CC BY
18
Anq ACTA BIOLOGICA SIBIRICA
# % Ajournai on the biodiversity of Siberia and
^^^^^^^^^^ the adjacent lands
RESEARCH ARTICLE
Acta Biologica Sibirica 9: 113-125 (2023) doi: 10.5281/zenodo.7725474 http://journal.asu.ru
Composition and structure of the fauna of ground beetles (Coleoptera, Carabidae) of the Zerafshan Range
Fazlitdin Khalimov1, 2
1 Gulistan State University, 4th microdistrict, Gulistan, 120100, Uzbekistan
2 Samarkand State University, 15 University Boulevard, Samarkand, 140100, Uzbekistan
Corresponding author: Fazlitdin Khalimov (xalimov1968@list.ru)
Academic editor: R. Yakovlev | Received 8 February 2023 | Accepted 23 February 2023 | Published 14 March 2023
http://zoobank.org/C53D0BCB-9C63-4605-A9F3-C791652C2830
Citation: Khalimov F (2023) Composition and structure of the fauna of ground beetles (Coleoptera, Carabidae) of the Zerafshan Range. Acta Biologica Sibirica 9: 113-125. https://doi.org/10.5281/zenodo.7725474
Abstract
The article presents the results of studying the fauna of ground beetles in the western part of the Zerafshan Range (Uzbekistan). The Carabidae list includes 81 species from 41 genera, 18 tribes and nine subfamilies. The ecological diversity of the ground beetle fauna is analyzed on three parts of the ridge. The greatest diversity of Carabidae is noted in the Karatepo Mountains (74 species), and the smallest in the Zirabulak-Ziyadin Mountains (54 species). The fauna of the ground beetles of the Chakalikalyan and Karatepa Mountains are closer to each other (Jaccard index - 0.79, Chekanovsky-S0rensen coefficient - 0.89). The lowest value of the similarity coefficients was noted between the Chakalikalyan and Zirabulak-Ziadin Mountains (0.64; 0.78). The most numerous are Anchomenus dorsalis (10.46%), Calathus ambiguus (8.23%), Amara aenea (7.48%) and Harpalus rufipes (6.03%).
Keywords
Degree of dominance, ground beetles, species abundance, species composition, Zeravshan Range
Introduction
Ground beetles are a very large and diverse taxon among beetles, and about 100 new species are discovered every year (Kotze et al. 2011). They are an important link
Copyright Fazlitdin Khalimov. 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.
in food chains and thus play a huge role in energy transformation (Schirmel et al. 2012). They are sensitive to environmental changes, showing strong habitat specificity and low ability to quickly spread to neighboring sites (Work et al. 2008; Koivula 2011). Therefore, ground beetles have been widely used for ecological monitoring (Skalski et al. 2016; Kosewska et al. 2016; Bell et al. 2017; K^dzior 2020).
Many species of ground beetles, as universal predators with a wide food spectrum, can act as an essential component of biological regulation (Symondson et al. 2002; Stiling and Cornelissen 2005). And in agricultural landscapes, they play a very significant role in reducing the number of pests (Koval and Guseva 2008, Halimov 2020) and weeds (Talarico et al. 2016, Deroulers and Bretagnolle 2019).
The beetle fauna of Central Asia is very diverse and is characterized by many endemic forms. So, in the Tien Shan mountain system, within the republics of Kazakhstan, Kyrgyzstan, Tajikistan and Uzbekistan, 819 species of ground beetles were noted. And the fauna of ground beetles of the Pamir-Alai mountain system, located in the southeastern part of Middle Asia, includes 510 species (Kryzhanovsky et al. 1995).
The invertebrate fauna of the Zeravshan Range is still poorly understood, although in recent years some works have appeared on individual groups (Khalimov 2020, Narzullaev 2022).
