, leg. N. Zalesskaya; 2 juv. (ZMUM), Mountainous Karabakh, Drmbon ca 30 km WSW of Mardakert, 800 – 850 m a.s.l., Quercus , Acer etc. forest, litter, 1 – 2.VI.1987; 4 males, 5 females (ZMUM), Mountainous Karabakh, Chilisa ca 7 km N of Kelbadjar, 1450 – 1500 m a.s.l., Quercus , Carpinus , Acer etc. forest, litter and under bark, 31.V.1987; 10 males, 5 females, 5 juv. (ZMUM), Mountainous Karabakh, Nadirkhanly ca 12 km NE of Kelbadjar, 1200 m a.s.l., Fraxinus and Juglans regia stand on rock, litter and under stones, 1.VI.1987; 3 females (ZMUM), Mountainous Karabakh, Lesser Istisu halfway between Kelbadjar and Istisu, 1550 m a.s.l., Quercus and Acer scrub on slope, litter, 31.V.1987, all leg. S. Golovatch and K. Eskov; 5 males, 2 females (ZMUM), Istisu ca 8 km SW of Masally, 80 – 140 m a.s.l., Quercus , Carpinus , Acer etc. forest, litter, under bark and stones, 19 – 20.X.1983, leg. S. Golovatch; 1 female (ZMUM), Djar near Zakataly, forest, litter, 18.IV.1986, leg. N. Loginova; 1 juv. (ZMUM), Perzivan SE of Zakataly, 600 m a.s.l., Quercus , Fagus etc. forest along stream, litter, 3.V.1987, leg. S. Golovatch and K. Eskov; 3 females, 8 juv. (ZMUM), Zakataly Nature Reserve, Katekhchay Valley, 700 m a.s.l., 23.V.1981, leg. S. Golovatch and J. Martens; 6 females, 4 juv. (ZMUM), Zakataly Nature Reserve, 1000 m a.s.l., forest, litter, 13.VIII.1989; 1 female, 2 juv. (ZMUM), Zakataly Nature Reserve, Agkemal, 2200 m a.s.l., forest, litter, 15.VIII.1989, all leg. S. Dashdamirov and H. Aliev; 1 female (ZMUM), Zakataly Distr., Djar, along Tarykchay River, steep slope at 45°, Fagus forest, litter, 8.VI.1986, leg. K. Mikhailov; 1 male, 1 female (ZMUM), Zakataly Distr., Djar, 18.IV.1986, leg. N. Loginova; 1 female (ZMUM), Zakataly Nature Reserve, X.1970, leg. L. Malinina; 8 males, 3 females, 7 juv. (IZB), Kedabek (= Gədəbek), montane forest, VIII.1947, leg. A. Bogachev; 12 males, 8 females (ZMUM), Gey-Giol Nature Reserve, ca 50 km S of Kirovabad (= Ganja), 1500 – 1900 m a.s.l., Fagus , Carpinus , Quercus etc. forest, litter, logs and under stones, 3 – 4.V.1983, leg. S. Golovatch; 2 males, 1 female, 2 juv. (ZMUM), ca 15 km WSW of Mardakert, 1100 m a.s.l., Quercus , Fagus , Acer etc. forest, litter, 2.VI.1987; 1 male, 5 females (ZMUM), NW above Bash-Layski ca 20 km NNW of Sheki, 1250 m a.s.l., Fagus , Carpinus , Acer etc. forest, litter, 3.V.1987, all leg. S. Golovatch and K. Eskov; 3 males, 14 females (ZMUM), AR Nakhichevan, S of Pass Bichenek, W of Shakhbuz, Quercus forest, 1900 m a.s.l., litter, 22.IV.1983, leg. S. Golovatch; 8 males, 7 females (ZMUM), Yarymdja SE of Altyagach, 1320 – 1350 m a.s.l., Fagus , Quercus , Carpinus etc. forest, litter and under bark, 20, 24 – 25.IV.1987; 1 female (ZMUM), Altyagach, 1050 – 1100 m a.s.l., Quercus , Fagus , Carpinus etc. forest, litter, 20 and 26.IV.1987, all leg. S. Golovatch and K. Eskov; 9 males, 1 female, 3 juv. (ZMUM), Altyagach, Yarymdja, 1200 – 1300 m a.s.l., forest, litter, 28 – 31.V.1998, leg. S. Dashdamirov; 5 males, 2 females (ZMUM), Shemakha, Pirkuli, 24.V.1988, leg. N. Loginova; 1 subadult male, 2 females (ZMUM), Shemakha Distr., Pirkuli Nature Reserve, near Observatorium, 1200 – 1250 m a.s.l., Quercus , Acer , Taxus etc. forest, litter, 30.IV.1987, leg. S.

