Colobognatha millipedes in the Caucasus (Diplopoda: Polyzoniida, Platydesmida, Siphonocryptida) Author Golovatch, Sergei Author Evsyukov, Aleksandr Author Reip, Hans S. text Zootaxa 2015 3972 2 250 266 journal article 10.11646/zootaxa.3972.2.6 eee2db07-ee84-4161-87af-54c590a748cb 1175-5326 233469 8D47A1B5-5AB3-4570-BC70-4CD81B340A87 Hirudisoma roseum ( Victor, 1839 ) Figs 1–3 , 9 Leiosoma rosea (recte: roseum ) Victor, 1839 : 46 (D). Polyzonium roseum : Lignau, 1907 : 199 (R); Muralewicz, 1911 : 3 , 10 (M). Leiosoma roseum : Lignau, 1911 : 54 (D, R); 1915: 389 (D, R); Lohmander, 1936 : 170 (M), 1939: 144 (D); Kobakhidze, 1965 : 395 (R); Ghilarov, 1965 : 143 , 144 (R). Leiosoma ponticum Lohmander, 1939 : 142 (D), syn. n. Leiosoma ( Polyzonium ) roseum : Lang, 1959 : 1792 (M). Hirudisoma roseum : Lokšina and Golovatch, 1979 : 382 (M); Talikadze, 1984 : 143 (R). Hirudisoma ponticum : Enghoff, 2006 : 178 (M). Material examined . Neotype (here designated): male ( ZMUM ρ2634), Georgia , Lagodekhi Nature Reserve, 600– 700 m a.s.l., Fagus , Fraxinus , Acer etc. forest., litter, under bark & stones, 5–6.V.1983 , leg. S. Golovatch. 4 females , 4 juveniles ( ZMUM ρ2591), RUSSIA , Krasnodar Prov., near Gelendjik, 28.IV.1984 , leg. A. Viktorov; 2 females ( ZMUM ρ2592), Krasnodar Prov., Goryachiy Klyuch, 10 km S of Fanagorskoye, near Cave Fanagoriyskaya, mixed Fagus , Quercus , Picea etc. forest, litter, under stones & in rotten logs, 30.X.1981 ; 3 females ( ZMUM ρ2593), same locality, Fagus , Acer etc. forest, litter & logs, 19.V.1983 ; 1 female ( ZMUM ρ2594), Krasnodar Prov., Severskaya Distr., 2–10 km S of Ubinskaya, Quercus , Fagus , Carpinus etc. forest, 300–450 m a.s.l., litter & under bark, 3–4.VII.1986 ; 1 male ( ZMUM ρ2595), Krasnodar Prov., Sochi, Dagomys, 250 m a.s.l., Quercus , Carpinus , Fagus etc. shrub, litter, logs, 18.V.1983 , all leg. S. Golovatch; 2 males ( ZMUM ρ2596), environs of Sochi, Krasnaya Polyana, 6–8.Vl.1978, leg, V. Dolin; 4 females ( ZMUM ρ2597), Krasnodar Prov., 5.5 km NE of Krasnaya Polyana, lower course of Achipse River, ca 600 m a.s.l., N 43 43' 10", E 40 15' 25", mixed deciduous forest, 19–23.VIII.2014 , leg. K. Makarov & A. Matalin; 1 male , 1 female ( ZMUC ), Krasnodar Prov., ca 4 km NW of Krasnaya Polyana, SE slopes of Mt Achishko, N43°42.516’ , E40°10.534’ , 1000 m a.s.l., Castanea & Fagus forest, sifted litter, 19.VII.2011 , leg. A. Solodovnikov & M. Kaae; 1 male ( ZMUC ), same locality, N43°42’23” , E40°09’41” , 1150 m a.s.l., Fagus & Rhododendron forest, sifted litter, 19.VII.2011 , leg. A. Solodovnikov, 1 male ( ZMUM ρ2598), Krasnodar Prov., Caucasian Nature Reserve, near Pslukh, ca 20 km E of Krasnaya Polyana, Mt Kogot, ca 1000 m a.s.l., Fagus & Abies forest, litter, under bark & stones, 18–20.V.1985 ; 4 females ( ZMUM ρ2599), same locality, Fagus , Quercus , Abies , Picea etc. forest, 1000 m , litter, 11–12.VIII.1986 ; 1 female ( ZMUM ρ2600), same locality, 1400 m , litter, under bark & stones 18–20.V.1985 ; 5 males , 15 females , 2 juveniles ( ZMUM ρ2601), Krasnodar Prov., Caucasian Nature Reserve, Pasture Abago near Guzeripl, 1100 m a.s.l., Abies & Fagus forest, Rhododendron thicket, litter, under bark & stones, 24–26.V.1985 ; 1 male , 4 females ( ZMUM ρ2602), same locality, Abies & Fagus forest, 1350–1400 m , 24–26.V.1985 ; 5 