Information about the fauna of the ground beetles of the Zarafshan valley can be found in general faunal studies (Alimdzhanov and Bronshtein 1956) or for the southern slopes of the Zarafshan ridge, located on the territory of Tajikistan (Mikhailov 1998). We previously reported the distribution of 49 species of ground beetles in this area (Khalimov 2020).
The purpose of our research was to study the species diversity and taxonomic composition of the ground beetles of the Zerafshan Range within the Republic of Uzbekistan.
Materials and methods
The studies were carried out in the period 2018-2022. The Zarafshan range on the territory of Uzbekistan is divided into four parts: the Chakilkalyan, Karatepa, Zirab-ulak and Ziadin mountains (Fig.1). The Chakilkalyan Mountains are the eastern part of the Zerafshan Range within Uzbekistan. The average height of the Chakil-kalyan Mountains is 1000-2000 m. To the west of the Chakilkalyan Mountains are the Karatepa Mountains (the average height is 1000-2000 m). The western part of the Zerafshan ridge is formed by the Zirabulak and Ziadin mountains, which have the character of low mountains, the average height of which is 400-900 m (Zhu-maev 1989).
Each of the above-mentioned parts of the Zeravshan Range has a peculiar hydrothermal regime and vegetation cover.
Figure 1. Research map (1-Chakalikalyan mountains, 2-Karatepa mountains, 3-Zirabulak and Ziadin mountains).
The beetles were collected using soil traps, light traps, an exhauster, and manual collection. The ground beetle system is given according to the Cataloque of Life (Lorenz 2021). The examined specimens are kept in the Entomological collection of Samarkand State University.
The degree of dominance was determined according to the Renkonen scale (Renkonen 1944), where more than 5% are dominant species, from 2% to 5% are subdominant species, 1-2% are few, and less than 1% are rare species.
Statistical calculations of biodiversity were carried out according to the guidance of E.A. Dunaev (1997).
Result
Over the years of research, 2686 beetle specimens belonging to nine subfamilies, 18 tribes, 41 genera and 81 species were collected (Table 1).
Table 1. Taxonomic composition of ground beetles of the Zerafshan Range
Subfamily Tribe № Species name
Nebriinae Nebriini 1. Nebria psammophila Solsky, 1874
2. Notiophilus sublaevis Solsky 1873
Cicindelinae Cicindelini 3. Cicindela turkestanica Ballion, 1871
Subfamily Tribe № Species name
4. Cicindela fischeri M.Adams, 1817
Omophroninae Omophronini 5. Omophron rotundatum (Fabricius, 1777)
Carabinae Carabini 6. Calosoma sycophanta Linnaeus, 1758
7. Calosoma auropunctatum Gilber, 1833
8. Carabus fedtschenkoi Solsky, 1874
9. Carabus staudingeri Ganglbauer, 1886
Brachininae Brachinini 10. Brachinus bayardi Dejean, 1831
11. Brachinus brevicollis Motschulsky, 1844
12. Brachinus explodens Duftschmid, 1812
13. Mastax thermarum Steven, 1806
Scaritinae Clivinini 14. Clivina collaris Herbst, 1784
Scaritini 15. Scarites basiplicatus Heyden, 1884
16. Scarites procerus Fischer von Waldheim, 1828
17. Scarites subcylindricus Chaudoir, 1843
18. Scarites terricola Bonelli, 1813
Broscinae Broscini 19. Broscus asiaticus Ballion, 1871
20. Broscus punctatus Dejean, 1828
21. Craspedonotus margellanicus Kraatz, 1884
