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250
Accepted by W. Shear: 4 May 2015; published: 11 Jun. 2015
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
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Copyright © 2015 Magnolia Press
Zootaxa 3972 (2): 250
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Article
http://dx.doi.org/10.11646/zootaxa.3972.2.6
http://zoobank.org/urn:lsid:zoobank.org:pub:8D47A1B5-5AB3-4570-BC70-4CD81B340A87
Colobognatha millipedes in the Caucasus (Diplopoda: Polyzoniida,
Platydesmida, Siphonocryptida)
SERGEI GOLOVATCH
1,4
, ALEKSANDR EVSYUKOV
2
& HANS S. REIP
3
1
Institute for Problems of Ecology and Evolution, Russian Academy of Science, Leninsky pr. 33, Moscow 119071, Russia
2
Lyceum No. 1 “Classical”, Balakireva Str. 32, Rostov-on-Don 344004, Russia
3
Senckenberg Museum of Natural History, Görlitz, Am Museum 1, D-02826 Görlitz, Germany
4
Corresponding author: E-mail: sgolovatch@yandex.ru
Abstract
The superorder Colobognatha is represented in the Caucasus by three genera and species, one each in the orders Polyzoni-
ida, Platydesmida and Siphonocryptida. Hirudisoma roseum (Victor, 1839) (Hirudisomatidae, Polyzoniida) is especially
widespread, ranging from S Russia, Abkhazia, Georgia (with a neotype designated and described from Kakhetia, E Geor-
gia) and NW Azerbaijan to E Turkey, and also including H. ponticum (Lohmander, 1939) (junior subjective synonym, syn.
nov.). Fioria hyrcana Golovatch, 1980 (Andrognathidae, Platydesmida) is endemic to the Hyrcanian parts of the Republic
of Azerbaijan and NW Iran along the western and southern coasts of the Caspian Sea. Due to the finding of Hirudicryptus
abchasicus sp. nov. (Siphonocryptidae, Siphonocryptida), from a single locality in Abkhazia, NW Caucasus, the order
Siphonocryptida is new to the fauna of the region. A key to all four species of the trans-Palaearctic genus Hirudicryptus
is given. All three Caucasian species of Colobognatha are described in due detail and abundantly illustrated, and their dis-
tributions mapped.
Key words: Polyzoniida, Platydesmida, Siphonocryptida, taxonomy, new species, new synonymy, neotype, key, distribu-
tion, map, Russia, Abkhazia, Georgia, Azerbaijan, Turkey, Iran
Introduction
The diplopod superorder Colobognatha Brandt, 1833 is known to comprise four small orders: Polyzoniida,
Platydesmida, Siphonocryptida and Siphonophorida (e.g. Shear 2011). Only the former two orders contain not only
tropical or subtropical, but also temperate species. Moreover, one of the species of the boreal genus Angarozonium
Shelley, 1997 (Polyzoniidae, Polyzoniida) is especially widespread across Siberia and represents perhaps the
northernmost range limit for the whole class Diplopoda (Mikhaljova 2004, Shelley and Golovatch 2011).
Only two genera and species of Colobognatha have hitherto been reported from the Caucasus and adjacent
parts of NW Iran: Hirudisoma roseum (Victor, 1839) (Hirudisomatidae, Polyzoniida) and Fioria hyrcana
Golovatch, 1980 (Andrognathidae, Platydesmida). The present paper provides their richly illustrated redescriptions
and numerous new faunistic records, also summarizing and updating the nomenclatural history and distributions of
both these species. In addition, the relict order Siphonocryptida is found in the region for the first time, being
represented there by a new species.
Material and methods
A large material of both Hirudisoma roseum and Fioria hyrcana has been amassed by the first author, now mostly
housed in the Zoological Museum of the Moscow State University (ZMUM), Moscow, Russia. A few additional
samples have been accumulated by the second and third authors, now retained in their private collections in
Rostov-on-Don and Jena, referred to below as (AE) and (HR), respectively. One sample belongs to the Institute of
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COLOBOGNATHA MILLIPEDES IN THE CAUCASUS
Animal Systematics and Ecology of the Siberian Branch of the Russian Academy of Sciences, Novosibirsk (ZMN),
a few more to the Natural History Museum of Denmark, Copenhagen (ZMUC, det. H. Enghoff) and the Biology
Department of the University of Belgrade, Serbia (UBS, det. D. Antić). Unexpectedly, a small sample of a third
species has also been found.
