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Methodology for a local fauna study of ground beetles (Coleoptera, Carabidae) in the forest-tundra zone of the Polar Urals, Russia
expand article infoOD Kovalev, NA Zubriy§, BYu Filippov
‡ Northern (Arctic) Federal University, Archangelsk, Russia
§ Federal Research Centre for Integrated Studies of the Arctic of the Russian Academy of Sciences, Arkhangelsk, Russia
Open Access

Abstract

An important step in research planning is the choice of methodology. This is especially important for territories which are difficult to access such as in the Arctic, where successive repetitions of field works require significant resources. The methodology utilizing the local fauna has been used over the past twenty years. It provides comparable data on the structure of fauna and species richness for different territories. The purpose of the present study was to assess the “local fauna method” with respect to fauna studies of ground beetles in the Arctic forest-tundra zone. The research was conducted from June 18 2017 to August 30 2017 within the Polar Urals (10 km from the Harp settlement in the Yamalo-Nenets Autonomous Okrug, which is a state of Russia). Carabids were sampled by using pitfall traps on 20 sites. This article will also include the results of our previous research concerning the structure of some local faunas from the forest-tundra zone of Nenets Autonomous Okrug (settlements Nes’, Oma, Khorey-Ver). The results of this study demonstrate the following: 1) the local fauna of the Polar Urals has 85 species of ground beetles from 25 genera, which is 77% of species lists of carabids for a 70-year period of research within the Polar Urals; 2) the local fauna of the Polar Urals has 29% similarity of list species with local faunas from the European part of the Arctic, with similar compositions of zoogeographical groups and life forms; 3) in one research period there was 90% detection of carabids species in the forest-tundra local fauna using the sampling method of pitfall traps within a period of 40 days, conducted at 15 sites, with the predominance of southern types of plant communities (meadows, forests).

Keywords

ground beetles, Coleoptera (Carabidae), local fauna, forest-tundra, Arctic

Introduction

Studies of ground beetle faunas (Coleoptera, Carabidae) in the Polar Urals have been systematically conducted over the last 60 years (Stebaev 1959, Olshvang 1980, Korobeinikov 1984, Zinovyev and Olshvang 2003, Uzhakina and Dolgin 2007). At the same time, ground beetle species from different geographic locations in Polar Urals range from 26 to 110 species (Stebaev 1959; Olshvang, 1980, Lomakin and Zinoviev 1997, Kozyrev et al. 2018). This is due to the different approaches to material collecting, the number of study sites and the research periods. Collected data characterize the species richness of ground beetles of the Polar Urals area in general but this is not applicable to comparative analysis due to its methodological heterogeneity.

In recent years the local fauna concept has been used in faunistic research (Penev 1996, Bolotov and Danilevich 2005, Filippov 2008, Makarov and Matalin 2009, Kolosova and Potapov 2010, Tatarinov and Kulakova 2010). The term “local flora” was proposed by Tolmachev (Tolmachev 1931) and this was later borrowed by Chernov (Chernov 1975) in zoological studies in the form of “local fauna”. Penev (Penev 1996) clarified this concept for a zoological approach as in a territory of 100 km2 in a certain biome. Using the local fauna approach it is possible to obtain comparable data on species richness and fauna structures of soil invertebrates with the determination of the maximum species compositions for a certain territory. This method was used for studying ground beetle local fauna of semi-deserts in the Volga Region (Makarov and Matalin 2009) and the European sector of the Russian Arctic (Filippov and Shuvalov 2005, Filippov 2008, Markov 2011, Zubriy and Filippov 2015). The main aim of the present research was to study the ground beetle local fauna of the Polar Urals forest-tundra zone and to estimate the effectiveness of species richness determination.

Materials and methods

Local fauna field works of ground beetles from the Polar Urals forest-tundra zone were conducted at a distance of 10 km from Kharp settlement (Yamalo-Nenets Autonomous Okrug, Russia) from 18 June 2017 to 30 August 2017 (see Fig. 1). Within 20 model sites, the main types of plant communities were selected: gramineous-herb meadows, upland sphagnous, sedge and dwarf birch bogs, moss-lichen and shrub tundras, and also larch forests, spruce forests and alder forests (see Table 1).

Fig. 1.

