The millipede family Polydesmidae in the Caucasus (Diplopoda: Polydesmida)

Abstract

The family Polydesmidae is represented in the Caucasus by two genera and 11 species: Polydesmus abchasius Attems, 1898, P. lignaui Lohmander, 1936, P. muralewiczi Lohmander, 1936 and P. mediterraneus Daday, 1889 (all confined to the NW and W Caucasus, but of which only the former three are endemic to the region, whereas the latter species is an introduction), as well as Brachydesmus assimilis Lohmander, 1936 (endemic to most of the region, except Hyrcania), B. pigmentatus Attems, 1951 (= B. pereliae Golovatch, 1976, syn. n.) (subendemic to Hyrcania), B. superus Latzel, 1884 (a cosmopolitan introduction), B. furcatus Lohmander, 1936 (= B. furcatus exiguus Strasser, 1970, syn. n.) (endemic to the NW Caucasus), B. kalischewskyi Lignau, 1915 (= B. karawajewi Lohmander, 1928, = B. ferrugineus Lohmander, 1936, = B. talyschanus Lohmander, 1936, = B. bidentatus Golovatch, 1976, all syn. n.) (a highly polymorphous and widespread species, apparently in a stage of active speciation, subendemic to the entire region), B. kvavadzei sp. n., from Ajaria, Georgia, and B. simplex sp. n., from Abkhazia and Sochi, Krasnodar Province, Russia. All known Caucasian species of Polydesmidae are described in due detail, abundantly illustrated and keyed, and their distributions mapped.

Full text

Accepted by W. Shear: 11 Jan. 2016; published: 1 Mar. 2016
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Copyright © 2016 Magnolia Press
Zootaxa 4085 (1): 001
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Article
1
http://doi.org/10.11646/zootaxa.4085.1.1
http://zoobank.org/urn:lsid:zoobank.org:pub:7F819439-FD77-4EF2-8E26-7B46F9B5FAAB
The millipede family Polydesmidae in the Caucasus (Diplopoda: Polydesmida)
SERGEI GOLOVATCH
1,4
, ALEKSANDR EVSYUKOV
2
& HANS 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, 02826 Görlitz, Germany
4
Corresponding author. E-mail: sgolovatch@yandex.ru
Table of contents
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .1
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
Material and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Taxonomic part . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3
Polydesmus (s. str.) abchasius Attems, 1898 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Polydesmus (s. str.) lignaui Lohmander, 1936 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Polydesmus (s. str.) muralewiczi Lohmander, 1936 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Polydesmus (Nomarchus) mediterraneus Daday, 1889. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
Polydesmus (s. str.) monticola Latzel, 1884 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
Brachydesmus (s. str.) assimilis Lohmander, 1936 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
Brachydesmus (s. str.) pigmentatus Attems, 1951 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
Brachydesmus (Eubrachydesmus) superus Latzel, 1884 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Brachydesmus (Lophobrachydesmus) furcatus Lohmander, 1936 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
Brachydesmus (Haplobrachidesmus) kalischewskyi Lignau, 1915 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .21
Brachydesmus (Haplobrachidesmus) kvavadzei sp. n. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
Brachydesmus (Haplobrachidesmus) simplex sp. n. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .45
Key to Polydesmidae species presently known to occur in the Caucasus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .50
Abstract
The family Polydesmidae is represented in the Caucasus by two genera and 11 species: Polydesmus abchasius Attems,
1898, P. lignaui Lohmander, 1936, P. muralewiczi Lohmander, 1936 and P. mediterraneus Daday, 1889 (all confined to
the NW and W Caucasus, but of which only the former three are endemic to the region, whereas the latter species is an
introduction), as well as Brachydesmus assimilis Lohmander, 1936 (endemic to most of the region, except Hyrcania), B.
pigmentatus Attems, 1951 (= B. pereliae Golovatch, 1976, syn. n.) (subendemic to Hyrcania), B. superus Latzel, 1884 (a
cosmopolitan introduction), B. furcatus Lohmander, 1936 (= B. furcatus exiguus Strasser, 1970, syn. n.) (endemic to the
NW Caucasus), B. kalischewskyi Lignau, 1915 (= B. karawajewi Lohmander, 1928, = B. ferrugineus Lohmander, 1936,
= B. talyschanus Lohmander, 1936, = B. bidentatus Golovatch, 1976, all syn. n.) (a highly polymorphous and widespread
species, apparently in a stage of active speciation, subendemic to the entire region), B. kvavadzei sp. n., from Ajaria, Geor-
gia, and B. simplex sp. n., from Abkhazia and Sochi, Krasnodar Province, Russia. All known Caucasian species of
Polydesmidae are described in due detail, abundantly illustrated and keyed, and their distributions mapped.
Key words: Polydesmidae, taxonomy, new species, new synonymy, key, distribution, map, polymorphism, Russia, Abk-
hazia, Georgia, Azerbaijan, Turkey, Iran

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Introduction
The largely Holarctic family Polydesmidae is one of the most common and quite diverse among the Diplopoda in
the Caucasus (e.g. Lohmander 1936, Lokšina and Golovatch 1979). No fewer than 16 species or subspecies from
two genera, Polydesmus Latreille, 1803, and Brachydesmus Heller, 1858, have hitherto been reported from the
region, but the fauna still remains poorly-known.
Polydesmus is primarily an amphi-Palaearctic genus, of which three species of Polydesmus s. str., i.e. P.
abchasius Attems, 1898, P. lignaui Lohmander, 1936, P. m u r a l e w i c z i Lohmander, 1936, are endemic to the NW
and W Caucasus. One more, the eastern Mediterranean P. mediterraneus Daday, 1889, seems to be an introduced
element, whereas another species has been reported from the region by mistake. The basically Euro-Mediterranean
genus Brachydesmus is more diverse and widespread in the Caucasus, being represented there by four subgenera.
Among them, the subendemic Haplobrachidesmus Lohmander, 1928, dominates the fauna (five descrived species:
B. kalischewskyi Lignau, 1915, B. karawajewi Lohmander, 1928, B. ferrugineus Lohmander, 1936, B. talyschanus
Lohmander, 1936, and B. bidentatus Golovatch, 1976) and blankets the whole region, including the adjacent parts
of N and NE Turkey and NW Iran. Brachydesmus s. str., a very large subgenus which includes dozens of species or
subspecies (e.g. Attems 1940), is represented in the Caucasus by two species, including one endemic (B. assimilis
Lohmander, 1936) and one subendemic (B. pigmentatus Attems, 1951). One more, the ubiquitous anthropochore B.
superus Latzel, 1884, is from the similarly diverse subgenus Eubrachydesmus Attems, 1912. In contrast, the
oligotypic eastern Mediterranean subgenus Lophobrachydesmus Attems, 1912, contains in the Caucasus only two
formal subspecies of a single endemic species: B. furcatus furcatus Lohmander, 1936, and B. furcatus exiguus
Strasser, 1970. In general, the eastern range limits of Brachydesmus lie in the Caucasus s.l., except for one outlier
species reported from the W Kopetdagh Mountains, SW Turkmentistan, while Polydesmus spp. occur only in the
NW and W Caucasus, totally declining further to the east to reappear only in Japan, Hongkong and N Vietnam.
There seems to be no need for an extensive historical overview of the explorations of the Polydesmidae in the
Caucasus, as all relevant records and references are given in the catalogue sections under each species concerned
and when appropriate, also supplied with necessary remarks. In addition, quite detailed historical accounts,
however incomplete and outdated, are available in Muralewicz (1911) and Lohmander (1936).
The first polydesmid which could have been documented from the Caucasus was “Polydesmus maculicornis
Brandt, Nob.”, but Brandt failed to describe it. The “type series” of his nomen nudum which derives from “Tiflis”
is still kept in the St. Petersburg Museum, treated below amongst Brachydesmus assimilis samples. The truly first
species of Polydesmidae to be described from the region was Polydesmus abchasius Attems, 1898, from Abkhazia
(Attems 1898). Lignau (1915) added Brachydesmus kalischewskyi Lignau, 1915, and Polydesmus minor Lignau,
1915, to the Abkhazian list, but unfortunately the latter species was a nomen praeoccupatum and had to be
renamed. Lohmander (1936), in his milestone treatment of the entire diplopod fauna of the Caucasus, dedicated
that species to N. Lignau: P. lignaui Lohmander, 1936.
Indeed, the contributions by H. Lohmander to the study of the millipede fauna of the former Soviet Union were
outstanding. Not only had he summarized the Caucasian fauna (Lohmander 1928, 1932, 1936), bringing it to a total
of 79 species, including nine in Polydesmidae, but he had also identified lots of material he had failed to publish
because of his untimely death in 1961. Thus, Kobakhidze’s (1965) paper which reviewed the fauna of Georgia,
Transcaucasia was largely based on unpublished samples identified by H. Lohmander. Similarly, a large, but only
partly identified collection of Diplopoda, including that from the Caucasus, which had been sent on loan to H.
Lohmander by M. Ghilarov (of the former Institute of Evolutionary Animal Morphology and Ecology, USSR
Academy of Sciences in Moscow) was returned to him soon after Lohmander’s death. This material is now housed
in the Moscow Museum. The rather few Polydesmidae samples, Lohmander’s identifications of which were also
verified by the Polish specialist C. Dziadosz, are being put on record here for the first time. The same concerns the
few samples retained by Lohmander in the Gothenburg Museum.
Since Attems’ (1940) global review of several families of Polydesmida, including Polydesmidae, very modest
progress in the knowledge of Caucasian polydesmids has been achieved. A few papers provided more or less
deficient checklists and/or faunistic records: entire Caucasus (Lang 1959, Lokšina and Golovatch 1979), Colchis
(Talikadze 1984), Hyrcania (Samedov et al. 1972; Rakhmanov 1972), Georgia (Kobakhidze 1965, Golovatch
1981a, Barjadze et al. 2015), Azerbaijan (Bababekova 1996), the Stavropol Province (Zuev 2014), Caucasian
Biosphere Nature Reserve, Krasnodar Province and the Republic of Adygea (Chumachenko 2012), as well as

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individual species descriptions, redescriptions or records (Strasser 1970, Golovatch 1976, 1981b, Loginova 1993,
1995, Kokhia 2011).
The present paper provides a complete review and a partial revision of the Polydesmidae in the fauna of the
Caucasus s. l. It continues our recent efforts in summarizing and updating the taxonomy and distributions of
individual larger groups of Caucasian Diplopoda (Golovatch et al. 2015, Evsyukov et al. 2016). Only rather brief
descriptive notes are provided for most of the known species, but always accompanied by sufficiently abundant
illustrations to make them easily recognizable. In addition, the paper is concluded by a general key to all species of
Polydesmidae encountered in the region.
Material and methods
A large material of Caucasian Polydesmidae has been amassed by the first author, now largely 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. A few samples, mainly type material, belong to the
Zoological Institute, Russian Academy of Sciences, St. Petersburg (ZISP); a few more, mainly unidentified, to the
Institute of Animal Systematics and Ecology, Siberian Branch, Russian Academy of Sciences, Novosibirsk (ZMN),
the Hungarian Natural History Museum, Budapest (HNHM), the Institute of Zoology, National Academy of
Sciences of Azerbaijan, Baku (IZB), the Naturhistorisk Museum Göteborg (NHMG), the Senckenberg Staatliches
Museum für Naturkunde, Görlitz (SMNG), and the Museum and Institute of Zoology, Polish Academy of Sciences,
Warsaw (MIZW), as indicated below.
In the catalogue sections, D stands for a description or descriptive notes, R for new or repeated records, while
M for a mere mention. Only records in the Caucasus are considered for alien species.
Taxonomic part
Polydesmus (s. str.) abchasius Attems, 1898
Figs 1, 3B, 3C, 4A, Map 1
Polydesmus abchasius Attems, 1898: 437 (D).
Polydesmus abchasius—Lignau, 1903: 106, 132 (R); 1915: 372 (R, D); Muralewicz, 1907: 340, 349 (R); 1911: 4, 9 (M); 1913:
220 (R); Issaev, 1911: 297 (R); Lohmander, 1936: 11 (M); Lang, 1959: 1790 (M); Kobakhidze, 1965: 391 (R); Lokšina and
Golovatch, 1979: 384 (M); Talikadze, 1984: 143 (R); Chumachenko, 2012: 452 (R).
Polydesmus complanatus—Muralewicz, 1907: 340, 349 (R); 1911: 9 (M).
Polydesmus (Polydesmus) abchasius—Attems, 1940: 21 (D).
NON Polydesmus (Polydesmus) abchasius—Bababekova, 1996: 90 (R).
Material examined. 4 males, 3 females, 2 juv. (ZMUM), RUSSIA, Krasnodar Prov., Goryachiy Klyuch, ca 12 km
SW of Fanagoriyskoye, near Cave Fanagoriyskaya, Fagus, Acer, Picea etc. forest, litter and under logs, 19.V.1983,
leg. S. Golovatch; 1 female (ZMUM), Krasnodar Prov., Sochi, 5.5 km NE of Krasnaya Polyana, lower course of
Achipse River, ca 600 m a.s.l., N43°43’10”, E40°15’25”, 19–23.VIII.2014, leg. K. Makarov and A. Matalin; 3
females (AE), Adygea, near Mt Oshten, lake Psenodakh, 10.VII.1997, leg. G. Bakhtadze; 2 males, 1 female (AE),
Adygea, Maikop, 14.III.2008, leg. M. Nartbiev; 4 females (AE), Adygea, near Guzeripl, Abies forest, 19.VI.2011,
leg. D. Khisametdinova and E. Bortnikov; 1 male (AE), Adygea, Partizanskaya Polyana, Armyanka River valley,
1.VII.2011, leg. D. Khisametdinova and V. Shmatko; 7 females (AE), same locality, 4.VII.2011, leg.
Khisametdinova; 1 male, 1 juv. (AE), Adygea, near Nikel, Belaya River valley, under stones, 23.VI.2007, leg. E.
Terskov; 2 males (AE), same locality, on tree trunks, 29.VI.2010, leg. I. Gorbenko; 1 female (ZMUM), Adygea,
Caucasian Nature Reserve, Pasture Abago S of Guzeripl, 1350–1400 m a.s.l., Abies and Fagus forest, litter, under
bark and stones, 24–26.V.1985, leg. S. Golovatch; 1 female (ZMUM), Karachaevo-Cherkessia, near Arkhyz,
1450–1600 m a.s.l., Abies and Fagus forest, under stones and bark, litter, 3.VIII.2015, leg. A. Evsyukov; 1 male
(AE), Krasnodar Prov., Zazulin, forest, 20.VI.2007; 3 males, 2 females (AE), Krasnodar Prov., Khadyzhensk,