males , 17 females , 6 juveniles ( ZMUC ), Krasnodar Prov., ca 9 km SW of Mt Fisht, Babuk-Aul, N43°53’26” , E39°49’11” , 560 m a.s.l., Castanea & Fagus forest, sifted leaf litter, 11.VII.2011 ; 2 females , 5 juveniles ( ZMUC ), same locality, N43°54.214’ , E39°50.507’ , 1200 m a.s.l., Fagus & Rhododendron forest, sifted leaf litter, 14.VII.2011 , all leg. A. Solodovnikov; 1 female (HR), Krasnodar Prov., Urushten River valley: right side, 1 km upstream of bridge at Kordon Chernorechye, Acer – Fagus forest with Sambucus , Corylus , Alnus , Ulmus in understorey, N43.9319 E40.6778 , 850 m a.s.l., 18.IX.2012 ; 18.IX.2012 ; 1 male (AE), Krasnodar Prov., near Mezmai, Zauda Mt., under stones & barks, 23.VII.2011 , leg. D. Khisametdinova & Yu. Kotchetov; 1 male , 1 female (AE), near Mezmai, under stones & barks, 23.VII.2011 , leg. D. Khisametdinova & Yu. Kotchetov; 1 female (HR), Adygea, Guzeripl: Molchepa riverside 6 km SE (right tributary of Belaya River), N41.6494 E41.7731 , 730 m a.s.l., 01.X.2012 , all leg. F. Walther; 24 males , 24 females , 4 juveniles (AE), Adygea, Kamyshovaya Polyana, under stones, 24.VI.2011 , leg. D. Khisametdinova & S. Kurshakov; 1 female (AE), Adygea, near Nikel, rotten logs, 10.V.2013 , leg. D. Khisametdinova & Yu. Kotchetov; 1 female ( ZMUM ρ2603), Karachaevo-Cherkessia, ca 30 km S of Kurdjinovo, 4 km N of Damkhurts, Laba River Valley, 1050–1100 m a.s.l., Fagus , Acer , Picea etc. forest, litter & bark, 4.VIII.1986 ; 1 female ( ZMUM ρ2604), Kabardino-Balkaria, Chegem Distr., 5 km S of Upper Chegem, 1700 m a.s.l., Betula , Pinus & Juniperus forest, litter, 12.VII.1986 ; 3 females ( ZMUM ρ2605), North Ossetia, S of Ordzhonikidze (now Vladikavkaz), between Chmi & Baltik, Quercus & Alnus on slope, litter & under stones, 2.VI.1982 , all leg. S. Golovatch; 1 male , 1 female (AE), North Ossetia, near Dzinaga, Urukh Canyon, under stones & barks, 12.VIII.2010 , leg. A Evsyukov & D. Khisametdinova. 2 females ( ZMUM ρ2610), ABKHAZIA , Gumista Nature Reserve, forest litter, 6.VI.1982 , leg. J. Boháč; 2 males , 4 females ( ZMUM ρ2606), Sukhumi Distr., Lake Amtkel ca. 16 km N of Tsebelda, 550 m a.s.l., Alnus forest, litter & under bark, 19.VIII.1986 ; 2 females ( ZMUM ρ2607), Lake Ritsa, 950–1100 m , Fagus , Abies , Picea , Acer etc. forest, litter, under bark & stones, 13–14.VIII.1986 ; 4 males ( ZMUM ρ2608, ρ2609), Sukhumi Distr., Bzyb River valley, Pskhu, 700–950 m , Fagus , Quercus , Castanea etc. forest, litter, under bark & stones, 15– 16.VIII.1986 , all leg. S. Golovatch; 2 males ( ZMUM ρ2611), near Shroma, under bark, 1.V.1971 , leg. N. Zalesskaja. 1 juvenile ( ZMUM ρ2612), GEORGIA , Ajaria, E of Kobuleti, 3 km SE of Chakhati, deciduous forest near spring, litter & under stones, 14.X.1981 ; 1 male ( ZMUM ρ2613), Ajaria, Zelenyi Mys, Batumi Botanical Garden, 13.X.1978 , all leg. S. Golovatch; 3 males , 5 females (ZMN), same locality, Betula & Rhododendron forest, 8– 10.IV.1988 , leg. D. Logunov; 1 male , 1 female , 1 juvenile ( ZMUM ρ2614), same locality, 10.X.1981 , leg. J. Boháč; 8 males , 10 females ( ZMUM ρ2615), same locality, 20–150 m , 30.V.