Golovatch and K. Eskov; 6 males (ZMUM), Laza near Kusary, 10.VIII.1985, leg. H. Aliev; 2 males, 1 female (ZMUM), Talysh Mts, Lerik Distr., Zuvand, Galabyn, 1700 – 2000 m a.s.l., under stones on slopes, 10 – 11.X.1983, leg. S. Golovatch; 1 female (ZMUM), Lerik Distr., Farm Azerbaijan, forest, litter, 21.VII.1970, leg. N. Loginova; 19 males, 10 females (ZMUM), Talysh Mts., Yardymly Distr., Allar, 1700 – 1800 m a.s.l., Quercus , Carpinus , Acer etc. sparse forest, litter, rotten wood and under stones, 9.X.1983; 1 juv. (ZMUM), Talysh Mts., Lerik Distr., Zuvand, Gilidara, 1800 m a.s.l., semi-desert, 8.X.1983, all leg. S. Golovatch; 1 female (ZMUM), Talysh Mts., Zuvand, Djoni, 1500 m a.s.l., 28 – 29.V.1976, leg. V. Dolin; 5 males, 1 female (ZMUM), Agdash Distr., Turianchay Nature Reserve, orchard, 20.X.1988, leg. S. Dashdamirov; 2 males, 1 female (ZMUM), Lenkoran Distr., Hyrcan Nature Reserve, Apo ca 6 km SW of Alekseevka, Quercus , Acer , Carpinus, Parrotia etc. forest, litter, under stones and bark, 14 and 16.X.1983; 2 juv. (ZMUM), Lenkoran Distr., Hyrcan Nature Reserve, Alekseevka, 50 m a.s.l., Quercus, Parrotia , Carpinus etc. forest, litter, rotten wood, 13.X.1983, all leg. S. Golovatch; 4 females (ZMUM), Lenkoran Distr., Hyrcan Nature Reserve, near Alekseevka, 18.IV.1985; 2 males (ZMUM), Avrora, lowland forest, 21.IV.1985; 4 females (ZMUM), same locality, water reservoir, 18.IV.1985, all leg. E. Kupriyanova; 3 males (ZMUM), same locality, forest, litter, 26.I – 4.II.1985, leg. A. Druk; 1 male (ZMUM), Astara Distr., forest, 12.V.1969, leg. N. Loginova; 2 males, 10 females (HR), Təzə Alvadı, 1.6 road-km to Masallı, N39.075º, E48.6304º; 0 m a.s.l., 25.III.2015; 3 males, 3 females (HR), Hyrcan Nature Reserve, Piran, northwestern town exit; N38.734º, E48.6441º, 230 m a.s.l., 26.III.2015; 1 male, 2 females (HR), Hyrcan Nature Reserve, road Lenkoran — Lerik at km 32, N38.7638º, E48.5819º, 400 m a.s.l., 26.III.2015; 2 males (HR), Hyrcan Nature Reserve, Günəşli, 2 road-km to Lerik, N38.7975º, E48.4644º, 710 m a.s.l., 26.III.2015; 2 males, 4 females (HR), Hyrcan Nature Reserve, SW of Zünqüləş, N38.4493º, E48.7623º, 60 m a.s.l., 27.III.2015; 1 male, 2 females (HR), Hyrcan Nature Reserve, SW of Zünqüləş, N38.448º, E48.7597º, 130 m a.s.l., 27.III.2015; 4 females (HR), Hyrcan Nature Reserve, Daştatük, 1.3 road-km Xanbulan Reservoir, N38.6747º, E48.7622º, 110 m a.s.l., N38.6747º, E48.7622º, 27.III.2015; 1 female (HR), Hyrcan Nature Reserve, SW of Aşağı Apu, N38.6726º, E48.7362º, 180 m a.s.l., 27.III.2015; 1 female (HR), Qurbanəfəndi 1.3 road-km to Xanəgah, N40.8614º, E48.1132º, 640 m a.s.l., 30.III.2015, all leg. H. Reip and D. Antić.

