Lignau, 1915 : 370 Brachydesmus ( Haplobrachydesmus ) Lohmander, 1936 : 18 Attems, 1940 : 137 Lang, 1959 : 1790 Lokšina and Golovatch, 1979 : 384 Brachydesmus Kobakhidze, 1965 : 390 Golovatch, 1981a : 110 Talikadze, 1984 : 143 Enghoff, 2006 : 192 Chumachenko, 2012 : 452 Barjadze et al ., 2015 : 21 Lohmander, 1928 : 530 Brachydesmus ( Haplobrachydesmus ) Lohmander, 1936 : 28 Brachydesmus ( Haplobrachydesmus ) Attems, 1940 : 136 Lokšina and Golovatch, 1979 : 384 Brachydesmus Lang, 1959 : 1790 Kobakhidze, 1965 : 390 Talikadze, 1984 : 143 Chumachenko, 2012 : 452 Zuev, 2014 : 348 Brachydesmus karawaewi Bababekova, 1996 : 90 Lohmander, 1936 : 22 Brachydesmus ( Haplobrachydesmus ) Attems, 1940 : 137 Lang, 1959 : 1790 Lokšina and Golovatch, 1979 : 384 Brachydesmus Kobakhidze, 1965 : 390 Ceuca, 1975 : 91 Golovatch, 1981a : 110 Talikadze, 1984 : 143 Loginova, 1993 : 151 Enghoff and Moravvej, 2005 : 69 Enghoff, 2006 : 192 Kokhia, 2011 : 28 Barjadze et al ., 2015 : 21 Lohmander, 1936 : 26 Brachydesmus ( Haplobrachydesmus ) Attems, 1940 : 137 Lang, 1959 : 1790 Lokšina and Golovatch, 1979 : 384 Brachydesmus Kobakhidze, 1965 : 391 Samedov et al ., 1972 : 1245 Rasulova and Rakhmanov, 1973 : 520 Talikadze, 1984 : 143 Bababekova, 1996 : 90 Golovatch, 1976 : 1568 Brachydesmus Lokšina and Golovatch, 1979 : 384 Bababekova, 1996 : 90 The millipede family Polydesmidae in the Caucasus (Diplopoda: Polydesmida) Golovatch, Sergei Evsyukov, Aleksandr Reip, Hans Zootaxa 2016 2016-03-01 4085 1 1 51 MVY5 Lignau, 1915 Lignau 1915 [151,942,1196,1222] Diplopoda Polydesmidae Brachydesmus GBIF Animalia Polydesmida 20 21 Arthropoda species kalischewskyi Haplobrachidesmus     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).    Brachydesmuskalischewskyi—  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).    Brachydesmuskarawajewi—  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).    Brachydesmusferrugineus—  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).    Brachydesmustalyschanus—  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.    Brachydesmusbidentatus—  Lokšina and Golovatch, 1979: 384(M);  Bababekova, 1996: 90(R).   Material examined.3 males, 2 females (ZMUM), RUSSIA, Krasnodar Prov., Tuapse Distr., 15 kmSE of Novomikhailovskii, Psebe, deciduous forest, litter, under bark and stones, 29.X.1981; 1 female (ZMUM), Krasnodar Prov., Sochi, Dagomys,  Quercusforest, 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,  Taxusand  Buxusforest, litter, 28.X.1981, all leg. S. Golovatch; 1 male, 2 females (ZMUM), same locality,  Taxusand  Buxusforest, III.2006, leg. Yu. Chumachenko; 2 females, 1 subadult female (ZMUM), Krasnodar Prov., Sochi, 5.5 kmNE of Krasnaya Polyana, lower course of Achipse River, ca 600 ma.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 ma.s.l., N43.638º, E40.2481º, 17.06.1997, leg. I. Belousov; 1 male (ZMUM), Krasnodar Prov., Tuapse Distr., Georgievskoe Forestry,  Castaneaforest, 6.V.1954, leg. K. Arnoldi, det. H. Lohmander, det. C. Dziadosz, 1968; 3 juv. (ZMUM), Krasnodar Prov., Caucasian Nature Reserve, Pslukh, ca 20 kmE of Krasnaya Polyana, 1000 ma.s.l.,  Fagus, Quercus, Abies, Piceaetc. forest, litter, 11 –12.VIII.1986; 5 males, 3 females, 3 juv. (ZMUM), Krasnodar Prov., Goryachiy Klyuch, 12 kmSW of Fanagoriyskoye, Cave Fanagoriyskaya, 19.V.1983; 7 juv. (ZMUM), Krasnodar Prov., Mostovskoy Distr., Yaroslavskaya, 300 ma.s.l.,  Quercus, Acer, Corylusetc. 