Trechinae Bembidiini 22. Asaphidion flavicorne Solsky, 1874
23. Bembidion abbreviatum Solsky, 1874
24. Bembidion almum J. Sahlberg, 1900
25. Bembidion insidiosum Solsky, 1874
26. Bembidwn kirgisorum Netolitzky, 1934
27. Bembidion lampros (Herbst 1784)
28. Bembidion luridicorne Solsky, 1874
29. Bembidion quadrimaculatum Linnaeus, 1761
30. Bembidion quadripustulatum Audinet-Serville, 1821
31. Bembidion piceocyaneum Solsky, 1874
32. Bembidion saxatile flavipalpe Netolitzky, 1930
Tachyini 33. Elaphropus tetraspilus (Solsky, 1874)
34. Tachyura euphratica Reitter, 1885
35. Tachys turkestanicus Csiki, 1928
Trechini 36. Trechus quadristriatus Schrank, 1781
37. Chlaenius circumscriptus (Duftschmid, 1812)
38. Chlaenius festivus Panzer, 1796
39. Chlaenius flavicornis Fischer, 1842
40. Chlaenius extensus Mannerheim, 1825
Subfamily Tribe № Species name
41. Chlaenius semicyaneus Solsky, 1874
42. Chlaenius tenuilimbatus Ballion, 1871
Harpalini 43. Acinopus laevigatus Menetries,1832
44. Acinopus striolatus Zoubkoff, 1833
45. Acupalpus parvulus (Sturm, 1825)
46. Anisodactylus binotatus Fabricius, 1787
47. Carenochyrus titanus Solsky,1874
48. Chilotomus usgentensis Schauberger, 1932
49. Ditomus calydonius Rossi, 1790
50. Dixus eremita Dejean, 1825
51. Dixus semicylindricus Piochard de la Brûlerie, 1872
52. Eocarterus chodshenticus Ballion, 1871
53. Harpalus distinguendus Duftschmid, 1812
54. Harpalus griseus Panzer, 1796
55. Harpalus rufipes De Geer, 1774
56. Harpulus rubripes Duftschmid, 1812
57. Harpalus tenebrosus Dejean 1829
58. Parophonus hirsutulus (Dejean, 1829)
59. Stenolophus abdominalis Mannerheim, 1844
Lebiini 60. Cymindis andreae Ménétriés, 1832
61. Cymindis quadrisignata Menetries, 1848
62. Lebia cyonocephala Linnaeus, 1758
63. Lebia festiva Faldermann, 1836
64. Lebia scapularis (Geoffroy, 1785)
Platynini 65. Agonum viridicupreum Goeze, 1777
66. Anchomenus dorsalis Pontoppidan, 1763
Pterostichini 67. Poecilus cupreus Linnaeus,1758
68. Poecilus liosomus Chaudoir, 1876
69. Poecilus longiventris Solsky, 1874
70. Poecilus sp.
71. Pterostichus niger Schaller, 1783
Sphodrini 72. Calathus ambiguus Paykull, 1790
73. Calathus melanocephalus Linnaeus, 1758
74. Calathus peltatus Kolenati, 1845
75. Dolichus halensis Schaller, 1783
76. Pseudotaphoxenus juvencus (Ballion, 1871)
77. Taphoxenus goliath Faldermann, 1836
Subfamily Tribe № Species name
Zabrini 78. Amara aenea De Geer,1774
79. Amara ovata Fabricius, 1792
80. Amara similata Gyllenhal, 1810
81. Zabrus morio Mandrias, 1832
The most representative in terms of the number of species is the subfamily Har-palinae, which includes 45 species (55.6% of the total species diversity). In terms of the number of individuals, the subfamily Harpalinae accounts for 79.3% of all collected ground beetles. Among the representatives of this subfamily, the tribe Har-palini has the largest number of species, which is represented by 17 species from 11 genera. Among them, Acinopus laevigatus, Eocarterus chodshenticus, Harpalus rufipes and Harpalus griseus are numerous.
The tribe Sphodrini is represented by six species from four genera, the most numerous of which are Calathus ambiguus and Pseudotaphoxenus juvencus. The tribe Pterostichini is represented by five species from two genera (Pterostichus niger and four species from the genus Poecilus).