Study sites (see Table 1 for numbers of sampling sites)

Ground beetles were collected using pitfall traps (Heydemann, 1956) – 500 ml plastic cups with a trap hole diameter of 93 mm. In each site 10–25 traps were installed in parallel lines with a distance of 10 m between traps and lines. For fixing insects 4% formaldehyde solution was used. The material was sampled from the traps once every ten days. Carabids near water bodies were collected using the exhaustion trapping method. As result, 375 traps were installed for a total of 24,852 trap-days and 6,409 specimens of imago were sampled (see Table 1).

The list of Russian ground beetles (Makarov et al. 2016) and the Catalogue of Palaearctic ground beetles (Löbl and Löbl 2017) were used for taxonomic classification.

Information about ground beetles of Fennoscandia was used for fauna the zoogeographical analysis (Lindroth 1992). Carabid habitat preferences in the tundra zone were determined by using the authors’ own data and also by using published data (Lindroth 1992, Dudko et al. 2010, Khobrakova et al. 2014). Categorization of the species in respect of their life forms was made according to the classification of Sharova (1981).

For determining the full species richness in the forest-tundra zone of the Polar Urals, field work periods and the number of sites in different habitat types were estimated by using the authors’ own previous data on local faunas in the European Arctic sector (see Table 2).

Carabid species similarity among local faunas of the North Europe and Polar Urals was assessed by using the Jaccard index in a dendrogram of cluster analysis via the simple average link algorithm in the BioDiversityPro software (McAleece et al. 1997). Species accumulation curves were estimated as a function of research periods, number of sample sites and habitat types of local fauna of the forest-tundra zone of the northern part of Europe and the Polar Urals. The species richness cumulate rates for local fauna and different communities were estimated by rarefaction procedure in the EstimateS software version 9.1.0 (Colwell 2013). The Akaike Information Criterion (AIC) was applied to a selection of models and describes the forecast trend lines for new species appearance with an increase in samples (Hammer 2015).

Sampling sites in the Polar Urals

No. Site Pitfall traps (number) Pitfall trap-days Imago specimens (number)
1 Sphagnum-ledum bog 20 1,350 196
2 Meadow grass 20 1,469 303
3 Green-mossy bog with dwarf birch 20 1,364 372
4 Forest glade 10 699 145
5 Meadow grass 20 1,418 364
6 Meadow grass 10 679 321
7 Birch-fir forest 20 1,370 949
8 Moss-shrub tundra 20 1,270 443
9 Rocky moss-lichen tundra with larch 20 1,400 27
10 Fir-wood ledum-green-mossy 20 1,420 140
11 Rocky moss-shrub tundra 20 1,420 153
12 Thickets of alders with grass 20 1,341 318
13 Meadow grass 20 1,380 608
14 Larch forest with moss and shrubs 20 1,331 139
15 Meadow grass 15 1,035 891
16 Moss-lichen-shrub tundra 20 1,380 198
17 Sphagnum-ledum bog with dwarf birch 20 1,311 208
18 Moss-ledum bog 20 1,216 47
19 Rocky moss-shrub tundra 20 1,216 420
20 Grass bog 20 698 38
by exhauster (waterside sites) 129
Total: 375 24,852 6,409

Results

For ground beetle local fauna of Polar Urals forest-tundra zone, 85 species from 25 genera were established (see Table 3). The taxonomic structure demonstrated the highest participation of genera representatives: Pterostichus (15 species), Bembidion (12 species), Amara (11 species) and Agonum (8 species).

The carabid species belonged to 4 zoogeographical categories. The greatest presence was exemplified by Palearctic species - 56.5%; the Northern Holarctic faunal type accounted for 31.8%; the Euro-Siberian faunal type accounted for 8.2%; and the European faunal type accounted for 3.5% (see Table 3).

Representatives of the class Zoophagous predominated in imago life forms of ground beetle species – 78.8% (see Table 4).

The life forms of the ground beetle species in this study belonged to 12 groups. Most of the zoophages belonged to the subclass Stratobios: surface and litter-dwelling stratobionts (30.5%), litter-dwelling stratobionts (17.6%). Among the mixophytophagous species, the haploid geohortobionts predominated (12.9%).