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forest, 15.VI.2007, 1 juv. (AE), Krasnodar Prov., Arkhipo-Osipovka, mountain slope, forest, under stones and bark,
29.VIII.2008; 2 males, 1 female (AE), same locality, 30.VIII.2008; 1 female, 5 juv. (AE), same locality,
31.VIII.2008, all leg. A. Evsyukov and D. Khisametdinova; 1 female (SMNG 33576), Adygea, Sakhrai, Kuna
Valley, 20.V.2004; 1 female (SMNG 34133), same locality, 20.VIII.2005; 1 male (SMNG 34158), same locality,
30.VIII.2005; 1 male, 3 females (SMNG 33581/2), Adygea, Polyana Shostakova, 23.V.2004; 1 female, 1 juv.
(SMNG 34141), same locality, 24.VIII.2005; 1 female, 2 juv. (SMNG 34136), Adygea, Koryto, 22.VIII.2005; 1
female (SMNG 34147), Adygea, Muly, Trash, 25.VIII.2005, all leg. K. Voigtländer; 1 male, 1 juv. (HR), Krasnodar
Prov., Solokhaul, 2.5 road-km to Dagomys, 260 m a.s.l., N43.7997º, E39.6481º, 11.VIII.2012; 3 males, 1 female
(HR), Krasnodar Prov., Lagonaki Plateau, source area of Kurdships River, N44.0792º, E40º, 16.VIII.2012; 1
female (HR), Krasnodar Prov., Yatyrgvarta Ridge, hut N of Gora Armovka, 1830 m a.s.l., N43.8908º, E40.6631º,
19.VIII.2012, all leg. F. Walther.
1 male, 3 females (ZMUM), ABKHAZIA, Gagra Distr., Arabika Massif, Gelgeluk, under stones,
29.VIII.2014, leg. I. Turbanov; 8 females (SMNG 14685), Transcaucasus, Riza-See (= Lake Ritsa), 20.VI.1984,
leg. W. Dunger.
2 males, 6 females, 1 juv. (ZMUM), GEORGIA, Ajaria, Kintrish Nature Reserve, Zeraboseli, 450–600 m
a.s.l., deciduous forest, 1–3.VI.1981, leg. S. Golovatch and J. Martens; 1 male (ZMN), Ajaria, near Batumi,
Koromistavi, Fagus and Castanea forest, 1000–1200 m a.s.l., X.1987, leg. D. Logunov; 1 female (HR), Ajaria,
Khala, Chakvistskhali River valley, 1 road-km upstream, entrance to Mtirala National Park, 90 m a.s.l., N41.6961º,
E41.8228º, 28.IX.2011; 1 female (HR), Ajaria, Khorolistavi to Mt Mta Mtirala, 730 m a.s.l., N41.6494º,
E41.7731º, 1.X.2012; 1 male (HR), Samegrelo and Zeno Svaneti, Taleri, Tekhuri River valley 6.7 road-km to
Lebarde, 380 m a.s.l., N42.5989º, E42.3497º, 10.X.2011; 1 juv. (HR), Samegrelo and Zemo Svaneti, Mukhuri,
N42.6372º, E42.1778º, 22.VIII.2014; 1 juv. (HR), Guria, Chkhakoura to Bakhmaro, N41.89º, E42.3678º,
21.VIII.2014, all leg. F. Walther; 1 female, 6 juv. (HR), Ambrolauri to Djava, E of Oni, NE of Pipileti, N42.5679,
E43.4995, 1.VI.2014; 3 males, 1 female, 2 juv. (HR), Ambrolauri to Djava, Oni E, Pipileti NE, N42.5681,
E43.4953, 1.VI.2014; 1 male, 1 female (HR), Ambrolauri to Djava, E of Oni, N of Pipileti, N42.5823, E43.4986,
1.VII.2014; 1 female (HR), Baghdati S to Akhaltsikhe, N of Abastumani, N41.8564, E42.7901, 1.VI.2014, all leg.
L. Mumladze; 1 juv. (ZMUM), Tkibuli Distr., between Tkibuli and Nikortsminda, Abies and Fagus forest, litter,
22.X.1981; 5 females (ZMUM), Ambrolauri Distr., Cave Nikortsminda near Nikortsminda, 1100 m a.s.l., 40–50 m
deep, on walls and rotten wood, 22.X.1981, all leg. S. Golovatch; 1 male (ZMUM), Sataplia Nature Reserve, 400
m a.s.l., Fagus and Buxus forest, litter, 5.VI.1981, leg. S. Golovatch and J. Martens; 1 male, 1 female, 2 juv.
(ZMUM), Racha, Oni Distr., N of Utsera, deciduous forest, near spring, rock, litter, 20.X.1981, leg. S. Golovatch.
Descriptive notes. Adult body with 20 segments (collum plus 17 podous plus one apodous body rings, plus
telson). Length ca 13–19 mm (male, female), width of midbody metazonae 1.9–3.3 (male) or 2.4–3.5 mm (female),
metazonite to prozonite width ratio > 2.0 (Fig. 1). Live coloration usually uniformly intense red to bright pink, in
alcohol often brown to faded light yellow-brown; legs light (Fig. 1).
Tegument smooth and moderately shining. Antennae long, reaching behind body segment 2 when stretched
dorsally, slightly clavate, usually infuscate towards a pallid tip. In width, head < collum < segment 2 < 3 < 4 < 5=15
or 16, thereafter body gradually to rapidly tapering towards telson (Fig. 1). Collum transversely sublanceolate.
Paraterga very strongly developed and lying at about upper ¼ of body height, mostly subhorizontal, only in collum
and 2–3 following segments more (male) or less (female) clearly upturned (Figs 1A, 3B). Caudolateral corner of
collum almost pointed, but obtuse-angled, of midbody paraterga subrectangular (Figs 1, 3C), of following
paraterga increasingly acute, but clearly drawn behind rear tergal margin only in segments 17–19 (Fig. 1). Lateral
margins of paraterga straight or nearly so, at most only slightly rounded (Fig. 1). Collum and central parts of
following metaterga with three transverse rows of faint, polygonal, flat, nearly obliterate, setigerous bosses (Figs 1,
3C). Tergal setae very small, only a little longer in front row on collum and in caudal row on segment 19. Paraterga
with three or four faint incisions at lateral margin of poreless and pore-bearing segments, repectively. Stricture
between pro- and metazonae wide, shallow and nearly smooth, at most faintly rugulose. Limbus microcrenulate-
microdentate. Pleurosternal carinae wanting. Epiproct short, conical, subtruncate at tip, pre-apical lateral papillae
evident (Fig. 1). Hypoproct subtrapeziform, paramedian setiferous papillae evident, well-separated and lying near
caudal edge.
Legs densely setose, long, in male crassate and longer (Fig. 1A), prefemora only faintly bulging laterad; all
postfemora, tibiae and tarsi typically with sphaerotrichomes ventrally.

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FIGURE 1. Habitus of Polydesmus abchasius Attems, 1898. A: male from Gelgeluk, Abkhazia, lateral view. B: female from
near Krasnaya Polyana, Krasnodar Prov., dorsal view. C: female ca 18 mm long from an unspecified locality in Abkhazia,
dorsal view. Pictures A and B by K. Makarov, taken not to scale; watercolour C after Lignau (1915).
Gonopod (Fig. 4A) exomere (ex) strongly subgeniculate at about distal ¼ and densely spinulate apically, with
a strong tooth (d) proximal to geniculation; endomere (en) with a large hairy pulvillus (pu) at base of a slender,
simple, apically curved rod (r) which is about half as long as ex.
Remarks. Muralewicz’s (1907, 1927) records of the eastern European P. complanatus (Linnaeus, 1758), in
Abkhazia, W Caucasus and at Vladikavkaz, N Caucasus, respectively, are clearly erroneous, the former actually
being referred to P. abchasius, the latter probably to P. muralewiczi or P. lignaui (cf. Lohmander 1936).
Bababekova’s (1996) record of P. abchasius in the Caucasus Major within Azerbaijan is certainly wrong as well,
since this species is confined to the NW and W Caucasus.

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Polydesmus (s. str.) lignaui Lohmander, 1936
Figs 2, 3D–3G, 4B, Map 1
Polydesmus minor Lignau, 1915: 374 (D).
Polydesmus lignaui Lohmander, 1936: 11 (M), nom. nov. pro P. minor Lignau, 1915, non P. m i no r Bollman, 1888.
Polydesmus (Polydesmus) lignaui lignaui—Attems, 1940: 21 (M).
Polydesmus lignaui—Attems, 1940: 20 (D); Lang, 1959: 1790 (M); Kobakhidze, 1965: 391 (R); Lokšina and Golovatch, 1979:
384 (M); Talikadze, 1984: 143 (R); Zuev, 2014: 348 (R, D).
Polydesmus minor—Lang, 1959: 1790 (M).
Material examined. 1 male (ZMUM), RUSSIA, Krasnodar Prov., Sochi, Lazarevskoe, garden, pitfall traps, 7–
17.VII.1986; 1 female (ZMUM), same locality, 21.VIII–4.IX.1986, all leg. Yu. Zayats; 1 female, 2 juv. (ZMUM),
Krasnodar Prov., Pass Mikhailovskii, Fagus forest, decaying log, 5.VII.1956; 1 female (ZMUM), Krasnodar Prov.,
Gelenjik, Fagus forest, 5.VII.1956; all leg. K. Arnoldi, all det. H. Lohmander, det. C. Dziadosz, 1968; 2 males, 1
female, 1 juv. (ZMUM), Krasnodar Prov., Severskaya Distr., 2–10 km S of Ubinskaya, 300–450 m a.s.l., Quercus,
Fagus, Carpinus etc. forest, litter and under bark, 3–4.VII.1986, leg. S. Golovatch; 4 males, 14 females, 1 juv.
(ZMUM), Karachaevo-Cherkessia, Teberda Nature Reserve, Dombai, 1700–1800 m a.s.l., floodplain Abies, Fagus,
Picea, Betula, Acer etc. forest, litter, under bark and stones, 11.V.1985; 1 female, 4 juv. (ZMUM), same locality, Mt
Mussa-Achitara, 2700–2800 m a.s.l., alpine meadow, under stones, 29.VII.1986; 1 male, 4 females (ZMUM),
Karachaevo-Cherkessia, Teberda Nature Reserve, Gonachkhir Canyon between Teberda and Dombai, road to
Klukhor Pass, 1700–1900 m a.s.l., Abies, Fagus, Acer etc. forest, 1.VI.1985, all leg. S. Golovatch; 1 male, 1
female, 1 subadult female (AE), Karachaevo-Cherkessia, Teberda Nature Reserve, Fagus and Carpinus forest,
litter, under logs, 26.VII.2012; 1 male, 3 females, 10 juv. (AE), same locality, Teberda River valley, meadow with
Fagus and Carpinus forest, 26.VII.2012, leg. D. Khisametdinova and Y. Kochetov; 3 males, 3 females (ZMUM),
Kabardino-Balkaria, near Mt Elbrus, 8 km off month of Adyl-Su River, Djan-Tugan, 2150–2200 m a.s.l.,
xerophytous meadow, pitfall traps, 11–20.VII.1987, leg. A. Kulikov.
3 males, 2 juv. (ZMUM), ABKHAZIA, SE of Lake Ritsa, Pass Anchkho, 2000 m a.s.l., Betula and
Rhododendron, litter, 16.VIII.1986, leg. S. Golovatch; 1 male, 1 female, 3 juv. (ZMUM), Sukhumi, University
Campus, N42°58,4’ E41°04,04’, ca 50 m a.s.l., Encopytus, tree shaking, litter, 7.XII.2003, leg. Yu. Marusik and G.
Antonova; 2 males, 3 females, 2 juv. (ZMUM), Sukhumi Botanical Garden, litter, 20.VIII.1986; 1 male, 3 females
(ZMUM), Sukhumi Distr., Bzyb Valley, Pskhu, 700–950 m a.s.l., Fagus, Quercus, Castanea etc. forest, litter, under
bark and stones, 15–16.VIII.1983, all leg. S. Golovatch; 1 female, 4 juv. (ZMUM), Kodor Mt. Ridge, Tkvarcheli
Distr., SSE of Khojali, 1750 m a.s.l., 28.VI.1986, leg. I. Ushakov.
1 female (ZMUM), GEORGIA, Mingrelsky (= Egrissky) Mt. Range, NW slope of Mt Tsekhuri, 1500–1800 m
a.s.l., Fagus forest, litter, 18–19.VIII.1988, leg. I. Belousov; 1 female (ZMUM), 10 km N of Djvari, 800 m a.s.l.,
Buxus, Fagus, Picea, Taxus etc. forest, litter, 20–21.VIII.1986; 3 females (ZMUM), Svanetia, Mestia, 1500 m a.s.l.,
Betula and Rhododendron on moraine, litter and under stones, 5 and 16.IX.1986; 1 female, 1 juv. (ZMUM), same
locality, under stones, 22.X.1979, all leg. S. Golovatch; 1 juv. (ZMUM), Svanetia, Mestia Distr., Lukhi N of
Khaishi, Nenskra Valley, 800 m a.s.l., forest, litter, 2.IX.1986, leg. A. Ryvkin; 22 males, 14 females, 6 juv.
(ZMUM), Svanetia, ca 40 km W of Mestia, Kherkhvashi E of Nakra (= Naki), 1200–1700 m a.s.l., Quercus, Fagus,
Carpinus, Picea, Abies etc. forest, litter and under bark, 21.VIII.–21.IX.1986; 5 males, 3 females, 2 juv. (ZMUM),
same locality, above Kherkhvashi, 1900–2200 m a.s.l., timberline (Rhododendron, Picea, Abies) and subalpine
meadows, 21.IX.1986, all leg. S. Golovatch.
Descriptive notes. Length ca 10–13 mm (male, female), width of midbody metazonae 1.25–1.7 (male) or 1.4–
2.1 mm (female), metazonite to prozonite width ratio < 2.0 (Fig. 2). Live coloration usually uniformly red or red-
brown to pink, in alcohol brown to faded light brown or pinkish; legs light (Fig. 2).
All characters as in P. abchasius (Figs 2, 3D–3G, 4B), except as follows. Collum about as broad as head (Fig.
2A, 2B, 2F). Paraterga 2–4 in female usually not upturned, subhorizontal (Fig. 3G). Gonopod (Fig. 4B) endomere
rod typically axe-shaped apically.

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FIGURE 2. Habitus of Polydesmus lignaui Lohmander, 1936. A: male from Kherkhvashi, Svanetia, Georgia, dorsal view. B–
E: anterior, middle and caudal parts of body, and entire body, respectively, of male from Sukhumi University Campus,
Abkhazia, dorsal (B–D) and lateral views (E). F: male ca 12 mm long from an unspecified locality in Abkhazia, dorsal view.
Pictures A–E by K. Makarov, taken not to scale; watercolour F after Lignau (1915).

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FIGURE 3. Habitus of Polydesmus muralewiczi Lohmander, 1936 (A, H–J), P. lignaui Lohmander, 1936 (D–G) and P.
abchasius Attems, 1898 (B, C). A, H–J: male from near Kursavka, Stavropol Prov., Russia, entire body, anterior body part, left
half of metatergite 10 and caudal body part, respectively, dorsal views. B, C: male from Pasture Abago, Caucasian Nature
Reserve, Adygea, Russia, anterior part of body, lateral view, and left half of metatergite 10, dorsal view, respectively. D–F:
male from Dombai, Karachaevo-Cherkessia, Russia, anterior body part, left half of metatergite 10 and caudal part of body,
respectively, dorsal views. G: female from Dombai, anterior part of body, lateral view. Picture A by K. Makarov, taken not to
scale. Scale bar: 1.0 mm (B–J).

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FIGURE 4. Left gonopods of Polydesmus abchasius Attems, 1898, P. lignaui Lohmander, 1936 and P. muralewiczi
Lohmander, 1936 (A–C), and right gonopod of P. muralewiczi Lohmander, 1936 (D). A: male from near Lake Ritsa, Abkhazia,
mesal view. B: male from Sukhumi Botanical Garden, Abkhazia, mesal view. C: male from near Kursavka, Stavropol Prov.,
Russia, mesal view. D: male lectotype from near Psebai, Krasnodar Prov., Russia (after Lohmander 1936). Designations: en,
endomere; ex, exomere; d, exomere tooth; pu, hairy pulvillus; r, endomere rod. Scale bar: 0.2 mm (A–C), not to scale (D).

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MAP 1. Distribution of Polydesmus species in the Caucasus: P. abchasius (red square), P. lignaui (blue circle), P. muralewiczi
(green star) and P. mediterraneus (pink triangle).
Polydesmus (s. str.) muralewiczi Lohmander, 1936
Figs 3A, 3H–3J, 4C, 4D, Map 1
Polydesmus complanatus—Muralewicz, 1927: 7 (R).
Polydesmus (Polydesmus) muralewiczi Lohmander, 1936: 9 (D).
Polydesmus (Polydesmus) lignaui muralewiczi—Attems, 1940: 21 (D).
Polydesmus muralewiczi—Lang, 1959: 1790 (M); Kobakhidze, 1965: 391 (R); Lokšina and Golovatch, 1979: 384 (M);
Talikadze, 1984: 143 (R); Zuev, 2014: 348 (R, D).
Material examined. 1 male (lectotype, designated herewith), 1 female (paralectotype, designated herewith),
“Typen” (ZISP), RUSSIA, Kuban Region, Station Psebai, under stones during ascent to Hill Gerisgel, 20.V.1911,
leg. D. Volnukhin; 4 juv. (ZMUM), Krasnodar Prov., Mostovskoy Distr., Shedok, N of Psebai, karst funnel, wet
Fagus, Acer, Quercus etc. forest, 650 m a.s.l., 4.VIII.1986; 1 male (ZMUM), Stavropol Prov., E of Georgievsk,
Quercus, Carpinus etc. forest, litter and under bark, 28 and 31.V.1982; 1 male, 2 females (ZMUM), Stavropol
Prov., Kursavska, Robinia and Rosa hedge along road, 30.V.1982, all leg. S. Golovatch.