Six species from the tribe Chlaeniini belonging to the genus Chlaenius were identified. The most common are Chlaenius extensus. The tribe Lebiini is represented by five species and the tribe Zabrini by four species (three species from the genus Amara and one species from the genus Zabrus). Amara aenea and Zabrus morio are the dominant species of carabid fauna.
In the Karabidocomplex of the Zerafshan Range, 15 species belonging to three tribes and five genera are found from the subfamily Trechinae. The tribe Trechini is represented by a single species, Trechus quadristriatus, and the tribe Tachyini by three species. The tribe Bembidiini includes 11 species (genus Asaphidion - one species, genus Bembidion - ten species). Although these species are widespread and numerous in agrolandscapes, their abundance in the studied ridge is low. In general, representatives of the subfamily Trechinae account for 10.46% of all ground beetles and 18.5% of the total species diversity.
Five species of beetles from the subfamily Scaritinae belonging to the genera Scarites (four species) and Clivina (one species) were identified. The subfamilies Carabinae and Brachininae are represented by four species each (the genus Cara-bus and Calosoma each have two species, three species from the genus Brachinus and one species from the genus Mastax). The subfamily Broscinae is represented by three species from the genera Broscus and Craspedonotus.
In the complex of ground beetles of the studied territories, two representatives each have the subfamilies Nebriinae and Cicindelinae, and the only species is the subfamily Omophroninae.
The dominant species of the carabid fauna of the Zerafshan Range are An-chomenus dorsalis (10.46%), Calathus ambiguus (8.23%), Amara aenea (7.48%),
Harpalus rufipes (6.03%). The habitats of subdominant species are rich in Zabrus morio (4.77%), Eocarterus chodshenticus (4.54%), Pseudotaphoxenus juvencus (4.47%), and Harpalus griseus (4.36%).
Very few are Brachinus bayardi, Broscus punctatus, Chlaenius circumscriptus, Chlaenius tenuilimbatus, Chilotomus usgentensis, Craspedonotus margellanicus, Ditomus calydonius, Notiophilus sublaevis and Taphoxenus goliath, whose dominance is less than 0.1% (Table 2).
Table 2. Diversity and abundance of Carabidae in three mountains of the Zeravshan Range
№ Species name Chakil-kalyan Karatepa Zirabulak-Ziadin Total Degree of dominance, %
1. Acinopus laevigatus 32 53 16 101 3.76
2. Acinopus striolatus 12 6 4 22 0.82
3. Acupalpus parvulus 3 2 - 5 0.19
4. Agonum viridicupreum - 4 2 6 0.22
5. Amara aenea 65 112 24 201 7.48
6. Amara ovata 1 4 2 7 0.26
7. Amara similata 2 4 6 12 0.45
8. Anchomenus dorsalis 94 119 68 281 10.46
9. Anisodactylus binotatus - 4 - 4 0.15
10. Asaphidion flavicorne 9 12 9 30 1.12
11. Bembidion abbreviatum 3 2 1 6 0.22
12. Bembidion almum 5 3 2 10 0.37
13. Bembidion insidiosum 6 4 - 10 0.37
14. Bembidion lampros 7 5 - 12 0.45
15. Bembidion luridicorne 13 9 9 31 1.15
16. Bembidion piceocyaneum 2 2 1 5 0.19
17. Bembidion quadrimaculatum 6 7 5 18 0.67
18. Bembidion quadripustulatum 2 3 1 6 0.22
19. Bembidion saxatile flavipalpe 12 7 2 21 0.78
20. Bembidwn kirgisorum 3 1 2 6 0.22
21. Brachinus brevicollis 8 10 12 30 1.12
22. Brachinus explodens 19 38 12 69 2.57
23. Brachinus bayardi 2 - - 2 0.07
