Analysis of the ways of formation of the entomofaunistic complexes in the Northwest Caucasus based on the material on coleopterous insects (Coleoptera)

By the example of 14 coleopterous insect families, namely Gyrinidae, Haliplidae, Noteridae, Dytiscidae, Carabidae, Hydrophilidae, Staphylinidae, Lucanidae, Trogidae, Scarabaeidae, Elateridae, Alleculidae, Tenebrionidae, and Chrysomelidae, regularities and the basic sources of formation of the most typical landscapecoenotic complexes in the Northwest Caucasus are discussed. The total number of the species included in the material analyzed amounts to about 2000. The maximal species diversity (854 species) is registered for the deciduous forests and dry open woodlands. In total, 11 types of the chorological complexes and distributional ranges (chorotypes) of the regional beetle fauna are distinguished. Significant concordance of the chorologic patterns in some groups of beetles is established. In this respect, Carabidae and Tenebrionoidea seem to be the closest on the one hand, and also Elateridae and Scarabaeoidea, on the other. The zonal fauna of the Northwest Caucasus is formed basically by species possessing Boreal ranges, while the azonal fauna, predominantly by species with the Ancient Mediterranean ranges. Allocation of coleopterous insects within regional zoochorones is investigated as well. It is established, that in different zoochorones of the upland part of the region the majority of coenofaunas show significant similarity of the arealogical pattern, even though being composed frequently by different taxa. The fauna of agrarian landscapes of the Northwest Caucasus is also examined. It includes 382 beetle species. The overwhelming majority of them belongs to the ground beetles (229 species), leaf beetles (78), and Scarabaeoidea (30). This fauna reveals the maximal similarity with the coenofaunas of the lowland steppe and meadows. It is established that the major role in formation of the agrocenoses beetle fauna belongs to the natural plain and upland steppe and treeless fields, foothill broad-leaf forests and open woodlands, and also to the floodplain and lowland forests. The bulk of the species occurring in the agrocenoses possesses wide ranges, predominantly of the Boreal type. Some peculiarities of the regional endemism are discussed as well.     

Recent records of steppe species in Belarus,first indications of a steppe species invasion?

BELARUS IS SITUATED AT A CROSSROAD OF NATURAL BORDERS OF SPECIES DISTRIBUTIONS: the NE part is situated in a taiga zone, whereas the other part of terrain is in the European forest zone. The distance of Belarus to the steppe zone is about 330 kilometers. This geographical position and the extensive knowledge of its fauna can be used to monitor changes in the distribution of different species. An intensive study of open habitat ground beetles was carried out from 1975-2008 in Belarus, using pitfall traps, quadrate-sampling methods, hand collecting, netting and light traps. In total, more than 130 000 specimens of ground beetles belonging to 169 species were collected from 62 fields and 11 meadows of different types. 217 specimens of Calosoma investigator (Illiger 1798), 2 specimens of Calosoma denticolle (Gebler 1833), and one specimen of Harpalus subcylindricus (Dejean, 1829), Harpalus honestus (Duftschmid 1812) and Zabrus tenebrioides (Goeze 1777) were present in this material. All specimens were macropterous and exclusively caught at fields and waste grounds on sandy soil. Nowadays Belarus is the northernmost location for these species in Eastern Europe. Steppe species most probably migrated to SE Belarus from NE Ukraine, using Dnieper and its river valleys. The shift in the geographic distribution of steppe species during the last thirty years in Belarus have been attributed to a higher frequency of warmer and wetter summers in the last few decades.     

What do we know about winter active ground beetles (Coleoptera,Carabidae) in Central and Northern Europe?

This paper summarizes the current knowledge on winter active Carabidae in Central and Northern Europe. In total 73 winter active species are listed, based on literature and own observations. Ground beetles are among the three most numerous Coleoptera families active during the autumn to spring period. The winter community of Carabidae is composed both of larvae (mainly autumn breeding species) and adults, as well as of epigeic species and those inhabiting tree trunks. Supranivean fauna is characterized by lower species diversity than the subnivean fauna. The activity of ground beetles decreases in late autumn, is lowest during mid-winter and increases in early spring. Carabidae are noted as an important food source in the diet of insectivorous mammals. They are also predators, hunting small winter active invertebrates.     