In the catalogue sections, D stands for a description or descriptive notes, R for new records, while M for a mere
mention.
Taxonomic part
Order Polyzoniida
Family Hirudisomatidae
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
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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, Tax u s 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.
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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. 2B, D), each ocellus
with a long seta at base (Fig. 2A). Antennae short, erect, rod-shaped, only very slightly clavate; antennomeres 2–6
subequal in length, 6
th
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, 2A–C).
Sterna large, but coxae almost in touch medially (Fig. 1B, 2B, D). Legs 6-segmented. Male coxae 1 and 2 each
with a ventral brush of short, but stiff setae (Fig. 3A, B). Male coxa 2 with a short, tube-shaped, simple, basal,
caudomediad directed gonapophysis (g) (Fig. 3B). Claws often, but not always phylloid, devoid of accessory
structures (Fig. 3A, B). Coxal sacks starting from leg 3.
Both anterior (a) and posterior (p) gonopods (Fig. 3C–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.
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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.
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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.
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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.
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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.
Order Platydesmida
Family Andrognathidae
Fioria hyrcana Golovatch, 1980
Figs 4, 5, 9
Fioria sp.: Lokšina & Golovatch, 1979: 382 (M).
Fioria hyrcana Golovatch, 1980: 459 (D).
Fioria hyrcana: Golovatch, 1981: 425 (R); Bababekova, 1996: 90 (M); Enghoff & Moravvej, 2005: 62 (M).
Material examined. 1 female (ZMUM ρ2637), AZERBAIJAN, Istisu ca 8 km SW of Masally, Quercus, Acer,
Carpinus etc., 80–140 m a.s.l., litter, under bark & stones, 19–20.X.1983; 1 female (ZMUM ρ2638), Lenkoran
Distr., Apo ca 8 km SW of Alekseevka, Hyrcan Nature Reserve, Quercus, Acer, Carpinus, Parrotia etc. forest,
250–370 m a.s.l., litter, rotten wood & under stones, 14–16.X.1983; 9 males, 4 females, 2 juveniles (ZMUM
ρ2639, ρ2640), Lenkoran Distr., Alekseevka, Hyrcan Nature Reserve, 50 m a.s.l., Quercus, Parrotia, Carpinus etc.
forest, litter, rotten wood, 13.X.1983, all leg. S. Golovatch; 1 female (ZMUM ρ2641), same locality, litter, 26.I.–
4.II.1985, leg. A. Druk; 1 male, 1 juvenile (HR), Lerik Distr., Hyrcan Nature Reserve, Piran, northwestern town
exit, N38.7340, E48.6441, 230 m a.s.l., pasture with single trees (Parrotia, Zelkova, Crataegus), under logs,
26.III.2015; 1 male, 2 females (HR), one female (UBS), Lerik Distr., Hyrcan Nature Reserve, road Lǝnkǝran–Lerik
at road-km 32, N38.7638, E48.5819, 400 m a.s.l., small side valley, forest of Parrotia trees with some Quercus, in
thick leaf litter, 26.III.2015; 4 males, 1 female, 1 juvenile (HR), one male, one female and one juvenile (UBS),
Astara Distr., Hyrcan Nature Reserve, SW of Zünqüləş, N38.4480, E48.7597, 130 m a.s.l., end of small valley,
steep slope, Parrotia, Quercus, Acer trees, in leaves, under tree trunks, 27.III.2015; 2 females (HR), Astara Distr.,
Hyrcan Nature Reserve, W of Zünqüləş, N38.4575, E48.7477, 50 m a.s.l., open valley with single Parrotia &
Populus trees, meadow, under logs, 27.III.2015; 3 males, 4 females, 1 juvenile (HR), Lǝnkǝran Distr., Hyrcan
Nature Reserve, SW of Aşağı Apu, N38.6726, E48.7362, 180 m a.s.l., Quercus forest, in leaves and rotten wood,
27.III.2015, all leg. H. Reip, D. Antić & F. Walther.
Descriptive notes. Length of adults 4.5–10.0 mm, width.0.8–1.0 mm, with 23–43 podous + 3–1 apodous
segments + telson, regardless of sex. Length to width ratio (except for smallest juveniles) > 8:1.