6 males, 17 females, 21 juv. (SMF 5949, identified as B. ferrugineus , published by Golovatch 1981b), IRAN, East Azerbaijan Prov., Arasbaran Wildlife Refuge, Makidi, secondrary Quercus bush, 4.VI.1978, leg. J. Martens and H. Pieper.

Descriptive notes. Length ca 9 – 21 mm, width of midbody metazonae 0.9 – 2.7 mm (male, female), metazonite to prozonite width ratio ca 1.4 – 1.6 ( Fig. 10 A – 10 D ). Live coloration usually uniformly light brown to reddish or dark brown.

All characters as in B. assimilis ( Figs 10 – 27 ), except as follows. Paraterga from poorly to well developed, largely subhorizontal to slightly declivous, only rarely the first few postcollar ones faintly upturned; fore margins and anterolateral corners largely broadly and regularly rounded, only rarely angular; caudolateral corner mostly obtuse-angled until segment 4, largely pointed or nearly pointed starting with segment 5, drawn behind rear tergal margin starting with segments 6 – 14. Collum ovoid to, more rarely, transversely sublanceolate. Tergal setae mostly very short and subclavate. Metatergal sculpture usually rough, only in segments 18 or 17 and 18 nearly obliterate. Male prefemora bulging laterad, sphaerotrichomes present at least on tarsi. Gonopod ( Figs 13 , 14 , 16 , 17 , 19 , 21 – 25 , 27 ) bipartite, slightly falcate, like in B. assimilis , but with a typically bifid endomere (en) carrying a large, conspicuous, densely pilose field, while endomere (ex) basically with three distal outgrowths: teeth e and i, and lobe l.

At least six morphs are recognized here. Their more detailed characteristics are given in Table 1.

Remarks. Amongst the larger to largest Brachydesmus occurring in the Caucasus and adjacent parts of E Turkey and NW Iran, this highly polymorphous species poses certainly the greatest problems for a taxonomic treatment. Both Lignau (1915) and, later, Lohmander (1936), when describing their B. kalischewskyi , B. karawajewi , B. ferrugineus and B. talyschanus as new species, had only very few samples in their hands. In contrast to Lignau’s (1915) material which came from relatively small Abkhazia alone, Lohmander’s (1936) stemmed from various, often widely separated places scattered across the entire Caucasus. More recently, Ceuca (1975) reported B. ferrugineus from E Turkey, whereas Golovatch (1976) added his B. bidentatus . Had he been aware of the then very fresh paper by Ceuca, he would never have dared describe it as a new species, because the gonopods of “ ferrugineus ” from Turkey and of “ bidentatus ” from the central Caucasus Major were nearly identical! Enghoff (2006) not only reiterated Ceuca’s (1975) record of B. ferrugineus , but he also reported B. kalischewskyi from two localities in E Turkey.

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9) 7 Golovatch (1981b) recorded B. ferrugineus in the East Azerbaijan Province of Iran, reiterated by Enghoff and Moravvej (2005). That sole Iranian sample has again been borrowed for the present study.

Since our collections are indeed very large and densely blanket the entire region in question, we believe all those “species” actually represent one, B. kalischewskyi by priority. We shall try to bring enough evidence to support this conclusion, based on numerous measurements taken and abundant illustrations provided.