小兴安岭阔叶红松林地表甲虫Beta多样性

Beta多样性用来衡量集群内物种组成的变异性,可以被分解为空间物种转换和物种集群镶嵌两个组分,是揭示群落构建机制的重要基础。目前开展了较多的地上生态系统beta多样性研究,然而地下生态系统beta多样性进展缓慢。以小兴安岭凉水和丰林自然保护区为研究地区,于2015年8、10月采用陷阱法对阔叶红松林进行调查,揭示地表甲虫(步甲科、隐翅虫科、葬甲科)的beta多样性。结果表明:(1)凉水共发现39种、856只地表甲虫,丰林共发现43种、1182只地表甲虫。8月凉水明显具有较高的全部甲虫(三个科的总和)物种多样性和丰富度,10月正好相反。(2)凉水和丰林之间地表甲虫beta多样性的差异仅发现于8月的步甲科和葬甲科之间。(3)凉水和丰林地表甲虫的beta多样性主要由空间物种转换组成,物种集群镶嵌对beta多样性的贡献很小,说明地表甲虫物种组成变异主要由本地物种之间较高的转换引起。研究表明小兴安岭阔叶红松林地表甲虫的beta多样性主要由空间物种转换组成,在揭示群落构建机制过程中,其内部物种交换和环境调控不容忽视。     

Fauna and population structure of springtails (Hexapoda,Collembola) in zonal forest-steppe biotopes of the Middle Volga River basin

A total of 92 springtail species were revealed in forest-steppe areas of the Privolzhskaya Lesostep nature reserve. The considerable part of the species (25–40%) are inhabitants of steppe and open sites with the predominance of soil forms. The specific features of the chernozem soil fauna are determined by the rare species occupying deep soil layers. In different forest-steppe biotopes, the springtail communities are similar in the species composition but differ in the structure. Steppe and forest types of communities were distinguished. When the steppe is forested, the most drastic changes in the collembolan population are revealed in the transitional shrub zone. The marginal community is characterized by the most even structure and a variable set of dominants.     

The influence of habitat type on the population dynamics of ground beetles (Coleoptera: Carabidae) in marshland

This study presents qualitative and quantitative research on communities of ground beetles (Carabidae) occurring in 2001 and 2003 in three types of habitats located near Bydgoszcz city, Poland (UTM CD19), on the Vistula river floodplain: elm and ash riparian forest (RF), poplar monoculture (P), and unintensively-farmed meadows (M). During both studied years 57 species of ground beetles were found in riparian forests (RF), 56 species in plot P and 47 in plot M. In 2001 from all sample plots 3845 beetles were collected; in 2003- 9589. In both years most specimens came from plot RF (3123 in 2001 and 6224 in 2003). From plot P 2656 specimens of ground beetles were collected and from plot M 1430 specimens. Statistically significant differences were found in the abundance of Carabidae, among sampling areas and the years of study.     

Species composition and landscape-zonal distribution of horseflies (Diptera,Tabanidae) in the territory of Belgorod Province

A list of 30 species of the family Tabanidae recorded in Belgorod Province is provided, and data on the landscape-zonal distribution, typical biotopes, and dates of flight in the region are given for each species. The greatest number of horsefly species (24) was recorded in the southern forest-steppe subzone, and smaller numbers, in the central forest-steppe subzone (18) and the steppe zone (16). Half of the forest and forest-steppe species extend into the steppe zone while some southern species penetrate into the forest-steppe zone. The pattern of horsefly distribution is determined by the biotopic conditions. The boundaries of distribution of individual species do not distinctly correspond to those of the landscape subzones.     