Live coloration uniformly red-brown to yellow, alcohol material usually somewhat faded (Fig. 4). Body only
moderately flattened dorsoventrally, vermiform, regularly and modestly broadened until segment 4, similarly
regularly and modestly narrowing towards telson on last 3 segments. Head small, subtriangular, only slightly
narrower than collum and longer than broad, labrum broadly and regularly rounded, without a rostrum (Fig. 4A, C).
Ocelli absent. Antennae short, C-shaped, strongly clavate; antennomere 6 largest (Figs 4A, 5A).
Collum ovoid, paraterga small, broadly rounded, dorsal surface with two transverse rows of 2+2 large, rounded
tubercles. Following paraterga much larger, mostly set at about half of body height, broadly rounded and obtuse-
angled anteriorly, more narrowly rounded and acute-angled posteriorly, somewhat drawn behind tergal margin only
in penultimate segment (Fig. 4). Each metatergum likewise clearly domed and with two transverse rows of 2+2
(segments 2–4) or 3+3 (all following segments before telson) large, rounded tubercles. Tegument extremely
densely micro-hirsute and dull all over. Ozopores lateral, small, inconspicuous, lying at bottom of a small incision
at about half of paratergal length, starting with segment 5. Telson coniform, completely exposed in dorsal view
(Fig. 4A, F).
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Sterna large, but coxae almost in touch medially, mostly divided by a small, but evident, elongated elevation
(Fig. 4E). Legs 6-segmented. Male coxa 2 with a short, tube-shaped, simple, basal gonapophysis directed
caudomesad. Claws usually simple, only a few anteriormost ones each with a minute accessory claw at base. Coxal
sacks starting with leg 3.
FIGURE 4. Fioria hyrcana Golovatch, 1980), female (A) and male (B–F) topotypes. A: habitus, lateral view. B, C: anterior
part of body, dorsal and ventral views, respectively. D, E: middle part of body, dorsal and ventral views, respectively. F:
posterior part of body, dorsal view. Pictures by K. Makarov, taken not to scale.
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FIGURE 5. Fioria hyrcana Golovatch, 1980, male holotype. A: antenna, lateral view. B, C: anterior gonopods, anterior and
posterior views, respectively. D: left posterior gonopod, anterior view. Scale bars: A, 0.3 mm; B–D, 0.1 mm. After Golovatch
(1980).
Anterior gonopods (Fig. 5B, C) plate-shaped, 6-segmented, strongly incrassate and stout. Posterior gonopods
(Fig. 5D) 7-segmented, much slenderer, tarsus longest, with a long, midway, spiniform filament and a slightly
expanded tip.
Remarks. This species, originally described from the Hyrcan Nature Reserve in SE Azerbaijan (Golovatch
1980), seems to be endemic to Hyrcania within both the Republic of Azerbaijan’s southeastern part and
northwestern Iran (Golovatch 1980, 1981).
Order Siphonocryptida
Family Siphonocryptidae
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Hirudicryptus abchasicus sp. nov.
Figs 6–10
Material examined. Holotype male (ZMUM ρ2583), ABKHAZIA, Gumista Nature Reserve, forest litter,
4.VI.1982, leg. J. Boháč. Paratype female (ZMUM ρ2584), same data, together with holotype.
Name: To emphasize Abkhazia, the terra typica; adjective.
Diagnosis: Distinguished from all congeners but H. canariensis (Loksa, 1967) by the much shorter antennae
which are only about as long as collum width, as well as by the smaller and much shorter collum which only forms
a quite inconspicuous casque-like shield to cover the head. Differs from H. canariensis by the concave last tergum
and less numerous, 9–11 + 9–11 vs 14–15 + 14–15, tubercles at the caudal margin of midbody terga. See also Key
below.
Description: Length about 8.5 (holotype) or 7 mm (paratype), width ca 0.9 and 1.0 (holotype) or 0.8 and 0.9
mm (paratype) on midbody pro- and metazonae, respectively. Holotype with 48 podous + 1 apodous segment +
telson, paratype with 43p + 1a + telson. Length to width ratio > 10:1.
