Dynamics of the abundance of cuckoo wasps (Hymenoptera, Chrysididae) in the conditions of the altitudinal-belt heterogeneity of the North Caucasus

The work provides data on the dynamics of the abundance of 205 taxa from 19 genera: Cleptes, Colpopyga, Elampus, Omalus, Philoctetes, Pseudomalus, Hedychridium, Hedychrum, Holopyga, Chrysidea, Chrysis, Euchroeus, Chrysura, Pseudochrysis, Spinolia, Spinolia, including species rare and new for the fauna of the North Caucasus and Russia. By the nature of the population dynamics, five groups of cuckoo wasps were identified: spring – represented by 7 species from 4 genera; spring-summer – 76 species from 10 genera; summer – 113 species from 16 genera; summer-autumn – 4 species from 2 genera and spring-summer-autumn – 5 species from 3 genus. According to the duration of flight, eurychronous species of cuckoo wasps were identified, which are found throughout the season from spring to autumn; mesochronous – the years of which affect the end of May and summer months and stenochronous – confined to a short period of time. Eurychronic accounted for 2.4%, mesochronous 24.9%, stenochronic 72.7%. The number of generations of cuckoo wasps is related to the dynamics of the number of the host. In the mountains, the peaks of activity shift towards the middle of summer, most species have one peak of activity; due to the frequent changes in weather conditions in the mountains, the phenological characteristics of the cuckoo wasps and their hosts do not coincide in terms with the lowland populations and the periods of activity can be extended in time. Phenological characteristics of rare and new species for the fauna of the North Caucasus and Russia are presented for the first time. The data obtained expand the understanding of the biodiversity and dynamics of the abundance of cuckoo wasps in the North Caucasus and can be used for environmental protection measures and rational nature management in the south of Russia.


Introduction
Cuckoo wasps (Chrysididae: Cleptinae, Chrysidinae) is a widespread group of hymenoptera; in the fauna of Russia there are 340 species and 13 subspecies from 23 genera [1]. Being entomophages, in biocenoses they can influence the number of their hosts -wild bees (Megachilidae) and some other wasps: Crabronidae; Vespidae (Eumeninae).
Information on the seasonal dynamics of flight and the phenological characteristics of the cuckoo wasps are not numerous and relate to the flat part of the Northwestern Black Sea region [2], the Interfluve of the Prut and Dniester [3] and Central Ciscaucasia [4]. For the first time, data on the dynamics of the abundance of wasp wasps in mountainous and foothill landscapes North Caucasus are presented. It is known that mountain landscapes have their own characteristics, where the parameters of the biota naturally change in all three dimensions of space, depending on the biological effect.
Taking into account the current state of the of the interaction of mountains and adjacent plains, factors of zoning in the plain and zonality in the mountains [5], taking into account the current state of the recreational load on the natural landscapes of the North Caucasus, and the expansion of monitoring studies, knowledge of the patterns of the dynamics of the abundance of components of biocenoses is important.
In this regard, new data on the dynamics of the population of cuckoo wasps, taking into account the altitudinal-belt structure of the landscapes of the North Caucasus, are relevant and can be used to solve practical and scientific issues.
The aim of this work was to study the dynamics of the abundance of cuckoo wasps (Hymenoptera, Chrysididae) in the conditions of the altitudinal-belt heterogeneity of the North Caucasus.

