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

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Brachydesmus (Haplobrachidesmus) kalischewskyi Lignau, 1915
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Brachydesmus (Haplobrachidesmus) kalischewskyi Lignau, 1915 

Figs 11 – 27View FIGURE 11View FIGURE 12View FIGURE 13View FIGURE 14View FIGURE 15View FIGURE 16View FIGURE 17View FIGURE 18View FIGURE 19View FIGURE 20View FIGURE 21View FIGURE 22View FIGURE 23View FIGURE 24View FIGURE 25View FIGURE 26View FIGURE 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 DView FIGURE 10). Live coloration usually uniformly light brown to reddish or dark brown.

All characters as in B. assimilis  ( Figs 10 – 27View FIGURE 10View FIGURE 11View FIGURE 12View FIGURE 13View FIGURE 14View FIGURE 15View FIGURE 16View FIGURE 17View FIGURE 18View FIGURE 19View FIGURE 20View FIGURE 21View FIGURE 22View FIGURE 23View FIGURE 24View FIGURE 25View FIGURE 26View FIGURE 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 13View FIGURE 13, 14View FIGURE 14, 16View FIGURE 16, 17View FIGURE 17, 19View FIGURE 19, 21 – 25View FIGURE 21View FIGURE 22View FIGURE 23View FIGURE 24View FIGURE 25, 27View FIGURE 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. 11View FIGURE 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. 23View FIGURE 23), Azerbaijan or Armenia ( Fig. 24View FIGURE 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 26View FIGURE 26 and 10A – 10DView FIGURE 10), the males also by their disparate gonopod structure (cf. Figs 27View FIGURE 27 and 10E – 10GView FIGURE 10).

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.

ZISP

Zoological Institute, Russian Academy of Sciences

ZMUM

Zoological Museum, University of Amoy

SMNG

Senckenberg Museum fuer Naturkunde Goerlitz

Kingdom

Animalia

Phylum

Arthropoda

Class

Diplopoda

Order

Polydesmida

Family

Polydesmidae

Genus

Brachydesmus

Loc

Brachydesmus (Haplobrachidesmus) kalischewskyi Lignau, 1915

Golovatch, Sergei, Evsyukov, Aleksandr & Reip, Hans 2016
2016
Loc

Brachydesmus karawaewi

Bababekova 1996: 90
1996
Loc

Brachydesmus

Bababekova 1996: 90
Loksina 1979: 384
1979
Loc

Brachydesmus

Barjadze 2015: 21
Chumachenko 2012: 452
Enghoff 2006: 192
Talikadze 1984: 143
Golovatch 1981: 110
Kobakhidze 1965: 390
1965
Loc

Brachydesmus

Barjadze 2015: 21
Kokhia 2011: 28
Enghoff 2006: 192
Enghoff 2005: 69
Loginova 1993: 151
Talikadze 1984: 143
Golovatch 1981: 110
Ceuca 1975: 91
Kobakhidze 1965: 390
1965
Loc

Brachydesmus

Bababekova 1996: 90
Talikadze 1984: 143
Rasulova 1973: 520
Samedov 1972: 1245
Kobakhidze 1965: 391
1965
Loc

Brachydesmus

Zuev 2014: 348
Chumachenko 2012: 452
Talikadze 1984: 143
Kobakhidze 1965: 390
Lang 1959: 1790
1959
Loc

Brachydesmus

Loksina 1979: 384
Attems 1940: 136
1940
Loc

Brachydesmus

Loksina 1979: 384
Lang 1959: 1790
Attems 1940: 137
1940
Loc

Brachydesmus

Loksina 1979: 384
Lang 1959: 1790
Attems 1940: 137
1940
Loc

Brachydesmus

Loksina 1979: 384
Lang 1959: 1790
Attems 1940: 137
Lohmander 1936: 18
1936
Loc

Brachydesmus

Lohmander 1936: 28
1936
Loc

Brachydesmus

Lohmander 1936: 22
1936
Loc

Brachydesmus kalischewskyi

Lignau 1915: 370
Loc

Brachydesmus karawajewi

Lohmander 1928: 530
Loc

Brachydesmus (Haplobrachydesmus) (sic!) talyschanus

Lohmander 1936: 26
Loc

Brachydesmus (Haplobrachydesmus) bidentatus Golovatch, 1976 : 1568

Golovatch 1976: 1568