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 ma.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 ma.s.l.,  Abiesand  Fagusforest, litter, under bark and stones, 24 –26.V.1985; 5 females (ZMUM), same locality, 1700 –1850 ma.s.l.,  Abiesand  Fagusforest,  Rhododendronthicket, litter, 24 –26.V.1985; 2 females (ZMUM), Krasnodar Prov., Caucasian Nature Reserve, Mt Aishkho-2 E of Pslukh, ca 28 kmE of Krasnaya Polyana, S slope, 1600 ma.s.l.,  Abies, Picea, Fagus, Betulaetc. forest, litter, 12.VIII.1986; 1 male, 1 female (ZMUM), Karachaevo-Cherkessia, ca 30 kmS of Kurdjinovo, Laba Valley, 4 kmN of Damkhurts, 1050 –1100 ma.s.l.,  Fagus, Acer, Piceaetc. forest, litter and under bark, 4.VIII.1986; 1 juv. (ZMUM), Karachaevo-Cherkessia, Teberda Nature Reserve, above Teberda, Mt Malaya Khatipara, 2400 ma.s.l., subalpine meadow, under stones, 29 –30.V.1985; 1 female (ZMUM), same locality, Kizgich Canyon, N of Arkhyz, 1550 –1600 ma.s.l.,  Abies, Picea, Pinus, Fagus, Betulaetc. forest, litter, under bark and stones, 5.VI.1985; 27 males, 1 female, 3 juv. (ZMUM), Karachaevo-Cherkessia, Lower Teberda S of Karachaevsk, 1000 ma.s.l.,  Quercus, Fagus, Aceretc. forest, litter, 3.VIII.1986, all leg. S. Golovatch; 2 females (ZMUM), Karachaevo-Cherkessia, Teberda, ca 1200 –2000 ma.s.l., VI –VII.1908, leg. A.P. Zolotarev; 31 juv. (ZMUM), Kabardino-Balkaria, Lower Chegem, 800 ma.s.l.,  Fagusforest, litter, 14.VII.1986; 18 males, 2 females (ZMUM), North Ossetia, Novo-Georgievskoye near Mozdok,  Quercus, Acer, Alnus, Fraxinusetc. 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,  Quercusand  Alnusscrub on slope, litter and under stones 2.VI.1982; 12 males, 5 females (ZMUM), North Ossetia, S of Alagir, 200 ma.s.l.,  Acer, Fagusetc. forest, litter, 17.X.1987; 3 males, 4 females, 9 juv. (ZMUM), Ingushetia, Assa Valley, ca 9 kmSSW of Muzhichi, 800 ma.s.l.,  Fagus, Alnus, Carpinusetc. forest, litter, under bark and stones, 15.VII.1986; 2 females, 2 juv. (ZMUM), Chechnya, 5 kmW of Shaami-Yurt, ca 26 kmW of Groznyi,  Acerforest, litter, 6.VI.1982; 1 female (ZMUM), Dagestan, Chiragchay Valley, Sardarkent between Kasumkent and Khiv, 650 ma.s.l.,  Crataegus, Rosa,  Carpinus, Juglansetc. scrub, litter, 24.X.1987; 1 male (ZMUM), Dagestan, Samur Valley, Garakh ca 35 kmSW of Magaramkent, 700 –800 ma.s.l.,  Crataegus, Quercus, Acer, Rosaetc. scrub, litter, 23.X.1987, all leg. S. Golovatch; 10 males, 8 females (ZMUM), Dagestan, Kaytag Distr., Karatsan, 6.X.2012, leg. P. Gamilova.   FIGURE 12.Morph A of  Brachydesmus kalischewskyiLignau, 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 kalischewskyiLignau, 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). 1 female (ZISP), SOUTH OSSETIA, near Kvaisi, under stones, 1700 ma.s.l., 19.X.1956, leg. K. Gorodkov; 1 male, 1 female (ZMUM), 15 kmE of Kvaisi, Surami Mt. Ridge, Pass Ertso, ca 1000 ma.s.l., deciduous brush, along stream, litter and stones, 20.X.1981; 4 males, 2 females (ZMUM), Lower Roki S of Pass Rokskii, 1500 ma.s.l.,  Fagus, Betulaetc. forest, litter, 17.X.1987, all leg. S. Golovatch.   FIGURE 14.Morph B of  Brachydesmus kalischewskyiLignau, 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. karawajewiLohmander, 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 kalischewskyiLignau, 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 kalischewskyiLignau, 1915. A–C:male syntype of  B. ferrugineusLohmander, 1936from 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. ferrugineusLohmander, 1936from 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. ferrugineusfrom 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).  