Diversity of phytophagous beetles (Coleoptera: Chrysomeloidea,Curculionoidea) in steppe communities in the forest-steppe of the High Trans-Volga region

A total of 551 species from 10 families of phytophagous beetles (superfamilies Chrysomeloidea and Curculionoidea) have been recorded in the steppe communities of the High Trans-Volga forest-steppe zone, 62% of which belong to the steppe landscape-geographic complex. The vast majority of the species belong to three families: Curculionidae (254 species; 46%), Chrysomelidae (186 species; 34%), and Apionidae (51 species; 9%). The regional steppe fauna has a complex arealogical structure combining the Western Scythian and Sarmatian elements, as well as relict species of the Angarian origin and some Eastern European ones. The highest diversity of phytophagous beetles is observed in the zonal meadow steppe communities (299 species; 54%). The most specific are the petrophilous steppe complexes which also include the greatest number of relict forms. Xerophilous Kazakhstan- Turanian species are mostly concentrated in the saline steppe communities of the southernmost forest-steppe zone. The species diversity is also high (249 species) in the open ruderal communities dominated by species characteristic of the pioneer stages of the steppe successions. Some steppe reservations include over 50% of the entire regional fauna of phytophagous beetles. The species diversity in the steppe localities decreases northward due to a gradual decrease in the number and proportion of the Scythian species: from 55% in the middle of the foreststeppe zone to 36% near its northern boundary. The species diversity of the slope communities in the forest-steppe zone is at least not lower than that in the similar areas of the steppe zone. The ratio of the number of Curculionoidea species to that of plant species is very high, reaching about 3: 4 in the model steppe localities of the High Trans-Volga forest-steppes, as compared to 1: 2 in the zonal steppe communities (according to the published data). The data considered confirm the significance of the forest-steppe zone as an important biodiversity center in the East European Plain and as a reservation of relicts of different origins.     

Unexpected visitors: flightless beetles in window traps

Ancient woodlands are highly important for biodiversity. Flightless beetles and window traps may first appear to be incompatible as a study group and a study method. Little is known about the actual capability of window traps in monitoring of entire forest arthropod communities. We try to determine the possibility of using window traps (designed as tree traps) as an additional method to trap flightless beetles of the genera Carabus (Carabidae), Acalles and Trachodes (Curculionidae), as vulnerable representatives of wildlife forest fauna with high conservation value. From the environmental point of view, the occurrence of the studied flightless beetles was significantly more influenced by altitude than tree species as the studied environmental variables. Several beetles also showed positive significant species to species relationships during their observed phenological activity. Our results showed that the studied flightless beetles may reappear in secondary forests if these reach their optimal structure. On the other hand, they may be sensitive to large scale disturbances and not only to those caused by commercial forestry. The results also showed that window traps designed as tree traps could be used to successfully trap assemblages of forest fauna including flightless species.     

The ground beetle fauna (Coleoptera,Carabidae) of Southeastern Altai

Long-term studies of the ground beetle fauna of Southeastern Altai (SEA) revealed 33 genera and 185 species; 3 and 15 species are reported for the first time from Russia and SEA, respectively. The following genera are the most diverse: Bembidion (47 species), Amara and Harpalus (21 each), Pterostichus (14), and Nebria (13). The subarid (35%) and boreal (32%) species prevail in the arealogical spectrum, while the mountain endemics comprise 13% of the fauna. The carabid fauna of SEA is heterogeneous in composition and differs significantly from that of the Western and Central Altai. The boreal mountain component mostly comprises tundra species with circum-boreal or circum-arctic ranges, while the subarid component (typical Mongolian together with Ancient Mediterranean species) forms more than one-half of the species diversity in the mountain basins. The species diversity increases from the nival mountain belt (15 species, predominantly Altai-Sayan endemics) to moss-lichen tundras (40, mostly boreal, species). The variety of habitats and constant hydrothermal regimes in the intrazonal valley communities determine high taxonomic diversity of Carabidae, including both tundra or meadow-steppe species typical of the region and some forest ones. Under the condition of moisture deficiency, many steppe species also concentrate in river floodplains.     