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Descriptive notes. Length ca 11–12 mm, width of midbody metazonae 1.2–1.25 mm (male, female),
metazonite to prozonite width ratio < 1.7 (Fig. 3A, 3H–3J). Live coloration usually uniformly light brown, not
reddish, about same as in alcohol (Fig. 3A).
All characters as in P. abchasius (Figs 3A, 3H–3J, 4C, 4D), except as follows. Collum clearly narrower than
head (Fig. 3A, 3H). Paraterga not upturned, subhorizontal, mostly acute-angled and drawn behind rear tergal
margin starting with segment 14 or 15 (Fig. 3A, 3H–3J). Gonopod (Fig. 4C–4D) with a shorter endomere rod and a
less heavily spinulate exomere tip.
Remarks. Although the original description was clearly based only on the male (Lohmander, 1936), the type
series, “Typen”, actually consists of two specimens (male and female) which are to be considered as syntypes. To
secure that the species is based on male material, the male is designated herewith as lectotype, the female as
paralectotype.
Polydesmus (Nomarchus) mediterraneus Daday, 1889
Fig. 5, Map 1
Polydesmus mediterraneus oertzeni—Talikadze, 1984: 143 (R).
Material examined. 3 males, 9 females (ZMUM), ABKHAZIA, Sukhumi Distr., Yashtukha, tree nursery,
29.III.1986, leg. A. Markossian; 2 subadult females (ZMUM), Sukhumi Botanical Garden, litter, 2.X.1978; 1 male
(slide only) (ZMUM), same locality 9.X.1978; all leg. S. Golovatch.
Descriptive notes. Length ca 8–16 mm, width of midbody metazonae 1.5–2.0 mm (male, female), metazonite
to prozonite width ratio < 1.6 (Fig. 5A). Live coloration usually uniformly light brown to brown, about same as in
alcohol (Fig. 5A).
All characters as in P. abchasius (Fig. 5), except as follows. Collum much narrower than head, transversely
oval in shape (Fig. 5A). Paraterga not upturned, subhorizontal, much more strongly rounded laterally, at least until
segment 9 obtuse-angled and drawn behind rear tergal margin starting with segment 14 or 15 (Fig. 5A). Male
prefemora strongly bulging laterad (Fig. 5B). Gonopod (Fig. 5C) with a stout and stump-shaped endomere (en) and
only a slightly curved and apically strongly bifid exomere (ex).
Remarks. The only previous record in the region (Talikadze 1984) concerned “Abkhazia”, without a precise
locality. This eastern Mediterranean species/subspecies is quite common in the adjacent Crimea, apparently being
introduced to the Caucasus. New illustrations are provided to facilitate recognition (Fig. 5).
Polydesmus (s. str.) monticola Latzel, 1884
Polydesmus monticolus (sic!)—Lang, 1959: 1795 (R).
Remark. The record of this eastern Alpine species (Kime and Enghoff 2011) from near Ubinskaya, Severskaya
Distr., Krasnodar Prov. (Lang 1959) is certainly based on a misidentification, likely referred to P. abchasius or P.
lignaui.
Brachydesmus (s. str.) assimilis Lohmander, 1936
Figs 6, 7, Map 3
Brachydesmus (Brachydesmus) assimilis—Lohmander, 1936: 12 (D).
Brachydesmus (Brachydesmus) assimilis—Attems, 1940: 104 (D).
Brachydesmus assimilis—Lang, 1959: 1790 (M); Kobakhidze, 1965: 390 (M); Lokšina and Golovatch, 1979: 384 (M); Zuev,
2014: 349 (R, D).

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FIGURE 5. Polydesmus mediterraneus Daday, 1889: A: male from Gurzuf, Crimea, habitus, dorsal view. B–C: male from
Yashtukha, Abkhazia, leg 7, lateral view, and left gonopod, mesal view, respectively. Designations: en, endomere, ex, exomere.
Picture A by K. Makarov. Scale bars: 0.2 mm (B), 0.1 mm (C), not to scale (A).
Material examined. 1 female, 1 juv. (ZMUM), RUSSIA, Adygea, Sakhrai near Novoprokhladnoye, S of Maikop,
700 m a.s.l., Fagus and Alnus forest, litter, 7.VII.1986; 4 males, 2 females (ZMUM), Stavropol Prov., E of
Novopavlovsk, Quercus, Salix, Alnus etc. forest along stream, 28.V.1982; 8 juv. (ZMUM), Kabardino-Balkaria,
Baxan Valley, Bedyk between Tyrnyauz and Zhankhoteko, 900 m a.s.l., Corylus, Fagus and Carpinus scrub, litter
and under stones, 20.VII.1986; 6 males, 2 females, 1 juv. (ZMUM), Ingushetia, Barsuki near Nazran, Crataegus,
Fraxinus, Rosa etc. bush along road, litter and under stones, 6.VI.1982; 1 female, 2 juv. (ZMUM), Ingushetia, Assa
Valley, ca 9 km SSW of Muzhichi, 800 m a.s.l., Fagus, Alnus, Carpinus etc. forest, litter, under bark and stones,
15.VII.1986; 1 male, 1 female, 5 juv. (ZMUM), Chechnya, Argun Valley, 5 km N of Sovetskoye, 750 m a.s.l.,
Corylus, Fagus, Carpinus etc. forest, litter, under bark and stones, 18.VII.1986; 3 females (ZMUM), Chechnya, 14
km WSW of Gudermes, Quercus forest, litter, 6.VI.1982; 4 juv. (ZMUM), Chechnya, Karachoy SE of Vedeno, 950

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m a.s.l., Fagus, Carpinus etc. forest, litter, under bark and stones, 17.VI.1986; 2 juv. (ZMUM), Dagestan, Botlikh
Distr., Pass Kharami, 2150 m a.s.l., subalpine meadow, 22.IV.1989; 6 juv. (ZMUM), Dagestan, N of Kurush, ca 35
km NNW of Usukhchay, 2500–2800 m a.s.l., alpine meadow, 7–13.VII.1989; 1 male, 1 female, 1 fragment
(ZMUM), Dagestan, Sergokala Distr., near Degva, Quercus forest on slope, litter, 9.VI.1982; 26 males, 46 females,
26 juv. (ZMUM), Dagestan, Upper Gunib, 1700 m a.s.l., Betula and Pinus forest, litter, under bark and stones, 8–
9.VI.1982; 4 males, 4 females, 3 juv. (ZMUM), same locality, 1500–1600 m a.s.l., Betula and Pinus forest,
25.VI.1989; 6 males, 6 females (ZMUM), Dagestan, Chiragchay Valley, Khiv, 900–950 m a.s.l., Carpinus, Fagus,
Corylus, Rhododendron etc. forest, litter and under stones, 24.X.1987; 25 juv. (ZMUM), Dagestan, Akhty Distr.,
above Mirakh, ca 16 km SSW of Usukhchay, 1800 m a.s.l., Betula forest with Salix and Alnus, at timberline,
10.VII.1989; 9 males, 6 females (ZMUM), North Ossetia, 10 km NW of Mozdok, Robinia stand along field, litter,
28.V.1982, all leg. S. Golovatch.
1 female (NHMG), GEORGIA, Tbilisi, 10.VI.1957, leg. H. Lohmander; 13 males, 14 females (ZMUM),
Mariamdjvari Nature Reserve, ENE of Sagaredjo, 1150–1250 m a.s.l., Fagus, Carpinus, Acer, Pinus etc. forest,
litter, under bark and stones, 13–14.V.1987; 2 females (ZMUM), Batsaro Nature Reserve, ca 20 km N of Akhmeta,
800–850 m a.s.l., Fagus, Castanea, Acer etc. forest, litter, 5–6.V.1987; 1 male, 2 females (ZMUM), Pass
Magalakhari between Akhmeta and Tianeti, 1200 m a.s.l., Fagus and Carpinus forest, litter and under bark,
6.V.1987; 1 male, 3 females (ZMUM), N of Kvareli, 700–750 m a.s.l., Fagus, Carpinus, Quercus etc. forest, litter
and under bark, 4.V.1987, all leg. S. Golovatch and K. Eskov; 3 males, 3 juv. (ZMUM), Kazbegi Distr., S of
Gudauri near Pass Krestovyi, 1800 m a.s.l., Rhododendron bush, litter, 4.VI.1982; 12 males, 10 females, 2 juv.
(ZMUM), Lagodekhi Nature Reserve, Fagus, Fraxinus, Acer etc. forest, 600–700 m a.s.l., litter, under bark and
stones, 5–6.V.1983, all leg. S. Golovatch; 1 male, 2 females (HR), Samzhe-Djavachetia, Abastumani
Oberservatorium on Mt Kanobili, 1580 m a.s.l., N41.7544, E42.8203, 28.X.2009, leg. F. Walther; 12 males, 10
females, “Polydesmus maculicornis Brandt” nomen nudum (ZISP), Tiflis (= Tbilisi), date and collector unknown.
4 males, 8 females (ZMUM), ARMENIA, Gegasar ca 5 km W of Spitak, Pambak Valley, Salix forest along
stream, 1650 m a.s.l., 13.XI.1985, leg. S. Golovatch; 2 juv. (ZMUM), Gandakar S of Idjevan, 1000 m a.s.l.,
Quercus and Corylus scrub, 28.V.1987; 1 juv. (ZMUM), Pass Khamza between Idjevan and Berd, 1900 m a.s.l.,
subalpine meadow, 26.V.1987, all leg. S. Golovatch and K. Eskov.
1 male, 5 females (ZMUM), AZERBAIJAN, Perzivan SE of Zakataly, 600 m a.s.l., Quercus, Fagus etc. forest
along stream, litter, 3.V.1987, leg. S. Golovatch and K. Eskov; 1 male, 3 females (ZMUM), Djar near Zakataly,
20.X.1987; 6 males, 9 females, 1 juv. (ZMUM), Perzivan SE of Zakataly, 600 m a.s.l., forest, 19.IV.1987, all leg.
N. Loginova; 1 male (ZMUM), Belakany W of Zakataly, 600 m a.s.l., garden, 24.V.1981; 1 male, 2 females
(ZMUM), Zakataly Nature Reserve, Katekhchay Valley, 700 m a.s.l., 23.V.1981; 1 female, 8 juv. (ZMUM),
Zakataly Nature Reserve, Agkemal, 1800–2100 m a.s.l., forest, 24–27.V.1981; 1 male, 1 female, 1 fragment
(ZMUM), same locality, Katekh-Chay Valley, 23.V.1981, all leg. S. Golovatch and J. Martens; 16 males, 6 females
(ZMUM), above Ajikent, S of Kirovabad (now Ganja), 1400 m a.s.l., meadow, under stones, 3.V.1983, leg. S.
Golovatch; 9 males, 17 females (ZMUM), NW above Bash-Layski ca 20 km NNW of Sheki, 1250 m a.s.l., Fagus,
Carpinus, Acer etc. forest, litter, 3.V.1987; 7 males, 8 females, 4 juv. (ZMUM), ca 5 km N of Kutkashen, 1150–
1200 m a.s.l., Fagus and Carpinus forest, litter and rotten wood, 2.V.1987; 2 males, 1 female (ZMUM), SW of
Kuba, 750 m a.s.l., Fagus, Quercus, Carpinus etc. forest, litter and under bark, 23.IV.1987; 4 females (ZMUM),
Nabran ca 30 km NW of Khachmas, sea-level, Quercus, Carpinus, Acer etc. forest, litter and under bark, 21–
22.IV.1987, all leg. S. Golovatch and K. Eskov; 8 males, 4 females (ZMUM), Laza near Kussary, 10.VIII.1985,
leg. H. Aliev.
Descriptive notes. Adult body with 19 segments (collum plus 16 podous plus one apodous body rings, plus
telson). Length ca 10–13 mm, width of midbody metazonae 1.0–1.5 (male, female), metazonite to prozonite width
ratio < 1.8 (Fig. 6). Live coloration usually uniformly yellowish to brown, about same in alcohol (Fig. 6).
Tegument smooth and moderately shining. Antennae long, reaching behind body segment 3 when stretched
dorsally, slightly clavate. In width, collum << head ≤ segment 2 < 3 < 4 < 5=15 or 16, thereafter body gradually to
quite rapidly tapering towards telson (Fig. 6). Collum transversely ovoid. Paraterga well-developed and lying at
about upper ¼ of body height, mostly subhorizontal to only faintly declivous, never upturned (Fig. 6). Caudolateral
corner of collum broadly rounded, of midbody and following paraterga increasingly acute and pointed, but
especially clearly drawn behind rear tergal margin only in segments (14)15–17 (Fig. 6). Fore margins of paraterga
largely straight, anterolateral corner angular (Fig. 6). Central parts of postcollum metaterga with three transverse

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rows of evident, roundish to polygonal, setigerous bosses or tubercles (Fig. 6). Tergal setae very small, mostly
subclavate, often considerably abraded, usually only a little longer in front row on collum and in caudal row on
segment 19. Paraterga with three or four faint setigerous incisions at lateral margin of poreless and pore-bearing
segments, repectively. Stricture between pro- and metazonae wide, shallow and nearly smooth, at most faintly
rugulose. Limbus microcrenulate-microdentate. Pleurosternal carinae wanting. Epiproct short, conical, subtruncate
at tip, pre-apical lateral papillae evident (Fig. 6). Hypoproct subtrapeziform, paramedian setiferous papillae
evident, well-separated and lying near caudal edge.
FIGURE 6. Habitus of Brachydesmus assimilis Lohmander, 1936. A: male from Belakany, Azerbaijan. B–H: male (B), female
(C–E) and male (F–H) from Upper Gunib, Dagestan, Russia. C, F: anterior part of body. D, G: right half of segment 10. E, H:
caudal part of body. Pictures A and B by K. Makarov, taken not to scale. Line drawings C–H by S. Dashdamirov, scale bar: 1.0
mm.

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FIGURE 7. Gonopods of Brachydesmus assimilis Lohmander, 1936. A–B: male from Mariamdjvari Nature Reserve, Georgia,
lateral and mesal views, respectively. C: male from near Pass Krestovyi, Georgia. D: male syntype from Tbilisi, Georgia (after
Lohmander 1936). Designations: en, endomere; ex, exomere; s, spine on exomere; t, tooth near base of exomere; k, spine at
base of endomere; b, tooth at base of femorite. Scale bar: 0.1 mm (A–C), not to scale (D).

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Legs densely setose, long, in male crassate and longer (Fig. 6A, 6B), prefemora not bulging laterad;
postfemora sometimes, tibiae and tarsi always with sphaerotrichomes ventrally.
Gonopods (Fig. 7) subfalcate, bipartite, exomere (ex) axe-shaped, suberect, its acuminate apex clearly curved
ventrad, with a strong distomesal spine (s) and a midway ventral tooth (t); endomere (en) short and stout, stump-
shaped, with only a very short finger-shaped rod placed distal to a well-developed hairy pad; another, ventral spine
(k) located at base of en; still one more, ventral tooth (b) usually present at base of femorite.
Remark. This species shows some variation not only in body size and in the shape of paraterga, but also in
gonopod structure (Fig. 7).
Brachydesmus (s. str.) pigmentatus Attems, 1951
Fig. 8, Map 3
Brachydesmus pigmentatus Attems, 1951: 407 (D).
Brachydesmus pigmentatus—Golovatch, 1981b: 426 (D); Enghoff and Moravvej, 2005: 69 (R).
Brachydesmus pereliae Golovatch, 1976: 1567 (D), syn. n.
Brachydesmus pereliae—Lokšina and Golovatch, 1979: 384 (M); Golovatch, 1979: 1323 (M); Read and Golovatch, 1994: 65
(M).
Material examined. 1 male (ZMUM), AZERBAIJAN, Talysh Mts, Farm Azerbaijan ca 10 km SE of Lerik, 550
m a.s.l., Quercus, Carpinus, Acer etc. forest along stream, litter and rotten wood, 12.X.1983; 4 juv. (ZMUM),
Talysh Mts., Lerik Distr., Zuvand, foot of Mt Kyumyurkey, Oranchay Valley, xerophytous montane grassland,
2100–2200 m a.s.l., under stones, 9.X.1983, all leg. S. Golovatch; 1 juv. (ZMUM), Istisu SW of Astara, Quercus,
Parrotia, Carpinus, Acer etc. forest, X.1978, leg. A. Druk; 1 male (ZMUM), Lenkoran Distr., Hyrcan Nature
Reserve, forest, rotten wood, 19.III.1971, leg. N. Loginova; 1 male (ZMUM), Lenkoran Distr., Hyrcan Nature
Reserve, litter, 26.I.–4.II.1985; 2 males, 2 females (ZMUM), same locality and data, all leg. A. Druk; 1 female
(ZMUM), Azerbaijan, Lenkoran Distr., Hyrcan Nature Reserve, near Alekseevka, 21.IV.1985; 1 male, 2 females
(ZMUM), same locality, Khan-Bulan River, 22.IV.1985; 1 female (ZMUM), same locality, Avrora, lowland forest,
21.IV.1985, all leg. E. Kupriyanova; 1 female (ZMUM), same locality, forest, litter, 26.I.–4.II.1985, leg. A. Druk; 1
female, 2 juv. (ZMUM), same locality, 50 m a.s.l., Quercus, Parrotia, Carpinus etc. forest, 13.X.1983; 4 males
(ZMUM), same locality, 8 km SW of Alekseevka, Apo, Quercus, Acer, Carpinus, Parrotia etc. forest, litter, under
stones and bark, 16.X.1983; 4 females (ZMUM), same locality, Apo ca 6 km SW of Alekseevka, Quercus, Acer,
Carpinus, Parrotia etc. forest, litter, under stones and bark, 14 and 16.X.1983; 1 female (ZMUM), Lenkoran Distr.,
Avrora, Moscow Forest, 50 m a.s.l., 1.VI.1996; 1 female (ZMUM), Lenkoran, Apo below Bilasar, 350 m a.s.l., 8–
9.VI.1996, all leg. S. Golovatch.
Descriptive notes. Length ca 13–21 mm, width of midbody metazonae 1.2–2.1 mm (male, female), but the
above samples ranging between 13 and 16 mm in length, and 1.8 and 2.1 mm in width; metazonite to prozonite
width ratio < 1.6 (Fig. 8A–8D). Live coloration usually uniformly grey-brown to brown, in alcohol sometimes
faded to light brown (Fig. 8A).
All characters as in B. assimilis (Fig. 8), except as follows. Paraterga subhorizontal to poorly declivous, mostly
only slightly rounded laterally; caudolateral corner obtuse-angled only until segment 6, pointed or nearly pointed
and increasingly well drawn behind rear tergal margin starting with segment 8 or 9 (Fig. 8A–8D). Fore margins of
paraterga mostly straight, anterolateral corner usually angular. Tergal setae a little longer, largely pointed. Male
prefemora strongly bulging laterad. Gonopod (Fig. 8E, 8F) subfalcate, bipartite, with a long, very simple, finger-
shaped, ventrad only slightly and regularly curved, apically acuminate exomere (ex), this being tightly appressed
ventrally to a similar, also finger-shaped, but somewhat shorter endomere (en), the latter at base carrying a small,
medioventral spine (k); en representing a solenomere devoid of a hairy pulvillus; a small tooth (t) located
medioventrally at base of femorite.
Remarks. This species has hitherto been considered as subendemic to Hyrcania within N and NW Iran alone
(Elburs and Zagros mountains) (Attems 1951; Golovatch 1981b). The above records are thus new to the fauna of
the Republic of Azerbaijan, in general also to that of the Caucasus.
Given such a considerable range of variation in body size in B. pigmentatus, we do not hesitate to formally
synonymize here this species with B. pereliae Golovatch, 1976, the latter taxon known from the W Kopetdagh

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Mountains, SW Turkmenistan. Indeed, the holotype of B. pereliae was said to be 10 mm long and 1.17 mm wide,
whereas the gonopods of B. pigmentatus and B. pereliae have been noted as being virtually identical (Golovatch
1976). Hence the new synonymy advanced: B. pigmentatus Attems, 1951 = B. pereliae Golovatch, 1976, syn. n.
FIGURE 8. Brachydesmus pigmentatus Attems, 1951, males from Hyrcan Nature Reserve, Azerbaijan. A: habitus, entire
body, dorsal view. B: anterior part of body, dorsal view. C: right half of metatergite 10, dorsal view. D: posterior part of body,
dorsal view. E: gonopods in situ, ventral view. F: right gonopod, mesal view. Designations: en, endomere; ex, exomere; k,
spine at base of endomere; t, tooth near base of femorite. Pictures A and E by K. Makarov, taken not to scale. Line drawings B–
D by S. Dashdamirov, scale bar: 1.0 mm (B–D), 0.3 mm (F).