On a dendrogram of species similarity, the Polar Urals carabid cluster has relatively low association (29.3%) with North Europe local faunas of the forest-tundra zone (see Fig. 2). The local faunas of Nenets Autonomous Okrug ground beetles are 40–50% similar in terms of species composition.

Fig. 2.

Dendrogram of species similarity Carabidae local faunas in the Nenets Autonomous Okrug: I – Naryan-Mar, II – Oma (Markov, 2011), III – Khorey-Ver, IV – Nes (Filippov, 2008), V – Polar Urals

Local faunas of the Nenets Autonomous Area

Settlement Research period (ten-day periods) Number of sites, n Number of species: pitfall traps/total, s Number of carabids individuals, n
Nes 27 Jun – 23 Aug 2002 (5) 8 74/74 7,728
Oma 3 Jun – 1 Sep 2008 (7) 12 63/80 6,361
Khorey-Ver 12 Jun – 22 Aug 2013 (6) 6 50/50 1388

List of Carabidae beetles, established for Polar Urals local fauna

No. Species 1Zoogeographical groups 2Life forms of imago Number of specimens
1 Trachypachus zetterstedtii (Gyllenhal, 1827)* PAL 1.1.1 1
2 Pelophila borealis (Paykull, 1790) HOL 1.2.1 5
3 Leistus terminatus (Hellwig in Panzer, 1793) PAL 1.2.2 1
4 Nebria rufescens (Ström, 1768) HOL 1.2.1 6
5 N. nivalis (Paykull, 1790) PAL 1.2.1 1
6 Notiophilus aquaticus (Linnaeus, 1758) HOL 1.2.1 99
7 N. biguttatus (Fabricius, 1779) E-SI 1.2.1 10
8 N. reitteri (Späth, 1899) PAL 1.2.1 48
9 Carabus nitens (Linnaeus, 1758) EUR 1.1.1 22
10 C. truncaticollis (Eschscholtz, 1833) HOL 1.1.1 57
11 C. henningi (Fischer von Waldheim, 1817) PAL 1.1.1 32
12 C. odoratus (Motschulsky, 1844) PAL 1.1.1 81
13 Cychrus caraboides (Linnaeus, 1758) EUR 1.1.1 7
14 Diacheila arctica (Gyllenhal, 1810) HOL 1.1.1 3
15 Diacheila polita (Faldermann, 1835) PAL 1.1.1 169
16 Blethisa multipunctata (Linnaeus, 1758) HOL 1.2.1 17
17 B. catenaria (Brown, 1944) HOL 1.2.1 1
18 Elaphrus angusticollis (R.F. Sahlberg, 1844) E-SI 1.2.1 21
19 E. lapponicus (Gyllenhal, 1810) PAL 1.2.1 33
20 Loricera pilicornis (Fabricius, 1775) HOL 1.2.1 29
21 Clivina fossor (Linnaeus, 1758) HOL 1.3.1 20
22 Dyschiriodes globosus (Herbst, 1783) PAL 1.3.1 6
23 D. melancholicus (Putzeys 1866) PAL 1.3.1 1
24 D. nigricornis (Motschulsky, 1844) PAL 1.3.1 2
25 D. neresheimeri (H. Wagner, 1915) E-SI 1.3.1 1
26 Miscodera arctica (Paykull, 1798) HOL 1.2.3 18
27 Bembidion properans (Stephens, 1829) PAL 1.2.1 50
28 B. bipunctatum (Linnaeus, 1761) PAL 1.2.1 9
29 B. transparens (Gebler, 1829) PAL 1.2.1 2
30 B. quadrimaculatum (Linnaeus, 1761) HOL 1.2.1 86
31 B. fellmanni (Mannerheim, 1823) PAL 1.2.1 15
32 B. andreae (Fabricius, 1787) PAL 1.2.1 1
33 B. saxatile Gyllenhal, 1827) PAL 1.2.1 27
34 B. prasinum (Duftschmid, 1812) PAL 1.2.1 53
35 B. dauricum (Motschulsky, 1844) HOL 1.2.1 18
36 B. crenulatum (R.F.Sahlberg, 1844) PAL 1.2.1 3
37 B. obscurellum (Motschulsky, 1844) HOL 1.2.1 1
38 B. semipunctatum (Donovan, 1806) PAL 1.2.1 1