–7.VI.1981 ; 1 male , 2 females ( ZMUM ρ2616), Ajaria, between Tsikhisdziri & Batumi, gardens, 100 m a.s.l., 30.V.1981 , all leg. S. Golovatch & J. Martens; 3 females ( ZMUM ρ2617), Ajaria, Kintrish Nature Reserve, Khekpara River Canyon, 12.X.1984 ; 3 females ( ZMUM ρ2618), same locality, 2300 m a.s.l., X.1984 , all leg. E. Kvavadze; 2 males , 1 female ( ZMUM ρ2619), Kintrish Nature Reserve, Zeraboseli, Rhododendron thicket, 600–800 m a.s.l., 2.VI.1981 , leg. S. Golovatch & J. Martens; 3 males , 3 females , 7 juveniles ( ZMUM ρ2620, ρ2621), same locality, Zeraboseli, 800 m a.s.l., Rhododendron thicket, litter, 13.X.1981 ; 3 males , 4 females ( ZMUM ρ2622, ρ2623), same locality, Zeraboseli, 450–600 m , deciduous forest, litter & under stones, 13.X.1981 , all leg. S. Golovatch; 2 females ( ZMUM ρ2624), Ajaria, 10 km E of Chakva, near Chakvistavi, litter, 8.V.1987 , leg. S. Kurbatov; 1 juvenile (inadvertently lost), Ajaria, near Keda, Picea , Quercus , Fagus etc. forest, litter, X.1975 , leg. A. Druk; 1 female ( ZMUM ρ2625), Kutaissi Distr., environs of Tskhaltubo, Sataplia Nature Reserve, near entrance to Cave Sataplia I, litter, 27.I.1984 , leg. V. Dushenkov; 1 female ( ZMUM ρ2626), Sataplia Nature Reserve, forest litter, 27.I.1984 leg. K. Makarov; 1 female ( ZMUM ρ2627), same locality, forest litter & under stones, 25.X.1981 , leg. S. Golovatch; 2 females ( ZMUM ρ2628), same locality, IV.1973 , leg. A. Krivolutsky; 4 males , 4 females ( ZMUM ρ2629), Kutaissi, old Pinus platations on slope, litter, 1.II.1987 , leg. K. Makarov; 2 males , 2 females ( ZMUM ρ2630), Svanetia, Nenskra River Canyon, Luki N of Khaishi, ca 800 m a.s.l., forest litter, 2.IX.1986 , leg. A. Ryvkin; 2 males , 2 females (HR), Samegrelo-Zemo Svaneti, Djvari: road Zugdidi–Mestia at km 33/165, N42.7514 E42.0472 , 470 m a.s.l., 26.IX.2012 , leg. F. Walther; 14 males , 13 females , 6 juveniles ( ZMUM ρ2631), Georgia , Surami Mt Ridge, Pass Djvari between Gomi & Sachkhere, Alnus , Fagus , Rhododendron etc. forest, 850 m a.s.l., litter, 7.V.1987 , leg. S. Golovatch & K. Eskov; 1 male , 4 females ( ZMUM ρ2632), Georgia , environs of Mtskheta, 10 km N of Djvari, 800 m a.s.l., Buxus , Fagus , Picea, Taxus etc. forest, litter, 20–21.VIII.1986 , leg. S. Golovatch; 2 females ( ZMUM ρ2633), Georgia , Mukhura ca 15 km E of Tkibuli, 700–800 m a.s.l., Castanea , Fagus , Carpinus etc. forest, litter, under bark & stones, 7–9.V.1987 , leg. S. Golovatch & K. Eskov; 1 female (HR), Racha- Lechkhumi and Kvemo Svaneti, Isunderi, road Tsageri–Tskhaltubo, from junction to Isunderi 4.3 rkm towards Tskhaltubo, N42.5119 E42.6372 , 320 m a.s.l., 23.IX.2011 , leg. F. Walther; 1 male , 4 females ( ZMUM ρ2635, ρ2636), Lagodekhi Nature Reserve, 600–700 m a.s.l., Fagus , Fraxinus , Acer etc. forest., litter, under bark & stones, 5–6.V.1983 , leg. S. Golovatch. 1 male ( ZMUM ρ2589), AZERBAIJAN , Zakatali Nature Reserve, 700 m Katekh-Chay River valley, 23.V.1981 , leg. S. Golovatch & J. Martens; 1 female ( ZMUM ρ2590), NW above Bash-Layski, ca 20 km NWW of Sheki, 1250 m a.s.l., Fagus , Carpinus , Acer etc. forest, litter. 