The influence of matrix and edges on species richness patterns of ground beetles (Coleoptera: Carabidae) in habitat islands

Aim The aim of this study was to analyse whether, and how, the inclusion of habitat specialists and edge‐preferring species modifies the species–area relationship predictions of the island biogeography theory for an insect group (ground beetles, Coloptera: Carabidae) living in natural fragments. Species–habitat island area relationships applied to terrestrial habitat islands can be distorted by the indiscriminate inclusion of all species occurring in the fragments. Matrices surrounding terrestrial habitat fragments can provide colonists that do not necessarily distinguish the fragment from the matrix and can survive and reproduce there. Edge‐preferring species can further distort the expected relationship, as smaller fragments have larger edge:core ratios. Location Nineteen forest fragments were studied in the Bereg Plain, Hungary, and SW Ukraine. This area contains natural forest patches, mainly of oak and hornbeam, and supports a mountain entomofauna. Methods Ground beetles (Carabidae) present in the 19 forest patches were categorized into generalists, forest specialists and edge‐preferring species. We analysed the relationship between species richness and fragment area using species richness in the different categories. Results The assemblages contained a high share of generalist species (species that occur also in the surrounding matrix). Forest patch size and the number of generalist species showed a marginally significant negative relationship, indicating that generalist species were more important in smaller patches. Forest specialist species richness was correlated positively with patch area. Edge‐preferring species were shown to influence the species–area relationship: the number of edge‐preferring species increased with the edge:area ratio. Main conclusions Both generalist and edge‐preferring species can considerably distort the species–area relationship. Island biogeography theory can be applied to habitat islands only if the habitat islands are defined correctly from the viewpoint of the target species.     

小兴安岭森林地表鞘翅目成虫对不同干扰梯度的响应

为探讨小兴安岭凉水自然保护区森林生态系统中地表鞘翅目成虫群落对不同人为干扰梯度的响应,于2015年7月、8月和10月分别对轻度干扰[原始阔叶红松林(KY)和谷地云冷杉林(YL)]、中度干扰[阔叶红松择伐林(ZF)和次生白桦林(BH)]及重度干扰[落叶松人工林(RL)和红松人工林(RHS)]的6个林型进行取样调查。结果表明:(1)整个采样周期共捕获地表鞘翅目成虫879只,隶属9科44物种;其中轻度干扰生境共捕获6科29种251只(KY捕获5科21种150只,YL捕获4科20种101只),中度干扰生境捕获6科27种276只(ZF捕获3科20种144只,BH捕获6科23种132只),重度干扰生境捕获6科29种352只(RL捕获4科22种232只,RHS捕获5科17种120只)。(2)7月和8月步甲科和葬甲科占据数量优势,10月步甲科和葬甲科成虫数量明显减少而隐翅虫科数量占优势;不同林型及不同干扰梯度地表鞘翅目成虫物种总数和总个体数于7、8、10月均表现为下降趋势,且群落多样性也呈不同程度下降。(3)林型和月份对地表鞘翅目成虫群落结构具有显著影响,干扰梯度对群落结构无显著影响;6个林型之间地表鞘翅目成虫个体数量具有显著差异,但在物种组成上无显著差异;不同干扰梯度间地表鞘翅目成虫个体数、物种数无显著差异,且随干扰梯度变化没有明显的梯度性规律;相似性系数和聚类分析表明,属于同一干扰梯度的两个林型没有表现出高度的相似性。本研究表明干扰梯度不是决定各林型间地表鞘翅目成虫群落多样性存在差异的主要原因,凉水森林生态系统地表鞘翅目成虫对不同干扰梯度的响应不符合中度干扰假说,林型和时间则是影响地表鞘翅目成虫群落组成的显著因素,本实验为地表生物多样性保护和森林生态系统管理提供数据支撑。     

Red deer (<Emphasis Type="Italic">Cervus elaphus</Emphasis> Linnaeus, 1758) increasing density effects on species assemblage of ground beetles (Coleoptera: Carabidae) in Alpine forests

