Diversity and distribution of the millipedes (Diplopoda) of Georgia, Caucasus

Abstract

The diplopod fauna of Georgia, Transcaucasia, is very rich given the country’s relatively small territory; it presently comprises 103 species from 44 genera, 12 families, and 7 orders. Most of the Diplopoda known from Georgia (86 species, or 83%) demonstrate Caucasian distribution patterns, 36 and 46 species, as well as 8 and 9 genera being endemic or subendemic to the country, respectively. A single Holarctic family, Anthroleucosomatidae (order Chordeumatida), contains 44 Caucasian species and 20 genera, of which 27 species and 14 genera are endemic or subendemic to Georgia. Likewise, all species from the orders Polyzoniida, Siphonocryptida, Glomerida and Chordeumatida, as well as most species of Julida and Polydesmida are native, also endemic or subendemic to the Caucasus, but the genera and families they represent are widely distributed at least across the Euro-Mediterranean Realm. Most of the presumed troglobionts in the Caucasus appear to be confined to western Georgia’s karst caves (14 species, 5 genera). Within Georgia, the fauna of the western part (= Colchis) is particularly rich and diverse, while that of the central and eastern parts of the country grows increasingly depauperate inland following the gradual climatic aridisation from west (Black Sea coast) to east (Armenia and Azerbaijan). The vertical distribution of the Diplopoda in Georgia, as well as the Caucasus generally, shows the bulk of the fauna restricted to forested lowland to mountain biomes or their remnants. Only very few Chordeumatida and Julus species seem to occur solely in the subalpine to alpine environments and thus may provisionally be considered as high-montane elements. Ongoing and future research on the millipedes of the Caucasus, especially in cave and montane environments, will undoubtedly allow for many more novelties and details of the diversity and distribution of Georgia’s Diplopoda to be revealed or refined.

Full text

Diversity and distribution patterns of the millipedes of Georgia 199
Diversity and distribution of the millipedes
(Diplopoda) of Georgia, Caucasus
Mzia S. Kokhia1, Sergei I. Golovatch2
1 Institute of Zoology, Ilia State University, K. Cholokashvili Ave., 3/5, Tbilisi 0162, Georgia 2 Institute for
Problems of Ecology and Evolution, Russian Academy of Sciences, Leninsky prospekt 33, Moscow 119071, Russia
Corresponding author: Sergei I. Golovatch (sgolovatch@yandex.ru)
Academic editor: Z. Korsós|Received 21 October 2019|Accepted 10 January 2020|Published 28 April 2020
http://zoobank.org/3D90FD4C-DC71-4F68-987D-8F24DE2141A8
Citation: Kokhia MS, Golovatch SI (2020) Diversity and distribution of the millipedes (Diplopoda) of Georgia,
Caucasus. In: Korsós Z, Dányi L (Eds) Proceedings of the 18th International Congress of Myriapodology, Budapest,
Hungary. ZooKeys 930: 199–219. https://doi.org/10.3897/zookeys.930.47490
Abstract
e diplopod fauna of Georgia, Transcaucasia, is very rich given the country’s relatively small territory; it
presently comprises 103 species from 44 genera, 12 families, and 7 orders. Most of the Diplopoda known
from Georgia (86 species, or 83%) demonstrate Caucasian distribution patterns, 36 and 46 species, as
well as 8 and 9 genera being endemic or subendemic to the country, respectively. A single Holarctic
family, Anthroleucosomatidae (order Chordeumatida), contains 44 Caucasian species and 20 genera, of
which 27 species and 14 genera are endemic or subendemic to Georgia. Likewise, all species from the
orders Polyzoniida, Siphonocryptida, Glomerida and Chordeumatida, as well as most species of Julida
and Polydesmida are native, also endemic or subendemic to the Caucasus, but the genera and families
they represent are widely distributed at least across the Euro-Mediterranean Realm. Most of the presumed
troglobionts in the Caucasus appear to be conned to western Georgia’s karst caves (14 species, 5 genera).
Within Georgia, the fauna of the western part (= Colchis) is particularly rich and diverse, while that of
the central and eastern parts of the country grows increasingly depauperate inland following the gradual
climatic aridisation from west (Black Sea coast) to east (Armenia and Azerbaijan). e vertical distribution
of the Diplopoda in Georgia, as well as the Caucasus generally, shows the bulk of the fauna restricted to
forested lowland to mountain biomes or their remnants. Only very few Chordeumatida and Julus species
seem to occur solely in the subalpine to alpine environments and thus may provisionally be considered as
high-montane elements. Ongoing and future research on the millipedes of the Caucasus, especially in cave
and montane environments, will undoubtedly allow for many more novelties and details of the diversity
and distribution of Georgia’s Diplopoda to be revealed or rened.
ZooKeys 930: 199–219 (2020)
doi: 10.3897/zookeys.930.47490
http://zookeys.pensoft.net
Copyright Mzia S. Kokhia, Sergei I. Golovatch. This is an open access article distributed under the terms of the Creative Commons Attribution License
(CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
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Mzia S. Kokhia & Sergei I. Golovatch / ZooKeys 930: 199–219 (2020)
200
Keywords
checklist, Colchis, endemism, fauna, Myriapoda
Introduction
Georgia is one of the main countries in the Caucasus, lying between western Asia and
Eastern Europe. It is bounded in the west by the Black Sea, in the north by Russia, in the
south by Turkey, and in the southeast and east by Armenia and Azerbaijan (Fig. 1). e
area is mainly montane to high montane, situated between 41° and 44°N, and 40° and
47°E. e Greater Caucasus Mountain Range, or Caucasus Major, forms the northern
border of Georgia, while the southern border is bounded by the Lesser Caucasus Moun-
tains, or Caucasus Minor. e Caucasus Major is much higher in elevation (more than
5000 m a.s.l.) than the plateau-like Caucasus Minor, both being connected by the sub-
meridional Surami (= Likhi) Mountain Range which divides Georgia into the western and
central + eastern parts. Both parts are quite varied in climate and biota. Western Georgia’s
landscape ranges from lowland marsh-forests, swamps, and temperate rainforests within
the Colchis Plain to eternal snows and glaciers, while the eastern part of the country even
contains a small segment of semi-arid plains. Forests cover around 40% of Georgia’s ter-
ritory, while the alpine/subalpine zone accounts for approximately 10% of the land. e
climate of Georgia is extremely diverse, considering the nation’s small size, but is largely
mild to warm. ere are two main climatic zones, roughly corresponding to the eastern
and western parts of the country. e Greater Caucasus Mountain Range plays an impor-
tant role in moderating Georgia’s climate and protects the nation from the penetration
of colder air masses from the north. e Lesser Caucasus Mountains partially protect the
region from the inuence of dry and hot air masses from the south (Bondyrev et al. 2015).
e millipede fauna of Georgia has recently been reviewed and shown to comprise 95
species from 42 genera, 12 families, and 7 orders (Kokhia and Golovatch 2018). A few rel-
evant faunistic papers have, or will have, appeared since (Golovatch 2018, Golovatch and
Turbanov 2017, Antić et al. 2018, Evsyukov et al. 2018, 2020, Vagalinski and Lazányi
2018, Short et al. 2020), allowing for the previous checklist to be rectied and updated,
as well as the previous reference list to be considerably shortened. e present checklist
contains 103 species from 44 genera, 12 families, and 7 orders (Table 1). Data on the
elevations at which the species occur, both within and beyond Georgia, are also added,
representing the basic information for our analysis of millipede vertical distributions.
Material and methods
Only described species and published records are considered in our paper, while dubi-
ous taxa and those not identied to the species level have been omitted both from the
checklist and reference list. Only one important exception has been made: Calyptophyl-
lum sp. as the only record of this genus in the Caucasus (Table 1).