Treating the kalischewskyi complex requires recognition of several morphs which crudely correspond to the more or less typical forms of several ex-species. To avoid the use of Latin names to denominate these forms, the typical morph kalischewskyi will be referred to as A, while the morphs karawajewi , ferrugineus and talyschanus as B, C and D, respectively. Neither the variety batumensis, proposed by Lohmander (1936) as an Ajarian form of kalischewskyi , nor bidentatus are to be treated as separate morphs, as they sufficiently clearly represent morphs A and B, respectively. Instead, we distinguish two more morphs which show stable morphological characters and more or less coherent distributions. Thus, morph E corresponds to the populations from Svanetia, NW Georgia, whereas morph F to most of the populations inhabiting Armenia, Nakhichevan and Mountainous Karabakh. It must be emphasized that, for the time being, none of the morphs seem to be sufficiently clear-cut to warrant the status of a separate species. There are numerous morphological intergradations and transitions observed at least between the neighbouring morphs, but in certain cases at least size differences in co-occurring morphs are dramatic ( Fig. 11 ), especially when material is scant. However, everything merges when sufficiently large samples are available.

Sometimes the attribution of a sample to this or that morph is by necessity arbitrary, as certain (micro)populations appear to be disjunct morphologically. The choice in such cases is mainly determined by geographical proximity. Thus, it is often very hard to distinguish morph B from morph D. For instance, we have only located the latter in a single place in Georgia, whereas Lohmander identified two samples of morph D in that country ( Kobakhidze, 1965). Furthermore, the diversity of morphs could easily be extended, e.g., for several disjunct (micro)populations in Ajaria, Krasnodar Province ( Fig. 23 ), Azerbaijan or Armenia ( Fig. 24 ).

Molecular data, bar-coding for one thing, would certainly be most helpful in further refining the picture, but at the moment such information is unavailable.

Only in very few places such as near Sochi and within Adygea does morph B of B. kalischewskyi , represented there by smaller individuals about 9 – 11 mm in length, occur not only sympatrically, but even syntopically together with the superficially similar B. furcatus . However, these species are easily distinguished by angular versus broadly rounded anterolateral corners of the paraterga, and by a more distinct versus a strongly obliterated metatergal sculpture, respectively (cf. Figs 26 and 10A – 10D ), the males also by their disparate gonopod structure (cf. Figs 27 and 10E – 10G ).

Table 1 shows size variations in the main six morphs of B. kalischewskyi , coupled with short descriptive notes and numerous illustrations to facilitate their recognition. The main characters such as size, gonopod structure and the shape of the terga seem to form more or less random mosaics as well. There can be no doubt that the kalischewskyi complex is in a stage of active speciation. It shows mixed sympatric ( Bolnick and Fitzpatrick 2007; Fitzpatrick et al. 2008), mosaic or simple parapatric speciation modes (see synopsis in Mallet et al. 2009). We cannot rule out that some populations at the range peripheries might prove to represent proper species which genetically evolve independently in the modern sense of the species concept ( Queiroz 2007), yet with gene flows in the middle of the distribution area. Such a pattern does not automatically contradict the recognition of only a single species because gene flow is not so uncommon in animals ( Nosil 2008). Therefore, B. kalischewskyi as we treat it could become a model species for further evolutionary studies.

Map 2 roughly depicts the distribution of B. kalischewskyi in the Caucasus and adjacent parts of E Turkey and NW Iran. Each basic morph has its own colour, whereas syntopic, sometimes mixed morphs are shown in both colours involved. As one can easily see, the mosaic is really impressive. This is another strong argument for regarding the kalischewskyi complex as actually being represented by a single, highly polymorphous species. Some of the morphs do reveal coherent patterns. Thus, morph F (orange dots in Map 2) seems to be more or less restricted to a relatively small area in Svanetia, NW Georgia. Morph B (green dots in Map 2), apparently the most widespread in the region, seems to be the sole inhabitant of the eastern half of the Caucasus Major, including its northern macro-slope. However, most of the morphs tend to show no coherent patterns, being scattered across the region and even beyond in what seems to be a more or less random way.