Red deer (Cervus elaphus) increasing density may trigger several modifications at the ecosystem level. Positive and negative impact of red deer on the invertebrate fauna is known in northern Europe, whereas it is currently unknown in the Alps. In this paper, we tested the impact of red deer density on ground beetles’ (Coleoptera: Carabidae) richness and species distribution in forests of the central-eastern Italian Alps (Stelvio National Park). Carabid beetles were sampled using pitfall traps in nine forest sites with different density of red deer. Carabid species richness was related to elevation, habitat type and red deer density in summer and winter using generalised linear mixed models and canonical correspondence analysis. The high winter red deer density affected positively the carabid species richness. Overgrazing seemed to increase presence of generalist species of dried and unshaded soils and decreasing of specialised species typical of well-preserved Rhododendro-Vaccinietum cover. Accordingly, the presence of patches with different red deer density seems to contribute to maintaining high levels of carabid beetles’ richness, albeit overgrazing needs to be monitored because it is likely to cause a local decrease of specialised forest species.     

Distribution of carabid beetles (Coleoptera: Carabidae) across a forest-grassland ecotone in southwestern China

This paper studied the occurrence of carabid beetles (Coleoptera: Carabidae) in the forest edge, the adjacent forest interior, and the surrounding grassland in southwestern China. Beetles were collected with pitfall traps along five replicated transects. Forest species rarely penetrated into the grassland from the forest interior, and the grassland specialists were not found in the forest interior. The forest edge hosted additional species from the adjacent grassland that increased its overall species richness. Nearly all forest species (23 of 24 species) and grassland species (13 of 15 species) can be found in the forest edge. Carabids of the forest edge were more similar to those of the forest interior than to those of the grassland by ordination and cluster analysis. Based on the specificity and fidelity, carabids can be distinguished into five species groups: habitat generalists, grassland-associated species, forest generalists, forest specialists, and edge-associated species. Multiple linear regression analysis showed that canopy cover and/or shrub cover were the most important factors in determining the richness, abundance, and diversity of carabids. The forest edge may serve as a transition zone for dispersal and re-colonization of carabid beetles from adjacent habitats and therefore is important for natural conservation.     

The fauna of the rove beetles (Coleoptera,Staphylinidae) in the southern Meshchera Lowland

The fauna of rove beetles of the sandur landscape in southern Meshchera (Ryazan Province), noted for the contrasting mesorelief forms, was studied for the first time. The regional fauna was found to comprise 178 species, of which 143 are listed for the first time. The core of the faunistic complex of rove beetles consists of Holarctic, Trans-Eurasian, and Euro-Siberian species, most of which commonly occur in the forest zone of European Russia. At the same time, some species distributed in the valley landscapes of Eastern Europe (Bledius tenenbaumi, Xantholinus dvoraki) have been recorded in southern Meshchera.     

Changes in Vegetation and Earthworm Populations under Free Grazing European Bison (<Emphasis Type="Italic">Bison bonasus</Emphasis>) in Broad-Leaved Forests of the Kaluzhskie Zaseki State Nature Reserve

This paper studies the impact of a free-ranging bison population on the vegetation and soil fauna in spatially adjacent biotopes (meadow-transition zone-surrounding forest) in the Kaluzhskie Zaseki State Nature Reserve. It is noted that bison maintains meadows in the absence of haymaking. Analysis of phytosociological relevés shows that the forest vegetation has been significantly transformed by bison over 10–15 years, as evident from (1) differences in the ecological-coenotic structure of the vegetation in the different biotopes, (2) a larger ecological-coenotic diversity of communities in the transition zones in comparison with surrounding forests, and (3) an increase in the participation of nitrophilous species in the vegetation. A fairly high earthworm species diversity (a total of seven species in the biotopes) is found; Aporrectodea caliginosa dominates in the earthworm density and biomass. The total earthworm density and biomass became lower in the transition zone than in neighboring forests and meadows, and the differences in earthworm density between the transition zone and the meadows were statistically significant.     