Diversity and distribution patterns of the millipedes of Georgia 201
Figure 1. Geographic division of Georgia.
ree zigzag transects chosen to grossly reect the north-to-south lie of the macro re-
lief of Georgia, extending from the Caucasus Major in the north to the Caucasus Minor
in the south (Figs 2–5), have been drawn, one each for the western, central and eastern
parts of the country (Fig. 1). e transect across western Georgia connects Pitsunda –
Arabika Plateau – Khaishi – Bagdati – Batumi (427 km long), that in central Georgia
connects Roki Tunnel – Tskhinvali – Tbilisi – Tsalka Reservoir – Ninotsminda – Ja-
vakheti National Park (275 km), and the eastern Georgia one connects Omalo – Tianeti
– Akhmeta – Shilda – Kvareli – Lagodekhi – Tamariani (186 km) (Fig. 2). Both at the
bottom of the maps and on the maps themselves, each transect is accompanied by the re-
spective altitudes given for each of the turn localities and thus provides a clear generalized
picture of the macro relief (Figs 3–5). ese three transects thus cover all major variations
in millipede vertical distribution across entire Georgia. is novel approach to a graphic
presentation of faunistic results allows us to combine the horizontal and vertical distribu-
tions of millipedes in the easiest and most vivid way on the same map. Mapping largely
concerns endemic or subendemic species and concerns only the territory of Georgia.
Most of the colour maps were generated using Google Earth Pro version 7.3.2.5495
and Adobe Photoshop CS6. e nal images were processed with Adobe Photoshop CS6.
Results
e diplopod fauna of the Caucasus region, including Georgia, is basically Euro-Medi-
terranean in its composition (Table 1). is also concerns the relatively few widespread,