24. Broscus asiaticus 15 6 3 24 0.89
25. Broscus punctatus 1 1 - 2 0.07
26. Calathus ambiguus 121 60 40 221 8.23
27. Calathus melanocephalus 7 11 4 22 0.82
28. Calathus peltatus 14 12 8 34 1.27
№ Species name Chakil-kalyan Karatepa Zirabulak-Ziadin Total Degree of dominance, %
29. Calosoma auropunctatum 5 7 - 12 0.45
30. Calosoma sycophanta 2 2 - 4 0.15
31. Carabus fedtschenkoi 9 11 - 20 0.74
32. Carabus staudingeri 1 3 - 4 0.15
33. Carenochyrus titanus 4 2 2 8 0.30
34. Chilotomus usgentensis - 2 - 2 0.07
35. Chlaenius festivus 4 2 2 8 0.30
36. Chlaenius circumscriptus - 2 - 2 0.07
37. Chlaenius flavicornis 12 7 5 24 0.89
38. Chlaenius semicyaneus 1 2 - 3 0.11
39. Chlaenius tenuilimbatus 1 1 - 2 0.07
40. Chlaenius extensus 29 10 21 60 2.23
41. Cicindela turkestanica 9 10 7 26 0.97
42. Cicindela fischeri - 4 - 4 0.15
43. Clivina collaris 8 9 3 20 0.74
44. Craspedonotus margellanicus - 1 - 1 0.04
45. Cymindis quadrisignata 9 17 26 52 1.94
46. Cymindis andreae 3 4 3 10 0.37
47. Ditomus calydonius - - 2 2 0.07
48. Dixus eremita 15 19 8 42 1.56
49. Dixus semicylindricus 3 3 - 6 0.22
50. Dolichus halensis 11 9 9 29 1.08
51. Eocarterus chodshenticus 37 54 31 122 4.54
52. Harpalus distinguendus 14 13 23 50 1.86
53. Harpalus griseus 42 21 54 117 4.36
54. Harpalus rufipes 40 64 58 162 6.03
55. Harpulus rubripes 11 10 4 25 0.93
56. Harpalus tenebrosus 3 - - 3 0.11
57. Lebia cyonocephala 6 14 - 20 0.74
58. Lebia scapularis - 8 - 8 0.30
59. Lebia festiva 3 - - 3 0.11
60. Mastax thermarum - 2 2 4 0.15
61. Nebria psammophila 5 3 - 8 0.30
62. Notiophilus sublaevis 2 - - 2 0.07
63. Omophron rotundatum - - 3 3 0.11
64. Parophonus hirsutulus - 5 3 8 0.30
65. Poecilus cupreus 7 7 6 20 0.74
№ Species name Chakilkalyan Karatepa Zirabulak-Ziadin Total Degree of dominance, %
66. Poecilus liosomus 14 16 10 40 1.49
67. Poecilus longiventris 14 17 11 42 1.56
68. Poecilus sp. 4 8 1 13 0.48
69. Pseudotaphoxenus juvencus 44 58 18 120 4.47
70. Pterostichus niger 17 12 11 40 1.49
71. Scarites basiplicatus 4 8 - 12 0.45
72. Scarites procerus - - 11 11 0.41
73. Scarites subcylindricus - 3 - 3 0.11
74. Scarites terricola 6 3 4 13 0.48
75. Stenolophus abdominalis 22 17 3 42 1.56
76. Tachys turkestanicus 4 3 5 12 0.45
77. Tachyura euphratica - 3 1 4 0.15
78. Tachyura tetraspila 4 4 - 8 0.30
79. Taphoxenus goliath - 2 - 2 0.07
80. Trechus quadristriatus 35 21 46 102 3.80
81. Zabrus morio 45 59 24 128 4.77
Number of individuals 973 1063 650 2686 100.00
Number of species 66 74 54 81 -
Species richness, DMg 9.45 10.47 8.18 10.13 -
Species richness, DMn 2.12 2.27 2.12 1.56 -
Diversity indicator Shannon, H 3.54 3.55 3.39 3.60 -
Shannon equitability index, Eh 0.84 0.83 0.85 0.82 -
Simpson's Diversity Index, D (S\) 0.045 0.046 0.047 0.043 -
Berger - Parker dominance index, d 0.124 0.112 0.10 0.105 -
As noted above, the Zeravshan Range is divided into the Chakilkalyan, Karatepa, Zirabulak and Ziadin mountains. The greatest diversity of ground beetles was noted on the Karatepa mountains (74 species), and the least on the Zirabulak and Ziadin mountains (54 species). The Chakilkalyan mountains occupies an intermediate position (66 species). However, there were no significant differences in the indices of ecological diversity of the beetle fauna in these territories (Table 2).