5.V.1987 , leg. S. Golovatch & K. Eskov. Descriptive notes. Length of adults 6.0–13.0 mm, width 1.6–2.7 mm , with 26–49 podous + 3–2 apodous segments + telson, regardless of sex. Neotype male (from Lagodekhi Nature Reserve, NE Georgia ) with 33p + 2a + T, length ca 7.0 mm, maximum width 1.6 mm . Length to width ratio ≤ 6:1. Live coloration often pink, alcohol material usually light yellow-brown with a very broad yellowish axial stripe and lighter paraterga ( Figs 1 , 2 ). Only antennae darker, usually rusty to purplish brown. Body hirudiform, very clearly flattened dorsoventrally, strongly, but regularly broadened until segment 6 or 7, strongly, but regularly narrowing towards telson on last 6 or 7 segments. Head small, about half as broad as collum, subtriangular, only slightly longer than broad, rostrum subtruncate and short. Ocelli black, convex, 2+ 2 in smaller/younger specimens or 3+ 3 in larger/older samples, on each side of head arranged in a nearly longitudinal line ( Fig. 2 B, D), each ocellus with a long seta at base ( Fig. 2 A). Antennae short, erect, rod-shaped, only very slightly clavate; antennomeres 2–6 subequal in length, 6th thickest. Collum biconvex, clearly transverse, paraterga almost sharp. Following paraterga mostly subrectangular and narrowly rounded, each with a narrow, slightly sinuate, lateral peritreme delimited by a weak sulcus only dorsally. Tegument smooth, shining, glabrous and translucent. Caudal margin of metaterga very slightly elevated. Ozopores entirely dorsal, small, inconspicuous, lying lateral to peritremal sulcus at about 2/3 of paratergal length, starting from segment 5. Telson coniform, completely concealed from above by plate-like and medially fused last paraterga ( Figs 1 , 2 A–C). Sterna large, but coxae almost in touch medially ( Fig. 1 B, 2B, D). Legs 6-segmented. Male coxae 1 and 2 each with a ventral brush of short, but stiff setae ( Fig. 3 A, B). Male coxa 2 with a short, tube-shaped, simple, basal, caudomediad directed gonapophysis ( g ) ( Fig. 3 B). Claws often, but not always phylloid, devoid of accessory structures ( Fig. 3 A, B). Coxal sacks starting from leg 3. Both anterior ( a ) and posterior ( p ) gonopods ( Fig. 3 C–E) 5-segmented. Anterior ones incrassate and stout, tarsus with a short, acuminate, tooth-shaped tip forming a caudomedial gutter for accommodation of a particularly long, subflagelliform, distally serrate posterior gonopod tarsus. Remarks. This species was originally described from “Natlim-Zéméli en Kahétie” by Victor I. Motschoulsky under his pseudonym T. Victor (Victor 1839) . Eventually, this locality represents a slightly misspelled “Natlismtsemeli”, in Georgian meaning “St. John’s”. There are numerous localities across Georgia named “Natlismtsemeli”. In Kakhetia alone, which is the eastern part of Georgia , at least four such localities can easily be traced (https: // maps.google.com/). It is thus impossible to state where exactly Motschoulsky collected the type or types except that it or they came from eastern Georgia . Since the type material of H. roseum is absent from the Motschoulsky Collection, currently kept in ZMUM ( Golovatch 2014 ), a neotype designation is required from among the near-topotypes stemming from Kakhetia. This is especially necessary so as to be able to verify the species’ identity versus the samples described from Abkhazia by Lignau (1911) . Thus, Lohmander (1936) noted the present uncertainty by referring to this species as “ Leiosoma roseum (Victor) Lignau ”. Since the name Leiosoma Victor, 1839 , established for L. roseum by monotypy, is preoccupied by Leiosoma Stephens, 1829 (Coleoptera) ( Jeekel 1971 ) , all subsequent mentions of L. roseum (e.g. Lignau 