Mzia S. Kokhia & Sergei I. Golovatch / ZooKeys 930: 199–219 (2020)
202
Table 1. A revised checklist of the Diplopoda of Georgia, with data on species distributions, both within
and beyond the country, their statuses, and the main relevant literature sources. Designations: i – intro-
duced; G – entire Georgia; W – western Georgia; C – central Georgia; E – eastern Georgia; R – Russian
Caucasus; T – Turkey; Ar – Armenia; Az – Azerbaijan; Cr – Crimean Peninsula; (+) – present; e – endemic
to Georgia; se – subendemic to Georgia; t – presumed troglobiont; sc – subcosmopolitan; EuM – Euro-
Mediterranean; M – Mediterranean; EM – eastern Mediterranean; EE – eastern European; Ca – Caucasian.
Fauna G R T Ar Az Cr Elevations
(m a.s.l.) and
status
Distribution
pattern
Main relevant
references
Class Diplopoda
Order Polyxenida
Family Polyxenidae
Genus Polyxenus Latreille, 1803
1. Polyxenus lagurus (Linnaeus,
1758)
W + + 20–1700, i sc Issaev 1911, Short
et al. 2020
2. Polyxenus lankaranensis Short,
Vahtera, Wesener & Golovatch,
2020
E + + 100–800 Ca Short et al. 2020
Genus Propolyxenus Silvestri, 1948 W
3. Propolyxenus argentifer (Verhoe,
1921)
G + + + + + 20–1700 EM Short et al. 2020
Family Lophoproctidae
Genus Lophoproctus Pocock, 1894
4. Lophoproctuscoecus Pocock, 1894 G + 20–900 EM Short 2015, Short
et al. 2020
Order Polyzoniida
Family Hirudisomatidae
Genus Hirudisoma Fanzago, 1881
5. Hirudisoma roseum (Victor, 1839) G + + + 20–1100, se EM Golovatch et al.
2015
Order Siphonocryptida
Family Siphonocryptidae
Hirudicryptus Engho & Golovatch, 1985
6. Hirudicryptus abchasicus
Golovatch, Evsyukov & Reip, 2015
W + 600–1500, se Ca Golovatch et al.
2015, Zuev 2017
Order Glomerida
Family Glomeridae
Genus Hyleoglomeris Verhoe, 1910
7. Hyleoglomeris awchasica (Brandt,
1840)
W + 20–2100, se Ca Golovatch 1975,
1976a, 1989b
8. H. specialis Golovatch, 1989 E + 500–1400, se Ca Golovatch 1989b
Genus Trachysphaera Heller, 1858
9. Trachyspaera costata (Waga, 1857) G + + + + + 20–2000 EuM Golovatch 1990,
2008
10. T. fragilis Golovatch, 1976 G + 80–460, t, e Ca Golovatch
1976c,1990,
Golovatch and
Turbanov 2017
11. T. minuta Golovatch, 1976 G + + + 20–1700, se Ca Golovatch 1976c,
1990
12. T. orientalis Golovatch, 1976 W 800–1100,
t, e
Ca Golovatch 1976c,
1990
13. T. radiosa (Lignau, 1911) W + 20–1800, se Ca Golovatch 1976a,
1990
14. T. solida Golovatch, 1976 W, C 20–2020, se Ca Golovatch 1976c,
1976c, 1990, 1993

Diversity and distribution patterns of the millipedes of Georgia 203
Fauna G R T Ar Az Cr Elevations
(m a.s.l.) and
status
Distribution
pattern
Main relevant
references
Family Glomeridellidae
Genus Typhloglomeris Verhoe, 1898
15. Typhloglomeris lohmanderi
(Golovatch, 1989)
C, E + + 600–1450, se Ca Golovatch 1989a,
2003
16. Typhloglomeris palatovi
Golovatch & Turbanov, 2018
W 650, t, e Ca Golovatch and
Turbanov 2017
Order Chordeumatida
Family Anthroleucosomatidae
Genus Acanthophorella Antić & Makarov, 2016
17. Acanthophorella barjadzei Antić
& Makarov, 2016
W 1120–1200,
t, e
Ca Antić and Makarov
2016
Genus AdshardicusGolovatch, 1981
18. Adshardicus strasseri Golovatch,
1981
W + 20–530, se Ca Engho 2006,
Antić and Makarov
2016
Genus Alpinella Antić & Makarov, 2016
19. Alpinella waltheri Antić &
Makarov, 2016
E 2860, e Ca Antić and Makarov
2016
Genus Brachychaetosoma Antić & Makarov, 2016
20. Brachychaetosoma turbanovi
Antić & Makarov, 2016
W 300, t, e Ca Antić and Makarov
2016
Genus Caucaseuma Strasser, 1970
21. Caucaseuma kelasuri Antić &
Makarov, 2016
W 190, e Ca Antić and Makarov
2016
22. C. variabile Antić & Makarov,
2016
C + 100–2500, se Ca Antić and Makarov
2016
Genus Cryptacanthophorella Antić & Makarov, 2016
23. Cryptacanthophorella manubriata
Antić & Makarov, 2016
W, C 800–1700, e Ca Antić and Makarov
2016
Genus Dentatosoma Antić & Makarov, 2016
24. Dentatosoma denticulatum Antić
& Makarov, 2016
W 400–900, e Ca Antić and Makarov
2016
25. D. magnum Antić & Makarov,
2016
W + 20–2200, se Ca Antić and Makarov
2016
26. D. zeraboseli Antić & Makarov,
2016
W 20–1700, e Ca Antić and Makarov
2016
Genus Georgiosoma Antić & Makarov, 2016
27. Georgiosoma bicornutum Antić &
Makarov, 2016
W 2000, t, e Ca Antić and Makarov
2016
Genus Herculina Antić & Makarov, 2016
28. Herculina oligosagittae Antić &
Makarov, 2016
W 1500–1700, e Ca Antić and Makarov
2016
29. H. polysagittae Antić & Makarov,
2016
C 1750, e Ca Antić and Makarov
2016
Genus Heterocaucaseuma Antić & Makarov, 2016
30. Heterocaucaseuma deprofundum
Antić & Makarov, 2018
W 2000–2100,
t, e
Ca Antić et al. 2018
31. H. longicorne Antić & Makarov,
2016
W 100–350, t, e Ca Antić and Makarov
2016, Antić et al.
2018
32. H. mauriesi (Golovatch &
Makarov, 2011)
W 215, t, e Ca Golovatch and
Makarov 2011,
Antić and Makarov
2016, Antić et al.
2018