2 males, syntypesof  B. ferrugineus( ZISP),  ABKHAZIA, “W Transcaucasia, Olginskoe, Tsebelda, 20.V.2012, leg. V. Lindholm”; 1 male, syntypeof  B. ferrugineus(gonopods missing) ( ZISP), “Black Sea Gouvernement, Novyi Afon, 12.V.1912, leg. V. Lindholm”; 1 female( ZISP), “ 1 femalejuv., Polydesmussp., det. C. Dziadosz, 1967”, Gagra, Mamsdyshkha, ca 2000 m, VIII.1911, leg. V. Isaev?; 1 female( ZMUM), Lake Ritsa, 1 800 ma.s.l.,  Abiesand  Fagusforest, litter, 13.IX.1985, leg. I. Ushakov; 2 males, 1 female, 1 juv.( ZMUM), Lake Ritsa, 950 –1100 ma.s.l.,  Fagus, Abies, Picea, Aceretc. forest, litter, under bark and stones, 13 –14.VIII.1986, leg. S. Golovatch; 1 male, 1 juv.( SMNG14685), 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 kmN of Tsebelda, 550 ma.s.l.,  Alnusforest, litter and under bark, 19.VIII.1986; 1 juv.( ZMUM), Sukhumi Distr., Tsebelda, 300 ma.s.l.,  Carpinus, Acerand  Buxusscrub, litter, 19.VIII.1986; 2 males, 1 juv.( ZMUM), Sukhumi Botanical Garden, 9.X.1978, leg. S. Golovatch; 1 female, 1 juv.( ZMUM), 2 kmN of Duripsh N of Gudauta,  Faguslog, 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 ma.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 kmSSW of Nabeglavi, 2000 ma.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 ma.s.l., deciduous forest, litter and under stones, 13.X.1981; 3 females (ZMUM), same locality, 800 ma.s.l.,  Rhododendronthicket, litter, 13.X.1981, all leg. S. Golovatch; 5 males, 6 females (ZMUM), same locality, 450 –600 ma.s.l., deciduous forest, 1 –3.VI.1981; 2 males, 1 female (ZMUM), same locality, 600 –700 ma.s.l.,  Rhododendronthicket, 2.VI.1981, all leg. S. Golovatch and J. Martens; 17 juv. (ZMUM), Ajaria, between Kobuleti and Ureki, seashore  Pinusforest, litter, 5 ma.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,  Platanusforest, litter, 9.X.1981; 9 males, 2 females, 2 juv. (ZMUM), Ajaria, Khulo, 900 ma.s.l.,  Quercus, Abies, Alnusetc. forest, litter, 11.X.1981; 4 juv. (ZMUM), Ajaria, E of Khulo, 950 ma.s.l., deciduous forest on rock, near spring, litter, 10.X.1981; 1 juv. (ZMUM), Ajaria, 6 kmW of Khulo, 800 ma.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 ma.s.l., gardens, 30.V.1981; 7 males, 20 females (ZMUM), Ajaria, Batumi Botanical Garden, 20 –150 ma.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 ma.s.l.,  Piceaforest, under stones, litter, umder bark, 2.IX.1999, leg. L. Penev; 1 juv. (ZMUM), Ajaria, E of Pass Goderdzi, ca 1620 ma.s.l.,  Abiesand  Pinusforest, N41º38’42.6", E42º35’45.5", 22.07.2012, leg. Y. Marusik; 6 males, 12 females (ZMUM), 15 kmW of Adigeni, 1500 –1700 ma.s.l,  Abies, Picea, Fagus, Aceretc. forest, litter, logs, under stones, 14 –15.V.1983, leg. S. Golovatch; 4 females (ZMUM), W of Adigeni, 1500 –1700 ma.s.l.,  Abies, Picea, Fagusetc. forest, 21 –22.VI.1978, leg. V. Dolin; 13 females (ZMUM), Borzhomi Nature Reserve, Baniskhevi Valley, 800–900 ma.s.l.,  Picea, Fagus, Carpinusetc. forest, litter, under bark and stones, 12 and 16.V.1983; 1 male (ZMUM), S of Bakuriani,  Pinusand  Fagusforest, 1750 ma.