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Brachydesmus (Eubrachydesmus) superus Latzel, 1884
Fig. 9, Map 3
Brachydesmus superus—Muralewicz, 1907: 340, 349 (R); 1927: 6 (R); Zuev, 2014: 349 (R).
FIGURE 9. Brachydesmus superus Latzel, 1884. A: habitus, entire body, dorsal view, male from Stavropol, Russia. B: left
gonopod, male from Zheleznovodsk, Stavropol Prov., Russia. Designation: pu, hairy pulvillus. Picture A taken by R. Zuev.
Scale bars: 1.0 mm (A), 0.1 mm (B).

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Material examined. 5 males, 11 females (ZMUM), RUSSIA, Stavropol Prov., Zheleznovodsk, park at foot of Mt
Zheleznaya, litter and under stones, 30.V.1982, leg. S. Golovatch.
5 males, 7 females (ZMUM), ABKHAZIA, Nizhnyaya Yashtukha near Sukhumi, tobacco plantation,
29.III.1987, leg. A. Markossian.
Descriptive notes. Length ca 6.5–10 mm, width of midbody metazonae 0.8–1.2 mm (male, female),
metazonite to prozonite width ratio < 1.6 (Fig. 9A). Live coloration usually uniformly light grey-brown to pallid.
All characters as in B. assimilis (Fig. 9), except as follows. Tegument particularly shiny (Fig. 9A). Paraterga
mostly poorly declivous, only slightly rounded laterally; caudolateral corner obtuse-angled only until segment 6,
pointed or nearly pointed and increasingly well drawn behind rear tergal margin starting with segment 11 or 12
(Fig. 9A). Fore margins of paraterga largely straight, anterolateral corner angular. Tergal setae a little longer,
mostly sharp. Male prefemora strongly bulging laterad. Gonopod (Fig. 9B) unipartite, slender, only slightly falcate,
ventrally with several teeth or spines and a prominent hairy pulvillus (pu) in distal part.
Remarks. Zuev’s (2014) claim for the first record of this ubiquitous anthropochore species in the Caucasus (at
Stavropol) is wrong, as he had overlooked Muralewicz’s (1907, 1927) reports from Vladikavkaz, from between
Balta and Lars, N Ossetia, and from Sukhumi, Abkhazia. However, those two latter records are not mapped here
pending verification (Map 3).
Brachydesmus (Lophobrachydesmus) furcatus Lohmander, 1936
Fig. 10, Map 3
Brachydesmus (Lophobrachydesmus) furcatus Lohmander, 1936: 16 (D).
Brachydesmus (Lophobrachydesmus) furcatus—Attems, 1940: 134 (D); Lang, 1959: 1795 (R).
Brachydesmus furcatus furcatus—Strasser, 1970: 204 (D); Lokšina and Golovatch, 1979: 384 (M); Talikadze, 1984: 143 (R).
Brachydesmus (Lophobrachydesmus) furcatus exiguus Strasser, 1970: 204 (D), syn. n.
Brachydesmus furcatus exiguus—Lokšina and Golovatch, 1979: 384 (M); Talikadze, 1984: 143 (R).
Brachydesmus furcatus—Chumachenko, 2012: 452 (R);
Material examined. 1 male, 5 juv. (ZMUM), RUSSIA, Krasnodar Prov., Gelendjik, Mikhailovskii Pass, Fagus
forest, 26.V.1956; 3 females (ZMUM), same locality and Pass, Fagus forest, 5.VII.1956; 1 female, 3 juv. (ZMUM),
same locality, Fagus forest, under bark, 26.V.1956; 1 female, 2 juv. (ZMUM), Gelendjik, forest, 28.IV.1955; 1
female (ZMUM), Krasnodar Prov., Tuapse Distr., Georgievskoe Forestry, Castanea forest, 7.V.1954; 1 female
(ZMUM), same locality, SSW slope, Quercus forest, 18.V.1954; 3 males, 6 juv. (ZMUM), Krasnodar Prov., Tuapse
Distr., Cherepsinskoye Forestry, Quersus forest, S slope, 15.V.1954, all leg. M. Ghilarov, all det. H. Lohmander,
det. C. Dziadosz, 1968; 1 female (ZMUM), Sochi, Khosta, Buxus forest, 29.VI.1956, leg. M. Ghilarov; 2 females
(ZMUM), same locality, Caucasian Nature Reserve, Tax us and Buxus forest, 15.V.1985, leg. S. Golovatch; 1 male,
1 female (ZMUM), same locality, Taxus and Buxus forest, III.2006, leg. Yu. Chumachenko; 1 female (ZMUM),
Adygea, Guzeripl, Caucasian Nature Reserve, Pasture Abago S of Guzeripl, 1700–1850 m a.s.l., Abies and Fagus
forest, Rhododendron thicket, litter, 24–26.V.1985, leg. S. Golovatch; 1 male, 2 females (AE), Adygea, near
Guzeripl, Abies forest, 19.VI.2011, leg. D. Khisametdinova and E. Bortnikov; 1 female (ZMUM), Krasnodar Prov.,
Sochi, Krasnaya Polyana, 6–8.VI.1976; 1 female (ZMUM), same locality, 1900 m a.s.l., under stones, 29.VI.1956;
1 female (ZMUM), same locality, 600 m a.s.l., Castanea forest, litter, 1.VI.1956, all leg. M. Ghilarov; 1 female
(ZMUM), Krasnodar Prov., Sochi, Dagomys, 250 m a.s.l., Quercus, Corylus, Fagus etc. scrub, litter, logs,
18.V.1983; 2 females (ZMUM), Krasnodar Prov., Sochi, Caucasian Nature Reserve, Pslukh ca 20 km E of
Krasnaya Polyana, Mt Aishkho-2, S slope, 1600 m a.s.l., Abies, Fagus, Betula etc. forest, 12.VIII.1986; 1 male, 2
females (ZMUM), same locality, Mt Kogot, Abies and Fagus forest up to timberline, 1650–1850 m a.s.l., litter,
under bark and stones, 18–20.V.1985; 1 female (ZMUM), Krasnodar Prov., Caucasian Nature Reserve, Krasnaya
Polyana, 6–8.VI.1978, all leg. S. Golovatch; 2 females, 39 juv. (ZMUM), Krasnodar Prov., Sochi, Mt Bolshoy
Akhun, Cave Akhunskaya, 19.VII.2015, leg. I. Turbanov; 9 juv. (ZMUM), Krasnodar Prov., Kabardinka, scrub,
2.VI.1956, leg. M. Ghilarov; 9 juv. (ZMUM), Krasnodar Prov., Severskaya Distr., 2–10 km S of Ubinskaya, 300–
450 m a.s.l., Quercus, Fagus, Carpinus etc. forest, litter and under bark, 3–4.VII.1986; 2 females, 22 juv. (ZMUM),
same locality, Mt Derbiy ca 15 km SW of Ubinskaya, 800–850 m a.s.l., Quercus, Fagus, Fraxinus etc. forest, litter
and under bark, 2.VII.1986, all leg. S. Golovatch; 6 juv. (ZMUM), same locality, Ubinskoye Forestry,

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FIGURE 10. Brachydesmus furcatus Lohmander, 1936. A: habitus, entire body, dorsal view, male from Mount Kogot,
Caucasian Nature Reserve, Krasnodar Prov., Russia. B–D: male from near Goryachiy Klyuch, Krasnodar Prov., Russia, B:
anterior part of body, dorsal view; C: right half of metatergite 10, dorsal view; D: posterior part of body, dorsal view. E–F:
male holotype of B. furcatus, right and left gonopods, lateral and mesal views, respectively (after Lohmander 1936); G: male
holotype of B. furcatus exiguus Strasser, 1970, left gonopod, lateral view (after Strasser 1970). Designations: en, endomere; ex,
exomere; t, tooth at base of femorite. Picture A by K. Makarov, taken not to scale. Line drawings B–D by S. Dashdamirov,
scale bar: 1.0 mm (B–D), not to scale (E–G).
Quercus forest, 16.VI.1953, leg. M. Ghilarov; 2 males, 3 females (ZMUM), Krasnodar Prov., Goryachiy Klyuch,
Cave Fanagoriyskaya, ca 12 km SW of Fanagoriyskoye, 30.X.1981; 1 male, 3 females, 3 juv. (ZMUM), same
locality, Fagus, Acer, Picea etc. forest, litter and under logs, 19.V.1983, all leg. S. Golovatch; 1 female (ZMUM),
Krasnodar Prov., Goryachiy Klyuch, without data; 1 male (ZMUM), same locality, Kotkh Mt. Range, Quercus
forest, soil, 18.V.1956; 11 juv. (ZMUM), same locality, Pshaf Mt. Range, forest on limestone, 20.V.1956; 3 females
(ZMUM), Pshaf Mt. Range, near “Belaya Latka”, Quercus forest, 17.VI.1956; 3 females, 8 juv. (ZMUM), Pshaf

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Mt. Range, near “Belaya Latka”, S slope, Quercus forest, 18.VI.1956; 8 females, 18 juv. (ZMUM), same locality,
Kotkh Mt. Range, summit, S slope, Quercus forest, 20.VI.1956; 2 females, 2 juv. (ZMUM), Kotkh Mt. Range,
summit, S slope, Quercus forest, 21.VI.1956; 1 male, 1 female (ZMUM), Kotkh Mt. Range, Quercus forest,
18.VI.1956; 1 female (ZMUM), same locality, Canyon Bukovoe, 5.IV.1954; 7 juv. (ZMUM), Bolnichi, Mt
Petushok, NE slope, Carpinus forest, 4.VIII.1955; 1 juv. (ZMUM), Krasnodar Prov., Mt Ostraya, 1.VIII.1954; 1
female (ZMUM), same locality, 3.VIII.1954; 1 juv. (ZMUM), Kuban, Azovskaya, under Tkachovskoye, Quercus
forest belt, 24.VII.1955, all leg. M. Ghilarov, all det. H. Lohmander, det. C. Dziadosz, 1968.
2 males, 3 females (ZMUM), ABKHAZIA, Myussera Nature Reserve, 20–130 m a.s.l., Castanea, Alnus etc.
forest, 8–10.IV.1983, leg. S. Golovatch; 1 male (SMNG 14685), Transcaucasus, Riza-See (= Lake Ritsa),
20.VI.1984, leg. W. Dunger.
Descriptive notes. Length ca 8.5–12 mm, width of midbody metazonae 0.8–1.3 mm (male, female),
metazonite to prozonite width ratio < 1.4 (Fig. 10A–10D). Live coloration usually uniformly pallid to light brown.
All characters as in B. assimilis (Fig. 10), except as follows. Paraterga poorly-developed, their fore margins
and anterolateral corner largely broadly and regularly rounded, not angular; caudolateral corner mostly obtuse-
angled, pointed or nearly pointed and drawn behind rear tergal margin only in segments 16–18 (Fig. 10A–10D).
Tergal setae mostly very short and subbacilliform. Metatergal sculpture especially superficial. Male prefemora not
bulging laterad. Gonopod (Fig. 10E–10G) bipartite, strongly falcate, much as in B. pigmentatus, but exomere (ex)
with a conspicuous densely pilose strip, endomere (en) simple and bare, while tooth (t) at base of femorite very
prominent and strongly rounded.
Remarks. The differences of B. furcatus exiguus Strasser (1970), described from a cave near Sochi, such as a
slightly smaller and narrower body, a somewhat shorter endomere, a smaller pilose strip on the exomere and a less
prominent tooth at the base of the gonofemorite (Fig. 10G) as compared to the sympatric nominate subspecies,
seem to represent only individual variations. Hence we do not hesitate to formally synonymize B. furcatus exiguus
under B. furcatus, syn. n. Moreover, the name B. f. exiguus Strasser, 1970 has long been jeopardized by the much
older homonym, B. exiguus Brölemann, 1894, from Italy (Brölemann 1894).
Given the striking similarities observed in gonopod structure between B. (s. str.) pigmentatus and B.
(Lophobrachydesmus) furcatus, the status of the latter subgenus is to be questioned.
Brachydesmus (Haplobrachidesmus) kalischewskyi Lignau, 1915
Figs 11–27, Map 2
Brachydesmus kalischewskyi Lignau, 1915: 370 (D).
Brachydesmus (Haplobrachydesmus) (sic!) kalischewskyi—Lohmander, 1936: 18 (R. D); Attems, 1940: 137 (D); Lang, 1959:
1790 (M); Lokšina and Golovatch, 1979: 384 (M).
Brachydesmus kalischewskyi—Kobakhidze, 1965: 390 (R); Golovatch, 1981a: 110 (R); Talikadze, 1984: 143 (R); Enghoff,
2006: 192 (R); Chumachenko, 2012: 452 (R); Barjadze et al., 2015: 21 (R).
Brachydesmus karawajewi Lohmander, 1928: 530 (D), syn. n.
Brachydesmus (Haplobrachydesmus) (sic!) karawajewi—Lohmander, 1936: 28 (R, D).
Brachydesmus (Haplobrachydesmus) karawajewi—Attems, 1940: 136 (D); Lokšina and Golovatch, 1979: 384 (M).
Brachydesmus karawajewi—Lang, 1959: 1790 (M); Kobakhidze, 1965: 390 (M); Talikadze, 1984: 143 (R); Chumachenko,
2012: 452 (R); Zuev, 2014: 348 (R, D).
Brachydesmus karawaewi (sic!)—Bababekova, 1996: 90 (R).
Brachydesmus (Haplobrachydesmus) (sic!) ferrugineus—Lohmander, 1936: 22 (D), syn. n.
Brachydesmus (Haplobrachydesmus) ferrugineus—Attems, 1940: 137 (D); Lang, 1959: 1790 (M); Lokšina and Golovatch,
1979: 384 (M).
Brachydesmus ferrugineus—Kobakhidze, 1965: 390 (R); Ceuca, 1975: 91 (D); Golovatch, 1981a: 110 (R); 1981b: 426 (D);
Talikadze, 1984: 143 (R); Loginova, 1993: 151 (R); 1995: 103 (R); Enghoff and Moravvej, 2005: 69 (R); Enghoff, 2006:
192 (R); Kokhia, 2011: 28, 50, 80 (R); Barjadze et al., 2015: 21 (R).
Brachydesmus (Haplobrachydesmus) (sic!) talyschanus Lohmander, 1936: 26 (D), syn. n.
Brachydesmus (Haplobrachydesmus) talyschanus—Attems, 1940: 137 (D); Lang, 1959: 1790 (M); Lokšina and Golovatch,
1979: 384 (M).
Brachydesmus talyschanus—Kobakhidze, 1965: 391 (R); Rakhmanov, 1972 : 116 (R); Samedov et al., 1972: 1245 (R);
Rasulova and Rakhmanov, 1973: 520 (R); Talikadze, 1984: 143 (R); Bababekova, 1996: 90 (R).
Brachydesmus (Haplobrachydesmus) bidentatus Golovatch, 1976: 1568 (D), syn. n.
Brachydesmus bidentatus—Lokšina and Golovatch, 1979: 384 (M); Bababekova, 1996: 90 (R).