MAP 2. Distribution of Brachydesmus kalischewskyi : morphs A – F shown in blue (A), green (B), red (C), pink (D), orange (E) and brown (F) circles, respectively; mixed or intermediate populations in two respective colours. The westernmost locality lying in north-central Turkey, which Ceuca (1975) misspelled as “Yalnarcen gecidii (Yalnircan silsilesi), 2500 m ”, apparently supports morph B and actually corresponds to Ilgazdaggecidi: Ilgazdag Pass, 15 km N of Tosya, N 41.144611º, E 34.062862º. It is omitted from Map 2.

No doubt this pretty crude picture requires further refinements through bringing in more evidence, both morphological and, especially, molecular-based. Discrimination or revival of some local species within the kalischewskyi complex is also possible in the future. At least some species names like karawajewi , ferrugineus or talyschanus are long available from a taxonomic point of view.

Polymorphism and distrubution mosaics like the ones demonstrated by B. kalischewskyi have long been reported in Polydesmidae . Thus, Epanerchodus polymorphus Mikhaljova and Golovatch, 1981 , from the Far East of Russia ( Mikhaljova and Golovatch 1981), based both on somatic and gonopod characters, shows two stable morphs in the male, but a complete transitional series in the female. Exactly the same pattern of dimorphic males, but polymophous females seems to be observed in another few highly localized cave species of Epanerchodus in Japan (e.g. Murakami 1970; Murakami and Kawasawa 1975). In contrast, the widespread East Asian E. orientalis Attems, 1901 shows only a gradual change in gonopod structure from north to south along Japan ( Nishikawa and Murakami 1993), but further southerly, in Taiwan and southern China, the morphological variation and distribution seem to grow increasingly sporadic and mosaic ( Golovatch et al. 2011, 2012).

Polymorphism and very complex mosaic distributions have also been reported in some other millipede groups, e.g. Xystodesmidae (Polydesmida) in North America ( Shelley and Whitehead, 1986). They concern not only likely infraspecific variations like those we face in Brachydesmus kalischewskyi , but also what Mesibov (2003) denominated as lineage mosaics, i.e. spatial mosaics of closely related species which are the result of differentiation in form and geographical space within monophyletic clades.

Figures

FIGURE 10. Brachydesmus furcatus Lohmander, 1936. A: habitus, entire body, dorsal view, male from Mount Kogot, Caucasian Nature Reserve, Krasnodar Prov., Russia. B – D: male from near Goryachiy Klyuch, Krasnodar Prov., Russia, B: anterior part of body, dorsal view; C: right half of metatergite 10, dorsal view; D: posterior part of body, dorsal view. E – F: male holotype of B. furcatus, right and left gonopods, lateral and mesal views, respectively (after Lohmander 1936); G: male holotype of B. furcatus exiguus Strasser, 1970, left gonopod, lateral view (after Strasser 1970). Designations: en, endomere; ex, exomere; t, tooth at base of femorite. Picture A by K. Makarov, taken not to scale. Line drawings B – D by S. Dashdamirov, scale bar: 1.0 mm (B – D), not to scale (E – G).

FIGURE 11. Size differences in a syntopic population of morphs A (larger) and B (smaller) of Brachydesmus kalischewskyi Lignau, 1915, males from Shatili, Argun River valley, Georgia. Scale bar: 1.0 mm. Picture by H. Reip.

FIGURE 12. Morph A of Brachydesmus kalischewskyi Lignau, 1915. A: male (?) from an unspecified locality in Abkhazia, length 15 mm. B: male from Khosta, Krasnodar Prov., Russia. C: male from Pass Mamisoni, W Georgia. Watercolour A after Lignau (1915); pictures B and C by K. Makarov, taken not to scale.