1911 ; Lohmander 1936 ; Kobakhidze 1965 ) are incorrect. Brandt (1840) and Latzel (1884) listed Leiosoma Victor, 1839 as a questionable synonym of Polyzonium Brandt, 1837 ; hence a few references to Polyzonium roseum or Leiosoma ( Polyzonium ) roseum . Gervais (1847) listed L. roseum among the junior synonyms of the common European Polyzonium germanicum Brandt, 1837 . In contrast, Lignau (1911 , 1915 ) believed that L. roseum was a senior subjective synonym of both Heterozonium hirsutum Verhoeff, 1901 and Polyzonium pallidum (Fanzago, 1875) . The former species had been quite poorly described from near Istanbul, Turkey ( Verhoeff 1901 ), now also known in Greece and Bulgaria ( Enghoff 2006 ; Kime and Enghoff 2011 ). The latter species had been beautifully redescribed from Italy by Berlese (1883) . Lohmander (1939) rejected Lignau’s synonymies and described the second Leiosoma species, from northeastern Turkey : L. ponticum Lohmander, 1939 . In addition, he emphasized the synonymy of Heterozonium Verhoeff, 1901 under Hirudisoma Fanzago, 1881 , which Verhoeff (1941) also accepted. The use of Leiosoma Victor, 1839 , being banned since Jeekel (1971) , the oldest available generic name to incorporate L. roseum and L. ponticum is Hirudisoma . Regrettably, neither has been considered in the latest review of Hirudisoma (see Mauriès 1964 ). Lokšina & Golovatch (1979) and Enghoff (2006) seem to have been the first to transfer L. roseum and L. ponticum to Hirudisoma , respectively. FIGURE 1. Hirudisoma roseum (Victor, 1839) , a large-sized female from near Krasnaya Polyana, Russia. A, B: habitus, dorsal and ventral views, respectively. Pictures by K. Makarov, taken not to scale. FIGURE 2. Hirudisoma roseum (Victor, 1839) , habituses of two medium-sized females from Batumi Botanical Garden, Ajaria, Georgia (A–C) and anterior part of body of a male from near Mezmai (D), lateral, dorsal, dorsal and ventral views, respectively. Pictures A–C by K. Makarov, picture D by A. Evsyukov, taken not to scale. FIGURE 3. Hirudisoma roseum (Victor, 1839) , male from Pskhu, Abkhazia (1, 2), male from Kinstrish Nature Reserve, Ajaria, Georgia (C, D) and male neotype from Lagodekhi Nature Researve, E Georgia (D). A, B: legs 1 and 2, respectively, caudal views. C: left anterior gonopod, anterior view. D, E: both left and right gonopods together, respectively, caudal views. Scale bar: 0.1 mm. Designations: g , gonapophysis; a , anterior gonopod; p , posterior gonopod. If Lohmander (1939) was definitely right when rejecting the synonymy of roseum with hirsutum , at least because the latter species is hirsute as explicitly implied by its name, his creation of ponticum seems to have been ill-founded. Thus, the characters used for the separation of ponticum from roseum were stated to lie in the former’s more strongly rounded paraterga, simple (non-phylloid) claws, shorter antennae, as well as a lower inner lobe of the coxa and a slightly differently shaped tarsus of the anterior gonopod. Apparently, because Lohmander based his ponticum on a single (and incomplete) male holotype , whereas our abundant material from the Caucasus shows all possible variations and transitions in the above features, we are inclined to formally consider H. ponticum as a junior subjective synonym of H. roseum , syn. nov.