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 kmE of Tkibuli,  Quercus, Fagus, Castaneaetc. forest, litter and under stones along spring, 23.X.1981, leg. S. Golovatch; 1 female (ZMUM), Mukhura ca 15 kmE of Tkibuli, 700 –800 ma.s.l.,  Castanea, Fagus, Carpinusetc. forest, litter, under bark and stones, 7 –9.V.1987, leg. S. Golovatch and K. Eskov; 8 juv. (ZMUM), ca 10 kmE of Tkibuli, Mukhura, 1010 ma.s.l.,  Carpinusand  Fagusforest with  Buxusthickets, N42°20’05,8”, E43°02’38,5”, 23.VII.2012, leg. Yu. Marusik; 1 male, 1 female, 3 juv. (ZMUM), Svanetia, ca 40 kmW of Mestia, Kherkhvashi E of Nakra (= Naki), 1200 –1700 ma.s.l.,  Quercus, Fagus, Carpinus, Picea, Abiesetc. forest, litter and under bark, 21.VIII –21.IX.1986; 4 males, 1 female (ZMUM), Svanetia, Mestia, 1500 ma.s.l.,  Betulaand  Rhododendronon 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 kmW of Mestia, above Kherkhvashi, 1900 –2200 ma.s.l., timberline (  Rhododendron, Picea, Abies) and subalpine meadows, 21.IX.1986; 5 males, 5 females (ZMUM), Ambrolauri Distr., Cave Nikortsminda near Nikortsminda, 1100 ma.s.l., 40 –50 mdeep, on walls and rotten wood, 22.X.1981; 1 male (ZMUM), same locality, 1150 ma.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 ma.s.l.,  Fagusand  Buxusforest, 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 kmNE of Shovi, Gurshevi near Pass Mamisoni, 2000 –2200 ma.s.l.,  Abies, Fagus, Alnusetc. forest, litter and under stones, 21.X.1981; 3 males, 3 females (ZMUM), Oni Distr., SW of Pass Mamisoni, 2500 ma.s.l.,  Salixbush along spring in subalpine meadow, litter and under stones, 21.X.1981; 1 male, 4 females (ZMUM), Oni Distr., Shovi,  Fagus, Alnus, Abiesetc. 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 ma.s.l.,  Fagus, Picea, Aceretc. forest, litter and under bark, 16 –18.V.1987, leg. S. Golovatch and K. Eskov; 1 female (ZMUM), Algeti State Reserve, Manglisi,  Quercusforest, 12.VI.1984, leg. E. Kvavadze; 4 males, 4 females (ZMUM), Saguramo Nature Reserve, NE of Mtskheta, Zedazeni, 1100 –1200 ma.s.l.,  Fagus, Carpinus, Aceretc. forest, litter and under bark, 20.V.1987; 1 female (ZMUM), Lake Shaori, mixed forest, litter, N42º23’46.6", E43º01’47.7", ca 1180 ma.s.l., 19.07.2012, leg. Y. Marusik; 2 males, 4 females (ZMUM), Batsaro Nature Reserve, ca 20 kmN of Akhmeta, 800 –850 ma.s.l.,  Fagus, Castanea, Aceretc. forest, litter, 5 –6.V.1987; 4 males, 2 females (ZMUM), Pass Magalakhari between Akhmeta and Tianeti, 1200 ma.s.l.,  Fagusand  Carpinusforest, litter and under bark, 6.V.1987; 9 juv. (ZMUM), Lagodekhi Nature Reserve, trail to meteorological station, ca 1835 ma.s.l.,  Carpinusforest, N41º51’38.0", E46º20’26.5", 27.07.2012, leg. Yu. Marusik; 4 males, 4 females (ZMUM), Kartli Mt. Ridge, Pass Sabaduri ca 40 kmNNE of Tbilisi, 1400 ma.s.l.,  Fagusforest, litter and under bark, 6.V.1987; 2 males, 1 female (ZMUM), Surami Mt. Ridge, SE of Pass Djvari, between Gomi and Sachkhere, 850 ma.s.l.,  Alnus, Fagus, Rhododendronetc. forest, litter, 7.V.1987; 1 male, 2 females (ZMUM), N of Kvareli, 700 –750 ma.s.l.,  Fagus, Carpinus, Quercusetc. 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 kmNE of Akhmeta, near Babaneuri, 500 ma.s.l.,  Fagus, Quercus,  Carpinusetc. forest, litter, 4 –5.V.1987; 1 female, 4 juv. (ZMUM), 10 kmN of Djvari, 800 ma.s.l.,  Buxus, Fagus, Picea, Taxusetc. forest, litter, 20 –21.VIII.1986; 1 male, 3 females, 1 juv. (ZMUM), Kazbegi, 2000 ma.s.l.,  Pinusand Betulaforest 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 ma.s.l.,  Rhododendronbush, 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 ma.s.l., VIII./ IX.1999, leg. O. Tietz; 1 male (HR), Tusheti, slope between Omalo and Khenako, 1810 ma.s.l., N42.3719º, E45.6444º, 4.IX.2009; 1 male (HR), Tusheti, pass between Omalo and Diklo, 2270 ma.