FIGURE 13. Morph A of Brachydesmus kalischewskyi Lignau, 1915: A – C: female from Pass Ertso, South Ossetia, anterior part of body, right half of metatergite 10 and caudal part of body, respectively, dorsal views. D – F: male from Pass Ertso, South Ossetia, anterior part of body, right half of metatergite 10 and caudal part of body, respectively, dorsal views. G – H: gonopods of a male from near Batumi, Ajaria, Georgia, ventral and mesal views, respectively. Designations: en, endomere; ex, exomere; i, mesal apical tooth of exomere; e, lateral apical tooth of exomere; l, subapical lobe of exomere. Line drawings A – F by S. Dashdamirov; G – H after Lohmander (1936). Scale bar: 1.0 mm (A – F), not to scale (G – H).

FIGURE 14. Morph B of Brachydesmus kalischewskyi Lignau, 1915: A: male from Yarymdja, E Caucasus Major, Azerbaijan, habitus, lateral view. B: male from Khosrov Nature Reserve, Caucasus Minor, S Armenia, habitus, dorsal view. C: left gonopod of male holotype of B. karawajewi Lohmander, 1928, from Teberda, Karachaevo-Cherkessia, Russia, mesal view. D – E: male from Verkhniy Agdan (now Gandzakar), Idjevan Distr., Armenia (not Georgia, as mistakenly stated by Lohmander 1936), ventral and mesal views, respectively. Designations: en, endomere; ex, exomere; i, mesal subapical tooth of exomere; e, lateral apical tooth of exomere; l, subapical lobe of exomere. Pictures A and B by K. Makarov; C – E after Lohmander (1928, 1936). Not to scale.

FIGURE 15. Morph C of Brachydesmus kalischewskyi Lignau, 1915. A: male from Lower Roki, South Ossetia, habitus, lateral view. B: male from near Krasnaya Polyana, Krasnodar Prov., Russia, habitus, dorsal view. Pictures by K. Makarov, taken not to scale.

FIGURE 16. Morph C of Brachydesmus kalischewskyi Lignau, 1915. A – C: male syntype of B. ferrugineus Lohmander, 1936 from near Borzhomi, Georgia, anterior part of body, right half of metatergite 10 and caudal part of body, respectively, dorsal views. D – F: male syntype of B. ferrugineus Lohmander, 1936 from Novyi Afon, Abkhazia, anterior part of body, right half of metatergite 10 and caudal part of body, respectively, dorsal views. G – H: gonopods of syntype of B. ferrugineus from Novyi Afon, Abkhazia, ventral and mesal views, respectively. Designations: en, endomere; ex, exomere; i, mesal apical tooth of exomere; e, lateral apical tooth of exomere; l, subapical lobe of exomere. Line drawings A – F by S. Dashdamirov; G – H after Lohmander (1936). Scale bar: 1.0 mm (A – F), not to scale (G – H).

FIGURE 17. Morph D of Brachydesmus kalischewskyi Lignau, 1915. A: male from Talysh Mts, Lerik Distr., Farm Azerbaijan, habitus, dorsal view; B: male from Talysh Mts, Lerik Distr., Zuvand, Galabyn, habitus, dorsal view. Pictures by K. Makarov, taken not to scale.

FIGURE 18. Morph D of Brachydesmus kalischewskyi Lignau, 1915, male holotype of B. talyschanus Lohmander, 1936: A – C: anterior part of body, right half of metatergite 10 and caudal part of body, respectively, dorsal views. D – E: left gonopod, mesal and ventral views, respectively. Designations: i, mesal subapical tooth of exomere; e, lateral apical tooth of exomere; l, subapical lobe of exomere. Line drawings A – C by S. Dashdamirov; D – E after Lohmander (1936). Scale bar: 1.0 mm (A – C), not to scale (D – E).

FIGURE 19. Morph E of Brachydesmus kalischewskyi Lignau, 1915, males from near Mestia, Svanetia, Georgia. A – B: habitus, dorsal views. C: leg 7, lateral view. D – E: right gonopod, lateral and mesal views, respectively. Designations: i, mesal subapical tooth of exomere; e, lateral apical tooth of exomere; l, subapical lobe of exomere. Pictures A and B by K. Makarov. Scale bars: 0.2 mm (C – E), not to scale (A – B).