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FIGURE 11. Size differences in a syntopic population of morphs A (larger) and B (smaller) of Brachydesmus kalischewskyi
Lignau, 1915, males from Shatili, Argun River valley, Georgia. Scale bar: 1.0 mm. Picture by H. Reip.
Material examined. 3 males, 2 females (ZMUM), RUSSIA, Krasnodar Prov., Tuapse Distr., 15 km SE of
Novomikhailovskii, Psebe, deciduous forest, litter, under bark and stones, 29.X.1981; 1 female (ZMUM),
Krasnodar Prov., Sochi, Dagomys, Quercus forest, litter, 28.X.1981; 1 male, 1 female (ZMUM), Krasnodar Prov.,
Sochi, Lazarevskoye, Saloniki, deciduous forest, litter, 28.X.1981; 3 males (ZMUM), Krasnodar Prov., Sochi,
Khosta, Taxu s and Buxus forest, litter, 28.X.1981, all leg. S. Golovatch; 1 male, 2 females (ZMUM), same locality,
Taxu s and Buxus forest, III.2006, leg. Yu. Chumachenko; 2 females, 1 subadult female (ZMUM), Krasnodar Prov.,
Sochi, 5.5 km NE of Krasnaya Polyana, lower course of Achipse River, ca 600 m a.s.l., N43°43’10”, E40°15’25”,
19–23.VIII.2014, leg. K. Makarov and A. Matalin; 2 males, 5 females (ZMUM), Krasnodar Prov., Aibga Mt.
Range, sources of Tikhaya Rechka River, 1700–1900 m a.s.l., N43.638º, E40.2481º, 17.06.1997, leg. I. Belousov; 1
male (ZMUM), Krasnodar Prov., Tuapse Distr., Georgievskoe Forestry, Castanea forest, 6.V.1954, leg. K. Arnoldi,
det. H. Lohmander, det. C. Dziadosz, 1968; 3 juv. (ZMUM), Krasnodar Prov., Caucasian Nature Reserve, Pslukh,
ca 20 km E of Krasnaya Polyana, 1000 m a.s.l., Fagus, Quercus, Abies, Picea etc. forest, litter, 11–12.VIII.1986;
5 males, 3 females, 3 juv. (ZMUM), Krasnodar Prov., Goryachiy Klyuch, 12 km SW of Fanagoriyskoye, Cave
Fanagoriyskaya, 19.V.1983; 7 juv. (ZMUM), Krasnodar Prov., Mostovskoy Distr., Yaroslavskaya, 300 m a.s.l.,
Quercus, Acer, Corylus etc. scrub, 4.VIII.1986, all leg. S. Golovatch; 1 female (ZMUM), Krasnodar Prov., Tuapse
Distr., Georgievskoye Forestry, Kazachye, forest, 7.V.1954, leg. M. Ghilarov; 1 female (ZMUM), Adygea, Maikop,
garden, V.2011; 1 male, 1 male fragment (without segments 14–19), 1 male fragment (without head and segments
1–10) (ZMUM), Adygea, Caucasian Nature Reserve, Pasture Abago S of Guzeripl, 1800 m a.s.l., subalpine
meadow, pitfall traps, 3–29.VIII.2009, all leg. Yu. Chumachenko; 1 male, 2 females, 1 juv. (SMNG 34137),
Adygea, Koryto, 22.VIII.2005; 1 male (SMNG 34144), Adygea, Polyana Shostakova, 25.VIII.2005; 3 males
(SMNG 34158), Adygea, Sakhrai, Kuna Valley, 30.VIII.2005, all leg. K. Voigtländer. 2 females (AE), Krasnodar
Prov., Khadyzhensk, 15.VI.2007, leg. A. Evsyukov and D. Khisametdinova; 4 females (ZMUM), same locality,
1350–1400 m a.s.l., Abies and Fagus forest, litter, under bark and stones, 24–26.V.1985; 5 females (ZMUM),
same locality, 1700–1850 m a.s.l., Abies and Fagus forest, Rhododendron thicket, litter, 24–26.V.1985; 2 females
(ZMUM), Krasnodar Prov., Caucasian Nature Reserve, Mt Aishkho-2 E of Pslukh, ca 28 km E of Krasnaya
Polyana, S slope, 1600 m a.s.l., Abies, Picea, Fagus, Betula etc. forest, litter, 12.VIII.1986; 1 male, 1 female
(ZMUM), Karachaevo-Cherkessia, ca 30 km S of Kurdjinovo, Laba Valley, 4 km N of Damkhurts, 1050–1100 m
a.s.l., Fagus, Acer, Picea etc. forest, litter and under bark, 4.VIII.1986; 1 juv. (ZMUM), Karachaevo-Cherkessia,
Teberda Nature Reserve, above Teberda, Mt Malaya Khatipara, 2400 m a.s.l., subalpine meadow, under stones, 29–
30.V.1985; 1 female (ZMUM), same locality, Kizgich Canyon, N of Arkhyz, 1550–1600 m a.s.l., Abies, Picea,
Pinus, Fagus, Betula etc. forest, litter, under bark and stones, 5.VI.1985; 27 males, 1 female, 3 juv. (ZMUM),

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

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FIGURE 13. Morph A of Brachydesmus kalischewskyi Lignau, 1915: A–C: female from Pass Ertso, South Ossetia, anterior
part of body, right half of metatergite 10 and caudal part of body, respectively, dorsal views. D–F: male from Pass Ertso, South
Ossetia, anterior part of body, right half of metatergite 10 and caudal part of body, respectively, dorsal views. G–H: gonopods
of a male from near Batumi, Ajaria, Georgia, ventral and mesal views, respectively. Designations: en, endomere; ex, exomere;
i, mesal apical tooth of exomere; e, lateral apical tooth of exomere; l, subapical lobe of exomere. Line drawings A–F by S.
Dashdamirov; G–H after Lohmander (1936). Scale bar: 1.0 mm (A–F), not to scale (G–H).
Karachaevo-Cherkessia, Lower Teberda S of Karachaevsk, 1000 m a.s.l., Quercus, Fagus, Acer etc. forest, litter,
3.VIII.1986, all leg. S. Golovatch; 2 females (ZMUM), Karachaevo-Cherkessia, Teberda, ca 1200–2000 m a.s.l.,
VI–VII.1908, leg. A.P. Zolotarev; 31 juv. (ZMUM), Kabardino-Balkaria, Lower Chegem, 800 m a.s.l., Fagus
forest, litter, 14.VII.1986; 18 males, 2 females (ZMUM), North Ossetia, Novo-Georgievskoye near Mozdok,
Quercus, Acer, Alnus, Fraxinus etc. forest, floodplain of Terek River, litter, logs, under stones, 27.V.1982; 1 male, 7
females, 6 juv. (ZMUM), North Ossetia, S of Ordzhonikidze (= Vladikavkaz), between Chmi and Baltik, Quercus
and Alnus scrub on slope, litter and under stones 2.VI.1982; 12 males, 5 females (ZMUM), North Ossetia, S of
Alagir, 200 m a.s.l., Acer, Fagus etc. forest, litter, 17.X.1987; 3 males, 4 females, 9 juv. (ZMUM), Ingushetia, Assa
Valley, ca 9 km SSW of Muzhichi, 800 m a.s.l., Fagus, Alnus, Carpinus etc. forest, litter, under bark and stones,
15.VII.1986; 2 females, 2 juv. (ZMUM), Chechnya, 5 km W of Shaami-Yurt, ca 26 km W of Groznyi, Acer forest,
litter, 6.VI.1982; 1 female (ZMUM), Dagestan, Chiragchay Valley, Sardarkent between Kasumkent and Khiv, 650

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m a.s.l., Crataegus, Rosa, Carpinus, Juglans etc. scrub, litter, 24.X.1987; 1 male (ZMUM), Dagestan, Samur
Valley, Garakh ca 35 km SW of Magaramkent, 700–800 m a.s.l., Crataegus, Quercus, Acer, Rosa etc. scrub, litter,
23.X.1987, all leg. S. Golovatch; 10 males, 8 females (ZMUM), Dagestan, Kaytag Distr., Karatsan, 6.X.2012, leg.
P. Gamilova.
1 female (ZISP), SOUTH OSSETIA, near Kvaisi, under stones, 1700 m a.s.l., 19.X.1956, leg. K. Gorodkov; 1
male, 1 female (ZMUM), 15 km E of Kvaisi, Surami Mt. Ridge, Pass Ertso, ca 1000 m a.s.l., deciduous brush,
along stream, litter and stones, 20.X.1981; 4 males, 2 females (ZMUM), Lower Roki S of Pass Rokskii, 1500 m
a.s.l., Fagus, Betula etc. forest, litter, 17.X.1987, all leg. S. Golovatch.
FIGURE 14. Morph B of Brachydesmus kalischewskyi Lignau, 1915: A: male from Yarymdja, E Caucasus Major, Azerbaijan,
habitus, lateral view. B: male from Khosrov Nature Reserve, Caucasus Minor, S Armenia, habitus, dorsal view. C: left gonopod
of male holotype of B. karawajewi Lohmander, 1928, from Teberda, Karachaevo-Cherkessia, Russia, mesal view. D–E: male
from Verkhniy Agdan (now Gandzakar), Idjevan Distr., Armenia (not Georgia, as mistakenly stated by Lohmander 1936),
ventral and mesal views, respectively. Designations: en, endomere; ex, exomere; i, mesal subapical tooth of exomere; e, lateral
apical tooth of exomere; l, subapical lobe of exomere. Pictures A and B by K. Makarov; C–E after Lohmander (1928, 1936).
Not to scale.

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

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FIGURE 16. Morph C of Brachydesmus kalischewskyi Lignau, 1915. A–C: male syntype of B. ferrugineus Lohmander, 1936
from near Borzhomi, Georgia, anterior part of body, right half of metatergite 10 and caudal part of body, respectively, dorsal
views. D–F: male syntype of B. ferrugineus Lohmander, 1936 from Novyi Afon, Abkhazia, anterior part of body, right half of
metatergite 10 and caudal part of body, respectively, dorsal views. G–H: gonopods of syntype of B. ferrugineus from Novyi
Afon, Abkhazia, ventral and mesal views, respectively. Designations: en, endomere; ex, exomere; i, mesal apical tooth of
exomere; e, lateral apical tooth of exomere; l, subapical lobe of exomere. Line drawings A–F by S. Dashdamirov; G–H after
Lohmander (1936). Scale bar: 1.0 mm (A–F), not to scale (G–H).
2 males, syntypes of B. ferrugineus (ZISP), ABKHAZIA, “W Transcaucasia, Olginskoe, Tsebelda, 20.V.2012,
leg. V. Lindholm”; 1 male, syntype of B. ferrugineus (gonopods missing) (ZISP), “Black Sea Gouvernement,
Novyi Afon, 12.V.1912, leg. V. Lindholm”; 1 female (ZISP), “1 female juv., Polydesmus sp., det. C. Dziadosz,
1967”, Gagra, Mamsdyshkha, ca 2000 m, VIII.1911, leg. V. Isaev?; 1 female (ZMUM), Lake Ritsa, 1800 m a.s.l.,
Abies and Fagus forest, litter, 13.IX.1985, leg. I. Ushakov; 2 males, 1 female, 1 juv. (ZMUM), Lake Ritsa, 950–
1100 m a.s.l., Fagus, Abies, Picea, Acer etc. forest, litter, under bark and stones, 13–14.VIII.1986, leg. S.
Golovatch; 1 male, 1 juv. (SMNG 14685), Transcaucasus, Riza-See (= Lake Ritsa), 20.VI.1984, leg. Dunger; 1
female (ZMUM), Sukhumi Distr., near Cave Kelassuri, litter, 11.IV.1983; 1 male (ZMUM), same locality, near

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Verkhnyaya Kelassuri, deciduous bush on rock, slope, litter, 27.X.1981, all leg. S. Golovatch; 6 males (ZMUM),
Shroma near Sukhumi, 27.IV.1971, leg. N. Zalesskaya; 1 male (ZMUM), Sukhumi, I.1986, leg. V. Dolin; 1 juv.
(ZMUM), Sukhumi Distr., Lake Amtkel, ca 16 km N of Tsebelda, 550 m a.s.l., Alnus forest, litter and under bark,
19.VIII.1986; 1 juv. (ZMUM), Sukhumi Distr., Tsebelda, 300 m a.s.l., Carpinus, Acer and Buxus scrub, litter,
19.VIII.1986; 2 males, 1 juv. (ZMUM), Sukhumi Botanical Garden, 9.X.1978, leg. S. Golovatch; 1 female, 1 juv.
(ZMUM), 2 km N of Duripsh N of Gudauta, Fagus log, under bark, 8.IX.1985, leg. I. Ushakov; 1 female (ZMUM),
Shroma near Sukhumi, 2.V.1971, leg. N. Zalesskaya; 1 male (ZMUM), Sukhumi, Canyon Alekseevskoe, in house,
3.XII.1953; 1 female (ZMUM), same locality, 8.X.1953, all leg. E. Borutzky; 1 male, 1 female (ZMUM), Sukhumi,
University Campus, N42°58,4’ E41°04,04’, ca 50 m a.s.l., Encopytus, tree shaking, litter, 7.XII.2003, leg. Yu.
Marusik and G. Antonova; 1 male (ZMUM), Myussera Nature Reserve, 1.II.1979, leg. L. Malinina.
1 male, syntype of B. ferrugineus (ZISP), GEORGIA, “Tiflis Gouvernement, Canyon Banis-khevi-tskhali
[near Borzhomi], 13.VI.1909, leg. L. Berg”; 1 male (ZMUM), Lanchkhuti Distr., upper reaches of Supsa River,
15.X.1983; 2 juv. (ZMUM), Chokhatauri Distr., Bakhmaro ca 40 km SSW of Nabeglavi, 2000 m a.s.l., IV.1984; 3
females (ZMUM), Ajaria, Kintrish Nature Reserve, Zeraboseli, Khekhpara Valley, 12.X.1984, all leg. E.
Kvavadze; 21 males, 13 females (ZMUM), same locality, 450–600 m a.s.l., deciduous forest, litter and under
stones, 13.X.1981; 3 females (ZMUM), same locality, 800 m a.s.l., Rhododendron thicket, litter, 13.X.1981, all leg.
S. Golovatch; 5 males, 6 females (ZMUM), same locality, 450–600 m a.s.l., deciduous forest, 1–3.VI.1981; 2
males, 1 female (ZMUM), same locality, 600–700 m a.s.l., Rhododendron thicket, 2.VI.1981, all leg. S. Golovatch
and J. Martens; 17 juv. (ZMUM), Ajaria, between Kobuleti and Ureki, seashore Pinus forest, litter, 5 m a.s.l.,
N41º55’58.1", E41º45’53.0", 21.07.2012, leg. Y.M. Marusik; 5 males, 1 female, 1 juv. (ZMUM), Ajaria, Keda
Distr., Magutseti, Platanus forest, litter, 9.X.1981; 9 males, 2 females, 2 juv. (ZMUM), Ajaria, Khulo, 900 m a.s.l.,
Quercus, Abies, Alnus etc. forest, litter, 11.X.1981; 4 juv. (ZMUM), Ajaria, E of Khulo, 950 m a.s.l., deciduous
forest on rock, near spring, litter, 10.X.1981; 1 juv. (ZMUM), Ajaria, 6 km W of Khulo, 800 m a.s.l., deciduous
forest on rocky slope, litter, 11.X.1981, all leg. S. Golovatch; 1 male, 1 female (ZMUM), Ajaria, Tsikhisdziri N of
Batumi, 100 m a.s.l., gardens, 30.V.1981; 7 males, 20 females (ZMUM), Ajaria, Batumi Botanical Garden, 20–150
m a.s.l., 30.V.–7.VI.1981, all leg. S. Golovatch and J. Martens; 3 males, 1 juv. (ZMUM), Ajaria, Meskheti Mt.
Range, Pass Goderdzi, 1800 m a.s.l., Picea forest, under stones, litter, umder bark, 2.IX.1999, leg. L. Penev; 1 juv.
(ZMUM), Ajaria, E of Pass Goderdzi, ca 1620 m a.s.l., Abies and Pinus forest, N41º38’42.6", E42º35’45.5",
22.07.2012, leg. Y. Marusik; 6 males, 12 females (ZMUM), 15 km W of Adigeni, 1500–1700 m a.s.l, Abies, Picea,
Fagus, Acer etc. forest, litter, logs, under stones, 14–15.V.1983, leg. S. Golovatch; 4 females (ZMUM), W of
Adigeni, 1500–1700 m a.s.l., Abies, Picea, Fagus etc. forest, 21–22.VI.1978, leg. V. Dolin; 13 females (ZMUM),
Borzhomi Nature Reserve, Baniskhevi Valley, 800–900 m a.s.l., Picea, Fagus, Carpinus etc. forest, litter, under
bark and stones, 12 and 16.V.1983; 1 male (ZMUM), S of Bakuriani, Pinus and Fagus forest, 1750 m a.s.l., litter
and logs, 13.V.1983, all leg. S. Golovatch; 1 female (ZISP), Bakuriani, Bakuriani River valley, 11.V.1986, leg. L.
Zhiltsova; 6 males, 5 females (ZMUM), Mukhura, 14 km E of Tkibuli, Quercus, Fagus, Castanea etc. forest, litter
and under stones along spring, 23.X.1981, leg. S. Golovatch; 1 female (ZMUM), Mukhura ca 15 km E of Tkibuli,
700–800 m a.s.l., Castanea, Fagus, Carpinus etc. forest, litter, under bark and stones, 7–9.V.1987, leg. S.
Golovatch and K. Eskov; 8 juv. (ZMUM), ca 10 km E of Tkibuli, Mukhura, 1010 m a.s.l., Carpinus and Fagus
forest with Buxus thickets, N42°20’05,8”, E43°02’38,5”, 23.VII.2012, leg. Yu. Marusik; 1 male, 1 female, 3 juv.
(ZMUM), Svanetia, ca 40 km W of Mestia, Kherkhvashi E of Nakra (= Naki), 1200–1700 m a.s.l., Quercus, Fagus,
Carpinus, Picea, Abies etc. forest, litter and under bark, 21.VIII–21.IX.1986; 4 males, 1 female (ZMUM),
Svanetia, Mestia, 1500 m a.s.l., Betula and Rhododendron on moraine, litter and under stones, 5 and 16.IX.1986; 1
male (ZMUM), same locality, under stones, 22.X.1979; 7 males, 3 females (ZMUM), Svanetia, ca 40 km W of
Mestia, above Kherkhvashi, 1900–2200 m a.s.l., timberline (Rhododendron, Picea, Abies) and subalpine meadows,
21.IX.1986; 5 males, 5 females (ZMUM), Ambrolauri Distr., Cave Nikortsminda near Nikortsminda, 1100 m a.s.l.,
40–50 m deep, on walls and rotten wood, 22.X.1981; 1 male (ZMUM), same locality, 1150 m a.s.l., on logs,
19.X.1987; 1 juv. (ZMUM), Tskhaltubo, near Cave Belaya, deciduous forest on rock, litter and under stones,
26.X.1981; 1 juv. (ZMUM), Kutaissi Distr., Sataplia Nature Reserve, mixed deciduous forest, 8.VIII.1974; 1 juv.
(ZMUM), Orpiri NE of Kutaissi, rotten wood, 11.VIII.1974, all leg. S. Golovatch; 1 female (ZMUM), Sataplia
Nature Reserve, 400 m a.s.l., Fagus and Buxus forest, litter, 5.VI.1981, leg. S. Golovatch and J. Martens; 2 females
(ZMUM), Sataplia Nature Reserve, IV.1973, leg. D. Krivolutsky; 8 males, 7 females, 1 juv. (ZMUM), Oni Distr.,
ca 10 km NE of Shovi, Gurshevi near Pass Mamisoni, 2000–2200 m a.s.l., Abies, Fagus, Alnus etc. forest, litter and