Mzia S. Kokhia & Sergei I. Golovatch / ZooKeys 930: 199–219 (2020)
204
Fauna G R T Ar Az Cr Elevations
(m a.s.l.) and
status
Distribution
pattern
Main relevant
references
Genus Metamastigophorophyllon Ceuca, 1976
33. Metamastigophorophyllon giljarovi
(Lang, 1959)
W + 20–1850, se Ca Antić and Makarov
2016
34. M. hamatum Antić & Makarov,
2016
W + 150–2200, se Ca Antić and Makarov
2016
35. M. lamellohirsutum Antić &
Makarov, 2016
W 700–800, e Ca Antić and Makarov
2016
36. M. torsivum Antić & Makarov,
2016
G + 800–1700, se Ca Antić and Makarov
2016
Genus Paranotosoma Antić & Makarov, 2016
37. Paranotosoma attemsi Antić &
Makarov, 2016
W 1500–1800, e Ca Antić and Makarov
2016
38. P. cordatum Antić & Makarov,
2016
W 20–800, e Ca Antić and Makarov
2016
39. P. subrotundatum Antić &
Makarov, 2016
W, C + 350–850, se Ca Antić and Makarov
2016
Genus Pseudoagellophorella Antić & Makarov, 2016
40. Pseudoagellophorella eskovi Antić
& Makarov, 2016
C, E + + 100–2080, se Ca Antić and Makarov
2016
41. P. mirabilis Antić & Makarov,
2016
W 20–130, e Ca Antić and Makarov
2016
42. P. papilioformis Antić &
Makarov, 2016
E + 850–2100, se Ca Antić and Makarov
2016
Genus Ratcheuma Golovatch, 1985
43. Ratcheuma excorne Golovatch,
1985
W 1180, t, e Ca Golovatch 1984/85,
Antić and Makarov
2016
Order Julida
Family Blaniulidae
Genus Cibiniulus Verhoe, 1927
44. Cibiniulus phlepsii (Verhoe,
1897)
W + 20–130 EuM Engho 1984, 2006
Genus Nopoiulus Menge, 1851
45. Nopoiulus brevipilosus Engho,
1984
W 130, t, e Ca Engho 1984,
Golovatch and
Engho 1990
46. N. densepilosus Engho, 1984 W + + 1500–1700 Ca Engho 1984,
Golovatch and
Engho 1990
47. N. golovatchi Engho, 1984 W + 20–130, se Ca Engho 1984, 1990
48. N. kochii (Gervais, 1847) G + + + + 10–2200, i? sc Engho 1984,
Golovatch and
Engho 1990
Family Nemasomatidae
Genus Nemasoma C.L. Koch, 1847
49. Nemasoma caucasicum
(Lohmander, 1932)
G + + + + 20–2000, se Ca Kobakhidze 1965,
Engho 1985
Family Julidae
Genus Archileucogeorgia Lohmander, 1936
50. Archileucogeorgia abchasica
Lohmander, 1936
W 130, t, e Ca Lohmander 1936
51. A. satunini Lohmander, 1936 W 130, e Ca Lohmander 1936
Genus Brachyiulus Berlese, 1884
52. Brachyiulus lusitanus Verhoe,
1898`
C + 100, i M Lohmander 1936

Diversity and distribution patterns of the millipedes of Georgia 205
Fauna G R T Ar Az Cr Elevations
(m a.s.l.) and
status
Distribution
pattern
Main relevant
references
Genus Byzantorhopalum Verhoe, 1930
53. Byzantorhopalum rossicum
(Timotheew, 1897)
W? + + + 30–1500 EE Lohmander 1936,
Vagalinski and
Lazányi 2018
Genus Catamicrophyllum Verhoe, 1900
54. Catamicrophyllum caucasicum
(Attems, 1901)
G + + + 700–2000, se Ca Lohmander 1936,
Engho 1995
Genus Calyptophyllum Brolemann, 1922
55. Calyptophyllum sp. W 100? ? Lohmander 1936,
Engho 1995
Genus Chaetoleptophyllum Verhoe, 1898
56. Chaetoleptophyllum exum
Golovatch, 1979
G + 15–2200, se Ca Golovatch 1979,
Evsyukov et al.
(2020)
Genus Cylindroiulus Verhoe, 1894
57. Cylindroiulus bellus (Lignau,
1903)
W? + + 100 EM Lignau 1903,
Read 1992,
Chumachenko
2016
58. C. crassiphylacum Read, 1992 W, C + 600–1700, se Ca Read 1992
59. C. kacheticus Lohmander, 1936 E + 500–1250, se Ca Lohmander 1936,
Read 1992
60. C. olgainna Read, 1992 W 300–1100, e Ca Read 1992
61. C. parvus Lohmander, 1928 C, E + 500–2100, se Ca Lohmander 1936,
Read 1992
62. C. placidus (Lignau, 1903) W, C + 20–2200, se Ca Lignau 1903, Read
1992
63. C. pterophylacum Read, 1992 W, C + 20–1600, se Ca Read 1992, Zuev
2014
64. C. quadrus Read, 1992 W, C 700–1000, e Ca Read 1992
65. C. ruber (Lignau, 1903) W + 100–2000, se Ca Lignau 1903, 1915,
Read 1992
66. C. schestoperovi Lohmander,
1936
W + 400–1800, se Ca Lohmander 1936,
Read 1992
67. C. truncorum (Silvestri, 1896) W + + 130, i sc Read 1992
Genus Grusiniulus Lohmander, 1936
68. Grusiniulus redikorzevi
Lohmander, 1936
C 800–900, e Ca Lohmander 1936,
Vagalinski and
Lazányi 2018
Genus Julus Linnaeus, 1758
69. Julus colchicus Lohmander, 1936 G + + 20–2850, se Ca Lohmander 1936,
Engho 2006,
Evsyukov et al.
2018
70. J. kubanus Lohmander, 1936 W, E + 300–2100, se Ca Lohmander 1936,
Kobakhidze 1965,
Evsyukov et al.
2018
71. J. lignaui Verhoe, 1910 W + 1500–2800,
se
Ca Evsyukov et al.
2018
72. J. lindholmi Lohmander, 1936 W + 450–2200, se Ca Lohmander 1936,
Evsyukov et al.
2018

