Corresponding author: Sergei I. Golovatch (
Academic editor: Z. Korsós
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
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.
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.
Geographic division of Georgia.
The millipede fauna of Georgia has recently been reviewed and shown to comprise 95 species from 42 genera, 12 families, and 7 orders (
Only described species and published records are considered in our paper, while dubious taxa and those not identified to the species level have been omitted both from the checklist and reference list. Only one important exception has been made:
Three zigzag transects chosen to grossly reflect the north-to-south lie of the macro relief of Georgia, extending from the Caucasus Major in the north to the Caucasus Minor in the south (Figs
Most of the colour maps were generated using Google Earth Pro version 7.3.2.5495 and Adobe Photoshop CS6. The final images were processed with Adobe Photoshop CS6.
The diplopod fauna of the Caucasus region, including Georgia, is basically Euro-Mediterranean in its composition (Table
A revised checklist of the
Fauna | G | R | T | Ar | Az | Cr | Elevations (m a.s.l.) and status | Distribution pattern | Main relevant references |
---|---|---|---|---|---|---|---|---|---|
Class |
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Order |
|||||||||
Family |
|||||||||
Genus |
|||||||||
1. |
W | + | + | 20–1700, i | sc | ||||
2. |
E | + | + | 100–800 | Ca |
|
|||
Genus |
W | ||||||||
3. |
G | + | + | + | + | + | 20–1700 | EM |
|
Family |
|||||||||
Genus |
|||||||||
4. |
G | + | 20–900 | EM | |||||
Order |
|||||||||
Family |
|||||||||
Genus |
|||||||||
5. |
G | + | + | + | 20–1100, se | EM |
|
||
Order |
|||||||||
Family |
|||||||||
6. |
W | + | 600–1500, se | Ca | |||||
Order |
|||||||||
Family |
|||||||||
Genus |
|||||||||
7. |
W | + | 20–2100, se | Ca | |||||
8. |
E | + | 500–1400, se | Ca |
|
||||
Genus |
|||||||||
9. |
G | + | + | + | + | + | 20–2000 | EuM | |
10. |
G | + | 80–460, t, e | Ca | |||||
11. |
G | + | + | + | 20–1700, se | Ca | |||
12. |
W | 800–1100, t, e | Ca | ||||||
13. |
W | + | 20–1800, se | Ca | |||||
14. |
W, C | 20–2020, se | Ca | ||||||
Family |
|||||||||
Genus |
|||||||||
15. |
C, E | + | + | 600–1450, se | Ca | ||||
16. |
W | 650, t, e | Ca |
|
|||||
Order |
|||||||||
Family |
|||||||||
Genus |
|||||||||
17. |
W | 1120–1200, t, e | Ca |
|
|||||
Genus |
|||||||||
18. |
W | + | 20–530, se | Ca | |||||
Genus |
|||||||||
19. |
E | 2860, e | Ca |
|
|||||
Genus |
|||||||||
20. |
W | 300, t, e | Ca |
|
|||||
Genus |
|||||||||
21. |
W | 190, e | Ca |
|
|||||
22. |
C | + | 100–2500, se | Ca |
|
||||
Genus |
|||||||||
23. |
W, C | 800–1700, e | Ca |
|
|||||
Genus |
|||||||||
24. |
W | 400–900, e | Ca |
|
|||||
25. |
W | + | 20–2200, se | Ca |
|
||||
26. |
W | 20–1700, e | Ca |
|
|||||
Genus |
|||||||||
27. |
W | 2000, t, e | Ca |
|
|||||
Genus |
|||||||||
28. |
W | 1500–1700, e | Ca |
|
|||||
29. |
C | 1750, e | Ca |
|
|||||
Genus |
|||||||||
30. |
W | 2000–2100, t, e | Ca |
|
|||||
31. |
W | 100–350, t, e | Ca | ||||||
32. |
W | 215, t, e | Ca | ||||||
Genus |
|||||||||
33. |
W | + | 20–1850, se | Ca |
|
||||
34. |
W | + | 150–2200, se | Ca |
|
||||
35. |
W | 700–800, e | Ca |
|
|||||
36. |
G | + | 800–1700, se | Ca |
|
||||
Genus |
|||||||||
37. |
W | 1500–1800, e | Ca |
|
|||||
38. |
W | 20–800, e | Ca |
|
|||||
39. |
W, C | + | 350–850, se | Ca |
|
||||
Genus |
|||||||||
40. |
C, E | + | + | 100–2080, se | Ca |
|
|||
41. |
W | 20–130, e | Ca |
|
|||||
42. |
E | + | 850–2100, se | Ca |
|
||||
Genus |
|||||||||
43. |
W | 1180, t, e | Ca | ||||||
Order |
|||||||||
Family |
|||||||||
Genus |
|||||||||
44. |
W | + | 20–130 | EuM | |||||
Genus |
|||||||||
45. |
W | 130, t, e | Ca | ||||||
46. |
W | + | + | 1500–1700 | Ca | ||||
47. |
W | + | 20–130, se | Ca | |||||
48. |
G | + | + | + | + | 10–2200, i? | sc | ||
Family |
|||||||||
Genus |
|||||||||
49. |
G | + | + | + | + | 20–2000, se | Ca | ||
Family |
|||||||||
Genus |
|||||||||
50. |
W | 130, t, e | Ca |
|
|||||
51. |
W | 130, e | Ca |
|
|||||
Genus |
|||||||||
52. |
C | + | 100, i | M |
|
||||
Genus |
|||||||||
53. |
W? | + | + | + | 30–1500 | EE | |||
Genus |
|||||||||
54. |
G | + | + | + | 700–2000, se | Ca | |||
Genus |
|||||||||
55. |
W | 100? | ? | ||||||
Genus |
|||||||||
56. |
G | + | 15–2200, se | Ca | |||||
Genus |
|||||||||
57. |
W? | + | + | 100 | EM | ||||
58. |