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under stones, 21.X.1981; 3 males, 3 females (ZMUM), Oni Distr., SW of Pass Mamisoni, 2500 m a.s.l., Salix bush
along spring in subalpine meadow, litter and under stones, 21.X.1981; 1 male, 4 females (ZMUM), Oni Distr.,
Shovi, Fagus, Alnus, Abies etc. forest, litter and under stones near spring, 21.X.1981, all leg. S. Golovatch; 2
males, 2 females (ZMUM), Algeti Nature Reserve, W of Manglisi, 1400–1450 m a.s.l., Fagus, Picea, Acer etc.
forest, litter and under bark, 16–18.V.1987, leg. S. Golovatch and K. Eskov; 1 female (ZMUM), Algeti State
Reserve, Manglisi, Quercus forest, 12.VI.1984, leg. E. Kvavadze; 4 males, 4 females (ZMUM), Saguramo Nature
Reserve, NE of Mtskheta, Zedazeni, 1100–1200 m a.s.l., Fagus, Carpinus, Acer etc. forest, litter and under bark,
20.V.1987; 1 female (ZMUM), Lake Shaori, mixed forest, litter, N42º23’46.6", E43º01’47.7", ca 1180 m a.s.l.,
19.07.2012, leg. Y. Marusik; 2 males, 4 females (ZMUM), Batsaro Nature Reserve, ca 20 km N of Akhmeta, 800–
850 m a.s.l., Fagus, Castanea, Acer etc. forest, litter, 5–6.V.1987; 4 males, 2 females (ZMUM), Pass Magalakhari
between Akhmeta and Tianeti, 1200 m a.s.l., Fagus and Carpinus forest, litter and under bark, 6.V.1987; 9 juv.
(ZMUM), Lagodekhi Nature Reserve, trail to meteorological station, ca 1835 m a.s.l., Carpinus forest,
N41º51’38.0", E46º20’26.5", 27.07.2012, leg. Yu. Marusik; 4 males, 4 females (ZMUM), Kartli Mt. Ridge, Pass
Sabaduri ca 40 km NNE of Tbilisi, 1400 m a.s.l., Fagus forest, litter and under bark, 6.V.1987; 2 males, 1 female
(ZMUM), Surami Mt. Ridge, SE of Pass Djvari, between Gomi and Sachkhere, 850 m a.s.l., Alnus, Fagus,
Rhododendron etc. forest, litter, 7.V.1987; 1 male, 2 females (ZMUM), N of Kvareli, 700–750 m a.s.l., Fagus,
Carpinus, Quercus etc. forest, litter and under bark, 4.V.1987, all leg. S. Golovatch and K. Eskov; 1 male, 1 juv.
(ZMUM), Babaneuri Nature Reserve ca 16 km NE of Akhmeta, near Babaneuri, 500 m a.s.l., Fagus, Quercus,
Carpinus etc. forest, litter, 4–5.V.1987; 1 female, 4 juv. (ZMUM), 10 km N of Djvari, 800 m a.s.l., Buxus, Fagus,
Picea, Taxus etc. forest, litter, 20–21.VIII.1986; 1 male, 3 females, 1 juv. (ZMUM), Kazbegi, 2000 m a.s.l., Pinus
and Betula forest and subalpine meadows, litter and under stones, 2–6.VI.1982; 1 male, 2 females (ZMUM),
Kazbegi Distr., S of Gudauri near Pass Krestovyi, 1800 m a.s.l., Rhododendron bush, litter, 4.VI.1982, all leg. S.
Golovatch; 1 male (SMNG), Kazbegi, N42.69º, E44.52º, 28.VII.1985, leg. B. Seifert; 1 male (SMNG), Polyana
Firsova, 1400 m a.s.l., VIII./IX.1999, leg. O. Tietz; 1 male (HR), Tusheti, slope between Omalo and Khenako,
1810 m a.s.l., N42.3719º, E45.6444º, 4.IX.2009; 1 male (HR), Tusheti, pass between Omalo and Diklo, 2270 m
a.s.l., N42.4044º, E45.6239º, 5.IX.2009; 3 males, 1 female (HR), Khevsureti, Khone Skali River valley, between
Khone and Khone Chala, 1870 m a.s.l., N42.5675º, E45.2203º, 10.IX.2009; 1 female (HR), Khevsureti, Churo
Valley, mouth of Khone Skali River, 1980 m a.s.l., N42.5683º, E45.2322º, 10.IX.2009; Khevsureti, Khone Skali
River valley, between Khone and Khone Chala, 1870 m a.s.l., N42.5675º, E45.2203º, 10.IX.2009; 3 males, 1
female (HR), Khevsureti, ca 200 m downstream of Shatili, 1415 m a.s.l., N42.6594º, E45.1653º, 12.IX.2009; 4
males, 1 female (HR), Khevsureti, Argun River valley, between Shatili and Georgi Tsvinda, 1580 m a.s.l.,
N42.6397º, E45.1425º, 13.IX.2009; 2 males (HR), Shida Kartli, Ormotsi, Tana River valley, 16.6 road-km
upstream of Didi Ateni, 1110 m a.s.l., N41.8611º, E43.9444º, 16.IX.2011; 1 male (HR), Samtskhe–Djavakheti,
Borzhomi to Chitakhevi, junction to Mtsvane Monastery, 850 m a.s.l., N41.8039º, E43.3181º, 17.IX.2011; 1 male
(HR), Samtskhe-Djavakheti, Bakuriani, 7.5 road-km to Tskhratskaro Ugheltekhili Pass, 2080 m a.s.l., N41.7067º,
E43.5058º, 17.IX.2011; 1 female, 1 juv. (HR), Samtskhe-Djavakheti, Vardzia, 2 road-km from bridge to Tmogvi,
N41.3823º, E43.2982º, 25.VIII.2014; 1 male (HR), Racha-Lechkhumi and Kvemo Svaneti, Khotevi, 1 road-km to
Nikortsminda, 930 m a.s.l., N42.4636º, E43.1236º, 22.IX.2011; 1 male (HR), Racha-Lechkhumi and Kvemo
Svaneti, Zubi, road between Tsageri and Tkhaltubo, junction to Zubi, 430 m a.s.l., N42.5761º, E42.6808º,
23.IX.2011; 1 male (HR), Ajaria, Machakhlispiri, Chorokhi River valley, 2.7 road-km upstream of Acharistskhali,
80 m a.s.l., N41.5278, E41.7211, 29.IX.2011; 3 males (HR), Ajaria, NE of Dologani, 700 m, 100 m a.s.l.,
N41.5525º, E41.8019º, 29.IX.2011; 1 male (HR), Mtskheta–Mtianeti, Saguramo, 1.2 road-km towards Zedazeni
Monastery, 770 m a.s.l., N41.8889º, E44.7731º, 3.X.2011; 2 males, 1 female, 1 juv. (HR), Mtskheta–Mtianeti,
Tskhinvali, 6 road-km to Tianeti, 1120 m a.s.l., N42.1392º, E44.8167º, 5.X.2011; 1 male (HR), Samegrelo-Zemo
Svaneti, Nodashi, 1100 m a.s.l., N43.0586º, E42.4164º, 29.IX.2012; 1 male (HR), 3 males, 1 female (HR),
Mtskheta–Mtianeti, Stepantsminda, Darial Gorge, 4.5 road-km to border (near Gveleti), 1470 m a.s.l., N42.7058º,
E44.6267º, 6.X.2012; 3 females (HR), Mtskheta–Mtianeti, Andaki River valley 2.2 km upstream of its confluence
with Argun River, N42.5006º, E45.2156º, 8.VIII.2014; 1 male (HR), Mtskheta–Mtianeti, Darial Gorge at road-km
8/131, N42.7039º, E44.6264º, 10.VIII.2014; 1 male, 1 female (HR), Mtskheta–Mtianeti, 1.1 km SW of Kobi,
N42.5553, E44.4973º, 12.VIII.2014; 1 male (HR), Imereti, Nakeralos Ugheltekhili Pass, 730 m SSE, N42.3708,
E43.0428º, 14.VIII.2014; 1 male (HR), Samegrelo and Zemo Svaneti, Ushguli, entrance to Enguri (= Inguri) River
canyon, 2 km W of Ushguli, N42.9147º, E42.9747º, 19.VIII.2014, all leg. F. Walther.

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

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FIGURE 18. Morph D of Brachydesmus kalischewskyi Lignau, 1915, male holotype of B. talyschanus Lohmander, 1936: A–
C: anterior part of body, right half of metatergite 10 and caudal part of body, respectively, dorsal views. D–E: left gonopod,
mesal and ventral views, respectively. Designations: i, mesal subapical tooth of exomere; e, lateral apical tooth of exomere; l,
subapical lobe of exomere. Line drawings A–C by S. Dashdamirov; D–E after Lohmander (1936). Scale bar: 1.0 mm (A–C),
not to scale (D–E).
3 males, 1 female (ZMUM), ARMENIA, Azizbekov Distr., Zangegur Mt. Ridge, Pass Voratan, subalpine
meadow, 2100–2200 m a.s.l., under stones, 21.IV.1983; 1 male (ZMUM), near Azizbekov, 1200 m a.s.l., Artemisia
grassland, under stones, 21.IV.1983; 2 males, 6 females (ZMUM), Ararat Distr., Khosrov Nature Reserve, 1450–
1550 m a.s.l., Juniperus, Quercus, Crataegus etc. bush along river, litter and under stones, 19–20.IV.1983, all leg.
S. Golovatch; 4 males, 2 females (ZMUM), Ararat Distr., Khosrov Nature Reserve, 2000–2200 m a.s.l., subalpine
meadow, under stones, 11.V.1984; 5 males, 2 females, 1 juv. (ZMUM), Megri Distr., Nyuvadi, xerophytous
canyon, under stones, 25.IV.1983, all leg. V. Januschev; 7 males, 4 females, 4 juv. (ZMUM), Megri Distr., 6 km N
of Shvanidzor, sparse Quercus forest, 1200–1300 m a.s.l., litter, under stones and bark, 24.IV.1983; 8 males, 13
females, 3 juv. (ZMUM), environs of Megri, xerophytous bare canyon, 1000 m a.s.l., sparse Juniperus and Paliurus
bush, under stones, 24.IV.1983; 13 males, 18 females (ZMUM), Megri Distr., SSE of Lichk, Megri River valley,
1530 m a.s.l., Quercus forest, litter, under stones and in rotten wood, 25.IV.1983; 17 males, 15 females (ZMUM),
Megri Distr., above Kuris, 1500 m a.s.l., Quercus, Acer etc. forest, litter, under bark and stones, 26.IV.1983; 7
males, 4 females, 4 juv. (ZMUM), Megri Distr., 6 km N of Shvanidzor, sparse Quercus forest, 1200–1300 m a.s.l.,
litter, under stones and bark, 24.IV.1983; 23 males, 10 females (ZMUM), Kafan Distr., near Kadjaran, Megri Mt.
Ridge, N of Pass Tashtun, 2000 m a.s.l., Quercus forest on steep slope, litter and logs, 27.IV.1983; 38 males, 29
females (ZMUM), Kafan Distr., Shikahoh Nature Reserve, Shikahoh, 900–950 m a.s.l ., Quercus, Fagus, Carpinus
etc. forest along stream, litter, under logs and stones, 28.IV.1983; 7 males, 7 females (ZMUM), Kafan Distr.,
Shikahoh Nature Reserve, Shishkert, 1700–1800 m a.s.l., Quercus, Fagus, Carpinus forest, litter and under stones,
29.IV.1983; 2 males, 6 females (ZMUM), same locality and data; 8 males, 1 female, 1 juv. (ZMUM), Kafan Distr.,