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 ma.s.l., N42.5675º, E45.2203º, 10.IX.2009; 1 female (HR), Khevsureti, Churo Valley, mouth of Khone Skali River, 1980 ma.s.l., N42.5683º, E45.2322º, 10.IX.2009; Khevsureti, Khone Skali River valley, between Khone and Khone Chala, 1870 ma.s.l., N42.5675º, E45.2203º, 10.IX.2009; 3 males, 1 female (HR), Khevsureti, ca 200 mdownstream of Shatili, 1415 ma.s.l., N42.6594º, E45.1653º, 12.IX.2009; 4 males, 1 female (HR), Khevsureti, Argun River valley, between Shatili and Georgi Tsvinda, 1580 ma.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 ma.s.l., N41.8611º, E43.9444º, 16.IX.2011; 1 male (HR), Samtskhe–Djavakheti, Borzhomi to Chitakhevi, junction to Mtsvane Monastery, 850 ma.s.l., N41.8039º, E43.3181º, 17.IX.2011; 1 male (HR), Samtskhe-Djavakheti, Bakuriani, 7.5 road-km to Tskhratskaro Ugheltekhili Pass, 2080 ma.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 ma.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 ma.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 ma.s.l., N41.5278, E41.7211, 29.IX.2011; 3 males (HR), Ajaria, NE of Dologani, 700 m, 100 ma.s.l., N41.5525º, E41.8019º, 29.IX.2011; 1 male (HR), Mtskheta–Mtianeti, Saguramo, 1.2 road-km towards Zedazeni Monastery, 770 ma.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 ma.s.l., N42.1392º, E44.8167º, 5.X.2011; 1 male (HR), Samegrelo-Zemo Svaneti, Nodashi, 1100 ma.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 ma.s.l., N42.7058º, E44.6267º, 6.X.2012; 3 females (HR), Mtskheta–Mtianeti, Andaki River valley 2.2 kmupstream 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 kmSW of Kobi, N42.5553, E44.4973º, 12.VIII.2014; 1 male (HR), Imereti, Nakeralos Ugheltekhili Pass, 730 mSSE, N42.3708, E43.0428º, 14.VIII.2014; 1 male (HR), Samegrelo and Zemo Svaneti, Ushguli, entrance to Enguri (= Inguri) River canyon, 2 kmW of Ushguli, N42.9147º, E42.9747º, 19.VIII.2014, all leg. F. Walther.   FIGURE 17.Morph D of  Brachydesmus kalischewskyiLignau, 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 kalischewskyiLignau, 1915, male holotype of  B. talyschanusLohmander, 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). 3 males, 1 female (ZMUM), ARMENIA, Azizbekov Distr., Zangegur Mt. Ridge, Pass Voratan, subalpine meadow, 2100 –2200 ma.s.l., under stones, 21.IV.1983; 1 male (ZMUM), near Azizbekov, 1200 ma.s.l.,  Artemisiagrassland, under stones, 21.IV.1983; 2 males, 6 females (ZMUM), Ararat Distr., Khosrov Nature Reserve, 1450 –1550 ma.s.l.,  Juniperus, Quercus, Crataegusetc. 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 ma.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 kmN of Shvanidzor, sparse  Quercusforest, 1200 –1300 ma.s.l., litter, under stones and bark, 24.IV.1983; 8 males, 13 females, 3 juv. (ZMUM), environs of Megri, xerophytous bare canyon, 1000 ma.s.l., sparse J uniperusand Paliurusbush, under stones, 24.IV.1983; 13 males, 18 females (ZMUM), Megri Distr., SSE of Lichk, Megri River valley, 1530 ma.s.l.,  Quercusforest, litter, under stones and in rotten wood, 25.IV.1983; 17 males, 15 females (ZMUM), Megri Distr., above Kuris, 1500 ma.s.l.,  Quercus, Aceretc. forest, litter, under bark and stones, 26.IV.1983; 7 males, 4 females, 4 juv. (ZMUM), Megri Distr., 6 kmN of Shvanidzor, sparse  Quercusforest, 1200 –1300 ma.