Species richness and zonal features of partial faunas of phytophagous beetles (Coleoptera,Chrysomeloidea, Curculionoidea) on grassy slopes in the east of the Russian Plain and the Cis-Ural Region

Comparative analysis of the fauna of phytophagous beetles (Chrysomeloidea, Curculionoidea) on grassy slopes in the east of the Russian Plain and in the Cis-Ural Region was performed. The species richness, taxonomic and chorological structure, and specificity of 22 partial faunas were assessed along the latitudinal gradient from the taiga to the steppe zone. The slope faunas are shown to have extrazonal features and to function as refuges of rare species. The species richness of slope communities of phytophagous beetles increases 2–3-fold (from 105–174 to 283–303 species) from the taiga to the forest-steppe, largely due to an increase in the number and share of representatives of the steppe landscape-geographic complex. The ratio of the number of species of Curculionidae to that of Chrysomelidae changes from 1: 1 to 1.5: 1.     

Biotopes and spatial organization of arthropod communities

Plant, spider, beetle, and ground beetle communities were studied in different agrolandscape biotopes (a wheat field, a field margin, and a forest belt) of Krasnodar Territory. In these biotopes, borders between plant communities were distinct. On the contrary, arthropod communities were not usually confined to the certain biotopes. The high degree of mobility of arthropod species resulted in a decrease in their association with the biotope. Representatives of some species (especially, beetles) were revealed in all the biotopes. In relation to season and some other factors, they frequently concentrated in different biotopes. Analysis of arthropod assemblages only within a single biotope gives us a rather impoverished knowledge of the whole population. It is necessary to study these assemblages not only within the basic biotope, but also in adjacent ones.     

Effects of forest fragmentation on species richness and composition of ground beetles (Coleoptera: Carabidae and Brachinidae) in urban landscapes

To clarify the effects of forest fragmentation in urban landscapes on the abundance, species richness, dominance, and species composition of ground beetles (Coleoptera: Carabidae and Brachinidae), we compared the beetles collected in 12 pitfall traps from April to July and from September to November between three continuous suburban forests and eight isolated urban forests (0.06–1.02 ha), most of which were in the precincts of shrines and temples in Hanshin District, Honshu, Japan. A total of 28 species and 4178 individuals of ground beetles were collected. Segregation of urban forests from continuous suburban forests has changed the species composition and resulted in the loss of some large‐sized forest species and the addition of some non‐forest species. Simpson's index of dominance (λ) also increased in the urban forests. The richness of forest species markedly decreased with the reduction in forest area but not with the distance from continuous forests, although the species richness of non‐forest species did not change with them. Also, species composition changed only with forest area. These findings indicate that continuous forests do not necessary serve as a “mainland” for urban forest species and that every urban habitat, however small in size, acts as a temporary reservoir of species. In comparison with populations of small‐sized species, populations of large‐sized forest species appeared to decline more readily during forest fragmentation.     

Introduced and native ground beetle assemblages (Coleoptera: Carabidae) along a successional gradient in an urban landscape

According to the intermediate disturbance hypothesis (IDH), species diversity should be higher at sites with intermediate levels of disturbance. We tested this hypothesis using ground beetles (Coleoptera: Carabidae) collected in pitfall traps from sites that varied in time since last disturbance. This successional gradient was embedded in an urban landscape near Montreal, Quebec. We predicted that diversity in young forests and old fields would be higher than in agricultural fields and old forests. Fifty-five species (2932 individuals) were found in 2003 and 46 species (2207 individuals) in 2004. In both years, species richness was highest from traps placed in agricultural fields. We collected nine introduced species; these had higher catch rates than the native species in both years (64.8% of total catch). When introduced species were removed from the Nonmetric Multidimensional Scaling ordination analysis, the assemblages from agricultural fields were less distinct compared to those of the other habitats, suggesting the introduced fauna is important in structuring carabid assemblages from the agricultural fields. Introduced species may play a significant role in the community composition of ground beetles in urban landscapes, and their influence may be the cause of the lack of support found for the IDH.