39 Patrobus assimilis (Chaudoir, 1844) E-SI 1.2.2 19
40 P. septentrionis (Dejean, 1828) HOL 1.2.2 89
41 Pterostichus nigrita (Paykull, 1790) PAL 1.2.3 1
42 Pt. strenuus (Panzer, 1797) PAL 1.2.2 2
43 Pt. brevicornis (Kirby, 1837) HOL 1.2.2 814
44 Pt. adstrictus (Eschscholtz, 1823) HOL 1.2.3 5
45 Pt. kaninensis (Poppius, 1906) EUR 1.2.2 4
46 Pt. macrothorax (Poppius, 1906) PAL 1.2.2 40
47 Pt. middendorffi (J.Sahlberg, 1875) PAL 1.2.2 223
48 Pt. pinguedineus (Eschscholtz, 1823 HOL 1.2.2 22
49 Pt. dilutipes (Motschulsky, 1844) PAL 1.2.3 15
50 Pt. eximius (A. Morawitz, 1862) PAL 1.2.3 22
51 Pt. kokeili (Poppius, 1907) E-SI 1.2.3 213
52 Pt. montanus (Motschulsky, 1844) PAL 1.2.3 1199
53 Pt. haematopus (Dejean, 1831) HOL 1.2.2 11
54 Pt. rubripes (Motschulsky, 1860) HOL 1.2.2 185
55 Pt. vermiculosus (Menetries, 1851) HOL 1.2.2 98
56 Calathus melanocephalus (Linnaeus, 1758) PAL 1.2.2 306
57 C. micropterus (Duftschmid, 1812) PAL 1.2.2 11
58 Agonum dolens (C.R. Sahlberg, 1827) PAL 1.2.1 2
59 A. quinquepunctatum (Motschulsky, 1844) PAL 1.2.1 305
60 A. ericeti (Panzer, 1809) PAL 1.2.1 4
61 A. viduum (Panzer, 1797) PAL 1.2.1 2
62 A. consimile (Gyllenhal, 1810) PAL 1.2.1 7
63 A. fuliginosum (Panzer, 1809) E-SI 1.2.1 24
64 A. gracile (Sturm, 1824) PAL 1.2.1 17
65 A. exaratum (Mannerheim, 1853) HOL 1.2.2 5
66 Amara plebeja (Gyllenhal, 1810) PAL 2.1 7
67 A. bifrons (Gyllenhal, 1810) PAL 2.3 110
68 A. brunnea (Gyllenhal, 1810) HOL 2.2 11
69 A. erratica (Duftschmid, 1812) HOL 2.3 9
70 A. ingenua (Duftschmid, 1812) PAL 2.3 14
71 A. lunicollis (Schiødte, 1837) PAL 2.2 21
72 A. praetermissa (C.R. Sahlberg, 1827) PAL 2.2 499
73 A. quenseli (Schönherr, 1806) PAL 2.3 255
74 A. municipalis (Duftschmid, 1812) PAL 2.3 4
75 A. interstitialis (Dejean, 1828) HOL 2.2 3
76 A. reitteri (Tschitscherine, 1894 PAL 2.3 1
77 Curtonotus torridus (Panzer, 1797) PAL 2.3 1
78 C. alpinus (Paykull, 1790) PAL 2.3 691
79 C. hyperboreus (Dejean, 1831) HOL 2.3 96
80 Dicheirotrichus cognatus (Gyllenhal, 1827) HOL 2.2 5
81 D. mannerheimi (R.F.Sahlberg, 1844) PAL 2.2 1
82 Harpalus torridoides (Reitter, 1900) E-SI 2.3 5
83 H. nigritarsus (C.R.Sahlberg, 1827) HOL 2.3 48
84 Cymindis vaporariorum (Linnaeus, 1758 PAL 1.2.4 23
85 Paradromius ruficollis (Motschulsky, 1844) PAL 1.2.5 3
Total: 6409

Carabid sampling periods in four local faunas of the forest-tundra zone of the Polar Urals and North Europe were conducted at different times of the vegetation seasons from June to September (see Table 2). During those periods, more than 90% of the ground beetle species were collected during the first two to four ten-day periods of the field studies (see Fig. 3A). The appearance of new species in samples collected over the study periods is specific for each local fauna, due to phenological variance and the number of sites for each territory.