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 ma.s.l.,  Quercusforest on steep slope, litter and logs, 27.IV.1983; 38 males, 29 females (ZMUM), Kafan Distr., Shikahoh Nature Reserve, Shikahoh, 900 –950 ma.s.l.,  Quercus, Fagus, Carpinusetc. forest along stream, litter, under logs and stones, 28.IV.1983; 7 males, 7 females (ZMUM), Kafan Distr., Shikahoh Nature Reserve, Shishkert, 1700 –1800 ma.s.l.,  Quercus, Fagus, Carpinusforest, 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 ma.s.l.,  Quercus, Fagus, Carpinusetc. 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  Platanusstand along river, litter, rotten wood, under stones, 30.IV.1983; 3 males, 1 female, 1 juv. (ZMUM), Legvaz, 4 kmNWN of Megri,  Juniperus, Quercusetc. scrub with Paliurusand Rosa, litter and under stones, 1000 ma.s.l., 24 –25.IV.1983; 3 females (ZMUM), Stepanavan, foot of Mt Archasar, 1400 ma.s.l.,  Quercusforest 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 ma.s.l.,  Quercus, Acer, Fagusetc. forest, litter, 22.V.1987, leg. S. Golovatch and K. Eskov; 2 males (ZMUM, ex Coll. HNHM), Karashamb, Agveran, 1500 ma.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 ma.s.l.,  Carpinusforest, litter, 24.V.1987, leg. S. Golovatch and K. Eskov; 2 females (ZMUM), Dilizhan Nature Reserve, Akhnabat Valley near Salakh, 1400 –1500 ma.s.l.,  Fagus, Taxus, Carpinusetc. forest, litter, under bark and stones, 17.IV.1983, leg. S. Golovatch; 1 female (ZMUM), Dilizhan Nature Reserve, Agartsyn, 1350 –1400 ma.s.l.,  Fagus, Aceretc. forest, litter, 28 –29.V.1987; 1 male, 7 females (ZMUM), W of Shamshadyn, halfway between Berd and Idjevan, 1500 –1600 ma.s.l.,  Fagus, Carpinus, Aceretc. forest, litter and under bark, 26 –27.V.1987; 6 males, 4 females (ZMUM), Berdavan ca 10 kmS of Noemberian, 900 –950 ma.s.l.,  Quercus, Carpinus, Aceretc. 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 kmWNW of Dashbulag, near Badara,  Quercus, Carpinusetc. forest, 850–900 ma.s.l., litter, under bark and stones, 2.V.1983; 9 males, 5 females (ZMUM), Mountainous Karabakh, Dashalty near Shusha, 1100 –1300 ma.s.l.,  Quercus, Carpinusetc. 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 kmWSW of Mardakert, 800 –850 ma.s.l.,  Quercus, Aceretc. forest, litter, 1 –2.VI.1987; 4 males, 5 females (ZMUM), Mountainous Karabakh, Chilisa ca 7 kmN of Kelbadjar, 1450 –1500 ma.s.l.,  Quercus, Carpinus, Aceretc. forest, litter and under bark, 31.V.1987; 10 males, 5 females, 5 juv. (ZMUM), Mountainous Karabakh, Nadirkhanly ca 12 kmNE of Kelbadjar, 1200 ma.s.l.,  Fraxinusand  Juglans regiastand on rock, litter and under stones, 1.VI.1987; 3 females (ZMUM), Mountainous Karabakh, Lesser Istisu halfway between Kelbadjar and Istisu, 1550 ma.s.l.,  Quercusand  Acerscrub on slope, litter, 31.V.1987, all leg. S. Golovatch and K. Eskov; 5 males, 2 females (ZMUM), Istisu ca 8 kmSW of Masally, 80 –140 ma.s.l.,  Quercus, Carpinus, Aceretc. 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 ma.s.l.,  Quercus, Fagusetc. forest along stream, litter, 3.V.1987, leg. S. Golovatch and K. Eskov; 3 females, 8 juv. (ZMUM), Zakataly Nature Reserve, Katekhchay Valley, 700 ma.s.l., 23.V.1981, leg. S. Golovatch and J. Martens; 6 females, 4 juv. (ZMUM), Zakataly Nature Reserve, 1000 ma.s.l., forest, litter, 13.VIII.1989; 1 female, 2 juv. (ZMUM), Zakataly Nature Reserve, Agkemal, 2200 ma.