An analysis of the similarity of the fauna of the studied territories shows that the fauna of the ground beetles of the Chakalikalyan and Karatepa mountains are closer to each other (Jaccard index - 0.79, Chekanovsky-Sorensen coefficient - 0.89) (Table 3). The lowest value of the similarity coefficients was noted between the Chakalikalyan and Zirabulak-Ziadin mountains (0.64; 0.78).
Table 3. Similarity of species composition of Carabidae in three areas of the Zeravshan Range (Jaccard / Chekanovsky-Sorensen indices)
Mountains Karatepa Chakilkalyan Zirabulak-Ziadin
Karatepa - 0.89 0.8
Chakilkalyan 0.79 - 0.78
Zirabulak-Ziadin 0.66 0.64 -
The species Ditomus calydonius, Omophron rotundatum and Scarites procerus are unique for the Zirabulok-Ziyadin Mountains and are found only in this part of the Zerafshan Range. Eight species of ground beetles were found only in the Karatepa Mountains (Anisodactylus binotatus, Chilotomus usgentensis, Chlaenius circumscriptus, Cicindela fischeri, Craspedonotus margellanicus, Lebia scapularis, Scarites subcylindricus and Taphoxenus goliath). Brachinus bayardi, Harpalus tene-brosus, Lebia festiva and Notiophilus sublaevis were found only in the Chakalikalyan Mountains.
In these three areas of the Zeravshan Range, the degree of dominance of individual species can vary significantly. Thus, Calathus ambiguus (12.44%) is the most numerous in the Chakalikalyan Mountains. The habitats of dominant species are also Anchomenus dorsalis (9.66%) and Amara aenea (6.68%). Subdominants also have a high degree of dominance: Zabrus morio (4.62%), Pseudotaphoxenus juven-cus (4.52%), Harpalus griseus (4.32%) and Harpalus rufipes (4.11%).
In the Karatepa Mountains, the dominant species are located in the following order: Anchomenus dorsalis (11.19%), Amara aenea (10.54%), Harpalus rufipes (6.02%), Calathus ambiguus (5.64%), Zabrus morio (5.55%), Pseudotaphoxenus ju-vencus (5.46% ), Eocarterus chodshenticus (5.08%) and Acinopus laevigatus (4.99%). Thus, Eocarterus chodshenticus and Acinopus laevigatus join the dominant species, and Harpalus rufipes is excluded from the dominant species.
And on the Zirabulak-Ziadin mountains, the dominant species are located in the following order: Anchomenus dorsalis (10.46%), Harpalus rufipes (8.92%), Harpalus griseus (8.31%), Trechus quadristriatus (7.08), Calathus ambiguus (6.15%), Eocarterus chodshenticus ( 4.77%) and Cymindis quadrisignata (4.0%). Here, there are no dominant species of Zabrus morio and Pseudotaphoxenus juvencus, but Trechus quadristriatus and Cymindis quadrisignata appear among the dominant species. However, in this section of the Zeravshan Range, the number of ground beetles, which are widely distributed in the agricultural landscapes Harpalus rufipes, Harpalus griseus and Trechus quadristriatus, is increasing.