Mzia S. Kokhia & Sergei I. Golovatch / ZooKeys 930: 199–219 (2020)
206
Fauna G R T Ar Az Cr Elevations
(m a.s.l.) and
status
Distribution
pattern
Main relevant
references
Genus Kubaniulus Lohmander, 1936
73. Kubaniulus gracilis Lohmander,
1936
W + 20–700, se Ca Lohmander 1936,
Evsyukov et al.
2020
Genus Leptoiulus Verhoe, 1894
74. Leptoiulus hastatus Lohmander,
1932
C + 800–1530, se Ca Lohmander 1936,
Engho 2006,
Evsyukov et al.
2020
75. L. tanymorphus (Attems, 1901) C, E + + + 80–1800, se Ca Lohmander 1936,
Evsyukov et al.
2020
Genus Leucogeorgia Verhoe, 1930
76. Leucogeorgia longipes Verhoe,
1930
W 170, t, e Ca Verhoe 1930,
Barjadze et al. 2019
77. L. rediviva Golovatch, 1983 W 330, t, e Ca Golovatch 1983,
Barjadze et al. 2019
Genus Megaphyllum Verhoe, 1894
78. Megaphyllum dioscoriadis
(Lignau, 1915)
W + 130–1400, se Ca Lignau 1915,
Lohmander 1936,
Kobakhidze 1965,
Chumachenko
2016, Vagalinski
and Lazányi 2018
79. M. hercules (Verhoe, 1901) W + 20, i EM Lazányi and
Vagalinski 2013
80. M. spathulatum (Lohmander,
1936)
W? ? ? Ca Lohmander 1936,
Lazányi and
Vagalinski 2013
Genus Omobrachyiulus Lohmander, 1936
81. Omobrachyiulus adsharicus
(Lohmander, 1936)
W 20–30, e Ca Lohmander 1936,
Vagalinski and
Lazányi 2018
82. O. brachyurus (Attems, 1899) G + + + + 20–2500 EM Lohmander 1936,
Kobakhidze 1965,
Engho 2006,
Vagalinski and
Lazányi 2018
83. O. curvocaudatus (Lignau, 1903) W + 30–1700, se Ca Lohmander 1936,
Kobakhidze 1965,
Vagalinski and
Lazányi 2018
84. O. divaricatus (Lohmander,
1936)
G + 600 –2000, se Ca Lohmander 1936,
Kobakhidze 1965,
Vagalinski and
Lazányi 2018
85. O. hortensis (Golovatch, 1981) W 150, e Ca Golovatch 1981,
Vagalinski and
Lazányi 2018
86. O. implicitus Lohmander, 1936
(= O. i. ritsensis (Golovatch, 1981))
W + 400–1800, se Ca Lohmander 1936,
Chumachenko
2016, Vagalinski
and Lazányi 2018,
Vagalinski in litt.

Diversity and distribution patterns of the millipedes of Georgia 207
Fauna G R T Ar Az Cr Elevations
(m a.s.l.) and
status
Distribution
pattern
Main relevant
references
87. O. macrourus (Lohmander,
1928) (= O. m. abchasicus
(Lohmander, 1936))
W, C 130–2000, e Ca Lohmander 1936,
Kobakhidze 1965,
Vagalinski and
Lazányi 2018,
Vagalinski in litt.
Genus Pachyiulus Berlese, 1883
88. Pachyiulus avipes (C.L. Koch,
1847)
W + 30, i M Lohmander 1936
89. P. krivolutskyi Golovatch, 1977 W + 20–1800, se Ca Golovatch 1977,
Evsyukov 2016
Genus Syrioiulus Verhoe, 1914
90. Syrioiulus adsharicus
(Lohmander, 1936)
W 120, e Ca Lohmander 1936,
Golovatch 2018
91. S. georgicus (Lohmander, 1932) C 800–900, e Ca Lohmander 1932,
Golovatch 2018
Order Polydesmida
Family Paradoxosomatidae
Genus Oxidus Cook, 1911
92. Oxidus gracilis (C.L. Koch,
1847)
W + 20–100, i Ca Lignau 1915,
Lohmander 1936,
Chumachenko
2016
Genus Strongylosoma Brandt, 1833
93. Strongylosoma kordylamythrum
Attems, 1898
G + + 20–2200 Ca Lohmander 1936,
Kobakhidze 1965,
Evsyukov et al. 2016
94. S. lenkoranum Attems, 1898 C + + + 80–1650 Ca Lohmander 1936,
Kobakhidze 1965,
Evsyukov et al. 2016
Family Polydesmidae
Genus Brachydesmus Heller, 1858
95. Brachydesmus assimilis
Lohmander, 1936
C, E + + + 600–2800, se Ca Golovatch et al.
2016
96. B. furcatus Lohmander, 1936 W + 20–1900, se Ca Golovatch et al.
2016
97. B. kalischewskyi Lignau, 1915 G + + + + 50–2400, se Ca Golovatch et al.
2016
98. B. kvavadzei Golovatch,
Evsyukov & Reip, 2016
W 70–1520, e Ca Golovatch et al.
2016
99. B. simplex Golovatch, Evsyukov
& Reip, 2016
W + 20–1100, se Ca Golovatch et al.
2016
100. B. superus Latzel, 1884 W + 150–450, i sc Golovatch et al.
2016
Genus Polydesmus Latreille, 1803
101. Polydesmus abchasius Attems,
1899
W, C + 10–2230, se Ca Golovatch et al.
2016
102. P. lignaui Lohmander, 1936 W + 100–2200, se Ca Golovatch et al.
2016
103. P. mediterraneus Daday, 1889 W + 100, i EM Golovatch et al.
2016