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Shikahoh Nature Reserve, Tsav, 1000 m a.s.l., Quercus, Fagus, Carpinus etc. forest along stream, litter, under logs
and stones, 29.IV.1983; 1 female (ZMUM), Kafan, log, under bark, 27.IV1983; 1 male (ZMUM), Kafan Distr.,
Shikahoh Nature Reserve, Nerkin And, old Platanus stand along river, litter, rotten wood, under stones,
30.IV.1983; 3 males, 1 female, 1 juv. (ZMUM), Legvaz, 4 km NWN of Megri, Juniperus, Quercus etc. scrub with
Paliurus and Rosa, litter and under stones, 1000 m a.s.l., 24–25.IV.1983; 3 females (ZMUM), Stepanavan, foot of
Mt Archasar, 1400 m a.s.l., Quercus forest near spring, 14.XI.1985, all leg. S. Golovatch; 1 male, 1 female
(NHMG), Armenia, Jelal-Ogly (= now Stepanavan), 5.VIII.1920, leg. A. Shelkovnikov, det. H. Lohmander, vid. S.
Golovatch; 4 females (ZMUM), Kirovakan (= Vanadzor), 1600 m a.s.l., Quercus, Acer, Fagus etc. forest, litter,
22.V.1987, leg. S. Golovatch and K. Eskov; 2 males (ZMUM, ex Coll. HNHM), Karashamb, Agveran, 1500 m
a.s.l., shrubby hillside, under stones, 16.IX.1982, leg. O. Merkl and L. Ronkay; 28 males, 13 females, 3 juv.
(ZMUM), SW of Shnokh, halfway between Alaverdi and Bagratashen, 750 m a.s.l., Carpinus forest, litter,
24.V.1987, leg. S. Golovatch and K. Eskov; 2 females (ZMUM), Dilizhan Nature Reserve, Akhnabat Valley near
Salakh, 1400–1500 m a.s.l., Fagus, Taxus, Carpinus etc. forest, litter, under bark and stones, 17.IV.1983, leg. S.
Golovatch; 1 female (ZMUM), Dilizhan Nature Reserve, Agartsyn, 1350–1400 m a.s.l., Fagus, Acer etc. forest,
litter, 28–29.V.1987; 1 male, 7 females (ZMUM), W of Shamshadyn, halfway between Berd and Idjevan, 1500–
1600 m a.s.l., Fagus, Carpinus, Acer etc. forest, litter and under bark, 26–27.V.1987; 6 males, 4 females (ZMUM),
Berdavan ca 10 km S of Noemberian, 900–950 m a.s.l., Quercus, Carpinus, Acer etc. forest, litter and under stones,
24–25.V.1987, all leg. S. Golovatch and K. Eskov.
Male holotype of B. talyschanus (ZISP), AZERBAIJAN, “Baku Gouvernement, Lenkoran Distr., Lerik,
12.V.1912, leg. A. Kirichenko”; 13 males, 18 females (ZMUM), Mountainous Karabakh, Askeran Distr., ca 6 km
WNW of Dashbulag, near Badara, Quercus, Carpinus etc. forest, 850–900 m a.s.l., litter, under bark and stones,
2.V.1983; 9 males, 5 females (ZMUM), Mountainous Karabakh, Dashalty near Shusha, 1100–1300 m a.s.l.,
Quercus, Carpinus etc. forest, litter, logs, under stones, 1.V.1983, all leg. S. Golovatch; 1 juv. (ZMUM),
Mountainous Karabakh, environs of Firуuza, Domy, near Cave Glenkar, in soil, 16–17.VIII.1977, leg. N.
Zalesskaya; 2 juv. (ZMUM), Mountainous Karabakh, Drmbon ca 30 km WSW of Mardakert, 800–850 m a.s.l.,
Quercus, Acer etc. forest, litter, 1–2.VI.1987; 4 males, 5 females (ZMUM), Mountainous Karabakh, Chilisa ca 7
km N of Kelbadjar, 1450–1500 m a.s.l., Quercus, Carpinus, Acer etc. forest, litter and under bark, 31.V.1987; 10
males, 5 females, 5 juv. (ZMUM), Mountainous Karabakh, Nadirkhanly ca 12 km NE of Kelbadjar, 1200 m a.s.l.,
Fraxinus and Juglans regia stand on rock, litter and under stones, 1.VI.1987; 3 females (ZMUM), Mountainous
Karabakh, Lesser Istisu halfway between Kelbadjar and Istisu, 1550 m a.s.l., Quercus and Acer scrub on slope,
litter, 31.V.1987, all leg. S. Golovatch and K. Eskov; 5 males, 2 females (ZMUM), Istisu ca 8 km SW of Masally,
80–140 m a.s.l., Quercus, Carpinus, Acer etc. forest, litter, under bark and stones, 19–20.X.1983, leg. S.
Golovatch; 1 female (ZMUM), Djar near Zakataly, forest, litter, 18.IV.1986, leg. N. Loginova; 1 juv. (ZMUM),
Perzivan SE of Zakataly, 600 m a.s.l., Quercus, Fagus etc. forest along stream, litter, 3.V.1987, leg. S. Golovatch
and K. Eskov; 3 females, 8 juv. (ZMUM), Zakataly Nature Reserve, Katekhchay Valley, 700 m a.s.l., 23.V.1981,
leg. S. Golovatch and J. Martens; 6 females, 4 juv. (ZMUM), Zakataly Nature Reserve, 1000 m a.s.l., forest, litter,
13.VIII.1989; 1 female, 2 juv. (ZMUM), Zakataly Nature Reserve, Agkemal, 2200 m a.s.l., forest, litter,
15.VIII.1989, all leg. S. Dashdamirov and H. Aliev; 1 female (ZMUM), Zakataly Distr., Djar, along Tarykchay
River, steep slope at 45°, Fagus forest, litter, 8.VI.1986, leg. K. Mikhailov; 1 male, 1 female (ZMUM), Zakataly
Distr., Djar, 18.IV.1986, leg. N. Loginova; 1 female (ZMUM), Zakataly Nature Reserve, X.1970, leg. L. Malinina;
8 males, 3 females, 7 juv. (IZB), Kedabek (= Gədəbek), montane forest, VIII.1947, leg. A. Bogachev; 12 males, 8
females (ZMUM), Gey-Giol Nature Reserve, ca 50 km S of Kirovabad (= Ganja), 1500–1900 m a.s.l., Fagus,
Carpinus, Quercus etc. forest, litter, logs and under stones, 3–4.V.1983, leg. S. Golovatch; 2 males, 1 female, 2 juv.
(ZMUM), ca 15 km WSW of Mardakert, 1100 m a.s.l., Quercus, Fagus, Acer etc. forest, litter, 2.VI.1987; 1 male,
5 females (ZMUM), NW above Bash-Layski ca 20 km NNW of Sheki, 1250 m a.s.l., Fagus, Carpinus, Acer etc.
forest, litter, 3.V.1987, all leg. S. Golovatch and K. Eskov; 3 males, 14 females (ZMUM), AR Nakhichevan, S of
Pass Bichenek, W of Shakhbuz, Quercus forest, 1900 m a.s.l., litter, 22.IV.1983, leg. S. Golovatch; 8 males, 7
females (ZMUM), Yarymdja SE of Altyagach, 1320–1350 m a.s.l., Fagus, Quercus, Carpinus etc. forest, litter and
under bark, 20, 24–25.IV.1987; 1 female (ZMUM), Altyagach, 1050–1100 m a.s.l., Quercus, Fagus, Carpinus etc.
forest, litter, 20 and 26.IV.1987, all leg. S. Golovatch and K. Eskov; 9 males, 1 female, 3 juv. (ZMUM), Altyagach,
Yarymdja, 1200–1300 m a.s.l., forest, litter, 28–31.V.1998, leg. S. Dashdamirov; 5 males, 2 females (ZMUM),
Shemakha, Pirkuli, 24.V.1988, leg. N. Loginova; 1 subadult male, 2 females (ZMUM), Shemakha Distr., Pirkuli
Nature Reserve, near Observatorium, 1200–1250 m a.s.l., Quercus, Acer, Taxus etc. forest, litter, 30.IV.1987, leg. S.

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Golovatch and K. Eskov; 6 males (ZMUM), Laza near Kusary, 10.VIII.1985, leg. H. Aliev; 2 males, 1 female
(ZMUM), Talysh Mts, Lerik Distr., Zuvand, Galabyn, 1700–2000 m a.s.l., under stones on slopes, 10–11.X.1983,
leg. S. Golovatch; 1 female (ZMUM), Lerik Distr., Farm Azerbaijan, forest, litter, 21.VII.1970, leg. N. Loginova;
19 males, 10 females (ZMUM), Talysh Mts., Yardymly Distr., Allar, 1700–1800 m a.s.l., Quercus, Carpinus, Acer
etc. sparse forest, litter, rotten wood and under stones, 9.X.1983; 1 juv. (ZMUM), Talysh Mts., Lerik Distr.,
Zuvand, Gilidara, 1800 m a.s.l., semi-desert, 8.X.1983, all leg. S. Golovatch; 1 female (ZMUM), Talysh Mts.,
Zuvand, Djoni, 1500 m a.s.l., 28–29.V.1976, leg. V. Dolin; 5 males, 1 female (ZMUM), Agdash Distr., Turianchay
Nature Reserve, orchard, 20.X.1988, leg. S. Dashdamirov; 2 males, 1 female (ZMUM), Lenkoran Distr., Hyrcan
Nature Reserve, Apo ca 6 km SW of Alekseevka, Quercus, Acer, Carpinus, Parrotia etc. forest, litter, under stones
and bark, 14 and 16.X.1983; 2 juv. (ZMUM), Lenkoran Distr., Hyrcan Nature Reserve, Alekseevka, 50 m a.s.l.,
Quercus, Parrotia, Carpinus etc. forest, litter, rotten wood, 13.X.1983, all leg. S. Golovatch; 4 females (ZMUM),
Lenkoran Distr., Hyrcan Nature Reserve, near Alekseevka, 18.IV.1985; 2 males (ZMUM), Avrora, lowland forest,
21.IV.1985; 4 females (ZMUM), same locality, water reservoir, 18.IV.1985, all leg. E. Kupriyanova; 3 males
(ZMUM), same locality, forest, litter, 26.I–4.II.1985, leg. A. Druk; 1 male (ZMUM), Astara Distr., forest,
12.V.1969, leg. N. Loginova; 2 males, 10 females (HR), Təzə Alvadı, 1.6 road-km to Masallı, N39.075º,
E48.6304º; 0 m a.s.l., 25.III.2015; 3 males, 3 females (HR), Hyrcan Nature Reserve, Piran, northwestern town exit;
N38.734º, E48.6441º, 230 m a.s.l., 26.III.2015; 1 male, 2 females (HR), Hyrcan Nature Reserve, road Lenkoran—
Lerik at km 32, N38.7638º, E48.5819º, 400 m a.s.l., 26.III.2015; 2 males (HR), Hyrcan Nature Reserve, Günəşli, 2
road-km to Lerik, N38.7975º, E48.4644º, 710 m a.s.l., 26.III.2015; 2 males, 4 females (HR), Hyrcan Nature
Reserve, SW of Zünqüləş, N38.4493º, E48.7623º, 60 m a.s.l., 27.III.2015; 1 male, 2 females (HR), Hyrcan Nature
Reserve, SW of Zünqüləş, N38.448º, E48.7597º, 130 m a.s.l., 27.III.2015; 4 females (HR), Hyrcan Nature Reserve,
Daştatük, 1.3 road-km Xanbulan Reservoir, N38.6747º, E48.7622º, 110 m a.s.l., N38.6747º, E48.7622º,
27.III.2015; 1 female (HR), Hyrcan Nature Reserve, SW of Aşağı Apu, N38.6726º, E48.7362º, 180 m a.s.l.,
27.III.2015; 1 female (HR), Qurbanəfəndi 1.3 road-km to Xanəgah, N40.8614º, E48.1132º, 640 m a.s.l.,
30.III.2015, all leg. H. Reip and D. Antić.
6 males, 17 females, 21 juv. (SMF 5949, identified as B. ferrugineus, published by Golovatch 1981b), IRAN,
East Azerbaijan Prov., Arasbaran Wildlife Refuge, Makidi, secondrary Quercus bush, 4.VI.1978, leg. J. Martens
and H. Pieper.
Descriptive notes. Length ca 9–21 mm, width of midbody metazonae 0.9–2.7 mm (male, female), metazonite
to prozonite width ratio ca 1.4–1.6 (Fig. 10A–10D). Live coloration usually uniformly light brown to reddish or
dark brown.
All characters as in B. assimilis (Figs 10–27), except as follows. Paraterga from poorly to well developed,
largely subhorizontal to slightly declivous, only rarely the first few postcollar ones faintly upturned; fore margins
and anterolateral corners largely broadly and regularly rounded, only rarely angular; caudolateral corner mostly
obtuse-angled until segment 4, largely pointed or nearly pointed starting with segment 5, drawn behind rear tergal
margin starting with segments 6–14. Collum ovoid to, more rarely, transversely sublanceolate. Tergal setae mostly
very short and subclavate. Metatergal sculpture usually rough, only in segments 18 or 17 and 18 nearly obliterate.
Male prefemora bulging laterad, sphaerotrichomes present at least on tarsi. Gonopod (Figs 13, 14, 16, 17, 19, 21–
25, 27) bipartite, slightly falcate, like in B. assimilis, but with a typically bifid endomere (en) carrying a large,
conspicuous, densely pilose field, while endomere (ex) basically with three distal outgrowths: teeth e and i, and
lobe l.
At least six morphs are recognized here. Their more detailed characteristics are given in Table 1.
Remarks. Amongst the larger to largest Brachydesmus occurring in the Caucasus and adjacent parts of E
Turkey and NW Iran, this highly polymorphous species poses certainly the greatest problems for a taxonomic
treatment. Both Lignau (1915) and, later, Lohmander (1936), when describing their B. kalischewskyi, B.
karawajewi, B. ferrugineus and B. talyschanus as new species, had only very few samples in their hands. In
contrast to Lignau’s (1915) material which came from relatively small Abkhazia alone, Lohmander’s (1936)
stemmed from various, often widely separated places scattered across the entire Caucasus. More recently, Ceuca
(1975) reported B. ferrugineus from E Turkey, whereas Golovatch (1976) added his B. bidentatus. Had he been
aware of the then very fresh paper by Ceuca, he would never have dared describe it as a new species, because the
gonopods of “ferrugineus” from Turkey and of “bidentatus” from the central Caucasus Major were nearly
identical! Enghoff (2006) not only reiterated Ceuca’s (1975) record of B. ferrugineus, but he also reported B.
kalischewskyi from two localities in E Turkey.

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

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FIGURE 20. Morph F of Brachydesmus kalischewskyi Lignau, 1915. A: male from near Kadjaran, Megri Mt. Range, Armenia.
B: male from Shikahoh Nature Reserve, Megri Mt. Range, Armenia. C: male from Shnokh, Armenia, habitus, dorsal, lateral
and dorsal views, respectively. Pictures by K. Makarov, taken not to scale.
Golovatch (1981b) recorded B. ferrugineus in the East Azerbaijan Province of Iran, reiterated by Enghoff and
Moravvej (2005). That sole Iranian sample has again been borrowed for the present study.
Since our collections are indeed very large and densely blanket the entire region in question, we believe all
those “species” actually represent one, B. kalischewskyi by priority. We shall try to bring enough evidence to
support this conclusion, based on numerous measurements taken and abundant illustrations provided.
Treating the kalischewskyi complex requires recognition of several morphs which crudely correspond to the
more or less typical forms of several ex-species. To avoid the use of Latin names to denominate these forms, the
typical morph kalischewskyi will be referred to as A, while the morphs karawajewi, ferrugineus and talyschanus as
B, C and D, respectively. Neither the variety batumensis, proposed by Lohmander (1936) as an Ajarian form of
kalischewskyi, nor bidentatus are to be treated as separate morphs, as they sufficiently clearly represent morphs A
and B, respectively. Instead, we distinguish two more morphs which show stable morphological characters and
more or less coherent distributions. Thus, morph E corresponds to the populations from Svanetia, NW Georgia,

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whereas morph F to most of the populations inhabiting Armenia, Nakhichevan and Mountainous Karabakh. It must
be emphasized that, for the time being, none of the morphs seem to be sufficiently clear-cut to warrant the status of
a separate species. There are numerous morphological intergradations and transitions observed at least between the
neighbouring morphs, but in certain cases at least size differences in co-occurring morphs are dramatic (Fig. 11),
especially when material is scant. However, everything merges when sufficiently large samples are available.
Sometimes the attribution of a sample to this or that morph is by necessity arbitrary, as certain
(micro)populations appear to be disjunct morphologically. The choice in such cases is mainly determined by
geographical proximity. Thus, it is often very hard to distinguish morph B from morph D. For instance, we have
only located the latter in a single place in Georgia, whereas Lohmander identified two samples of morph D in that
country (Kobakhidze, 1965). Furthermore, the diversity of morphs could easily be extended, e.g., for several
disjunct (micro)populations in Ajaria, Krasnodar Province (Fig. 23), Azerbaijan or Armenia (Fig. 24).
FIGURE 21. Morph F of Brachydesmus kalischewskyi Lignau, 1915. A–C: male from Agveran, Armenia, anterior part of
body, right half of metatergite 10 and caudal part of body, respectively, dorsal views. D–E: male from near Shvanidzor, Megri
Mt. Range, Armenia, right gonopod, ventral and mesal views, respectively. F: male from near Badara, Mountainous Karabakh,
Azerbaijan, right gonopod, mesal view. Line drawings A–C by S. Dashdamirov. Scale bars: 1.0 mm (A–C), 0.2 mm (D–F).

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

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

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FIGURE 24. Examples of gonopods of Brachydesmus kalischewskyi Lignau, 1915. A–C: male from Farm Azerbaijan, Talysh
Mts, Azerbaijan, morph D, dorsal, ventral and mesal views, respectively. D–F: male from Shishkert, Shikahoh Nature Reserve,
Armenia, morph B?, mesal, lateral and ventral views, respectively. Designations: i, mesal, subapical tooth of exomere; e, lateral
apical tooth of exomere; l, subapical lobe of exomere. Scale bar: 0.1 mm.
Molecular data, bar-coding for one thing, would certainly be most helpful in further refining the picture, but at
the moment such information is unavailable.
Only in very few places such as near Sochi and within Adygea does morph B of B. kalischewskyi, represented
there by smaller individuals about 9–11 mm in length, occur not only sympatrically, but even syntopically together
with the superficially similar B. furcatus. However, these species are easily distinguished by angular versus broadly
rounded anterolateral corners of the paraterga, and by a more distinct versus a strongly obliterated metatergal
sculpture, respectively (cf. Figs 26 and 10A–10D), the males also by their disparate gonopod structure (cf. Figs 27
and 10E–10G).
Table 1 shows size variations in the main six morphs of B. kalischewskyi, coupled with short descriptive notes
and numerous illustrations to facilitate their recognition. The main characters such as size, gonopod structure and
the shape of the terga seem to form more or less random mosaics as well. There can be no doubt that the
kalischewskyi complex is in a stage of active speciation. It shows mixed sympatric (Bolnick and Fitzpatrick 2007;
Fitzpatrick et al. 2008), mosaic or simple parapatric speciation modes (see synopsis in Mallet et al. 2009). We
cannot rule out that some populations at the range peripheries might prove to represent proper species which
genetically evolve independently in the modern sense of the species concept (Queiroz 2007), yet with gene flows in

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the middle of the distribution area. Such a pattern does not automatically contradict the recognition of only a single
species because gene flow is not so uncommon in animals (Nosil 2008). Therefore, B. kalischewskyi as we treat it
could become a model species for further evolutionary studies.
FIGURE 25. Examples of gonopods of Brachydesmus kalischewskyi Lignau, 1915. A–C: male from Farm Azerbaijan, Talysh
Mts, Azerbaijan, morph D, lateral, mesal and ventral views, respectively. D: male from Arasbaran Wildlife Refuge, East
Azerbaijan Prov., Iran, morph D, mesal view. Designations: i, mesal subapical tooth of exomere; e, lateral apical tooth of
exomere; l, subapical lobe of exomere. Scale bars: 0.2 mm.