The number of dominant species can vary significantly at different points within the same mountain, depending on the height of the area. So, as the height of the area increases, the abundance of Pseudotaphoxenus juvencus and Eocarterus chod-shenticus increases, while the other dominant species show an opposite trend, that is, as the height increases, a decrease in abundance is observed. It should be noted
that altitudinal belts affect not only the diversity and abundance of ground beetles, but also the morphometric features of individual species (Zokirova and Khalimov 2022).
Conclusion
For the first time, a list of ground beetles of the Zeravshan Range was compiled, including 81 species. A comparative analysis of the ecological diversity of the car-abid fauna of the Chakaliyan, Karatepa and Zirabulak mountains, which are part of the ridge, was carried out. Dominant and rare species were identified. Among rare beetle species in need of protection are Brachinus bayardi, Broscus punctatus, Chlaenius circumscriptus, Chlaenius tenuilimbatus, Chilotomus usgentensis, Craspe-donotus margellanicus, Ditomus calydonius, Notiophilus sublaevis and Taphoxenus goliath. However, further research is required to fully understand the state of the populations of these species.
Acknowledgements
The author expresses his sincere gratitude to I. Kabak (St. Petersburgh) and R. Dud-ko (Novosibirsk) for their help in species identification.
References
Alimdjanov RA, Bronshteyn TG (1956) Invertebrate animals of the Zarafshan valley. A systematic list of species indicating useful and harmful forms. Publishing House of the Academy of Sciences of the UzSSR, Samarkand, 348 pp. [In Russian] Bell AJ, Phillips ID, Nielsen SE, Spence JR (2017) Species traits modify the species area relationship in ground-beetle (Coleoptera: Carabidae) assemblages on islands in a boreal lake. PLoS One 12 (12), e0190174. https://doi.org/10.1371/journal.pone.0190174 Deroulers P, Bretagnolle V (2019) The consumption pattern of 28 species of carabid beetles (Carabidae) to a weed seed, Viola arvensis. Bulletin of Entomological Research 109 (2): 229-235. http://dx.doi.org/10.1017/S0007485318000457 Dunaev EA (1997) Methods of ecological and entomological research. MosgorSYUN, Moscow, 44 pp. [In Russian] Halimov F (2020) Seasonal dynamics of dominant species of soil predators (Coleoptera: Carabidae,Staphylinidae) in agrolandscapes and their potential gluttony. Travaux du Muséum National d'Histoire Naturelle "Grigore Antipa" 63 (2): 175-187. https://doi. org/10.3897/travaux.63.e54116
K^dzior R, Szwalec A, Mundala P, Skalski T (2020) Ground beetle (Coleoptera, Carabidae) life history traits as indicators of habitat recovering processes in postindustrial areas. Ecological Engineering 5 (142): 105615. https://doi.org/10.1016/j.ecoleng.2019.105615 Khalimov F (2020) The ground beetles (Coleoptera, Carabidae) of the Karatepa and Chakilkalyan mountains (west part of Zarafshan Mountains Range, Uzbekistan). Biosystems Diversity 28 (3): 265-271. http://dx.doi.org/10.15421/012035 Koivula MJ (2011) Useful model organisms, indicators, or both? Ground beetles (Coleoptera, Carabidae) reflecting environmental conditions. Zookeys 100: 287-317. https:// doi.org/10.3897/zookeys.100.1533 Kosewska A, Nietupski M, Nijak K, Sklaski T (2016) Effect of plant protection on assemblages of ground beetles (Coleoptera, Carabidae) in pea (Pisum L.) and lupine (Lupi-nus L.) crops. Periodicum Biologorum 118 (3): 213-222. http://dx.doi.org/10.18054/ pb.2016.118.3.3911