Mzia S. Kokhia & Sergei I. Golovatch / ZooKeys 930: 199–219 (2020)
208
likely introduced species from the orders Polyxenida, Julida and Polydesmida that oc-
cur in the Caucasus. Even among the few unquestioned introductions, only Oxidus
gracilis (C.L. Koch, 1847) is an Oriental or East Asian alien element.
All species of Polyzoniida, Siphonocryptida, Glomerida and Chordeumatida, as
well as most species of Julida and Polydesmida appear to be native, endemic or sub-
endemic, but the genera and families they represent are widely distributed across the
Euro-Mediterranean Realm. As a result, endemism is profound at the species and, to a
lesser degree, generic levels. Most of the species (86, or 83%) show a Caucasian distri-
bution pattern, thus being endemic or subendemic to the Caucasus region. e same
pattern was found at the generic level, with 18 genera being endemic or subendemic
to the Caucasus, including all 14 genera of the order Chordeumatida that inhabit the
region (Antić and Makarov 2016, Antić et al. 2018). ere are neither families nor
orders of Diplopoda that are conned to the Caucasus region alone.
Our analysis of the distribution of Georgia’s millipedes is largely based on strictly
endemic and subendemic species (36 and 46, respectively: Table 1) and genera (8
and 9, respectively: Alpinella, Brachychaetosoma, Cryptacanthophorella, Georgiosoma,
Grusiniulus, Herculina, Leucogeorgia and Ratcheuma, vs. Adshardicus, Acanthophorella,
Archileucogeorgia, Caucaseuma, Dentatosoma, Heterocaucaseuma, Omobrachyiulus, Par-
anotosoma and Pseudoagellophorella). It shows that western Georgia, including Ab-
khazia and Ajaria – which are shown separately (Figs 9, 10) to more clearly depict
the localities/distributions and thus to avoid an “overcrowded” picture – supports the
richest and most diverse fauna (Figs 7–10). is is also the area where all 14 presumed
troglobionts are found in Georgia, all conned to karst caves (Barjadze et al. 2019).
Abkhazia, northwestern Georgia, is the richest subregion both in epigean and troglo-
bitic Diplopoda (Figs 7–9), hosting, among others, Heterocaucaseuma deprofundum
Antić & Makarov, 2018. is species is the world’s deepest record of a millipede, found
at 60–1980 m below the surface in the Krubera-Voronja and Sarma caves, Arabika
Massif, Abkhazia (Fig. 3). Both these caves are among the deepest globally and support
the second and third deepest subterranean invertebrate communities, respectively. Fur-
thermore, both harbour still one more diplopod species, a yet undescribed Leucogeorgia
sp. (Antić et al. 2018).
is picture is hardly surprising, as due to the proximity to the Black Sea the
climate of western Georgia is largely humid warm temperate. More easterly, the
climate is increasingly dry and hot, already dominating eastern Georgia (Bondyrev
et al. 2015). Following this trend, the millipede fauna is increasingly depauperate:
at least 79 diplopod species occur in western Georgia (= Colchis), but this number
drops down to 37 in the central and to 25 in the eastern parts of Georgia (Table 1,
Figs 7–12). Millipedes are mainly conned to forests in the Caucasus and in Georgia
reecting their terrestrial, meso- to hygrophilous, largely also calciphilous, arthropod
relationships which are historically, trophically and ecologically closely associated with
forested biomes (Golovatch and Kime 2009). Dry steppes and arid light forests in
central and eastern Georgia (Table 2), as well as the Colchidan swamps of western
Georgia support only very few millipede species. Especially tolerant to xeric conditions

Diversity and distribution patterns of the millipedes of Georgia 209
Figure 2. Map of Georgia with three transects (light blue), one each in the western, central and eastern
parts of the country, to crudely show both horizontal and vertical distributions of millipedes endemic or
subendemic to the country.
Figure 3. Map of western Georgia with its transect (light blue), Pitsunda – Arabika Plateau – Khaishi –
Bagdati – Batumi, and macro relief (bottom).