FIGURE 20. Morph F of Brachydesmus kalischewskyi Lignau, 1915. A: male from near Kadjaran, Megri Mt. Range, Armenia. B: male from Shikahoh Nature Reserve, Megri Mt. Range, Armenia. C: male from Shnokh, Armenia, habitus, dorsal, lateral and dorsal views, respectively. Pictures by K. Makarov, taken not to scale.

FIGURE 21. Morph F of Brachydesmus kalischewskyi Lignau, 1915. A – C: male from Agveran, Armenia, anterior part of body, right half of metatergite 10 and caudal part of body, respectively, dorsal views. D – E: male from near Shvanidzor, Megri Mt. Range, Armenia, right gonopod, ventral and mesal views, respectively. F: male from near Badara, Mountainous Karabakh, Azerbaijan, right gonopod, mesal view. Line drawings A – C by S. Dashdamirov. Scale bars: 1.0 mm (A – C), 0.2 mm (D – F).

FIGURE 22. Examples of gonopods of Brachydesmus kalischewskyi Lignau, 1915. A: male from Pasture Abago, Caucasian Nature Reserve, Adygea, morph A. B: male from Khosrov Nature Reserve, S Armenia, morph B. C: male from Pass Ertso, South Ossetia, morph A. D: male from Nadirkhanly, Mountainous Karabakh, Azerbaijan, morph F. E: male from Pass Voratan, Zangezur Mt. Range, Armenia, morph B. F: male from near Pass Krestovyi, Georgia, morph B. Designations: i, mesal subapical tooth of exomere; e, lateral apical tooth of exomere; l, subapical lobe of exomere. Scale bar: 0.2 mm.

FIGURE 23. Examples of gonopods of Brachydesmus kalischewskyi Lignau, 1915. A – C: male from Psebe, Krasnodar Prov., Russia, morph A?, lateral, mesal and ventral views, respectively. D – F: male from Zeraboseli, Ajaria, Georgia, morph A?, sublateral, dorsal and ventral views, respectively. Designations: i, mesal subapical tooth of exomere; e, lateral apical tooth of exomere. Scale bars: 0.1 mm.

FIGURE 24. Examples of gonopods of Brachydesmus kalischewskyi Lignau, 1915. A – C: male from Farm Azerbaijan, Talysh Mts, Azerbaijan, morph D, dorsal, ventral and mesal views, respectively. D – F: male from Shishkert, Shikahoh Nature Reserve, Armenia, morph B?, mesal, lateral and ventral views, respectively. Designations: i, mesal, subapical tooth of exomere; e, lateral apical tooth of exomere; l, subapical lobe of exomere. Scale bar: 0.1 mm.

FIGURE 25. Examples of gonopods of Brachydesmus kalischewskyi Lignau, 1915. A – C: male from Farm Azerbaijan, Talysh Mts, Azerbaijan, morph D, lateral, mesal and ventral views, respectively. D: male from Arasbaran Wildlife Refuge, East Azerbaijan Prov., Iran, morph D, mesal view. Designations: i, mesal subapical tooth of exomere; e, lateral apical tooth of exomere; l, subapical lobe of exomere. Scale bars: 0.2 mm.

FIGURE 26. Brachydesmus kalischewskyi Lignau, 1915, morph A male from Pasture Abago, Adygea, Russia. A: habitus, lateral view. B – D: anterior, middle and caudal parts of body, respectively, dorsal views. Pictures by K. Makarov, taken not to scale.

FIGURE 27. Brachydesmus kalischewskyi Lignau, 1915, morph B, males from Pasture Abago and Arkhyz, Adygea, Russia, respectively. A: leg 7, lateral view. B: left gonopod, mesal view. C: leg 7, lateral view. D: left gonopod, mesal view. Scale bars: 1.15 mm (A), 0.2 mm (B and D), 0.1 mm (C).