Kotze J, Brandmayr P, Casale A, Dauffy-Richard E, Dekoninck W, Koivula M, Lovei B, Mossakowski D, Noordijk J, Paarmann W, Pizzoloto R, Saska P, Schwerk A, Serrano J, Szyszko J, Palomares A, Turin H, Venn S, Vermeulen R, Zetto Brandmayr T (2011) Forty years of carabid beetle research in Europe - from taxonomy, biology, ecology and population studies to bioindication, habitat assessment and conservation. ZooKeys 5 (100): 55-148. https://doi.org/10.3897/zookeys.100.1523 Koval AG, Guseva OG (2008) Structure of the ground beetle (Coleoptera, Carabidae) complexes in potato field agrocenoses on different soils in Northwestern Russia. Entomological Review 88: 269-276. https://doi.org/10.1134/S0013873808030020 Kryzhanovskij OL, Belousov IA, Kabak II, Kataev BM, Makarov KV, Shilenkov VG (1995) A checklist of the ground-beetles of Russia and adjacent lands (Insecta, Coleoptera, Carabidae). Pensoft, Sofia - Moscow. http://dx.doi.org/10.13140/RG.2.1.1128.3688 Lorenz W (2021) Carabcat database. In: Catalogue of Life Checklist (v.03 (08/2021)). https://
doi.org/10.48580/dfqf-3dk Mihaylov VA (1998) Ground beetles (Coleoptera, Carabidae) of the southeast of Central Asia (fauna, ecology, values in biocenoses). Abstract of the dissertation for the degree of Doctor of Biological Sciences. St. Petersburg, 35 pp. [In Russian] Narzullayev SB (2022) New data on the vertical distribution of nematode communities in mountain ecosystems of Mount Zarafshan, Uzbekistan. Biodiversitas Journal of Biological Diversity 23 (8): 3967-3975. https://smujo.id/biodiv/article/view/11451/6011 Renkonen O (1944) Die Carabiden und Staphylinidenbestánde eines Seeufers in SW-Finn-
land. Annales Entomologici Fennici 9 (10): 23-31. Schirmel J, Blindow I, Buchholz S (2012) Life-trait and functional diversity patterns of ground beetles and spiders along a coastal headland successional gradient. Basic And Applied Ecology 13: 606-614. https://doi.org/10.1016/j.baae.2012.08.015 Skalski T, K^dzior R, Wyzga B, Radecki-Pawlik A, Plesiñski K, Zawiejska J (2016) Impact of incision of gravel-bed rivers on ground beetle assemblages. River Research and Applications 32: 1968-1977. https://doi.org/10.1002/rra.3027
Stiling P, Cornelissen T (2005) What makes a successful biocontrol agent? A metaanalysis of biological control agent performance. Biological Control 34 (3): 236-246. https://doi. org/10.1016/j.biocontrol.2005.02.017 Symondson WOC, Sunderland KD, Greenstone MH (2002) Can generalist predators be effective biocontrol agents? Annual Review of Entomology 47 (1): 561-594. https://doi. org/10.1146/annurev.ento.47.091201.145240 Talarico F, Giglio A, Pizzolotto R, Brandmayr P (2016) A synthesis of feeding habits and reproduction rhythm in Italian seed-feeding ground beetles (Coleoptera: Carabidae). European Journal of Entomology 113 (1): 325-336. http://dx.doi.org/10.14411/ eje.2016.042
Work TT, Koivula M, Klimaszewski J, Langor C, Spence JR, Sweeney J, Hebert C (2008) Evaluation of carabid beetles as indicators of forest change in Canada. Canadian Entomologist 140: 393-414. https://doi.org/10.4039/n07-LS07 Zhumaev T (1989) Mountains of Uzbekistan. Mehnat, Tashkent, 217 pp. [In Russian] Zokirova DF, Khalimov FZ (2022) Morphometric features of the beetle Acinopus (Acino-pus) laevigatus Menetries, 1832 (Coleoptera, Carabidae) in the mountain ecosystems of Uzbekistan. Bulletin of the Iraq natural History Museum 17 (2): 141-153. https://doi. org/10.26842/binhm.7.2022.17.2.0141