According to the accumulative curve which is beginning to reach an asymptote, sampling was successfully conducted for only one local fauna – the forest-tundra zone of the Polar Urals (Fig. 3B). Species richness was incomplete for the sampled ground beetle local faunas of Nes, Oma and Khorey-Ver settlements, and the accumulative curves show no signs of leveling off (Fig. 3B).

Fig. 3.

Species accumulation curves of Carabidae: A – by period of research works; B – by number of sites. Dotted lines – 95% confidence intervals, solid lines – trend lines

Carabidae beetle life forms of the Polar Urals local fauna

Life form Number of species, s Percentage of species, %
Zoophagous: 67 78.8
walking epigeobionts 8 9.4
running epigeobionts 4 4.7
surface and litter-dwelling stratobionts 26 30.6
litter-dwelling stratobionts 15 17.6
litter and soil-dwelling stratobionts 6 7.05
crevice-dwelling stratobionts 1 1.2
litter and bark-dwelling stratobionts 1 1.2
digging geobionts 5 5.9
running and digging geobionts 1 1.2
Mixophytophagous: 18 21.2
stratohortobionts 1 1.2
crevice-dwelling stratobionts 6 7.05
geohortobionts 11 12.9

Structures of forest-tundra ground beetle community assemblages depend on local fauna. For instance, in the forest-tundra Polar Urals 12 species of ground beetles were established for all types of habitat and 56 species were collected from two or three habitat types. Rare species, which were sampled in only one habitat type, belong to 17 species of carabids. More than half of the rare species (9 species) were collected from meadows; 5 species in tundra and 3 species in forest habitats. Thus, under the sampling design used in the present study approximately one third of the sites should have selected specimens from meadow communities. Carabid species richness in meadow has significant differences with respect to tundra, forest and bog habitats according to rarefaction analysis whereby the confidence intervals are overlapping (see Fig. 4A).

In the forest-tundra zone of the Nes, Oma and Khorey-Ver settlements in the Polar Urals the tendency was to provide the smallest contribution of zonal community types (bogs, tundra) in the ground beetle species richness that was determinated (see Fig. 4). For local faunas of the Nenets Autonomous Okrug no significant differences were estimated in the collected number of carabid species between forest and meadow communities (see Fig. 4). Only for Nes was the maximum number of carabids species collected in forest habitats (see Fig. 4B).

Fig. 4.

Species abundance of Carabidae communities: A – Polar Urals, B – Nes, C – Oma, D – Khorey-Ver. Dotted lines – 95% confidence intervals

Discussion

The aforementioned “local fauna method” was used, and to our knowledge this was the first time, in the forest-tundra of the Polar Urals and within one vegetation season this allowed the collecting of 77% of the regional carabid fauna. Ground beetle species richness in the fauna of the present study is comparable with local fauna (81 species) of the Kanin and Timan highlands in the forest-tundra zone of the European part of the Russian Arctic (Markov, 2011).

Seventeen (17) sampling sites were studied with respect to the collecting of 90% (77 species) carabid species richness in one local fauna of the forest-tundra of the Polar Urals. According to the estimated trend (logarithmic function) for the forest-tundra of Nenets Autonomous Okrug 90% carabid species richness was determined for the local faunas of Oma settlement (66 species) – with 13 sites, Nes (86 species) – with 14 sites and Khorey-Ver (57 species) – with 13 sites. Thus, for identifying the complete species richness of the local fauna in the forest-tundra zone, at least 15 sampling sites were used.

The zoogeographical group compositions for the local fauna of the Polar Urals with the Kanin-Timan tundra, Nes and Oma settlements is similar to the local faunas of the European forest-tundra (Filippov 2008, Markov 2011). A high proportion of Holarctic species in the carabid local fauna of the Polar Urals at 31.8% differs to that of Kanin (19.8%) and Oma (22.9%) in the forest-tundra zone of the European North (Filippov 2008, Markov 2011).