Coloration in alcohol rather uniformly red- or grey-brown to yellowish with a narrow light axial line (alcohol
material), anterior part of body in paratype darker, marbled grey; metatergal bosses/tubercles at caudal margin and
antennae mostly dark grey-brown while a very wide mid-dorsal stripe lighter both in holo- and paratype (Fig. 6A,
C). Ocelli inside a black ocular field. Body very clearly flattened dorsoventrally, hirudiform, regularly, but strongly
broadened until segment 6 or 7, similarly regularly and clearly narrowing towards telson on last 4 or 5 segments.
Head very small and rather narrow, coniform, about 1/3 as broad as collum and about twice as long as broad;
rostrum narrowly rounded (Figs 6A, 7A). Ocelli large, convex, 2+2 on each side of head arranged in a nearly
longitudinal line, each ocellus with a long seta at base. Antennae short, stout, erect, rod-shaped, not clavate, about
as long as collum width; antennomere 6 largest (Figs 6A, B, 7A).
Collum biconvex, short and broad (Fig. 7A), like head characteristically strongly inclined forward to form a
small casque-like shield, also distinctly separated from next tergum (Fig. 6B, D); paraterga on collum small,
strongly declined, subvertical, each delimited by an oblique sharp bend; dorsal surface rather flat, only slightly
impressed medially; caudal margin with a transverse row of several small grains/tubercles (Fig. 7A) and a
considerable lighter gap between paramedian pair (Fig. 6). Axial suture inconspicuous, but visible starting with
collum. Following paraterga nearly acute at caudal corner, especially clearly drawn behind tergal margin only in 2
nd
and a few caudalmost segments, paraterga of penultimate segment fused medially into a broad, terminally slightly
concave plate with two caudolateral denticles and a vestigial axial suture (Figs 6C, 7C). Lateral peritremes on
paraterga evident, delimited by a sulcus both dorsally and, to lesser degree, ventrally. Metaterga slightly, but clearly
elevated caudally, each with 9–11 + 9–11 small bosses/tubercles at caudal margin, like collum with a considerable
lighter gap between paramedian pair. Surface of collum and following metaterga faintly rugulose and
microtuberculate, dull all over. Ozopores lateral, starting with segment 5, small, inconspicuous, each lying at
bottom of a small oblong groove in front of caudolateral corner of a clearly thickened, nearly porostele-shaped
peritreme (Fig. 7B). Telson small, completely concealed in dorsal view by caudal plate of penultimate segment’s
fused paraterga (Figs 6A, C, D, 7C).
Sterna about 2/3 as wide as coxae, the latter almost in touch medially (Fig. 6A, C, E). Legs 6-segmented. Male
coxa 2 with a short, tube-shaped, simple, basal gonapophysis directed caudomesad. Claws simple (Fig. 8A). Coxal
sacks starting with leg 3.
Anterior gonopods (Fig. 8B) apparently 5-segmented, C-shaped, strongly incrassate and stout. Tarsus with a
strong apical stylet and a gutter for accommodation of posterior gonopod tarsus (Fig. 8B, C). Posterior gonopods
(Fig. 8D) apparently 6-segmented, much slenderer; tarsus longest, flagelliform, very simple.
Remarks. The order Siphonocryptida has hitherto been known to contain only six species in two genera and a
single family (Enghoff and Golovatch 1995; Korsós et al. 2008, 2009; Enghoff 2010). The group’s highly peculiar
distribution only emphasizes its relictual status (Shelley and Golovatch 2011). Thus, the genus Siphonocryptus
Pocock, 1894, comprises three species, one in Sumatra, Indonesia, the other two in continental Western Malaysia
(Enghoff and Golovatch 1995; Enghoff 2010). In contrast, the distribution pattern of Hirudicryptus Enghoff &
Golovatch 1995 is trans-Palaearctic (Fig. 10). The type species, H. canariensis (Loksa, 1967), occurs only on
Madeira and the Canaries, where it is largely confined to the relict laurisilva biome. The second species, H.
taiwanensis Korsós, Enghoff & Chang, 2008, is endemic to Taiwan, whereas the third, H. quintumelementum
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FIGURE 6. Hirudicryptus abchasicus sp. nov., male holotype (A–C) and female paratype (D, E). A, D, E: habitus, mainly
sublateral, lateral and mainly ventral views, respectively. B: anterior part of body, subventral view. C: anterior and posterior
parts of body, dorsal view. Pictures by K. Makarov, taken not to scale.