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°,  Fagusforest, 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 kmS of Kirovabad (= Ganja), 1500 –1900 ma.s.l.,  Fagus, Carpinus, Quercusetc. forest, litter, logs and under stones, 3 –4.V.1983, leg. S. Golovatch; 2 males, 1 female, 2 juv. (ZMUM), ca 15 kmWSW of Mardakert, 1100 ma.s.l.,  Quercus, Fagus, Aceretc. forest, litter, 2.VI.1987; 1 male, 5 females (ZMUM), NW above Bash-Layski ca 20 kmNNW of Sheki, 1250 ma.s.l.,  Fagus, Carpinus, Aceretc. 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,  Quercusforest, 1900 ma.s.l., litter, 22.IV.1983, leg. S. Golovatch; 8 males, 7 females (ZMUM), Yarymdja SE of Altyagach, 1320 –1350 ma.s.l.,  Fagus, Quercus, Carpinusetc. forest, litter and under bark, 20, 24 –25.IV.1987; 1 female (ZMUM), Altyagach, 1050 –1100 ma.s.l.,  Quercus, Fagus, Carpinusetc. 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 ma.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 ma.s.l.,  Quercus, Acer, Taxusetc. 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 ma.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 ma.s.l.,  Quercus, Carpinus, Aceretc. sparse forest, litter, rotten wood and under stones, 9.X.1983; 1 juv. (ZMUM), Talysh Mts., Lerik Distr., Zuvand, Gilidara, 1800 ma.s.l., semi-desert, 8.X.1983, all leg. S. Golovatch; 1 female (ZMUM), Talysh Mts., Zuvand, Djoni, 1500 ma.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 kmSW of Alekseevka,  Quercus, Acer, Carpinus, Parrotiaetc. forest, litter, under stones and bark, 14 and 16.X.1983; 2 juv. (ZMUM), Lenkoran Distr., Hyrcan Nature Reserve, Alekseevka, 50 ma.s.l.,  Quercus, Parrotia, Carpinusetc. 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 ma.s.l., 26.III.2015; 1 male, 2 females (HR), Hyrcan Nature Reserve, road Lenkoran —Lerik at km 32, N38.7638º, E48.5819º, 400 ma.s.l., 26.III.2015; 2 males (HR), Hyrcan Nature Reserve, Günəşli, 2 road-km to Lerik, N38.7975º, E48.4644º, 710 ma.s.l., 26.III.2015; 2 males, 4 females (HR), Hyrcan Nature Reserve, SW of Zünqüləş, N38.4493º, E48.7623º, 60 ma.s.l., 27.III.2015; 1 male, 2 females (HR), Hyrcan Nature Reserve, SW of Zünqüləş, N38.448º, E48.7597º, 130 ma.s.l., 27.III.2015; 4 females (HR), Hyrcan Nature Reserve, Daştatük, 1.3 road-km Xanbulan Reservoir, N38.6747º, E48.7622º, 110 ma.s.l., N38.6747º, E48.7622º, 27.III.2015; 1 female (HR), Hyrcan Nature Reserve, SW of Aşağı Apu, N38.6726º, E48.7362º, 180 ma.s.l., 27.III.2015; 1 female (HR), Qurbanəfəndi 1.3 road-km to Xanəgah, N40.8614º, E48.1132º, 640 ma.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  Quercusbush, 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 eand 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  Brachydesmusoccurring in the Caucasus and adjacent parts of E Turkeyand 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. ferrugineusand  B. talyschanusas new species, had only very few samples in their hands. In contrast to Lignau’s (1915) material which came from relatively small Abkhaziaalone, Lohmander’s (1936) stemmed from various, often widely separated places scattered across the entire Caucasus. More recently, Ceuca (1975)reported  B. ferrugineusfrom 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 Turkeyand 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. kalischewskyifrom two localities in E Turkey.   