Life form groups of imago ground beetles life forms are typical for the subarctic forest-tundra zone on the border area of Europe and Asia. For instance, in the forest-tundra zone of Nenets Autonomous Okrug (Nes settlement) the subclass Stratobios is dominates, as it does in the Polar Urals: surface and litter-dwelling stratobionts (31.0%) and litter-dwelling stratobionts (14.7%), but the proportion of haploid geohortobionts is slightly higher in Nes with 18.9% (Filippov 2008).

In total, 85 species were collected during one vegetation period. According to the published data (Kozyrev et al. 2018), 84 species of ground beetles were recorded for the Salekhard and Labytnangi regions and 110 species for the forest-tundra zone of the Polar Urals within a study period of the last 60 years. In addition, in this study the species Trachypachus zetterstedtii was collected for the first time in the forest-tundra of the Polar Urals. We obtained data from the forest-tundra zone under study and the results of the “local fauna method” in sampling material during the one vegetation season is no less effective than using data collected in a multi-year research project using irregular sampling methods (Makarov and Matalin 2009).

We propose that when planning to study carabid local fauna of the forest-tundra zone, at least 15 model sites should be used for at least a 40-day period. Further, two-thirds of the studied sites should be of the intrazonal community type (forest and meadow), and the rest should be of the interfluve type (tundra and bog).

Acknowledgements

This study was supported by the Federal Agency for Scientific Organizations (no. 0409-2016-0022), program of the Russian Academy of Sciences Presidium (no. 55 «Arctic»), Ministry of Education and Science of Russian Federation (no. 6.2343.2017/4.6). We also would like to acknowledge Professor, Dr.Sc V.D. Bogdanov, and PhD V.G. Shtro (Institute of Plant and Animal Ecology RAS) for the invaluable support and help in field research on the basis of the Arctic Research Station (Labytnangi). The authors would like to thank Professor, Dr.Sc K.V. Makarov (Moscow Pedagogical State University) for determining the taxonomic status of ground beetle species.