Mzia S. Kokhia & Sergei I. Golovatch / ZooKeys 930: 199–219 (2020)
210
Table 2. Vertical zonation of Georgia’s vegetation belts.
Vegetation belts Western Georgia, altitude
(m a.s.l.)
Eastern Georgia, altitude
(m a.s.l.)
deserts, dry steppes and arid light forests 150–600
forests 0–1900 600–1900
subalpine 1900–2500 1900–2500
alpine 2500–3100 2500–3000
subnival and nival 3100–3600 and > 3600 3000–3500 and > 3500
Figure 4. Map of central Georgia with its transect (light blue), Roki Tunnel – Tskhinvali – Tbilisi –
Tsalka Reservoir – Ninotsminda – Javakheti National Park, and its macro relief (bottom).
seems to be Leptoiulus tanymorphus (Attems, 1901) (Fig. 12), whereas both Hirudisoma
roseum (Victor, 1839) and Julus colchicus Lohmander, 1936 (Fig. 6), as well as several
Chordeumatida tend to represent particularly hydrophilous epigean species. Nearly all
cavernicoles (e.g., Leucogeorgia spp.) are likewise highly hydrophilous.
As noted above, due to the quite extensive karsts that blanket much of western
Georgia, in particular Abkhazia, Samegrelo, Racha Lechkhumi and Imereti, a large
proportion of the total fauna is taken up by true cavernicoles (14 species, or 13%). e
bulk, however, remains forest-dwelling millipedes and their woody habitats mainly
are more or less montane. Present-day Georgia enjoys a remarkable network of nature
reserves and national parks, with more than 1/3 of the entire national territory still

Diversity and distribution patterns of the millipedes of Georgia 211
Figure 5. Map of eastern Georgia with its transect (light blue), Omalo – Tianeti – Akhmeta – Shilda –
Kvareli – Lagodekhi – Tamariani, and its macro relief (bottom).
Figure 6. Map showing the distributions of four particularly widespread millipedes endemic or suben-
demic to Georgia. Designations: yellow ball (Hirudisoma roseum), green ball (Chaetoleptophyllum exum),
pink square (Metamastigophorophyllon torsivum), blue star (Julus colchicus).

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212
Figure 7. Map of western Georgia (= Colchis) showing the distributions of some endemic or subendemic
species. Designations: orange ball (Hyleoglomeris awchasica), red ball (Nopoiulus golovatchi), green triangle
(Cylindroiulus pterophylacum), red ring (Cylindroiulus ruber), white ball (Polydesmus abchasius), blue ball
(Trachysphaera fragilis), green star (Trachysphaera radiosa), pink ring (Cylindroiulus schestoperovi), yellow
ring (Hirudicryptus abchasicus).
covered with mountain forests. In contrast, its lowland woodlands have largely been
destroyed and long replaced by agri- or sylvicultures, as well as orchards and vineyards
(https://apa.gov.ge/en/protected-areas/national-park).
Following Gulisashvili (1964) and Nakhutsrishvili (2013), the altitudinal nature
zonation of Georgia can crudely be presented in a tabular form (Table 2). e zonation
varies quite clearly in dierent parts of Georgia (Fig. 1) in relation to climatic gradi-
ents. Central Georgia (Figs 1, 4), which is climatically closer to the eastern part of the
country, warrants recognition as a separate entity based at least on the distribution of
several endemic or subendemic species of Diplopoda (Fig. 11).
No transects are contained in Figures 6–12 to avoid an “overcrowded” presentation
of the numerous species distributions; however, these are easy to extrapolate from the
gures and thus to follow the general trends and variations in the macro relief of the
corresponding parts of Georgia. Only relatively few millipedes occur in subalpine to
alpine environments (usually ≥ 2200 m a.s.l.) in Georgia (Table 2). Yet nearly none of
them can be considered as being characteristic of the high altitudes, because the same
species appear to populate lower elevations as well, down to almost sea-level: Caucase-
uma variabile Antić & Makarov, 2016, Dentatosoma magnum Antić & Makarov, 2016,