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

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FIGURE 27. Brachydesmus kalischewskyi Lignau, 1915, morph B, males from Pasture Abago and Arkhyz, Adygea, Russia,
respectively. A: leg 7, lateral view. B: left gonopod, mesal view. C: leg 7, lateral view. D: left gonopod, mesal view. Scale bars:
1.15 mm (A), 0.2 mm (B and D), 0.1 mm (C).
Map 2 roughly depicts the distribution of B. kalischewskyi in the Caucasus and adjacent parts of E Turkey and
NW Iran. Each basic morph has its own colour, whereas syntopic, sometimes mixed morphs are shown in both
colours involved. As one can easily see, the mosaic is really impressive. This is another strong argument for

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regarding the kalischewskyi complex as actually being represented by a single, highly polymorphous species. Some
of the morphs do reveal coherent patterns. Thus, morph F (orange dots in Map 2) seems to be more or less
restricted to a relatively small area in Svanetia, NW Georgia. Morph B (green dots in Map 2), apparently the most
widespread in the region, seems to be the sole inhabitant of the eastern half of the Caucasus Major, including its
northern macro-slope. However, most of the morphs tend to show no coherent patterns, being scattered across the
region and even beyond in what seems to be a more or less random way.
MAP 2. Distribution of Brachydesmus kalischewskyi: morphs A–F shown in blue (A), green (B), red (C), pink (D), orange (E)
and brown (F) circles, respectively; mixed or intermediate populations in two respective colours. The westernmost locality
lying in north-central Turkey, which Ceuca (1975) misspelled as “Yalnarcen gecidii (Yalnircan silsilesi), 2500 m”, apparently
supports morph B and actually corresponds to Ilgazdaggecidi: Ilgazdag Pass, 15 km N of Tosya, N 41.144611º, E 34.062862º.
It is omitted from Map 2.
No doubt this pretty crude picture requires further refinements through bringing in more evidence, both
morphological and, especially, molecular-based. Discrimination or revival of some local species within the
kalischewskyi complex is also possible in the future. At least some species names like karawajewi, ferrugineus or
talyschanus are long available from a taxonomic point of view.
Polymorphism and distrubution mosaics like the ones demonstrated by B. kalischewskyi have long been
reported in Polydesmidae. Thus, Epanerchodus polymorphus Mikhaljova and Golovatch, 1981, from the Far East
of Russia (Mikhaljova and Golovatch 1981), based both on somatic and gonopod characters, shows two stable
morphs in the male, but a complete transitional series in the female. Exactly the same pattern of dimorphic males,
but polymophous females seems to be observed in another few highly localized cave species of Epanerchodus in

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Japan (e.g. Murakami 1970; Murakami and Kawasawa 1975). In contrast, the widespread East Asian E. orientalis
Attems, 1901 shows only a gradual change in gonopod structure from north to south along Japan (Nishikawa and
Murakami 1993), but further southerly, in Taiwan and southern China, the morphological variation and distribution
seem to grow increasingly sporadic and mosaic (Golovatch et al. 2011, 2012).
Polymorphism and very complex mosaic distributions have also been reported in some other millipede groups,
e.g. Xystodesmidae (Polydesmida) in North America (Shelley and Whitehead, 1986). They concern not only likely
infraspecific variations like those we face in Brachydesmus kalischewskyi, but also what Mesibov (2003)
denominated as lineage mosaics, i.e. spatial mosaics of closely related species which are the result of
differentiation in form and geographical space within monophyletic clades.
Brachydesmus (Haplobrachidesmus) kvavadzei sp. n.
Fig. 28, Map 3
Material examined. Holotype male (MIZW), GEORGIA, Ajaria, Zelenyi Mys near Batumi, Botanical Garden,
30.X.–6.XI.1963, leg. B. Pisarski. Paratypes: 3 males, 2 fragments (MIZW), 1 male (ZMUM p3035), same data,
together with holotype. Non-type: 1 female (ZMUM p3036), Ajaria, Khulo Distr., 3 km W of Danisparauli,
deciduous forest, litter, 10.X.1981, leg. S. Golovatch.
Diagnosis. Differs from B. kalischewskyi, the only consubgener remaining accepted, in the particularly small
body, the nearly fully obliterate metatergal sculpture, the absence of sphaerotrichomes, and the presence of only a
single apical tooth on the gonopod exomere; from B. simplex sp. n. primarily by the particularly slender male legs,
the narrower paraterga and the considerably shorter exomere of the gonopod.
Name. To commemorate the late Dr. Eristo Sh. Kvavadze, a prominent specialist in the earthworms of the
Caucasus and a close friend of the first author.
Description. Length 5–6 mm, width of midbody metazonae 0.65–0.7 mm (male), metazonite to prozonite
width ratio ca 1.3 (Fig. 26A). Coloration uniformly pallid to light brown.
All characters as in B. assimilis (Fig. 28), except as follows. In width, collum < 2 = 3 < 4 < head = 5. Paraterga
poorly developed, their fore margins and anterolateral corner largely broadly and regularly rounded, not angular;
caudolateral corner mostly acute-angled, pointed or nearly pointed and drawn behind rear tergal margin only in
segments 17 and 18 (Fig. 28A). Tergal setae mostly very short and subbacilliform. Metatergal sculpture nearly
fully obliterated. Male prefemora not bulging laterad, slender; sphaerotrichomes wanting (Fig. 28B). Gonopod
(Fig. 28C–28D) bipartite, stout, endomere (en) typically bifid and partly pilose, but exomere (ex) long, with only a
single, apical, rounded tooth (e).
Brachydesmus (Haplobrachidesmus) simplex sp. n.
Fig. 29, Map 3
Material examined. Holotype male (ZMUM p3037), ABKHAZIA, Gagra Distr., Lake Ritsa, 950–1100 m a.s.l.,
Fagus, Abies, Picea, Acer etc. forest, litter, under bark and stones, 13–14.VIII.1986, leg. S. Golovatch. Paratypes:
5 males (ZMUM p3038), same data, together with holotype; 1 male, 6 females (ZMUM p3039), Myussera Nature
Reserve, 20–130 m a.s.l., Castanea, Alnus etc. forest, 8–10.IV.1983, leg. S. Golovatch; 3 males (MIZW), 1 male
(ZMUM p3040), RUSSIA, Krasnodar Prov., Sochi, open Carpinus and Quercus forest, loam, 7.XII.1963; 1 male
(MIZW), same locality, dendrarium and city park, under stones, 9.XII.1963, all leg. P. Pisarski.
Diagnosis. See Diagnosis of B. kvavadzei sp. n., as well as Key below.
Name. To emphasize the particularly simple gonopod exomere.
Description. Length 6–7.5 mm, width of midbody metazonae 0.8–1.0 mm (male, female), metazonite to
prozonite width ratio ca 1.5–1.6 (Fig. 29A). Coloration uniformly pallid to light yellow.
All characters as in B. assimilis (Fig. 29A), except as follows. In width, collum < 2 < 3 < head = 4 < 5.
Paraterga well-developed, their fore margins nearly straight, while anterolateral corners clearly angular;
caudolateral corner mostly acute-angled and dentiform starting with segment 5, subspiniform and drawn behind
rear tergal margin starting with segment 15 (Fig. 29A). Tergal setae mostly very short and subclavate. Metatergal

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sculpture indistinct, but at least middle and caudal rows of flat small tubercles visible. Male legs clearly incrassate
compared to female, but male prefemora not bulging laterad; sphaerotrichomes absent (Fig. 29B). Gonopod (Fig.
29C–29G) bipartite, stout, exomere with only a single, small, apical tooth (e).
FIGURE 28. Brachydesmus kvavadzei sp. n. A: male paratype, habitus, dorsal view. B: leg 7, lateral view. C–D: right
gonopod, mesal and lateral views, respectively. Designations: en, endomere; ex, exomere; e, apical tooth of exomere. Picture A
by K. Makarov, taken not to scale. Scale bars: 0.25 mm (B), 0.1 mm (C–D).

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FIGURE 29. Brachydesmus simplex sp. n., male paratype from Sochi, Krasnodar Prov., Russia. A: habitus, dorsal view. B: leg
7, lateral view. C–D: left gonopod, ventral and dorsal views, respectively. E: right gonopod telopodite of male paratype from
near Lake Ritsa, Abkhazia, lateral view. F–G: left gonopod of male paratype from Myussera Nature Reserve, Abkhazia, dorsal
and ventral views, respectively. Designation: e, lateral apical tooth of exomere. Scale bar: 0.1 mm (B–G), not to scale (A).
Key to Polydesmidae species presently known to occur in the Caucasus
1. Adults with 20 segments, including telson. Collum often as broad as or broader than head. Gonopod telopodite often elongate
and subgeniculate distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polydesmus, 2
- Adults with 19 segments, including telson. Head broader than collum. Gonopod telopodite stout and never subgeniculate dis-
tally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brachydesmus, 5

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2. Live coloration usually light brown to brown (Figs 3A, 5A). Paraterga not upturned, subhorizontal to slightly declivous, dor-
sum slightly, but clearly convex in the middle (Figs 3A, 3H–3J, 5A). Collum clearly narrower than head (Figs 3A, 3H, 5A). .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
- Coloration largely red to pinkish. Paraterga very broad, held high, about level to a flat mid-dorsal region, a few first postcollum
paraterga slightly, but clearly upturned (Figs 2E, 3B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3. Male prefemora strongly bulging laterad (Fig. 5B). Gonopod telopodite stout (Fig. 5C) . . . . . . . . . Polydesmus mediterraneus
- Male prefemora only slightly bulging laterad. Gonopod telopodite strongly elongate, subgeniculate distally (Fig. 4C, 4D) . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. muralewiczi
4. Length ca 10–13 mm (male, female), width of midbody metazonae 1.25–1.7 (male) or 1.4–2.1 mm (female), metazonite to
prozonite width ratio < 2.0 (Fig. 2). Collum about as broad as head (Fig. 2A, 2B, 2F) . . . . . . . . . . . . . . . . . . . . . . . . . P. lignaui
- Length ca 13–19 mm (male, female), width of midbody metazonae 1.9–3.3 (male) or 2.4–3.5 mm (female), metazonite to pro-
zonite width ratio > 2.0 (Fig. 1). Collum clearly broader than head (Fig. 1) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. abchasius
5. Paraterga poorly-developed, their fore margins and anterolateral corners largely broadly and regularly rounded, not angular
(Fig. 10C). Metatergal sculpture strongly obliterate (Fig. 10A–10D). Gonopod exomere (ex) with a conspicuous densely pilose
strip (Fig. 10E–10G). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brachydesmus furcatus
- Paraterga variously developed, but their fore margins and anterolateral corners mostly clearly angulate (e.g. Figs 6, 9A, 12,
13B, 13E, 15B, 16B, 16E). Metatergal sculpture often more strongly developed. Gonopod exomere without a pilose strip, but
endomere often with densely pilose patches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6. Tergal setae sharp (Fig. 9A). Gonopod (Fig. 9B) unipartite, slender, only slightly falcate, ventrally with several teeth or spines.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. superus
- Tergal setae mostly bacilliform or subclavate, only rarely sharpened. Gonopod telopodite bipartite, with a distinct exomere and
a smaller, but evident endomere.
7. Adults < 8 mm long and ≤ 1.0 mm wide. Gonopod exomere with only a single apical tooth (Figs 28C–28D, 29C–29G).
Colchis, Black Sea coast area . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
- Adults ≥ 9 mm long and ≥ 1.0 mm wide. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
8. Length 6–7.5 mm, width of midbody metazonae 0.8–1.0 mm, metazonite to prozonite width ratio ca 1.5–1.6 (Fig. 29A). Male
legs slightly, but clearly incrassate (Fig. 29B). Gonopod exomere longer (Figs 29C–29G). Ajaria, Georgia. . B. simplex sp. n.
- Length 5–6 mm, width of midbody metazonae 0.65–0.7 mm, metazonite to prozonite width ratio ca 1.3 (Fig. 28A). Male legs
slender (Fig. 28B). Gonopod exomere shorter (Figs 28C–28D). Abkhazia and Sochi, Krasnodar Prov., Russia . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .B. kvavadzei sp. n.
9. Gonopod endomere (en) partially densely pilose and clearly bifid; exomere (ex) with 1–2 apical teeth e and i (the latter often
subapical), often also with another subapical lobule or denticle l (Figs 13G–13H, 14C–14E, 16G–16H, 18D–18E, 19D–19E,
21D–21F, 22–25, 27B, 27D). Extremely widespread (Map 2) and variable in all other characters, morphism outlined in Table
1. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. kalischewskyi
- Gonopods subfalcate, endomere (en) simple, more or less finger-shaped (Figs 7, 8E–8F) . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10. Tergal setae mostly subclavate (Fig. 6C–6H). Gonopod exomere (ex) more elaborate (Fig. 7). Much of both Caucasus Major
and Caucasus Minor, but not Hyrcania (Map 3). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. assimilis
- Tergal setae sharp (Fig. 9A). Gonopod exomere (ex) simple (Fig. 8F). Hyrcania (Map 3).. . . . . . . . . . . . . . . . . . B. pigmentatus
Conclusions
In the Caucasus, the Polydesmidae can be regarded as a fairly diverse family which, like most of the Diplopoda,
largely contains forest-dwellers. Apart from two obvious introductions, Polydesmus mediterraneus and
Brachydesmus superus, the fauna is represented by endemic or subendemic species. Furthermore, the subgenera
Haplobrachidesmus and Lophobrachydesmus of Brachydesmus are subendemic to the region together with the
adjacent parts of N and NE Turkey and NW Iran. For the present, the fauna may be considered as quite well-
known.
The distribution of Polydesmidae in the Caucasus (Maps 1–3) shows a number of obvious patterns. Only very
few species are local in distribution, e.g. B. kvavadzei sp. n. or B. simplex sp. n., whereas most are widespread. The
few high-montane records (above 2000 m a.s.l.) are only found amongst the latter category: Polydesmus lignaui,
Brachydesmus assimilis, B. pigmentatus and, especially, B. kalischewskyi. All indigenous Polydesmus species are
confined to the NW and W Caucasus (Map 1), thus warranting the distribution pattern to be termed Colchidan. The
same concerns B. furcatus, B. kvavadzei sp. n. and B. simplex sp. n. The polymorphous B. kalischewskyi is
particularly widely distributed, blanketing the whole region except for most of its arid parts (Map 2). At the
opposite end, B. pigmentatus can be considered as showing a Hyrcanian distribution pattern, being subendemic to
Hyrcania both within the Republic of Azerbaijan and NW and N Iran (Elburs and Zagros mountains) (Map 3).
Strict syntopy among the indigenous Polydesmidae of the Caucasus seems to be absent in the Polydesmus
abchasius—P. lignaui pair, as well as in Brachydesmus furcatus, B. assimilis and B. pigmentatus which totally fail

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to co-occur (Maps 1, 3). Instead, at least in some places each of the Caucasian species, including both new ones,
shows sympatry or even syntopy with B. kalischewskyi (Map 2).
MAP 3. Distribution of the remaining species of Brachydesmus in the Caucasus region and adjacent parts of Iran: B. assimilis
(blue circle), B. furcatus (pink triangle), B. pigmentatus (red square, to be extended to SW Turkmenistan in the east), B. superus
(green star), B. kvavadzei sp. n. (black cross) and B. simplex sp. n. (brown diamond).
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 all or much of it at our choice. We are greatly obliged to the following
curators for the loans of material under their care: Kirill Mikhailov and Elena Kudryavtseva (both ZMUM), Olga
Ovchinnikova and Viktor Krivokhatsky (both ZISP), Dmitri Logunov and Galina Azarkina (both ZMN), Zoltán
Korsós (HNHM), Halid Aliev (IZB), Ted von Proschwitz (NHMG), Karin Voigtländer (SMNG) and Jolanta
Wytwer (MIZW). Henrik Enghoff (ZMUC) and William A. Shear (Hampden-Sydnet, Virginia, U.S.A.) kindly
made several important corrections of and suggestions to advanced drafts. Selvin Dashdamirov (now Düsseldorf,
Germany) very skillfully executed several line drawings, while Kirill Makarov (Moscow, Russia) and Roman Zuev
(Stavropol, Russia) most helpfully took most of the pictures. One of the collectors, F. Walther (Hamburg,
Germany), is grateful to VolkswagenStiftung for funding his several recent excursions to the Caucasus (within the
project 'Biogeography of the land molluscs of the Caucasus region').

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