References

  • Bolotov IN, Danilevich EA (2005) Local faunas of diurnal lepidopterans (Lepidoptera, Diurna) in the European North of Russia: the Solovetsky Islands. Vestnik Pomorskogo universiteta. Seriya: estestvennye i tochnye nauki 2(8): 79–93.
  • Chernov YuI (1975) Prirodnaya zonal'nost' i zhivotnyj mir sushi. Mysl', 222 pp.
  • Colwell RK (2013) EstimateS: Statistical estimation of species richness and shared species from samples. http://purl.oclc.org/estimates [accessed 21 June 2018]
  • Dudko RYu, Matalin AV, Fedorenko DN (2010) The Ground Beetle Fauna (Coleoptera, Carabidae) of Southeastern Altai. Entomological Review 90(8): 968–988.
  • Filippov BYu (2008) Puti adaptatsii i ehkologicheskie zakonomernosti osvoeniya zhuzhelitsami (Coleoptera, Carabidae) severa Russkoj ravniny: Аvtoref. dis. … dokt. biolog. nauk. M., 41 pp.
  • Filippov BYu (2008) Puti adaptacii i ehkologicheskie zakonomernosti osvoeniya zhuzhelicami (Coleoptera, Carabidae) severa Russkoj ravniny : dis. ... dokt. biolog. nauk. Moscow, 367 pp.
  • Hammer Ø (2015) PAST: Paleontological Statistics Version 3.06: Reference manual. Natural History Museum University of Oslo, Oslo, 225 pp.
  • Heydemann B (1956) Über die Bedeutung der “Formalinfallen” für die zoologische Landesforschung. Faun. Mitt. Norddeutschl 6: 19–24.
  • Khobrakova LTs, Shilenkov VG, Dudko RYu (2014) Zhuki – zhuzhelitsy (Coleoptera, Carabidae) Buryatii. BNTSSORАN, Ulan – Udeh, 300 pp.
  • Korobeinikov YuI (1984) Zhuzhelicy Priobskogo Severa. Problemy pochvennoj zoologii. Ashkhabad, 149–150.
  • Kolosova YuS, Potapov GS (2010) Local faunas of bumblebees (Hymenoptera: Apidae, Bombini) of the European North of Russia: the Pechora river delta and the Kolguev island. Vestnik Pomorskogo universiteta. Seriya: estestvennye i tochnye nauki 3: 69–75.
  • Lomakin DE, Zinoviev EV (1997) Fauna zhuzhelic (Coleoptera, Carabidae) poluostrova Yamal. Materialy po istorii i sovremennomu sostoyaniyu fauny severa Zapadnoj Sibiri. Chelyabinsk, 3–15.
  • McAleece N, Gage JD, Lambshead J, Patterson GLJ (1997) Biodiversity Professional. The Natural History Museum and The Scottish Association for Marine Science. http://www.sams.ac.uk
  • Makarov KV, Kryzhanovskij OL, Belousov IA, Kabak II, Kataev BM, Shilenkov VG, Matalin AV, Fedorenko DN, Komarov EV (2016) A Taxonomic List of the Ground-Beetles (Carabidae) of Russia. http://www.zin.ru/Animalia/Coleoptera/rus/car_rus.htm [Last updated 8 November 2016]
  • Makarov KV, Matalin AV (2009) Ground-beetle communities in the Lake Elton region, southern Russia: a case study of a local fauna (Coleoptera, Carabidae). In: Babenko AB, Matveeva NV, Makarova OL, Golovach SI (Eds) Species and Communities in Extreme Environments. Festschrift towards the 75th Anniversary and a Laudatio in Honour of Academician Yuri Ivanovich Chernov. Pensoft Publishers & KMK Scientific Press Ltd., Sofia – Moscow, 357–384.
  • Markov SA (2011) Species composition of ground beetles (Coleoptera, Carabidae) of Kanin-Timan forest-tundra local fauna. Vestnik Severnogo federal'nogo universiteta. Estestvennye nauki 3: 55–61.
  • Lindroth CH (1992) Ground Beetles (Carabidae) of Fennoscandia. A zoogeographic Study. Part I. Speciphic Knowledge Regarding the Species. Washington: Amerind Publishing Co Pvt. Ltd., 630 pp.
  • Löbl I, Löbl D (2017) Catalogue of Palaearctic Coleoptera Vol. 1 Revised and Updated Edition): Archostemata – Myxophaga – Adephaga. Brill, Leiden – Boston, 1443 pp.
  • Olshvang VN (1992) Struktura i dinamika naseleniya nasekomykh Yuzhnogo Yamala. Nauka, Ekaterinburg, 104 pp.
  • Penev L (1996) Large-Scale Variation in Carabid Assemblages, With Special Reference to the Local Fauna Concept. Ann. Zool. Fenn. 33: 49–63.
  • Sharova IKh (1981) Zhiznennye formy zhuzhelits. Nauka, 360 pp.
  • Stebaev IV (1959) Pochvennye bespozvonochnye Salekhardskih tundr i izmenenie ih gruppirovok pod vliyaniem zemledeliya. Zoologicheskij zhurnal 10: 1559–1572.
  • Tatarinov AG, Kulakova OI (2010) Lokal faunas of butterflies (Lepidoptera: Papilionoidea, Hesperioidea) in the European north of Russia: Padimey lakes, head reaches of the Bolshaya Rogovaya river. Vestnik Pomorskogo universiteta. Seriya: estestvennye i tochnye nauki 1: 72–80.
  • Tolmachev АI (1931) K metodike sravnitel'no-floristicheskikh issledovanij. Ponyatie o flore v sravnitel'noj floristike. Zhurnal RBO 16(1): 111–124.
  • Uzhakina OА, Dolgin MM (2007) Obzor fauny zhuzhelits (Coleoptera, Carabidae) tundrovykh ehkosistem evropejskogo Severo-Vostoka Rossii. Bespozvonochnye evropejskogo Severo-Vostoka Rossii. Syktyvkar, 267–286.
  • Zinoviev EV, Olshvang VN (2003) Zhuki severa Zapadno-Sibirskoj ravniny, Pripolyarnogo i Polyarnogo Urala. Biologicheskie resursy Polyarnogo Urala. Nauch. vestnik 3(2): 37–60
  • Zubriy NA, Filippov BYu (2015) Carabid beetles local fauna (Coleoptera, Carabidae) of the typical tundra of the Yugor Peninsula. Vestnik Severnogo federal'nogo universiteta. Seriya: Estestvennye nauki 2: 46–55.