Materials and methods
The collection of material was carried out in 1987 -2020: in Ciscaucasia -Stavropol Territory, the main part of which is from the territory of the specially protected ecological-resort region of the Caucasian Mineral Waters; in the Western Caucasus -Krasnodar Territory, Utrish Nature Reserve, Caucasian Nature Reserve, Sochi National Park, in the Republic of Adygea, the vicinity of Maikop, Lagonaki Plateau; in the Central Caucasus -the Kabardino-Balkarian high-mountain reserve, the Elbrus National Park; in Karachay-Cherkessia -Teberdinsky reserve and its Arkhyz section; in Dagestan -the Dagestan nature reserve with its sites: Sarykum dunes, Kizlyar Bay and «Samurskiy Les» [6][7][8][9][10][11][12][13][14][15][16][17][18][19], also used materials from the collection of the Zoological Institute of the Russian Academy of Sciences (St. Petersburg).
For the collection of insects, the methods generally accepted in entomology were used: mowing with an entomological net, individual trapping and colored traps by Mörike [20][21][22]. The taxonomic position of species is given according to the system that has developed to date [23][24][25][26][27][28][29]. Zoogeographic regionalization of the Caucasus is given according to Sokolov, Tembotov [30].

Results and Discussion
Shiny wasps live in the nesting places of their owners, where they actively manifest themselves as entomophages, regulating their numbers. The paper provides data on the dynamics of the abundance of 205 taxa from 19 genera, including rare, poorly studied, and new species for the fauna of the North Caucasus.
The sustainable development of biocenoses is influenced by many factors, of which hydrothermal conditions and the presence of ecological niches for their inhabitants can be decisive. As shown by the research results, 205 taxa from 19 genera were recorded in the dynamics of the abundance of cuckoo wasps in the Northern Caucasus: Cleptes, Colpopyga, Elampus, Omalus, Philoctetes, Pseudomalus, Hedychridium, Hedychrum, Holopyga, Chrysidea, Chrysis, Euchroeus, Chrysura Spintarina, Stilbum, Trichrysis, Parnopes.
According to seasonal activity, 5 phenological groups were identified: spring, spring-summer, summer, springsummer-autumn and summer-autumn (tabl., fig. 1-3). We fully admit that in other variants of the zonation of landscapes on the territory of the North Caucasus, some species from this group may also occur in early summer. For example, the cuckoo wasp C. frivaldzkyi sparsepunctata was recorded for the first time in the North Caucasus, it is rare, and additional information on the dynamics of numbers can clarify its phenological characteristics. Chr. trimaculata has not been previously registered in the North Caucasus. One or two peaks of activity can be observed in the population dynamics. The species C. gracillima gracillima has 2 peaks (Fig. 1). The increase in numbers is observed at the end of May and reaches a peak at the beginning and at the end of June and gradually decreases by the middle of summer. The species C. comta has 2 peaks of activity, the maximum falls on the middle of July (Fig. 2).  By the nature of seasonal activity, five phenological groups of cuckoo wasps were distinguished: springrepresented by 7 species from 4 genera; spring-summer -76 species from 10 genera; summer -113 species from 16 genera; summer-autumn -4 species from 2 genera and spring-summer-autumn -5 species from 3 genus.
According to the duration of flight, eurychronous species of cuckoo wasps were identified, which are found throughout the season from spring to autumn; mesochronous -the years of which affect the end of May and summer months and stenochronous -confined to a short period of time. Eurychronic accounted for 2.4%, mesochronous 24.9%, stenochronic 72.7%.
The number of generations of cuckoo wasps is related to the dynamics of the host population. In the mountains, the peaks of activity shift towards the middle of summer, most species have one peak of activity; due to the frequent changes in weather conditions in the mountains, the phenological characteristics of the cuckoo wasps and their hosts do not coincide in terms with the lowland populations andthe periods of activity can be extended in time. Phenological characteristics of rare and new species for the fauna of the North Caucasus and Russia are presented for the first time.
The mountains have their own complex of species that rarely descend into the lower altitudinal belts, among them there are species that occur from May to August, such as Chr. ignifrons. The peak of C. viridula activity is in the middle of summer.
A new species for the fauna of Russia, Spintarina vagans, was noted by us only in the Central Caucasus at an altitude of 1000 to 2000 m above sea level.
The data obtained expand the understanding of the biodiversity and dynamics of the abundance of cuckoo wasps in the North Caucasus and can be used for environmental protection measures and rational nature management in the south of Russia.