B. kalischewskyi ! # # " % " & ’ (") * ++, + " % ­) * ++, + " & ’ (+ +. + ’ + " 0 ­ +,. + + "))1 & ’ ( 3 3) + +. +, + " & ’ ( " ") * ++,. + + " & ’ 0( % %) * + +., + " & ’ ( #. #) * / 6 *) 7 " ­ 0 e i7 ) 8 7) l i) 7 " 3 0 e7) i) ) 8 l7) )1)1 " 3 0 % e i7 ) 8 7) l7  i) , % e7) i) l7) " ­ 0 e i7 7) l )1 ­ e i7 7  i7) l 9) 7 Golovatch (1981b)recorded  B. ferrugineusin the East  AzerbaijanProvinceof Iran, reiterated by Enghoff and Moravvej (2005). That sole Iranian sample has again been borrowed for the present study.   FIGURE 19.Morph E of  Brachydesmus kalischewskyiLignau, 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 kalischewskyiLignau, 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. Since our collections are indeed very large and densely blanket the entire region in question, we believe all those “species” actually represent one,  B. kalischewskyiby priority. We shall try to bring enough evidence to support this conclusion, based on numerous measurements taken and abundant illustrations provided. Treating the  kalischewskyicomplex 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  kalischewskyiwill be referred to as A, while the morphs  karawajewi,  ferrugineusand  talyschanusas B, C and D, respectively. Neither the variety batumensis, proposed by Lohmander (1936)as an Ajarian form of  kalischewskyi, nor  bidentatusare 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, Nakhichevanand 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, KrasnodarProvince ( Fig. 23),  Azerbaijanor Armenia( Fig. 24).   FIGURE 21.Morph F of  Brachydesmus kalischewskyiLignau, 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 kalischewskyiLignau, 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 kalischewskyiLignau, 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 kalischewskyiLignau, 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. 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 Adygeadoes morph B of  B. kalischewskyi, represented there by smaller individuals about 9 –11 mmin 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 26and 10A –10D), the males also by their disparate gonopod structure (cf. Figs 27and 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  kalischewskyicomplex 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. kalischewskyias we treat it could become a model species for further evolutionary studies.   FIGURE 25.Examples of gonopods of  Brachydesmus kalischewskyiLignau, 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 kalischewskyiLignau, 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 kalischewskyiLignau, 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). Map 2 roughly depicts the distribution of  B. kalischewskyiin the Caucasus and adjacent parts of E Turkeyand 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  kalischewskyicomplex 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 kmN 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  kalischewskyicomplex is also possible in the future. At least some species names like  karawajewi,  ferrugineusor  talyschanusare long available from a taxonomic point of view. Polymorphism and distrubution mosaics like the ones demonstrated by  B. kalischewskyihave 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  Epanerchodusin Japan(e.g. Murakami 1970; Murakami and Kawasawa 1975). In contrast, the widespread East Asian  E. orientalisAttems, 1901shows only a gradual change in gonopod structure from north to south along Japan( Nishikawa and Murakami 1993), but further southerly, in Taiwanand 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.