W, C | + | 600–1700, se | Ca |
|
||||
59. |
E | + | 500–1250, se | Ca | |||||
60. |
W | 300–1100, e | Ca |
|
|||||
61. |
C, E | + | 500–2100, se | Ca | |||||
62. |
W, C | + | 20–2200, se | Ca | |||||
63. |
W, C | + | 20–1600, se | Ca | |||||
64. |
W, C | 700–1000, e | Ca |
|
|||||
65. |
W | + | 100–2000, se | Ca | |||||
66. |
W | + | 400–1800, se | Ca | |||||
67. |
W | + | + | 130, i | sc |
|
|||
Genus |
|||||||||
68. |
C | 800–900, e | Ca | ||||||
Genus |
|||||||||
69. |
G | + | + | 20–2850, se | Ca | ||||
70. |
W, E | + | 300–2100, se | Ca | |||||
71. |
W | + | 1500–2800, se | Ca |
|
||||
72. |
W | + | 450–2200, se | Ca | |||||
Genus |
|||||||||
73. |
W | + | 20–700, se | Ca | |||||
Genus |
|||||||||
74. |
C | + | 800–1530, se | Ca | |||||
75. |
C, E | + | + | + | 80–1800, se | Ca | |||
Genus |
|||||||||
76. |
W | 170, t, e | Ca | ||||||
77. |
W | 330, t, e | Ca | ||||||
Genus |
|||||||||
78. |
W | + | 130–1400, se | Ca | |||||
79. |
W | + | 20, i | EM |
|
||||
80. |
W? | ? | ? | Ca | |||||
Genus |
|||||||||
81. |
W | 20–30, e | Ca | ||||||
82. |
G | + | + | + | + | 20–2500 | EM | ||
83. |
W | + | 30–1700, se | Ca | |||||
84. |
G | + | 600 –2000, se | Ca | |||||
85. |
W | 150, e | Ca | Golovatch 1981, |
|||||
86. |
W | + | 400–1800, se | Ca | |||||
87. |
W, C | 130–2000, e | Ca | ||||||
Genus |
|||||||||
88. |
W | + | 30, i | M |
|
||||
89. |
W | + | 20–1800, se | Ca | |||||
Genus |
|||||||||
90. |
W | 120, e | Ca | ||||||
91. |
C | 800–900, e | Ca | ||||||
Order |
|||||||||
Family |
|||||||||
Genus |
|||||||||
92. |
W | + | 20–100, i | Ca | |||||
Genus |
|||||||||
93. |
G | + | + | 20–2200 | Ca | ||||
94. |
C | + | + | + | 80–1650 | Ca | |||
Family |
|||||||||
Genus |
|||||||||
95. |
C, E | + | + | + | 600–2800, se | Ca |
|
||
96. |
W | + | 20–1900, se | Ca |
|
||||
97. |
G | + | + | + | + | 50–2400, se | Ca |
|
|
98. |
W | 70–1520, e | Ca |
|
|||||
99. |
W | + | 20–1100, se | Ca |
|
||||
100. |
W | + | 150–450, i | sc |
|
||||
Genus |
|||||||||
101. |
W, C | + | 10–2230, se | Ca |
|
||||
102. |
W | + | 100–2200, se | Ca |
|
||||
103. |
W | + | 100, i | EM |
|
All species of
Our analysis of the distribution of Georgia’s millipedes is largely based on strictly endemic and subendemic species (36 and 46, respectively: Table
This 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 (
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%). The 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 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 (
Following
No transects are contained in Figures
At the present, the only exception that may possibly be referred to as a high-montane element in the fauna of Georgia, as well as the entire Caucasus, seems to be
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.
Map of western Georgia with its transect (light blue), Pitsunda – Arabika Plateau – Khaishi – Bagdati – Batumi, and macro relief (bottom).
Map of central Georgia with its transect (light blue), Roki Tunnel – Tskhinvali – Tbilisi – Tsalka Reservoir – Ninotsminda – Javakheti National Park, and its macro relief (bottom).
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 |
Map of eastern Georgia with its transect (light blue), Omalo – Tianeti – Akhmeta – Shilda – Kvareli – Lagodekhi – Tamariani, and its macro relief (bottom).
Map showing the distributions of four particularly widespread millipedes endemic or subendemic to Georgia. Designations: yellow ball (
Map of western Georgia (= Colchis) showing the distributions of some endemic or subendemic species. Designations: orange ball (
Map of western Georgia (= Colchis) showing the distributions of some other endemic or subendemic species. Designations: green diamond (
Map of Abkhazia showing the distributions of some endemic or subendemic species. Designations: red triangle (
Map of Ajaria showing the distributions of some endemic or subendemic species. Designations: blue ball (
Map of central Georgia showing the distributions of some endemic or subendemic species. Designations: blue ring (
Map of eastern Georgia showing the distributions of some endemic or subendemic species. Designations: green ball (
Ongoing research on the diplopod fauna of Georgia will undoubtedly reveal many more species and refine their distributions. This particularly concerns several genera of
This paper was executed in the framework of the project “Fauna of Georgia – Current Trends and Conservation”. The 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.