Diversity and distribution patterns of the millipedes of Georgia 213
Figure 8. Map of western Georgia (= Colchis) showing the distributions of some other endemic or
subendemic species. Designations: green diamond (Georgiosoma bicornutum), white ball (Trachysphaera
minuta), orange diamond (Trachysphaera orientalis), red ball (Trachysphaera solida), red star (Typhloglom-
eris palatovi), red ring (Paranotosoma cordatum), blue star (Cylindroiulus quadrus), blue ball (Cylindroiulus
placidus), yellow square (Pachyiulus krivolutskyi), green star (Acanthophorella barjadzei), pink ring (Meta-
mastigophorophyllon lamellohirsutum), blue ring (Paranotosoma attemsi), yellow ring (Ratcheuma excorne),
white ring (Nemasoma caucasicum), green ring (Leucogeorgia longipes).
Metamastigophorophyllon hamatum Antić & Makarov, 2016, Chaetoleptophyllum exum
Golovatch, 1979, Cylindroiulus placidus (Lignau, 1903), Strongylosoma kordylamyth-
rum Attems, 1898, Brachydesmus assimilis Lohmander, 1936, B. kalischewskyi Lignau,
1915, Polydesmus abchasius Attems, 1899 and P. lignaui Lohmander, 1936. e same
concerns Omobrachyiulus brachyurus (Attems, 1899) and Catamicrophyllum caucasicum
(Attems, 1901), both of which occur also at ≤ 2500 m a.s.l. in the Caucasus Minor of
Armenia and Azerbaijan; the former species also in Dagestan, Russia, Caucasus Major
(personal observations). Nopoiulus kochii (Gervais, 1847) is a subcosmpolitan species,
common also throughout the Caucasus (10–2200 m a.s.l., Table 1), but because the
entire genus Nopoiulus is particularly diverse in the Caucasus region, the latter could
well have also been the origin centre of N. kochii (Golovatch and Engho 1990).
At the present, the only exception that may possibly be referred to as a high-montane el-
ement in the fauna of Georgia, as well as the entire Caucasus, seems to be Alpinella waltheri
Antić & Makarov, 2016 (2860 m a.s.l., Table 1, Map 12). Even though some species of Ju-
lus, i.e., Julus colchicus Lohmander, 1936 (20–2850 m a.s.l.), J. kubanus Lohmander, 1936
(300–2100 m a.s.l.) and J. lindholmi Lohmander, 1936 (450–2200 m a.s.l., Table 1, Figs 9,
12), mostly occur over a wide range of altitudes, J. lignaui Verhoe, 1910 (1500–2800 m
a.s.l.) is perhaps the sole congener that seems to be inclined to dwelling in high-mountain
environments. However, the paucity or even absence of unequivocally high-mountain ele-
ments in the Caucasus generally, and in Georgia in particular, requires conrmation, as our
knowledge of the millipede fauna of the regions concerned is still far from complete.

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214
Figure 10. Map of Ajaria showing the distributions of some endemic or subendemic species. Designa-
tions: blue ball (Adshardicus strasseri), red diamond (Brachydesmus kvavadzei), green ball (Dentatosoma den-
ticulatum), orange star (Dentatosoma zeraboseli), yellow square (Omobrachyiulus adsharicus), white triangle
(Omobrachyiulus divaricatus), yellow ball (Paranotosoma cordatum), yellow triangle (Syrioiulus adsharicus).
Figure 9. Map of Abkhazia showing the distributions of some endemic or subendemic species. Designa-
tions: red triangle (Brachychaetosoma turbanovi), blue square (Caucaseuma kelasuri), orange star (Archile-
ucogeorgia abchasica), pink star (Omobrachyiulus implicitus), orange square (Cylindroiulus olgainna), blue
ring (Paranotosoma subrotundatum), yellow star (Julus lindholmi), green star (Leucogeorgia rediviva), green
ring (Dentatosoma magnum), pink ring (Heterocaucaseuma deprofundum), orange diamond (Metamastigo-
phorophyllon giljarovi), white ring (Kubaniulus gracilis), blue star (Metamastigophorophyllon hamatum), red
ring (Pseudoagellophorella mirabilis), red diamond (Megaphyllum dioscoriadis), yellow square (Nopoiulus
brevipilosus), yellow triangle (Archileucogeorgia satunini), orange ball (Heterocaucaseuma longicorne), red
ball (Omobrachyiulus hortensis), blue ball (Brachydesmus furcatus), green ball (Brachydesmus simplex), yellow
ball (Polydesmus lignaui), green square (Heterocaucaseuma mauriesi).

Diversity and distribution patterns of the millipedes of Georgia 215
Figure 11. Map of central Georgia showing the distributions of some endemic or subendemic spe-
cies. Designations: blue ring (Brachydesmus kalischewskyi), yellow ring (Caucaseuma variable), green Ring
(Catamicrophyllum caucasicum), red ball (Cylindroiulus crassiphylacum), orange ring (Cylindroiulus ptero-
phylacum), white ring (Grusiniulus redikorzevi), yellow ball (Herculina oligosagittae), blue ball (Herculina
polysagittae), pink star (Leptoiulus hastatus), red diamond (Metamastigophorophyllon martensi), yellow star
(Omobrachyiulus macrourus (= O. m. abchasicus)), white star (Syrioiulus georgicus).
Figure 12. Map of eastern Georgia showing the distributions of some endemic or subendemic species.
Designations: green ball (Alpinella waltheri), red ball (Brachydesmus assimilis), blue triangle (Cylindroiulus
kacheticus), yellow ball (Cylindroiulus parvus), yellow ring (Hyleoglomeris specialis), orange square (Julus
kubanus), red diamond (Leptoiulus tanymorphus), white ball (Pseudoagellophorella eskovi), red ring (Pseu-
doagellophorella papilioformis).

Mzia S. Kokhia & Sergei I. Golovatch / ZooKeys 930: 199–219 (2020)
216
Conclusion
Ongoing research on the diplopod fauna of Georgia will undoubtedly reveal many more
species and rene their distributions. is particularly concerns several genera of Julidae,
including new cavernicolous and epigean ones (D. Antić, A. Evsyukov, B. Vagalinski,
personal communications). As a result, the present paper must only be taken as provi-
sional, marking the present state of the art and is certain to be updated in the near future.
Acknowledgements
is paper was executed in the framework of the project “Fauna of Georgia – Current
Trends and Conservation”. e second author was supported by the Presidium of the
Russian Academy of Sciences, Program No. 41 “Biodiversity of natural systems and
biological resources of Russia”.
We are particularly grateful to A.S. Kandaurov (Tbilisi) who helped us generate the
maps. Special thanks go to both reviewers, Hans Reip and Alexandr Evsyukov, for so
critically and positively evaluating our work.
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