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FIGURE 7. Hirudicryptus abchasicus sp. nov., male holotype. A: head, collum and segment 2, oral view. B: midbody
paraterga, lateral view. C: caudal end of body, dorsal view. Scale bar: 0.5 mm.
Korsós, Geoffroy & Mauriès, 2009, is only known from a few females collected at a 2500 m elevation in Nepal,
Himalayas (Enghoff and Golovatch 1995; Korsós et al. 2008, 2009). It may well be that the distribution pattern
under consideration dates back at least to the Oligocene times of the so-called “Warm Earth” to have highly
probable explanations rooted in palaeobotanical evidence. These imply a gradual shrinkage and disruption ever
since of the previously dominating and continuous subtropical biome (Golovatch 1997; Zherikhin 2003). Being so
vastly disjunct, the present-day distribution of Siphonocryptida is best accounted for by extinction events (Shelley
and Golovatch 2011).
The discovery of H. abchasicus sp. nov. in the Caucasus very nicely bridges the huge gap between
Macaronesia and the Himalayas (Fig, 10). This species is clearly a highly relictual element in the Caucasian fauna.
Luckily, it stems from a nature reserve, thus demanding no other special measures of protection.
The following key can be offered to separate all four presently known species of Hirudicryptus.
1. Caudal margin of last tergum straight, devoid of lateral prongs. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
- Caudal margin of last tergum clearly concave, flanked by small lateral prongs (Figs 6C, 7C) . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2. Midbody terga with 14–15 + 14–15 tubercles at caudal margin. Macaronesia . . . . . . . . . . . . . . . . . . . . . . . . . . . H. canariensis
- Midbody terga with 9–11 + 9–11 tubercles at caudal margin. Nepal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. quintumelementum
3. Midbody terga with 6–8 + 6–8 tubercles at caudal margin Adults larger: length 10.5–16.8 mm (males), up to 19.4 mm
(females), width 1.2–2.0 or 1.3–2.5 mm, respectively. Taiwan. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. taiwanensis
- Midbody terga with 9–11 + 9–11 tubercles at caudal margin (Fig. 6C). Adults smaller, 7–8.5 mm long, 0.9–1.0 mm wide. Cau-
casus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .H. abchasicus sp. nov.
Conclusion
The distribution of Colobognatha in the Caucasus (Fig. 9) shows clear-cut inclinations of these millipedes to
dwelling in reasonably warm and humid forest habitats, both lowland and mid-montane. Hardly surprisingly, this
small animal group is only represented there by three species, genera, families and orders which, being
unquestioned humicoles, are mostly confined to areas located close enough to warm temperate coastal regions of
the Black and Caspian seas. Harsher environments such as high mountains and drier terrain seem to be fully devoid
of representatives of this diplopod superorder. This holds especially true for the mostly high-montane, colder and/
or arid Caucasus Minor within Armenia and Azerbaijan, as well as extra-Hyrcanian Iran. With increasing climate
continentality, even Hirudisoma roseum, the most widespread species, steadily declines from the Black Sea coast
eastward along the Caucasus Major, especially strongly so on its northern, drier macro slope. Fioria hyrcana is
strictly confined to Hyrcania while Hirudicryptus abchasicus sp. nov. to a near-coastal forest patch in the NW
Caucasus within Colchis.
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FIGURE 8. Hirudicryptus abchasicus sp. nov., male holotype. A: leg 11, caudal view. B: anterior gonopods, anterior view; C:
tip of right anterior gonopod, anterodorsal view. D: right posterior gonopod, ventrolateral view. Scale bar: 0.1 mm.
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FIGURE 9. Distribution of Colobognatha in the Caucasus and adjacent parts of Turkey and Iran. Blue square—Hirudisoma
roseum (Victor, 1839); red asterisk—Fioria hyrcana Golovatch, 1980; green triangle—Hirudicryptus abchasicus sp. nov.
FIGURE 10. Distribution of the genus Hirudicryptus.
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Acknowledgements
We are most grateful to all relevant collectors whose material is used in the present paper for placing it at our
disposal, as well as for allowing us to deposit their material at our choice. Henrik Enghoff (ZMUC) generously
shared with us information concerning the Caucasian Colobognatha housed in the ZMUC collection. Mzia Kokhia
(Tbilisi, Georgia) kindly helped us solve a toponymic problem.
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